Threatened Amphibians

Threatened Amphibians of the World

of the World
Simon Stuart
Michael Hoffmann
Janice Chanson
Neil Cox
Richard Berridge
Pavithra Ramani
Bruce Young

Simon Stuart, Michael Hoffmann, Janice Chanson, Neil Cox,

Amphibians are facing an extinction crisis, on a scale that scientists are only just beginning to understand.
Threatened Amphibians of the World is a visual journey through the first-ever comprehensive assessment of

Richard Berridge, Pavithra Ramani, and Bruce Young

the conservation status of the world’s 6,000 known species of frogs, toads, newts, salamanders, and caecilians.
Some 1,900 species known to be threatened with extinction are covered, including a description of threats to
each species and an evaluation of conservation measures in place or needed. Each entry includes a photograph
or illustration of the species where available, a distribution map, and information on range, population and habitat
and ecology. Introductory chapters present a detailed analysis of the results, complemented by a series of short
essays written by many of the world’s leading herpetologists. Appendices include annotated lists of lower risk
species and a country-by-country listing of threatened amphibians.
Threatened Amphibians of the World presents a snapshot in time of the dire status of an entire class of organisms,
and in so doing poses searching questions about the health of our environment and what we are going to do
about it.
THREATENED AMPHIBIANS OF THE WORLD
S.N. Stuart, M. Hoffmann, J.S. Chanson, N.A. Cox, R.J. Berridge, P. Ramani, and B.E. Young (eds.)

THREATENED AMPHIBIANS OF THE WORLD

Cover: Hylomantis lemur, Endangered. © Joel Sartore / www.joelsartore.com
Back cover: Agalychnis callidryas, Least Concern. © Kenji Nishida
Page 1: Ichthyophis kohtaoensis, Least Concern. © Danté Fenolio
Page 135: Atelopus ignescens, Extinct. © Michael and Patricia Fogden
Page 145: Leptopelis vermiculatus, Vulnerable. © Maik Dobiey
Page 609: Epipedobates bassleri, Near Threatened © Maik Dobiey

Recommended citation:
Stuart, S.N., Hoffmann, M., Chanson, J.S., Cox, N.A., Berridge, R.J., Ramani, P., and Young, B.E. (eds.) (2008). Threatened Amphibians of the World. Lynx Edicions, Barcelona, Spain;
IUCN, Gland, Switzerland; and Conservation International, Arlington, Virginia, USA.

Published as a partnership between IUCN, Conservation International and Lynx Edicions.

First edition: July 2008

© Lynx Edicions – Montseny 8, 08193 Bellaterra, Barcelona (Spain)

© Texts: introductory matter: authors / IUCN and Conservation International; species accounts: IUCN, Conservation International and NatureServe
© Photographs: credited photographers

Printed by Ingoprint S.A.

DL: B-32.689-2008
ISBN: 978-84-96553-41-5

All rights reserved. No form of reproduction, distribution, public communication or transformation of this work may be carried out without the authorization of its copyrights holders, except that foreseen by the law. Those
needing to photocopy or electronically scan any part of this work should contact Lynx Edicions.
 Threatened Amphibians of the World is dedicated to

The 500 herpetologists from around the world

who devoted their knowledge, intellect and time
to the Global Amphibian Assessment, and without
whom this book could not have been written

and especially to

George Rabb for his visionary leadership

and commitment to confronting the
amphibian extinction crisis, which gives us
hope in an otherwise bleak situation.

Dendrobates azureus, Vulnerable. © Albert Popalis

 CONTENTS

FOREWORDS IX

PREFACES XI

ACKNOWLEDGEMENTS XIII

INTRODUCTORY CHAPTERS 1
Chapter 1. An Introduction to the Amphibians 2
Chapter 2. Why Save Amphibians? 23
Chapter 3. The Global Amphibian Assessment (GAA): History, Objectives and Methodology 30
Chapter 4. The State of the World’s Amphibians 33
Chapter 5. Amphibians of the Afrotropical Realm 53
Chapter 6. Amphibians of the Australasian Realm 65
Chapter 7. Amphibians of the Indomalayan Realm 74
Chapter 8. Amphibians of the Nearctic Realm 85
Chapter 9. Amphibians of the Neotropical Realm 92
Chapter 10. Amphibians of the Palaearctic Realm 106
Chapter 11. Amphibian Conservation – Responding to the Global Decline of Amphibians 114

EXTINCT AND EXTINCT IN THE WILD AMPHIBIAN SPECIES 135

GLOBALLY THREATENED AMPHIBIAN SPECIES 145

NEAR THREATENED AMPHIBIAN SPECIES 609

BIBLIOGRAPHY 645

APPENDICES 693
Appendix Ia. The IUCN Red List Categories (IUCN 2001) 693
Appendix Ib. Summary of the five criteria (A-E) used to evaluate if a species belongs
in a Category of Threat (Critically Endangered, Endangered or Vulnerable) 693
Appendix II. Definitions for the initial data collection regions 694
Appendix III. A summary of IUCN Red List status by genus 694
Appendix IV. List of rapidly declining species 696
Appendix V. Species listed by territory 699
Appendix VI. Amphibian Conservation Summit Declaration:
an Amphibian Conservation Action Plan 702
Appendix VIIa. The scale of conservation required for globally threatened amphibians 703
Appendix VIIb. The nature of the ecological process for which
landscape-scale conservation action is required 723
Appendix VIII. List of Critically Endangered and Endangered species for which
captive breeding is an immediately necessary ex-situ conservation action 725
Appendix IX. Amphibians listed as Critically Endangered (Possibly Extinct)
on the 2007 IUCN Red List of Threatened Species 727
Appendix X. Amphibians listed as Vulnerable D2 on the 2007 IUCN Red List
of Threatened Species 728
Appendix XI. List of Least Concern species listed on the 2007 IUCN Red List
of Threatened Species, with country of occurrence 729
Appendix XII. List of Data Deficient species, with country of occurrence 739
Appendix XIII. The differences in IUCN Red List Status for some Brazilian species 744
Appendix XIV. Glossary of some key terms used 745
Appendix XV. Websites and amphibian-related resources 745

INDEX OF EXTINCT, THREATENED AND NEAR THREATENED

SPECIES ACCOUNTS 747

FOREWORDS
The first signs of trouble for amphibians arose around three decades ago. IUCN, Conservation International and
NatureServe made great efforts to understand what was happening, and in 2004, the Global Amphibian Assessment
was produced with this shocking result: one-third of amphibian species are threatened with extinction. In addition,
we were lacking data on many species which are potentially threatened. New studies show that many of the species
that are listed as ‘Data Deficient’ on the IUCN Red List of Threatened Species™ are indeed threatened. One-third of
an entire distinct group of species at risk of extinction is not a trivial figure.
This book is another step in our attempt to alert the world about the extinction crisis that toads, frogs, salamanders
and caecilians are currently facing. It is a landmark book comprising all what needs to be known about amphibian
conservation, and as such I am very happy to see its publication and to write this foreword. I would, however, be
even happier if the amphibian extinction crisis was not taking place. I am concerned that the size of this book is a
reflection of the number of threatened species that it comprises. When reading it one should not forget that only
threatened amphibians are included in the species accounts that occupy the bulk of its length.
Many of us probably do not give much attention to amphibians and will probably be surprised to see how diverse,
unique, colourful, and fascinating they are. Many species have developed very specific evolutionary characteristics
and adaptations. We might think that they all lay eggs in water, but some give birth to live young, some do not need
water, some species carry their eggs, others carry their young. Two Australian frogs incubated their eggs in their
stomachs and both are already Extinct. As a result, an entire evolutionary lineage has been lost, along with up to
164 other species. These reproductive traits are the results of millions of years of evolution. Who knows if such
adaptations would not have been key to the survival of species in response to climate change?
Amphibians are sending us a very strong signal that something is seriously wrong with the way we take care of
our environment. They have been considered as the ‘canaries in the coal mine’; their permeable skin makes them
sensitive to many forms of pollution; they are also extremely sensitive to climate change and diseases. The sharp
decline in amphibian populations could prefigure, if we are not careful, what other groups of species might undergo
in the future. Amphibians provide us with a critical challenge if we are to meet the target that all countries agreed
upon: a significant reduction in the rate of biodiversity loss by 2010.
Amphibians are a key part of ecosystems and the food chain; they control the number of many invertebrates, and
are prey for many species of birds or mammals. Some species are also a source of revenue for human communities
who eat them, use them as medicines, or supply them to the international pet trade. The skins of certain species
contain molecules that are very promising in terms of developing new medicines for people. Who knows, a little frog
hiding in the understorey of a tropical rainforest might bring the solution to the AIDS pandemic?
After reading this book and contemplating all these pictures, no one will look at amphibians with the same eyes
again.
Julia Marton-Lefèvre
Director General
IUCN, the World Conservation Union
Eight years ago, I was invited to write a foreword to BirdLife International’s landmark publication Threatened Birds
of the World. At the time, I commented how encouraging it was that at least one organization had managed to hone
its focus down to the level of sites and species across the planet. Writing this foreword for what is, in essence, the
sister publication to BirdLife’s earlier work, I am struck by our progress since then.
In 2003, at the Fifth World Parks Congress in Durban, South Africa, the results of a global analysis revealed the
tremendous gaps in coverage of the existing protected areas network. While not all threats to biodiversity can be
mitigated via habitat protection, habitat loss remains the overwhelming threat to our planet’s unique life forms. It is
encouraging, then, to see the expansion of the Important Bird Areas concept, pioneered by BirdLife International, to
all taxa under the umbrella of the Key Biodiversity Areas approach, which aims to identify globally important sites
for the conservation of biodiversity. Such initiatives, aided by the efforts of the Alliance for Zero Extinction, which
prioritizes that subset of sites known to hold the last remaining population of a highly threatened species, provide
conservation planners, funders and practitioners with a valuable lens on where, and what form of, conservation action
is most appropriate in order to avoid the loss of global biodiversity.
Such fine-scale resolution would be impossible without the foundation provided by the data in the IUCN Red List,
the recognized authority for the conservation status of global biodiversity. Before the launch of the Global Amphibian
Assessment in September 2004, only a few hundred hand-picked frogs, toads and salamanders appeared on the
IUCN Red List. Today, we have detailed, up-to-date information on the distribution, population status, threats, and
conservation status of nearly 6,000 amphibians.
Threatened Amphibians of the World is a visual exploration of the results of the Global Amphibian Assessment
– a compelling synthesis of herpetological knowledge and a yardstick for amphibian conservation. In tandem with the
Amphibian Conservation Action Plan, produced by the IUCN/SSC Amphibian Specialist Group, it puts the amphibian
agenda center-stage. And it does so with tremendous visual impact, presenting us with a face on an unfolding crisis
like nothing that has gone before.
The plight of the world’s amphibians is not just theirs, but ours as well. We should care that one-third of amphibians
stand a high risk of extinction in the wild – and probably many more – because the threats that imperil them are those
that stem from our actions and will be no less detrimental to our own livelihoods: pollutants, invasive species and
disease, global climate change, and rampant habitat loss. Responding to these threats must be a societal concern.
Let us hope that a second edition of this book does not require two volumes…
Russell A. Mittermeier
President, Conservation International
Steering Committee member, IUCN Species Survival Commission
IUCN Council member 2004-2008, IUCN – World Conservation Union
IX
“Round up the usual suspects,” as Captain Renault famously said in the film Casablanca, did not seem to apply. Some-
thing appeared to be going on with amphibians that could not be wholly explained by the standard litany of ecological
threats, such as habitat degradation and loss. The declines that were first noticed in earnest in the late 1980s and
early 1990s raised the possibility that something was seriously amiss: a crisis that might start with environmentally
sensitive creatures such as amphibians, but ultimately affect other strands of life, including humans.
NatureServe is a non-profit organization dedicated to providing the scientific basis for effective conservation
action, with a deep appreciation for the role that research plays in addressing environmental challenges. We were
pleased to join with IUCN and Conservation International in carrying out the Global Amphibian Assessment (GAA) as
a way to document and draw attention to the amphibian declines now underway.
NatureServe scientists began seriously delving into the plight of amphibians through our coordination of RANA
(Research and Analysis Network for Neotropical Amphibians), a U.S. National Science Foundation-funded effort.
Through RANA, researchers with an interest in understanding amphibian declines in Latin America came together to
share information, approaches, and results, and to train a new generation of Latin American herpetology students.
The project has also helped to document some of the causes that appear to be behind these declines, including fungal
diseases and shifts in climate. Building on our work with RANA, as well as our long history of assessing conservation
status for North American and Latin American species, I am pleased that NatureServe was able to contribute to the
GAA by coordinating the Western Hemisphere portion of the initiative, and creating a website for disseminating the
project’s data and results to the broader community (www.globalampihibians.org).
The GAA partnership has succeeded admirably in bringing the dire condition of the world’s amphibian species into
focus, and in motivating the scientific and conservation communities to take action. The present volume provides an
important summary of what is known about amphibians around the world, and what will be required to ensure their
future. While the work to do so will be difficult, and success is not assured, only by committing to the approaches
outlined in the recent Amphibian Conservation Action Plan will we have a hope for preserving the many and varied
faces of this ancient lineage.
Mary L. Klein
President and CEO, NatureServe
Amphibians, be they frogs, toads, salamanders or caecilians, are thought to be amongst the most sensitive ecologi-
cal indicators, warning us of unsafe environmental conditions that are not only impacting our health, but that of
the global ecosystem. With at least 32% of the described amphibian species threatened with extinction, and 500
species probably beyond the point of no return, the global amphibian extinction crisis represents one of the greatest
conservation challenges of our times.
A direct result of mankind’s insatiable appetite for natural resources, the rapid decline in amphibian numbers
demonstrates the sensitivity of not only species, but of entire ecosystems. The current crisis exemplifies how, if
unmanaged, actions of every-day human consumption and waste can compound to form ripple-like reactions resulting
in entire populations disappearing, and all within our lifetime. In 2004, a global collaboration of work from 500 of the
world’s most respected amphibian specialists from over 60 nations was released: the Global Amphibian Assessment.
Citing the possible loss of 122 species since 1980 alone, this unprecedented collaborative research effort highlighted
the pace of modern amphibian declines. Since the first release of the Global Amphibian Assessment, there has been
constant updating of information, including the addition of 160 new species to the IUCN Red List of Threatened
SpeciesTM nine of which are Critically Endangered, 17 Endangered, and 12 Vulnerable. However, a large percentage
of species remain in the Data Deficient category, emphasizing the need for more amphibian assessment work.
At present, little is known of the deadly fungal disease that is wiping out some amphibians faster than can be
recorded. With fears that this could worsen with global climate change, there is no better time than the present to
act with the knowledge we have, improve shortcomings in our knowledge, and continue work with our current and
future collaborative efforts to raise the awareness for amphibian conservation. The recent release of the Amphibian
Conservation Action Plan by IUCN/SSC’s Amphibian Specialist Group and the publication of this book, are tributes to
the enormous efforts dedicated teams are undertaking to tackle the global loss of species. Onwards to 2008, aptly
named “the Year of the Frog”.
Holly T. Dublin
Chair, IUCN Species Survival Commission

PREFACES
Imagine a world without frogs. No spring choruses of calling males. No tadpoles for kids to play with. No brightly
coloured jewels perched on rocks and leaves throughout the world’s tropical forests. We are currently witnessing the
global extinction of amphibians – they are facing extinctions at levels unprecedented in any other group of organisms in
human history. In light of this, we have some serious questions to address: How extensive is the amphibian extinction
crisis? How much worse can it get? Why is this happening now? Will it affect humans? Is there any hope?
As early as 1989, researchers exchanged anecdotal reports of declines and disappearances of amphibian species
from many areas around the world. Species, such as Costa Rica’s Golden Toad (Bufo periglenes), went missing from
pristine protected areas, often in the absence of familiar threats such as habitat loss and pollution (e.g., Crump et al.
1992). Further complicating matters, data on species distributions and population fluctuations were lacking for most
amphibian species, thus preventing our ability to make accurate estimates of endangerment. In response, the IUCN
Species Survival Commission formed The Declining Amphibian Population Task Force (DAPTF; and see Essay 11.1) to
guide, sponsor, and fund crucial research into the phenomenon of global amphibian declines.
There are currently (as of the time of writing) just over 6,000 recognized species of amphibians and this number
has increased by 48% in the past 20 years (Frost et al. 2006). This reflects both increased access to, and ease of
collecting data in, remote locations, as well as the growth of active scientific communities in a few megadiverse
countries (see Essay 1.1). Unfortunately, this increase in knowledge of amphibian diversity is offset by massive
population declines and growing numbers of species extinctions (Stuart et al. 2004; Lips et al. 2005). Simply
put, amphibian species are disappearing from the planet perhaps faster than we can discover them. According
to the results of the Global Amphibian Assessment (GAA; Stuart et al. 2004), some 43% of amphibian species
are declining in abundance, 32% of amphibians (one-third of all species) are threatened with extinction, and 122
species likely have become extinct since 1980 (with 34 species confirmed Extinct). Worse yet, these numbers are
certainly underestimates because 23% of all known species are so poorly known that their threat status could not
be classified at all. Statistics of this sort represent a near-categorical extinction event perhaps on the scale of the
disappearance of the dinosaurs.
The GAA also reviewed causes of declines, and identified familiar threats such as habitat loss, chemical con-
tamination, and over-harvesting as being involved in some cases. However, the terrifying realization was that novel
threats related to emerging infectious diseases and climate change accounted for many of the disappearances.
Conservationists have worked for decades to limit habitat loss, pollution, and over-harvesting in natural areas, but
our current well-intentioned conservation programmes are insufficient to prevent extinctions from such insidious
threats as emerging infectious diseases and climate change.
Amphibian declines and extinctions are real and the recently discovered chytrid fungal pathogen (Batrachochytrium
dendrobatidis) is a primary cause of many catastrophic losses (Lips et al. 2006). We can detect the pathogen using
a non-lethal skin swab (Annis et al. 2004), predict its potential ecogeographical distribution (Ron 2005), predict local
movement, and foresee the catastrophic effects on local amphibian faunas (Lips et al. 2006). Species most threatened
with disease-induced extinction are those in high-elevation, riparian habitats, with small geographic ranges (Lips
et al. 2003). High endemism of amphibians in upland areas throughout the tropics suggests that climate changes
with or without associations with chytrid may also be devastating amphibians. The scope of the amphibian crisis is
massive, and the local and global effects of these losses are numerous and widespread (Whiles et al. 2006). When
amphibians disappear, we lose their roles in maintaining ecosystem function, and also any potential benefits they
may offer to humans.
Important questions remain: How can ecosystems continue to function in the absence of amphibians? What can
be done to preserve some amphibian species for future generations? How can we stop these extinctions? Research
and conservation organizations do not have the answers to these questions at this time. Nevertheless, a large-scale
commitment to truly maverick, but coordinated, conservation efforts is in order (Mendelson et al. 2006). Because
multiple causes are involved, diverse and innovative solutions must be implemented. For those species affected by
habitat loss, we need to continue current efforts aimed at safeguarding important sites. Understanding direct and
indirect effects of chemicals on amphibians requires extensive additional levels of attention, as do contaminant-
monitoring programmes. While over-harvesting does not affect most amphibian species, it represents a real problem
for some, such as the Asian giant salamanders (Andrias spp.; see Essay 4.5). Unfortunately, effective solutions to
global problems, such as climate change and emerging infectious diseases, do not currently exist. Responses to
threats related to climate change will require global commitment to alternative fuels and reduced emissions, and
reconfiguring protected areas to encompass broad elevational ranges, in order to account for potential shifts in spe-
cies distributions. Solutions to emerging infectious diseases will require significant levels of funding and aggressive
research to better understand the epidemiology and pathology of the diseases, and to develop vaccines and/or
resistance in wild populations.
Creating and implementing global solutions to these global threats requires significant levels of commitment,
organization, funding, and – especially – time. In cases of predicted catastrophic losses, the only conservation tool
we have at our disposal is the establishment of ex-situ survival assurance colonies (Mendelson and Rabb 2006).
Captive programmes are not a solution to the actual problems, but they may preserve living colonies of individuals,
increase numbers through breeding, and potentially serve as research colonies to enable timely studies of pathology
and evolution of resistance. The challenge again is a matter of scale: there are currently fewer than 40 amphibian
species in managed ex-situ programmes (Zippel 2005), yet at least five times that number require urgent ex-situ
conservation action (Appendix VII). These numbers represent a major challenge for zoos, gardens, and aquariums
(see Essay 11.5).
Many species need immediate attention to prevent their extinction, and potentially thousands of others remain to
be discovered or described. It is not uncommon to encounter several new species per trip to certain regions of Latin
America, especially upland areas, which are most affected by threats related to climate and disease. As a result, we
are losing species we never knew existed. Among the current authors, JRM has described about 40 new species of
Latin American amphibians in the past 15 years, but already half of them are likely extinct. Similarly, KRL has observed
the annihilation of the amphibian faunas at five different sites in Latin America in the past 15 years. Of course, we
hope our experiences are the exceptions, or even inaccurate, but we’re very afraid that they are close to reality.
Karen R. Lips and Joseph R. Mendelson III
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The Global Amphibian Assessment sharpened the scientific community’s focus on both the nature and extent of threats
to amphibians worldwide. Now is the time to act on new knowledge regarding the causes of an ongoing amphib-
ian extinction event. Clear and internationally coordinated options for thwarting further extinctions of threatened
amphibians must be developed with parties capable of implementing actions. The Amphibian Conservation Summit
was called in September 2005 to design and promote a response to this global crisis. To this end, the participants
in the summit and now the newly formed IUCN/SSC Amphibian Specialist Group (ASG) commends the Amphibian
Conservation Action Plan (ACAP) to governments, the business sector, civil society, and the scientific community for
urgent and immediate adoption and implementation.
Participants in the Amphibian Conservation Summit set priorities for conservation and research actions within 12
areas relevant to amphibian conservation: 1) Designing a network of conservation sites for amphibians – Key Biodi-
versity Areas; 2) Freshwater resources and associated terrestrial landscapes; 3) Climate change, biodiversity loss, and
amphibian declines; 4) Infectious diseases; 5) Over-harvesting of amphibians; 6) Evaluating the role of environmental
contamination in amphibian population declines; 7) Captive programmes; 8) Reintroductions; 9) The continuing need
for assessments, making the Global Amphibian Assessment an ongoing process; 10) Systematics and conservation;
and 11) Bioresource banking efforts in support of amphibian conservation. The ACAP embodies this work, although
we recognize that additional themes may need to be addressed in future versions of the document.
A declaration (Appendix VI) was released following the Summit urging four kinds of intervention that are needed
to conserve amphibians, all of which should be initiated with immediate effect: 1) Expanding our understanding of
the causes of declines and extinctions; 2) Continuing to document amphibian diversity, and how it is changing; 3)
Developing and implementing long-term conservation programmes; and 4) Responding to emergencies and immediate
crises. The full text of the Amphbian Conservation Summit Declaration and ACAP is at (www.amphibians.org).
The amphibian extinction crisis requires a global response at an unprecedented scale. The ACAP requires the
international community to enter uncharted territory and to take great risks. But the risks of inaction are even
greater. The ACAP calls on all governments, corporations, civil society, and the scientific community to respond.
There needs to be unparalleled commitment to developing and implementing the ACAP with accompanying changes
in international and local environmental policies that affect this class of vertebrate animals – as they truly are the
proverbial canaries in the global coal mine. This document offers practical, large-scale, creative, innovative and
realistic actions that will be required to halt the present tide of extinctions of amphibian species and includes an
ambitious yet realistic budget.
A unified global strategy incorporating survival assurance colonies, disease research, and habitat protection forms
the focus of this new plan to save amphibians. We must, of course, also remain vigilant and act on other threats,
including climate change, over-harvesting, and toxins. Lessons learned from confronting the amphibian crisis will be
transferable to other groups and ecosystems. We have many other potential crises-in-the-making, such as coral reef
collapses, fisheries collapses, emerging human diseases such as Ebola, SARS (Severe Acute Respiratory Syndrome),
Nipah virus, and our poor record of managing freshwater resources that will certainly lead to global shortages of
clean drinking water with great negative consequences. Many, if not all, of these environmental challenges are the
result to varying degrees of the same human footprint that our species is leaving on this planet.
The road to success must include a broad set of stakeholders who help implement the ACAP. This is important
because there are many issues that are beyond the simple realm of “amphibian conservation work.” Addressing
many of the underlying causes of this crisis will help us avert the next global environmental challenge. Helping curb
unsustainable wildlife use would not only decrease some of the threats to particular amphibian species, but also
help us apply these same solutions to other species. Similarly, tackling climate change, although a huge task in its
own right, will make a major contribution to the continued survival of all species as well as to the sustainability of
ecosystems in general.
Claude Gascon
Co-Chair IUCN/SSC Amphibian Specialist Group Secretariat
Executive Vice President, Conservation International, Virginia
James P. Collins
Co-Chair IUCN/SSC Amphibian Specialist Group Secretariat
Professor, School of Life Sciences, Arizona State University, Arizona

ACKNOWLEDGEMENTS
This book is a product of the Global Amphibian Assessment, and therefore owes everything to those who contributed
directly to that process, and indeed who continue to do so. In particular, this product would not have been possible
without the expertise, knowledge and enthusiasm of the more than 500 herpetologists who contributed their time
to the data collation and review process. This book is a testimony to the dedicated efforts of these individuals and
their unstinting passion for amphibian conservation.
Although we once again express our sincere thanks to the many individuals that contributed to the GAA below, it
is also necessary here to extend particular thanks to those who helped make publication of Threatened Amphibians
of the World a reality. Undertaking and producing a book of this scope has relied greatly on the dedicated help and
willingness of a very large number of people, as borne out by the acknowledgements that follow.
Firstly, as with any enterprise of this nature, the publication of Threatened Amphibians of the World would not
have been possible without the support of Conservation International providing generous financial support, in large
part through the Gordon and Betty Moore Foundation, to IUCN – The International Union for the Conservation of
Nature to cover staff time in the Biodiversity Assessment Unit. In particular we thank Gustavo Fonseca and Russell
Mittermeier for their courage and vision in making this happen.
It is necessary to extend a special word of thanks to several individuals for their help, time and advice in the plan-
ning and production of this book. Ali Stattersfield, Stuart Butchart and Martin Sneary provided initial useful insights
into the production of this book, based on their own experiences producing its sister publication, Threatened Birds
of the World. Our fellow staff in the IUCN Species Programme, particularly Jane Smart, Jean-Christophe Vié and Jim
Ragle, provided much needed network support, and often had to shoulder the burden on other projects while we were
editing amphibian species accounts. We trust they are as happy to see this project completed as we are! In particular,
we take this opportunity to single out the heroic efforts of the staff in the IUCN Red List Unit, Craig Hilton-Taylor and
Caroline Pollock, for their efforts in ensuring the quality of the data feeding into the IUCN Red List. Vineet Katariya is
thanked for her assistance with a number of GIS-related tasks and analyses, including producing Figures 7 and 10 in
Chapter 4. We also extend thanks to The Shared Earth Foundation and its President and CEO, Caroline D. Gabel, and
to George Meyer and Maria Semple, for making valuable contributions to the printing costs of this book.
At Conservation International (CI), the real herpetologists, Don Church, Claude Gascon and Robin Moore, provided
much valued advice and suggestions and also wrote several essays and contributed their expertise to several chapters.
Conservation International provided valuable network support through the entire duration of the project, and we are
particularly indebted to a number of current and former staff in CI’s Center for Applied Biodiversity Science for ideas,
fruitful discussions and help in other ways, including: Charlotte Boyd, Daniel Brito, Thomas Brooks, Naamal de Silva,
Graham Edgar, Matt Foster, David Knox, Thomas Lacher, Penny Langhammer, John Pilgrim, Ana Rodrigues, Will Turner
and Peter Paul van Dijk. Noura Bakkour kindly helped with translations of several materials from French to English, and
vice versa. Noura and Sarah Wyatt provided support with several cumbersome administrative tasks. Kellee Koenig
provided valuable cartographic help with the production of the maps for essays 7.2 and 11.3.
Our thanks, too, to Julia-Marton Lefèvre, Russell Mittermeier, Mary Klein and Holly Dublin for agreeing to write
the forewords, and to Jim Collins, Claude Gascon, Karen Lips and Joseph Mendelson III for writing the prefaces.
A number of colleagues kindly agreed to review drafts of the introductory chapters, and their comments and
input helped improve the overall content and accuracy of these chapters immeasurably. In particular, we would like
to thank: Darrel Frost, Taran Grant, Ron Heyer, David Wake and George Zug (Introduction to Amphibians); Teresa
Mulliken (Why Save Amphibians?); and Ariadne Angulo, Charlotte Boyd, Peter Riger, Lee Hannah and Michael Wai
Neng Lau (Amphibian Conservation). Several people also provided useful inputs into the regional chapters, and they
are acknowledged accordingly at the end of each chapter. Notwithstanding these valued and important inputs, we
stress that any errors in fact or judgment remain ours alone. The editors would also like to thank all the authors of the
regional chapters for their hard work, for keeping to the deadlines, and for their constructive comments in reviewing
the drafts – we hope that the final product meets the approval of all!
In addition to the regional chapter authors, several people devoted time to writing essays to draw attention to
particular issues or to highlight particular themes or trends. We wish to extend our thanks to the following for their
excellent contributions:
Alan Channing, Alex Ngari, Ana S.L. Rodrigues, Andrés R. Acosta-Galvis, Andrew A. Cunningham, Andrew Turner,
Angel Alcala, Annika Hiller, Arvin Diesmos, Ben D. Bell, Bhoopathy, Bryan l. Stuart, Catherine H. Graham, Célio F.B.
Haddad, Christopher J. Schneider, Claude Gascon, Claude Miaud, Claudia Corti, David B. Wake, David J. Gower, David
Knox, Débora Leite Silvano, Djoko T. Iskandar, Don R. Church, Enrique La Marca, Esteban O. Lavilla, Feng Xie, Franco
Andreone, Frank Glaw, Franky Bossuyt, G.K. Bhatta, Gabriela Parra-Olea, Geoffrey A. Hammerson, Gerry Marantelli,
Gracia Syed, Graeme R. Gillespie, Herilala Randriamahazo, Ian G. Bride, Ignacio De la Riva, Indraneil Das, Ismail Ugurtas,
James Hanken, James P. Collins, Jean-Louis Amiet, Jeanne E. McKay, Jelka Crnobrnja Isailovic, Jen M. Germano,
Jodi J.L. Rowley, Johannes Penner, John D. Lynch, Jörn Köhler, Joseph C. Mitchell, Joseph R. Mendelson III, Julián
Faivovich, Justin Gerlach, Karen R. Lips, Kelum Manamendra-Arachchi, Kevin Buley, Kevin Zippel, Kim Howell, Louise
A. Rollins-Smith, Madhava Meegaskumbura, Maj de Poorter, Mark Wilkinson, Mark-Oliver Rödel, Martin Wegmann,
Masafumi Matsui, Matt Foster, Melizar V. Duya, Michael J. Tyler, Michael Wai Neng Lau, Michele Menegon, Michelle
S. Koo, Miguel Vences, Mirza D. Kusrini, Naamal de Silva, P. Joël Adeba, Pablo A. Menéndez-Guerrero, Penny Lang-
hammer, Peter Daszak, Peter Paul van Dijk, Petros Lymberakis, Philip J. Bishop, Rachunliu G Kamei, Rafael Marquez,
Rafe M. Brown, Raffael Ernst, Raoul H. Bain, Richard A. Griffiths, Richard Gibson, Rick Relyea, Robert C. Lacy, Robert
F. Inger, Robert Johnson, Robert Puschendorf, Robin D. Moore, Robinson Mugo, Rohan Pethiyagoda, Ross A. Alford,
Ruth Grace Ambal, S.D. Biju, S. Blair Hedges, S. Gilles A. Nago, Santiago R. Ron, Sherif Baha el Din, Stefan Lötters,
Steve Richards, Susanne Schick, Tahar Slimani, Taran Grant, Thomas S.B. Akre, Tim Davenport, Tim Halliday, Vance T.
Vredenburg, Varad Giri, Vineet Katariya, Wang Xiaoming, Wolfgang Böhme, Yehudah Werner, and Zhang Ke-jia.
Don Church and Kevin Zippel led the valuable analysis of Appendix VIII that forms part of Chapter 11; Appendix
VIIa and VIIb are products of an analysis by Charlotte Boyd and colleagues recently published in the journal Con-
servation Letters.
That this book is anywhere near as colourful as we intended is largely a tribute to the 350-odd people and
institutions who generously provided or made available use of photographs of threatened amphibians to bring the
species accounts to life, as well as those persons who provided images for use elsewhere in the book. We received
many more images than we could use, and in some cases we had difficult decisions to make regarding which image
would be most appropriate. For anyone who contributed a photograph hoping that it would be used, and who finds
this not to be the case, we hope you will not hold this against us! We have not been successful in securing images
for every threatened species. That we were able to secure as many as we did, owes a great deal to the generosity
and understanding of those who helped out. We would especially like to thank Esteban Lavilla, Vanessa Verdade,
Simon Chan, Andres Charrier, Federico Bolaños, Jahson Alemu, and Sergio Potsch de Carvalho e Silva for their help
in sourcing photographs from friends and colleagues.
Despite having been meticulous in the copyrighting of photographs, we apologize for any errors and omissions
regarding photo credits and hope that any instances in this regard are minimal. We emphasize that the identification
of the species in the photographs is the responsibility of the photographers, and we claim no expertise of our own in
ensuring that these are correct. The names of the photographers are provided alongside each photograph, and are not
listed here. However, we must mention the following people who provided particularly large numbers of images that
were used in the book: Jean-Louis Amiet, Franco Andreone, Marion Anstis, César Luis Barrio-Amorós and Fundación
AndígenA, S.D. Biju, Thomas Bille, Rafe Brown, Tim Burkhardt, Ignacio de la Riva, Bill Duellman, S. Blair Hedges,
Frank Glaw, K. Jayaram, Twan Leenders, John Lynch, Kelum Manamendra-Arachchi, Madhava Meegaskumbura,
Rohan Pethiyagoda, and Miguel Vences.
XIII
Throughout the compilation of Threatened Amphibians of the World, we have been collaborating with ARKive
– www.arkive.org, to source images of threatened amphibian species in order to illustrate the book and increase the
number of species profiled by ARKive. ARKive is a not-for-profit initiative of Wildscreen, a UK-based charity, gathering
together films and photographs of the world’s threatened fauna and flora into one centralized digital library. ARKive
is leading the ‘virtual’ conservation effort, compiling comprehensive and enduring multi-media species profiles for the
16,000-plus species threatened with extinction, according to the IUCN Red List of Threatened Species. We encourage
any photographers who would like to make their images of threatened species available for educational and awareness
raising purposes to contact ARKive directly. We would also like to say a special thank you to Michelle Lindley, ARKive
Media Researcher, for her enthusiasm and assistance in locating, cataloguing and ensuring the smooth passage of
images between the photographers, ARKive and the GAA; and to Harriet Nimmo, Wildscreen Chief Executive, for
suggesting this exciting and valuable collaboration, and for eagerly supporting it.
Doan Nguyen provided tremendous support by helping scan many slides that were sent to us for use in the book,
while David A. Sanchez took on the formidable task of going through the immense slide collection of John Lynch
and scanning slides of threatened species for the book. Breck Bartholomew, at Bibliomania, kindly made available
high-resolution scans of the images of Craugastor milesi and Craugastor chrysozetetes by James McCranie, which
appear in Guia de Campo de los Anfibios de Honduras by James R. McCranie & Franklin Castañeda. We also extend
thanks to the following photographers for making their superb images available for use on the front and back covers,
as well as the half-title pages: Joel Sartore (cover); Kenji Nishida (back cover); Danté Fenolio, Michael and Patricia
Fogden, and Maik Dobiey (Half-title pages).
In addition, we would also like to acknowledge Kraig Adler and the Society for the Study of Amphibians and Reptiles
for their permission to reproduce selected paintings by David M. Dennis from the excellent “Hylid Frogs of Middle
America”, by William E. Duellman. Likewise, we were kindly granted permission to reproduce illustrations from the
“Atlas of Amphibians of China” by Fei et al. (1999), for which we acknowledge the permission of Prof. Liang Fei, and
the two artists who contributed to the book, Yisheng Wang and Jian Li. Finally, we would also like to acknowledge
Ted Kahn and César Landazábal for permission to use their illustrations published in Ranas Arlequines, published
by Conservation International, and we thank José Vicente Rodriguez Machecha of CI-Colombia for making these
illustrations available to us, and for his assistance in securing permission to reproduce other images that appear
in this publication.
A project of this magnitude is never without sacrifices made by family and loved ones. Simon Stuart thanks his
wife Ann, and daughters Claire and Jyoti, for all their love and support throughout the Global Amphibian Assessment
project, and the writing of this book, even though it took him away from home far too often. Michael Hoffmann would
like to thank his parents, Francois and Cheryl, for their unstinting support and prayers, particularly over the last four
years while being 8,000 miles from home, and also his fellow editors for putting up with his continuous harassment
to meet project deadlines. Janice Chanson is extremely grateful for the love and support of her husband John
(including moving continents twice), and the happiness and love that baby Sacha whom arrived midway through the
writing of this book brings to our lives. Neil Cox thanks his parents and grandparents for giving him a love of both the
countryside and traditional natural history, and he thanks Kerry and Megan for their love, patience and understanding
while he spent far too much time away from home. Richard Berridge thanks his parents (and especially his mum,
Eileen) for supporting his interest in mantellas, including putting up with a house that smells like a brewery (on ac-
count of breeding fruit flies in the laundry cupboard), and for rescuing frogs on the stairs during dinner parties; and
also to his much loved grandfather, Norman, for starting off his interest in natural history in the first place. Pavithra
Ramani thanks her family and friends for their tremendous support during the production of this book. They have all
kept an open mind and have shown an eagerness to learn, which she heartfully appreciates. Bruce Young is grateful
for the tremendous kindness and patience shown by his wife Cynthia and daughters Emily and Amanda during the
compilation of the Global Amphibian Assessment.
Finally, the last word of thanks must go to the ever remarkable publication house that is Lynx Edicions, and
particularly to Josep del Hoyo for constant guidance, encouragement and patience, and Susanna Silva for leading
on the production side with great efficiency and creativity.
GLOBAL AMPHIBIAN ASSESSMENT ACKNOWLEDGEMENTS
Donors
The Moore Family Foundation and the Gordon and Betty Moore Foundation, through Conservation International, provided
the core financial support for the Global Amphibian Assessment. The MAVA Foundation, the US Department of State,
the Regina Bauer Frankenberg Foundation for Animal Welfare, the National Science Foundation (DEB-0130273 and
INT-0322375), the Critical Ecosystem Partnership Fund, George Meyer, the European Commission, Ben Hammett, and
the Disney Foundation provided additional major support. The Kadoorie Farm and Botanic Garden, WWF Australia, the
Taipei Zoological Foundation, the Chicago Zoological Society, the Society for Wildlife and Nature, the Columbus Zoo and
the IUCN Centre for Mediterranean Cooperation also provided generous support. Claude Gascon and Jorgen Thomsen
in particular assisted us with fundraising. Any opinions, findings, and conclusions or recommendations expressed in
this material are entirely those of the authors and do not necessarily reflect the views of any donor.
Conservation Partners
Special mention must be made of George Rabb, who was the first to recognize the enormity of the global amphibian
conservation crisis, and who has mobilized both scientists and conservationists to address this challenge. Out of
respect for his visionary leadership, we dedicate this book to him.
Darrel Frost of the American Museum of Natural History provided extensive assistance on taxonomic and nomen-
clatural issues, without which it would have been much more difficult to implement the Global Amphibian Assessment.
David Wake of the Museum of Vertebrate Zoology at the University of California at Berkeley gave us privileged access
to the AmphibiaWeb database. We are most grateful to both of these people for their unfailing support.
We received assistance and advice in ways too numerous to mention from the IUCN/SSC Amphibian Specialist
Group (formerly the IUCN/SSC Declining Amphibian Populations Task Force), in particular from Don Church, Jim Collins,
Claude Gascon, Tim Halliday, Jim Hanken, Jeanne McKay, Robin Moore and John Wilkinson.
The following people provided local logistical support for the GAA workshops: Jean-Marc Hero (Australia), Zhong
Shengxian (China), Barasa Johnson (Kenya), Sanjay Molur and Sally Walker (India), Rosa Mary Saengsanthitham
(Thailand), Enrique Lahmann (Costa Rica), Sabrina Cowan and Allen Allison (Hawaii), Adriano Paglia, Jose Maria
Cardoso da Silva and Luis Paulo de Souza Pinto (Brazil), Paul and Sara Salaman and José Vicente Rodríguez (Ecuador),
Doreen Zivkovic (Switzerland), Esteban Lavilla (Argentina), Juan Carlos Ortiz (Chile), David Gower and Mark Wilkinson
(UK), Sixto Inchaustegui (Dominican Republic), and Sonsoles San Román and Jamie Skinner (Spain). We also thank
Craig Hilton-Taylor, Sanjay Molur, Bob Inger, Arvin Diesmos, Matt Foster, Mike Hoffmann, Penny Langhammer, Don
Church and David Knox who assisted in facilitating working groups during GAA workshops.
John Pilgrim worked diligently on adding all the amphibians that were described since the 2004 release of the
data, including creating range maps for each of these species.
Gustavo Fonseca, Claude Gascon, Russell Mittermeier, Tom Lacher, Tom Brooks, Larry Master, Bruce Stein and
Georgina Mace provided guidance and encouragement throughout the project, and we wish to express a special debt
XIV Threatened Amphibians of the World

of gratitude to them. Rob Waller, Vineet Katariya and Mark Denil provided extensive, high-quality GIS support. Leslie Fei Liang John Palis
Honey set up and manages the Global Amphibian Assessment website on http://www.globalamphibians.org/. Feng Fernando Castro John Poynton
Xie provided invaluable assistance in many ways, and in particular by helping us with the Chinese data, and giving Fernando Nogales S John Wilkinson
us access to scientific literature in Chinese. Don Church and Allison Parker did a very large amount of work locating Firoz Ahmed Jon Loman
missing bibliographic references and entering them into the database. Noura Bakkour, Laara Manler and Andrew Flavio Baldisseri Jorge A. Cespedez
Mitchell provided important logistical and administrative support. Flora Acuña Juncá Jorge Luis Martinez Ruiz
The distribution maps used for U.S. species were adapted from several sources, including NatureServe’s Central Franco Andreone Jörn Köhler
Databases and the United States Amphibian Atlas Database. The NatureServe data were developed in collaboration Frank Glaw Jose Antonio Mateo Miras
with its Natural Heritage member programmes, a leading source of information about rare and endangered species, Frank Lemckert Jose Hernández
and threatened ecosystems. The United States Amphibian Atlas Database was compiled at Ball State University Frank Solís Jose Langone
by Priya Nanjappa, Laura M. Blackburn, and Michael J. Lannoo and supported in part by grants and/or matching Franklin Casteñeda José Nuñez
funds from the National Fish and Wildlife Foundation, United States Fish and Wildlife Service, and Disney Wildlife Franky Bossuyt José P. Pombal Jr.
Conservation Fund. François Tron Jose Padial
Fred Kraus José Vicente Rueda Almonacid
GAA Contributing Scientists Fred Parker Joseph Mitchell
G.K. Bhat Joseph R. Mendelson III
The GAA was entirely dependent on the more than 500 herpetologists who generously gave of their time and Gabriel Skuk Joseph T. Collins
knowledge. The enthusiasm and commitment of these people has enabled us to generate a comprehensive global Gabriela Parra Josiah Townsend
picture of amphibian status and trends for the first time. We record our thanks to the following people, asking for Gad Degani Juan Formas
forgiveness from anyone whose name is inadvertently omitted or misspelled: Gajanan Buddhe Juan Manuel Guayasamin
Gary Fellers Juan Manuel Renjifo
A. Vivek Gour-Broome Chou Wenhao Genevieve Gee Judit Vörös
Abraham Mijares-Urrutia Christian Marty Geng Baorong Julian Faivovich
Abyerami Balasubramanian Christopher Austin Geoffrey A. Hammerson Julian Harrison
Adolfo Amezquita Christopher Beachy George Zug Julian Lee
Agnieszka Ogrodowczyk Christopher J. Raxworthy Georgina Santos Barrera Justin Gerlach
Ahmad Mohammed Mousa Disi Christopher Magin Gerardo Chavez Kaniskha Ukuwwela
Alain Dubois Christopher Pearl Ghazi S.M. Asmat Karen Lips
Alan Channing Christopher Phillips Gilda Andrade Karl-Heinz Jungfer
Alan Pounds Clare Morrison Gláucia Moreira Drummond Karthikeyan Vasudevan
Alberto Cadena Claude Gascon Gopalakrishna Bhatta Kartik Shanker
Alberto R. Estrada Claude Miaud Graeme Gillespie Katie Hampson
Alberto Veloso Claudia Azevedo-Ramos Gu Hui-qing Keith McDonald
Aldrin Mallari Claudia Cortez Fernandez Guarino R. Colli Kelly Irwin
Alex Kupfer Claudia Corti Guinevere Wogan Kelum Manamendra-Arachchi
Alfonso Miranda Leiva Conrad Hoskin Gunther Köhler Ken Aplin
Alfredo Salvador Cristina Arzabe Gustavo Cruz Kenneth Dodd
Allen Allison Cynthia Dolino Güven Eken Kim Howell
Alonso Quevedo Gil César Aguilar Puntriano Hal Cogger Kristiina Ovaska
Alvin Braswell César Jaramillo Hans-Werner Herrmann Juan Antonio Oliver Valls
Ana Almendariz César Luis Barrio Amorós Harold Cogger Juan Carlos Ortiz
Ana Carolina Queiroz Carnaval Dale Jackson Harry Hines Juan Elias Garcia-Perez
Ana Maria Paulino Telles de Carvalho e Silva Dale Roberts Hartwell Welsh Krushnamegh Kunte
Anand Padhye Dan Cogalniceanu Helen Díaz-Páez Kumthorn Thirakhupt
Andrea Herman Daniel R. Neira Herrera Helen Gerson Kurtulus Olgun
Andreas Schmitz Dante Pavan Hellen Kurniati Larry David Wilson
Andres Acosta David Beamer Hendrik Muller Leong Tzi Ming
Andres Merino-Viteri David Bickford Herman Nuñez Leslie Minter
Andrew Gardner David Bradford Hinrich Kaiser Letty Afuang
Andrew Turner David Cannatella Ho Thu Cuc Li Cheng
Angel Alcala David Donaire-Barroso Idriz Haxhiu Li Pipeng
Anna Wong David Gower Ignacio J. De la Riva Li Shengquan
Annamaria Nistri David Hunter Indraneil Das Liang Gang
Annemarie Ohler David Moyer Iñigo Martínez-Solano Lic. José A. Langone
Anslem De Silva David Newell Irina Maslova Lily Rodriguez
Antonio Muñoz Alonso David Price Irwin García Linda LaClaire
Antonio W. Salas David Tarkhnishvili Ishmail H. Ugurtas Liz Dovey
Argelia Rodríguez David Vieites Ismael E. di Tada Liza Paguntalan
Ariadne Angulo David Wake István Kiss Lovemore C.J. Mazibuko
Ariel Rodríguez Gómez Debjani Roy Ivan Sazima Lu Shunqing
Arne Schiotz Debora Silvano J. W. (Pim) Arntzen Luciana Barreto Nascimento
Aroshana Weerasinghe Deepthi Wickramasinghe Jaime Bertoluci Lucy Aquino
Arvin C. Diesmos Denis Vallan Jaime Bosch Lue Kuangyang
Atherton de Villiers Denise de C. Rossa-Feres James Hanken Luis Canseco
Avital Gasith Derek Brown James Harrison Luis Coloma
Axel Kwet Dick Watling James McCranie Luis J. Villanueva-Rivera
Baldwin Gutierrez Diego Almeida James Menzies Luis M. Diaz
Beatrice Ibéné Diego Baldo James Rorabaugh Luke Shoo
Ben D. Bell Diego F. Cisneros-Heredia James Vonesh Luis Zambrano
Benedetto Lanza Dilip Bhagwanrao Sawarkar Jan Meerman M.A. Bangoura
Benedikt Schmidt Diva Ma. Borges-Nojosa Jarujin Nabhitabata M.S. Ravichandran
Bexel Ayyasamy Daniel Djoko Iskandar Javier Icochea Monteza Madhava Meegaskumbura
Blair Hedges Dolores Huacaz Jay Savage Mae Leonida Diesmos
Boris Blotto Don Church Jean Lescure Magno V. Segalla
Boris Tuniyev Donald Shepard Jean Raffaeli Maklarin Lakim
Bosco Chan Dondi Ubaldo Jean-Louis Amiet Malcolm Largen
Brad Shaffer Dwight Lawson Jean-Luc Perret Mandy Tocher
Brandon Anthony Ed Meyer Jean-Marc Hero Manfred Beier
Brian Kubicki Edgar Lehr Jeanne McKay Manuel Acevedo
Bruce Bury Edif. Newton Plaza Jeet Sukumaran Manuel Morales
Bruce Waldman Eduardo Toral Jef Jaeger Marc Cheylan
Bruce Young El Hassan El Mouden Jelka Crnobrnja-Isailovic Marcelino Hernandez
Bruno Pimenta Eldad Elron Jenny Daltry Marcelo Napoli
Bryan Stuart Eli Greenbaum Jerry Hardy Marcos Vaira
Byron Wilson Eliza Maria Xavier Freire Jesus H. Cordova Santa Gadea Marga Born
Carlos Alberto Gonçalves da Cruz Elizabeth Scott Jesus Manzanilla Margaret Considine
Carlos Davidson Emilio Balletto Jiang Jianping Maria Cristina Ardila-Robayo
Carlos Frederico D. da Rocha Engin Gem Jigme Palden Maria Isabel Herrera
Carmen A. Ubeda Enrique La Marca Joan Mayol Serra Maria Ogielska
Carmen Díaz Paniagua Erik N. Smith Johannes Foufopoulos Mariela Osorno
Celio Haddad Erik R. Wild John Clarke Marinus S. Hoogmoed
Celsa Senaris Erin Muths John E. Cadle Mario Garcia Paris
Channa Bambaradeniya Ernesto Fernández-Badillo John Fa Mario Yánez- Muñoz
Charles Msuya Ernesto Recuero Gil John Jensen Marisol Pedregosa
Charles Painter Esteban O. Lavilla John Lynch Marius Burger
Chelmala Srinivasulu Federico Bolaños John Measey Mark Bailey
 Acknowledgements XV

Mark Wilkinson Oswaldo Luiz Peixoto Robert P. Reynolds Susan Koenig

Mark-Oliver Roedel Pablo Menendez Robert Powell Sushil Dutta
Martha Crump Paola Mosig Reidl Robert Puschendorf Sylvana Mass
Martha Patricia Ramirez P. Patricia Narvaes Robert Schabetsberger Tahar Slimani
Martin Bustamante Patrick Malonza Robert Stuebing Tanya Chanard
Martin Pickersgill Paul Chippindale Robert Webb Taran Grant
Marvalee Wake Paul Edgar Roberta Lecis Tassanee Eamkamon
Mary Gartshore Paul Horner Roberto Ibanez Tatjana Dujsebayeva
Mason Ryan Paul Imbun Yambun Rogelio Cedeño Vázquez Ted Kahn
Mathias Behangana Paul Moler Rogério Pereira Bastos Tej Kumar Shrestha
Mathieu Denoël Paul Walker Rohan Pethiyagoda Terry Schwaner
Matthew Parris Paula Cabral Eterovick Roman Khalikov Theodore Papenfuss
Mattias Stoeck Paulino Ponce-Campos Ronald A. Nussbaum Thierry Fretey
Masafumi Matsui Paulo Christiano de Anchietta Garcia Ronald Crombie Thomas Uzzell
Maureen A. Donnelly Paulo Sá-Sousa Ronald Heyer Tibor Kovács
Max Sparreboom Pedro Beja Ross Alford Tim Davenport
MD. Kamrul Hasan Dolon Per Nyström Ross D. MacCulloch Tim Halliday
Meren Ao Perry Ong Roy Swain Tom Mann
Michael Adams Peter Beerli Riyad A. Sadek Travis Ryan
Michael Cunningham Peter Brown Rob Grasso Trent Garner
Michael Harvey Peter Paul Van Dijk Robert Drewes Trevor Beebee
Michael Mahony Peter Robertson Roy W. McDiarmid Tuba Kiliç
Michael Sredl Peter Vogel Rubén Albornoz Ulisses Caramaschi
Michael Wei Neng Lau Petros Lymberakis Ruth Adriana Maldonado-Silva Ulisses Galatti
Michel Breuil Phil Bishop Ruth Amanda Estupinan Ulrich Sinsch
Michele Menegon Philippe Gaucher S. Bhupathy Uriel Barrantes
Miguel Lizana Avia Philippe Geniez S. P. Vijayakumar V.A. Prasanna Samarawickrama
Miguel Tejedo Pierre-André Crochet S.D. Biju Valentín Pérez Mellado
Miguel Trefaut Rodrigues Pradeep Kumara S.R.M Swarnapali Samaradiwaka Vance Vredenburg
Miguel Vences Pranabes Sanyal Sabitry Choudhury Bordoloi Ulrich Joger
Miklós Puky Pranjalendu Ray Saibal Sengupta Vanessa Verdade
Milan Vogrin Querube Fuenmayor Sanjay Molur Varad Giri
Mills Tandy Rafael Joglar Santiago Ron Varol Tok
Min Mi-Sook Rafael Lajmanovich Sarah May Víctor R. Morales
Mohammad Ali Reza Khan Rafael Márquez Sarig Gafny Victorino Molina
Mohammed Firoz Ahmed Rafe Brown Sean Blomquist Vidana Arachchilage Samarawickrama
Mohini Mohan Borah Rainer Guenther Senarathge Weerawardhena Vladimir Ischenko
Monique Van Sluys Rainer Schulte Sergio Potsch de Carvalho e Silva Wang Xiuling
Mozafar Sharifi Rajehdra Vyas Sergius Kuzmin Wang Yuezhao
Muhammad Sharif Khan Ramón Formas Sherif Baha El Din Wenhua Lu
Mumpuni Randy Jennings Shi Haitao Wichase Khonsue
Murat Sevinç Randy McCranie Simon Loader Wieslaw Babik
Murray Littlejohn Ranjit Daniels Sixto Inchaustegui Wilfredo Arizabal
Naomi Doak Raoul Bain Slimani Tahar Will Osborne
Nasrullah Rastegar-Pouyani Rastko Ajtic Sohrab MD Sarkar Uddin William E. Duellman
Natalia B. Ananjeva Raymond Brereton Somsak Panha Wilmar Bolivar
Neftalí Ríos-Lopez Renato Neves Feio Souad Hraoui- Bloquet Wolfgang Böhme
Néstor G. Basso Renaud Boistel Srinivasulu Chelmala Wu Guanfu
Nguyen Quang Truong Richard Griffiths Stefan Lötters Xie Feng
Nick Arnold Richard Highton Steffen Reichle Yang Datong
Nikolai Orlov Richard Retallick Stephen Richards Ye Changyuan
Nimal Dharmapriya Rathnayake Richard Siegel Stephen Richter Yehudah. L. Werner
Noami Doak Richard Thomas Steven C. Anderson Yodchaiy Chuaynkern
Norman Scott Richard Tinsley Steven Karsen Yolanda Matamorros
Norsham Suhaina Yaakob Richard Zweifel Steven Morey Yoshio Kaneko
Olivier Pauwels Rita Pascolini Steven Swan Yuan Zhigang
Oommen V. Oommen Robert Herrington Suh Y. Yang Zhao Ermi
Oscar Flores-Villela Robert Inger Suleima Santiago Zhao Wenge
Oscar Fuentes-Ramos Robert Jehle Surya Adoor Zheng Zhonghua
Introductory Chapters
 CHAPTER 1. AN INTRODUCTION TO THE AMPHIBIANS
Neil Cox, Simon Stuart, Janice Chanson, Michael Hoffmann,
David Gower, Mark Wilkinson, Don Church and Robin Moore
Figure 1. Species richness map for amphibians in the Order Anura, with dark red colours corresponding to regions of higher richness. Colour
scale based on 10 quantile classes. Maximum richness equals 142 species.
Figure 2. Species richness map for amphibians in the Order Caudata, with dark red colours corresponding to regions of higher richness. Colour
scale based on 10 quantile classes. Maximum richness equals 23 species.
Figure 3. Species richness map for amphibians in the Order Gymnophiona, with dark red colours corresponding to regions of higher richness.
Colour scale based on 8 quantile classes. Maximum richness equals 8 species.
AN OVERVIEW OF AMPHIBIAN CLASSIFICATION
AND DIVERSITY
Amphibians are familiar to most people as frogs and toads, salamanders and newts, and to
a much smaller group of people as caecilians. All amphibians are members of the tetrapod
vertebrate Class Amphibia. There are more than 6,000 currently recognized species of extant
amphibians, with representatives present in virtually all terrestrial and freshwater habitats,
but absent from the coldest and driest regions, and from the most remote oceanic islands.
The number of recognized species of amphibians has grown enormously in recent years,
with a nearly 50% increase between 1985 and 2004 (Frost 1985, 2004) and an increase
in species numbers of 25% in the years between 1992 and 2003 (Köhler et al. 2005) (and
see Essay 1.1). This unprecedented growth largely reflects an increase in collecting work
in previously remote locations, a significant growth of active herpetological communities
in a few megadiverse countries, and the application of complementary techniques, such as
molecular genetics, to support more traditional taxonomic methods. Even countries such
as Sri Lanka, in which biodiversity inventories were deemed to be relatively complete (see
Essay 1.2), are revealing startling levels of previously undocumented and unsuspected
diversity. Unfortunately, as this book demonstrates, our rapid increase in the knowledge
of amphibian species diversity and biology is coincident with a massive global decline in
amphibian populations.
Frogs, Salamanders and Caecilians –
the Amphibian Orders
The Amphibians are divided between three higher-level ranks or Orders: Gymnophiona,
Anura and Caudata, with Anura and Caudata being more closely related to each other than
either is to the Gymnophiona.
Anura is comprised of the frogs (and their subgroup, the toads), and is by far the largest
Order, with 5,208 living species currently recognized1. Anurans are globally distributed,
being found on every continent with the exception of Antarctica (Figure 1). While tropical
habitats are richest in anuran diversity, frogs and toads may be encountered in many
different environments ranging from dry deserts, through tropical and temperate regions
to areas as far north as the Arctic Circle and as far south as Tierra del Fuego at the tip of
South America. The elevation at which frogs have been recorded ranges from sea level
(or even below it in some cases) to as high as 5,244m asl in the Peruvian Andes (Seimon
et al. 2006). Although anurans have a wide variety of body shapes, adult frogs and toads
are always tailless and have four limbs. The have a constrained and specialized body
form including a very short body, an elongated pelvis, relatively large head, and two
well-developed hind legs used for jumping and/or swimming. The frogs and toads range
greatly in size. The giant Goliath Frog Conraua goliath of West Africa may grow to over
30cm (snout-vent length), while mature adults of the Flea-frog Brachycephalus didactylus
of south-eastern Brazil and Eleutherodactylus iberia and E. limbatus of Cuba may be less
than 1cm (snout-vent length). Virtually all species have external fertilization and most
have aquatic larvae (tadpoles).
The Caudata (salamanders and newts) contains 535 described living species and is the
second largest of the three amphibian Orders, although representing only about 10% of
the number of species as frogs. Salamanders and newts are less widely distributed than
the frogs and toads, with families mostly concentrated in the north temperate regions
(Figure 2); but about 40% of the species occur in the New World tropics. There are no
salamander or newt species in Sub-Saharan Africa, Madagascar, the Indian Subcontinent
south of the Himalayas, insular Southeast Asia, and Australasia, and only a few in South
America and mainland Southeast Asia. The southern Appalachian Mountains of the eastern
United States is the centre of species diversity for the Caudata (see Essay 8.1). Around 60
species of the family Plethodontidae (lungless salamanders) are found in this mountain
range, including many locally restricted species. Most salamanders and newts have the
primitive tetrapod body shape, with four limbs and a long flattened tail, but there is also
a tendency for body elongation and limb reduction or loss. They generally move over land
by walking and aquatic species use their long tail for rapid swimming in water. Within
the Caudata there are many well-known paedomorphic forms (sexually mature animals
retaining juvenile characteristics). The highly threatened Axolotl Ambystoma mexicanum
of the Valley of Mexico is probably the most familiar paedomorphic species. The largest
of the Caudata, and indeed of modern amphibians, is the rapidly declining Chinese Giant
Salamander Andrias davidianus, which may grow to around 200cm in total length; the
smallest may be the salamander Thorius arboreus of Mexico, which has an average total
length of only around 1.7cm. Most salamanders have internal fertilization via a spermato-
phore and aquatic larvae, but direct development of terrestrial eggs without an aquatic
life-stage whatsoever is also common.
The Gymnophiona (caecilians) occur in the tropics of the Americas, Africa and Asia
(Figure 3). In addition, caecilians are present on the Seychelles, but absent from Madagascar,
New Guinea and Australasia; their apparent absence from much of the Congo Basin may
be due to lack of survey effort. With only 172 described species, this is the smallest of the
three amphibian Orders, even though the group is at least as old as all of the diversity of
salamanders and frogs taken together. Because they tend to be rare animals in collections,
the taxonomic status of many caecilian species is uncertain. According to our current
understanding of caecilian taxonomy, the two most species-rich genera are Ichthyophis
(34 species) of South and Southeast Asia, and Caecilia (33 species) occurring mainly in
South America. Caecilians are elongated and limbless, and they are sometimes mistaken
for snakes, eels or earthworms. The largest of the caecilians is the poorly known Caecilia
thompsoni of Colombia, which may exceed 150cm in length. The smallest caecilian is prob-
ably the threatened Grandisonia brevis of the Seychelles Islands, which grows little larger
than 11cm. In general, caecilians are ‘burrowing’ species and are most often encountered
in leaf-litter or under soil. A few are aquatic species (such as the Rubber Eel Typhlonectes
natans). Caecilian species show a relatively wide range of reproductive modes. All species
seem to have internal fertilization and at least 31 of the known species are live-bearing.
Oviparous caecilians lay their eggs on land have either direct development or an aquatic
larva (and see Essay 1.3).
 Chapter 1. An Introduction to the Amphibians 3

Amphibian classification at the family level Scolecomorphidae. This poorly known family of caecilians comprises two genera, the
West African Crotaphatrema and East African Scolecomorphus, with three species in
At a slightly finer taxonomic level, amphibians are divided up among 48 families, some of each. We know very little about Crotaphatrema, as only seven individuals have ever been
which, like the recently named Nasikabatrachidae (Biju and Bossuyt 2003), are monotypic (in- seen, all from western Cameroon. The diversity of the genus Scolecomorphus is centred
cluding only a single species), whereas others, particularly the Hylidae and Leptodactylidae, on eastern Tanzania. Two of the species are known to be live-bearing, and the third is
are represented by hundreds of species, the latter including the genus Eleutherodactylus, presumed to be so.
the most speciose vertebrate genus with more than 600 species (see Essay 1.4). Amphibian
classification was relatively stable for much of the 20th century, but the very roots of the
amphibian tree have been shaken by a new classification proposed by Frost et al. (2006).
Frost and associates showed that a number of traditionally recognized groups (including
entire families) are para- or polyphyletic (in other words, they are not natural groups in terms
of evolutionary history). The new classification proposed by Frost et al. (2006) appeared
while this book was in preparation. Thus, we use the traditional classification of amphibian
families (following Frost 2004). However, in this chapter, we outline the new classification of
Frost et al. (2006), as amended by Grant et al. (2006) and Glaw and Vences (2006), to show
some of the major taxonomic changes that are now under discussion. We also discuss the
implications of Frost et al. (2006) in the regional introductory chapters.

The Traditional View of Amphibian Families

This section presents the traditionally recognized amphibian families, alphabetically within
each order.

Caecilians – Gymnophiona Uraeotyphlidae. This small family of caecilians contains only five species all of which
are in the genus Uraeotyphlus. The family is entirely restricted to the Western Ghats and
Caeciliidae. This is the largest of the caecilian families, containing 113 species, distributed surrounding parts of southern India. The breeding biology of the family is poorly known, but
within 21 genera. Many of these species are known from fewer than 10 specimens and little they are probably all oviparous species with terrestrial eggs and aquatic larvae.
is known of species ranges and biological attributes. In this treatment, the semi-aquatic
to aquatic species of South America, often recognized as the Family Typhlonectidae, are
included in the Caeciliidae. Representatives are widely, but disjunctly, distributed throughout
much of the tropics, being found in South and Central America, West and East Africa, on the
Indian Subcontinent and also on the Seychelles Islands (uniquely, there are more caecilian
species in the Seychelles than there are frogs). Although the breeding biology of most
caeciliid species remains poorly known at present, larval-developing, direct-developing
and live-bearing species have all been recorded.

Salamanders and Newts – Caudata

Ambystomatidae – Mole Salamanders. This family of 30 species (all within the genus
Ambystoma) is strictly North American (occurring south to central Mexico), where they range
widely throughout much of the continent. Animals are largely terrestrial and breed in slow
or still waters but some are stream-dwellers; all species undergo larval development. The
family includes a number of paedomorphic species that retain larval characteristics (such
as gills) as adults, including the famous Axolotl Ambystoma mexicanum.
Ichthyophiidae. This poorly known family of caecilians is widely distributed in South
Asia and Southeast Asia. The 39 recognized species are divided between the two very
poorly delimited genera Ichthyophis and Caudacaecilia. However, many of the currently
named species are poorly circumscribed and require taxonomic confirmation. All taxa are
egg-layers. The eggs are deposited in underground chambers and they are guarded by the
mother at least until hatching. The larvae may be encountered in waterlogged soil, streams,
ponds and other waterbodies.

Amphiumidae – Amphiumas. This family contains three species, all in the genus
Amphiuma, and is endemic to the coastal plain of the eastern and south-eastern United
States. These are relatively large, eel-like salamanders, which are specialized for a mostly
aquatic lifestyle. The eggs of these species are laid in a moist cavity on land close to the
water; the female guards the nest until the larvae hatch and are washed into the adjacent
water. An interesting feature of the larvae is their external gills, which disappear once
the lungs develop.
Rhinatrematidae. This poorly known family of caecilians is restricted to northern South
America, with most species known from the tropical Andes. The genus Epicrionops has
eight species and Rhinatrema one. All species are presumed to lay eggs on land, with the
larvae living in waterbodies.
4 Threatened Amphibians of the World
The Pickerel Frog Rana palustris (Least Concern), found over much of eastern North America,
is a typical representative of the very diverse amphibian Order Anura, which includes the
frogs and toads. © Don Church
Cryptobranchidae – Giant Salamanders. This small family is entirely aquatic. Crypto-
branchidae are very large salamanders placed in three species in two genera (Andrias and
Cryptobranchus). The family has a disjunct distribution with species present only in East Asia
and the eastern United States. The species undergo larval development, with paired strings
of externally fertilized eggs deposited under stones or between rocks within streams and
other waterbodies. This family contains the largest (~200cm) and the heaviest (~50kg) of the
modern day amphibians, the Chinese Giant Salamander Andrias davidianus.
Dicamptodontidae – Pacific Mole Salamanders. This small family of relatively large,
larval-developing salamanders is restricted to western parts of the United States, with
one species occurring marginally in Canada. All four members are in the single genus
Dicamptodon. The eggs are deposited in clear streams, where they are attached singly to
rocks and other cover. The stream-adapted larvae take between 2 and 4.5 years to reach
metamorphosis and may achieve sexual maturity without metamorphosing. One species,
Dicamptodon copei, is exclusively paedomorphic.
Hynobiidae – Asian Salamanders. These salamanders are Asian with a centre of diversity
in Japan, with only the very widespread Siberian Salamander Salamandrella keyserlingii also
present in extreme eastern Europe. There are 46 species divided between the seven genera,
with Hynobius being the most speciose (27 species; see Essay 10.1). The salamanders of this
family are highly aquatic, living mostly in streams and pools, and have external fertilization,
and the eggs are laid in two egg-sacs attached to rocks or submerged vegetation. In some
species, the male is known to guard the eggs.
Plethodontidae – the Lungless Salamanders. This is the largest of the salamander
families, with the vast majority of the 365 species being found in the New World (see
Essay 9.3). Outside of this region there are seven species known from Europe, and a single,
recently discovered Asian species (Karsenia koreana) currently known only from the middle
of the Korean Peninsula. Plethodontids are the only salamanders that have a substantial
presence in the New World tropics, where more than 60% of the species are found, most
of them in Central America. The family is generally present in a wide variety of habitats
including damp terrestrial areas, waterbodies (both stagnant and flowing), cave and similar
habitats, and arboreal sites (such as within bromeiliads). Most of the species have direct
development, unique to this family among salamanders; however, at least 50 species
undergo larval development.
Proteidae – Mudpuppies and Waterdogs. This small family of salamanders comprises
five species in the genus Necturus, found in the streams of eastern North America, and the
single strange species, the Olm Proteus anguinus, which is restricted to subterranean aquatic
habitats within the Dinaric Alps of south-eastern Europe. All of the species are both aquatic
and paedomorphic. The breeding biology of this family is poorly known; however, female Olms
have been recorded laying ~70 eggs on rocks, and, in addition, the species is occasionally
live-bearing with fully formed young born after a period of about two years.
Rhyacotritonidae – Torrent Salamanders. This small family of four species, all in the
genus Rhyacotriton, is endemic to the Coastal and Cascade Ranges of the western United
States. Populations of these salamanders are usually closely associated with cold streams
in old-growth coniferous forest. All four species undergo larval development.
 Chapter 1. An Introduction to the Amphibians
Salamandridae – Newts and Relatives. This family is widespread with representa-
tives present in Europe, North Africa, Asia and North America. Most of the species within
Salamandridae are aquatic breeders, laying eggs in ponds, streams and other suitable
waterbodies. A small number of the species, including the well-known Fire Salamander
Salamandra salamandra, are live-bearing.
Sirenidae – the Sirens. This small family has two genera (Siren and Pseudobranchus)
with four species, and is restricted to the south-eastern United States and northern Mexico.
Animals are mostly aquatic; the eggs are deposited singly or in small numbers, and are
attached to submerged vegetation.
Frogs and Toads – Anura
Allophrynidae. This monotypic family is generally widespread in northern South America,
where it ranges from Venezuela, through much of the Guiana region and into north and
eastern Brazil2. The single species is largely associated with water bodies, where explosive
breeding and larval development take place. The classification of the family, and the single
recognized species, Allophryne ruthveni, has been problematic. Allophryne has previously
been assigned to a number of families, including Allophrynidae, Bufonidae, Centrolenidae,
Hylidae and Leptodactylidae.
Arthroleptidae – Squeakers or Screaming Frogs. This family is dominated by the
direct-developing genus Arthroleptis (34 species), which is widely distributed in more
mesic environments of the African mainland. Together with Schoutedenella, it is one of
the most taxonomically confused groups of frogs in the world, with many species in Sub-
Saharan Africa being of uncertain identity, and with large numbers of species remaining
to be described. The third genus in the family, Cardioglossa (16 species), is represented by
stream-associated, larval-developing frogs in West and Central Africa, with the centre of
known diversity in Cameroon.
5
Ascaphidae – Tailed Frogs. The small family Ascaphidae contains only two species, both
in the genus Ascaphus. The family exhibits a number of primitive characteristics, and is
unique among the frogs and toads in that animals posses a copulatory organ (the eponymous
“tail”) that allows internal fertilization to take place in the fast-flowing streams they inhabit.
The unpigmented eggs are attached as strings to the underside of rocks within the stream;
the larvae also develop within this environment. The family is endemic to North America,
where it is distributed in the north-western USA and south-western Canada.
Astylosternidae. This family of stream-associated, larval-developing frogs has its centre
of diversity in Cameroon, with a few species occurring more widely within the equato-
rial forest belt. Two genera, Astylosternus (11 species) and Leptodactylodon (15 species)
dominate the family, with three monotypic genera, one of which, Trichobatrachus, is the
famous Hairy Frog.
Bombinatoridae – Fire-bellied Toads. This small family of only 10, often colourful,
species is widely distributed in Europe and Asia. There are two genera, Bombina (eight
species), which mostly ranges from Western Europe into East Asia, and the poorly known
genus Barbourula, that has only two known species, one present in the Philippines and one
in Borneo. Bombina are all believed to undergo larval development; the reproductive mode of
Barbourula is currently unknown, but they are suspected to be direct-developing species.
Brachycephalidae – Saddle-back Toads or Pumpkin Toadlets. This small family of
toads is restricted to the humid forests of the Atlantic coast of south-eastern Brazil. There is
a single genus, Brachycephalus, containing the eight known species3 - five of these having
been described since 1998. These small toads live in leaf litter on the forest floor, and are all
believed to deposit a few large, terrestrial eggs and to undergo direct development.
6 Threatened Amphibians of the World
Bufonidae – True Toads. Members of the Bufonidae are near global in occurrence, being absent
naturally from Madagascar, New Guinea, Australia and surrounding islands. Species occupy a
wide variety of habitats, from very arid conditions to humid tropical rainforest. It is a speciose
family and although the vast majority of the species have larval development, there are some
direct-developing and live-bearing species. The genus Nimbaphrynoides (two species), includes the
only fully viviparous frog species (with the embryos being nourished internally by the female).
This Spotted Salamander Hynobius naevius
(Least Concern) is one of sixteen Hynobius
salamander species found only in Japan.
The only other representatives of the Family
Hynobiidae are found in mainland China (six
species), the island of Taiwan (three), and the
Korean Peninsula (three). © Henk Wallays
Centrolenidae – Glass Frogs. This speciose Neotropical family ranges from southern
Mexico to Bolivia and north-eastern Argentina. The family is overwhelmingly concentrated
in the tropical Andean countries (Colombia, Ecuador, Peru and Venezuela), where 124 species
(90% of the total) occur. Most species are arboreal to some degree, and typically lay small
clutches of eggs on vegetation or rocks overhanging water. On hatching, the larvae fall into
the water below and complete their development. Members of this family are unusual, in
that the dorsal part of the body is commonly a shade of green, whereas the ventral side is
often transparent, allowing many of the internal organs to be seen.
Dendrobatidae – Poison Frogs. The dendrobatids are almost exclusively confined to
the tropics of Central and South America. The greatest diversity is in the tropical Andean
countries, where 190 species (81% of the total) occur. A single species occurs in the Lesser
Antilles (on Martinique). Only a few species are the familiar brightly coloured animals often
seen in the pet trade (especially in the genera Dendrobates, Epipedobates and Phyllobates);
the majority are less colourful, including most species of the largest genus Colostethus (131
species). Nearly all species have a larval phase. In some species an adult guards the eggs
at the deposition site (such as a bromeliad), before carrying the hatched larvae on its back
to water (usually a stream) where development is completed.
Discoglossidae – Painted Frogs and Midwife Toads. This small family has its distribu-
tion centred on the western Mediterranean Basin, with one species extending as far north
as northern Germany, and a single species known from Israel (the now extinct Hula Painted
Frog Discoglossus nigriventer). The 12 species are divided between two genera, Alytes and
Discoglossus. Frogs of both genera undergo larval development in suitable waterbodies. The
breeding behaviour of the genus Alytes is notable in that the males carry the strings of eggs
wrapped around their back legs, taking them to the water when they are ready to hatch.
Heleophrynidae – Ghost Frogs. This small relict family of stream-associated, larval-
developing frogs is endemic to southern Africa. All members are in the genus Heleophryne.
These frogs are dependent on permanent streams because their tadpoles require two
years to develop.
Hemisotidae – Snout-burrowers. This small family is widespread through much of
the Sub-Saharan African mainland. All members are in the genus Hemisus. Eggs are laid
underground in a nest cavity, and the tadpoles subsequently move into water where they
complete their development (the females sometimes digging channels for them to move
from the nest to water).
Hylidae – True Treefrogs. This family is the second largest clade of amphibians and consists
of three subfamilies: Hylinae (Americas and northern Eurasia); Phyllomedusinae (tropical Ameri-
cas); and Pelodryadinae (Australia, New Guinea and other islands). The family is particularly
diverse in the New World, with only a few hylid genera occurring elsewhere (see Essay 1.5).
Hylids occur throughout the Americas (with the exception of the extreme north and south),
and are also widespread in Australia and New Guinea. Elsewhere, there are a few species
in Europe and northern Asia, but the family is absent from most of tropical Asia, and all of
Sub-Saharan Africa and Madagascar. Many species are arboreal, although there are excep-
tions, with terrestrial and aquatic species also. The vast majority of species undergo larval
development. The subfamily Hemiphractinae, which included only species that undergo direct
development, was recently shifted to the family Leptodactylidae (Faivovich et al. 2005).
Hyperoliidae – African Treefrogs and Reed Frogs. This large family comprises 18
genera, including Hyperolius (125 species), Leptopelis (51 species), Afrixalus (32 species) and
Kassina (13 species). Hyperoliids can be found in most habitats throughout the Afrotropical
region, including Madagascar and the Seychelles Islands. Nearly all the members of the
family undergo larval development, but a wide variety of reproductive strategies occur
within the family. Under certain conditions, females of at least one species, Hyperolius
viridiflavus, can change into fully functional males (Grafe and Linsenmair 1989). Hyperoli-
ids are closely related to arthroleptids and astylosternids (with some authors suggesting
that Leptopelis is, in fact, closer to these groups than to other hyperoliids), sharing many
characteristics with them.
 Chapter 1. An Introduction to the Amphibians 7

Leiopelmatidae – New Zealand Frogs. This endemic New Zealand family comprises Megophryidae – Asian Spadefoots. These often-large Asian frogs range from India and
four largely terrestrial species in the single genus Leiopelma. The female lays large un- Pakistan through much of Southeast Asia, to the Philippines, Borneo and the Sunda Islands.
pigmented eggs in damp locations on the ground, which undergo direct development. The The family is quite diverse with 10 genera, the most species-rich of these being Xenophrys
males of three of the species (Leiopelma archeyi, L. hamiltoni and L. pakeka) actively guard (31 species). All of the species undergo larval development in streams or similar habitats.
the eggs and transport the developing young. Leiopelmatidae is particularly interesting in
the apparent retention of a number of primitive characteristics for frogs (see Essay 6.2),
mostly shared with Ascaphidae.

Microhylidae – Narrow-mouthed Toads. This diverse, large Anuran family is globally

Leptodactylidae – the Southern Frogs. This is by far the most speciose of all the
amphibian families, with the genus Eleutherodactylus alone containing 610 species (and
see Essay 1.4). The Leptodactylidae are found throughout South and Central America, with
a few species ranging as far north as the southern United States. The family is diverse
morphologically, with some species exclusively adapted to terrestrial, arboreal or aquatic
lifestyles. Even though the largest group in the family (Eleutherodactylinae) breed by direct
development, many are larval developing and at least one species, the Critically Endangered
Golden Coqui Eleutherodactylus jasperi, is live-bearing.
widely distributed, and may be encountered in many different habitats. The 69 recognized
genera include many highly specialized forms, with a number of species adapted to either
a largely fossorial or arboreal lifestyle. Both larval-developing and direct-development
reproductive modes have been recorded. Although many of the direct-developing species
are found on New Guinea, there are direct-developing genera outside of this region such
as Breviceps in Africa and Myersiella in Brazil.

Limnodynastidae. This family of 50 species is distributed from the Aru Islands of Indo-
nesia, through New Guinea and much of Australia, to the island of Tasmania. Five genera
(Adelotus, Heleiporus, Neobatrachus, Notaden and Philoria) are endemic to Australia, and
only five species occur outside of Australia, with four endemic to New Guinea. All of the
eight genera are larval developing, with some genera (such as Limnodynastes) depositing
eggs in floating foam nests. The Tusked Frog, Adelotus brevis, is unusual in that adults
have small tusk-like teeth sticking up from the bottom jaw; these might possibly be used in
defence of the nest (Duellman and Trueb 1994).
Hemiphractus fasciatus (Near Threatened) from Panama, Colombia and Ecuador, is one of six
species contained within in the newly proposed Family Hemiphractidae (currently included
in Leptodactylidae). All species are highly specialized treefrogs from primary rainforest that
feed only on other frog species. © Edgargo Griffith

Mantellidae. This family is endemic to Madagascar, and is dominated by the genera

Mantidactylus (86 species) and Boophis (53 species). However, by far the best-known
members of the family are the brightly coloured Madagascar poison frogs of the genus
Mantella (15 species). There are two other smaller genera. The family includes both larval-
and direct-developing species.
8 Threatened Amphibians of the World

Myobatrachidae. This family is distributed in southern New Guinea and much of Australia,
including the island of Tasmania. The family comprises 71 species and 11 genera, with the
three largest genera being Uperoleia (24 species), Crinia (15 species) and Pseudophryne
(13 species). All species are endemic to Australia except for Crinia remota, which is found
in northern Australia as well as the southern lowlands of New Guinea (both Indonesia and
Papua New Guinea), and Uperoleia lithomoda which occurs in northern Australia as well
as the southern lowlands of Papua New Guinea. Members are for the most part terrestrial,
with a number of burrowing species (e.g., Arenophryne). There are both larval-developing
and direct-developing species within the family. The monotypic genus Assa is unusual in
that the hatching larvae are carried in brood pouches on the hips of the male.
Pelobatidae – Western Palaearctic Spadefoots. This small family consists of only four
species, all in the genus Pelobates, ranging from Morocco and Iberia in the west, through
Europe, Southwest Asia and the Caucasus Mountains to western Kazakhstan. All four species
are larval developing, with adults generally spawning in stagnant temporary waterbodies.
The adults are adapted to digging in soil, spending much of the year underground, and
generally require uncultivated sandy habitats.

Pelodytidae – Parsley Frogs. This small family, consisting of three species all in the single
genus Pelodytes, has a disjunct distribution in south-western Europe and the Caucasus
Mountains. All species undergo larval development, with eggs laid in short strings.

Nasikabatrachidae – Indian Burrowing Frog. This recently described, currently mono-

typic, family is known only from a few localities in the Western Ghats of southern India. For
much of the year animals are fossorial, living up to 3.7m below ground. The species undergoes
larval development, with the adults coming to the surface for a few weeks each year to
breed and deposit their eggs in both temporary and permanent waterbodies. This new family
appears to be most closely related to the Sooglossidae of the Seychelles Islands.

Petropedetidae. This family is widespread in Africa, and is dominated by the puddle frogs
of the genus Phrynobatrachus (66 species). There are 13 genera in total, eight of which
are monotypic. Other genera are Cacosternum (10 species), Arthroleptella (7 species),
Petropedetes (7 species) and Arthroleptides (3 species). The family exhibits a wide range
of reproductive modes, although the great majority are larval developers.

The most familiar of amphibian larvae are

the free-swimming ‘tadpoles’ of the frogs
and toads. The body shape is roughly ovoid,
with a long laterally compressed tail used for
swimming. Pictured here is a larval Wood
Frog Rana sylvatica (Least Concern). © Twan
Leenders
 Chapter 1. An Introduction to the Amphibians
Pipidae – Tongueless Frogs. This small, distinct family of highly aquatic frogs is restricted
to South America and Sub-Saharan Africa. The Pipidae is unique among frogs in that species
do not have a tongue. In African genera, such as Xenopus, small pigmented eggs are laid
in water and undergo larval development. In the South American genus Pipa, the eggs are
laid into the females back and in some cases (such as in the Surinam Toad Pipa pipa) these
eggs develop directly into frogs without a larval stage.
Ranidae – True Frogs. This is the third largest Anuran family, and with a few exceptions,
for instance most of Australia and New Zealand, it has a very cosmopolitan distribution.
Members of this family may be found in a wide variety of arid, temperate and humid tropical
habitat types. While most reproduce through larval development, there are a number of
direct-developing genera such as Platymantis. The Ranidae contains the largest living anuran,
the Goliath Frog Conraua goliath, which can grow to over 30cm and weigh over 3kg.
Rhacophoridae – Asian Treefrogs. This large family of mostly arboreal frogs is widely
distributed in Asia and Africa. Of the nine genera, the most species-rich are Philautus (146
species) and Rhacophorus (64 species), both of which are restricted to Asia. The family
includes a diversity of both larval- and direct-developing species. Some species of the genus
Rhacophorus have extensively webbed toes that enable them to glide between trees.
Rheobatrachidae – Gastric-brooding Frogs. This family was restricted to eastern
Queensland, Australia, and is now considered to be extinct. There were only two spe-
cies in the family, Rheobatrachus silus and R. vitellinus, both of which shared a unique
reproductive mode. The eggs and larvae were brooded within the stomach of the female,
with the larvae feeding off of the egg yolk before emerging from their mother’s mouth as
fully formed frogs (see Essay 6.1). The status of this family has long been controversial,
with a number of authors including it as a subfamily within either the Limnodynastidae or
the Myobatrachidae.
9
Rhinodermatidae – Darwin’s Frogs. This small terrestrial family from the temperate
forests of Chile and neighbouring southern Argentina contains only two species, both within
the genus Rhinoderma. The small clutches of unpigmented eggs are laid in leaf litter. In
Rhinoderma rufum, the male takes up the hatching larvae in its mouth and transfers them
to water to complete their development. In Darwin’s Frog, Rhinoderma darwinii, the male
not only takes the larvae into its mouth, but also broods the larvae in the vocal sacs until
development into froglets is completed.
Rhinophrynidae – Burrowing Toad. This monotypic family is largely restricted to the
coastal lowlands of Central America. The single species, Rhinophrynus dorsalis, is for the
most part fossorial, coming only to the surface to mate during the rainy season; the eggs
and larvae develop in temporary pools formed by heavy rains.
Scaphiopodidae – North American Spadefoots. This small family of seven species,
divided between the two genera Scaphiopus and Spea, has been included by some authors in
the family Pelobatidae. All of the species tend to burrow in loose soil, with adults emerging
to breed in temporary pools during the rainy season. In general, the larvae develop quickly
before the seasonal desiccation of their habitat.
Sooglossidae – Seychelles Frogs. This family is endemic to the Seychelles Islands, and
comprises four species in two genera. Three species breed by direct development, and in
one the larvae are carried on the adults’ backs.
10
Direct development is one of the three basic
amphibian breeding strategies. In this photo-
graph, the young of the Bornean shrub-frog
Philautus amoenus (Vulnerable) can be seen
developing into frogs without undergoing a
free-living larval stage. © Andreas & Christel
Nöllert
Threatened Amphibians of the World
Ichthyophiidae. Frost et al. (2006) include the Uraeotyphlidae in this Indomalayan family,
but its content is otherwise unchanged.
Rhinatrematidae. This family remains unchanged.
Salamanders and Newts – Caudata
Cryptobranchidae. This family remains unchanged.
Hynobiidae. This family remains unchanged.
Rhyacotritonidae. This family remains unchanged.
Amphiumidae. This family remains unchanged.
The Proposed Revisions to Amphibian Families Plethodontidae. This family remains unchanged.
For some time it has been recognized that the traditional classification of amphibian families, Proteidae. This family remains unchanged.
especially among the frogs, does not reflect their evolutionary relationships (e.g., Darst and
Cannatella 2004; Faivovich et al. 2005; Grant et al. 2006). Frost et al. (2006) proposed a new Sirenidae. This family remains unchanged.
taxonomy of living amphibians, based on analysis of the interrelationships of 522 species
selected to the best of their ability to reflect the entire amphibian tree, using molecular Ambystomatidae. Frost et al. (2006) include the Dicamptodontidae in this North American
genetic data and also taking into account anatomical features, especially of larvae, follow- family, but its content is otherwise unchanged.
ing Haas (2003). The hypothesized evolutionary tree of amphibians, which was modified
somewhat by Grant et al. (2006) is summarized to the level of families in Essay 1.6. The
merits and demerits of this new taxonomy are currently the subject of considerable debate
among herpetologists. One group (e.g., Frost et al. 2006) emphasizes the need to move to
a monophyletic taxonomy but retain traditional nomenclature which has been more-or-less
universally employed for 250 years. The other group (e.g., Cannatella and Hillis 2004; Hillis
2007) emphasize the need for a monophyletic taxonomy, but promote abandoning traditional
nomenclature for a new approach that they argue better reflects understanding of phylogeny,
while simultaneously arguing that caution is needed before adopting any changes prior to
more thorough investigation. Frost et al. (2006) acknowledge that their proposal will be sub-
ject to changes as more information becomes available, as has already happened in the case
of Grant et al. (2006). In addition, there are differences of opinion among authors regarding
how data are to be analysed, probably the most serious issue at hand. Although this will
be a period of intense instability, the end result should be a much improved understanding
of amphibian evolutionary relationships and a taxonomy that reflects this. Whatever the
outcome of this debate, it is clear that: a) the traditional classification of amphibian families,
which forms the basis of the analysis in this book, will not survive; and b) although there
will certainly be some changes to our understanding of the amphibian tree in coming years, Salamandridae. This family remains unchanged.
much of what is proposed by Frost et al. (2006) is likely to be adopted.
Much of the current debate concerning Frost et al. (2006) is focused on the generic, rather Frogs and Toads – Anura
than the family level. This debate focuses in particular on the splitting up of three large
genera, Eleutherodactylus, Bufo and Rana, but we do not discuss this further. It should be Leiopelmatidae. Frost et al. (2006) include the Ascaphidae in this family, returning to the
noted that the genus Craugastor was separated from Eleutherodactylus by Crawford and taxonomy prior to Ford and Cannatella (1993), but its content is otherwise unchanged. As
Smith (2005), and we do follow this change. In addition, the formerly large genus Hyla was a result, this family now contains six species, and its distribution is in New Zealand and
split by Faivovich et al. (2005), a change that we have included. the Pacific Northwest of North America, with the New Zealand species breeding by direct
Below, we go through the Frost et al. (2006) proposed new family structure for amphibians, development, and the North American ones by larval development.
as amended by Grant et al. (2006), in the order of the amphibian tree (Essay 1.6). Although
there are many changes at the family level, the most far-reaching and complex have to do
with the splitting up the Leptodactylidae and Ranidae, and the changes to these two families
also impact several families that were previously recognized.
Caecilians – Gymnophiona
Caeciliidae. Frost et al. (2006) include the Scolecomorphidae in this pantropical family,
but its content is otherwise unchanged.
Pipidae. This family remains unchanged.
Rhinophrynidae. This family remains unchanged.
 Chapter 1. An Introduction to the Amphibians
Alytidae. This is the revised name for the family Discoglossidae, the content of which
otherwise remains unchanged.
Bombinatoridae. This family remains unchanged.
Megophryidae. This family remains unchanged.
Pelobatidae. This family remains unchanged.
Pelodytidae. This family remains unchanged.
Scaphiopodidae. This family remains unchanged.
Heleophrynidae. This family remains unchanged.
Sooglossidae. Frost et al. (2006) include the Nasikabatrachidae in this family, consistent
with the phylogenetic placement of this taxon by Biju and Bossuyt (2003), but its content is
otherwise unchanged. As a result, this primitive family now contains five species, and its
distribution is in the Seychelles Islands and the Western Ghats of southern India.
Batrachophrynidae. Frost et al. (2006) separate this family from the Leptodactylidae
(consistent with work by San Mauro et al. 2005). It consists of three small genera (collectively
comprising just six species), Batrachophrynus, Caudiverbera and Telmatobufo, from southern
Chile and north into southern Andean Peru and Bolivia. These species are highly aquatic and
breed by larval development. Frost et al. (2006) suggested that Batrachophrynus might not
be in this group; should Batrachophrynus be found to be close to Telmatobius, the name for
this family will become Calyptocephalellidae.
Limnodynastidae. Frost et al. (2006) transfer the genus Mixophyes from this family to
the Myobatrachidae, but otherwise this family is unchanged. There are about 45 described
species from Australia and New Guinea.
11
Myobatrachidae. Frost et al. (2006) include the Rheobatrachidae in this family, and transfer
the genus Mixophyes to this family from the Limnodynastidae. There are almost 80 described
species from Australia and New Guinea.
Hemiphractidae. Frost et al. (2006) separate this family from the Leptodactylidae and
partitioned it further to address its non-monophyly as also suggested by Darst and Cannatella
(2004). It consists of one small genus, Hemiphractus (comprising six species), ranging from
Panama to the upper Amazon Basin. These species are highly specialized treefrogs from
primary rainforest that feed only on other species of frog. They breed by direct development,
with the eggs being carried on the female’s back.
Brachycephalidae. The nomenclatural effect of Frost et al. (2006) showing that Brachyc-
ephalus (Brachycephalidae) is imbedded within the Eleutherodactylinae (genera Adelophryne,
Atopophrynus, Barycholos, Dischidodactylus, Craugastor, Eleutherodactylus, Euparkerella,
Geobatrachus, Holoaden, Ischnocnema, Phrynopus, Phyllonastes and Phyzelaphryne) is
that the name of the combined large group becomes Brachycephalidae, a new grouping
of nearly 800 species covering all of South and Central America (except Chile and most of
Argentina, Paraguay and Uruguay) and the Caribbean islands, with a few species ranging
as far north as the southern United States. With the exception of Eleutherodactylus jasperi,
which is live-bearing, all members of this family so far examined lay eggs that develop
directly without a larval stage.
Cryptobatrachidae. Frost et al. (2006) separate this family from the Leptodactylidae. It
consists of two genera (collectively comprising 21 species), Cryptobatrachus and Stefania
confined to moderate to high elevations in northern South America (the Colombian Andes, the
Sierra Nevada de Santa Marta, and the Guianan Shield in Guyana, Venezuela and northern
Brazil). They exhibit direct development, with the eggs being carried on the female’s back.
12 Threatened Amphibians of the World

Oreophryne wapoga (Data Deficient) is a Leptodactylidae4. Frost et al. (2006) split the Leptodactylidae into nine families (also
microhylid frog from New Guinea in which comprising the traditionally recognized Brachycephalidae and Rhinodermatidae). These nine
the eggs develop directly without a free-living families are: Amphignathodontidae; Batrachophrynidae; Brachycephalidae; Ceratophryidae;
larval stage. The male parent guards the eggs Cryptobatrachidae; Cycloramphidae; Hemiphractidae; Leptodactylidae; and Thoropidae. How-
on the ground, and transports the juveniles on ever, Grant et al. (2006) merged the Thoropidae into the Cycloramphidae, split the Hylodidae
its back, as shown here. © Rainer Günther out of the Cycloramphidae, and split the remaining Leptodactylidae into Leptodactylidae
and Leiuperidae. Under this arrangement, the Leptodactylidae contains just four genera
(Hydrolaetare, Leptodactylus [including former Adenomera and Lithodytes], Paratelmatobius
and Scythrophrys) and about 75 species that breed in water with larval development (with
some species building foam nests). The family ranges widely in South and Central America
and the Caribbean islands, north to the southern United States, but is absent from Cuba,
Chile and southern Argentina.

Amphignathodontidae. Frost et al. (2006) separated this family from the Hemiphractidae.
It consists of two genera, Flectonotus and Gastrotheca (collectively comprising around
60 species), ranging from Costa Rica south to northern Argentina, southern Brazil, and
Trinidad and Tobago. These species, collectively termed marsupial frogs, possess a dorsal
pouch for brooding their eggs (which can develop with or without a free-living larval stage,
depending on the species).

Cycloramphidae. Frost et al. (2006) separate this family from the Leptodactylidae, and to
render it monophyletic included the traditionally recognized Rhinodermatidae within it. Sub-
sequently, Grant et al. (2006) added the genus Thoropa to this family, but removed the genera
Crossodactylus, Hylodes, and Megaelosia to the family Hylodidae. With these changes, the
Cycloramphidae consists of 12 genera (collectively comprising around 90 species), Alsodes,
Crossodactylodes, Cycloramphus, Eupsophus, Hylorina, Limnomedusa, Macrogenioglottus,
Odontophrynus, Proceratophrys, Rhinoderma, Thoropa and Zachaenus, occurring in southern
tropical and temperate South America. Most species breed by larval development, though in
some cases the larvae are terrestrial, or live in the splash zones of waterfalls.

Hylidae. This family remains unchanged.

Centrolenidae. Frost et al. (2006) include the Allophrynidae in this family, but its content
and overall distribution is otherwise unchanged.

Leiuperidae. Grant et al. (2006) separate this family from the Leptodactylidae. It consists
of five genera (collectively comprising around 70 species), Edalorhina, Physalaemus,
Pleurodema, Pseudopaludicola and Somuncuria, ranging from Mexico south to Argentina.
They breed by larval development, and some species make foam nests.

Ceratophryidae. Frost et al. (2006) separate this family from the Leptodactylidae. It consists
of seven genera (collectively comprising around 80 species), namely Atelognathus, Batra-
chyla, Ceratophrys, Chacophrys, Insuetophrynus, Lepidobatrachus, and Telmatobius, ranging
from Colombia south to Chile and Argentina. They breed by larval development.

Bufonidae. This family remains unchanged.

Hylodidae. Grant et al. (2006) separate this family from the Cycloramphidae. It consists
of three genera (collectively comprising nearly 40 species), Crossodactylus, Hylodes, and
Megaelosia, mainly restricted to southern Brazil, but ranging into nearby Argentina and
Paraguay. They undergo larval development.
 Chapter 1. An Introduction to the Amphibians 13

Aromobatidae. Grant et al. (2006) separate this family from the Dendrobatidae. Grant et al.’s Hyperoliidae. Frost et al. (2006) remove the genus Leptopelis from this family from Africa,
revision of the Dendrobatidae and Aromobatidae results in great changes in the allocation of Madagascar and the Seychelles, but its content is otherwise unchanged.
species to the traditionally recognized genera. The Aromobatidae consists of nearly 90 species
in five genera and three subfamilies, occurring in tropical South America, north to Nicaragua,
with one species on Martinique in the Lesser Antilles. The reproductive modes of these species
are the same as described for the traditionally recognized Dendrobatidae above.

Brevicipitidae. Frost et al. (2006) separate this family from the Microhylidae, following
Loader et al. (2004). There are five genera (Balebreviceps, Breviceps, Callulina, Probreviceps,
and Spelaeophryne) and 24 species occurring in eastern and southern Africa, from Ethiopia
south to South Africa. Where known, all species breed by direct development.
Dendrobatidae. Grant et al. (2006) separate the Aromobatidae from this family, leaving
nearly 160 species in 11 genera and three subfamilies, occurring widely in tropical South
America, north to Nicaragua. The reproductive strategies of these species are the same as
described for the traditionally recognized Dendrobatidae above.

Hemisotidae. This family remains unchanged.

Ptychadenidae. Frost et al. (2006) separate this family from the Ranidae. There are three
genera (Hildebrandtia, Lanzarana, Ptychadena) and 51 species occurring widely in Sub-Sa-
Microhylidae. Frost et al. (2006) removed the African family, Brevicipitidae, from this haran Africa and Madagascar. All species reproduce by larval development in water.
family, but otherwise its content remains unchanged, though it is now much less diverse in
Africa. The Microhylidae consists of at least seven very distinct subfamilies: Asterophryninae
(southern Philippines and Sulawesi to northern Australia; direct developing); Cophylinae
(Madagascar; non-feeding larvae); Dyscophinae (Madagascar; larval developing); Gas-
trophryninae (the Americas; larval developing); Melanobatrachinae (Eastern Arc Mountains of
Tanzania and the Western Ghats of India; larval developing); Microhylinae (South, Southeast
and East Asia; larval developing); Scaphiophryninae (Madagascar; larval developing), with
several genera still unassigned in Africa and Asia.

Ceratobatrachidae. Frost et al. (2006) separate this family from the Ranidae as phylogeneti-
cally distinct. There are six genera (Batrachylodes, Ceratobatrachus, Discodeles, Ingerana,
Palmatorappia, Platymantis) and almost 80 species occurring from southern China, though
Myanmar, adjacent Thailand, Peninsular Malaysia, northern Borneo, the Philippines, New
Guinea, the Admiralty and Bismarck Archipelagos, the Solomon Islands, Fiji and Palau. All
species breed by direct development.

Arthroleptidae. Frost et al. (2006) merge the Astylosternidae into this family, and also
transfer the genus Leptopelis from the Hyperoliidae to this family, which consists of
around 130 species, widely distributed in Sub-Saharan Africa. It includes both direct- and
larval-developing species.
14 Threatened Amphibians of the World
Micrixalidae. Frost et al. (2006) separate this family from the Ranidae. There is a single
genus (Micrixalus) and 11 species restricted to the Western Ghats of southern India. All
species exhibit larval development.
Phrynobatrachidae. Frost et al. (2006) separate this family from the Petropedetidae as
phylogenetically distant. There is one genus Phrynobatrachus (including Dimorphognathus
and Phrynodon and excluding Ericabatrachus, which Scott [2005] transferred to what is
now Pyxicephalidae) and almost 70 species occurring widely in Sub-Saharan Africa. Most
species breed by larval development.
Petropedetidae. Frost et al. (2006) make major changes to the content of this family. All the
existing genera in the family, except for Petropedetes and Arthroleptides, are moved to two
new families endemic to the Afrotropics, Phrynobatrachidae and Pyxicephalidae. However,
two genera are transferred from Ranidae to Petropedetidae: Conraua, and Indirana (this
last genus of 10 species occurring only in the Western Ghats of southern India). Under this
arrangement, Petropedetidae becomes a small family of just 26 species, 16 in tropical Africa
and 10 in India. All species breed by larval development, and many have larvae associated
with the splash zones of waterfalls.
Pyxicephalidae. Frost et al. (2006) establish this new family from genera previously
assigned to Petropedetidae and Ranidae, largely consistent with a monophyletic group
first recognized by Van der Meijden et al. (2005). It comprises 13 genera (Amietia, Afrana,
Anhydrophryne, Arthroleptella, Aubria, Cacosternum, Ericabatrachus [transferred to this
group by Scott 2005], Microbatrachella, Natalobatrachus, Nothophryne, Poyntonia, Pyxi-
cephalus, Strongylopus, and Tomopterna) and 61 species that are widely distributed in
Sub-Saharan Africa. Most of the species breed by larval development, but some are
direct developers.
Dicroglossidae. Frost et al. (2006) establish this new family from genera previously
assigned to Ranidae, but found to be distantly related to that group. It includes 11 genera
(Annandia, Eripaa, Euphlyctis, Fejervarya, Hoplobatrachus, Limnonectes, Minervarya, Nan-
nophrys, Nanorana, Occidozyga, Ombrana, Quasipaa, and Sphaerotheca) and 143 species
that are widely distributed in South, East and Southeast Asia (as far east as the Philippines
and the Lesser Sunda Islands), with one species in Sub-Saharan Africa and another on the
Arabian Peninsula. All species breed by larval development.
Nyctibatrachidae. Frost et al. (2006) establish this new family from genera previously
assigned to Ranidae. It is composed of two genera (Nyctibatrachus, Lankanectes) and 13
species that are endemic to Sri Lanka and the Western Ghats of southern India. All species
breed in water with larval development.
Ranidae. Frost et al. (2006) split the Ranidae into eight families (also comprising the tradi-
tionally recognized Petropedetidae) to remedy the polyphyly of the Ranidae as traditionally
recognized. These eight families are: Ceratobatrachidae; Dicroglossidae; Micrixalidae;
Nyctibatrachidae; Petropedetidae; Ptychadenidae; Pyxicephalidae; and Ranidae. Under
this arrangement, the Ranidae contains just eight genera (Amnirana, Amolops, Huia,
Meristogenys, Pseudoamolops, Pterorana, Rana and Staurois, though Frost et al. [2006]
redelimited and recognized several new genera) and 310 species that breed in water with
larval development. The family ranges widely throughout Eurasia and tropical Asia, to north-
ern Australia and the Solomon Islands, throughout North America south to northern South
America, and also in tropical Africa. It is absent from the Caribbean islands, Madagascar
and New Zealand, and from most of southern Africa, Australia and most of South America
south of the Amazon basin.
Mantellidae. This family remains unchanged, although Glaw and Vences (2006) have
recently provided a completely revised taxonomy that bears little resemblance to what
preceded it.
Rhacophoridae. This family remains unchanged.
In summary, under the revisions of Frost et al. (2006) and Grant et al. (2006), nine tradition-
ally recognized families disappear (Scolecomorphidae, Uraeotyphlidae, Dicamptodontidae,
Ascaphidae, Nasikabatrachidae, Rheobatrachidae, Allophrynidae, Rhinodermatidae,
Astylosternidae), and 17 new families are established (Batrachophrynidae, Hemiphractidae,
Cryptobatrachidae, Amphignathodontidae, Ceratophryidae, Cycloramphidae, Leiuperidae,
Hylodidae, Aromobatidae, Brevicipitidae, Ptychadenidae, Ceratobatrachidae, Micrixalidae,
Phrynobatrachidae, Pyxicephalidae, Dicroglossidae, Nyctibatrachidae). The number of
amphibian families increases from 48 to 56.
Reproduction, Parental Care, and Metamorphosis
The characteristic most closely associated with amphibians is their ability to live in both
terrestrial and freshwater environments – they are amphibious. Movement of amphibians
between land and water is commonly linked to breeding activity. Adults move from terrestrial
habitats to spawn in water, the resulting free-living larvae are aquatic, and fully developed
young return to land following the process of metamorphosis. However, there are many ex-
 Chapter 1. An Introduction to the Amphibians
ceptions to this lifestyle, including species that complete their entire live cycles in terrestrial
habitats (such as many caecilians and the Puerto Rican Coqui Frog Eleutherodactylus coqui);
species that are wholly aquatic (e.g., frogs of the genus Pipa); and many species that undergo
direct development or are live-bearing, bypassing the free-living larval stage.
Amphibians use either internal or external fertilization. With very few exceptions, the
fertilization of eggs in frogs and toads is external, while the vast majority of salamanders
and newts, and probably all caecilians, have internal fertilization. Among the frogs and toads,
the only species known to exhibit forms of internal fertilization are members of the family
Ascaphidae, the live-bearing African toad genera Nimbaphrynoides and Nectophrynoides,
the African toad species Mertensophryne micranotis and the live-bearing frog from Puerto
Rico Eleutherodactylus jasperi. Within the salamanders and newts, the families Hynobi-
idae, Cryptobranchidae, and possibly the Sirenidae, have external fertilization (Duellman
and Trueb 1994).
The eggs of amphibians are commonly formed of one or several semi-permeable
gelatinous membranes surrounding the ovum. These membranes allow gasses and water
to pass freely through the egg and also protect the egg against damage, desiccation,
infection and predation. Eggs are most often deposited (oviposition) in water bodies or in
damp sites (such as underneath moist leaf-litter) where desiccation is unlikely to prevent
successful development.
The most familiar of amphibian larvae are the free-swimming ‘tadpoles’ of frogs
and toads. The body shape of most anuran larvae is roughly ovoid, with a long laterally
compressed tail used for swimming. The mouthparts are generally formed of upper and
lower horny beak-like jaws, with surrounding oral papillae. Anuran larvae are commonly
herbivorous with a gut that is often quite long in order to process plant matter. The larvae
of salamanders and newts tend to have a very similar morphology to that of the adults, with
noticeable external differences being their smaller size, external filamentous gills, flattened
tail fin, and the lack of a true tongue. The young of larval-developing caecilians are eel-like
and superficially similar in form to the adult animals. Caecilian larvae differ from the adults
in having a tail fin, a spiracle and gill slits, lateral line organs and expanded lips for suction
feeding. While some caecilians hatch with external gills, these are generally reabsorbed
into the body or shed within a few days.
Reproductive modes
While most amphibian species lay eggs (oviparous), there are in general three main breed-
ing modes: those that lay eggs and have a free-living larval stage; those that lay eggs and
undergo direct development, circumventing the larval stage; and the non-egg laying live-
bearing (viviparous and ovoviviparous) species, in which the young develop completely, or
to some large degree, within the female.
Having a larval stage is by far the most common, and perhaps the most familiar, amphibian
reproductive mode. This mode of reproduction has been recorded in more than two-thirds
of the world’s ~6,000 amphibian species (see Chapter 4), perhaps being completely absent
only from the families Brachycephalidae, Leiopelmatidae, Typhlonectidae and Scolecomor-
phidae (though in several families, such as Arthroleptidae, Leptodactylidae, Microhylidae,
Plethodontidae and Rhacophoridae, many species do not have a larval stage). In the majority
of species, the eggs are laid within a suitable water-body, which may range in size from a
leaf-axil to a large lake, and hatch into free-living aquatic larvae (tadpoles) that complete
their development and undergo metamorphosis before leaving the water.
In the majority of direct-developing species, a reproductive mode that is particularly
common within the families Arthroleptidae, Caeciliidae, Leptodactylidae, Microhylidae,
Plethodontidae and Rhacophoridae, the eggs are deposited in a damp or moist terrestrial
site (such as underneath leaf-litter, bark or rocks), where they complete development,
sometimes under the umbrella of paternal care, and hatch as fully formed young. Advantages
to a direct-development strategy include reduced vulnerability to aquatic predators and to
drying up of wetlands.
A very few amphibian species (less than 2% of all species) have a live-bearing re-
productive mode. Ovoviviparous species include all species in which the young undergo
development within the female obtaining nourishment from the yolk alone. In viviparous
species the developing young obtain additional nourishment from the female. In most
ovoviviparous or viviparous species very few well-developed young are born. Many of
the known live-bearing amphibian species are caecilians of the families Caeciliidae (28
species) and Scolecomorphidae (three species). Within the salamanders and newts there
are 12 live-bearing species in the genera Salamandra and Lyciasalamandra (some species
of Salamandra give birth to free-living larvae that complete their development in water).
There are few known live-bearing frog and toad species (14 species in total), relative to the
diversity of the Order as a whole. Most of these species, such as the Critically Endangered
Kihansi Spray Toad Nectophrynoides asperginis and Golden Coqui Eleutherodactylus jasperi
are ovoviviparous, with the equally highly threatened Nimbaphrynoides liberiensis and
Nimbaphrynoides occidentalis being the only known truly viviparous frogs.
For the purposes of analysis in this book, amphibian reproductive modes have been
combined into three general groups: larval developing - species that lay eggs, from which
free-living larvae hatch; direct developing – species that lay eggs in which the young develop
into the adult form, and undergo any metamorphosis prior to hatching; and live-bearing
– species that do not lay eggs, and give birth to young animals.
Parental Care
While most amphibian species show very little or no parental care after initial egg deposition,
there are a number of instances where the care of both eggs and larvae is highly developed.
Parental care can include the production of foam nests in which the eggs develop (Anura),
driving away predators (including conspecifics), moistening and aeration of the eggs, oscil-
lation (or turning) of eggs, the removal of damaged or infected eggs from the clutch, and
possibly the application of protective skin secretions to prevent pathogen growth (Duellman
and Trueb 1994; Stebbins and Cohen 1995). Some well-known examples of protection in-
clude: the African Bullfrog Pyxicephalus adspersus that will attack much larger animals and
people that come close to the nest site; the construction and guarding of shallow nesting
basins by males of the frog Hypsiboas rosenbergi; the large aquatic Hellbender Salamander
Cryptobranchus alleganiensis that defends eggs against predation; and, where known, the
terrestrial eggs of oviparous caecilians are attended by the female until they hatch into
aquatic larvae (Duellman and Trueb 1994; Stebbins and Cohen 1995).
In some instances the eggs or larvae are actively transported by one of the adults. Fol-
lowing amplexus, the strings of eggs of midwife toads (Alytes spp.) are collected on the
hind legs of the male, which are carried (with periodic moistening in water) until the larvae
begin to hatch. The male then transports them to water where they hatch and complete
their development. Many of the species of dendrobatid frogs from the Neotropics (including
a number of the well-known Poison Frogs) show quite advanced parental care. In addition
15
The young of the caecilian Boulengerula
to the adults guarding the terrestrial site of egg deposition, the larvae upon hatching are taitana (Least Concern) have been found
typically transported on the back of one of the adults to a stream or water-filled bromeliad to nourish themselves by stripping and
where development is completed. Similar behaviour has also been recorded in two ranid frogs eating the outer layer of the mother’s skin.
(Limnonectes finchi and L. palavanensis) from Borneo, the sooglossid Sooglossus seychel- © Alexander Kupfer
lensis (in which larval development is completed on the adults’ back), in the Leiopelmatidae
(in which the males of Leiopelma archeyi, L. hamiltoni and L. pakeka transport the young on
their back), and most recently in the New Guinea microhylids Liophryne schlaginhaufeni,
Sphenophryne cornuta, Oreophryne cf. wapoga and the newly described Callulops pullifer
(Inger et al. 1986; Inger and Voris 1988; Duellman and Trueb 1994; Stebbins and Cohen 1995;
Günther et al. 2001; Bickford 2002; Günther 2006).
Forms of brooding developing eggs within pouches or body cavities have evolved
in a number of amphibian species. For example, the eggs of the well-known aquatic
toads of the genus Pipa develop on the back of the female, with either larvae or toadlets
emerging from under the skin. Several leptodactylid genera (Hemiphractus, Gastrotheca,
Flectonotus, Stefania) contain species in which the eggs are carried in pouches, or are
glued to the back of the adult. Depending on the species, the eggs either develop directly
into froglets, or are released from the adult as well-developed non-feeding larvae. The
hatching larvae of the monotypic Australian genus Assa climb into hip pouches on the male,
where they may remain for around two months feeding on the yolk sac before emerging
as fully developed froglets.
The two species of the genus Rhinoderma from southern Argentina and Chile are the
only known examples of vocal sac brooding amphibians. The male Darwin’s Frog Rhinoderma
darwinii collects recently hatched larvae into the vocal sac where they complete their
development, eventually emerging from the male’s mouth as froglets. Similarly, the two,
now extinct gastric-brooding species of Rheobatrachus from eastern Australia were unique
in that the female picked up the eggs that then completed their development to froglets
entirely within the female’s stomach (see Essay 6.1). As with Darwin’s Frog, Rheobatrachus
froglets emerged from the mouth once fully developed.
There are a few instances known of amphibians actively providing nourishment to the
developing larvae. Females of a number of species in the genus Dendrobates regularly lay
unfertilized eggs into the bromeliads and leaf axils in which their larvae are developing.
The larvae rely on these eggs as a source of nutrition, and may not survive if the eggs
are not deposited. The developing larvae of the Critically Endangered Mountain Chicken
Leptodactylus fallax feed exclusively on unfertilized eggs provided by the female; with as
many as 10,000 to 25,000 unfertilized eggs deposited in total (Gibson and Ley 2004). The
young of the caecilian Boulengerula taitana have recently been found to nourish themselves
by stripping and eating the outer layer of their mother’s skin. This form of parental care
might be common in direct-developing caecilians, but is not known from other amphibians
(Kupfer et al. 2006).
Metamorphosis
The process of metamorphosis from the larval stage to adult form is often dramatic in
amphibians. In general, an internal release of the hormone thyroxine prepares the organs
and tissues of the larval amphibian for metamorphosis. The timing of the thyroxine release
is largely determined by both environmental factors (such as overcrowding or predation
levels) and chemical factors, including a reduction in the corticoid hormones that inhibit
thyroxine release (Zug et al. 2001).
The transformation is most striking in the frogs and toads. During metamorphosis, the
larvae begin to develop all their limbs (including the strong hind legs), larval mouthparts
are replaced by true jaws and teeth, the tail is reabsorbed, the glandular outer skin of the
adult develops, the gut changes to process a more carnivorous diet, the lungs continue their
development, and the skeleton hardens. As salamander young often closely resemble the
adults, metamorphosis appears to be less remarkable. However, significant changes do take
place, including the reabsorbtion of the external gills and the tail fin, and the development
of a true tongue. In some salamanders and newts, adults may retain juvenile characteristics
(such as external gills); this phenomenon is termed paedomorphism. Caecilian larvae are
superficially similar to adult animals, but metamorphic changes may be substantial including
16 Threatened Amphibians of the World
This image of a larval Marbled Salamander
Ambystoma opacum (Least Concern) shows
clearly the large filamentous gills used for
respiration. © Henk Wallays
a thickening of the skin and development of scales, changes in the cranium and glossal
skeleton and associated musculature, loss of labial folds and gill slits, and the growth of
sensory tentacles.
Skin, Respiration and Thermoregulation
The distinctive semi-permeable skin of amphibians performs a number of important biological
functions. The outer skin (epidermis) is covered with mucous glands and serous granular
glands. The mucous glands help to keep the skin moist and slippery by secreting mucus.
The granular glands secrete alkaloids or other chemicals that are often toxic to potential
predators. The skin helps to protect the animal against physical damage (such as abrasion),
and antimicrobial peptides covering the skin prevent infection by pathogens (further details
of the possible values of these skin secretions are discussed in Chapter 2).
Amphibians occur in a wide variety of colours, ranging from the subtle shades of green,
brown and black used to camouflage animals, to the bright warning colours of the familiar
posion frogs. Pigmentation cells within the skin largely determine colour and shading. Pig-
ments may be dispersed or concentrated through hormonal action, allowing skin colour and
patterns to quickly change in response to the animal’s environment.
The skin can be an important respiratory organ. Most amphibians have a semi-perme-
able skin with both a low level of keratinization, and an extensive network of below-skin
blood capillaries adapted to cutaneous gas exchange (oxygen absorption through the skin).
The importance of this varies between species, but for some groups, such as the lungless
plethodontid salamanders, cutaneous gas exchange may account for 85-90% of total gas
exchange (Duellman and Trueb 1994; Stebbins and Cohen 1995). Some amphibians, such as
the giant salamanders of China and Japan (Andrias spp.), have evolved skin folds that may
enhance gas exchange through an increased skin surface area. Hair-like filaments develop
on the body of male Hairy Frogs Trichobatrachus robustus during the breeding season,
and these may aid oxygen uptake during times of increased demand. A single species of
caecilian is entirely lungless; the enigmatic Atretochoana eiselti is a giant aquatic species
known from only two specimens.
Most amphibians at some point in their development use gills for gas exchange. Larval
amphibians often have filamentous gills that are the primary means of respiration. These gills
are usually reabsorbed into the body during metamorphosis, while the adult lungs develop.
The lungs of amphibian species are ventilated by a buccopharyngeal pump accompanied by
exaggerated ‘gulping’ movements of the floor of the mouth. Lungs vary greatly in size and
structure, and this variation may be related to a number of factors including the importance
of other respiratory organs (such as the skin), thermal characteristics, activity, and habitat
and evolutionary characteristics (Stebbins and Cohen 1995). In general, amphibians inhabit-
ing well-oxygenated cold environments (for instance, mountain streams) are more likely to
have smaller or reduced lungs; species from warmer habitats with low oxygen levels more
commonly have larger lungs with a more complex structure. Some aquatic caecilians have
very well-developed lungs extending almost the entire length of their bodies that may be
important in buoyancy as well as in respiration. Gas exchange across the lining of the buccal
cavity and pharynx (mouth and throat) appears to be particularly important to a number of
salamanders, most especially the lungless species.
Unlike ‘warm-blooded’ birds and mammals (endotherms), amphibians are ectotherms or
‘cold-blooded’ animals, with the outside environment largely determining body temperature.
Both behavioural and physiological mechanisms regulate body temperature. Behavioural
mechanisms are mostly associated with movements to warmer or cooler sites, or microhabi-
tats, within the animal’s environment. This includes moving to basking sites where the animal
can raise its body temperature through exposure to the sun, or movements to shaded areas
away from the heat. Physiological adaptations include increasing or decreasing evaporative
cooling through the skin; changes in skin pigmentation in response to UV exposure; and pos-
sibly changes in the amount of blood flow to the skin. While very few amphibians can persist
at temperatures above 43ºC (110ºF), several species, such as the North American Wood
Frog Rana sylvatica, can tolerate freezing conditions through conversion of liver glycogen to
glucose in response to extracellular ice formation. Animals recover when milder temperatures
arrive by greatly increasing blood glucose levels (Stebbins and Cohen 1995).
As with thermoregulation, body water regulation in amphibians is controlled by both
behavioural and physiological mechanisms. The generally highly water permeable skin
of amphibians allows movement of water both into the body through osmosis when
the animal enters the water or is exposed to a moist surface, and away from the body
through evaporative transpiration when the animal is outside a wet or damp environment.
Individual animals can control the intake of water through the skin by entering or leaving
water, and many animals regularly move between terrestrial and wet or damp sites to
rehydrate themselves.
Body water regulation is most commonly associated with preventing water loss, rather
than coping with an excess of water. Amphibians have developed several strategies to
prevent water loss, including: having a largely nocturnal lifestyle; inhabiting moist places
when away from water; burrowing or undergoing periods of aestivation to avoid dry periods
when desiccation is a possibility; a reduction of the body area available for evaporative
transpiration; the retention of urea in plasma (especially among species from arid habitats);
and, in a few species, the excretion of nitrogenous waste as up to 80% uric acid (uricotelic
species). In some instances, amphibians have to adapt to excess, rather than a loss, of
water. Amphibians that are largely, or wholly, aquatic in freshwater for a lengthy period
tend to have modified skin and kidneys to prevent or compensate for excessive osmosis
from the surrounding environment.
Because of the loss of body water through osmosis in saltwater, there are no truly marine
amphibians. The frog Fejervarya cancrivora from Southeast Asia occurs in the brackish water
of mangrove forest, and is the closest to a marine amphibian.
Diet
With few exceptions, the diet of adult amphibians consists of a wide range of freshwa-
ter and terrestrial invertebrates. Most amphibians appear to be opportunistic feeders,
with the types of prey captured often dependent on availability and seasonality. While
most adult amphibians only consume invertebrates, some larger amphibians, such as
the African Bullfrog Pyxicephalus adspersus, can also catch small vertebrates such as
mice, birds, reptiles or other amphibians. Amphibians have also been recorded feeding
opportunistically on dead carrion, pet food, organic waste, and similar scraps. There are a
few well-recorded instances of amphibians in which the adult also consumes vegetables
or fruits. Adults of the treefrog Xenohyla truncata regularly consume small fruits, and as
some of the defecated seeds germinate it seems possible that this frog may contribute to
plant dispersal (da Silva et al. 1989).
Amphibian larvae have a greater diversity of feeding mechanisms and diet than adult
amphibians. The larvae of most frogs and toads are highly specialized filter feeders, us-
ing organs called the branchial food traps and gill filters to collect food. Trapped food is
transported from these traps to the oesophagus by mucal threads. Anuran larvae typically
fall into two groups, those that feed on microscopic matter (microphagous), and those that
feed on much larger objects (macrophagous). Microphagous larvae feed on food items such
as suspended algae or protists, and tend to have large branchial food traps. Macrophagous
larvae generally have both reduced branchial food traps and keratinized mouthparts adapted
to scraping and biting food, including algal mats or animal prey (including other amphibian
larvae). In contrast to the frogs and toads, the vast majority of salamander and newt larvae
and probably all caecilian larvae are carnivorous, and feed on aquatic invertebrates.
Movement and Migration
In general, the daily movement of amphibian species is mostly restricted to the vicinity
of the animals’ home range. Although the home range size of most amphibian species
remains unstudied or poorly known, in many cases it is believed that home range sizes are
typically quite small, being a total of only a few square metres in size. However, it should
be noted that amphibians have a wide range of dispersal abilities, and in some instances
are known to move greater distances than previously anticipated (Smith and Green 2005).
Possibly the most commonly recognized amphibian movements are the more substantial
seasonal migrations of large numbers of animals from their individual home territories to
and from breeding sites. The sight of mass breeding congregations of amphibians in ponds,
lakes and other water bodies is generally familiar to most people, and may often be their
first encounter with amphibians. Seasonal breeding migrations tend to be triggered by a
number of environmental conditions; these can include changes in daylight, temperature, or
in moisture and precipitation levels. For example, Gascon et al. (2003) mention a breeding
aggregation of more than 5,000 adults of the Marbled Salamander Ambystoma opacum in
a cluster of three ponds at the base of the Blue Ridge Mountains following the torrential
rains of Hurricane Dennis in September of 1999; the breeding event gave rise to over 12,000
metamorphic salamanders that emigrated en masse from ponds in the following May.
Movements most often take place at night, probably reducing the risk of predation by birds
and other diurnal predators (Zug et al. 2001).
While many amphibians may move only relatively short distances from the surrounding
terrestrial habitat to and from the breeding site, it is possible for some species to travel
longer distances to their traditional breeding site. Experiments with displaced individuals
of Red-bellied Newts Taricha rivularis have demonstrated that animals were able to locate
their breeding sites from as far as eight kilometres away (Twitty et al. 1967).
Amphibian movements and migrations are guided by a number of homing and orientation
mechanisms (Zug et al. 2001). Visual landmarks within the home territory or migration route
help to direct individual animals, especially to familiar escape routes when disturbed. Many
species are guided by olfactory cues, including the characteristic scents, smells or odours
of the home territory or breeding site. The sounds of calling conspecifics provide auditory
cues for frogs defending territories or trying to locate a mate. Variation in polarized light
allows amphibians to differentiate between wet and dry areas; in addition, salamanders
are able to orient themselves through the use of the pineal body which acts as a polarized
light receptor (Zug et al. 2001). Many amphibians are sensitive to magnetic fields, and a
number of studies into this interesting field have demonstrated that animals can orientate
themselves through the use of a light sensitive magnetic compass (including Phillips 1977,
1986a,b; Sinsch 1987; Deutschlander 2000; Freake and Phillips 2005). Two magnetoreception
systems have been found: the first measures spatial variation in the magnetic field and uses
this information to derive a geographic map of the area, while the second system is sensitive
to the horizontal alignment (azimuth) of the magnetic field and is used for compass alignment
(Freake and Phillips 2005). Recent studies have confirmed the presence of orientation by
magnetic fields in both the Caudata and Anura (Freake and Phillips 2005).
Communication
Amphibians generally communicate through vocal, chemical or visual means, or through
a combination of these three systems. Of the three Orders of amphibian, only the frogs
and toads tend to make significant vocalizations. While some species of caecilians and
salamanders can produce soft clicking, squeaking or hissing sounds, in general there are
very few voluntary sounds made. Although there are a few exceptions, most species of frogs
and toads tend to force air from the lungs over the larynx where the vocal cords vibrate
creating sound. In many cases the calls are transmitted via vocal sacs, with some external
vocal sacs having a balloon-like appearance. While further research is needed into the role
of the vocal sacs, these are often considered to both intensify or modify the acoustic signal
and assist in radiating calls in all directions.
The volume and intensity of calls can vary greatly by species, and this often depends on
whether the call is intended for nearby conspecifics or for those that are far away. Environ-
mental conditions may also determine the volume and intensity of the call. For example,
the threatened species Amolops tormotus from eastern China has recently been discovered
to emit calls in ultrasonic frequencies; this is possibly an adaptation to the loud torrential
stream habitats in which the animal lives (Feng et al. 2006). The Bornean frog Metaphrynella
sundana has been shown to actively exploit the acoustic properties of partially water-filled
tree cavities (Lardner and bin Lakim 2002). By tuning their vocalizations to the resonant
frequency of the hole, the calling males enhance their chances of attracting a female.
 Chapter 1. An Introduction to the Amphibians 17

Most communication between individual caecilians and salamanders appears to be
largely through chemical secretions and visual signs. Perhaps the most commonly employed
chemical signals are pheromones produced by courtship glands, which may be used to
distinguish between species and also to locate conspecifics (Zug et al. 2001). Visual or
tactile signals in amphibians have been observed in all three Orders and can include animals
biting, nudging or butting their partner, tail-whipping (especially in some European newts),
or using a foot-flagging display and colour displays especially where vocalizations may not
easily be heard (Zug et al. 2001; Hödl 2000).
Amphibian communication methods are very diverse and have several different functions.
Advisement calls and signals are used to attract a mate, to aggressively warn others of ter-
ritorial boundaries, or are made in response to encountering other males. Reciprocation calls
and signals are made by females willing to mate in response to the courtship of a particular
male. Unreceptive females give release calls or signals during attempts at amplexus, as do
males that have been mistakenly identified as females. Distress signals may be low-pitched
shrieks or shrill cries, or specific movements or chemicals, which are produced by either sex
in response to disturbance or fright.
FURTHER GENERAL READING ON AMPHIBIANS
Duellman, W.E. and Trueb, L. 1994. Biology of Amphibians. McGraw-Hill, New York, USA.
Hofrichter, R (ed.) 2000. The Encyclopedia of Amphibians. Key Porter Books Limited, Toronto, Canada.
Stebbins, R.C. and Cohen, N.W. 1995. A Natural History of Amphibians. Princeton University Press,
Princeton, New Jersey, USA.
Zug, G., Vitt, L.J. and Caldwell, J.P. 2001. Herpetology: An Introductory Biology of Amphibians and Reptiles,
Second Edition. Academic Press, San Diego, USA.
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11. The Field Museum, Chicago, Illinois, USA.
Pombal, J.P., Jr. and Gasparini, J.L. 2006. A new Brachycephalus (Anura: Brachycephalidae) from the Atlantic
rainforest of Espírito Santo, southeastern Brazil. South American Journal of Herpetology 1(2):87-93.
San Mauro, D., Vences, M., Alcobendas, M., Zardoya, R. and Meyer, A. 2005. Initial diversification of living
amphibians predated the break-up of Pangaea. American Naturalist 165:590-599.
Scott, E. 2005. A phylogeny of ranid frogs (Anura: Ranoidea: Ranidae), based on a simultaneous analysis
of morphological and molecular data. Cladistics 21:507-574.
Seimon, T.A., Seimon, A., Daszak, P., Halloys, S.R.P., Schloegel, L.M., Aguilar, C.A., Sowell, P., Hyatt, A.D.,
Konecky, B. and Simmons, J.E. 2006. Upward range extension of Andean anurans and chytridiomycosis
to extreme elevations in response to tropical deglaciation. Global Change Biology 12:1-12.
Sinsch, U. 1987. Orientation behavior of toads (Bufo bufo) displaced from breeding sites. Journal of
Comparative Physiology A. 161:715-727.
Smith, M.A. and Green, D.M. 2005. Dispersal and the metapopulation paradigm in amphibian ecology and
conservation: are all amphibian populations metapopulations? Ecography 28:110-128.
Stebbins, R.C. and Cohen, N.W. 1995. A Natural History of Amphibians. Princeton University Press,
Princeton, New Jersey, USA.
Twitty V., Grant, D. and Anderson, O. 1967. Initial homeward orientation after long-distance displacements
in the newt Taricha rivularis. Proceedings of the National Academy of Sciences 57:342-348.
Van der Meijden, A., Vences, M., Hoegg, S. and Meyer, A. 2005. A previously unrecognized radiation
of ranid frogs in southern Africa revealed by nuclear and mitochondrial DNA sequences. Molecular
Phylogenetics and Evolution 37:674-685.
Zug, G., Vitt, L.J. and Caldwell, J.P. 2001. Herpetology: An Introductory Biology of Amphibians and Reptiles
Second Edition. Academic Press, San Diego, USA.

Paratelmatobius lutzii is an example of a

Brazilian frog that is currently listed as Data
Deficient, but which should probably be con-
sidered as Critically Endangered. It was once
common in an area of a few square metres on
the Alto do Itatiaia, at 2,200m asl in the Serra
da Mantiqueira, on the border of the states of
Minas Gerais and Rio de Janeiro. However, it
has not been seen since 1978, despite suit-
able habitat remaining and significant survey
efforts. © Ivan Sazima
18 Threatened Amphibians of the World

ESSAY 1.1. TRENDS IN RATES OF AMPHIBIAN SPECIES DESCRIPTIONS

Since the official implementation of zoological nomenclature by Carl von than three times the number of species descriptions compared with the same example, 223 species are officially known from Madagascar (December 2005),
Linné in 1758 there has been a steady increase in the number of recognized period 100 years earlier. The average annual number of amphibians described yet we have clear evidence (from morphology, bioacoustics and genetics) for
extant amphibian species. However, rates of species descriptions over time during the 1990s was about 81 species; the number of described species in the existence of at least 282 species and know of 45 further forms which
have been neither constant nor decreasing – as one would expect when 2005 alone exceeds 200, a record that was never reached before. One factor are likely to represent new species as well, indicating that about 100 new
assuming that the number of undiscovered species decreases with a longer contributing to this high rate of description was the publication of at least Madagascan species still await their description at the current time.
period of scientific investigation. Unlike other, well-known, vertebrate groups 35 new species from Sri Lanka (Manamendra-Arachi and Pethiyagoda 2005, Given this situation, and the strongly increasing description rates in many
such as mammals and birds,, amphibians are a class where species numbers Meegaskumbura and Manamendra-Arachi 2005). tropical areas of the world, we believe that an eventual doubling of the current
have increased exponentially, especially in recent decades (Glaw and Köhler New species descriptions can be attributed, in part, to known populations number of known amphibian species to around 12,000 is not inconceivable,
1998, AmphibiaWeb 2006). of described species that have been found to be genetically or bioacoustically though this global amphibian inventory may still require an additional 50 years
Since the early times of amphibian classification, the major works of a (in frogs) distinct, and may in many cases be recognized as different spe- to be completed. In the meantime, since many of our currently recognized
few active taxonomists contributed significantly to descriptions of novel taxa, cies. Modern molecular and bioacoustic techniques that provide increased amphibian “species” may actually prove to represent species complexes, this
specifically Josephus N. Laurenti, François Daudin, Johann Baptist von Spix, ‘resolution’ have revealed that in many cases several morphologically will have practical relevance for conservation. Taxonomic revisions will likely
André M. Constant Duméril and Gabriel Bibron, Albert C. L. G. Günther, Edward similar, and hence, cryptic species may be concealed within a single taxon have an important bearing on the Red List status of many species, particularly
D. Cope, Wilhelm Peters, George A. Boulenger and Oskar Boettger. With the name. However, compared with true first-hand discoveries in amphibians, if formerly widespread non-threatened species complexes are actually shown
increasing number of researchers dealing with the classification of amphib- removal from synonymy constitutes a relatively small portion (~14%) of to represent a number of smaller-ranged threatened species. In addition, the
ians and the intensified exploration of tropical regions, taxonomic work on newly recognized species (Köhler et al. 2005). Indeed, a large proportion non-cryptic ‘real’ discoveries of the future are far more likely to be taxa having
amphibians experienced its first major boost and description rates of species of new species are genuine new discoveries, as exemplified by the recent restricted ranges, with the result that these species are more likely to qualify
per period steadily increased until the end of the 19th century. For example, spectacular findings of a new frog family, genus and species in India, as threatened. These two factors suggest that future estimates of the degree
about 260 species of amphibians were described in the last decade of the 19th the Nasikabatrachidae (Biju and Bossuyt 2003), and of a new genus and of threat among amphibians will be much higher due to factors unrelated to
century, many of which are still considered to represent valid taxa. species of plethodontid salamander, Karsenia, in Korea – the first Asian habitat destruction and other human-induced declines.
Subsequently, a decrease in species description rates occurred during representative of this family (Min et al. 2005). A case study of mantellid frogs
two periods of the last century. These periods correlate with the First and in Madagascar indicated that newly discovered species since 1990 were Jörn Köhler, Frank Glaw and Miguel Vences
Second World Wars, respectively, with obviously low research activity (Glaw as genetically divergent as those described in previous research periods.
and Köhler 1998). However, the general long-term trend was still an increase Additionally, most had not been collected previously, indicating that the
in description rates. increase in new amphibian species on this island of endemism is not a sign References
More recently, a second major boost of new species discoveries and of taxonomic inflation due to exaggerated splitting approaches or different
descriptions has been in evidence. Whereas at the end of 1992, the number species concepts (Köhler et al. 2005). AmphibiaWeb: Information on amphibian biology and conservation. [web application].
of amphibians recognized stood at only 4,533 species (Duellman 1993, Glaw If the current trend in species descriptions continues, we probably face 2006. Berkeley, California: AmphibiaWeb. Available: http://amphibiaweb.org/.
et al. 1998), as of July 2006, this total stood at 6,041 species (AmphibiaWeb a decade (2000-2009) in which more than 1,500 amphibian species could be Biju, S.D. and Bossuyt, F. 2003. New frog family from India reveals an ancient biogeographi-
2006). This equates to an increase of more than 32% in only 13 years. The described. This raises the interesting question of just how long will this trend cal link with the Seychelles. Nature 425:711-714.
absolute number of newly described amphibian species per decade (not only continue and how many amphibian species are there on earth? Of course, Duellman, W.E. 1993. Amphibian species of the world: additions and corrections. University
the cumulative number of valid and described species) has been steadily nobody knows the final number and there is no available method to give an of Kansas Museum of Natural History, Special Publications 21:1-372.
increasing since the 1960s, with especially steep increases since the 1990s exact estimate. We suspect that at least several hundred new species remain Glaw, F. and Köhler, J. 1998. Amphibian species diversity exceeds that of mammals.
(Glaw and Köhler 1998, Köhler et al. 2005). No fewer than 810 amphibian to be described from throughout the tropical regions (including Madagascar, Herpetological Review 29:11-12.
species were described in the last decade of the 20th century; this is more India, Sri Lanka, Indonesia, Philippines, New Guinea, and South America). For Glaw, F., Köhler, J., Lötters, S. and Vences, M. 1998. Preliminary list and references
of newly described amphibian species and subspecies between 1993 and 1997.
Elaphe 6:1-24.
1,600
Köhler, J., Vietes, D.R., Bonett, R.M., Hita Garcia, F., Glaw, F., Steinke, D. and Vences, M.
2005. New amphibians and global conservation: a boost in species discoveries in a
highly endangered vertebrate group. BioScience 55:693-696.
Manamendra-Arachi, K. and Pethiyagoda, R. 2005. The Sri Lankan shrub-frogs of the genus
1,200
Philautus Gistel, 1848 (Ranidae: Rhacophorinae), with description of 27 new species.
Number of species

Raffles Bulletin of Zoology. Supplement No. 12:163-303.

Meegaskumbura, M. and Manamendra-Arachi, K. 2005. Description of eight new species
800 of shrub frogs (Ranidae: Rhacophorinae: Philautus) from Sri Lanka. Raffles Bulletin of
Zoology. Supplement No. 12:305-338.
Min, M.S., Yang, S.Y., Bonett, R.M., Vieites, D.R., Brandon, R.A. and Wake, D.B. 2005.
400 Discovery of the first Asian plethodontid salamander. Nature 435:87-90.

0
Figure 1. Descriptions of new amphibian species per decade until July
1758-59

1760-69

1770-79

1780-89

1790-99

1800-09

1810-19

1820-29

1830-39

1840-49

1850-59

1860-69

1870-79

1880-89

1890-99

1900-09

1910-19

1920-29

1930-39

1940-49

1950-59

1960-69

1970-79

1980-89

1990-99

2000-09

2006 on a global scale (including taxa considered to be valid species today

Research period only). A steady increase of the numbers is recognizable since 1940. The
Actual Predicted yellow bar shows the expectation for the period 2000-2009 if the rate of
2000-2006 continues.

ESSAY 1.2. AMPHIBIAN DIVERSITY AND THE CASE OF SRI LANKA’S BURGEONING INVENTORY

Biodiversity exploration in Sri Lanka, a 65,000-km2 continental island situated of these new species have been formally named (e.g., Manamendra-Arachchi
50
immediately south of India, began with the arrival of Portuguese colonists in and Pethiyagoda 2005; Meegaskumbura and Manamendra-Arachchi 2005)
1505. The country had, from pre-Roman times, been an important source of and as many as 50 more are in the process of description.
spices, including black pepper and cardamom. European demand for medicinal The vast majority of the novelties discovered in Sri Lanka belong to the 40
plants and spices was growing, and sporadic botanical investigations were Oriental shrub-frog genus Philautus. These small frogs are ubiquitous in the
Number of species described

not long in getting under way. Natural history studies gathered momentum very humid forests of tropical Asia, and their tinkling vocalizations figure
30
with transfer of the island to the Netherlands in the early 17th century, the prominently in the nocturnal forest chorus. Philautus are direct-developing
first botanic gardens being established in 1669. In 1747, Carolus Linnaeus frogs: their eggs, deposited on leaves or in shallow nests excavated in the
wrote Flora Zeylanica (1747), his only tropical flora, on the then-known forest floor, hatch directly into near-fully metamorphosed froglets, bypassing 20
plants of the island. Inheriting the island from the Dutch in 1796, the British the ‘conventional’ aquatic tadpole stage. Although direct-development rarely
institutionalized biodiversity exploration by establishing a botanic garden at features in popular texts on amphibian biology, about 20% of the world’s an-
10
Peradeniya in 1822, and a natural-history museum in 1877. uran species are reported to show this developmental mode (Thibaudeau and
By 1993, when the Wildlife Heritage Trust (WHT) – a non-profit orga- Altig 1999). Many Sri Lankan Philautus appear to have remained undiscovered
nization dedicated to the scientific exploration and documentation of Sri for so long because they had been assumed to belong to a small number of 0
Lanka’s biodiversity – began a survey of the island’s amphibian populations, polymorphic species. However, the WHT studies have shown that the new 1758-1799 1800-1849 1850-1899 1900-1949 1950-1992 1993-2005

the island’s vertebrate fauna was generally regarded as well known. Every species were consistently distinct, not only in morphology, but also with Research Periods
group had benefited from taxonomic reviews – Kirtisinghe (1957), the major respect to bioacoustics and genetics (Meegaskumbura and Manamendra-
taxonomic work on amphibians before the WHT explorations, recognized Arachchi 2005) (Figure 2).
35 species – and there was no hint in the regularly published checklists The island’s species count presently stands at 103, of which 19 are Figure 1. Number of descriptions of new amphibian species from Sri Lanka
that significant novelties were to be expected. Indeed, the accretion of new categorized as Extinct, 11 Critically Endangered, 35 Endangered and 6 from the time of Linnaeus to the present.
amphibian species had slowed to a trickle, just four having been discovered Vulnerable. While loss of habitat is clearly the most immediate threat,
in the previous half-century (Figure 1). habitat fragmentation combined with on-going climatic change is likely to
The initial purpose of the WHT survey was to identify key habitats for result in many more species being considered threatened, especially those
amphibian conservation, with the knowledge that by 1993 Sri Lanka’s rapidly restricted to high altitudes. Although there is no evidence of Sri Lankan asl) in the central mountains, average annual temperature increased by
growing human population was causing ever-increasing expanses of forest to amphibians having been victim to the synergistic effects of climate change 1.3°C and average annual precipitation decreased by ~20% in the period
be converted to agriculture. However, within just a short period of time, the and disease (e.g., Pounds et al. 2006), on-going climatic change could stress 1869-1995 (Schaefer 1998).
WHT team began uncovering many novelties, and in 1998 announced that the shrub-frog populations. Bahir et al. (2005) showed that Philautus breed only The irony of so many new amphibian species being discovered when
island’s amphibian fauna could reach an unprecedented 250 species (Pethiya- during periods of sustained rainfall and continuously high relative humidity amphibian populations worldwide are declining has not escaped attention
goda and Manmendra-Arachchi 1998), an estimate since revised downward (80-100%). However, meteorological data from across Sri Lanka show clear (e.g., Hanken 1999): some 1,000 newly discovered species were described
to ~140 by Meegaskumbura et al. (2002). Between 1993 and the present, 42 warming and desiccation trends. For example, at Nuwara Eliya (1,800 m worldwide in the period 1992-2003 (Köhler et al. 2005; and see Essay 1.1).
 Chapter 1. An Introduction
introduction to the Amphibians
amphibians 19

Figure 2. Although Philautus hoffmanni (A,C) and P. asankai (B,D) are distinc- A. E.
30
tively coloured, these ‘sister species’ are morphologically so similar that they
key out as a single species. Philautus hoffmanni is restricted to altitudes of
about 1,250m in Sri Lanka’s Knuckles Hills, while P. asankai occurs only at alti- 20

tudes of 800-1,800m in the central massif, about 100km to the south. The two
populations are separated by a deep valley. Despite their superficial similarity, 10

the species are distinguished by their vocalizations (E,F), which differ in call
length, pulse rate, dominant frequency, fundamental frequency, pulse length,

kU
0
and the number of pulses per call. They also differ from each other by a 12S
and 16S mt-DNA divergence of 1.01%, a cytochrome-b divergence of 6.04%, -10
and a suite of subtle, but consistent, morphological characters.
-20

-30
The case of Sri Lanka suggests that much amphibian diversity, which is 0 0.1 0.2 0.3 0.4 0.5 0.6
rapidly becoming impoverished, remains to be discovered, even in those
15
regions deemed to be historically ‘well studied’. This is already evidenced by B.
ongoing studies in other megadiverse tropical countries, such as the Philip-
pines (e.g. Brown and Guttman 2002; and see Essay 7.3). There is an urgent
need, therefore, for further exploration and survey work in under-sampled
regions, re-analysis of old museum collections, the application of new 10

taxonomic methodologies, including molecular and vocalization analyses,

kHz
and for the much-needed rejuvenation of neglected taxonomic collections
(and see Essay 11.10).
5
Rohan Pethiyagoda, Madhava Meegaskumbura, Kelum
Manamendra-Arachchi and Christopher J. Schneider

0
References 0 0.1 0.2 0.3 0.4 0.5 0.6

Bahir, M.M., Meegaskumbura, M., Manamendra-Arachchi, K., Schneider, C.J. and

Pethiyagoda, R. 2005. Reproduction and terrestrial direct development in Sri Lankan
shrub frogs (Ranidae: Rhacophorinae: Philautus). The Raffles Bulletin of Zoology, C. F.
30
Supplement No. 12:339-350.
Brown, R.M. and Guttman, S.I. 2002. Phylogenetic systematics of the Rana signata complex
of Philippine and Bornean stream frogs: reconsideration of Huxley’s modification of 20

Wallace’s Line at the Oriental-Australian faunal zone interface. Biological Journal of

the Linnaean Society 76:393-461. 10

Hanken, J. 1999. Why are there so many new amphibian species when amphibians are

kU
declining? Trends in Ecology and Evolution 14:7-8. 0
Kirtisinghe, P. 1957. The amphibian of Ceylon. Published by the author, Colombo, Ceylon.
Köhler, J., Vieites, D.R., Bonett, R.M., García, F.H., Glaw, F. Skeinke, D. and Vences, M. -10
2005. New amphibians and global conservation: a boost in species discoveries in a
highly endangered vertebrate group. BioScience 55:693-696. -20
Manamendra-Arachchi, K. and Pethiyagoda, R. 2005. The Sri Lankan shrub-frogs of the
genus Philautus Gistel, 1848 (Ranidae: Rhacophorinae), with description of 27 new
-30
species. The Raffles Bulletin of Zoology, Supplement No. 12:163-303. 0 0.1 0.2 0.3 0.3 0.3 0.6 0.7 0.8 0.9
Meegaskumbura, M., Bossuyt, F., Pethiyagoda, R., Manamendra-Arachchi, K., Bahir,
15
M.M., Schneider, C.J. and Milinkovitch, M.C. 2002. Sri Lanka: an amphbian hotspot.
Science 298:379.
Meegaskumbura, M. and Manamendra-Arachchi, K. 2005. Description of eight new species D.
of shrub frogs (Ranidae: Rhacophorinae: Philautus) from Sri Lanka. The Raffles Bulletin
of Zoology, Supplement No. 12:305-338. 10

Pethiyagoda, R. and Manamendra-Arachchi, K. 1998. Evaluating Sri Lanka’s amphibian

kHz

diversity. Occasional Papers of the Wildlife Heritage Trust 2:1-12.

Pounds, J.A., Bustamante, M.R., Coloma, L.A., Consuegra, J.A., Fogden, M.P.L., Foster, P.N.,
La Marca, E., Masters, K.L., Merino-Viteri, A., Puschendorf, R., Ron, S.R., Sánchez- 5
Azofeifa, G.A., Still, C.J. and Young, B.E. 2006. Widespread amphibian extinctions from
epidemic disease driven by global warming. Nature 439:161-167.
Schaefer, D. 1998. Climate change in Sri Lanka? Statistical analyses of long-term tem-
perature and rainfall records. In: M. Domroes and H. Roth (eds), Sri Lanka: past and
0
present—archaeology, geography, economics—selected papers on German research, 0 0.1 0.2 0.3 0.3 0.3 0.6 0.7 0.8 0.9
pp. 102-117. Margraf Verlag, Weikersheim, Germany.
Thibaudeau, G. and Altig, G. 1999. Endotrophic anurans: development and evolution. In:
R.W. Mcdiarmid and G. Altig (eds.), Tadpoles: the biology of anuran larvae, pp. 170-188.
University of Chicago Press, Chicago, USA.

ESSAY 1.3. CAECILIANS

Of the three orders of living amphibians, caecilians (Gymnophiona) are the
least familiar to both amateur and professional biologists. Caecilians are
found in most moist tropical regions of the world except in Madagascar and
Australasia, and they extend into some adjacent subtropical areas in Indochina
and South America. There are c. 170 recognized caecilian species in three,
four or six families, according to alternative taxonomies. Species taxonomy
is still dominated by the work of E.H. Taylor (e.g., Taylor 1968). Subsequent
work relegated many of Taylor’s taxa to synonyms, but also increased the
rate of description of new species (e.g., Wilkinson and Nussbaum 2006).
The current taxonomy of many caecilians is confused, and unpublished work
indicates the need for further synonymy, but also justifies the expectation of
an eventual rise in the number of caecilian species once taxonomic revisions
are complete and as discoveries and descriptions of new species continue
(Gower and Wilkinson 2005; Wilkinson and Nussbaum 2006).
All caecilians lack limbs and limb girdles, and tails are very short or absent.
Other notable external features include annulated skin, eyes covered with
skin and sometimes also bone, and a pair of, sensory tentacles in front of
the eyes. The name Gymnophiona (‘naked snakes’) seems like a rather ap-
propriate description given their superficial snake- (or worm-) like form and
lack of external scales. However, many species produce scales in pockets
associated with the skin’s annulation (Zylberberg et al. 1980).
Despite their superficial uniformity, caecilians are intriguingly diverse in
morphology and ecology, as might be expected from a group that may have
its origins in the Mesozoic. For example, adult caecilians range from less
than 10 cm to more than 150 cm in total length, with fewer than 70 to more
than 270 vertebrae. This reflects the substantial variation in the degree of
elongation of caecilians. They can be dull or vividly coloured, uniform, striped
or mottled. There are also notable variations in dentition, annulation, the
presence and distribution of scales, and in the structure of the skull, trunk
musculature, and sensory, urogenital and cardiovascular systems. The adults
of the vast majority of known caecilian species are terrestrial and burrow
in soil and/or leaf litter. Some are dedicated subterranean burrowers, while
others are more surface-cryptic (e.g., Gower et al. 2004b). In addition, there
are semi-aquatic and fully aquatic forms (replete with fins for swimming),
and one aquatic species, Atretochoana eiselti (DD), is the largest lungless
tetrapod (Nussbaum and Wilkinson 1995).
Caecilian reproductive modes include terrestrial oviparity, with aquatic
larvae or direct development, and viviparity. Aquatic larvae have a small tail
fin, lateral-line organs, labial folds and spiracles, and they undergo substan-
tial metamorphosis. As far as is known, all male caecilians have the cloaca
modified into an eversible copulatory organ (e.g., Gower and Wilkinson 2002),
fertilization is internal, and the mothers of all oviparous species guard egg
clutches in subterranean chambers. Different types of viviparity have been
distinguished by the degree of dependence of newborns, and the amount and
type of any post-parturition maternal care (Loader et al. 2003). The foetuses
of some viviparous species are thought to feed on the lining of their mother’s
oviducts (Parker 1956), while the young of some species peel and eat their
mothers’ specially modified skin (Kupfer et al. 2006).
Caecilians are generally not as well studied as frogs and salamanders.
This unfortunate lack of attention is perhaps primarily a consequence of
their largely tropical distribution and cryptic habits. Another impediment to
progress is that the distribution of experts and active researchers in caecilian
biology rarely coincides with that of their study taxa. Perhaps only in India
are there more than a couple of caecilian biologists studying native taxa, and
this situation has underpinned the relatively recent and rapid advancements
in knowledge of this fauna (Gower et al. 2004a).
An astonishing two-thirds of all caecilian species are classified as Data
Deficient, according to IUCN Red list categories and criteria (Gower and
Wilkinson 2005). The majority of these taxa are Data Deficient by virtue of
their poorly characterized taxonomy, and a lack of data on distribution, ecol-
ogy, abundance, and response to threats. Caecilian ecology often demands
special field techniques that are rarely employed by vertebrate biologists,
but substantial progress has been made in recent years (see Gower and
Wilkinson 2005; Gower et al. 2006). However, with a handful of notable
exceptions (e.g., Kupfer et al. 2004, 2006; Measey 2004; Measey et al. 2003),
there have been no detailed studies of reproductive ecology and population
20
density - important parameters for accurate and precise conservation assess-
ments. To underpin their scientific conservation, further studies are needed
of caecilian reproduction and ecological requirements, and their response to
habitat alteration and climate change. Simultaneously, remaining taxonomic
problems must be resolved.
Many caecilians are thought to inhabit moist forests, and it is known that
some species have a biphasic life history with aquatic larvae. Thus, these spe-
cies might be thought of as vulnerable to chytridiomycosis (not yet studied in
caecilians) and deforestation – factors that threaten some other amphibians.
Some caecilians are known to thrive in certain agricultural landscapes, but it
is likely that others are adversely affected by habitat alteration and changing
land use (Gower and Wilkinson 2005).
At present, no caecilians are known to have become extinct (at least those
species described in the last few hundred years), and only five species are
considered threatened. The taxonomy of these five taxa seems relatively
stable, and they all occur in relatively small regions (an isolated Kenyan
Mountain, Sri Lanka, and the Seychelles). The extent of occurrence is an
important parameter in conservation assessments, but what sets the five
threatened caecilians apart from other sympatric species is that they are
relatively infrequently recorded, and/or their known habitats (native forest
and agricultural plots) are threatened by habitat loss or pollution. Three of
the five species are believed to have larvae that are dependent on water.
Although the majority of caecilians are classified as Data Deficient, there is
Typhlonectes compressicauda (Least Concern) is a member of a South
American group of aquatic and semi-aquatic caecilians. This species has a
wide distribution across Amazonian Venezuela, Colombia, Peru, Brazil and
the Guyanas. © Alexander Kupfer
ESSAY 1.4. THE MEGADIVERSE GENUS ELEUTHERODACTYLUS
The genus Eleutherodactylus, once the most speciose radiation of vertebrates on
earth, is beginning to be dismembered. The dismemberment is partially driven
by new molecular data showing that some other genera are intercalated within
Eleutherodactylus sensu lato (Brachycephalus and Phrynopus being the initial two
cases). However, the urge to dismember Eleutherodactylus is also driven by its
very large size and a desire to have smaller genera at hand. Most recently, there
has been a trend to recognize the genus Craugastor, which is largely restricted
to Middle America, contains more than 100 species, and has been viewed as
monophyletic for about twenty years (confirmed by more recent molecular work;
Crawford and Smith 2005; Darst and Cannatella 2004; Frost et al. 2006).
Figure 1. Richness map of amphibian species in the genera Eleutherodactylus
and Craugastor, with dark red colours corresponding to regions of higher
richness. Colour scale based on 10 quantile classes. Maximum richness
equals 48 species.
Threatened Amphibians of the World
no room for complacency, and the group as a whole is threatened by lack
of knowledge.
David J. Gower and Mark Wilkinson
References
Gower, D.J. and Wilkinson, M. 2002. Phallus morphology in caecilians and its systematic
utility. Bulletin of the Natural History Museum, Zoology Series 68:143-154.
Gower, D.J. and Wilkinson, M. 2005. Conservation biology of caecilian amphibians.
Conservation Biology 19:45-55.
Gower, D.J., Bhatta, G., Giri, V., Oommen, O.V., Ravichandran, M.S. and Wilkinson, M.
2004a. Biodiversity in the Western Ghats: the discovery of new species of caecilian
amphibians. Current Science 87:739-740.
Gower, D.J., Loader, S.P., Moncrieff, C.B. and Wilkinson, M. 2004b. Niche separation and
comparative abundance of Boulengerula boulengeri and Scolecomorphus vittatus
(Amphibia: Gymnophiona) in an East Usambara forest, Tanzania. African Journal of
Herpetology 53:183-190.
Gower, D.J., Oommen, O.V. and Wilkinson, M. 2006. Marking amphibians with alpha
numeric fluorescent tags – caecilians lead the way. Herpetological Review 37: 302
Kupfer, A., Nabhitabhata, J. and Himstedt, W. 2004. Reproductive ecology of female
caecilians (genus Ichthyophis): a baseline study. Biological Journal of the Linnean
Ichthyophis cf. tricolor. Species of the two genera (Ichthyophis and Caudacae-
cilia) of ichthyophiid caecilians are distributed across South and Southeast
Asia, west of Wallace’s Line, and are the only caecilians occurring in the
latter region. Taxonomic difficulties mean that the precise identification
of this specimen is unclear. As far as is known, all ichthyophiids retain the
presumably ancestral caecilian reproductive mode of oviparity with aquatic
larvae that metamorphose into terrestrial adults. © Ashok Captain
In its larger sense, Eleutherodactylus species are natively distributed
from the south-western part of the United States (the southernmost parts of
Arizona, New Mexico, and Texas) and from the Bahamas and Cuba to the south
and east in an unbroken tapestry until the northernmost (both north-east and
north-west) parts of Argentina. Species are known from sea level to at least
4,350m asl (in the Central Cordillera of Colombia). Some taxonomic discontinu-
ity/ fragmentation is apparent across this broad distribution. Some species
from Texas to Guatemala are assigned to the subgenus Syrrhophus (about 25
species of small frogs). From the south-western United States south and east
to western Colombia, Ecuador, and Venezuela one finds the frogs of the genus
Craugastor. These two generic groupings account for nearly all species of
Eleutherodactylus sensu lato known from Middle America except for perhaps
a dozen species of another group found in Lower Middle America (but better
represented in South America). From the Bahamas and Cuba through all of
the Greater Antilles, one finds a different group (subgenus Euhyas), which
scarcely even enters the Lesser Antilles. Human intervention has delivered
a few species of Euhyas onto the mainland (south-eastern United States,
Mexico, Nicaragua). Another group of a half-dozen species (subgenus Pelo-
rius) is restricted to Hispaniola. The third grouping in the Antilles (subgenus
Eleutherodactylus) includes species on all of the major islands and through
the Lesser Antilles. Some of these are widely introduced elsewhere (Bermuda,
Hawaii, northern South America). In spite of being the most notable Neotropi-
cal radiation, few works have treated the genus (sensu lato or sensu stricto)
as a whole (Lynch 1976, 2001; Lynch and Duellman 1997)
In South America, Eleutherodactylus species abound in the northern Andes,
but barely penetrate Peru except as an assortment of species distributed
along the eastern flanks of the Andes that then continues south to northern
Argentina. These frogs are abundant in the western lowlands of Colombia
and Ecuador, patchily in the inter-Andean valleys of Colombia, and are once
again diverse in the western part of the Amazon Basin. However, east of the
western fringe of the Amazon Basin, Eleutherodactylus becomes a scarce
biological component (or even disappears entirely in the Llanos of Colombia
and Venezuela as well as in the Cerrado formations of Brazil), with only a minor
flare-up in the Guianas. In the once-forested Atlantic domain of eastern Brazil
(and south to north-eastern Argentina) there is another focus of diversity, in
sharp contrast to the situation over most of Brazil.
The South American fauna has not been fragmented with generic (or
subgeneric) names as was the case for the Antilles and Middle America.
Nonetheless, a number of clades are evident. The species of the Atlantic for-
ests of Brazil, Uruguay and Argentina represent a single clade (Lynch 2001),
Society 83:207-217.
Kupfer, A., Müller, H., Antoniazzi, M.M., Jared, C., Greven, H., Nussbaum, R.A. and
Wilkinson, M. 2006. Parental investment by skin feeding in a caecilian amphibian.
Nature 440:926-929.
Loader, S.P., Wilkinson, M., Gower, D.J. and Msuya, C.A. 2003. A remarkable young
Scolecomorphus vittatus (Amphibia: Gymnophiona: Scolecomorphidae) from the North
Pare Mountains, Tanzania. Journal of Zoology 259:93-101.
Measey, G.J. 2004. Are caecilians rare? An East African perspective. Journal of East
African Natural History 93:1-19.
Measey, G.J., Gower, D.J., Oommen, O.V. and Wilkinson, M. 2003. A pilot mark-recapture
study of the caecilian amphibian Gegeneophis ramaswamii (Amphibia: Gymnophiona:
Caeciliidae) in southern India. Journal of Zoology 261:129-133.
Nussbaum, R.A. and Wilkinson, M. 1995. A new genus of lungless tetrapod: a radically
divergent caecilian (Amphibia: Gymnophiona). Proceedings of the Royal Society.
Ser. B 261:331-335.
Parker, H.W. 1956. Viviparous caecilians and amphibian phylogeny. Nature 178:250-252.
Taylor, E.H. 1968. The Caecilians of the world: a taxonomic review. University of Kansas
Press, Lawrence, Kansas, USA.
Wilkinson M, and Nussbaum, R.A. 2006. Phylogeny and classification of caecilians. In:
J.-M. Exbrayat (ed.), Reproductive Biology and Phylogeny of Gymnophiona, pp 39-78.
Science Publishers, Inc., Enfield, New Hampshire, USA.
Zylberberg, L., Castanet, J., and de Ricqles. A. 1980. Structure of the dermal scales in
Gymnophiona (Amphibia). Journal of Morphology 165:41-54.
Herpele squalostoma (Least Concern) is a member of the Caeciliidae, the
largest and most taxonomically confused family of caecilians. This species
has its eyes covered by skin and bone, and probably spends the vast majority
of its life underground. H. squalostoma is one of the few caecilian species to
appear occasionally in the pet trade in the West. © Alexander Kupfer
although some species of this clade extend westward to Bolivia. Another
clade, present in the north-western part of the continent, is composed of
species with broad heads and chunky toad-like bodies. The remaining spe-
cies are grouped into the subgenus Eleutherodactylus, with their distribution
centered on the Andes (and immediately adjacent lowlands) of Colombia
and Ecuador. This heterogeneous unit extends into Lower Central America
(at least to Honduras), east to the mouth of the Amazon River, and south
to Bolivia. Superimposed upon this pattern of distribution one finds some
taxonomic elements that have been separated from Eleutherodactylus
sensu lato on what could be viewed as tenuous grounds, for example,
Phrynopus in the Andes from Colombia to Bolivia, separated because they
lack digital disks.
At least in terms of Colombia and Ecuador, there is an obvious pattern
to occurrence and diversity. In those areas with a negligible dry season,
Eleutherodactylus communities achieve impressive diversities (to more than
20 sympatric species). However, as the dry season becomes more and more
marked (and longer), diversity decays rapidly and only one or two species
persist. When the dry season increases to three or more months duration,
Eleutherodactylus disappear from the frog community or must retreat
into enclaves (caves or phyllotemata) so as to survive the dry conditions.
The cause of this pattern is almost certainly the pattern of reproduction
employed by all known species (save the apparently extinct E. jasperi, once
an ovoviviparous species). Adults, juveniles, and egg masses occupy the
same ecological space and the embryos and small juveniles are especially
susceptible to desiccation. Embryos and juveniles tend to occur in the leaf
litter and the dry season is especially severe in that layer of the habitat (adults
may retreat to phyllotemata or caves and by virtue of their smaller surface
area/ volume ratio resist more effectively the harshness of the dry season
than the smaller juveniles).
Members of the genus are smallish frogs. The smallest species have
body-size maxima of about 10 mm and these very small species (8-18mm
body size) are typically confined to the wettest parts of the landscape. The
other extreme is occupied by species of Craugastor that reach between 100
and 130 mm body-size. Most of these occupy very wet habitats (such as the
Chocó), but some occur in lands that are or appear to be dry (for example,
parts of the Mexican highlands extending into the south-western part of the
United States where I once collected Craugastor augusti amidst cactus). The
generic name Eleutherodactylus means free toes – in allusion to the absence
of inter-digital membranes (webs – and this is true for the vast majority of
the species now assigned to Eleutherodactylus. However, a few species have

apparently never read Dumeril and Bibron or Boulenger and have obviously
webbed feet (for example, the Puerto Rican E. karlschmidti and the Colombian
E. anatipes and E. zygodactylus).
For many reasons, Eleutherodactylus are confusing to the non-specialist.
The confusion stems from several factors: (1) the sheer number of sympatric
species in several areas; (2) the difficulty in distinguishing between adults and
juveniles, compounded by the first factor; (3) the marked sexual dimorphism,
where for most species adult males are only about 75% the length of adult
females; and (4) the widespread phenomenon of pattern polymorphism,
especially evident in the exposed visible parts of the animal (dorsal surfaces
of the body, head and limbs). When diversity is modest (fewer than a half
dozen sympatric species), even the non-specialist becomes slowly comfort-
ESSAY 1.5. HYLIDAE
Hylidae, the treefrogs, is among the most species-rich family of amphibians, with
more than 800 described species. These include some species quite familiar to
the general public, such as common Green Treefrog Hyla cinerea and leaf frogs,
but also an incredible diversity of less well-known groups. Hylids are found
mostly in South and Central America, as well as in the Australo-Papuan Region,
but some species also occur in the Caribbean, North America, and temperate
Eurasia, including extreme northern Africa and the Japanese Archipelago.
Our knowledge about the relationships and composition of Hylidae has
experienced dramatic changes in recent years, in part thanks to the revela-
tions of DNA-based studies. For example, marsupial frogs, for a long time
considered to be hylids, have been shown by several analyses to be quite
unrelated with treefrogs, and have been shown to be unrelated to this family
in recent analyses (Haas 2003; Darst and Cannatella 2004; Faivovich et al.
2005; Wiens et al. 2005; Frost et al. 2006)
As it stands, Hylidae is now composed of three subfamilies. Phyllom-
edusinae, whose species are sometimes called “monkey frogs” or “leaf
frogs”, includes the poster frog of the amphibian world, the Red-eyed
Treefrog Agalychnis callidryas (LC), but also other charismatic species such
as Phyllomedusa sauvagii (LC), and the quite uncommon, but astonishingly
beautiful, Cruziohyla craspedopus (LC). This subfamily is distributed from
the tropical areas of Mexico to northern Argentina. It currently comprises
56 species divided into seven genera. A very interesting characteristic of
Phyllomedusinae is that most of its species lay eggs outside water. Eggs are
most frequently adhered to leaves (sometimes even completely enveloped
by them) or tree trunks, where the earlier developmental stages are spent,
and the tiny larvae then fall into the water body below them (be it a pond,
stream, or even the water trapped in a tree hole), where they undergo larval
development as free-living feeding larvae.
The subfamily Pelodryadinae includes all the Australo-Papuan treefrogs.
Among their better known representatives are Litoria caerulea (LC), and the
White-lipped Treefrog Litoria infrafrenata (LC). Its more than 170 species
are included in three genera: most (126) occur in the genus Litoria, with the
remaining members included either Cyclorana or Nyctimystes5. Pelodryadines
are amazing in that they diversified in complete isolation, occupying an exten-
sive range of habitats, and evolving multiple adult and tadpole morphologies
that are quite reminiscent to those of the subfamily Hylinae.
The subfamily Hylinae includes the core of hylid diversity. Its more than
590 species are divided into 34 genera and four tribes. The tribe Cophomantini
is restricted to the Neotropics and includes five genera. This tribe includes,
among others, the “gladiator frogs”, so called because males of several of its
species engage in violent territorial combats, assisted by a pre-pollex (one of
the bones of the hand) that is modified as a spine of variable size and shape.
The tribe Dendropsophini, whose identity still requires additional research,
includes seven genera restricted to the Neotropics. This tribe includes the most
aquatic hylids (genera Lysapsus and Pseudis), so peculiar that for a long time
they were included in a separate family (Pseudidae). It also includes the two
most species-rich genera of the subfamily, the taxonomically complex Scinax
and Dendropsophus (about 90 species each, and doubtless much more awaiting
description), and possibly the only known frugivorous frogs (genus Xenohyla).
Phyllomedusa nordestina, a representative of the subfamily Phyllomedusinae, is known from north-eastern Brazil.
The species was only recently described in 2006, and has yet to be assessed according to IUCN Red List criteria.
© Célio F.B. Haddad
Chapter 1. An Introduction
introduction to the Amphibians
amphibians
able with these frogs.
John D. Lynch
References
Crawford, A.J. and Smith, E.N. 2005. Cenozoic biogeography and evolution in direct-devel-
oping frogs of Central America (Leptodactylidae: Eleutherodactylus) as inferred from
a phylogenetic analysis of nuclear and mitochondrial genes. Molecular Phylogenetics
and Systematics 35:536-555.
Darst, C.R. and Cannatella, D.C. 2004. Novel relationships among hyloid frogs inferred
from 12S and 16S mitochondrial DNA sequences. Molecular Phylogenetics and
Systematics 31:462-475.
Litoria daviesae (Vulnerable), a representative of the subfamily Pelodry-
adinae, is distributed on the eastern edge of the tablelands and the great
escarpment of the Great Dividing Range of New South Wales, Australia.
© Marion Anstis
The tribe Lophiohylini is also restricted to the Neotropics. This tribe
includes 10 genera, many of which include species characterized by heavily
modified skull architecture, including fusion of several bones and unusual
ornamentations, earning them the name of “casque-headed frogs”. An
interesting characteristic of this tribe is that many of its species depend on
bromeliads and/or tree holes for refuge and even for reproduction, with one
genus (Phyllodytes) and some of the species of other genera (Osteocephalus,
Osteopilus) laying eggs in the water trapped by the leaf axils of bromeliads,
where the tadpoles then spend their entire larval life. The members of the
genus Nyctimantis and some species of Trachycephalus are suspected, or
known, to lay their eggs in water trapped in tree holes.
The tribe Hylini includes 16 genera and is mostly concentrated in Central
America and North America, with species of the genus Hyla also present
in temperate Eurasia, including extreme northern Africa and the Japanese
Archipelago. This diverse tribe includes common species such as the Green
Treefrog Hyla cinerea (LC), the cricket frogs (genus Acris), the unusual and
diverse Plectrohyla, from the mountains of Mexico, Guatemala, and Honduras,
and the rarely seen fringe-limbed species of Ecnomiohyla.
According to the results of the Global Amphibian Assessment, only a
single hylid is classed as Extinct, namely Phrynomedusa fimbriata, which
has not been recorded since it was first collected in 1898 in south-eastern
Brazil. In addition, the Global Amphibian Assessment identified another 123
hylids as “Critically Endangered” or “Endangered”; interestingly, of these,
81 are from the tribe Hylini (roughly 50% of the known species in the tribe!),
Bokermannohyla luctuosa (Least Concern), a representative of the subfamily Hylinae, is a common species from
south-eastern Brazil. © Célio F.B. Haddad
21
Frost, D.R., Grant, T., Faivovich, J., Bain, R., Haas, A., Haddad, C.F.B., De Sá, R.O., Channing,
A., Wilkinson, M., Donnellan, S.C., Raxworthy, C., Campbell, J.A., Blotto, B.L., Moler, P.,
Drewes, R.C., Nussbaum, R.A., Lynch, J.D., Green, D. and Wheeler, W. The amphibian
tree of life. Bulletin of the American Museum of Natural History 297:1-370.
Lynch, J.D. 1976. The species groups of South American frogs of the genus Eleutherodactylus (Lep-
todactylidae). Occasional Papers Museum of Natural History University of Kansas 61:1-24.
Lynch, J.D. 2001. Four osteological synapomorphies within Eleutherodactylus (Amphibia:
Leptodactylidae) and their bearing on subgeneric classification. Revista Academia de
Ciencias Exactas, Físicas y Naturales 25:127-136.
Lynch, J. D. and Duellman, W.E. 1997. Frogs of the Genus Eleutherodactylus (Leptodac-
tylidae) in Western Ecuador: Systematics, Ecology, and Biogeography. University of
Kansas Natural History Museum Special Publication 23:1-236.
while only 43 belong to the remaining groups (4 phyllomedusines, 12 pelo-
dryadines, 2 Dendropsophini, 13 Cophomantini, and 7 Lophihylini).
Among those hylids classed in the Critically Endangered or Endangered
categories, the overwhelming threat is habitat loss (113 species). However,
whether the overwhelming majority of Hylini present on the list of Critically
Endangered and Endangered hylids is partially the consequence of differences
in the perceived threat posed by habitat destruction by different specialists,
or a genuine reflection of real destruction of the most diverse areas in Central
America and the Caribbean, deserves further study. One important problem
facing the conservation of hylids in many countries is our ignorance about the
geographic distribution of a large number of species, a problem that compli-
cates any assessment of the status of their populations. A further problem
stems from our plain ignorance about the taxonomy of several groups; there
are likely many dozens of species of hylids that are unknown to scientists
sitting on the shelves of biological collections, not to mention many species
still awaiting discovery in poorly surveyed regions.
In conclusion, Hylidae is a large and diverse family of amphibians, but, like
most other amphibian families, its members are facing population declines in
the face of a litany of threats (Blaustein et al. 1994; Beebee and Griffiths 2005).
Although actions to mitigate against threats such as habitat loss are undoubt-
edly necessary, the resolution of this environmental crises depends firstly on
an attitude change of the humans in relations to their environment.
Julián Faivovich and Célio F.B. Haddad
References
Beebee, T.J.C. and Griffiths, R.A. 2005. The amphibian decline crisis: A watershed for
conservation biology? Biological Conservation 125:271-285.
Blaustein, A.R., Wake, D.B. and Sousa W.P. 1994. Amphibians declines: judging stabil-
ity, persistence and susceptibility of populations to local and global extinctions.
Conservation Biology 8:60-71.
Darst, C.R. and Cannatella, D.C. 2004. Novel relationships among hyloid frogs inferred
from 12S and 16S mitochondrial DNA sequences. Molecular Phylogenetics and
Evolution 31:462-475.
Faivovich, J., Haddad, C.F.B., Garcia, P.C.A., Frost, D.R., Campbell, J.A. and Wheeler, W.C.
2005. Systematic review of the frog family Hylidae, with special reference to Hylinae:
phylogenetic analysis and taxonomic revision. Bulletin of the American Museum of
Natural History 294:1-240.
Frost, D.R., Grant, T., Faivovich, J., Bain, R., Haas, A., Haddad, C.F.B., de Sá, R.O., Channing,
A., Wilkinson, M., Donnellan, S.C., Raxworthy, C.J., Campbell, J.A., Blotto, B.L., Moler,
P., Drewes, R.C., Nussbaum, R.A., Lynch, J.D., Green, D. and Wheeler, W.C. 2006. The
amphibian tree of life. Bulletin of the American Museum of Natural History 297:1-370.
Haas, A. 2003. Phylogeny of frogs as inferred from primarily larval characters (Amphibia:
Anura). Cladistics 19:23-89.
Wiens, J.J., Fetzner, J.W., Jr., Parkinson, C.L. and Reeder, T.W. 2005. Hylid frog phylogeny
and sampling strategies for speciose clades. Systematic Biology 54:719-748+48
(Supp. Data).
22 Threatened Amphibians of the World

ESSAY 1.6. THE AMPHIBIAN TREE OF LIFE

Recent advances in the field of computational biology and high-throughput The taxonomic changes within frogs are more drastic, although the bulk of Ranoides (1,975 species – 38% of the frogs).
DNA sequencing have opened up possibilities for exploring the relatedness the Frost et al. (2006) results have been suggested previously and many can be The Hyloides is the dominant group of frogs in the Americas and Australia.
between large numbers of species in a much more rigorous way than was considered conventional, the taxonomy having just lagged behind the general Of the 18 families, 13 are endemic to the Americas, and two (Limnodynastidae
previously possible (Wheeler 1996; Wheeler et al. 1996-2003). As a result of understanding of frog evolution. The vast majority of the species continue and Myobatrachidae) are centred on Australia. Of the remaining families, the
these advances, Frost et al. (2006), presented the first ever comprehensive to belong to the Neobatrachia (see Figure). However, several groups branch Hylidae is overwhelmingly concentrated in the Americas and Australia (but
amphibian tree, and this was subsequently modified by Grant et al. (2006) (see off from the stem of the tree before we reach the Neobatrachia. The first with a few species in Eurasia), the Bufonidae is globally widespread except
Figure 1). The analysis by Frost et al. (2006) addressed the relationships of group to branch off comprises a single family, the Leiopelmatidae (the New in Australasia, and the Sooglossidae is restricted to the Seychelles Islands
522 species of amphibians, from all nominal families as well as a informative Zealand frogs and the tailed frogs of western North America), comprising and the Western Ghats of southern India.
selection of genera. Details of this very large tree are undoubtedly going to just six species. This is the sister group of all the other frogs (5,205 species). The Ranoides is the dominant group in Africa and Eurasia, to which 13 of
change as data on more species become available, but the broad picture is Traditionally, this family would be considered “primitive”, but there is no the 16 families are endemic. Of the remaining families, the Microhylidae is
now probably reasonably well delimited. particular reason to consider these frogs as being more or less “primitive” widespread through both the New World and Old World tropics, the Ranidae
The tree shows that caecilians are not only the most distinctive group of than any others. However, they do comprise the most phylogenetically isolated is globally widespread except in the southern parts of South America, Africa
amphibians; they are also the sister group of all other living amphibians (i.e., group of frogs in the world. and Australia, and the Ceratobatrachidae ranges from East Asia to the islands
Batrachia, the salamanders and the frogs). In other words, the 172 species of The next major branch among the bulk of the frogs puts two families, of the South Pacific.
caecilians form one ancient major branch in the tree, and the remaining 5,746 the Pipidae (30 highly aquatic species in tropical Africa and South America) An understanding of the relationships between these families is important
species of salamanders and frogs form the other major branch. Among the and Rhinophrynidae (one species from Mesoamerica) as the sister group to for conservation. In deciding conservation priorities, consideration needs
salamanders and frogs, the next major division is, not surprisingly, between all remaining species. The Alytidae (midwife toads and painted frogs – 12 to be given to the evolutionary distinctness of species, as well as to the
these two groups; they are sister groups of each other. species in Europe and the Mediterranean region) and the Bombinatoridae severity of threat to species. For example, using such considerations we
Within the salamanders, two families (Cryptobranchidae and Hynobiidae) (fire-bellied toads – ten species from western and eastern Eurasia and would devote much more conservation attention to the caecilians than
arguably form the sister group of all the other salamanders. According to Southeast Asia) are the next major branch to diverge from the tree, followed they currently receive. And within the frogs, we should focus much more on
Frost et al. (2006), these two families (collectively comprising 49 species) by the spadefoots (Megophryidae from East and Southeast Asia, Pelobatidae the New Zealand frogs and tailed frogs as the most evolutionarily distinct
are centred on East Asia, with one species of Cryptobranchidae occurring in and Pelodytidae from western Eurasia, and Scaphiopodiae from North and group of species.
North America and a single species of Hynobiidae just reaching Europe. The Central America), which form a distinct clade of 142 species. As suggested
remaining clade of salamanders (comprising seven families and 486 species) by Frost et al. (2006) and other authors (e.g., Savage 1973; Roelants and Simon N. Stuart
is, with the exception of the Salamandridae, overwhelmingly North American. Bossuyt 2005) spadefoots are the sister group of the Neobatrachia, which
The position of Sirenidae, placed as the sister taxon of Proteidae by Frost comprises the remaining 5,010 species. Acknowledgements
et al. (2006), is controversial as this family was traditionally considered to The first split within the Neobatrachia is between the Heleophrynidae We thank Darrel Frost and Taran Grant for their very helpful comments on
form the sister taxon of all other salamanders. We can only assume that (the six species of South African ghost frog) and the remaining species, an earlier draft of this essay.
this result will be either further confirmed or rejected as more data come to the Phthanobatrachia. The Phthanobatrachia consist of two familiar major
bear on the question. groupings, the Hyloides (3,157 species – 60% of all frog species) and the
References
Frost, D.R., Grant, T., Faivovich, J.N., Bain, R.H., Haas, A., Haddad, C.F.B., de Sá, R.O.,
Channing, A., Wilkinson, M., Donnellan, S.C., Raxworthy, C.J., Campbell, J.A., Blotto,
B.L., Moler, P., Drewes, R.C., Nussbaum, R.A., Lynch, J.D., Green, D.M. and Wheeler,
Caeciliidae Caecilians Salamanders Frogs and Toads W.C. 2006. The amphibian tree of life. Bulletin of the American Museum of Natural
Ichthyophiidae History 297:1-370.
Rhinadermatidae Grant, T., Frost, D.R., Caldwell, J.P., Gagliardo, R., Haddad, C.F.B., Kok, P.J.R., Means, D.B.,
Cryptobranchidae Noonan, D.P., Schargel, W.E. and Wheeler, W.C. 2006. Phylogenetic systematics of
Hynobiidae dart-poison frogs and their relatives (Amphibia: Athesphatanura: Dendrobatidae).
Rhyacotritonidae Bulletin of the American Museum of Natural History 299:1-262.
Amphiumidae Roelants, K. and Bossuyt, F. 2005. Archaeobatrachian paraphyly and Pangaean diversifica-
Plethodontidae tion of crown-group frogs. Systematic Biology 54:111-126.
Proteidae Savage, J.M. 1973. The geographic distribution of frogs: patterns and predictions. In J. L.
Sirenidae Vial (editor), Evolutionary Biology of the Anurans: Contemporary Research on Major
Ambystomatidae Problems: 351-445. Columbia, Missouri: University of Missouri Press.
Salamandridae Wheeler, W.C. 1996. Optimization alignment: the end of multiple sequence alignment in
Leiopelmatidae phylogenetics? Cladistics 12:1-9.
Pipidae Wheeler, W.C., Gladstein, D.S. and De Laet, J. 1996-2003. POY: Phylogeny reconstruc-
Rhinophrynidae tion via optimization of DNA data. New York: Computer software distributed by
Alytidae the authors and from the American Museum of Natural History: ftp://ftp.amnh.
Bombinatoridae org/pub/molecular/poy.
Megophryidae
Pelobatidae
Pelodytidae
Scaphiopodidae
Heleophrynidae
Sooglossidae
Batrachophrynidae
Limnodynastidae
Myobatrachidae
Hemiphractidae
Brachycephalidae
Neobatrachia Cryptobatrachidae
Amphignathodontidae
Hylidae
Hyloides
Centrolenidae
Ceratophrynidae
Leptodactylidae
Phthanobatrachia Cycloramphidaeeps
Leiuperidae
Bufonidae
Hylodidae
Aromobatidae
Dendrobatidae
Microhylidae
Arthroleptidae
Hyperoliidae
Brevicipitidae
Hemisotidae
Ptychadenidae
Ceratobatrachidae
Micrixalidae
Ranoides
Phrynobatrachidae
Petropedetidae
Pyxicephalidae
Dicroglossidae
Nyctibatrachidae
Ranidae
Mantellidae
Rhacophoridae Figure 1. The amphibian tree of life, showing the relationships between
amphibian Orders and Families, according to Frost et al. (2006), as modified
by Grant et al. (2006).
 CHAPTER 2. WHY SAVE AMPHIBIANS?
Neil Cox, Richard J. Berridge, Don Church, Peter Paul van Dijk, Mirza Kusrini,
Michal Lau, Thomasina Oldfield, Louise Rollins-Smith and Feng Xie
While scientists discuss the urgency of preventing further amphibian declines, it is clear
that many people are uncertain about why we should care about the loss of these species.
When the ongoing extinction crisis is discussed, the question often asked is “why save
amphibians?” This chapter provides a brief discussion of some of the reasons for saving
amphibians.
AMPHIBIANS AS FOOD
To many people, it may be surprising to hear that amphibians are a valued food source
throughout much of the world. The use of amphibians for food can range widely from the
consumption of animals for subsistence means, to their inclusion as essential ingredients
in gourmet dishes. The results of the GAA indicate that ~220 of the world’s amphibian
species are used for food.
The muscular back legs of the larger species of frogs, particularly members of the
families Ranidae and Leptodactylidae, are eaten in many cultures; in some regions, other
body parts of both frogs and salamanders are also consumed. Liner (2005) has helpfully
documented 193 recipes for frogs and toads, and 26 for salamanders (mostly recipes from
North America), and freely admits that this total does not include many of the expected
high number of recipes from Asia and Europe. In some instances, species that may seem
particularly unappetising are eaten; for example, the Campa Indians of central Peru are
reported to roast (to remove noxious skin alkaloids) and eat the poison frog Epipedobates
petersi (Rodríguez and Myers 1993).
Subsistence harvesting of amphibians for food is often quite small-scale or opportunistic,
and may have little overall impact on amphibian populations (with the exception of some
restricted-range species). The results of the GAA suggest that at least 212 amphibians are
used for subsistence food; however, the diversity of species consumed is probably under
recorded, and further studies will almost certainly reveal additional species not previously
identified as being used.
Commercial harvesting of wild amphibians for food operates at scales ranging from
the local collection and transportation of animals to regional or provincial markets, to the
extensive international trade in frog legs. Unfortunately, commercial collection of wild
amphibians has often been unsustainable, and in many cases appears to have resulted
in significant reductions in amphibian populations. Declines of formerly common species
have been reported from the United States (Jennings and Hayes 1985), Europe (Szilárd and
Csengele 2001; Ljubisavljevic et al. 2003; Török 2003), South Asia (Fugler 1983; Abdulali
1985; Niekisch 1986), China (Ye et al. 1993), Viet Nam (Nguyen Quang Truong 2000), and
several Andean countries (Lehr 2000, 2006).
From a global conservation perspective, amphibians with localized distributions, or those
that are in high demand, such as the Critically Endangered Chinese Giant Salamander Andrias
davidianus, have undergone even more alarming declines. Many restricted-range species that
are collected for food are now threatened with extinction (such as the frog Batrachophrynus
brachydactylus EN, confined to the region of Lake Junin in Peru, and the Mountain Chicken
Frog Leptodactylus fallax of the islands of Montserrat and Dominica [Rollins-Smith et al.
2005a]), and while some species such as the Chinese Giant Salamander are protected by
both national and international legislation (nationally protected in China since 1988; listed
on CITES Appendix I in 1997), unsustainable harvesting continues to threaten dwindling
populations (and see Essay 4.7).
The international trade in frog legs still largely depends on the collection of wild animals,
with the majority of animals in trade (all species) coming from wild sources (possibly up
to 85% in 2002; see FIRI 2005). While frog legs are sold globally, the largest market by
far is Western Europe, particularly France, Belgium, the Netherlands [almost exclusively
wholesale for re-export], Luxembourg, and Switzerland (Niekisch 1986; Kusrini and Alford
2006). The principal source of frog legs exported to these countries were initially India and
Bangladesh; however, concerns about the ecological damage resulting from the heavy harvest
of amphibians, in particular of the species Hoplobatrachus tigerinus (LC) and Euphlyctis
hexadactylus (LC), led to export bans in the mid-1980s, and the listing of both species on
CITES Appendix II in 1985, to establish controls on international trade aimed at keeping
it within sustainable levels. Exports of frog legs from Indonesia (mostly from Java) have
continued to increase since the first exports in 1969, and this country is now considered to
be the world’s primary exporter. In 2002, around 3,800t of frog legs, largely of the Mangrove
Frog Fejervarya cancrivora (LC), were exported from Indonesia, with 83% of these exports
going to Europe (Kusrini and Alford 2006). The impact of this heavy collection on Indonesia’s
frog populations is unclear, especially as the large domestic market still needs to be fully
investigated (see Essay 2.1). The results of the GAA given here indicate that the Fanged
River Frog Limnonectes macrodon of Indonesia now meets the criteria for the category of
Vulnerable, in part as a result of the frog legs trade.
In order to meet both international and domestic market demands for frogs, commercial
frog aquaculture has developed in some parts of the world. While frogs have generally
been considered difficult to farm, as they often require moving food and are relatively
slow to reach marketable size, suitable pelleted food has recently been developed for
feeding to both adults and larvae. Fishery statistics available from the FAO indicate that
farms in China alone annually produce substantial numbers of frogs; in 2004 these farms
are reported to have provided 73,837t of frogs (all species) (FAO-FIDI, c2004). In some
instances, it is possible to find example statistics for the cultivation of a single species of
amphibian. Approximately, 1,382t of American Bullfrogs Rana catesbeiana were report-
edly produced by farms in 2004 with most originating from Taiwan (Province of China)
(FAO-FIDI, c2004; FIRI 2005). Lau et al. (1999) report that more than six million Chinese
Edible Frogs Hoplobatrachus rugulosus were imported to Hong Kong from Thailand in
19991. Unfortunately, frog aquaculture has in many instances been detrimental to wild
stocks, with a key problem being the escape of large, often non-native amphibians (such
as the American Bullfrog) and their diseases, into surrounding environments (see Lanza
and Ferri 1997; Rueda-Almonacid 2000; Kiesecker et al. 2001; Mazzoni et al. 2003; Goris
and Maeda 2004) (and see Essay 4.8).
In much of the world, amphibians, most
especially frogs such as this mix of Fejervarya
cancrivora (Least Concern) and Limnonectes
macrodon (Vulnerable), provide an important
but often overlooked source of animal protein.
© Anisa Fitri
MEDICINAL USE OF AMPHIBIANS
While amphibians have long been recognized for their value in traditional medicines2, they
are increasingly also being acknowledged as an important potential source of chemical
substances for use in modern medicine. According to the GAA, at least 73 amphibians are
considered to have some kind of medicinal value.
In many parts of the world amphibians are still collected for use in traditional medicines by
tribal and local peoples, often to meet primary health needs. Aside from this more localized
use, there are some areas, notably East Asia, where commercial markets exist for the use of
amphibians in traditional medicinal practices. More than 30 species of amphibian have been
recorded in Traditional Chinese Medicine alone (Ye et al. 1993). Most of the animals used in
the various traditional medicines tend to come from wild sources, and although much of the
harvesting is of common or widespread species (and may often have little overall effect on
the conservation status of populations), there is concern that a number of restricted-range
species, such as the Critically Endangered Achoque Ambystoma dumerilii, a salamander from
Lake Patzcuaro in Mexico, are now significantly threatened by over-harvesting.
The potential value of amphibians to modern medicine is coming under increasing
scientific study, with the diverse skin secretions produced by amphibians being of particular
interest. The skin of an amphibian must protect the animal against many environmental
pressures, including both disease and predation. Chemical secretions identified so far fall
into the four categories of biogenic amines, bufodienolides (bufogenins), alkaloids and
steroids, and peptides and proteins (Clarke 1997; Pukala et al. 2006). Of these substances,
perhaps the most familiar are the alkaloids produced by the Neotropical poison frogs
(Dendrobatidae), which rank among the most powerful animal poisons known. Hundreds
of alkaloids have now been identified in extracts from amphibian skins, with many of
these considered to be potentially very useful chemicals (Daly 1995; Philippe and Angenot
2005). For example, the alkaloid epibatidine, isolated from the skin of the Ecuadorian
species Epipedobates tricolor (EN), is a potent non-addictive analgesic considered to
be around 100 to 200 times more effective than morphine (Badio et al. 1994). While a
number of ongoing studies have demonstrated that many of the alkaloids in amphibian
skin are not produced directly by the animals themselves, but are sequestered from the
arthropods that make up much of their diet (Daly 1995; Smith and Jones 2004; Saporito et
al. 2004), emerging evidence suggests that some species of Australian frogs in the genus
Pseudophryne are capable of directly biosynthesizing skin alkaloids (pseudophrynamines)
(Smith et al. 2002).
A number of important advances have recently been made in the study of the peptides
synthesized and typically stored in granular glands of amphibians. Several hundred peptides
have now been recorded from amphibians, and many of these are known to have antimicrobial
properties that act as an important natural defence against skin pathogens (Rollins-Smith
et al. 2005b). Antimicrobial peptides from amphibian skin have potentially very significant
applications for both human health and conservation. Exciting work by VanCompernolle
et al. (2005) has identified three amphibian antimicrobial peptides that inhibit Human Im-
munodeficiency Virus (HIV) infection of human immune system T cells. Importantly, these
peptides have also been found to be effective at preventing transmission of HIV from
dendritic cells to T cells (see Essay 2.2).
One important finding, particularly with regards to global conservation efforts, is that of
Rollins-Smith et al. (2002, 2003) who determined that a number of amphibian skin peptides
can inhibit the growth of the chytrid fungal pathogen Batrachochytridium dendrobatidis,
which is implicated as a major cause of ongoing global amphibian declines. Indeed, it now
appears that many amphibian species produce antimicrobial peptides that protect against
infection with the chytrid pathogen (Rollins-Smith et al. 2005b). However, as many amphib-
ians are suffering chytrid-associated population declines, it seems entirely plausible that in
many instances these natural defences have now been compromised by other environmental
factors (Rollins-Smith et al. 2005b).
THE INTERNATIONAL PET TRADE
Since the mid 20th century, a wide range of amphibian species has been represented in
the international pet trade. The GAA recorded that at least 278 species of amphibian were
in the pet trade alone, and it is likely that many additional species enter the trade on a
regular basis. The main centres of export for the amphibian pet trade are the wet tropics,
with species largely coming from the rainforests of Africa (including Madagascar), Central
America, South America and Southeast Asia. Comprehensive data covering all amphibians
traded are often hard to come by, especially as there is no global database or monitoring
system for the trade in non-CITES species (Schlaepfer et al. 2005). Very generally, the main
international markets for pet amphibians are considered to be within North America and
24 Threatened Amphibians of the World

Europe, and developed countries such as Japan. Schlaepfer et al. (2005) provide a valuable of 7-10 individuals per square metre in Red-backed Salamander Plethodon cinereus popula-
study of imports of wild reptiles and amphibians into the United States from 1998-2002; tions within the Shenandoah National Park, eastern USA; and estimates by Stewart and
during this period at least five million amphibians, reported as being from wild-caught Rand (1991) of as many as 20,570 adult Puerto Rico Coqui Frogs Eleutherodactylus coqui
sources, entered the US primarily for the purposes of the pet trade3. per single hectare of forest.
The export of mantella frogs (Mantellidae) from Madagascar has been particularly It is probable that amphibians can occur in such abundance because both adults and larvae
closely examined, and provides a helpful case study of the value of some amphibians in are often the immediate primary vertebrate predators of invertebrates (especially insects), in
the international pet trade. Data gathered by Rabemananjara et al. (in press) reveal that both freshwater and moist terrestrial environments (Stebbins and Cohen 1995). Additionally
over the 10-year period from 1994-2003 the top ten importers of mantella frogs were: the many larvae are primary consumers of plant matter, such as algae, and detritus and can
United States (71% of all mantellas exported from Madagascar), the Netherlands (5.5%), be frequently found at high densities in freshwater habitats. In agricultural landscapes,
Germany (5.2%), Canada (4.6%), Belgium (4.5%), France (2.2%), Japan (1.9%), Switzerland such as rice paddies, amphibians have been suggested as important agents in the control
(1.1%), Great Britain (1.3%), and Spain (1.0%). This trade comprised over 230,000 animals, of invertebrate pests (Abdulali 1985; Oza 1994). There is some concern that the removal of
and over a period of three years (2001-2003) was worth almost US$250,000 in foreign amphibians from these systems could lead to increases in these pests, and subsequently
currency revenue for Madagascar. Although these figures relate just to mantellas, and to an increase in the need for the application of insecticides and other agrochemicals. It
are therefore subject to certain influences4, the broad trends in these data, particularly in should be noted here that a number of amphibians, such as the Cane Toad Bufo marinus (LC),
terms of which countries have the highest demand for pet amphibians, are likely to apply have been widely introduced as biological controls of insect pests. In the majority of cases,
A number of brightly coloured amphibians across many other genera. naturalized populations of non-native amphibians are considered to have had damaging
are now regularly bred in captivity and kept The most common amphibians in the international trade are the frogs and toads, and ecological and/or socioeconomic consequences. Lever (2003) provides extensive details
as pets. Pictured here is the Dyeing Poison among these several groups of species predominate. The brightly coloured poison frogs on the effects of known amphibian introductions worldwide.
Frog Dendrobates tinctorius (Least Concern), (Dendrobatidae) and the equally attractive mantellas from Madagascar (Table 1) have Amphibians are one of the key links in many ecosystem food webs. As “cold-blooded”
a native species of the lowland forests of always been in high demand. Other popular anurans include similarly spectacular species, ectotherms, ingested energy from food is efficiently converted to high-quality new tissue,
the Guianas and adjacent Brazil. © Henk such as treefrogs (most often in the family Hylidae), fire-bellied toads (Bombinatoridae), with this biomass then becoming readily available for transfer to higher trophic levels
Wallays horned frogs (Ceratophrys, in the Leptodactylidae) and several aquatic frogs (often species (Pough 1983; Stebbins and Cohen 1995; Whiles et al. 2006). In both freshwater and ter-
in the genera Hymenochirus or Xenopus). Several salamander species are also traded, restrial ecosystems, amphibians as adults and larvae represent a major food resource for
with aesthetic appeal seeming to be the main criterion for inclusion. Popular taxa include many predators, including large invertebrates, fishes, other amphibians, reptiles (snakes in
the Asian fire-bellied newts of the genus Cynops, members of the North American family particular), birds, and mammals. Through the process of metamorphosis, they also provide
Plethodontidae, and other brightly coloured salamanders such as the European Fire Sala- a link in the transfer of nutrients from aquatic systems to terrestrial ones (including forest,
mander Salamandra salamandra (LC), and several newts of the Asian genus Tylototriton. see Gibbons et al. 2006).
Salamanders of the family Ambystomatidae are also in demand as pets, often because of From the few studies that have been undertaken into the potential consequences of
their large size, general ease of maintenance and, perhaps most particularly in the case of removing amphibians from an ecosystem, it appears that important changes can take place
the Axolotl Ambystoma mexicanum (CR), their fascinating ecology and breeding biology. in both ecosystem processes and services. Among the structure and functions affected by the
Caecilians tend to be comparatively rare in the international pet trade, with the easier to removal of amphibians are algal communities, invertebrate populations, predator dynamics,
observe fully aquatic species (such as Typhlonectes natans LC), being more readily available organic matter dynamics, patterns of primary production, leaf litter decomposition, and
than the predominantly subterranean forms. nutrient cycling (Whiles et al. 2006).
While the vast bulk of the international pet trade in amphibians comprises wild-caught The future impact of ongoing amphibian declines on the world’s ecosystems is still
specimens, in several instances captive-bred individuals feature more prominently. For uncertain with much further study needed. However, given the important role of amphibians
example, some captive-bred colour morphs of particular poison frogs (such as the pink and in many food webs it seems likely that significant measurable changes in both freshwater
yellow morph of Epipedobates tricolor) now appear to be commoner in the trade than their and terrestrial environments will eventually take place.
wild progenitors. A comprehensive study of the trade in dendrobatid frogs by Gorzula (1996)
revealed that an increasing proportion of the animals recorded in trade from 1987 to 1993
were reported as having been captive-bred; of nearly 16,000 poison frogs in international MONITORING ENVIRONMENTAL CHANGE
trade during this period, live captive-reared frogs represented 13.1% of the total traded5.
It seems possible that the implementation of various regulatory controls, such as CITES Amphibians have a number of physiological, ecological and life history characteristics that
(which has also had a marked effect on the availability of particular species), has provided make them susceptible to environmental changes. They are often considered to be valuable
a stimulus for the development of captive-breeding methods, which in turn have reduced bioindicators of environmental change in both aquatic and terrestrial environments, and
the need for wild-caught individuals. Several amphibians that are particularly popular as declines in amphibian populations or species diversity have frequently been directly linked
pets are also captive-bred in commercial quantities, such as the Ceratophrys horned frogs, to habitat degradation (Harfenist et al. 1989; Vitt et al. 1990; Blaustein and Wake 1995;
some treefrogs (such as Red-eyed Treefrog Agalychnis callidryas LC), and a number of Stebbins and Cohen 1995; Dodd 1997; Southerland et al. 2004).
poison frog species. And, indeed, the Axolotl Ambystoma mexicanum is more common in It seems that relatively few attempts have been made to explicitly estimate how good
captivity now than it is in the wild. amphibians are as indicators compared with other taxonomic groups. Welsh and Droege
There is potential scope for developing sustainable in situ harvesting, or farming, of (2001) argued that the low variation in counts of plethodontid salamanders among studies
some of the more desirable species, where their ecology is sufficiently well understood relative to Lepidoptera (butterflies and moths), passerine birds, small mammals, and other
for sustainable harvest quotas and practices to be established (see also Chapter 11). This amphibians suggests that these salamanders provide a statistical advantage over other
could have profound conservation benefits, as although most amphibian species kept as species for monitoring forest health6. In contradiction to Welsh and Droege (2001), Bailey
pets have been now bred in captivity on occasion, the great bulk of all animals in trade are et al. (2003, 2004a,b,c) explicitly estimated detection probability and temporary movement
still collected from wild populations. And even where the capture of wild specimens does in plethodontid salamander populations using marked individuals and found that counts of
not yet constitute a threat to a species, the development of in situ harvesting practices animals varied both spatially and temporally, thereby potentially confounding any studies
could provide the required incentive to motivate landowners to protect and even enhance that use simple counts by themselves as a surrogate to population size. In other words, the
the species’ habitat. A local, sustainable, production programme for species of relatively findings of Bailey et al. (2004a,b,c) suggest that counts that are not corrected by incorporating
high worth (e.g., mantella frogs or poison frogs) could also have the additional benefit of detection and temporary movement probabilities across temporal and spatial components
Table 1. Export of Mantella frogs between playing a role in poverty reduction strategies for some of the poorest parts of the world of a study area are likely to provide inaccurate and potentially biased indications of how
1994 and 2003 from Madagascar as reported (and see Chapter 11). populations and, thereby, the environment are faring. As further complication to the potential
in the CITES annual report data obtained from of plethodontids as indicator species, Bailey et al. (2004a) found that accounting for detection
the UNEP-WCMC trade database (data col- probability can only do a reasonable job of correcting counts if temporary movement from
lated on 2 May 2005). The species M. baroni, ECOSYSTEM ROLE OF AMPHIBIANS a population is a random process.
M. cowani, M. haraldmeieri, M. madagas- Although there remains some uncertainty concerning the overall value of amphibians
cariensis and M. pulchra have gone through Although generally unseen, amphibians can often be very abundant in some ecosystems. as bioindicators, especially with regards to monitoring more difficult to observe species,
stages of profound taxonomic rearrangements In both temperate and tropical environments, amphibians may exceed all other terrestrial the following text outlines some of the commonly recognized characteristics of amphibians
between 1988-1999, and these have been vertebrate classes (reptiles, mammals, and birds) in both the numbers of individuals and in that are believed to be of use in recording environmental change.
largely confounded also in the pet trade. The the proportion of total available biomass (Stebbins and Cohen 1995). Examples of particularly Physiologically, the skin and egg capsules of amphibians are often highly permeable
trade figures of these species therefore need high abundances of amphibians include findings by Gergits and Jaeger (1990) of densities to gases and liquids and may readily absorb pollutants such as toxic chemicals, endocrine
to be viewed with some caution (reproduced mimics, and other substances from the surrounding environment (Stebbins and Cohen 1995;
from Rabemananjara et al. in press). Dodd 1997). Contamination with many of these substances can often result in rapid declines
in amphibian populations, and, perhaps more pervasively, can frequently lead to abnormal
metamorphosis from the larval stage to the adult form (malformed animals may often be
Taxon 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003 TOTAL % observed in contaminated populations).
Mantella spp. 0 0 0 230 620 200 6,760 9,853 1,420 1,291 20,374 8.71 Amphibian larvae feed on a wide variety of plant and animal matter, in all parts of the
Mantella 100 11,965 16,693 17,406 3,194 8,850 11,445 10,335 4,780 2,681 116,196 49.68 water column from the benthic detritus to the surface film. The larvae are often susceptible
aurantiaca to ingesting persistent chemicals attached to particulate matter. The chemicals can readily
Mantella baroni 0 0 0 0 0 0 0 0 10 650 660 0.28 accumulate in fat deposits, and sequestered contaminants may also significantly affect
Mantella bernhardi 0 0 0 0 0 30 490 1,005 650 60 2,235 0.96 normal development or metamorphosis when the fat deposits are drawn upon. As adult
Mantella betsileo 0 0 0 1,000 435 175 150 4,040 1,215 1,465 8,480 3.63 amphibians largely feed on invertebrates, they are also vulnerable to bioaccumulation of
Mantella cowani 0 0 0 0 52 150 425 975 1,520 500 3,622 1.55 persistent chemical pollutants, such as polychlorinated biphenyls (PCBs) or dichloro-diphe-
Mantella crocea 0 0 0 0 395 250 1,157 1,750 630 100 4,282 1.83 nyl-trichloroethane (better known as DDT), in the food web (Hall et al. 1985; Stebbins and
Mantella expectata 0 0 0 100 624 105 1,260 1,790 2,585 1,100 7,564 3.23 Cohen 1995; Dodd 1997; Fagotti et al. 2005; DeGarady and Halbrook 2006).
Mantella 0 0 0 0 0 0 240 310 380 350 1,280 0.55 Amphibian populations can decline in response to changes in the acidity of aquatic
haraldmeieri habitats, through either a decrease or increase in pH levels. Sources of changes in pH include
Mantella laevigata 0 0 0 100 435 415 2,537 2,795 1,170 1,581 9,033 3.86 acidic precipitation, or point pollution, such as runoff from abandoned mines (Dodd 1997;
Mantella 0 0 0 125 2,182 1,535 6,195 8,805 5,945 4,848 29,635 12.67 Middlekoop et al. 1999). In general, eggs and developing larvae are the most sensitive life
madagascariensis stages to low pH (< 4.5) (Dodd 1997).
Mantella 0 0 0 0 0 0 0 0 1,270 1,780 3,050 1.30 Adult temperate and montane amphibians are often heliothermic, and may expose
milotympanum themselves to sunlight in order to increase their body temperatures. This characteristic may
Mantella nigricans 0 0 0 100 200 0 155 490 80 0 1,025 0.44 make animals more susceptible to increases in ultraviolet (UV) radiation, often associated
Mantella pulchra 0 0 0 0 784 905 3,277 4,430 2,990 2,560 14,946 6.39 with a thinning of the ozone layer (Stebbins and Cohen 1995). In addition, as amphibians are
Mantella viridis 0 0 0 125 690 385 1,951 3,825 2,495 2,040 11,511 4.92 moisture dependent ectotherms, they are generally sensitive to any changes in temperature
TOTAL 100 11,965 16,693 19,186 9,611 13,000 36,042 50,403 27,140 21,006 233,893 100 or precipitation levels. Unfavourable conditions may prevent reproduction, possibly for many
years (Stebbins 1995).

Increases in sedimentation or turbidity of streams and other waterbodies may be revealed
through changes in both the species richness, and population sizes, of amphibians present
(Welsh and Ollivier 1998). Many species, possibly most especially in their larval stage,
are sensitive to the loss of certain microhabitats or food sources resulting from increased
levels of particulates. Other species may be affected by a reduced efficiency of their gills
within turbid waters.
Unlike many vertebrates, such as birds, some amphibian species have a low inclination or
ability to disperse and tend to be largely confined to small areas of moist habitat (although
see Chapter 1) (Smith and Green 2005). These sites, and the amphibian populations present,
are often naturally fragmented and there can be little or no movement between popula-
tions. Consequently, the loss of one of these populations, for instance through a temporary
pollution event, does not necessarily mean that the site will be easily recolonized from any
nearby populations. The loss of amphibian populations in some species can, therefore, be
used to track ecological disturbance with some confidence that recolonization from adjacent
populations will not mask any notable changes to the environment.
It has sometimes been suggested that while some invertebrates have the ecological
characteristics that make amphibians helpful bioindicators, they generally differ in that
many species are short-lived and can have significant population fluctuations within a short
timeframe. However, amphibian populations may also exhibit unstable population dynamics
(e.g. Pechmann et al. 1991). To use them as bioindicators it is necessary to either monitor
their populations over long periods or establish a solid understanding of the relationships
among population vital rates, the environmental factors that influence these rates, and how
the environment is likely to change in the future so that stochastic population projections
can be performed.
While nearly all caecilian and salamander species make little, if any, noticeable vocaliza-
tions, the often loud calls by frog species are well known to most people. Communication
between individual frogs is a helpful means by which the general species composition of
the area, and relative density of the individuals for each species, can be roughly determined.
Auditory monitoring of frog vocalizations can help indicate any changes in either species
composition or density of individuals (which in turn may reflect changes in environmental
quality of the site) (Crouch and Paton 2002; Brandes et al. 2006).
LEARNING FROM AMPHIBIANS
For many people, one of their first experiences in handling amphibians is in dissecting a
frog during school biology classes. Frogs are most frequently used in these classes, as they
cost relatively little and are often readily available. The primary aim of such lessons, of
course, is to observe the animal’s major internal organs (such as the heart and the liver),
and thereby allow students to gain a better understanding of basic vertebrate (including
human) anatomy and physiology. As many biology classes require a course in dissection,
there is understandably a high demand for subject animals. Although the volume of frogs
specifically used in dissection classes is difficult to determine, an early study by Gibbs et
al. (1971) reported that approximately 326.5t of wild-caught animals (Leopard Frogs Rana
pipiens and Bullfrogs Rana catesbeiana) were annually supplied for educational and research
purposes in the United States7. Pandian and Marian (1986) estimated that 18 million frogs
were used in 1980 by the education sector in India; a significant rise on the 2.6 million animals
estimated to have been used in 1956. It is difficult to determine the current volume of animal
use in countries such as the United States, as supply companies are perhaps understandably
concerned about the confidentiality of these data. However, information kindly provided by
Carolina Biological Supply Company (USA) suggests an overall decline in the use of frogs
(both live and preserved) within the United States for dissection in education (D.E. James
pers. comm.). Most of the frogs now used in the classroom appear to originate from cultivated
stock, although small numbers of wild animals are still available8.
Chapter 2. Why Save Amphibians? 25
Amphibians are one of the key links in eco-
system food webs, providing a significant
food source for snakes and other predators.
This juvenile Pink-sided Treefrog Agalych-
nis litodryas (Vulnerable) has become prey
for the snake Leptodeira septentrionalis.
© H. Mauricio Ortega-Andrade
Today, modern technology means that there are now a number of interactive or ‘virtual’
programmes and software available for use in dissection and anatomy classes (Kinzie et al.
1996). Many of these programmes use virtual frogs as the dissection subject, and in some
instances provide an alternative to the use of animals in classes.
Outside the classroom, amphibians have contributed to important medical research and
discoveries. In fact, seven Nobel prizes have been awarded based on research done with
amphibians. While using a dissected frog, the 18th century Italian anatomist Luigi Galvani
found by chance that muscle and nerve cells produce electricity. During the discovery,
Galvani’s assistant accidentally touched an electrically charged scalpel to the sciatic nerve
of the dissected frog, causing the limbs to violently twitch and convulse. This observation
and subsequent experiments by Galvani helped to establish the basis for the study of the
nervous system through the scientific disciplines of neurophysiology and neurology.
Amphibian oocytes (a cell which may produce an egg [ovum] by meiotic division) are
commonly used in ‘patch clamp’ experiments for measuring the electrical properties of
biological cells and tissues. The Nobel Prize-winning team of Bert Sakmann and Erwin
Neher developed this technique, which is used to measure the movement of electrically
charged molecules through individual channels in the membranes of cells (Neher and
Sakmann 1976). To conduct an experiment, a very fine, polished glass pipette is placed in
contact with the membrane of a cell, and then slight suction is used to make a very tight
seal between the pipette and the ‘patch’ of cell membrane underneath the pipette’s tip.
The passage of electrically charged molecules through the cell’s membrane channels, both
into, and out of the cell, is then measured. Xenopus oocytes are commonly used for these
experiments because of their relatively large size (at approximately 1mm across they are 10
times larger than most animal cells), which makes their handling and manipulation easier.
Not only can patch clamping be used to examine the electrical properties of the channels
that are normally found in the membranes of Xenopus oocytes, but treatment with genetic
material from other organisms makes the oocytes ‘express’ other organisms’ membrane
channels too, making Xenopus oocytes an extremely valuable tool for neurological research.
Molecules that are temporary copies of DNA (mRNA - messenger RNA) can be extracted
from the cells or tissues of other organism, and then injected into the Xenopus oocyte, which
will manufacture the foreign proteins, and incorporate that organism’s foreign channels into
its own membranes, as if they were actually its own9.
Amphibian eggs have an especially important role in the field of embryology – the study
of the formation, early growth, and development of living organisms. The large size of am-
phibian eggs and their translucent jelly-like coat provide a helpful window into embryonic
development, with the African-clawed Frog Xenopus laevis (LC) most regularly used as the
model organism for embryology studies (Gurdon and Hopwood 2000). During the 1930s, it
was discovered that female Xenopus could be induced to ovulate when injected with the
urine, and hormones, of a pregnant woman. Additional research demonstrated that other
species of frogs, such as Rana pipiens and Bufo viridis, could also be used as pregnancy
tests (Miller and Wilberger 1949; Rabau and Szeinberg 1959). For some time during the
1940s and 1950s the use of Xenopus, or the Hogben test as it was called, was the only
pregnancy test readily available to hospitals. Once it was realized that large eggs suitable
for microsurgery could be obtained year-round from domesticated Xenopus, embryologists
no longer needed to rely on the seasonal collection of wild amphibian eggs for their experi-
ments. The eggs of Xenopus are now the leading choice for many embryology studies, as
they are ideal for microsurgery, and can be easily manipulated both morphologically and
genetically (Stebbins and Cohen 1995).
It may be of some interest that the world’s first successful cloning experiments relied on
the use of amphibians. In 1952, the two scientists Robert Briggs and Thomas King managed
to clone a Leopard Frog by carefully transferring early stage nuclei (from blastula cells)
into enucleated eggs (Briggs and King 1952). Although some of Brigg’s and Kings’ initial
attempts failed, they continued perfecting their methods and by the end of the project a
number of eggs had successfully developed into normal tadpoles. However, during their
26
Laboratory studies using the African Clawed
Frog Xenopus laevis (Least Concern) have
led to significant medical breakthroughs.
© Louis du Preez
It was once believed that the Common Fire
Salamander Salamandra salamandra (Least
Concern) was invulnerable to fire, mostly
because hibernating animals would quickly
emerge from firewood once it had been lit.
© Jelger Herder
People may often be surprised by how many
simple images of amphibians surround them
in daily life. For example, this baker’s shop
in Denmark sells tasty frog-shaped cakes.
© Thomas Brooks
Threatened Amphibians of the World
experiments, Briggs and King were unable to clone frogs using nuclei from older, differ-
entiated, embryo cells. The breakthrough came in 1962, when the scientist John Gurdon
finally demonstrated that successful cloning could be achieved from differentiated cells.
Gurdon transferred intestinal cell nuclei from Xenopus tadpoles into enucleated eggs and
managed to produce 10 normal tadpoles (Gurdon 1962). Although there was some initial
scepticism among scientists concerning Gurdon’s results, his later experiments confirmed
that a major advance in our understanding of cloning had indeed taken place. The potential
importance of the achievements of these cloning pioneers is difficult to overestimate. Their
experiments with amphibians have formed the scientific basis for later landmark events,
such as the cloning of Dolly the sheep, the first mammal to be successfully cloned from an
adult cell (Wilmut et al. 1997).
Unfortunately, although we are continuing to gain an appreciation for the value of amphib-
ians for medical advances and in research, it seems as though one important lesson may
have been lost forever. The two species of gastric-brooding frogs (Rheobatrachus silus and
R. vitellinus) from the rainforests of Queensland, Australia, had extraordinary reproductive
habits in which the females brooded young within the stomach and disgorged the fully
developed young through the mouth. Research by Tyler et al. (1983), when the species was
already rare, provided evidence that the tadpoles secrete a substance (prostaglandin E2) that
inhibits the secretion of gastric acid by the female. It certainly seems possible that further
studies of this remarkable reproductive mode could have resulted in medical advances in
the treatment of people who suffer from gastric ulcers. Sadly, neither of the Rheobatrachus
species has been seen since the mid-1980s, despite extensive field surveys, and both are
now considered to be extinct (see Essay 6.1).
THE CULTURAL USE OF AMPHIBIANS
Amphibians have long been represented in the cultures of many human societies. As it is
difficult to present a concise picture of the spiritual, material, intellectual and emotional uses
of amphibians, the following examples are intended to provide only a very brief overview
of their many contributions.
Within written records, amphibians are well represented in antiquity by the Ancient
Egyptians, most especially in the form of the goddess Heqet who symbolized birth and rebirth,
life and fertility. Heqet is depicted in hieroglyphs as a frog or as a woman with a frog’s head,
a reflection of the sudden abundance of calling frogs during and following Nile River floods.
An important cultural reference to both frogs and Ancient Egypt is found within the Biblical
book of Exodus and Sura 7 in the Quran, in which Moses leads the Israelites from Egypt.
In Exodus, 10 plagues are inflicted upon Egypt by God to convince Pharaoh to allow the
Israelites to leave. The second of these plagues was a horde of frogs that overran the land.
Pharaoh gave permission for the Israelites to leave if Moses would agree to remove the frogs;
however, once the frogs had died Pharaoh reneged on his word. God subsequently sent eight
more calamities to Egypt before the Pharaoh agreed to let the Israelites leave.
Frogs are not the only amphibians that have been associated with religion, spiritualism
or mysticism. Salamanders were long associated with the popular European myth that these
animals were invulnerable to fire. Indeed, it was often believed that salamanders preferred
the hottest or fiercest of fires. It seems as though this legend is almost certainly derived
from the frequent hibernation of Fire Salamanders Salamandra salamandra in stacks of
firewood. The Axolotl of Mexico provides another helpful example of a culturally important
salamander (see Essay 2.3).
There are several local festivals around the world in which frogs are the leading feature,
and a few examples are given here. During the Maguai (= frog) festival of the Zhuang
people of Guangxi, China, a frog is captured, honoured and then buried in order to ensure
good weather and an excellent harvest for the following year. Since 2003, the city of San
Fernando on Luzon Island in the Philippines has held the Pyestang tugak frog festival; with
the purpose of perpetuating the Kapampangan people’s traditional methods of catching and
cooking frogs. The City of Rayne in Louisiana, USA, declares that it is the “Frog Capitol of the
World”! People attending this annual festival can see frog races and jumping competitions,
as well as frogs dressed up in small uniforms.
In Japan, the frog is traditionally viewed as a symbol of good fortune endowed with magi-
cal powers. The word for frog in Japanese, ‘kaeru’, also means ‘to return’; this may include
a return from travel, the return of a fortune, or the return of youth. Small charms of frogs are
often carried, and are considered to be especially lucky if the frog’s mouth is open.
Amphibians are regularly found in both classical literature and fairy tales. Shakespeare
gives a familiar view of the toad in the comedy ‘As You Like It’ (“Sweet are the uses of
adversity, which, like the toad, ugly and venomous, Wears yet a precious jewel in his
head.”), and amphibians are an important ingredient of a witches brew (“eye of newt and
toe of frog”) in the tragedy ‘Macbeth’. Mark Twain wrote about the unfortunate gambler
Jim Smiley and his trained frog, Daniel Webster, in the short story ‘The Celebrated Jump-
ing Frog of Calaveras County’. The much-loved central characters of the children’s stories,
‘Jeremy Fisher’ by Beatrix Potter, ‘The Frog Prince’ by the Brothers Grimm and ‘The Wind in
the Willows’ by Kenneth Grahame, are all amphibians. Aesop included frogs in two of his
most popular fables ‘The Frog and the Ox’ and ‘The Frogs Who Desired a King’.
Images of amphibians, especially frogs, are often used for advertising or entertainment
in popular culture. International advertising campaigns for beer (Budweiser®) and telephone
ring-tones (the ‘Crazy Frog’) feature amphibians, as do advertisements and packaging for
products as diverse as shoe-polish (Erdal®) and chocolate (‘Freddo Frog’). Within entertain-
ment, amphibians are commonly found in video games, such as the classic arcade game
‘Frogger’, as puppets, such as the frog ‘Kaj’ in the popular Danish children’s TV series “Kaj
og Andrea”, and as characters in various cartoons and manga comics including: Michigan J.
Frog, an animated ‘Looney Tunes’ cartoon character; Sergeant Keroro, an anime character;
Flip the Frog, an animated cartoon character who starred in a series of cartoons from 1930
to 1933; and Baron Silas Greenback, the recurring villain in the British animated television
series, ‘Danger Mouse’. Finally, of course, perhaps the most widely recognized amphibian
in contemporary pop-culture is the puppet Kermit the Frog, famously associated with the
well-known television programmes Sesame Street and The Muppets.
THE BEAUTY OF AMPHIBIANS
In concluding this chapter on the value of amphibians to society, it is important to discuss
their aesthetic worth, and although this component of intrinsic value generally defies
complete quantification in monetary terms, it is important to recognize, and provides
perhaps the most important incentive for valuing amphibians (McCauley 2006a,b). In
particular, because amphibians are abundant and generally harmless, they may be one
of the first wild animals encountered by young children that can lead to an enduring
appreciation of nature.
Carl von Linnaeus, the founding father of modern taxonomy, famously described amphib-
ians as “ugly, disgusting creatures ... whose creator failed to try and make more of them”
(Hofrichter 2000). However, there is, in fact, a diversity of colour, form, appearance and
activity among amphibians to rival the aesthetic delights of anything else that the natural
world cares to dazzle us with. Colours range from one end of the spectrum to the other,
with every possible permutation in between, with reds, oranges, brilliant vermilions, blues,
greens, violets, pinks, black, and even white. Some are paradigms in post-modernism, be-
ing delicately dipped in just one shade, while others combine crazy rainbow combinations
with spots, stripes, hatches and stipples, madly finished off with tiger-striped legs. Yet others
amaze with their uniform bright, almost fluorescent colouration, impossibly tastefully well
offset by a crystal clear, shining bright black. No one who sees these species can fail to be
amazed by their outlandish, excessive, seemingly pointless beauty.
Yet, these colours serve a very important purpose. Bright colours may serve to advertise
fitness to a potential mate, and toxicity (to predators) in the dark confines of the rainforest.
A tasteful mix of browns and greys is just the right combination to render a dainty treefrog
identical, upon resting, to a very ordinary, quite unappetising bird dropping. Other species
produce amazing visual displays by hiding their brightest surprises until the situation de-
mands that they parade themselves in all their glory – the fire-bellied toads (Bombina spp.)
are camouflaged on their upper surface, being a mixture of browns, blacks, and greens, but
upon disturbance by a predator they arch their backs and reveal a stunning fiery belly, aflame
with bright oranges, reds, and yellows, warning any foolish predator to back off, lest they
ingest some of the poison the toads carry. The frog Physalaemus nattereri (LC) even reveals
startling eye-patches on its hind legs when being pursued by a predator; the eye patches
are actually inguinal venomous macroglands, but they contribute to the appearance of a
large face with a pair of impressive large eyes.
The particular characteristics or forms are often reflected in their scientific names.
The Latin name of the Golden Poison Frog Phyllobates terribilis (EN) is most apt when
one considers that it is the most toxic species of frog in the world (Myers and Daly 1983).
Another example is the Lake Titicaca Frog Telmatobius culeus (CR), whose names translates
as ‘aquatic scrotum’ on account of this species’ multiple folds of baggy skin, designed to
enhance oxygen uptake in its cold water habitat.
There is little doubt that the sheer diversity of form and appearance provides an endless
aesthetic treat for the interested observer. But perhaps the greatest appeal of amphibians
is not visual, but acoustic as anyone who has ever been privy to a chirping chorus of frogs
as they begin their courtship ritual in early spring will attest.
“Frogs do for the night what birds do for the day... they give it a voice. And that voice is
a varied and stirring thing that ought to be better known.”
These words written by the zoologist, Archie Carr (Carr 1973) reflect the thinking of many
of us who study amphibians. Sadly, such melodic rhapsodies are under threat; this book sets
out to argue that we cannot stand by while the frogs croak their last.
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28
ESSAY 2.1. FROGS FOR HUMAN CONSUMPTION
Walking down the aisle of a traditional market or supermarket in many cities in
Southeast Asia, you can find frogs sold for human consumption, either live or as
fresh skinless legs. Although they are not consumed as universally as are other
aquatic taxa such as fish and prawns, frogs have traditionally been used as food in
many cultures. People in most regions of the world, including Europe, Asia, Africa,
the Americas, and Australia, consume frogs. Usually, only the legs are eaten. Frogs’
legs are often considered to be delicacies, and are sometimes highly appreciated
as a unique dish, for example “les cuisses de grenouilles” of France.
The number of species exploited for frog legs varies among regions. For
instance, in Europe the species most harvested are the green frog Rana escu-
lenta (LC) complex and other species such as Rana dalmatina (LC) and Rana
ridibunda (LC), whereas in Asia the species harvested are mostly associated
with rice fields, such as Fejervarya cancrivora (LC) and Fejervarya limnocharis
(LC). Asians also harvest some stream-associated species such as Limnonectes
macrodon (VU) and Limnonectes blythii (NT). Most harvested species share
several attributes: they are large, with heavy muscled legs (for example,
Limnonectes macrodon exported from Indonesia have snout-vent lengths up to
15cm), are mostly in the family Ranidae, and most do not possess skin glands
that produce secretions that are highly toxic to humans. The largest extant frog
species, the African Bullfrog Pyxicephalus adspersus (LC), is consumed by many
traditional cultures in southern Africa; in this case, most of the frog carcass is
consumed (Okeyo 2004). On a local scale, which is probably too small to be of
conservation significance, some smaller frogs are also harvested in many areas
and sold for consumption by other animals, such as fish and pet snakes. In the
USA, and probably in other countries, smaller frog species, juvenile ranids, and
salamanders are also collected or raised and used or sold, usually locally, as fish
bait (e.g., Meronek et al. 1997). The extent of this trade appears to be small,
but it is of potential concern for its immediate impact on frog populations, the
potential to disseminate pathogens such as the amphibian chytrid skin fungus,
and for ethical reasons since most amphibians are used as live bait.
Most frogs consumed by humans are caught from the wild. Only one spe-
cies of frog is widely farmed, the American Bullfrog Rana catesbeiana (LC),
which has been introduced to many other countries, such as Indonesia, often
specifically to boost the production of frog legs for export (Iskandar 1998).
The introduction of bullfrogs may cause many problems (and see Essay 4.8).
Several studies in the USA have suggested that feral populations of Rana
catesbeiana outside the natural range of this species have caused declines
of native frog populations by acting as competitors or predators (Hayes and
Jennings 1986). This species also is likely to act as a vector and reservoir of
amphibian diseases, including chytridiomycosis (Mazzoni et al. 2003).
Harvesting for consumption has led to declines in populations of some
species, for example in the California Red-legged Frog (Rana aurora drayto-
nii) in the USA (Jennings and Hayes 1985). The combination of population
declines, ongoing demand, and expanding trade has led to the nomination of
some edible species for protected status. When local species are protected
in an area where frog consumption is traditional, the sources of frog legs
shift to countries that do not restrict frog harvesting. Frozen frog legs have
thus become an international commodity, in which developing countries in
Asia or South America are the producers and developed countries, mostly in
Europe, are the consumers. This has shifted the focus of consumption-related
conservation concern from the consuming countries to exporting countries.
In 1985 two major producing countries, India and Bangladesh, placed their
two edible species (Euphyctis hexadactylus, LC, and Holobatrachus tigerinus,
LC) on CITES Appendix II because of declines in their populations (Abdulali 1985;
6,000,000
5,000,000
4,000,000
Weight (kg)
3,000,000
2,000,000
1,000,000
0
1969 1970 1971 1972 1973 1974 1975 1976 1977 1978 1979 1980 1981 1982 1983 1984 1985 1986 1987 1988 1989 1990 1991 1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002
Year
ESSAY 2.2. ANTIMICROBIAL SKIN PEPTIDES OF AMPHIBIANS
The skin of frogs and toads has been a component of folk medicine of many
cultures since ancient times (Lazarus and Attila 1993). Each amphibian species
seems to have its own arsenal of compounds that are stored in the skin to
ward off predators and/or microorganisms. They include amines, alkaloids,
and peptides (Pukala et al. 2006). These stored materials are essentially a
“packet of poisons” to disarm predators along with a “first aid kit” containing
antibiotics and pain relievers to help with the healing processes.
Granular glands in the skin (also called serous glands or poison glands)
produce an array of bioactive substances, including peptides with antimicrobial
properties. The peptides are synthesized as larger proteins that are cleaved to
smaller active peptides and stored in high concentrations in membrane-enclosed
granules within each gland. When the animal is alarmed or injured, the entire
contents of the gland are pushed out onto the skin, and the peptides play their pro-
tective roles. Then the gland is reformed and new peptides are synthesized.
Threatened Amphibians of the World
Pandian and Marian 1986). As the number of frog legs exported from these
countries declined, Indonesian exports rose markedly; currently Indonesia is one
of the major exporters of frog legs in the world (Kusrini and Alford 2006). The
maximum level of exports from Indonesia occurred in 1992, when approximately
5,600 metric tonnes, valued at US$23,596,841, were exported (Kusrini and Alford
2006). This underestimates the total harvest substantially as it ignores the large
domestic market (Kusrini 2005). The total harvest of frogs in Indonesia may be
as high as 400 million individuals per annum (Kusrini 2005).
Concern regarding the impact of frog harvesting is related not only to the
viability of frog populations, but also to the fact that frogs provide ecosystem
and even economic services, in particular pest control in agricultural systems
(Abdulali 1985; Pandian and Marian 1986). It has been suggested that frog
harvesting and pesticide use may interact synergistically: decreases in frog
populations caused by harvesting may allow pest populations to increase,
leading to increased use of pesticides, which may then have greater negative
effects on frogs (Abdulali 1985; Pandian and Marian 1986).
The large number of frog legs exported from Indonesia has raised concern
regarding the sustainability of populations of its edible frogs. During the early
1990s several Indonesian Rana species were proposed for listing on CITES Ap-
pendix II (Martens 1991). This proposal was rejected, perhaps in part for political
reasons, but primarily because there was insufficient data on their population
status and the extent of the trade to justify a CITES listing (Favre 1989; Schmuck
2000). More recently, research has shown that there is no evidence that
harvesting has seriously depleted the populations of edible frogs, particularly
the species that inhabit rice fields. Kusrini (2005) showed that harvesting in
Indonesia is mostly accomplished using highly targeted manual techniques that
do not disturb the habitat or affect populations of non-target species. Harvesting
rates are constrained by the need to harvest during phases of the rice growing
cycle when frogs can be efficiently captured without interfering with the crops.
Kusrini suggested that populations of Indonesian rice field species have been
able to withstand continuous harvest pressure because of these constraints
on harvesting pressure, combined with their life history traits, which include
rapid growth, early maturation, and relatively high reproductive output, and
also with their ability to thrive in disturbed landscapes. This may not be true
for species harvested in other habitats or environments. Stream-breeding frogs,
for example, often appear to have lower reproductive capacities and slower
population turnover, which may make them more vulnerable to the effects
of harvesting. Temperate species that must concentrate their reproductive
activity into restricted habitats during short growing seasons may have lower
rates of replacement and be more vulnerable during their short, concentrated
reproductive episodes. There is a need to improve the quality and quantity of
information available on harvesting rates and trade in frogs and frog products,
and our understanding of the biology of the species harvested, so that develop-
ing problems can be averted before they become serious.
Critiques of frog harvesting have mostly neglected the human dimension.
Pandian and Marian (1986) briefly mentioned that the ban on Indian frog leg
exports resulted in the collapse of this industry and the loss of jobs for many
people. Since most frogs are caught from the wild, frog harvesting provides a
substantial income for unskilled labour in developing countries. In the case of
Indonesia, the trade has been well established for several decades and provides
employment and a livelihood for many local people. Although Kusrini (2005)
showed that the present Indonesian harvest appears to be sustainable, it may
be approaching the critical threshold of unsustainability. In the case of Indonesia,
it is of concern that the sizes and numbers of frogs harvested are not regulated,
Most amphibian antimicrobial peptides (AMPs) are composed of 10 to
46 amino acids. AMPs tend to contain positively charged amino acids and
hydrophobic amino acids and tend to form an -helix that folds in such a
way that the positively charged regions and the hydrophobic regions are in
discrete areas of the molecule (Yeaman and Yount 2003). This allows them to
remain water soluble and capable of interacting with biological membranes.
They kill microorganisms by disrupting the plasma membrane or by entering
the cell and interfering with vital cellular processes such as DNA replication
and protein synthesis.
Several hundred unique AMPs have been isolated from amphibian species,
and the number is rapidly growing (Conlon et al. 2004; Pukala et al. 2006). They
fall into a number of related families, but each species has its own repertoire
of peptides with very little species overlap. Most of the AMPs were isolated
and characterized by classical biochemical purification methods (high pressure
Hoplobatrachus rugulosus (Least Concern) for sale in a market in Phnom Penh.
© Jodi Rowley/Robert Puschendorf
as they are in many other countries such as Romania (Török 2003). Additionally,
exported frogs are often misidentified (Veith et al. 2000). In January 2007, the
specific international Harmonized System Code for frogs’ legs will be dropped
from the Tariff of the World Customs Organization, making monitoring of the
international trade more difficult. More thorough and evenly applied monitoring
is needed within harvesting and exporting countries to ensure that populations
are not being harvested at unsustainable levels, and the international trade
needs to be more, rather than less, thoroughly monitored, so that the sources
and destinations of traded frogs can be more reliably traced.
Mirza D. Kusrini and Ross A. Alford
References
Abdulali, H. 1985. On the export of frog legs from India. Journal of the Bombay Natural
History Society 2:347-375.
Favre, D.S. 1985. International trade in endangered species: A guide to CITES. Martinus
Nijhoff Pub, Netherlands. 415 pp.
Hayes, M.P. and Jennings, M.R. 1986. Decline of ranid frog species in western North America:
Are bullfrogs (Rana catesbiana) responsible? Journal of Herpetology 20:490-509.
Jennings, M.R. and Hayes, M.P. 1985. Pre-1900 overharvest of California red-legged frogs
(Rana aurora draytonii): The inducement for bullfrog (Rana catesbiana) introduction.
Herpetologica 41:94-103.
Iskandar, D.T. 1998. Amfibi Jawa dan Bali. Puslitbang Biologi-LIPI, Bogor, Indonesia. 132 pp.
Kusrini, M.D. 2005. Edible frog harvesting in Indonesia: Evaluating its impact and
ecological context. PhD thesis. School of Tropical Biology, James Cook University,
Townsville, Australia.
Kusrini, M.D. and Alford, R.A. 2006. Indonesia’s exports of frogs’ legs. Traffic Bulletin 21:13-24.
Mazzoni, R., Cunningham, A.A., Daszak, P., Apolo, A., Perdomo, E. and Speranza, G.
2003. Emerging pathogen of wild amphibians in frogs (Rana catesbeiana) farmed for
international trade. Emerging Infectious Diseases 9:995-998.
Meronek, T.G., Copes, F.A. and Coble, D.W. 1997. The Bait Industry in Illinois, Michigan,
Minnesota, Ohio, South Dakota, and Wisconsin. Technical Bulletin Series #105, USDA’s
Cooperative State Research, Education and Extension Service, USA.
Okeyo, D.O. 2004. The delicacy of Giant Bullfrog eating in Namibia. Science in Africa
http://www.scienceinafrica.co.za/2004/january/bullfrog.html
Pandian, T.J. and Marian, M.P. 1986. Production and utilization of frogs: An ecological
view. Proc. Indian Acad. Sci. (Anim. Sci.) 95:289-301.
Schmuck, J. 2000a. Trade and species conservation. In: R. Hofrichter (ed.), The encyclopedia
of amphibians, pp. 228-241. Key Porter Books Lim., Ontario, Canada.
Török, Z. 2003. Action plan for sustainable exploitation of Rana ridibunda stocks from the
Danube delta biosphere reserve (Romania). Froglog 60.
Veith, M., Kosuch, J., Feldmann, R., Martens, H. and Seitz, A. 2000. A test for correct
species declaration of frog legs imports from Indonesia into the European Union.
Biodiversity and Conservation 9:333-341.
Figure 1. Total export of Indonesian frogs legs from 1969 to 2000 (Source:
Foreign Trade Statistical Bulletin, BPS).
liquid chromatography, testing for antimicrobial activity, Edman degradation
sequencing of active peptides) (Conlon et al. 2004).
Most amphibian AMPs have been isolated for their antibacterial proper-
ties. They may be active against a broad spectrum of pathogens or active
against a limited subset. One example of a broad spectrum peptide is caerin
1.1 from Litoria caerulea (LC). Caerin 1.1 has been shown to inhibit Gram
positive and some Gram negative bacteria, inhibit growth of cancer cells,
inhibit viral infection of target cells [human immunodeficiency virus (HIV) and
herpes simplex virus], prevent growth of the malaria parasite, and kill nema-
todes (Apponyi et al. 2004; VanCompernolle et al. 2005). Another example
of a recently discovered antibacterial peptide is brevinin-2TSa isolated from
the Tsushima Brown Frog (Rana tsushimensis, LC) from Japan. This peptide
effectively killed several important Gram negative pathogens (Klebsiella pneu-
moniae, Enterobacter cloacae, Pseudomonas aeruginosa, Proteus mirabilis)

as well as problematic Gram positive bacteria (Staphylococcus aureus and
methicillin-resistant S. aureus, Staphylococcus epidermidis, and Enterococcus
faecalis). Yet, it was not harmful to red blood cells suggesting it would be a
good candidate for drug development (Conlon et al. 2006).
Fewer of the amphibian AMPs have been tested for their antiviral
activities. Because most of them are membrane active, it is very likely that
many of them will be effective agents to inhibit infection of target cells by
enveloped viruses such as HIV and influenza virus. Six amphibian AMPs
inhibited infectivity by a frog ranavirus (frog virus 3) and a fish herpesvirus
(channel catfish virus) (Chinchar et al. 2001, 2004) and 13 amphibian AMPs
were tested for their ability to inhibit transmission of HIV. Three peptides
from Australian frogs (caerin 1.1, caerin 1.9, and maculatin 1.1) were potent
inhibitors of viral infectivity and could interfere with the transfer of virus from
dendritic cells to lymphocytes (VanCompernolle et al. 2005). Brevinin-1 from
Daruma Pond Frog Rana porosa inhibited infection by herpes simplex virus
type 1 and type 2 (Yasin et al. 2000).
Another important set of pathogens are fungal pathogens. The chytrid
fungus, Batrachochytrium dendrobatidis (Bd), is thought to be the cause
of recent amphibian declines in the USA, Australia, and Central America
(Lips et al. 2006; and see Essay 4.5). Many amphibian AMPs potently inhibit
growth of Bd (Rollins-Smith and Conlon 2005), and those species whose
peptides are most active against Bd in the laboratory tend to survive better
in nature (Woodhams et al. 2006a,b). Many amphibian AMPs are also ef-
fective against the human fungal pathogen, Candida albicans (Conlon et al.
2004), which causes thrush.
A few amphibian AMPs have been tested for their ability to kill malarial
parasites. Caerin 1.8 from Litoria chloris (LC; Apponyi et al. 2004) and
dermaseptin S3 from Phyllomedusa sauvagii (LC) have been shown to have
activity against malaria parasites, and S3 selectively kills parasites within
host erythrocytes without harming the erythrocytes (Ghosh et al. 1997).
The mucous of frog skin is a rich environment for microorganisms, and
control of potential pathogenic organisms is essential for life. It seems
clear that the amphibian AMPs in the mucous play an essential role in
protection of the species against skin pathogens. The peptide families
are conserved, and the peptides can cooperate to be much more effec-
tive as mixtures.
What might this teach us that could benefit human health? Antimicrobial
peptides have potential for development as therapeutic agents for use on the
skin or at mucosal surfaces. Further, because most of the AMPs are small,
they can be chemically synthesized. While costs of chemical synthesis are
one barrier to the development of AMPs as effective drugs, safety and ef-
ficacy are also important concerns. It seems only reasonable that we should
learn all that we can from the amphibians about how these unique peptides
protect them and exploit this knowledge for the benefit of human health as
well as the health of amphibians.
Louise A. Rollins-Smith
ESSAY 2.3. THE CULTURAL IMPORTANCE OF THE MEXICAN AXOLOTL AMBYSTOMA MEXICANUM
The term axolotl (ajolote in Spanish) originates from the ancient Aztec language,
Nauatal (in which it is properly pronounced ‘ashólotl’). Amongst its many transla-
tions it means: water slave, water-servant, water sprite, water monster or water
dog (from atl for water and xolotl which has various meanings). These names
are derived from Aztec mythology where the axolotl was considered to be the
transfiguration of Xolotl, the twin brother of the famous feathered serpent deity
named Quetzalcoatl. As with Quetzalcoatl, the exact significance and attributes
of Xolotl have varied somewhat between Mesoamerican civilizations throughout
history. Xolotl has been known as the God of the Evening Star (the underworld)
who ruled over games, deformities, twins, and other irregularities in nature.
He was also attributed with powers over the dead and the resurrected, having
An early depiction of Xo-
chimilca farmers creat-
ing chinampas by piling
up mud, vegetation and
faeces inside corrals of
the water-loving willow
(Salix bonplandiana).
An early Spanish image of the Axolotl in the Florentine Codex. Fray Bernardino
de Sahagun described the Axolotl as a lizard-like animal with an eel-like tail
and ears on the neck.
Chapter 2. Why Save Amphibians?
References
Apponyi, M.A., Pukala, T.L., Brinkworth, C.S., Vaselli, V.M., Bowie, J.H., Tyler, M.J., Booker,
G.W., Wallace, J.C., Carver, J.A., Separovic, F., Doyle, J., and Llewellyn, L.E. 2004.
Host-defence peptides of Australian anurans: structure, mechanisms of action and
evolutionary significance. Peptides 25:1035-1054.
Chinchar, V.G., Wang, J. Murti, G., Carey, C., and Rollins-Smith, L. 2001. Inactivation of frog
virus 3 and channel catfish virus by esculentin-2P and ranatuerin-2P, two antimicrobial
peptides isolated from frog skin. Virology 288:351-357.
Chinchar,V.G., Bryan. L., Silphadaung, U., Noga E., Wade, D., Rollins-Smith L. 2004. Inactiva-
tion of viruses infecting ectothermic animals by amphibian and piscine antimicrobial
peptides. Virology 323:268-275.
Conlon, J.M., Al-Ghaferi, N., Abraham, B., Sonnevend, A., Coquet, L., Leprince, J.,
Jouenne, T., Vaudry, H., and Iwamuro, S. 2006. Antimicrobial peptides from the
skin of the Tsushima brown frog Rana tsushimensis. Comparative Biochemistry and
Physiology, Part C 143:42-49.
Conlon, J.M., Kolodziejek, J., and Nowotny, N. 2004. Antimicrobial peptides from ranid
frogs: taxonomic and phylogenetic markers and a potential source of new therapeutic
agents. Biochimica et Biophysica Acta 1696:1-14.
Conlon, J.M., Sonnevend, Á., Patel, M., Davidson, C., Nielsen, P.F., Pal, T., and Rol-
lins-Smith, L.A. 2003. Isolation of peptides of the brevinin-1 family with potent
candidacidal activity from the skin secretions of the frog Rana boylii. Journal of
Peptide Research 62:207-213.
Ghosh, J.K., Shaool, D., Guillaud, P., Ciceron, L., Mazier, D., Kustanovich, I., Shai, Y.,
and Mor, A. 1997. Selective cytotoxicity of dermaseptin S3 toward intraerythrocytic
Plasmodium falciparum and the underlying molecular basis. Journal of Biological
Chemistry 272:31609-31616.
Lazarus, L.H. and Attila, M. 1993. The toad, ugly and venomous, wears yet a precious
thought to be responsible for escorting newly deceased souls to the afterlife.
Conversely, he was also believed to have brought mankind and fire from the
underworld. One popular myth describes Xolotl attempting to escape death by
disguising himself into various forms which were either monstrous or double
in nature such as double-stalked corn or agave plants (Smith 1969). As each
disguise was uncovered by the Gods, Xolotl made one final attempt to escape
detection by transforming himself into the axolotl. It was in this state that he
was finally captured and sacrificed to nourish the sun and the moon, thereby
helping to ensure that the cycle of night and day would continue.
The Axolotl Ambystoma mexicanum is endemic to the canals of Lake
Xochimilco, at the southern edge of the valley of Mexico. Archaeological
excavations in the Xochimilco basin show that from at least 6000 BC Axolotls
were part of a rich lacustrine habitat where it was considered the top predator
(Shaffer 1989; Duhon 1997). Although the wetland was used for agricultural
purposes before Aztec occupation, the development of chinampas (man-made,
tree-lined corrals of agricultural land separated by canals) intensified during
the construction of the Aztec capital Tenochtitlan.
For thousands of years the Axolotl has been used as a food source (Smith
1989). When the Spanish arrived, it was still an important part of the local
economy, as the Franciscan missionary, Friar Bernardino de Sahagun, wrote
in his seminal manuscript, the Florentine Codex (Sahagun 1938):
“There are some creatures in the water that are called axolotl that have
feet and hands like small lizards, and they have the tail of an eel and the
body as well; they have a very wide mouth and whiskers at the neck. It is
very good to eat; it is the food of the lords.”
Indeed, the Axolotl was considered such good eating that celebrated
Spanish naturalist Francisco Hernandez (1959) thought it “provoked lechery”. It
has also been immortalized by the renowned artist Diego Rivera in his famous
mural depicting the axolotl as a product in the ancient market of Tenochtitlan,
and in Julio Cortazar’s well-known essay “Axolotl” in his collection of short
stories The End of the Game.
Today, the Axolotl continues to play a role in the local economy. Various
medicines and tonics that feature the Axolotl as the main ingredient are sold
in local markets and are believed to provide remedies for respiratory ailments,
such as bronchitis. Similarly, a captive-bred population of a close relative,
Ambystoma dumerilli (or Achoque), is harvested by nuns in Pascuaro, in north-
western Michoacán, to make a medicinal syrup. This is also believed to cure
lung illnesses and help children suffering from anaemia (McKay 2003). Many
of the medical powers accredited to the Axolotl may stem from its ability to
regenerate damaged or destroyed body parts, including limbs, parts of the
cornea, brain, and heart cells. These abilities have made the Axolotl the focus
of widespread medical research since the late 1800s.
29
jewel in his skin. Progress in Neurobiology 41:473-507.
Lips, K.R., Brem ,F., Brenes, R., Reeve, J.D., Alford, R.A., Voyles, J., Carey, C., Livo, L.,
Pessier, A.P., and Collins, J.P. 2006. Emerging infectious disease and the loss of
biodiversity in a Neotropical amphibian community. Proceedings of the National
Academy of Sciences USA 103:3165-3170.
Pukala,T.L., Bowie, J.H., Maselli, V.M., Musgrave, I.F., and Tyler, M.J. 2006. Host-defence
peptides from the glandular secretions of amphibians: structure and activity. Natural
Product Reports 23:368-393.
Rollins-Smith, L.A. and Conlon, J.M. 2005. Antimicrobial peptide defenses against chytrid-
iomycosis, an emerging infectious disease of amphibian populations. Developmental
and Comparative Immunology 29:589-598.
VanCompernolle, S.E., Taylor, R.J., Oswald-Richter, K., Jiang, J., Youree, B.E., Bowie,
J.H., Tyler, M.J., Conlon J.M., Wade, D., KewalRamani, V.N. Dermody T.S., Aiken,
C., Rollins-Smith L.A., and Unutmaz, D. 2005. Amphibian antimicrobial skin peptides
potently inhibit HIV infection and transfer of virus from dendritic cells to T cells.
Journal of Virology 79:11598-11606.
Woodhams, D.C. Rollins-Smith, L.A., Carey, C., Reinert, L., Tyler, M.J., and Alford, R.
2006a. Population trends associated with antimicrobial peptide defenses against
chytridiomycosis in Australian frogs. Oecologia 146:531-540.
Woodhams, D.C., Voyles, J., Lips, K.R., Carey, C., and Rollins-Smith, L.A. 2006b. Predicted
Disease Susceptibility in a Panamanian Amphibian Assemblage Based on Skin Peptide
Defenses. Journal of Wildlife Diseases 42:207-218.
Yasin, B., Pang, M., Turner, J.S., Cho, Y., Dinh, N.N., Waring, A.J., Lehrer, R.I., and
Wager, E.A. 2000. Evaluation of the inactivation of infectious Herpes simplex virus
by host-defense peptides. European Journal of Clinical Microbiology and Infectious
Diseases 19:187-194.
Yeaman, M.R., and Yount, N.Y. 2003. Mechanisms of antimicrobial peptide action and
resistance. Pharmacological Reviews 55:27-55.
Antimicrobial peptides in leopard
frog (Rana pipiens) skin secre-
tions dramatically reduce skin
microbes. The left panel shows
bacteria developing on growth
medium from skin sampled before
induction of skin secretions by
injection of norepinephrine. The
right panel shows growth from
skin of the same frog sampled 15
minutes after skin peptide induc-
tion. © Douglas C. Woodhams,
Vanderbilt University
Due to its diminished population and protected status in the wild, the
Axolotl is not utilized to the extent that it was in the past. Nevertheless,
they are still illegally harvested from the wild and are sold in markets for the
local pet trade, biomedical research, and as a source of food and medicine
(Griffiths et al. 2004). Due to a lack of trade data, it is difficult to measure
the impact of this offtake, mainly because the market operates clandestinely
and often involves a related species, Ambystoma tigrinum, which is easily
confused with the Axolotl.
Because of its mythological connections and prominent position within
the ancient lacustrine economy of the region, the Mexican Axolotl has
become something of a cultural icon. Therefore, despite its precarious
status in the wild, this remarkable species continues to represent not only
a significant symbol of the past, but also a sobering example of the need
for imaginative solutions required for the present. Indeed, with local people
increasingly engaging with current attempts to promote the Axolotl as a
flagship species for the conservation of Xochimilco’s natural and cultural
diversity (Griffiths et al. 2004), there is real hope that the species will
continue to provide an important catalyst for sustainable development
in the future.
Jeanne E. McKay, Richard A. Griffiths and Ian G. Bride
References
Duhon, S.T. 1997. The axolotl in its native habitat-yesterday and today. Axolotl News
26:14-17.
Griffiths, R.A., Graue. V., Bride, I.G and McKay, J.E. 2004. Conservation of the axolotl (Am-
bystoma mexicanum) at Lake Xochimilco, Mexico. Herpetological Bulletin 89:4-11.
Hernandez, F. 1959. Historia Natural de Nueva Espana, Coleccion Obras Completas Vol.
IV. Universidad Nacional Autonoma de México, México City, México.
McKay, J. 2003. An evaluation of captive breeding and sustainable use of the Mexican axol-
otl (Ambystoma mexicanum). MSc Dissertation. University of Kent, Canterbury, UK.
Sahagun, Fr. Bernardino de. 1938. Historia General de las Cosas de Nueva Espana Vol. III.
In: P. Robredo (ed.), pp. 195. México D.F., México.
Schaffer, H.B. 1989. Natural history, ecology and evolution of the Mexican axolotl.
Axolotl News 18:5-11.
Smith, H.M. 1969. The Mexican Axolotl: Some misconceptions and Problems. Bioscience
19:593-597.
Smith, H. 1989. Discovery of the axolotl and its early history in biological Research. In:
J.B. Armstrong and G.M. Malacinski (eds.), Developmental Biology of the Axolotl, pp.
3-12. Oxford University Press, New York, USA.

Participants at the China and the Koreas GAA
workshop in Chengdu, China, in March 2002.
Participants at the Southeast Asia GAA work-
shop in Bangkok, Thailand, in September-
October 2002.
Participants at the Mesoamerica GAA work-
shop in La Selva, Costa Rica, in November
2002.
CHAPTER 3. THE GLOBAL AMPHIBIAN ASSESSMENT (GAA):
HISTORY, OBJECTIVES AND METHODOLOGY
Janice Chanson, Simon Stuart, Neil Cox, Bruce Young and Michael Hoffmann
BACKGROUND
While there have always been some instances of localized amphibian declines, often in
response to changes in the local environment, scientists had started to become increasingly
concerned about the possibility of widespread and severe population declines when they
met during 1989 at the First World Congress of Herpetology. A review of historical data
suggested that significant amphibian declines had begun as early as the 1970s in the western
United States, Puerto Rico and north-eastern Australia (Czechura and Ingram 1990; Kagarise
Sherman and Morton 1993; Drost and Fellers 1996; Burrowes et al. 2004).
Subsequent reports from field sites revealed the true severity of the population declines.
At one site in Costa Rica, 40% of the amphibian fauna had disappeared over a short period in
the late 1980s (Pounds et al. 1997). In addition, sudden disappearances of montane species
had also been simultaneously noted elsewhere in Costa Rica, and in Ecuador, and Venezuela
(Pounds and Crump 1994; Pounds et al. 1997; Young et al. 2001; Ron et al. 2003). In some
regions, the declines were taking place in seemingly pristine habitats (Czechura and Ingram
1990; Kagarise Sherman and Morton 1993; Pounds and Crump 1994; Drost and Fellers 1996;
Pounds et al. 1997; Young et al. 2001; Ron et al. 2003; Burrowes et al. 2004).
Some scientists were initially skeptical about the reports; amphibian populations were
already well known to sometimes fluctuate widely from year to year (Pechmann and Wilbur
1994). However, by using statistical tests and models, it was shown that the declines were
far more widespread and severe than would be expected under natural conditions (Pounds
et al. 1997). This evidence, in addition to many further reports of declines in the 1990s
(Laurance et al. 1996; Lips 1998; Lynch and Grant 1998; Houlahan et al. 2000; Young et al.
2001), convinced most herpetologists that non-random and severe declines were taking
place in the populations of many amphibian species.
THE INITIATION OF THE GAA AND ITS OBJECTIVES
The lack of a comprehensive global picture on the extent and severity of amphibian declines
prompted IUCN, in partnership with Conservation International and NatureServe, to undertake
the Global Amphibian Assessment (GAA), which commenced in 2001. The over-arching goal
of the GAA was to stimulate concerted and well-targeted activities to halt the current wave
of amphibian extinctions, through the development of an information baseline on amphibian
status and conservation needs. The initial objectives of the GAA included:
• Identification of the most threatened amphibian species, using the IUCN Red List Cat-
egories and Criteria (see Appendix I) to determine which species were in need of urgent
conservation action.
• Identification of the most important geographic regions and habitats that need to be
conserved in order to prevent further amphibian extinctions.
• Identification of the most important threats to amphibians globally, specifically those
resulting in rapid declines, and to propose both mitigating measures and prioritized
conservation actions for these.
• Comparing the conservation needs for amphibians with those for mammals and birds,
thus making it possible to determine the extent to which conservation initiatives can
efficiently and simultaneously address the needs of all of these species’ groups.
• Increasing and building the capacity of the IUCN Species Survival Commission expert
network for amphibians, thereby allowing scientists to coordinate their conservation
activities at the global level, and providing a base of expertise from which information
collected during the GAA could be regularly updated.
To meet these ambitious objectives the GAA had to develop a sound methodology by
which it could engage the scientific community and gather data on the distribution, abun-
dance, population trends, habitat requirements, and threats for all of the ~6,000 described
species of amphibians.
METHODOLOGY
The GAA process involved three core phases of data compilation: initial data collection;
data review; and data correction and consistency check.
The task of collecting the initial data was divided into 33 geographic regions (Appendix
II) that were defined to cover the global distribution of all amphibians. A network of regional
coordinators – scientists with a broad knowledge of the region’s amphibian species – were
then identified and given the responsibility of collecting data on the amphibians within their
region. During the initial data collection phase, regional coordinators were asked to produce
distribution maps of each species’ current geographic range, which were then digitized in a
geographic information system (ArcView 3.x). These maps are most often in a standardized
form of polygons that join known point localities.
In addition to range maps, coordinators provided general information on habitat require-
ments, population status, major threats and conservation measures for each species, and
were also asked to make use of the IUCN Classification Schemes for coding up Countries of
Occurrence, Habitat Preferences, Major Threats, Conservation Measures, and Utilization1.
Finally, regional coordinators were asked to provide a provisional Red List assessment of the
conservation status of each species by applying the IUCN Red List Categories and Criteria
(Appendix I; and see later).
Once the initial data had been collected, they were subjected to a process of peer review.
More than 500 amphibian specialists and experts from over 60 countries participated in this
process. The means of undertaking reviews varied from region to region, but in most cases
(usually the more species-rich regions) the process was handled through expert workshops.
However, in other regions, peer-review of data was managed through individual review by
leading herpetologists. The following GAA workshops have been held to date:
• Australia – Hobart, Tasmania, 6-8 February 2001.
• China and the Koreas – Chengdu, China, 18-21 March 2002.
• Sub-Saharan Africa – Watamu, Kenya, 16-18 April 2002.
• South Asia – Coimbatore, India, 1-5 July 2002.
• Southeast Asia – Bangkok, Thailand, 30 September – 4 October 2002.
• Mesoamerica (Mexico south through Panama) – La Selva, Costa Rica, 11-15 November
2002.
• The Papuan Region – Hawaii, United States, 24-27 February 2003.
• Tropical South America East of the Andes (merging the Amazonian Brazil, Atlantic
Forest-Cerrado-Caatinga, Paraguay, and Guianas regions, and parts of the Colombia,
Ecuador, Peru, Bolivia and Venezuela regions) – Belo Horizonte, Brazil, 31 March – 4
April 2003.
• Tropical Andes (merging the remaining parts of the Venezuela, Colombia, Ecuador, Peru,
and Bolivia regions) – Tandayapa, Ecuador, 18-22 August 2003.
• Madagascar – Gland, Switzerland, 22-25 September 2003.
• Chile – Concepción, Chile, 3-4 October 2003.
• Argentina and Uruguay – Puerto Madryn, Argentina, 12-14 October 2003.
• Caecilians (the only taxonomic rather than geographically based workshop) – London,
23-25 February 2004.
• Caribbean – Santo Domingo, Dominican Republic, 19-21 March 2004.
• Mediterranean – Malaga, Spain, 13-17 December 2004.
Data review has been completed by correspondence with amphibian experts in the
following regions: Northern Eurasia; Seychelles Islands; Japan; Australia (a continuation
of the initial data collection workshop); New Zealand; West Asia; North America; Europe;
and North Africa.
 Chapter 3. The Global Amphibian Assessment (GAA): History, Objectives and Methodology 31

Following the review process, the GAA coordinating team undertook a final review of all
the data to ensure that there had been a consistent approach in the collection and quality
of the data between the different regions and taxonomic groups (members of the GAA
coordinating team also attended all of the workshops to help promote consistent approaches
during the review process). To the extent possible within the constraints of time and budget,
when significant problems or uncertainties were encountered in the data, the coordinating
team contacted the experts who contributed information about the species to ensure that
any changes were correctly made, or information clarified, in order to ensure accurate
and consistent conservation status assessments for each individual species. However, it
is the GAA coordinating team, not the individual data providers, that takes responsibility
for the content of each species account. The data providers are listed as “assessors” on
each species account, but because of the limitations in time and funding, they have not all
been given the opportunity to sign off on the content of each account to which their name
is attached as an “assessor”.

DATA LIMITATIONS AND CAVEATS

Taxonomy
As already noted under the introductory chapter, the taxonomic framework used for the
Global Amphibian Assessment follows Frost (2004). Regional coordinators were initially
asked to follow the systematic arrangement provided in Version 2.2 of Amphibian Species
of the World (2000), and were only permitted to depart from this classification in a few
well-justified circumstances. The GAA has continued to follow revisions to the ASW up to
Version 3.0 (Frost 2004). The significant changes given in the most recent update of the
ASW are discussed in the introductory chapter.

Distribution maps Participants at the Papuan region GAA workshop in Hawaii, United States, in February 2003.

The GAA process has endeavored to produce maps for all of the ~6,000 amphibian species;
however, in a very few instances (~100) it has been impossible to produce a meaningful
map, mostly for species that are currently known only from non-specific type localities. The
maps are in the form of polygons that join known locations. A species’ distribution map can
consist of more than one polygon where there is an obvious discontinuity in suitable habitat.
Data attached to each polygon include presence (e.g., extant, extirpated) and origin (native,
introduced, reintroduced). In some cases, polygons were clipped to geographical features,
such as rivers, or elevational contours.

IUCN Red List Categories and Criteria

The IUCN Red List of Threatened Species2 (hereafter referred to as the Red List) is the
accepted standard for species global threat status (Lamoreux et al. 2003; Rodrigues et
al. 2006). The identification of threatened species is of great importance to biodiversity
conservation, since it enables practitioners to target those species known to be at highest
risk of extinction. The first Red List assessments were largely subjective and qualitative,
and primarily focused on a few hand-picked species. However, in 1994, IUCN introduced
a system of categorical rankings employing quantitative criteria and representing several
advances, including: enabling consistent application by different people, being based around
a probabilistic assessment of extinction risk, the incorporation of a time-scale, and the ability
to handle uncertainty. The adoption in 2001 of the most recent version of the categories
and criteria (IUCN 2001; Appendix I), has enabled considerable improvement of the rigor,
justification, and transparency of Red List assessments.
Despite the development of objective criteria, consistency and subjectivity in the applica-
tion of these remains an issue. The Red List criteria are designed to handle uncertainty, for
example uncertainties in population size or trends and distribution (Akçakaya et al. 2000),
but when there is inadequate information to make an assessment of extinction risk, the
category Data Deficient must be used. In the course of undertaking a Red List assessment,
there is a risk that errors will be introduced, specifically errors of omission (a species that
is genuinely threatened is included in a non-threatened category, such as Data Deficient
or Least Concern) and commission (a species that is not genuinely threatened is classed
in a threatened category). The attitudes of assessors to such risk and uncertainty range
along a spectrum from evidentiary (demanding substantial evidence of threatened status
before allowing such a classification) to precautionary (allowing a species to be listed in Participants at the Tropical South America East
a non-threatened category only if there is strong evidence that it is not threatened). For a information that was published before that date but which was only made available to the of the Andes region GAA workshop in Belo
well-known species with extensive quantitative data, the effects of the assessors’ attitudes GAA Coordinating Team after that date, is also not included here (and, as already noted, Horizonte, Brazil, in March-April 2003.
on listings are generally small. However, when the level of uncertainty is high, attitudes can the overarching taxonomic classification follows Frost 2004). However, the information
easily cause the observed discrepancies. Assessors with strongly precautionary attitudes presented in the species accounts has been fully edited for content and style, and a number
are concerned mainly about omission errors, whereas assessors with strongly evidentiary of corrections have been made usually to correct inconsistencies in data or obvious data
attitudes are concerned mainly about commission errors. Seeking to minimize the risk of one errors. In a few exceptional instances, we have introduced footnotes, where we feel it
type of error inevitably increases the risk of the other type. IUCN is concerned about both is necessary to draw attention to important recent findings (for example, rediscovery of
types of error and, therefore, seeks to find a balance between the extremes. The purpose Conraua derooi in West Africa).
of the consistency check was to help ensure that the Red List category selected for each
species reflected this balance. Rapidly Declining Species
Improved training in the use of the Red List criteria, particularly at a regional level, and
assessor awareness of issues relating to criteria application can help ensure consistency and In view of the rapid changes that are taking places in the status of amphibian species in
minimize discrepancies between the global IUCN Red List and national Red Lists. many parts of the world, the GAA Coordinating Team attempted to define a group of spe-
cies as “rapidly declining”, based on an estimate of the likely IUCN Red List status of each
Brazilian species species in 1980. Drawing on the data in the 2004 and 2006 IUCN Red List assessments, in
particular on information on population trends, habitat decline trends, threatening processes
For 113 species endemic to Brazil, it was not possible to reach agreement on the Red List and conservation actions, one member of the GAA Team (Simon Stuart) determined the
category between the GAA Coordinating Team and the experts on the species in Brazil (Ap- likely Red List category for each species in 1980. This was estimated conservatively, with
pendix XIII). The Red List category for the individual species listed in this book are those the default position being that no change had occurred unless there was strong evidence.
that were agreed at the GAA Brazil workshop in April 2003. However, in the subsequent Small changes in status were not considered to be sufficient to allow Red List category
consistency check conducted by the GAA Coordinating Team, many of these were found to change. Changes in category were generally related to dramatic changes in the status of
be inconsistent with the approach adopted elsewhere in the world. It is these consistent Red the species. Examples include: the disappearance of most, or all, of the population due
List categories that are used in the actual analyses presented in the introductory chapters to disease or poorly understood processes; a catastrophic decline due to over-harvesting;
of this book, and in Appendix V; the consistent categories are presented as footnotes in the or severe habitat loss (especially for species that are not tolerant of habitat disturbance).
species accounts in the book, where relevant. Back-calculating of Red List categories has previously been done for birds in order to detect
genuine trends in status over time, though this has only been done from 1994 back to 1988
Underlying data (BirdLife International 2004).
Amphibians that moved to a higher Red List threat category between 1980 and 2006
The data upon which the analyses in this book are based, and the content presented in the were defined as “rapidly declining” species. The assignment of individual species to the
individual species accounts that comprise the bulk of this work, are essentially the same “rapidly declining” group clearly requires further review. However, for broad multi-species
as the information contained within the 2007 IUCN Red List of Threatened Species. Any geographic analyses, the results give what seems to be a reliable overall picture of the
species described or revalidated after 31 December 2005, is not included. In addition, any patterns of amphibian declines.
32 Threatened Amphibians of the World
Participants at the Tropical Andes region GAA workshop in Tandayapa, Ecuador, in August 2003.
Participants at the Caribbean GAA workshop in Santo Domingo, Dominican Republic, in March 2004.
Endnotes REFERENCES
1 http://www.iucnredlist.org/info/authority_files and
http://www.iucn.org/themes/ssc/sis/authority. Akçakaya, H.R., Ferson, S., Burgman, M.A., Keith, D.A., Mace, G.M. and Todd, C.R. 2000. Making Consistent
htm IUCN Classifications under Uncertainty. Conservation Biology 14:1001-1013.
2 www.iucnredlist.org BirdLife International. 2004. State of the World’s Birds: Indicators for our Changing World. BirdLife
International, Cambridge, UK.
Burrowes, P.A., Joglar, R.L. and Green, D.E. 2004. Potential causes for amphibian declines in Puerto Rico.
Herpetologica 60:141-154.
Czechura, G.V. and Ingram, G. 1990. Taudactylus diurnus and the case of the disappearing frogs. Memoirs
of the Queensland Museum 29:361-365.
Drost, C.A. and Fellers, G.M. 1996. Collapse of a regional frog fauna in the Yosemite area of the Sierra
Nevada. Conservation Biology 10:414-425.
Frost, D.R. 2004. Amphibian Species of the World: an Online Reference. Version 3.0 (22 August, 2004).
Electronic Database accessible at http://research.amnh.org/herpetology/amphibia/index.html. American
Museum of Natural History, New York, USA.
Houlahan, J.E., Findlay, C.S., Schmidt, B.R., Meyer, A.H. and Kuzmin, S.L. 2000. Quantitative evidence for
global amphibian population declines. Nature 404:752-755.
Kagarise Sherman, C. and Morton, M.L. 1993. Population declines of Yosemite toads in the eastern Sierra
Nevada of California. Journal of Herpetology 27:186-198.
Lamoreux, J., Akçakaya, R., Bennun, L., Collar, N.J., Boitani, L., Brackett, D., Bräutigam, A., Brooks, T.M.,
Participants at the Chile GAA workshop in Concepción, Chile in October 2003.
Participants at the Southern Cone GAA workshop in Puerto Madryn, Argentina, in October 2003.
Fonseca, G.A.B., Mittermeier, R.A., Rylands, A.B., Gärdenfors, U., Hilton-Taylor, C., Mace, G., Stein, B.A.,
and Stuart, S. 2003. Value of the IUCN Red List. Trends in Ecology and Evolution 18:214-215.
Laurance, W. F., McDonald, K.R. and Speare, R. 1996. Epidemic disease and the catastrophic decline of
Australian rain forest frogs. Conservation Biology 10:406-413.
Lips, K.R. 1998. Decline of a tropical montane amphibian fauna. Conservation Biology 12:106-117.
Lynch, J.D. and Grant, T. 1998. Dying frogs in western Colombia: catastrophe or trivial observation? Revista
de la Academia Colombiana de Ciencias Exactas, Físicas y Naturales 23:149-152.
Pechmann, J.H.K. and Wilbur, H.M. 1994. Putting declining amphibian populations in perspective: natural
fluctuations and human impacts. Herpetologica 50:65-84.
Pounds, J.A., Fogden, M.P.L., Savage, J.M. and Gorman, G.C. 1997. Tests of null models for amphibian
declines on a tropical mountain. Conservation Biology 11:1307-1322.
Pounds, J.A. and Crump, M.I. 1994. Amphibian declines and climate disturbance: the case of the golden
toad and the harlequin frog. Conservation Biology 8:72-85.
Rodrigues, A.S.L., Pilgrim, J.D., Lamoreux, J.L., Hoffmann, M. and Brooks, T.M. 2006. The value of the Red
List for conservation. Trends in Ecology and Evolution 21:71-76.
Ron, S.R., Duellman, W.E., Coloma, L.A. and Bustamante, M.R. 2003. Population decline of the Jambato Toad
Atelopus ignescens (Anura: Bufonidae) in the Andes of Ecuador. Journal of Herpetology 37:116-126.
Young, B.E., Lips, K.R., Reaser, J.K., Ibanez, R., Salas, A.W., Cedeno, J. R., Coloma, L.A., Ron, S., La Marca,
E., Meyer, J.R., Munoz, A., Bolanos, F., Chavez, G. and Romo, D. 2001. Population declines and priorities
for amphibian conservation in Latin America. Conservation Biology 15:1213-1223.
 CHAPTER 4. THE STATE OF THE WORLD’S AMPHIBIANS

Janice Chanson, Michael Hoffmann, Neil Cox and Simon Stuart in a threatened category. Although the assessment revealed that 44% of amphibian spe-
cies are currently not considered to be globally threatened (these are species listed in the Red List Category Number of species
During the course of the 20th Century the global human population increased from 1.65 billion categories Near Threatened and Least Concern), the populations of most of these species Extinct (EX) 34
to 6 billion people. In April 2005, the United Nations launched the results of the Millennium are still decreasing, albeit not currently at a rate sufficient to meet the criteria for listing in Extinct in the Wild (EW) 1
Ecosystem Assessment1, a four-year effort involving 1,360 scientists from 95 countries a higher category of threat (see later). Critically Endangered (CR) 455
that aimed to determine more precisely how a burgeoning human population is impacting A total of 34 species4 are listed as officially Extinct5; 19 of these species are from Sri Endangered (EN) 768
the planet’s biodiversity and ecosystem services, and, conversely, how those changes are Lanka6 (see Figure 2; and see Essay 4.1). One species, the Wyoming Toad, Bufo baxteri, is Vulnerable (VU) 670
affecting human well-being. According to the Millennium Ecosystem Assessment, the considered Extinct in the Wild, since the surviving wild population persists only due to annual Near Threatened (NT) 369
world’s population doubled between 1960 and 2000, while the global economy gre six-fols releases of thousands of captive-reared toadlets (see Essay 11.7). Furthermore, within the Least Concern (LC) 2,236
over the same time-period. Perhaps not surprisingly then, over the past 50 years, humans category Critically Endangered, 130 of the 455 species are flagged as being Possibly Extinct Data Deficient (DD) 1,382
have changed the planet’s ecosystems more rapidly and more extensively than in any other (see Appendix IX). These typically are species that have not been seen for an extended period Total Number of Species 5,915
comparable period of time in human history, largely to meet rapidly growing demands for of time, but for which there is still some lagging hope that they may yet be hanging on in
1% 8%
food, fresh water, timber, fibre, and fuel. The most important direct drivers of change in their native habitat (see Butchart et al. 2006).
ecosystems are habitat change, overexploitation, invasive alien species, pollution, and 23%
13%
climate change (Millennium Ecosystem Assessment 2005).
Many of the dramatic findings of the Millennium Ecosystem Assessment are telling CONSERVATION STATUS BY CRITERIA
indictments of the anthropogenic influences that a rapidly technologically evolving human
society has brought to bear on earth’s natural resources. More land was converted to crop- In order for a species to qualify for one of the three threatened Red List categories (CR,
land in the 30 years after 1950 than in the 150 years between 1700 and 1850. Cultivated EN, VU), it must satisfy one or more of the five Red List criteria: A: Population reduction; 11%
systems (areas where at least 30% of the landscape is in croplands, shifting cultivation, B: Geographic range size and decline; C: Small population size and decline; D: Very small
confined livestock production, or freshwater aquaculture) now cover one-quarter of Earth’s or restricted population; and E. Quantitative analysis of the probability of extinction (see
terrestrial surface. In the last 20 years, some regions have experienced very high rates of Appendix I). By summarizing across species the criteria that trigger the Red List categories, 6%
forest loss, particularly in the tropics: in Central America, Amazonia, the Congo Basin, the 38%
forests of eastern Madagascar, and south-east Asia (mainly in lowland regions), and it is
projected that a further 10-20% of grassland and forestland will be converted between
2000 and 2050 (primarily to agriculture). More than two-thirds of the area of two of the
world’s 14 major terrestrial biomes and more than half of the area of four other biomes had Figure 1. Summary of Red List categories
been converted by 1990, primarily to agriculture. The amount of water impounded behind for all amphibian species. The percentage
dams quadrupled since 1960, and three to six times as much water is held in reservoirs of species in each category is given in the
as in natural rivers; most water use (70% worldwide) is for agriculture. Since 1750, the pie chart above.
atmospheric concentration of carbon dioxide has increased by about 32% (from about 280
to 376 parts per million in 2003), primarily due to the combustion of fossil fuels and land
use changes. Approximately 60% of that increase (60 parts per million) has taken place
since 1959 (Millennium Ecosystem Assessment 2005).
From a human welfare perspective, such dramatic changes are of primary importance
because of the direct linkages between ecosystems and the essential provisioning, regulating
and cultural services that they provide to humanity. That such changes already are having
a dramatic effect on human society is undeniable. We already know, for example, that the One of four small families of amphibians at
impacts of the massive Tsunami that struck Aceh province in Indonesia on December 26, risk of extinction, the Leiopelmatidae of New
2004, could have been significantly less were it not for the loss of coastal mangroves and Zealand is represented by only four species,
offshore coral reef systems (e.g., Marris 2005). Similarly, the aftermath of Hurricane Katrina, including Leiopelma hochstetteri (Vulnerable)
which devastated the Gulf Coast of the United States, and in particular the city of New from North Island, which is at possible risk
Orleans, may rekindle interest in bolstering the wetlands south of New Orleans to provide from the chytrid fungus. © Paddy Ryan
more of a barrier to future hurricanes (Travis 2005).
Unfortunately, these same aforementioned drivers of change are leading to a significant,
irreversible loss of Earth’s biodiversity – the very foundation upon which these ecosystem
services are built. Although some species respond positively to anthropogenic pressures,
the great majority show only limited tolerance of the increasingly widespread and rapid
changes impacting ecosystems worldwide. There is already ample evidence that current
species extinction rates are 100-1,000 times higher than the normal rate of extinction (Pimm
et al. 1995; Baillie et al. 2004). At least in the last 500 years alone, nearly 800 documented
extinctions have occurred, clearly an underestimate since this figure is based only on those
species actually known to have gone extinct within this time frame (Baillie et al. 2004).
And, at least one recent study suggests that we could lose at least three times this number
of species within just a few decades (Ricketts et al. 2005).
The Global Amphibian Assessment is the first-ever attempt to undertake a comprehensive
assessment of the conservation status of the world’s ~6,000 amphibian species. Only 124
threatened amphibians appeared on the 1996 IUCN Red List (Baillie and Groombridge 1996),
and only a few more were added to this number in the 2000 IUCN Red List (Hilton-Taylor
2000). Other than amphibians, only two other vertebrate Classes have been globally and
comprehensively assessed: mammals were assessed globally for the first time in 1996
(Baillie and Groombridge 1996), and, at the time of writing, are the focus of a massive revi-
sion through the IUCN-led Global Mammal Assessment; birds, on the other hand, a Class
of ~10,000 species, have been assessed globally four times since 1988 (most recently in
2004), with a fifth major global assessment planned for completion in 2008.
A comprehensive assessment of this nature permits us a clear picture of the state
and distribution of global biodiversity. However, of greatest interest, particularly among
amphibians, is the opportunity it presents to investigate the causes of the marked and rapid
declines and disappearances that have been observed in amphibian populations the world
over, especially those populations that have vanished in seemingly pristine habitats where Figure 2. The distribution of Extinct (n=34) and Extinct in the Wild (n=1; Bufo baxteri) amphibians (since 1500). Numbers correspond as follows:
apparent environmental degradation or anthropogenic impacts are low. 1-19 – Philautus malcolmsmithi, Adenomus kandianus, Nannophrys guentheri, Philautus adspersus, Philautus eximius, Philautus hypomelas,
Philautus leucorhinus, Philautus nasutus, Philautus temporalis, Philautus variabilis, Philautus oxyrhynchus, Philautus halyi, Philautus dimbul-
lae, Philautus nanus, Philautus rugatus, Philautus extirpo, Philautus stellatus, Philautus zal, Philautus zimmeri; 20 – Philautus travancorius;
GLOBAL CONSERVATION STATUS 21 – Rheobatrachus vitellinus; 22-23 – Rheobatrachus silus, Taudactylus diurnus; 24 – Cynops wolterstorffi; 25 – Discoglossus nigriventer;
26 – Plethodon ainsworthi; 27 – Rana fisheri; 28 – Bufo baxteri; 29 – Eleutherodactylus milesi; 30 – Eleutherodactylus chrysozetetes; 31 – Bufo
The GAA undertook risk assessments for 5,915 amphibian species2, categorizing one-third periglenes; 32 – Atelopus vogli; 33-34 – Atelopus longirostris, Atelopus ignescens; 35 – Phrynomedusa fimbriata.
of all extant species (1,893 out of 5,881) as threatened with extinction3 (Figure 1). This is
noticeably higher compared with other groups that have been globally and comprehensively
assessed, specifically mammals (23%) and birds (12%) (Baillie et al. 2004). Not only are
amphibians more threatened than mammals or birds, but they also have a higher propor- Red List Category Number of species A % of category B % of species C % of category D % of category
in category in category
tion of species on the verge of extinction. Some 21% of amphibians are classed in the
categories Critically Endangered or Endangered, compared with 10.5% of mammals and Critically Endangered (CR) 455 189 42 316 69 8 2 5 1
5.4% of birds. Worryingly, it seems increasingly certain that this high level of threat is a Endangered (EN) 768 50 7 730 95 4 1 2 0.3
significant underestimate of the true threat status of the class, since there is currently insuf- Vulnerable (VU) 670 36 5 463 69 1 0.1 177 26
ficient information available to assess the conservation status of 23% of amphibians (these
species are placed in the category Data Deficient). These poorly known species (a number
of which have only recently been described), are often rare and have small distributions, Table 1. A summary of the Red List criteria triggering each Red List category. No amphibians are listed under Criterion E, as quantitative analysis
and with the availability of more information, particularly on threats, may warrant listing of extinction risk requires considerably more data over longer time periods than is usually available for threatened amphibians.
34 Threatened Amphibians of the World

we can potentially obtain a better understanding of the factors driving species towards
Order/Family Total EX EW CR EN VU NT LC DD Number % Threatened extinction (Table 1)7.
threatened or Extinct Among Critically Endangered amphibians, 42% of the species meet the criteria for this
Anura category due to their having undergone a substantial population reduction (Criterion A), and
Allophrynidae 1 0 0 0 0 0 0 1 0 0 0 69% of species meet the geographic range criterion (Criterion B). The majority of amphibians
Arthroleptidae 51 0 0 3 9 2 3 18 16 14 27 that are recognized as threatened on the basis of population reduction most often fall within
Ascaphidae 2 0 0 0 0 0 0 2 0 0 0 the category of Critically Endangered, with 69% of species meeting Criterion A being within
Astylosternidae 29 0 0 2 11 8 2 5 1 21 72* this highest category of threat. Very few amphibian species meet either Criteria C or D (the
Bombinatoridae 10 0 0 0 1 4 0 5 0 5 50 exception being Vulnerable species meeting Criterion D2; see Appendix X) as the information
Brachycephalidae 8 0 0 0 0 1 1 1 5 1 13 necessary to appropriately apply these criteria is usually not available. For most species, it is
Bufonidae 476 5 1 85 71 52 26 171 65 214 45* considerably easier to observe a decline in the population, or to make an educated inference
Caeciliidae 113 0 0 1 1 1 0 41 69 3 3* of the range size, than it is to estimate the actual number of individuals.
Centrolenidae 138 0 0 6 16 29 10 28 49 51 37
Dendrobatidae 234 0 0 20 29 16 14 58 97 65 28
Discoglossidae 12 1 0 0 0 2 4 5 0 3 25 CONSERVATION STATUS ACROSS TAXA
Heleophrynidae 6 0 0 2 0 0 0 4 0 2 33
Hemisotidae 9 0 0 0 0 1 0 4 4 1 11 There are significant differences among the three taxonomic Orders of amphibians in the total
Hylidae 804 1 0 71 64 47 27 431 163 183 23* number of species and conservation status (Table 2). The Anura (frogs and toads), by virtue
Hyperoliidae 253 0 0 1 19 29 17 133 54 49 19* of absolute numbers of species, greatly influence the average threat level for all amphibians,
Leiopelmatidae 4 0 0 1 1 2 0 0 0 4 100 with 32% either globally threatened or extinct. The less species-rich Caudata (salamanders
Leptodactylidae 1,238 2 0 145 247 172 61 351 260 566 46* and newts) are significantly more threatened than the global average, with 47% of spe-
Limnodynastidae 50 0 0 1 7 2 1 37 2 10 20 cies currently threatened or extinct. The smallest Order of amphibians, the Gymnophiona
Mantellidae 158 0 0 7 12 16 12 77 34 35 22* (caecilians), remain extremely poorly known, with two-thirds (66%) of caecilians currently
Megophryidae 128 0 0 3 14 27 13 40 31 44 34 assessed as Data Deficient. Only 3% of caecilians are known to be threatened.
Microhylidae 430 0 0 6 27 39 18 177 163 72 17* Significant variations in both the total number of species and threat status are also
Myobatrachidae 71 1 0 6 2 4 3 49 6 13 18* evident between the 48 amphibian families (Table 2). Very diverse families of frogs and
Nasikabatrachidae 1 0 0 0 1 0 0 0 0 1 100 toads that are significantly more threatened than the global average include the Bufonidae,
Pelobatidae 4 0 0 0 1 0 1 2 0 1 25 Leptodactylidae, and Rhacophoridae. Sadly, both species of the extraordinary Australian
Pelodytidae 3 0 0 0 0 0 0 3 0 0 0 endemic family Rheobatrachidae (the gastric-brooding frogs) are now Extinct (see Essay 6.1).
Petropedetidae 102 0 0 3 13 8 10 39 29 24 24 Four small families are at severe risk of disappearing altogether; the Leiopelmatidae of New
Pipidae 30 0 0 1 2 0 1 21 5 3 10* Zealand, with all four species in the family threatened (see Essay 6.2); the Rhinodermatidae
Ranidae 666 2 0 19 61 82 60 295 147 164 25* of Chile and Argentina with two threatened species, one of which (Rhinoderma rufum) has
Rhacophoridae 274 18 0 17 50 33 26 64 66 118 43* not been recorded since the late 1970s; the Sooglossidae, all endemics of the Seychelles,
Rheobatrachidae 2 2 0 0 0 0 0 0 0 2 100 and the monotypic family Nasikabatrachidae. Diverse families that are less threatened than
Rhinodermatidae 2 0 0 1 0 1 0 0 0 2 100 the global average include Ranidae, Microhylidae, and Hyperoliidae. Among the three most
Rhinophrynidae 1 0 0 0 0 0 0 1 0 0 0 species-rich salamander families, Hynobiidae and Plethodontidae exhibit much higher levels
Scaphiopodidae 7 0 0 0 0 0 1 6 0 0 0 of threat than Salamandridae.
Sooglossidae 4 0 0 0 0 4 0 0 0 4 100
TOTAL ANURA 5,208 32 1 400 658 581 311 2,028 1,197 1,672 32
Caudata
Ambystomatidae 30 0 0 9 2 2 1 13 3 13 43 CONSERVATION STATUS ACROSS BIOGEOGRAPHIC
Amphiumidae 3 0 0 0 0 0 1 2 0 0 0 REALMS
Cryptobranchidae 3 0 0 1 0 0 2 0 0 1 33
Dicamptodontidae 4 0 0 0 0 0 1 3 0 0 0 Biogeographic realms are the eight continent-scale terrestrial and freshwater regions
Hynobiidae 46 0 0 5 10 12 2 11 6 27 59* distinguished by characteristic biota that reflect shared evolutionary histories (Udvardy
Plethodontidae 365 1 0 36 82 58 37 91 60 177 48* 1975). We adapted the biogeographic realm classification of Olson et al. (2001) (Figure 3)
Proteidae 6 0 0 0 1 1 1 3 0 2 33 to allocate species to one of six biogeographic realms10. These realms are broadly similar
Rhyacotritonidae 4 0 0 0 0 1 2 1 0 1 25 to the six zoogeographic realms employed by Duellman (1999) in his regional comparison
Salamandridae 70 1 0 3 14 12 11 27 2 30 43 of amphibian distributions (see Fig 1:1 in Duellman 1999).
Sirenidae 4 0 0 0 0 0 0 4 0 0 0 The Neotropical Realm, which stretches from central Mexico south throughout South
TOTAL CAUDATA 535 2 0 54 109 86 58 155 71 251 47* America and includes all of the Caribbean Islands, is by far the leader in terms of overall
Gymnophiona species richness of amphibians, being home to almost half of the world’s amphibians, of
Ichthyophiidae 39 0 0 0 0 2 0 5 32 2 5* which 96% are endemic to the realm (Table 3). Two other realms have even higher levels of
Rhinatrematidae 9 0 0 0 0 0 0 4 5 0 0 endemism, the Australasian (99%) and Afrotropical (98%) realms. The relatively low level
Scolecomorphidae 6 0 0 0 0 0 0 3 3 0 0 of endemism in the Palaearctic is mainly due to the somewhat arbitrary border between
Uraeotyphlidae 5 0 0 0 0 0 0 0 5 0 0 the Palaearctic and Indomalayan realms in China, and similarly the level of endemism in
TOTAL GYMNOPHIONA 172 0 0 1 1 3 0 53 114 5 3* the Nearctic is also reduced by the arbitrariness of the division of Mexico between the
Nearctic and Neotropical realms.
The Neotropical Realm also has the greatest proportion of threatened and extinct
Table 2. Red List Status by Order and Family8. *The difference between these percentages and the percentages for amphibians as a whole species, having experienced significant habitat loss, and is home to many narrow range
is statistically significant9 endemics, especially in the Andes and Mexico. It is also the region that has been hardest
hit by chytridiomycosis. Although there are many species that are threatened or extinct as
a result of habitat loss, disease and introduced species in Australia and New Zealand, the
contribution of the relatively unthreatened amphibians of the island of New Guinea results
in the overall percentage of threatened species in the Australasian Realm being the lowest
of all the regions, with only 13% and significantly less than the global average of 33%. The
Afrotropics, Nearctic, and the Palaearctic are also regions with significantly less threatened
species than the global average. In-depth syntheses of amphibian distributions in each of
the six biogeographic realms are presented in Chapters 5-10.

SPECIES RICHNESS AND ENDEMISM

Nearly a decade has passed since the last major synthesis on the global distribution
patterns of amphibians (Duellman 1999). Prior to Duellman’s (1999) synthesis, the only
major contributions in the last 50 years that have taken a global perspective on amphibian
distributions are those of Savage (1973) and Duellman and Trueb (1986). There have, of
course, been many treatises focusing on regional distribution patterns on continents, such
as Central America (Savage 1982), China (Zhao and Adler 1993) and Europe (Gasc 1997),
as well as island faunas, such as the Caribbean (Hedges 1996) and the Philippines (Inger
1954). However, Duellman’s publication was the first to attempt a major synthesis of the
global distribution patterns of amphibians, describing differences and similarities among
continents, major islands and archipelagos, and discussing the implications of these patterns
of distribution relative to conservation.

Figure 3. Biogeographic Realms adapted from Olson et al. (2001).

Realm Total (endemics) % endemics EX EW CR EN VU NT LC DD Number threatened % Threatened or EX

Afrotropical 969 (954) 98 0 0 33 107 100 53 467 209 240 25*
Australasian* 544 (538) 99 3 0 16 21 29 10 287 178 69 13*
Indomalayan 999 (800) 80 20 0 32 134 143 104 311 255 329 33
Nearctic 337 (230) 68 2 1 10 24 46 40 189 25 83 25*
Neotropical 2,916 (2,808) 96 7 0 358 456 324 140 956 675 1,145 39*
Table 3. Red List status in each biogeographic Palaearctic 468 (260) 56 2 0 13 40 58 48 245 62 113 24*
region. *includes Oceania
 Chapter 4. The State of the World’s Amphibians 35

Leptopelis vermiculatus (Vulnerable) is a range-restricted species endemic to several

mountain blocks in the Eastern Arc chain of Tanzania, including the East and West Usam-
baras, Nguu, Udzungwa, Poroto and Mount Rungwe. Around two-thirds of amphibians have
distribution ranges less than 50,000km2. © Maik Dobiey

Figure 4. Global species richness for amphibians, with dark red colours corresponding to higher richness. Colour scale based on 10 quantile
One of the major advances of the GAA process is the collection of high resolution spatial classes. Maximum richness equals 144 species.
data that allow us to undertake analyses that expand considerably upon the work and
conclusions of these aforementioned earlier assessments, making comparisons possible
across both broad geographic regions and political units. Importantly, we can make these
comparisons across threatened species. Such an analysis of extent of occurrence data has
not previously been possible for amphibians beyond that of a single continent (e.g., Brooks
et al. 2001). It is also possible to compare these maps to human demographic data to gain
a better understanding of the challenges faced by amphibians in an increasingly human
dominated world (See Essay 4.2).

GEOGRAPHIC PATTERNS OF SPECIES RICHNESS

AND ENDEMISM
The most obvious pattern to emerge from an analysis of the overlay of all species distribu-
tions is visual evidence that the tropics hold much higher species richness than do temper-
ate regions (Figure 4). This is further supported by a summary of the number of species in
each five-degree latitudinal band, which shows that species richness is much higher in the
tropics than is to be expected based on area alone, and peaks around the equator (Baillie
et al. 2004; Mace et al. 2005).
Areas of high global species richness among amphibians include Central America, the
Andes and the Amazon Basin, the Atlantic forest of Brazil, the Upper Guinea forests of
West Africa, the Afromontane regions of Africa (including the Albertine Rift, the highlands
of Cameroon, and the Eastern Arc Mountains), eastern Madagascar, the Western Ghats,
and montane Southeast Asia. The only non-tropical area with high species richness is the
south-eastern United States, which is the global centre for salamander diversity. Species
richness is lowest in arid regions, and there are no amphibians at all in the high latitudes Figure 5. Richness map of threatened amphibian species, with dark red colours corresponding to higher richness. Colour scale based on 10
of the northern hemisphere, central Asia, North Africa, south-central Australia, Antarctica, quantile classes. Maximum richness equals 44 species.
and most of New Zealand. Of course, this map must be interpreted with some caution
when considering the problem of uneven survey efforts around the world; regions such as
Indonesia, New Guinea and the Congo Basin are especially likely to be under represented
on this map due to lack of adequate surveys, although the overall pattern of species richness
decreasing away from the poles holds true. Considering existing maps of species richness
among mammals and birds, there is an obvious degree of congruence in patterns of overall Order All species minus DD, EX and EW All threatened species
species richness among these groups (Baillie et al. 2004; Mace et al. 2005). Median area (km2) Number of species Median area (km2) Number of species
The global distribution of threatened amphibians (Figure 5) differs significantly from the Anura 19,118 3,978 1,090 1,639
map of overall species diversity. Major concentrations of threatened species are associated Caudata 4,378 462 417 249
with insular systems (montane environments, islands), where there has been severe habitat Gymnophiona 33,224 58 44 5
loss and exposure to the fungal disease chytridiomycosis, including the sierras and trans- All species 16,794 4,498 976 1,893
volcanic belt of Mexico, Central America, the tropical Andes, Brazil’s Atlantic Forest, the
Greater Antilles, the Upper Guinea forests of West Africa, the forest of western Cameroon
and eastern Nigeria, the Albertine Rift, the Eastern Arc Mountains of Tanzania and other Table 4. Median range size of species in each amphibian Order. Only extant non-Data Deficient species with a range map are included in
Afromontane outliers, the south-east coast of South Africa, Madagascar’s wet forests, the the analysis11.
Western Ghats and Sri Lanka, montane Southeast Asia, and eastern Australia. South and
East China also emerge quite strongly, although the pattern is driven to a large degree by
the formerly widespread, but now Critically Endangered, Chinese Giant Salamander.
In contrast then to published richness maps of threatened mammals (Baillie et al. 2004;
Grenyer et al. 2006) or birds (Baillie et al. 2004; BirdLife International 2004; Orme et al.
2005; Grenyer et al. 2006), most of the world is devoid of threatened amphibians. It has
already been established that there is a widespread correlation between a species’ range
size and extinction risk (Purvis et al. 2000), and geographic range is inherent in the Red List
criteria (see Appendix I). Since amphibians are largely sedentary species, their ranges tend
to be much smaller than more mobile species, such as birds (Baillie et al. 2004). Threatened
amphibians are more likely to have small range sizes (Table 4) and tend to occur more densely
in smaller areas than either mammals or birds.
Not only do most amphibians have small ranges, but these narrowly distributed species
tend to co-occur in ‘centres of endemism’ (Anderson 1994). These centres of endemism tend
to lie across isolated or topographically varied regions, and show extreme concentration in
the tropics (Figure 6). This is the geographical manifestation of ‘Rapoport’s rule’ (Rapoport
1982), which states that the mean latitude of a species’ range correlates with the species’
range size, although the generality of this ‘rule’ has been questioned (Gaston 1999). Previous
analyses have already demonstrated the remarkable degree of overlap between centres of
endemism across birds, mammals and amphibians (Baillie et al. 2004).

SPECIES RICHNESS AND ENDEMISM WITHIN

COUNTRIES
An important contribution of any analysis of geographic patterns of amphibian distributions Figure 6. Centres of endemism inhabited, respectively, by more than two overlapping species with global distributions of less than
is reporting on the distribution of species by political units, specifically countries. Particularly 50,000km² 12.
36 Threatened Amphibians of the World

Brazil Jamaica Colombia Haiti

Dominican
Colombia Seychelles Mexico Republic
Ecuador Madagascar Ecuador Jamaica

Peru Cuba Brazil Cuba

Mexico Australia China Puerto Rico

Sri Lanka
Indonesia Sri Lanka Peru
Guatemala
China Japan Guatemala
Mexico
Venezuela Philippines Venezuela
United States Seychelles
of America Puerto Rico India
Philippines
Papua New United States Costa Rica
Guinea of America
Honduras
India Chile Honduras
Ecuador
Madagascar Mexico Madagascar
Chile
Australia Papua New Panama
Guinea Japan
Congo, D.R. Brazil Cameroon
Turkey
Bolivia India Sri Lanka
Costa Rica
Malaysia China United States
of America Colombia
Camerooon Haiti Philippines Panama
Panama Venezuela Australia El Salvador
Costa Rica Colombia Cuba Taiwan, Province
of China
Tanzania Taiwan, Province Malaysia Trinidad
of China and Tobago
0 200 400 600 800 1000 40 50 60 70 80 90 100 0 50 100 150 200 250 0 20 40 60 80 100
Number of species Percentage of endemic species Number of threatened species Percentage of species threatened
Endemics Non-endemics Endemics Non-endemics
Figure 7 Figure 9 Figure 10 Figure 12

Figure 7. The 20 most diverse countries for

amphibians.
Figure 9. The 20 countries with the highest
percentage of country endemics. Countries
with less than 10 native species are excluded
(NB. The totals for China include species in
Hong Kong and Macau).
Figure 10. The 20 countries with the highest
number of threatened amphibians.
Figure 12. The 20 countries with the highest
percentage of threatened amphibians. Only
countries with 10 or more native amphibians
are included.
informative are the numbers of species endemic to each country, and especially the numbers
of endemic threatened species, because they can guide a “doctrine of ultimate responsibil-
ity” for each nation to ensure the persistence of global biodiversity (Mittermeier et al. 1998).
Although the number of threatened species per country provides a coarse measure of threats
to biodiversity, albeit heavily conflated by area (Balmford and Long 1995), dividing the number
of threatened species by total numbers of species per country does give a useful measure of
relative threat to a nation’s biodiversity. Further, the presentation of threatened species occur-
rences by country is also useful in providing a cross-check to national Red Lists, and vice versa
(Hilton-Taylor et al. 2000; Rodriguez et al. 2000), given the important implications of these for
national conservation policy. A detailed summary for all countries is provided in Appendix V.
When comparing amphibian richness amongst countries, Brazil, with 751 species currently
recognized, is first, followed closely by Colombia with 697 (Figure 7). These results need to be
considered with some caution, particularly bearing in mind the relative level of survey effort
in some countries. For example, both Colombia and Brazil have received extensive survey
efforts in recent decades, and although both countries can be expected to add significantly
to their totals (see Essay 9.5 and Essay 9.4, respectively), the level of increase is likely to
be less than in some of the other highly diverse countries such as Papua New Guinea or
Indonesia. In South America, Peru in particular is relatively poorly sampled and is almost
certain to experience substantial increases in its number of recorded species, and could
even surpass Ecuador. Nonetheless, the species richness in Ecuador is remarkable for a
country one-thirtieth the size of Brazil. Figure 8 provides an alternative illustration of the
relative significance of each country in terms of amphibian diversity.
The number and percentage of endemics within a country (Table 5 and Figure 9, respec-
tively), gives a slightly different perspective on the importance of countries in amphibian
conservation. When looking at the number of country endemics, the first two countries,
Brazil and Colombia, not surprisingly emerge yet again at the top of the list. Interestingly,
Ecuador with the third largest number of amphibians is only 11th in the list of percentage of
endemics, with only 36% of species endemic. To put some perspective on the contribution
these countries make to global amphibian species richness, the endemic species of the top
10 countries account for 40% of the world’s amphibian species, and the top 20 for 56%.
When considering the percentage of country endemics, it is perhaps not surprising to see
that the first nine countries listed are island states. Many of these have very small numbers
of species, with the exception of Madagascar with 226 species currently described, only one
of which is not endemic to the country. Similarly, Australia with a total of 214 species has
200 country endemics. The two countries with the highest number of endemics, Brazil and
Colombia, are only ranked 18th and 24th when looking at percentage of endemic species.
Among countries with the highest number of threatened amphibians (Figure 10), there are
some apparent differences to the most species-rich countries listed in Figure 7, suggesting
that either amphibians in some countries are more susceptible to threats, or that threats vary
between countries, or that there are other factors influencing the distribution of threatened
species. Figure 11 illustrates the relative importance of each country to threatened amphib-
ians. Colombia, the country with the second largest amphibian fauna, has the most threatened
species. The major threat to amphibians in Colombia is habitat loss, although there have also
been many as yet unexplained declines occurring (presumably due to chytridiomycosis). The
dramatic topography of the Andes means that many of the amphibians have very restricted
ranges thus making them more vulnerable to threatening processes.
Ranking of countries with the highest percentage of threatened species (Figure 12 is quite
different to the list of countries with the highest number of threatened species (Figure 10).
The first five countries13 with the highest percentage are all in the Caribbean, and over 70%
of all the amphibians in these countries are threatened. Compared with other regions, the
Caribbean region stands out with by far the highest percentage of threatened species. Most
notable in this regard is Haiti, with 46 of the 50 species native to this country threatened
with extinction, almost entirely due to rampant habitat loss.
In Mexico, a country ranked fifth for species richness, but second in terms of number of
threatened species, more than 50% of amphibians are threatened. Severe habitat loss is the
overwhelming threat to Mexican amphibians, but chytridiomycosis is also present.

Figure 8. The world according to amphib-

ians. The size of each country represents the
relative proportion of the world’s amphib-
ians found there. Map produced by Vineet
Katariya.
 Chapter 4. The State of the World’s Amphibians 37

Table 5. The 20 countries with the highest

Ranking Country Native Native Endemic % endemics Number threatened Number threatened % Species number of country endemics.
endemics Threatened
1 Brazil 751 489 65 110 106 15
2 Colombia 697 337 48 209 159 30
3 Mexico 363 246 68 198 162 55
4 Madagascar 226 225 99.6 55 55 24
5 Australia 214 200 93 47 47 22
6 United States of America 261 182 70 52 48 20
7 Peru 411 181 44 81 59 20
8 China 325 170 52 87 82 27
9 Papua New Guinea 244 164 67 10 8 4
10 Indonesia 347 161 46 33 23 10
11 Ecuador 447 159 36 163 102 36
12 Venezuela 298 155 52 69 62 23
13 India 236 151 64 63 61 27
14 Philippines 98 77 79 48 48 49
15 Sri Lanka 83 67 81 52 51 63
16 Tanzania 162 65 40 41 40 25
17 Cuba 59 57 97 47 47 80
18 Malaysia 203 56 28 47 34 23
19 Cameroon 196 55 28 53 38 27
=20 Bolivia 209 53 25 21 18 10
=20 Congo, Democratic Republic of the 211 53 25 13 4 6

HABITAT AND ECOLOGY

Habitat Preferences
Recording the habitat preferences of species from a standardized classification scheme (see
documentation on CD) is part of the required documentation in the Red List assessment
process. Almost 80% of amphibians are forest species (Table 6 and Figure 13) and 90% of
these species (4,251 species), in turn, occur in tropical forests. Of all forest species, 36%
are either threatened or extinct, and for those occurring in tropical forests the percentage
of threatened or extinct species is slightly higher at 37%. However, montane forest spe-
cies (very generally defined here as forest at altitudes above 1,000m asl) are much more
likely to be threatened than those occurring in lowland forest. This is probably because
montane species are more likely to have restricted ranges and hence are more susceptible
to threatening processes, and also because chytridiomycosis appears to be having a much
greater impact on species occurring at elevations above 1,000m asl.
Only a small percentage of amphibians occur in relatively dry habitats such as arid
and semi-arid habitats and savannah, 2% and 8%, respectively, and the percentages of
species in these habitats that are threatened are very low (9% and 3%, respectively). The
lack of species in these habitats is not surprising as amphibians are known to prefer moist
habitats. These drier habitats are also usually less varied topographically and are relatively
less impacted by anthropogenic disturbance, so it is to be expected that there will also be
fewer threatened species.
Although amphibians are classically considered as leading ‘dual lives’, occurring
in both land and water environments (hence the etymological origins of the word), in
fact only two-thirds of species (3,775) undergo a free-swimming larval development
stage in which they are dependent on a freshwater habitat. Those species that are
not dependent on freshwater habitats are predominantly direct-developing species,
which develop directly from eggs without a larval stage (see Chapter 1). It is the larval
stage of larval-developing species that most likely occurs within freshwater habitats. While almost 80% of amphibians are forest
In Table 6, occurrence in freshwater is divided into three subgroups: still, flowing, or species, only two-thirds undergo a free-swim-
swamp/marsh. Flowing freshwater habitats for amphibians usually include streams, ming larval development stage in which they
creeks or brooks; still freshwater habitats are often temporary rainpools or other small are dependent on some freshwater habitat
pools of freshwater (favoured breeding sites for many explosive-breeding species). (like Atelopus varius pictured here). Species
This distinction between freshwater habitats appears to have a significant influence that are not dependent on freshwater habitats
on the likelihood that a species is threatened. Of species that are associated with are predominantly direct-developing species,
flowing water, 36% are currently threatened or extinct, whereas only 15% of species which develop directly from eggs without a
associated with still water are currently threatened or extinct. This may be related to free-living larval stage. © Twan Leenders

Figure 11. The world according to threatened

amphibians. The size of each country repre-
sents the relative proportion of the world’s
threatened amphibians found there. Map
produced by Vineet Katariya.
38 Threatened Amphibians of the World

Table 6. The habitat preferences for all

amphibian species. “All tropical forest” Habitat type All species % of all Threatened % of all
encompasses both lowland and montane species or Extinct species in
species habitat that Number of species % of species in habitat that are threatened
tropical forest. are threatened Forest
or Extinct
Savannah
Forest 4,710 80 1,678 36 Grassland
All tropical forest 4,251 72 1,566 37
Shrubland
Lowland tropical forest 2,849 48 807 28
Montane tropical forest 2,507 42 1,142 46 Secondary terrestrial habitats
Savannah 484 8 13 3 Flowing freshwater
Grassland 987 17 217 22
Marsh/swamp
Shrubland 860 15 148 17
Still open freshwater
Secondary terrestrial habitats 1,496 25 258 17
Figure 13. The number of species occurring Flowing freshwater 2,506 42 897 36 Arid and semi-arid habitats
in a particular habitat, and the percentage of Marsh/swamp 722 12 113 16
0 1,500 3,000 4,500 0 10 20 30
species occurring within each habitat that Still open freshwater 1,793 30 275 15
are threatened. Arid and semi-arid habitats 94 2 8 9

the likelihood of exposure to disease increasing in flowing water, as the water may be southern Mexico, the Atlantic Forest of Brazil, New Guinea, Sri Lanka (see Essay 4.3), the
acting as a vector for disease. mountainous regions of the eastern United States, and the west coast of the United States.
Lesser concentrations are also found in east and central Africa, Madagascar, the Western
Reproductive Modes Ghats and the Philippines. The number and overall range of direct-developing species in
the Old World tropics is still greatly under-estimated, especially because large numbers of
As noted earlier (see Chapter 1), for the purposes of analysis in this book, amphibian species in the genera Philautus (Indomalayan) and Arthroleptis (Afrotropical) probably remain
reproductive modes have been combined into three general groups: larval developing to be described. Live-bearing species are only found in a few places: the Amazon, Central
– species that lay eggs, from which free-living larvae hatch; direct developing – species America, Europe, West Africa, the Eastern Arc Mountains and West Asia.
that lay eggs in which the larvae develop and metamorphose prior to hatching; and live- A map of the global distribution of all three reproductive modes (Figure 15) shows that
bearing – species in which the larvae develop inside the mother, with no eggs being laid the majority of the global range of amphibians is home only to larval-developing species.
(including viviparous and ovoviviparous species). By far the most common reproductive This is perhaps why most people think of amphibians as being freshwater, larval-developing
mode is larval development, with 4,004 species known or believed to use this mode. species only, and are often surprised to hear of the existence of other remarkable reproduc-
Direct development is also quite common, being present or strongly suspected in 1,742
species, whereas there are only 58 known or suspected live-bearing species. There are
111 species for which the reproductive mode remains completely unknown and cannot
readily be inferred from close relatives.
The global distribution of larval-developing, direct-developing, and live-bearing Family Larval-developing Direct-developing Live-bearing Unknown
amphibians is shown in Figure 14. The richness map of larval developers is very similar Anura
to the richness map for all amphibian species (Figure 4), which is not surprising consider- Allophrynidae 1 0 0 0
ing that over two-thirds of amphibians are larval developers. The few regions that differ Arthroleptidae 16 35 0 0
significantly are the Caribbean; a region extending from southern Mexico south to Ecuador Ascaphidae 2 0 0 0
and Venezuela; New Guinea; and the mountains of the eastern United States, which have Astylosternidae 29 0 0 0
very few larval-developing species compared with overall species richness. Compared with Bombinatoridae 8 0 0 2
larval developers, direct developers occur in relatively few areas. The main concentrations Brachycephalidae 0 8 0 0
are in the Caribbean, the Andes of South America extending through Central America to Bufonidae 426 24 13 13
Centrolenidae 138 0 0 0
Dendrobatidae 232 2 0 0
Discoglossidae 12 0 0 0
Heleophrynidae 6 0 0 0
Hemisotidae 9 0 0 0
Hylidae 800 0 0 4
Hyperoliidae 250 1 0 2
Leiopelmatidae 0 4 0 0
Leptodactylidae 394 836 1 7
Limnodynastidae 50 0 0 0
Mantellidae 125 20 0 13
Megophryidae 128 0 0 0
Microhylidae 192 233 0 5
Myobatrachidae 63 8 0 0
Nasikabatrachidae 1 0 0 0
a. Pelobatidae 4 0 0 0
Pelodytidae 3 0 0 0
Petropedetidae 90 10 0 2
Pipidae 25 3 0 2
Ranidae 585 80 0 1
Rhacophoridae 138 130 0 6
Rheobatrachidae 0 2 0 0
Rhinodermatidae 1 1 0 0
Rhinophrynidae 1 0 0 0
Scaphiopodidae 7 0 0 0
Sooglossidae 1 3 0 0
TOTAL ANURA 3,737 1,400 14 57
Caudata
Ambystomatidae 30 0 0 0
Amphiumidae 3 0 0 0
Cryptobranchidae 3 0 0 0
Dicamptodontidae 4 0 0 0
b. Hynobiidae 45 0 0 1
Plethodontidae 50 315 0 0
Proteidae 6 0 0 0
Rhyacotritonidae 4 0 0 0
Salamandridae 57 0 13 0
Sirenidae 4 0 0 0
TOTAL CAUDATA 206 315 13 1
Gymnophiona
Caeciliidae 8 27 28 50
Ichthyophiidae 39 0 0 0
Figure 14. Richness map of a) larval-de- Rhinatrematidae 9 0 0 0
veloping amphibians; b) direct-developing Scolecomorphidae 0 0 3 3
amphibians; and c) live-bearing amphibians. Uraeotyphlidae 5 0 0 0
Darker red colours correspond to regions TOTAL GYMNOPHIONA 61 27 31 53
of higher richness. Colour scale based on TOTAL ALL AMPHIBIANS 4,004 1,742 58 111
10, 10 and 5 quantile classes, respectively.
Maximum richness equals 127, 55 and 9 spe-
cies, respectively. c. Table 7. Variation in reproductive mode between taxonomic Orders and Families.
 Chapter 4. The State of the World’s Amphibians 39

Figure 15. Combined distribution of amphib-

ian reproductive modes.

Breeding Strategy
Larval developers only
Direct developers only
Larval and direct developers
Larval and live-bearing
Larval, direct and live-bearing

tive modes. Live-bearing species always occur sympatrically with larval-developing species,
and sometimes with direct-developing species. The regions where all three reproductive
modes co-exist are the Amazon Basin, Colombia to southern Mexico, West and Central
Africa, East Africa and north-central Italy and south-eastern France. Direct developers are
the only amphibians in New Zealand, Fiji, the Lesser Antilles, the Bahamas, Palau, the
Solomon Islands and various small islands off New Guinea, but otherwise are always found
in areas inhabited by larval developers.
The prevalence of the various reproductive modes varies between the different taxonomic
Orders (Table 7). All three Orders have species representative of each reproductive mode.
The most common strategy for frogs and toads is larval development (over 70% of species),
and the most common strategy for salamanders and newts is direct development (59%).
While the most common strategy for caecilians is larval development (35% of species), the
majority (54%) of live-bearing species are caecilians. There is still very little known about
most caecilian species, and the reproductive mode for nearly one-third (30%) of caecilians
remains unknown.
Larval development is a very common reproductive mode amongst amphibian families,
and all but four families have some larval-developing species. The three anuran families,
Brachycephalidae, Rheobatrachidae and Leiopelmatidae, contain direct-developing species
only, and the caecilian family Scolecomorphidae has three live-bearing species in one genus,
and the reproductive mode for the three species in the other genus are unknown. Compared
with larval development, direct development and live-bearing are reproductive modes
restricted to far fewer families. Only 19 families have species that are direct developers,
and the overwhelming majority of these are within the anuran family Leptodactylidae, and
in particular the largest amphibian genus Eleutherodactylus. Amongst other anuran families,
slightly more than half of the microhylid species, and almost half of the rhacophorid species
are direct developers. There is only one salamander family containing direct-developing
species, Plethodontidae, and only one caecilian family, Caecilidae.
Within the Caudata there are 13 live-bearing species all in the genera Salamandra and
Lyciasalamandra. There appear to be few live-bearing frog and toad species (14 species in
total), relative to the diversity of this Order. Most of these species, such as the Critically
Endangered Kihansi Spray Toad Nectophrynoides asperginis and Golden Coqui Eleuthero-
dactylus jasperi are ovoviviparous, with the equally highly threatened Nimbaphrynoides
liberiensis and Nimbaphrynoides occidentalis being the only truly viviparous anurans.
The reproductive mode of a particular species is related to whether or not it is likely
to be threatened with extinction (Table 8). Amongst direct developers, 45% of species are
currently threatened with extinction or are already extinct, which is considerably higher than
the global average of 33%. Twenty-one of the 34 extinct amphibians were direct developers.
Larval developers are relatively less threatened than the global average.
The median range size of amphibians employing each of the three different reproductive
modes is summarized in Table 9. When considering all extant, non-Data Deficient species,
the median for larval-developing species is much greater than that of direct-developing
species. The much smaller range size of direct developers makes these species more
susceptible to habitat loss and other threats compared to larval developers. This might
partly explain why direct-developing amphibians are proportionately more threatened than
larval-developing amphibians.
Major Threats
As part of the Global Amphibian Assessment, the major threats to all amphibians were
collated. Analyses of these data permit a quantitative perspective on the major causal
mechanisms not only driving amphibians to extinction, but also allowing us some insight
into the threats that are driving amphibians most rapidly to extinction.
Given the major finding of the Millennium Ecosystem Assessment that humans have
changed ecosystems more rapidly and extensively in the last 50 years than in any comparable
period of time in human history, it is not surprising that habitat loss remains the number
one threat to amphibians globally, with two-thirds (63%) of species affected, of which
almost nine out of every 10 species (87%) are threatened with extinction (Table 10; Figure
16). These figures for threatened amphibians are comparable with those for threatened
birds, where habitat loss and degradation affects 86% of species (BirdLife International
2004). These results take on more meaning when one considers that nearly three-quarters
of amphibians are inhabitants of tropical forest environments – the same habitats that
are subject to the highest rates of forest loss. The most common form of habitat loss is
clearance for crops, followed very closely by extractive processes, such as logging (be it
for subsistence purposes or due to selective and/or clear-cutting15), and urbanization and
industrial development (Table 10).
But just how high are rates of forest loss? In Indonesia, the vast majority of forest loss
has occurred over the last 30 years or so, mainly the result of commercial logging, followed
by massive agricultural projects (particularly rubber, oil palm, industrial timber for pulp pro-
duction, and so forth) and smallholder agriculture. One recent estimate alone indicates that
Kalimantan’s protected lowland forests declined by 56% between 1985 and 2001 primarily
due to logging (Curran et al. 2004). Another study combining high-quality remote sensing
applications and extensive field surveys across an area of 1.17 million ha in south-west
Sumatra determined that the single largest contiguous area of forest standing across their
study area had been nearly halved in the space of three decades (692,850 ha to 344,409
ha between 1972 and 2002), at an average rate per original forest cover of 1.69% per year
(Gaveau et al. 2007). In Madagascar, rates of forest loss have been estimated at slightly
less than 1% per annum (Harper et al. 2007).
One of the key consequences of habitat loss is habitat fragmentation. Decreasing
patch size and increasing isolation of patches results in an increased risk of demographic,
stochastic and genetic events, thereby increasing extinction risk by reducing demographic
and genetic input from immigrants (not to mention reducing the chance of recolonization
after local extinction) (Cushman 2006). Connectivity is therefore important in the regional
viability of amphibian populations, being predominantly affected by means of juvenile
dispersal. However, in fragmented landscapes, the survival of dispersers is often lower than
required for population viability, and even species with limited dispersal abilities are likely
to be imperilled by habitat loss and fragmentation over longer time periods (see Cushman

Reproductive mode All species EX EW CR EN VU NT LC DD Number threatened or EX % Threatened or EX

Larval-developing 4,004 11 1 266 412 410 262 1,775 867 1,100 27
Direct-developing 1,742 23 0 176 337 249 106 418 433 785 45
Live-bearing 58 0 0 7 9 4 1 18 19 20 34
Unknown 111 0 0 6 10 7 0 25 63 23 21 Table 8. The Red List status of species with
varying reproductive modes.

Reproductive mode All species (minus DD, EX and EW) All threatened species
Median area (km2) Number of species Median area (km2) Number of species
Larval development 41,999 3,125 1,671 1,088
Direct development 2,272 1,286 454 762
Live-bearing 1,755 39 287 20 Table 9. Median range size for the different
Not known 12,264 48 713 23 reproductive modes employed by amphibians.
Total 16,794 4,498 976 1,893 Only extant non-Data Deficient species with a
range map are included in the analysis14.
40 Threatened Amphibians of the World

Species of the genus Lyciasalamandra, such as
L. helverseni (Vulnerable) from the islands of
Karpathos and Kasos in Greece, are among the
only live-bearing salamanders, giving birth to
fully developed young. © Jan van der Voort
2006 for review). Among those amphibians currently listed as threatened, 45% are affected
by severe fragmentation of their habitat, particularly in regions such as the Western Ghats
and Sri Lanka, the Philippines, the coastal forests of Eastern Africa, and the Atlantic Forest
of Brazil (C. Boyd et al. unpubl.).
The second most significant threat to amphibians is pollution, affecting around one-fifth
(19%) of species, and 29% of threatened species. These percentages are much higher than
those recorded for mammals or birds (4% and 12%, respectively; Baillie et al. 2004), probably
due to the fact that a high proportion of amphibians are aquatic. Among the over-riding
causes of increasing pollution is the increasing use of agrochemicals (fertilizers, herbicides,
fungicides, pesticides), especially in developing countries; other types of pollution include
the discharge of industrial wastes into waterways, acid precipitation, and atmospheric
pollution that results in thinning of the ozone layer and increased ultra-violet (UV) radia-
tion. Most studies investigating the effects of pollutants and contaminants on amphibians
have focused on physiology (e.g., disruption of the endocrine system; Hayes et al. 2002),
behaviour (e.g., changes in activity patterns; Bridges 1997), and morphology (specifically,
morphological anomalies, such as ectromelia, apody or syndactyly; Oullet et al. 199716).
Although a foreboding body of evidence suggests that pollutants and contaminants can
compromise amphibian communities (see Essay 4.4), the relative contribution of contami-
nants to declines remains unclear. However, at least one recent study has demonstrated a
correlation between population declines and chemicals carried by winds from agriculture
in California (Davidson et al. 2002)17.
Although disease appears to be a relatively less significant threat for amphibians, for
those species affected it can cause sudden and dramatic population declines and disap-
pearances (see Essay 4.5). That disease is operating in synergy with global climate change
is now also strongly supported, with evidence suggesting that recent trends toward warmer
nights and increased daytime cloud cover have created optimum conditions for the chytrid
fungus, Batrachochytrium dendrobatidis, at mid-elevation sites (1,000-2,400m asl), which
is where the preponderance of amphibian extinctions and disappearances have occurred
(Pounds et al. 2006; and see Essay 4.6). In comparison, although habitat loss and degrada-
All habitat loss
tion affect a much greater number of species, the rate at which species decline is usually
much slower, and there are a number of strategies, such as the establishment of protected
Invasive species areas, available to counter this threat.
Utilization

Accidental mortality

Persecution
MAJOR THREATS ACROSS TAXA
Pollution The significance of a threatening process is influenced by a species’ ability to adapt to this
Natural disasters process. Some species are more able to adapt to certain threats than others; for example, species
that are able to persist, or even thrive, in degraded habitats are much less affected by habitat loss
Disease
than species that breed only in bromeliads. Comparing major threats to threatened species across
Human disturbance taxonomic groups (Table 11) can potentially provide some insight into how susceptible particular
Changes in native sp. dynamics groups of species are to certain threatening processes. Caution is needed in interpreting these
Fire
results, given the uneven global distribution of both threats and biodiversity. Hence, it is not
always possible to separate whether a particular taxonomic group is more or less susceptible
0 1,000 2,000 3,000 4,000 to a threatening process, or whether exposure to threats is influencing these figures.
Number of species Across all the Orders, habitat loss is still overwhelmingly the most significant threat.
Figure 16. The major threats to amphibians Threatened species Non-threatened species For frogs and toads, and salamanders and newts, habitat loss affects around three times
globally. more species than the next most significant threat, pollution. Disease is the third most
significant threat to frogs and toads, but only seventh in level of significance for salamanders
and newts. Disease is almost three times more prevalent amongst threatened frogs and
toads than salamanders and newts. To date, there have been no recorded incidences of
Major Threat All species % of all species Threatened species % of Threatened disease amongst caecilians.
species Looking at other major threats, utilization affects proportionally three times more
All habitat loss 3,700 63 1,655 87 threatened salamanders and newts, than frogs and toads, suggesting that salamanders
Agriculture – Crops 2,588 44 1,227 65 and newts appear to be relatively more susceptible to utilization (particularly for human
Agriculture – Tree Plantations 508 9 230 12 consumption and for the international pet trade). Salamanders and newts on average have a
Agriculture – Livestock 1,376 23 620 33 much longer generation length than the frogs and toads; in the Critically Endangered Chinese
Timber and other vegetation removal 2,529 43 1,169 62 Giant Salamander, a species that has been subject to extensive exploitation (see Essay 4.7),
Urbanization and industrial development 2,106 36 945 50 it takes two months for the eggs to hatch after fertilization. Hence the effect of removing an
Invasive species 456 8 207 11 individual from a wild population is likely to have a more significant impact.
Utilization 250 4 105 6 When comparing the different types of habitat loss affecting threatened amphibians, frogs
Accidental mortality 47 1 22 1 and toads are most significantly impacted by habitat loss for agricultural crops, followed
Persecution 3 0.05 1 0.05 closely by vegetation removal (mostly logging) and infrastructure development (mainly for
Pollution 1,122 19 544 29 human settlement) (Table 11). The relative significance of different types of habitat loss is
Natural disasters 220 4 128 7 slightly different for salamanders and newts, with vegetation removal affecting the most
Disease 451 8 362 19 species, followed closely by infrastructure development and agricultural crops.
Human disturbance 243 4 151 8 By comparing major threats to threatened species across taxonomic families it is possible
Changes in native species dynamics (excluding disease) 16 0.3 4 0.2 to get a finer scale understanding of the variation in the significance of different threats to
Fire 615 10 290 15 species (Table 12). Such an analysis can also assist species conservation by highlighting
groups of species which are more or less likely to be impacted by particular threats. For
example, 36% of threatened bufonids are affected by disease, compared with only 19% of
Table 10. The major threats to amphibians globally. Only present threats to species are tallied. “All species” excludes Extinct species. all threatened species. It can be predicted that bufonids in any newly infected region would
therefore have a high probability of being impacted by the disease.
Not surprisingly, across almost all families habitat loss affects the greatest proportion
of species. The only exceptions are the families Caeciliidae, Leiopelmatidae and Sooglos-
Major Threat Anura % of Anurans Caudata % of Caudata Gymnophiona % of Gymnophiona sidae. Two of the three threatened species in the family Caeciliidae, and all of the species
All habitat loss 1,440 88 212 85 3 60 in the Sooglossidae, are endemic to the Seychelles Islands and are listed on the basis of
Agriculture – Crops 1,092 67 132 53 3 60 their restricted range with a major threat being invasive species. Similarly, the four spe-
Agriculture – Tree Plantations 209 13 21 8 0 0 cies within the Leiopelmatidae are all endemic to New Zealand and are also threatened
Agriculture – Livestock 560 34 60 24 0 0 by invasive species. When looking at the families most affected by invasive species, it is
Timber and other vegetation removal 1,013 62 155 62 1 20 island endemics (for example, the Mantellidae of Madagascar and the leptodactylids of
Urbanization and industrial development 799 49 146 59 0 0 the Caribbean) that are most affected. While many threatened amphibians are impacted by
Invasive species 186 11 19 8 2 40 invasive alien species, some amphibians themselves can also be invasive, and in so doing
Utilization 72 4 33 13 0 0 become a threat in their own right (see Essay 4.8).
Accidental mortality 18 1 4 2 0 0 Pollution, the second most significant threat to all amphibians, also affects a significant
Persecution 0 0 1 0.4 0 0 number of species in almost all families, with no families in particular appearing to be more
Pollution 479 29 62 25 3 60 susceptible to this threat than others.
Natural disasters 97 6 30 12 1 20 The incidence of disease, the third most significant threat to all threatened species, does
Disease 343 21 19 8 0 0 seem to vary across families. Disease is only recorded in 15 of the 36 families containing
Human disturbance 129 8 22 9 0 0 threatened species, and in particular there are 10 families containing a higher proportion
Changes in native species dynamics 3 0.2 1 0.4 0 0 of species affected than the global average of 19%. These families are Leiopelmatidae
(excluding disease) (100%), Rhinodermatidae (100%), Discoglossidae (50%), Myobatrachidae (58%), Hylidae
Fire 220 13 68 27 2 40 (51%), Bufonidae (36%), Dendrobatidae (28%), Leptodactylidae (21%), Centrolenidae (20%),
and Limnodynastidae (20%). Of the 362 threatened species affected by disease, 286 (79%)
of them are found in just three of these families: Leptodactylidae (118 species), Hylidae (93
Table 11. The major threats to threatened amphibians globally in each amphibian Order. Only present threats to species are tallied. species), and Bufonidae (75 species).
 Chapter 4. The State of the World’s Amphibians 41

Family Number of All habitat loss Invasive species Utilization Accidental Persecution Pollution Natural Disease Human Changes in native Fire
threatened mortality disasters disturbance sp. dynamics
species in family (ex. disease)
Anura
Arthroleptidae 14 13 0 0 0 0 1 0 0 0 0 1
Astylosternidae 21 21 0 1 0 0 4 0 0 0 0 1
Bombinatoridae 5 5 0 0 5 0 3 1 0 1 0 0
Brachycephalidae 1 1 0 0 0 0 0 0 0 0 0 0
Bufonidae 208 167 21 3 0 0 79 13 75 14 1 31
Centrolenidae 51 48 8 0 0 0 29 4 10 4 0 4
Dendrobatidae 65 57 12 9 0 0 27 3 18 1 0 4
Discoglossidae 2 2 1 0 0 0 0 1 1 0 0 0
Heleophrynidae 2 2 2 0 0 0 1 0 0 1 0 2
Hemisotidae 1 1 1 0 0 0 0 0 0 0 0 0
Hylidae 182 169 21 4 1 0 66 18 93 16 0 30
Hyperoliidae 49 47 3 0 0 0 2 1 0 0 1 5
Leiopelmatidae 4 2 4 0 0 0 2 3 4 0 0 2
Leptodactylidae 564 490 28 13 2 0 128 19 119 67 0 75
Limnodynastidae 10 9 8 0 1 0 8 0 2 2 0 2
Mantellidae 35 35 22 10 0 0 4 1 0 0 1 14
Megophryidae 44 36 0 5 2 0 14 0 0 2 0 3
Microhylidae 72 63 16 2 0 0 9 2 0 6 0 19
Myobatrachidae 12 11 10 0 0 0 6 5 7 2 0 6
Nasikabatrachidae 1 1 0 0 0 0 0 0 0 0 0 0
Pelobatidae 1 1 1 0 0 0 1 0 0 0 0 0
Petropedetidae 24 23 4 0 0 0 6 0 3 0 0 2
Pipidae 3 2 1 0 0 0 1 0 0 0 0 0
Ranidae 162 143 19 24 6 0 51 13 9 9 0 7
Rhacophoridae 100 89 1 1 1 0 35 13 0 4 0 8
Rhinodermatidae 2 2 0 0 0 0 2 0 2 0 0 1
Sooglossidae 4 0 3 0 0 0 0 0 0 0 0 3
TOTAL ANURA 1,672 1,440 186 72 18 0 479 97 343 129 3 220
Caudata
Ambystomatidae 13 13 13 5 2 1 12 1 2 5 0 1
Cryptobranchidae 1 1 0 1 0 0 1 0 0 0 0 0
Hynobiidae 27 25 2 15 0 0 11 2 1 4 0 0
Plethodontidae 176 150 2 1 0 0 23 22 16 9 1 66
Proteidae 2 2 0 0 0 0 2 0 0 1 0 0
Rhyacotritonidae 1 1 0 0 0 0 0 0 0 0 0 0
Salamandridae 29 20 2 11 2 0 13 5 0 3 0 1
TOTAL CAUDATA 251 212 19 33 4 1 62 30 19 22 1 68
Gymnophiona
Caeciliidae 3 1 2 0 0 0 1 1 0 0 0 2
Ichthyophiidae 2 2 0 0 0 0 2 0 0 0 0 0
TOTAL GYMNOPHIONA 5 3 2 0 0 0 3 1 0 0 0 2

Table 12. The major threats to threatened amphibians globally in each amphibian Family.
Utilization, a major threat for 6% of all threatened species, is also only recorded as a
threat in 15 of the 36 families containing threatened species. There are seven families in
particular that contain a greater proportion of species affected by utilization compared with
the average: Hynobiidae (56%), Salamandridae (38%), Ambystomatidae (38%), Mantel- Threat type Afrotropical Australasian Indomalayan Nearctic Neotropical Palaearctic
lidae (29%), Ranidae (15%), Dendrobatidae (14%) and Megophryidae (11%). Species in All habitat loss 92 76 88 79 89 81
the Dendrobatidae, Mantellidae and Salamandridae are particularly popular in the pet Agriculture – Crops 80 6 61 29 71 37
trade, and those in the Ranidae and Megophryidae are quite popular, particularly in Asia Agriculture – Tree plantations 8 8 18 15 13 4
for providing food. Agriculture – Livestock 33 36 5 14 42 14
Timber and other vegetation removal 75 26 64 49 64 43
Urbanization and industrial development 74 44 46 39 47 53
MAJOR THREATS ACROSS BIOGEOGRAPHIC REALMS Invasive species 22 67 3 21 7 8
Utilization 6 2 8 8 3 37
Comparing threats across biogeographic realms, habitat loss is still by far the most significant Accidental mortality 0.4 3 3 8 0.3 4
threat, affecting at least over three-quarters of all threatened species in every region (Table Persecution 0 0 0 1 0 0
13). However, there is substantial variation in the relative importance of the various causes of Pollution 13 47 32 34 29 37
habitat loss across the regions. For example, in the Afrotropics, 80% of threatened species are Natural disasters 1 18 8 11 6 10
affected by expanding croplands, as are the majority of threatened species in the Neotropics Disease 2 36 0.3 18 28 2
and Indomalayan realms. In Australasia, on the other hand, only 6% of threatened species are Human disturbance 0.4 24 6 15 9 10
impacted by this threat, and only 29% and 37%, respectively, are affected in the Palaearctic Changes in native species dynamics (excluding disease) 1 0 0 1 0.1 0
and Nearctic. These differences between the regions probably reflect the timing of large-scale Fire 21 24 6 16 17 4
agricultural expansion, the three regions with the minority of threatened species affected being
those that have already converted most of their suitable land to agriculture. A similar trend,
albeit to a lesser extent, is evident when one looks at habitat loss that has taken place as a Table 13. The percentage of threatened species affected by each major threatening process across regions. Only present threats to species
result of timber and other vegetation removal. Again, the three regions most impacted are are tallied.
those that still have a more significant proportion of their original forest cover.
The significance of invasive species and disease as a major threat varies substantially
between regions. Invasive species is the second most significant threat in the Australasian
Realm, affecting 67% of threatened species, which is three times more than any other Threat type Larval % of larval Direct % of direct Live-bearing % of live-bearing
region, and much higher than the global average of 11%. The Afrotropics and Nearctic also All habitat loss 963 89 657 86 12 60
have a higher proportion of species than the global average affected by this threat (22% Agriculture – Crops 651 60 549 72 7 35
and 21%, respectively). In the other three realms, the importance of invasive species as a Agriculture – Tree plantations 133 17 96 13 0 0
threat is comparatively small. Australasia is also the region with the highest proportion of Agriculture – Livestock 351 32 256 34 1 5
threatened species affected by disease (36%), followed by the Neotropics with 28%, and Timber and other vegetation removal 631 58 510 67 9 45
the Nearctic with 18%. In comparison, the Afrotropics, Indomalayan and Palaearctic realms Urbanization and industrial development 586 54 336 44 9 45
have far fewer species affected, only 2%, 0.3% and 2%, respectively. Invasive species 168 15 32 4 1 5
Utilization is a relatively minor threat in all regions (affecting 8% or less of threatened Utilization 103 9 1 0.1 1 5
species), except for the Palaearctic Realm where 37% of threatened species are affected. Accidental mortality 21 2 1 0.1 0 0
This figure is mainly driven by the significant number of Chinese species that are utilized. The Persecution 1 0.1 0 0 0 0
relative importance of the different threatening processes within each region is discussed Pollution 396 36 142 19 1 5
further in Chapters 5-10. Natural disasters 68 6 60 8 0 0
Disease 250 23 110 14 2 10
Human disturbance 72 7 77 10 1 5
MAJOR THREATS ACROSS REPRODUCTIVE MODES Changes in native species dynamics (excluding disease) 3 0.3 1 0.1 0 0
Fire 140 13 146 19 1 5
Not surprisingly, habitat loss is the most significant threat across all three reproductive modes
(Table 14). When comparing the various types of habitat loss there is no difference between
larval and direct developers in the ranking of importance, although for direct developers Table 14. Major threats to threatened species employing different reproductive modes. Only present threats to species are tallied.
42 Threatened Amphibians of the World

Table 15. A summary of the population trends

for all extant amphibians, as well as across the Population Trend Number of extant % of all extant Afrotropical Australasian Indomalayan Nearctic Neotropical Palaearctic
different realms, given as a percentage of all amphibians globally species globally
species in the realm. Decreasing 2,500 42 44 10 52 31 45 53
Stable 1,567 27 23 53 16 48 25 27
Increasing 29 0.5 1 0.4 0.2 1 1 1
Unknown 1,788 30 32 37 31 20 29 19

Table 16. The number of rapidly declining

amphibians in each IUCN Red List category Threat Number of species EX EW CR EN VU NT Number threatened or Extinct % of rapidly declining that
categorized by the type of threat they are rapidly declining are threatened or Extinct
experiencing. Over-exploitation (A) 38 0 0 8 8 11 11 27 71
Reduced habitat (B) 206 1 0 66 33 29 77 129 63
Enigmatic decline (C) 226 8 1 156 38 14 9 217 96
All rapidly declining species 470 9 1 230 79 54 97 373 79

Figure 17. The number of “rapidly declining” 250

the percentage of species affected by expanding cropland, livestock and timber and other
amphibians in the IUCN Red List Categories, vegetation removal is slightly higher. For live-bearing species, it is difficult to identify any
broken into the major threat causing the trends as there are only 20 threatened species employing this reproductive mode.
Number of species that have changed category from 1980 to 2006

decline. Invasive species are a much greater threat to larval developers, affecting 15% of
200 threatened species. This is most likely because the vast majority of amphibians in Australia,
the country most impacted by invasive species, are larval developers (87%). Utilization is
primarily a threat affecting only larval-developing species, with 103 of the 105 species af-
150
fected employing this reproductive mode. Like invasive species, this trend in utilization is
most likely because the country that utilizes the most species, China, is also dominated by
larval developers (there are only nine direct-developing species in China).
Pollution is a more significant threat to larval-developing rather than direct-developing
100 species. Most larval developers rely on freshwater for breeding making them more sus-
ceptible to water pollution than direct developers that are more likely to be restricted to
terrestrial habitats. Disease is also more prevalent amongst larval developers (23% compared
with 14% for direct developers) and this may also be explained by larval developers being
50
more often linked to freshwater habitats, since it is suspected that chytrid is most likely
transmitted from one site to another through waterways.

0 Population Status and Trends

Extinct Extinct in the Wild Critically Endangered Endangered Vulnerable Near Threatened

Table 17. The number of rapidly declining IUCN Red List Category in 2006 By measuring trends in the status of biodiversity, we can gain a better understanding of
species in each taxonomic Order and Family. Over-exploitation Reduced habitat Enigmatic decline the impact that humans are having, and determine how successful we are at addressing
*these percentages differ significantly from biodiversity loss. This has become all the more important given the target set by the nations
the global average21. of the world for significantly reducing the rate of loss of biological diversity by 201018. Here,
we use IUCN Red List data for amphibians to present several insights into the overall trends
of amphibian species globally, including the use of an indicator, developed specifically to
Family Over-exploitation (A) Reduced habitat (B) Enigmatic decline (C) Total rapidly % of Family help monitor progress towards achieving this ambitious target.
declining species
Anura
Arthroleptidae 0 3 0 3 6 ESTIMATES OF POPULATION TRENDS
Bufonidae 0 26 81 107 22*
Centrolenidae 0 0 4 4 3 Red List data offer several means of estimating population trends. The most simple involves
Dendrobatidae 0 6 11 17 7 categorizing the current population trend for all extant amphibians as either decreasing,
Discoglossidae 0 3 0 3 25 stable, increasing or unknown. These trend data are not necessarily derived from population
Hemisotidae 0 1 0 1 11 data, and are often based on expert opinion related to the impact of threats, including, for
Hylidae 0 17 49 66 8 example, habitat loss, invasive species and the confirmation of disease within populations,
Hyperoliidae 0 6 0 6 2* as well as changes in range size.
Leiopelmatidae 0 0 1 1 25 At least 42% of amphibian species are believed to be declining, only 27% are considered
Leptodactylidae 9 48 53 110 9 to have generally stable populations, and fewer than 1% of species are believed to be
Limnodynastidae 0 4 3 7 14 increasing (Table 15). As many as 30% of species are so poorly known that it is difficult to
Mantellidae 1 0 0 1 1* give even a rough estimate of population trends. Unfortunately, it seems plausible that this
Megophryidae 0 4 0 4 3 30% will include many additional species that are undergoing population declines. These
Microhylidae 3 7 0 10 2* general figures give some indication of the magnitude of the decline in amphibian popula-
Myobatrachidae 0 2 7 9 13 tions globally. Although currently one-third of species are globally threatened, there are also
Pelobatidae 0 2 0 2 50 significant numbers of non-threatened species that have declining populations (albeit that are
Petropedetidae 0 7 0 7 7 not currently declining at a rate fast enough to qualify for listing under a threatened category).
Ranidae 17 31 6 54 8 It is inevitable that many of these species could eventually become globally threatened if
Rhacophoridae 0 9 0 9 3* threats are not mitigated and their populations continue to decline.
Rheobatrachidae 0 0 2 2 100* Although the small number of species with increasing populations includes populations
Rhinodermatidae 0 0 1 1 50 that are recovering from past declines, for the most part these species are often widespread
Scaphiopodidae 0 1 0 1 14 invasive animals that can represent a significant threat to native amphibian populations
TOTAL ANURA 30 177 218 425 8 through predation, competition, hybridization, or disease transmission.
Caudata The magnitude of the decline in amphibian populations varies across biogeographic
Ambystomatidae 2 4 0 6 20 realms (Table 15). The two regions with the highest proportion of species with decreasing
Cryptobranchidae 1 0 0 1 33 populations are the Indomalayan and Palaearctic realms. The Indomalayan Realm also
Hynobiidae 1 1 0 2 4 has the lowest proportion of species with a stable population. The Australasian Realm
Plethodontidae 0 16 8 24 7 has significantly more optimistic statistics, with only 10% of species considered to have
Proteidae 0 1 0 1 17 populations in decline, and 53% of species with stable populations, although this very likely
Salamandridae 4 7 0 11 16 is partly due to the stabilizing effect of New Guinea (where threats are still relatively minor).
TOTAL CAUDATA 8 29 8 45 8 This region also has the highest percentage of species for which the population trend is
TOTAL ALL AMPHIBIANS 38 206 226 470 16 unknown (37%); however, again, most of these lesser known species are from New Guinea
where populations are less likely to be in decline.

Biogeographic Realm Over-exploitation (A) % of rapidly Reduced habitat (B) % of rapidly Enigmatic decline (C) % of rapidly TOTAL % of species
declining species declining species declining species in the region
Afrotropical 3 10 26 87 1 3 30 3
Australasian 0 0 12 34 23 65 35 6
Indomalayan 20 34 37 64 1 2 58 6
Nearctic 1 4 14 58 9 38 24 7
Neotropics 12 4 99 32 194 64 305 10
Palaearctic 18 43 23 55 1 2 42 9

Table 18. The number of rapidly declining amphibians in each biogeographic realm, categorized by the type of threat they are experiencing.
 Chapter 4. The State of the World’s Amphibians 43

“RAPIDLY DECLINING” SPECIES

For amphibians, 2004 was the first time that all species were assessed using the Red List
Categories and Criteria. The first comprehensive assessment of all species in a group is a
measure of the current projected extinction risk of the group, but does not give any indica-
tion of how this is changing with time. Therefore, to estimate an overall trend in projected
extinction risk for amphibians, the information used to make the assessments in 2004 was
also used to assign a backcast Red List Category for 1980. These backcast categories were
determined by the GAA coordinating team and not by the assessors of the species in 2004.
They are not official Red List assessments, as these cannot be accepted retrospectively,
but are instead used here to identify species that appear to be in rapid decline. When
determining these backcast categories, it was assumed that the category in 1980 was the
same as in 2004 unless there was evidence to support a change in category. It is most likely
that this methodology for retrospectively assigning categories has resulted in a conservative
estimate of the change in status of amphibians, since undoubtedly there would have been
some changes in status for some species for which there was insufficient evidence available
to support a change in category.19
Rapidly declining species are defined here as species that have moved to a higher Red
List category of threat between 1980 and 2006. They have been categorized into three
groups according to the most dominant threat that is thought to be causing the decline:
over-exploitation (A); reduced habitat (B); or enigmatic decline (C).
Particular threats appear to be pushing amphibians to extinction more quickly than others.
Of the 470 species that are rapidly declining (see Appendix IV), 373 (79%) are considered
threatened or already Extinct (Table 16 and Figure 17). The most likely threat to push species
into a threatened category or to extinction is enigmatic decline (96% of enigmatic decline
species are threatened or already extinct). In contrast, only 63% of rapidly declining species

1.0

Baskets of frogskins from a market in Thailand.

0.9 suffering from reduced habitat are currently threatened or already extinct. For rapidly declin- The GAA recorded more than 220 species of
ing species experiencing over-exploitation, 71% of these species are currently threatened amphibians currently used for human con-
and there are no documented extinctions. sumption, although in some cases such use
RLI

0.8
Species in certain taxonomic groups are more likely to be experiencing a rapid decline than does not always represent a major threat to
other groups (Table 17). For example, there are no caecilians known to be in rapid decline at the species. © Peter Paul van Dijk
0.7 present, but around 8% of all frogs and toads and 8% of salamanders and newts are. Two
families of anurans, Bufonidae and Rheobatrachidae20, have significantly more species in
rapid decline compared with the average. There are currently 476 species described in the
0.6
Bufonidae, 107 (22%) of which are in rapid decline.
Amphibians in different regions of the world are experiencing different threats, and
0.5 these threats also vary in their severity (Table 18). For example, in the Australasian and
1980 1985 1990 1995 2000 2005 2010 Neotropical realms most rapidly declining species are experiencing enigmatic declines. For
Year the other four realms, reduced habitat is the reason for decline amongst the majority of Endnotes
rapidly declining species. In the Indomalayan and Palaearctic realms, over-exploitation is 1 www.maweb.org
also responsible for over one-third of the declines amongst rapidly declining species. The 2 This is the number of amphibians officially recog-
Neotropics overall has the most rapidly declining species, with 10% of amphibians in the nized as valid by the GAA as of December 2006.
Figure 18. Red List Index for amphibians. If all species are Least Concern the RLI is equal region currently considered to be in rapid decline. The Nearctic has the least number of At the time of going to press, 6,184 species were
to 1, and if all species are Extinct the RLI is equal to 0. amphibians in rapid decline, only 24; however, as a percentage of species richness in the officially listed on Amphibian Species of the
region, the Afrotropics have the least rapidly declining species with only 3%. World (v 5.0), and 6,308 on AmphibiaWeb (Feb
25, 2008).
3 A threatened species is one that is included in the
1.0
RED LIST INDEX IUCN Red List Categories Vulnerable, Endangered
or Critically Endangered.
0.9 The most robust means of looking at how the Red List status of large groups of species 4 At the time of going to press, there were reports
has changed over time is by means of the IUCN Red List index. The Red List Index (RLI) of the rediscovery of one EX species, namely
uses the Red List categories of species within a group, and how they change over time, to Philautus travancoricus from the Western Ghats
RLI

0.8
estimate an overall trend in projected extinction risk (Butchart et al. 2004, 2007). Every time a in India (S.D. Biju pers. comm.).
comprehensive assessment of all species in the group is completed, the Red List categories 5 A taxon is presumed Extinct, according to the IUCN
0.7 are then used to calculate a value for the RLI for this time period. For amphibians, the RLI Red List Categories and Criteria, when exhaustive
was calculated in 2004 (Baillie et al. 2004), but since then the methodology for calculating surveys in known and/or expected habitat, at
the RLI has been revised, a small number of amphibians have been reassessed, and some appropriate times (diurnal, seasonal, annual),
0.6
assessments have been corrected. Here, we present a calculation of the RLI for amphibians throughout its historic range have failed to record
using the revised formulation and use the most current categories for each species. an individual. Surveys should be over a time frame
0.5 Between 1980 and 2006 there has been a decrease in the RLI, as illustrated by a down- appropriate to the taxon’s life cycles and life form.
1980 1985 1990 1995 2000 2005 2010 ward trend in Figure 18, which indicates that the projected extinction risk for amphibians Species that are believed to have gone extinct prior
Year is increasing. In order to meet the Convention on Biological Diversity target of significantly to 1500 AD are not considered.
reducing the loss of biodiversity by 2010 (Secretariat of the Convention on Biological Diversity 6 At the time of going to press, Meegaskumbara et
AUS PAL NEAR AFR IND NEO
2003), the RLI must increase from the previous time period – i.e., the line in Figure 13 must al. (2007) had just described two additional extinct
show an upward trend. With only two data points for the amphibian RLI, at present it can species, Philautus pardus and P. maia, known only
Figure 19. Red List Index for amphibians in the different biogeographic realms. (Note: only be shown that the projected extinction risk is increasing, but as the global assessment from collections made in Sri Lanka prior to 1876;
species that occur in more than one realm contribute to the RLI in all the realms in which of amphibians continues on a regular basis, future data points will show whether or not the both species await formal assessment.
they occur.) Key to realms: AUS = Australasian; PAL = Palaearctic; NEAR = Nearctic; AFR = rate of increase is accelerating or decelerating. 7 When interpreting these results, it is necessary
Afrotropical; IND = Indomalayan; NEO = Neotropical. The Red List Index can also be disaggregated, for example, by biogeographic realm (Figure to bear in mind that a species can be listed in
19) and by reproductive mode (Figure 20). All six realms show an increase in the projected a category on the basis of one or more criteria,
extinction risk for amphibians from 1980 to 2006. However, the level of increase varies across and that a species may trigger criteria for a lower
realms, as does the average threatened status of amphibians. The current threat status of category of threat than that for which it is listed
1.0
amphibians is best (that is on average there are less threatened species) in the Australasian (these criteria are not recorded). For example, a
Realm and worst in the Neotropical Realm. For all regions, the RLI shows a downward trend species may qualify for Endangered on the basis
0.9 – i.e., the projected extinction risk has increased in 2006 compared with 1980, indicating of criterion B and for Vulnerable on the basis
that amphibian declines are a global problem. The rate of increase in projected extinction of criterion C. When listed as Endangered only
risk is lowest for the Afrotropics, and highest for the Neotropics. criterion B will be given.
RLI

0.8
The RLI was also calculated across the three different reproductive modes: larval develop- 8 As discussed elsewhere in this volume, the
ers, direct developers or live-bearing species. There are also quite a few species for which major reclassification by Frost et al. (2006) and
0.7 the reproductive mode is not known and these species were omitted from this RLI calculation. Grant et al. (2006) has dramatic implications for
The results show that the average threatened status of larval developers in 1980 and 2006 amphibian classification at the family level. The
is better than live bearers and direct developers. All three groups show a downward trend results presented here can be expected to undergo
0.6
in the RLI, indicating an increase in projected extinction risk, but the increase for direct substantial modification based on the conclusions
developers is less compared with larval developers and live bearers. of that study.
0.5 9 P<0.01 (binomial test)
1980 1985 1990 1995 2000 2005 2010 10 No amphibian species occur in the Antarctic Realm,
Year KEY FINDINGS while only three species occur in the Oceanian
Realm, so we include the Oceanian Realm in all
Larval developers Direct developers Live-bearing
• The GAA assessed 5,915 amphibian species globally and categorized one-third of all discussion on the Australasian Realm. For further
extant species (1,893 out of 5,881) as threatened with extinction. information on the boundaries between these
Figure 20. Red List Index for amphibians across the different reproductive modes. • There are 34 species currently listed as Extinct and one as Extinct in the Wild. regions, please refer to the regional summaries.
44
Plectrohyla dasypus (Critically Endangered)
from the Parque Nacional Cusuco in Honduras
is one of many species that have undergone
a rapid decline in status, in this case due
to the effects of chytridiomycosis. © Silviu
Petrovan
11 The range size for each species has been calcu-
lated using the area of the polygons representing
each species’ Extent of Occurrence.
12 The methodology for mapping these amphibian
“centres of endemism” follows that developed by
BirdLife International in identifying their Endemic
Bird Areas (Stattersfield et al. 1998). Around two-
thirds of amphibians have range sizes less than
50,000km² compared with around one-quarter of
birds.
13 Table 10 only includes countries with more than 10
species. If all countries are included, the country
with the highest percentage is New Zealand with
all four of its species, which are also all endemic,
threatened with extinction.
14 The range size for each species has been calcu-
lated using the area of the polygons representing
each species’ Extent of Occurrence.
15 Selective logging is generally defined as the
process by which only the most desirable trees,
often the largest and\or most commercially viable,
are logged from an area of forest; in contrast, clear-
cutting is generally defined to be a large-scale
and commercial process, by which all trees from
a given area are taken in a logging operation.
16 Although amphibian deformities can result from
exposure to contaminants, many cases of limb
deformities in nature have been linked to other
abiotic and biotic factors, such as UV radiation,
parasitic infections, retinoids, predation, and the
interaction of several of these factors (see Oullet
2000; Sessions 2003).
17 In this study, pesticides that inhibit acetyl-
cholinesterase were most strongly correlated
with declines. Such cholinesterase inhibitors (or
“anticholinesterase”) suppress the action of the
enzyme, and ultimately lead to death. Anticholin-
esterases are commonly used for killing insects in
organophosphate and other pesticides.
18 http://www.biodiv.org/2010-target/default.asp
19 In 2006, a small number of species were reas-
sessed, and the new information supporting these
reassessments was also used to better estimate
the categories in 1980, and where necessary
some categories were changed. The Red List
categories for all species in 2006 and the revised
1980 categories form the basis of the information
in the section on rapidly declining species.
20 The family Rheobatrachidae has only two species,
both of which are now Extinct.
21 P<0.01 (binomial test)
Threatened Amphibians of the World
• Within amphibian Orders, Caudata (Salamanders and Newts) contains the highest
proportion of threatened species (47%), followed by Anura (Frogs and Toads) (32%) and
Gymnophiona (Caecilians) (3%).
• Very diverse families of frogs and toads that are significantly more threatened than the
global average include the Bufonidae (45%), Leptodactylidae (46%), and Rhacophoridae
(43%). Diverse families that are less threatened than the global average include Ranidae
(25%), Microhylidae, (17%) and Hyperoliidae (19%).
• The Neotropical realm has the highest number of species (2,916, of which 2,808 are
endemic to the realm) and the greatest proportion of threatened species (39%).
• Major concentrations of threatened species include the sierras and trans-volcanic belt
of Mexico, Central America, the tropical Andes, Brazil’s Atlantic Forest, the Greater
Antilles, the Upper Guinea forests of West Africa, the forest of western Cameroon
and eastern Nigeria, the Albertine Rift, the Eastern Arc Mountains of Tanzania and
other Afromontane outliers, the south-east coast of South Africa, Madagascar’s wet
forests, the Western Ghats and Sri Lanka, montane Southeast Asia, and eastern
Australia.
• Brazil, with 751 species currently recognized, is the country with the most amphibian
species, followed closely by Colombia with 697. These two countries also have the
highest number of endemic species, 489 and 337, respectively.
• Four countries have more than 100 threatened species: first is Colombia with 209, second
is Mexico with 198, third is Ecuador with 163, and fourth is Brazil with 110.
• Almost 80% (4,710 species) of amphibians occur in forest habitats, and 90% (4,251
species) of these species occur in tropical forest. Species occurring in montane tropical
forests are more likely to be threatened (46% of species), than those occurring in lowland
tropical forest (28% of species).
• When comparing reproductive modes of amphibians, 45% of direct-developing spe-
cies, 28% of larval-developing species, and 34% of live-bearing species are currently
threatened with extinction or are already extinct.
• Habitat loss, primarily due to expanding croplands, vegetation removal (mainly logging),
and urbanization/industrial development, is the most pervasive threat to amphibians,
affecting 63% of all species and 87% of threatened species. Other threats that are
commonly recorded include pollution, invasive species, fire and disease. Disease is a
major threat for 8% of all species and for 19% of threatened species.
• At least 42% of amphibian species are believed to have declining populations, only
27% are considered to have generally stable populations, and less than 1% of species
are believed to be increasing. As many as 30% of species are so poorly known that it is
difficult to give even a rough estimate of population trends.
• There are 470 “rapidly declining“ species globally. Of these, 38 species are declining
primarily due to over-exploitation, 206 are declining due to reduced habitat, and 226
are experiencing “enigmatic declines” (probably due to the effects of chytridiomycosis
in combination with climate change).
• Calculation of the Red List Index for amphibians shows a worsening trend in extinction
risk for amphibians from 1980 to 2006.
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 Chapter 4. The State of the World’s Amphibians 45

ESSAY 4.1. PICKLED FROGS HELP BIODIVERSITY ASSESSMENT

At a time when amphibian populations globally are in decline, the recent extant Sri Lankan direct developers are not associated closely with freshwater References
discoveries of large numbers of new frog species on Sri Lanka, an island habitats, the role of waterborne diseases in Sri Lanka’s extinction of direct
from which the pathogenic chytrid fungus has not yet been reported, may developers may have been slight. Dalton, R. 2003. Natural history collections in crisis as funding is slashed. Nature 423:575.
seem heartening (Meegaskumbura et al. 2002). A total of 42 new species The large-scale extinction of many of the island’s amphibians might have Kumar, A., Pethiyagoda, R. and Mudappa, D. 2004. Western Ghats and Sri Lanka.
of anurans have been described since 1993, and many more species are in gone undetected if not for the historical collections in the world’s natural Pp. 152-157, in: R.A. Mittermeier, P. Robles-Gil, M. Hoffmann, J.D. Pilgrim, T.M.
the process of being formally described. Description of these new species history museums; unfortunately, old amphibian collections in Sri Lanka’s Brooks, C.G. Mittermeier, J.L. Lamoreux and G. Fonseca (eds.), Hotspots Revisited.
necessitated the examination of all available type material, as well as other National Museum have not survived the years to help facilitate this effort. Cemex. 392 pp.
preserved voucher material, for all Sri Lanka’s amphibians. As a result of the Clearly, the extraordinary value historical collections represent by way of Manamendra-Arachchi, K. and Pethiyagoda, R. 2005. The Sri Lankan shrub-frogs of the
country’s five-century history of colonial occupation, these specimens, col- biodiversity baselines against which present-day surveys may be assessed, genus Philautus Gistel, 1848 (Ranidae: Rhacophorinae), with description of 27 new
lected largely between 1850 and 1950, lie scattered amongst natural history remains to be widely appreciated (Fig. 1). This is especially relevant at a time species. The Raffles Bulletin of Zoology, Supplement No. 12:163-303.
museums in Europe, the United States and India. In particular, since Sri Lanka when the contemporary role and value of natural history collections is being Meegaskumbura, M., Bossuyt, F., Pethiyagoda, R., Manamendra-Arachchi, K., Bahir,
was a British colony from 1796 to 1948, most type specimens are housed in questioned, and with many museums facing financial crises (Dalton 2003; M.M., Schneider, C. J. and Milinkovitch, M.C. 2002. Sri Lanka: an amphbian hotspot.
The Natural History Museum (formerly the British Museum of Natural History), Suarez and Tsutsui 2004; Winker 2004). Furthermore, historical collections can Science 298:379.
in London. Sri Lanka’s own national museum contains no amphibian types be useful in several other ways, as exemplified most recently by the detec- Meegaskumbura, M. and Manamendra-Arachchi, K. 2005. Description of eight new species
whatsoever. Although locality information beyond ‘Ceylon’ is usually lacking tion of chytrid fungus in the skin of a museum specimen collected from near of shrub frogs (Ranidae: Rhacophorinae: Philautus) from Sri Lanka. The Raffles Bulletin
on the specimen labels, these specimens provide the only available baseline Monteverde, Costa Rica, and deposited in a museum well before declines of Zoology, Supplement No. 12:305-338.
of Sri Lanka’s amphibian fauna before the island’s primary rainforests were were documented in the area (Puschendorf et al. 2006). Puschendorf, R., Bolanos, F., and Chaves, G. 2006. Amphibian chytrid fungus along
reduced to their present-day extent of approximately 750km2. Interestingly, since most of the species that were recently described an altitudinal transect before the first reported declines in Costa Rica. Biological
The recent review of historical material served to highlight several key (Manamendra-Arachchi and Pethiyagoda 2005; Meegaskumbura and Mana- Conservation 132:136-142.
aspects. First, a number of species described in the 19th century, and since mendra-Arachchi 2005) were absent from early museum collections, it seems Ricketts, T.H., Dinerstein, E., Boucher, T., Brooks, T.M., Butchart, S.H.M., Hoffmann, M.,
relegated to synonymy, were shown to represent valid taxa. Second, several that past surveys of the island were not very thorough. The documentation Lamoreux, J.F., Morrison, J., Parr, M., Pilgrim, J.D., Rodrigues, A.S.L., Sechrest, W.,
‘new species’ lay silently undiscovered among the old collections (Manamen- of the extinction of so many species from such a limited collection suggests Wallace, G.E., Berlin, K., Bielby, J., Burgess, N.D., Church, D.R., Cox, N., Knox, D.,
dra-Arachchi and Pethiyagoda 2005; Meegaskumbura and Manamendra- that these extinct species may be only a fraction of a much larger extinction Loucks, C., Luck, G.W., Master, L.L., Moore, R., Naidoo, R., Ridgely, R., Schatz, G.E.,
Arachchi 2005). At the same time, the Wildlife Heritage Trust’s 1993-2005 event in Sri Lanka. The account of Sri Lanka’s amphibian declines is also novel Shire, G., Strand, H., Wettengel, W. and Wikramanayake, E. 2005. Pinpointing and
comprehensive survey of the island’s amphibian fauna provides a reliable in that it combines a recent comprehensive survey of the island’s amphibians preventing imminent extinctions. Proceedings of the National Academy of Sciences
record of the species still persisting on the island and their relative abundance. with a re-examination of almost all preserved material worldwide. Many other USA 102:18497-18501.
A comparison of the results of the recent biodiversity survey, together with tropical countries are similarly under-explored, and it is therefore likely that Stuart, S.N., Chanson, J.S., Cox, N.A., Young, B.E., Rodrigues, A.S.L., Fischman, D.L. and
the historically preserved specimens, revealed that 19 named anuran species the global tally of recent amphibian extinctions will rise significantly when Waller, R.W. 2004. Status and trends of amphibian declines and extinctions worldwide.
have apparently disappeared from the island (an additional two undescribed their historical collections are similarly evaluated and compared with current Science 306:1783-1786.
species also appear to be extinct). Given that these 19 species had not been assessments of amphibian diversity. Suarez, A.V. and Tsutsui, N.D. 2004. The value of museum collections for research and
reported since their original collections before 1940, and were not recorded society. BioScience 54:66-74.
during the recent surveys across the island’s remaining forests, they were Madhava Meegaskumbura, Rohan Pethiyagoda, Janice Winker, K. 2004. Natural history museums in a postbiodiversity era. BioScience 54:455-
formally declared Extinct according to the IUCN Red List categories and Chanson, David Wake and James Hanken 459.
criteria (GAA; Stuart et al. 2004). Apart from a single species each from the
endemic genera Adenomus and Nannophrys, the Extinct species all belong
to the Oriental shrub-frog genus Philautus (Ranidae: Rhacophorinae), which
shows a remarkable radiation in Sri Lanka. A total of 62 of the approximately The collections of the world’s natural history museums represent important biodiversity baselines. Here, preserved amphibian specimens collected in Sri
140 species currently recognized in this genus are endemic to Sri Lanka. Lanka ca. 1900 are stored on the shelves of The Natural History Museum in London © Barry Clarke/NHM
To put these findings in a global context, according to the results of the
Global Amphibian Assessment, 19 of 34 amphibian species confirmed as
Extinct worldwide are from Sri Lanka. Even though a further 122 species are
considered ‘possibly extinct’ globally, the Sri Lankan total is still remarkably
high. Given that the island accounts for only around 2% of the world’s anuran
species (which is high given its relatively small size: 65,000km2), it is surpris-
ing that more than half of the confirmed amphibian extinctions worldwide
have occurred on Sri Lanka.
This extinction phenomenon in Sri Lanka (19 of the 103 native species
described to date) appears to have been driven largely by the destruction of
suitable habitat. Since 1815, approximately 95% of the island’s 16,000km2 of
rainforest was lost to coffee and cinchona plantations, later to be replaced
by tea and rubber (Kumar et al. 2004). While large expanses of dry forest
and scrub persist elsewhere in the island, these habitats only support three
of the 68 extant endemic amphibian species, with only a single species,
Philautus regius (DD), restricted to dry forest. Of Sri Lanka’s 84 surviving
anuran species, 11 are Critically Endangered (seven of them Philautus) and
28 Endangered (20 Philautus). Many of these surviving amphibian species,
especially the Philautus, have extremely restricted ranges. Even where there
are larger tracts of contiguous forest, the ranges of many of these species
are restricted by altitude or vegetation, resulting in the total contemporary
range of some species being as small as 5km2. Some 17 species are known
only from a single site each (per Ricketts et al. 2005), with some sites, such
as Morningside and Knuckles Forest Reserve, being home to several endemic
and threatened species.
Interestingly, though not unexpectedly, the results of the GAA also sug-
gest that, among threatened direct-developing species worldwide, 65 out
of the 115 (57%) species that are closely associated with freshwater are
threatened by disease, compared with only 45 out of the 650 (7%) species
not closely associated with freshwater. Given that the major global threat
to direct developers appears to be habitat loss (Chapter 4), and most of the

ESSAY 4.2. AMPHIBIANS AND HUMANS SHARING ONLY ONE PLANET

To effectively conserve amphibian populations in the wild it is not only
necessary to understand the needs of the individual species, but also the
context in which conservation efforts will need to take place. By comparing
overlays of individual species range maps with recent and future human
demographic variables (such as human population density, population growth,
Gross National Income, and poverty), it is possible to elucidate the social
and economic context in which conservation action must be implemented.
Here, we investigate the relationships between amphibian species richness
and two of these variables, human population density and levels of poverty
– using Infant Mortality Rate data (CIESIN 2005) as a surrogate – not as a
means of looking for causal relationships, but rather to identify regions where
conservation may be more challenging.
Human population density
By comparing the current human population density in 2005 (LandScan 2005)
with the global distribution of all amphibian species it is possible to identify
the regions that are favoured by both humans and amphibians (Figure 1).
Regions with a large number of amphibian species and a high population
density are mostly found in Asia, in particular southern and eastern India,
south-east China, southern Sri Lanka, and Indonesia (especially Java), as well
as coastal parts of West Africa, the Ethiopian Highlands, and the Albertine
Rift of Central Africa. There are also similar regions in the eastern United
States, which is a hotspot for salamander diversity. In South America, the
Atlantic Forest is the most prominent region.
Considering that humans and amphibians are dependent on freshwater
for survival, the arid regions of the world are the places with the fewest
amphibians (often none at all) and very low human population densities.
For example, the deserts of North Africa, central Australia, central North
America, and Central Asia. The majority of amphibians prefer moist tropical
conditions, and many of the regions rich in amphibian species, but with low
human population densities, are the world’s tropical rainforests, for example
the Amazon, the Congo, the tropical Andes, Southeast Asia and northern
Queensland in Australia (the Queensland Wet Tropics). Regions with low
diversity of amphibians, but high human population densities, are in general
cooler regions such as northern India, northern China and much of Europe.
By comparing the diversity of threatened amphibians with human popula-
tion density it is possible to highlight regions where species conservation is
most likely to come into conflict with increasing demand by humans for natural
resources (Figure 2). Many of these regions are the same as those highlighted
in Figure 1, for example south-east China, Java (Indonesia), the Albertine Rift
of Central Africa, coastal West Africa, and the Ethiopian Highlands. Notable
additions are the larger islands of the Caribbean, southern Mexico, and the
Philippines. In India, the highest density of threatened amphibians is focused
on the Western Ghats. Parts of the eastern United States, Europe and India
that are rich in amphibian species and have high human population densities
are no longer highlighted as they have relatively few or no threatened species.
In Europe and the United States this may be because species susceptible to
habitat loss may have declined a long time ago, and perhaps also because
many of the resources on which humans in these areas rely on come from
other regions, for example the forests of Africa and Asia.
46 Threatened Amphibians of the World

Figure 1. Amphibian species richness compared with human population density. Each quarter-degree grid cell is
coded according to the combined value of the two variables. Amphibian species richness is represented by the number
of amphibian species per grid cell (Low: 0-15; Medium: 16-30; High: 30-144). Population density is classified by the
number of people per km2 (Low: 0-10; Medium: 11-100; High: >100).
Figure 2. Threatened amphibian species richness compared with human population density. Each quarter-degree
grid cell is coded according to the combined value of the two variables. Threatened amphibian species richness is
represented by the number of threatened amphibian species (Red List Categories Critically Endangered, Endangered
and Vulnerable) per grid cell (Low: 0; Medium: 1-10; High: >10). Population density is classified by the number of
people per km2; Low: 0-10; Medium: 11-100; High: >100.

Figure 3. Amphibian species richness compared with poverty. Each quarter-degree grid cell is coded according to
the combined value of the two variables. Amphibian species richness is represented by the number of amphibian
species per grid cell (Low: 0-15; Medium: 16-30; High: 30-144). Infant Mortality Rate data (CIESIN 2005) are used
as an index of poverty. The rates are per thousand of live births and the data are at the sub-national level. The data
were reclassed as: Low: 0-100; Medium: 100-250; High: >250.
Figure 4. Threatened amphibian species richness compared with poverty. Each quarter-degree grid cell is coded
according to the combined value of the two variables. Threatened amphibian species richness is represented by the
number of threatened amphibian species (Red List categories Critically Endangered, Endangered and Vulnerable) per
grid cell (Low: 0; Medium: 1-10; High: >10). Infant Mortality Rate data (CIESIN 2005) are used as an index of poverty.
The rates are per thousand of live births and the data are at the sub-national level. The data were reclassed as: Low:
0-100; Medium: 100-250; High: >250.

Human poverty
An additional challenge to biodiversity conservation is the welfare of local hu-
man populations, in particular the degree of poverty. Many of the world’s most
densely populated regions are also some of the poorest, for example, China,
India, mainland Southeast Asia and West Africa. When comparing the most
impoverished regions with amphibian diversity we find these same regions
emerging again, but also less densely populated regions are also highlighted,
such as central and southern Africa, Madagascar, northern Brazil, Peru, and
Bolivia (see Figure 3). The land surface area of the world’s wealthiest nations
is relatively small and these countries often have cooler climates not favoured
by amphibians, hence the proportion of amphibian species found in these
countries is relatively low. Exceptions are the eastern United States, coastal
eastern Australia and central Europe. However, the relatively high numbers
of species in these regions may be more a reflection of intense survey effort
when compared with much poorer countries.
By comparing the diversity of threatened amphibians with the level of
poverty it is possible to highlight those regions where biodiversity conser-
vation will most likely suceed if it also improves the welfare of the local
communities (Figure 4). In addition these regions will less likely be unable to
shoulder any of the costs of conservation programmes. Many of these regions
are the same as those highlighted when comparing threatened species with
human population density, for example south-east China, the Albertine Rift
of Central Africa, the Ethiopian Highlands, coastal West Africa, Puerto Rico,
southern Mexico, the Western Ghats of India, and the Philippines. Regions
with a high level of poverty that were not previously highlighted as also be-
ing densely populated include Peru, Bolivia, Guyana, the Solomon Islands,
Vietnam, Madagascar, and the Cameroon highlands.
Conclusion
In an increasingly human-dominated world it is impossible for conservation to
be successful without understanding the context in which it is to take place.
By comparing the distribution of amphibians, and in particular threatened
amphibians, with human demographic data, it is possible to gain a better
understanding of some of the challenges facing conservation.
Vineet Katariya and
Janice Chanson
References
Center for International Earth Science Information Network (CIESIN), Columbia University;
2005 Global subnational infant mortality rates [dataset]. CIESIN, Palisades, NY, USA.
Available at: http://www.ciesin.columbia.edu/povmap/ds_global.html
LandScanTM Global Population Database. Oak Ridge, TN: Oak Ridge National Laboratory.
Available at http://www.ornl.gov/landscan/.

ESSAY 4.3. THE USE OF PHYLOGENIES IN PREDICTING TRENDS FOR CONSERVATION: A CASE STUDY FROM SRI LANKA

The discovery of large numbers of new amphibian species in Sri Lanka and
elsewhere (Pethiyagoda and Manamendra-Arachchi 1998; Köhler et al.
2005; and see Essay 1.1), even as population declines accelerate worldwide,
presents profound challenges to conservation managers. Prioritizing conserva-
tion actions in the face of declines caused by a diversity of threats – including
disease and habitat fragmentation and loss – is obviously difficult. Frequently,
detailed natural history knowledge that would facilitate conservation, is
missing for many, if not most, species. Nonetheless, it is often necessary to
make decisions even in the absence of detailed information.
Phylogenetic analyses of species and genera can provide an important
source of additional information, particularly when coupled with existing
ecological data. In this respect, the anurans of Sri Lanka provide an interesting
case study. Based on morphological characters, taxonomists had previously
allocated the treefrogs (subfamily Rhacophorinae of the family Ranidae) of
Sri Lanka among several genera, such as Philautus (including a subgenus
Kirtixalus), Polypedates, Rhacophorus and Theloderma. However, recent
phylogenetic analyses based on mitochondrial DNA have largely refuted
these classifications and demonstrated that Sri Lankan treefrogs belong to
only two clades: a large and diverse group of direct-developing species in
the genus Philautus, and a much smaller clade of foam-nesting species in the
genus Polypedates (Meegaskumbura et al. 2002) (Figure 1A).
While information on breeding strategy is available for 19 species of
Sri Lankan rhacophorids, several lines of evidence suggest that the entire
radiation of Philautus in Sri Lanka is characterized by direct development (i.e.,
their embryos undergo complete development within the egg, from which
they emerge as fully developed froglets, enabling them to develop with the
tadpoles bypassing the ‘conventional’ aquatic larval stage). The fact that each
of the 19 species examined shows direct development suggests, at least,
that the most recent common ancestor of those taxa was a direct developer
(Figure 1A, 1B and 1C). The sister group of the Sri Lankan clade of Philautus
is a diverse Indian clade (Meegaskumbura et al. 2002; Bossuyt et al. 2004)
in which direct development has been observed for a number of species (Biju
and Bossuyt 2005). This suggests that the most recent common ancestor of
the Indian and Sri Lankan taxa was also a direct developer. Therefore, it is
parsimonious to infer that all Sri Lankan Philautus are direct developing. In
contrast, Polypedates, which has only five species in Sri Lanka, build foam
nests on vegetation overhanging water, into which emerging tadpoles fall
and continue to develop.
While all Sri Lankan Philautus are direct developers, they exhibit two
distinct reproductive behaviours: soil-nesting and leaf-nesting (Bahir et al.
2005) (Figure 1B, 1C). While in amplexus, females of soil-nesting species
excavate a pit in damp soil with their hands into which they deposit eggs,
which are fertilized by the males. The females then cover up the nest with
soil and leaf litter. Leaf-nesting females, on the other hand, attach their clutch
to the underside of a leaf (also while still in amplexus), usually 1-2 m above
 Chapter 4. The State of the World’s Amphibians 47

the forest floor. Egg masses on leaves are highly vulnerable to desiccation A B C
when relative humidity falls below about 90%, and so the species depend on
good-quality forests with dense shade and low wind speeds. The phylogeny Philautus - Sri Lanka/ Ind (100+)
suggests that leaf nesting, as a reproductive trait, is restricted to a single
clade (Figure 1B and 1C). Philautus - Ind (100+)
Despite Sri Lanka having been connected by a land bridge to India for 3
much of its history and as recently as 10,000 years ago, phylogenies of several Kurixalus (2)
groups of organisms indicate that there has been little recent biotic exchange
between the two landmasses – particularly for taxa restricted to rainforest Philautus - SE Asia (63)
habitats (Bossuyt et al. 2004). This is true for Sri Lankan Philautus, which
Polypedates (28) P. hallidayi
belong to an old lineage distinct from the Indian direct developers, except for
a small group that appears to be derived from a species that migrated back to
India over 9 million years ago. The Sri Lankan Philautus, therefore, encompass Chiromantis (4)
substantial phylogenetic diversity that necessitates their consideration as a
unique entity from a conservation perspective. Rhacophhorus (63) 3
In addition to conserving diversity per se, it is widely recognized that
conservation must aim also to preserve the processes that generate diversity. Theloderma (9)
A mt-DNA based phylogeny has shown how sister taxa are distributed across
montane-forest refugia in Sri Lanka, suggestive of speciation by vicariance Buergeria (5)
P. alto
during the contraction of cool and wet habitats during dry periods of the
Miocene and the Pliocene (Meegaskumbura and Manamendra-Arachchi 2005)
(Figure 1D). This pattern of geographic isolation of sister taxa in patches of
favourable habitat is seen also in several other major lineages of direct-de-
veloping frogs, including the Neotropical genus Eleutherodactylus (Savage
2002), and the Afrotropical Arthroleptella (Dawood and Channin 2000), and, in-
terestingly, also in the direct-developing plethodontid salamanders of Central
America (Garcia-Paris et al. 2000). These observations suggest that montane D 2
isolation may be an important mechanism of speciation in direct-developing
amphibians. However, this does not mean that habitat fragmentation should
be encouraged. In addition, phylogenies can also help to identify refugia that
survive as habitats through successive major climatic oscillations (Hewitt and
Nichols 2005). Such refugia can be identified because they will, for a given
species, contain greater genetic diversity than areas of recent population P. stuarti
expansion (Schneider et al. 1998) and they often contain a higher diversity
P. femoralis
of montane endemic species (Schneider and Williams 2005). P. viridis 4.37-6.93*
The rhacophorine phylogeny (see Figure 1A) also suggests that the more
derived terrestrial direct development is associated with a larger number P. demoralis
5.31-11.97*
of species. The more basal Buergeria, containing only five species, shows P. mooreorum
conventional, and plesiomorphic, anuran reproduction, with eggs and larvae
undergoing development entirely in water. On the other hand, ca. 200 species P. poppiae 3
of Philautus are presently recognized, with dozens more species awaiting
description (Biju 2001; Meegaskumbura et al. 2002). Direct development
releases amphibians from dependence on water bodies for reproduction.
This, in turn, appears to enable amphibians to colonize a wide array of moist
terrestrial habitats, as evidenced by the diversity of such habitats (e.g., forest
P. asankai
canopy, leaf litter, rock surfaces, and open grasslands) occupied by species of
both Philautus and Eleutherodactylus. However, greater habitat specialization 7.63-11.06*
and/or smaller habitat patches would make them susceptible to stochastic
P. steineri
population declines as well as putting them at risk due to climate change.
Phylogenetic information contributes to conservation in several important P. microtympanum
ways. First phylogenetic information is fundamental to identify biodiversity
(i.e., to diagnose species or to identify evolutionarily significant units or 5
endemic taxa). Second, phylogenies are necessary for identifying important P. decoris
adaptive diversity. For example, the Sri Lankan / Indian radiation of Philautus P. mittermeieri 4.00-5.80
appears to have evolved direct development independently from Asian Philau-
tus (Meegaskumbura et al. 2002). Separate instances of adaptive evolution are P. hoffmani
crucial for understanding adaptation and thus the Sri Lankan / Indian Philautus P. asankai
P. sp.
take on added importance for conservation. Third, knowledge of phylogenetic
relationships allows us to extend our understanding of ecology and behavior 4.15-6.04*
in a few species to clades of closely related species for which we have limited
data. For example, understanding phylogenetic relationships may allow us to
use a relatively abundant species as a surrogate to develop captive-breeding
protocols for a closely related, but Critically Endangered, species in urgent
need of ex situ conservation. Phylogenies are important on several levels and 3
now, with the widespread use of DNA sequencing and molecular phylogenet-
ics, are likely to play an increasing role in conservation biology.
P. wynaadensis

Madhava Meegaskumbura, Kelum Manamedra-Arachchi,

Rohan Pethiyagoda and Christopher J. Schneider
P. zorro

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Biju, S.D. and Bossuyt, F. 2005. Two new species of Philautus (Anura: Ranidae: Rhacoph-
orinae) from Ponmudi Hill in the Western Ghats of India. Copeia 1:29-37.
Bossuyt, F., Meegaskumbura, M., Beenaerts, N., Gower, D.J., Pethiyagoda, R., Roelants,
K., Mannaert, A., Wilkinson, M., Bahir, M.M., Manamendra-Arachchi, K., Ng, P.K.L.,
Schneider, C.J., Oommen, O.V. and Milinkovitch, M.C. 2004. Sri Lanka: a center of
faunal endemism in a Biodiversity Hotspot 21. Science 306:479-481.
Dawood, A. and Channing, A. 2000. A molecular phylogeny of moss frogs from the
Western Cape, South Africa, with a description of a new species. Journal of Her-
petology 34:375-379.
Garcia-Paris, M., Good, D.A., Parra-Olea, G. and Wake, D.B. 2000. Biodiversity of Costa
Rican Salamanders: implications of high levels of genetic differentiation and phylo-
geographic structure for species formation. Proceedings of the National Academy of
Sciences 97:1640-1647.
Hewitt, G.M. and Nichols, R.A. 2005. Genetic and evolutionary impacts of climate change.
In: T.E. Lovejoy and L. Hannah (eds.), Climate change and biodiversity, pp.176-192.
Yale University Press, New Haven, USA.
Figure 1. The mt-DNA phylogeny of Sri Lankan Philautus (modified from Meegaskumbura et al. 2002 and Meegaskumbura and Manamendra-Arachchi 2005).
1A: the genus level relationships of the Old World treefrogs shows that Buergeria is basal and shows aquatic breeding (light blue), while the more derived
groups are tree-hole breeding (black), foam nesting (green), tree-hole nesting (purple) with brood care and direct developing (Red – South East Asian and
Indian species, and Blue – Sri Lankan species). Number of species within each genus is depicted within brackets. 1B and 1C: the major clades within the
Sri Lankan radiation, with the number of species for which breeding information is available inside the triangles. The leaf nesting clade is circled. 1D: some
of the clades (highlighted by a *) restricted to cloud forest habitats and the sister taxa within these clades distributed on adjacent cloud-forest habitats
(K - Knuckles Hills, C - Central Hills; R - Rakwana Hills). In contrast, Philautus decoris – mittermeieri clade is distributed across an altitudinal gradient. This
information is useful for making conservation decisions.
Köhler, J., Vieites, D.R., Bonett, R.M., García, F.H., Glaw, F., Skeinke, D. and Vences, M. Pethiyagoda, R. and Manamendra-Arachchi, K. 1998. Evaluating Sri Lanka’s amphibian
2005. New amphibians and global conservation: a boost in species discoveries in a diversity. Occasional Papers of the Wildlife Heritage Trust 2:1-12.
highly endangered vertebrate group. BioScience 55:693-696. Savage, J. 2002. The Amphibians and Reptiles of Costa Rica: A Herpetofauna Between Two
Manamendra-Arachchi, K. and Pethiyagoda, R. 2005. The Sri Lankan shrub-frogs of the Continents Between Two Seas. University of Chicago Press, Chicago, Illinois, U.S.A.
genus Philautus Gistel, 1848 (Ranidae: Rhacophorinae), with description of 27 new Schneider, C.J., Cunningham, M. and Moritz, C. 1998. Comparative phylogeography
species. The Raffles Bulletin of Zoology, Supplement No. 12:163-303. and the history of endemic vertebrates in the Wet Tropics rainforests of Australia.
Meegaskumbura, M., Bossuyt, F., Pethiyagoda, R., Manamendra-Arachchi, K., Bahir, Molecular Ecology 7:487-498.
M.M., Schneider, C.J. and Milinkovitch, M.C. 2002. Sri Lanka: an amphibian hotspot. Schneider, C.J. and Williams, S.E. 2005. Effects of Quaternary Climate Change on
Science 298:379. Rainforest Diversity: Insights from Spatial Analyses of Species and Genes in
Meegaskumbura, M. and Manamendra-Arachchi, K. 2005. Description of eight new species Australia’s Wet Tropics. In: E. Bermingham, C.W. Dick, and C. Moritz (eds.), Tropical
of shrub frogs (Ranidae: Rhacophorinae: Philautus) from Sri Lanka. The Raffles Bulletin Rainforests: Past, Present, and Future, pp. 401-424. University of Chicago Press,
of Zoology, Supplement No. 12:305-338. Chicago, Ilinois, USA.
48
ESSAY 4.4. THE EFFECTS OF PESTICIDES ON AMPHIBIANS
Since the general recognition of global amphibian declines in the early
1990s, biologists have sought to determine the underlying causes (Alford and
Richards 1999; Stuart et al. 2004). During the past two decades, a number
of hypotheses have been developed and it seems likely that many declines
have a multitude of causes. One leading hypothesis is that some amphibian
populations are declining due to exposure to chemical contaminants including
the more than 10 billion kg of globally applied pesticides (U.S.E.P.A. 1997).
Understanding all of the mechanisms by which contaminants may impact
amphibian populations is certainly a difficult proposition because there are
more than 80,000 registered chemicals in the United States alone, and the
registration process does not require any tests on amphibians. Thus, the effect
of pesticides on amphibian populations remains a very large and open ques-
tion. Below, I very briefly review the evidence linking pesticides to amphibian
declines and then discuss the various mechanisms by which pesticides affect
amphibians and thereby potentially cause declining populations.
The evidence supporting a connection
between pesticides and amphibian declines
Documenting a definitive link between pesticides and the decline of any organ-
ism in nature is a monumental task because it requires long-term monitoring
data over large regions (because amphibian populations are normally quite
variable over time) and, ideally, the ability to conduct controlled experiments
at these same scales. Given the virtual impossibility of such a task, perhaps
the best hope is to document pesticide-related patterns of declines in nature
at large geographic scales and then identify the most likely mechanism(s)
using smaller, more feasible experimental scales. This is the approach that
has been employed by amphibian biologists during the past decade.
The most extensive regional data examining patterns of declines come
from the western United States where a number of amphibian species are
experiencing population declines across a large regional scale. By combining
detailed records of annual pesticide use in the Central Valley of California
with surveys of hundreds of amphibian populations in the Sierra Nevada
Mountains, Davidson et al. (2001, 2002) found a positive correlation between
population declines and the amount of agricultural land that is upwind.
Subsequently, Davidson (2004) found a correlation between declining popula-
tions and the actual amount of pesticides that are applied upwind (Figure 1).
Amphibians in these mountain sites contain pesticides in their tissues and
reduced acetylcholine esterase activity (i.e., a signal of insecticide exposure;
Datta et al. 1998, Sparling et al. 2001), further strengthening the hypothesis
Threatened Amphibians of the World
that pesticides may indeed be the underlying cause. If pesticides are a cause
of amphibian declines, we need to consider the possible mechanisms by
which this could occur.
Mechanism #1: Direct toxicity
The most obvious mechanism by which pesticides could cause amphibian
declines is via direct toxicity. There are two key issues concerning this mecha-
nism. First, because very high concentrations of pesticides are almost always
lethal to amphibians, we must consider the actual concentrations that an
amphibian would experience in nature. Second, to cause a population decline,
the amount of death due to the pesticide would have to be over and above
natural causes of death experienced by amphibians (which can be >90% for
amphibian larvae; reviewed in Relyea 2007). For some pesticides including
the herbicide Roundup® (Monsanto Corp., St. Louis, MO), researchers have
observed 70-100% mortality of larval amphibians at concentrations that have
been observed in nature (Relyea 2005; Relyea et al. 2005). However, for many
of the pesticides that have been tested, including some of the most common
insecticides (e.g., carbamates and organophosphates), the concentrations
that cause amphibian death are above those concentrations that are most
commonly found in nature (Boone and Semlitsch 2001; Relyea 2004) although
seasonal spikes in pesticide use may make these rare, high concentration
events quite important to amphibian populations. In either case, it is clear
that we should also consider alternative mechanisms for how pesticides
might cause amphibian declines.
Mechanism #2: Sublethal effects on amphibian
behaviour, physiology, and endocrinology
One of the most striking discoveries in recent years is that pesticides can have
some surprising and unintended effects on amphibians at very low, sublethal
concentrations. Analogous to the side effects of human drugs, pesticides are
now known to alter animal behavior in ways that can reduce feeding, impair
locomotion, and weaken predator avoidance (Weis et al. 2001). Pesticides can
also compromise the immune system, making amphibians more susceptible to
parasites and pathogens (Christin et al. 2003). Moreover, there appears to be
a correlation between the frequency of amphibian malformations (e.g., extra
or missing limbs) and the proximity of agricultural land (Taylor et al. 2005).
Perhaps the most striking discovery is that some pesticides (e.g., atrazine) at
very low concentrations (i.e., within drinking water standards) can mimic sex
< 0.2 kg/m 0.2 - 5.2 kg/m
69% present 45% present
hormones and cause adult male frogs to become feminized and grow eggs in
their testes (Hayes et al. 2002; Hayes 2004). While these effects are certainly
dramatic, it is currently unclear as to whether they can subsequently result
in amphibian population declines.
Mechanism #3: Synergistic effects between
pesticides and other stressors
One way in which low concentrations might impact amphibian populations
is by interacting with other natural stressors and becoming more lethal than
we appreciate. Traditional pesticide testing is done on a single species
under laboratory conditions that typically lack any natural environmental
stressors. In contrast, there are wide variety of stressors that occur in nature
including elevated temperature, reduced pH, increased ultraviolet radiation,
competition, and the threat of predation. To date, very few pesticides have
been tested on amphibians in combination with different natural stressors,
but the results thus far have been quite surprising; insecticides can become
more lethal at higher temperatures (Boone and Bridges 1999) and more
lethal when combined with the smell of predators in the water (Relyea
2003; Figure 2). The physiological mechanisms underlying these synergistic
interactions are not yet known, but it is clear that natural stressors can
make seemingly sublethal concentrations of pesticides become highly
lethal to amphibians.
Mechanism #4: Indirect effects through
a food web
Amphibians live in diverse ecological communities with a network of interac-
tions, yet traditional pesticide testing is conducted on single species in the
laboratory. As a result, we know relatively little about how low concentra-
tions of pesticides that cannot directly kill amphibians might indirectly affect
amphibians via changes in the food webs that contain amphibians. These
effects can initially be positive if pesticides reduce or remove populations of
amphibian predators whereas the changes can be negative when pesticides
affect the food available to amphibians (Boone and Semlitsch 2001; Relyea
2005). Reduced food availability commonly reduces amphibian growth which
can have long-term effects on the time to reproduction (i.e. generation time),
future survival, and reproductive output (i.e., the number of eggs that can be
laid; Relyea 2007). However, we do not yet know whether these indirect food
web effects impact the population sizes of amphibians.
5.2 - 99 kg/m > 99 kg/m
32% present 17% present
Figure 1. Statewide patterns of declines in populations of five amphib-
ian species (Rana muscosa, R. boylii, R. cascadae, R. draytonii and Bufo
canorus) throughout California and the amount of pesticide applied
upwind of each site. The general pattern suggests a connection between
pesticide use and amphibian declines (figure courtesy of Carlos Davidson
based on data from Davidson 2004).

Conclusions
Confirming the role of pesticides in the global decline of amphibians is an
extremely daunting task, but it is being rapidly addressed by researchers. Since
realizing that amphibians were declining globally, we have been accumulating
additional regional datasets on amphibian populations to assess their status
and to test for correlative patterns with regional pesticide use. At the same
time, experimentalists are rapidly delving into the wide range of possible
mechanisms whereby pesticides might affect amphibians and discovering
some mechanisms that were inconceivable only a decade ago. Unfortunately,
most pesticide research is being conducted in North America and Europe on a
small fraction of the 6,000-odd amphibian species that exist (and mostly on the
larval stage), making our current power of inference rather limited (Schiesari
et al. 2007). If pesticides are contributing to amphibian declines in nature, we
will continue to require a multi-faceted approach from many sources of data
to develop a solid case. A solid case is critical because it must withstand the
doubtless challenges that will follow from the manufacturers and applicators
of pesticides around the world. Importantly, while scientists can provide the
data demonstrating the impacts of pesticides on amphibians, only societies can
determine whether or not demonstrated pesticide impacts are acceptable.
Rick Relyea
References
Alford, R.A. and Richards, S.J. 1999. Global amphibian declines: A problem in applied
ecology. Annual Review of Ecology and Systematics 30:133-165.
Boone, M.D. and Bridges, C.M. 1999. The effect of temperature on potency of carbaryl
for survival of tadpoles of green frog (Rana clamitans). Environmental Toxicology and
Chemistry 18:1482-1484.
No pesticide
100%
80%
60%
Survival
40%
20%
0%
0 4 8 12 16
Day
ESSAY 4.5. CHYTRIDIOMYCOSIS: DRIVER OF AMPHIBIAN DECLINES AND EXTINCTIONS
Amphibian populations are declining at alarmingly rapid rates worldwide
and species are disappearing from entire regions. A greater proportion of
amphibians are at imminent risk of extinction than any other animal class,
including birds. Some have already been lost forever. While habitat loss,
overexploitation and the other “usual suspects” are partly to blame, the
factor driving amphibians most rapidly to extinction is the emergence of the
virulent fungal disease, chytridiomycosis. Chytridiomycosis is caused by a
fungus in the order chytridiales (the chytrid fungi) called Batrachochytrium
dendrobatidis. This fungus was unknown until 1998, when it was simultane-
ously discovered as the primary cause of catastrophic amphibian declines in
the rain forests of Australia and Central America (Berger et al. 1998). While
many chytrid fungi are parasites of invertebrates, B. dendrobatidis is the only
chytrid fungus known to infect vertebrates. Although the fungus is known
only to infect amphibians, it has low host specificity having thus far been
recorded infecting at least 14 families and over 100 species of amphibian
on five continents.
Although B. dendrobatidis was unknown until the late 1990s, retrospective
studies have shown it to have been a major cause of amphibian declines at
least as far back as the 1970s. Many amphibian declines considered to have
been due to other reasons at the time, such as predation or competition by
introduced species, or stress and disruption by human activities, are now
known to have been due – at least in part - to chytridiomycosis (Green and
Sherman 2001; Green et al. 2002; Muths et al. 2003; Carey et al. 2006).
The primary hypothesis for the emergence of chytrid-caused declines
is the exposure of naïve host populations to a newly introduced pathogen,
although other hypotheses (such as a change in a pre-existing host-patho-
gen relationship) have not yet been discounted (Cunningham et al. 2003;
Weldon et al. 2004). Some amphibian species, such as the North American
Chapter 4. The State of the World’s Amphibians
Boone, M.D. and Semlitsch, R.D. 2001. Interactions of an insecticide with larval den-
sity and predation in experimental amphibian communities. Conservation Biology
15:228-238.
Christin M.S., Gendron A.D., Brousseau P., Menard L., Marcogliese D.J., Cyr D., Ruby S.
and Fournier M. 2003. Effects of agricultural pesticides on the immune system of
Rana pipiens and on its resistance to parasitic infection. Environmental Toxicology
and Chemistry 22:1127-1133.
Datta, S., Hansen, L., McConnell, L., Baker, J., LeNoir, J. and Seiber, J.N. 1998. Pesticides
and PCB contaminants in fish and tadpoles from the Kaweah River basin, California.
Bulletin of Environmental Contamination and Toxicology 60:829-836.
Davidson, C. 2004. Declining downwind: Amphibian population declines in California and
historical pesticide use. Ecological Applications 14:1892-1902.
Davidson, C., Shaffer, H.B. and Jennings, M.R. 2001. Declines of the California red-legged
frog: Climate, UV-B, habitat, and pesticides hypotheses. Ecological Applications
11:464-479.
Davidson, C., Shaffer, H.B. and Jennings, M.R. 2002. Spatial tests of the pesticide drift,
habitat destruction, UV-B, and climate-change hypotheses for California amphibian
declines. Conservation Biology 16:1588-1601.
Hayes, T.B. 2004. There is no denying this: Defusing the confusion about atrazine.
Bioscience 54:1138-1149.
Hayes, T.B., Collins, A., Lee, M., Mendoza, M., Noriega, N., Stuart, A.A. and Vonk, A. 2002.
Hermaphroditic, demasculinized frogs after exposure to the herbicide atrazine at low
ecologically relevant doses. Proceedings of the National Academy of Sciences of the
United States of America 99:5476-5480.
Relyea, R.A. 2003. Predator cues and pesticides: A double dose of danger for amphibians.
Ecological Applications 13:1515-1521.
Relyea, R.A. 2004. Synergistic impacts of malathion and predatory stress on six species of
North American tadpoles. Environmental Toxicology and Chemistry 23:1080-1084.
Relyea, R.A. 2005. The impact of insecticides and herbicides on the biodiversity and
productivity of aquatic communities. Ecological Applications 15:618-627.
Relyea, R.A. 2007. Getting out alive: How predators affect the decision to metamorphose.
1.6 mg/l of carbaryl
100%
80%
60%
Survival
40%
20%
0%
0 4 8 12 16
Day
No predator odors Predator odors
Bullfrog (Rana catesbeiana) (Daszak et al. 2004) and the African Clawed
Frog (Xenopus laevis) (Weldon et al. 2004), are generally unaffected by B.
dendrobatidis infection and there has been speculation these species might
act as transport hosts, introducing the infection to new areas and to new host
species and populations (Mazzoni et al. 2003; Weldon et al. 2004; Garner
et al. 2006). The global trade in amphibians, which began in the 1930s with
the trade in Xenopus laevis for pregnancy testing, mushroomed in the 1970s
and 1980s with the widespread harvesting of wild frogs in India, Indonesia
and China for the international restaurant trade. The level of international
trade continues to rise with the development of commercial frog farms in
Asia and Latin America that export bullfrogs and other ranid species for the
North American and European markets. There is evidence that this trade
might be a major driver of the current chytridiomycosis pandemic (Mazzoni
et al. 2003; Weldon et al. 2004; Garner et al. 2006; Kusrini and Alford 2006;
Cronin et al. in press).
Infection with B. dendrobatidis is limited to the keratinized tissues of the
animal: the most superficial, keratinized, cells of the skin in metamorphosed
amphibians, and the mouthparts of anuran larvae (Daszak et al. 1999; Berger
et al. 2005). The infection is intracellular (i.e. the fungus grows within the
skin cell) and discharge tubes open onto the surface of the cell for the
release of zoospores, the infective stage of the fungus. The presence of the
fungus induces a number of host responses, including, in metamorphosed
animals, hyperkeratosis, irregular foci of hyperplasia, disordered epidermal
cell layers, spongiosis, erosion and ulceration of the skin. Areas of the skin
more frequently in close contact with the water, such as the digits and the
pelvic patch, are more severely affected (Berger et al. 1998). Inflammatory
responses in the dermis are mild, if at all present, and involve mononuclear
cells. Inflammation within the epidermis is rare. The lack of an immune
49
Oecologia 152:389-400.
Relyea, R.A., Schoeppner, N.M. and Hoverman, J.T. 2005. Pesticides and amphibians: The
importance of community context. Ecological Applications 15:1125-1134.
Schiesari, L., Grillitsch, B. and Grillitsch, H. 2007. Biogeographic biases in research
and their consequences for linking amphibian declines to pollution. Conservation
Biology 21:465-471.
Sparling, D.W., Fellers, G.M. and McConnell, L.L. 2001. Pesticides and amphibian
population declines in California, USA. Environmental Toxicology and Chemistry
20:1591-1595.
Stuart, S.N., Chanson, J.S., Cox, N.A., Young, B.E., Rodrigues, A.S.L., Fischman, D.L.
and Waller, R.W. 2004. Status and Trends of Amphibian Declines and Extinctions
Worldwide. Science 306:1783-1786.
Taylor, B., Skelly, D., Demarchis, L.K., Slade, M.D., Galusha, D. and Rabinowitz, P.M. 2005.
Proximity to pollution sources and risk of amphibian limb malformation. Environmental
Health Perspectives 113:1497-1501.
U.S.E.P.A. 1997. Pesticides Industry Sales and Usage. Report # 733-R-97-002. Wash-
ington, DC. USA.
Weis, J.S., Smith, G., Zhou, T., Santiago-Bass, C. and Weis, P. 2001. Effects of con-
taminants on behavior: Biochemical mechanisms and ecological consequences.
Bioscience 51:209-217.
Figure 2. The synergistic impacts of an insecticide (carbaryl) on bullfrog
tadpoles when combined with the presence or absence of predator odours
in the water. When no insecticide is present, there is low mortality with
or without the odour of predators. When the insecticide is present at high
concentrations (6.5 mg/L) there is high mortality with or without the odour of
predators. However, at intermediate concentrations (0.6 mg/L), there is low
mortality in the absence of predator odours, but 46 times higher mortality in
the presence of predator odours.
6.5 mg/l of carbaryl
100%
80%
60%
Survival
40%
20%
0%
0 4 8 12 16
Day
response might be due to a lack of stimulation of the host immune system
as the site of infection is both superficial and intracellular. Also, the degree
of tissue damage is very low. There is no evidence of immuno-suppression
either as an underlying factor to, or as a consequence of, B. dendrobatidis
infection (Berger et al. 1999).
Signs of the disease include lethargy, loss of righting reflex and abnormal
posture. Gross lesions are usually not apparent, but increased epidermal
sloughing, epidermal ulceration and reddening (hyperaemia) of the digital
and ventral skin have been reported (Daszak et al. 1999; Berger et al. 2000).
Diagnosis is by visualization of the characteristic intracellular zoosporangia
in the keratinized epidermis using either cytology of sloughed skin or
histopathology of skin samples (e.g., toe-clip biopsies or post-mortem skin
samples). Molecular techniques, such as the polymerase chain reaction (PCR)
and quantitative PCR, have largely superseded cytology and histopathology
for rapid diagnosis and for screening large numbers of animals.
Despite almost a decade of research on chytridiomycosis, there is still
much that remains unknown. The exact mechanism by which B. dendroba-
tidis causes mortality is not known, but there are currently three possible
mechanisms hypothesized. Firstly, the chytrid might release toxins, such
as proteolytic enzymes, which are absorbed to cause systemic poisoning.
Secondly, epidermal hyperplasia may impair cutaneous respiration and/or
osmoregulation (Berger et al. 1998, 2005). Thirdly, a combination of both of
these mechanisms might be involved (Daszak et al. 1999). The absence of any
specific internal lesions suggests that the ultimate cause of death probably
is either metabolic or toxic (Berger et al. 1999).
Although several life-history (e.g., low fecundity) and ecological (e.g.,
riparian, montane habitat) factors have been shown to correlate with popula-
tion declines due to chytridiomycosis, these might be solely indicators of a
50 Threatened Amphibians of the World

Batrachochytrium dendrobatidis infection in the superficial, keratinized layer
of skin sloughed from an adult North American bullfrog. Note focal thicken-
ing of the sloughed skin with numerous subspherical empty sporangia. One
sporangium still contains zoospores (arrow). © Andrew Cunningham, ZSL.
population’s ability to respond to any stochastic factor (amphibians in riparian,
montane habitats tend to have small and restricted ranges). Infection with B.
dendrobatidis is highly variable between species: some species (e.g., Rana
catesbeiana, Xenopus laevis) become infected, but show no evidence of
disease, while others (e.g., Atelopus spp.) appear to be highly susceptible
to infection, with very high mortality rates. However, within a species,
there are differences in response, depending on variables such as dose of
exposure, temperature and humidity. Therefore, a chytrid-positive population
might appear to be resistant, only to decline markedly following a change
in environmental conditions.
Addressing the threat of chytridiomycosis is not easy. We are used to
tackling species threats by introducing or increasing protection measures
of the animals and/or their habitats. Pathogens respect neither the law nor
protected areas. In fact, many of the most devastating effects of chytridio-
mycosis have occurred in pristine and protected areas, such as rainforest
reserves in Australia and Central America. Although B. dendrobatidis can
be killed by a range of antifungal medications, it is both impractical and
ecologically dangerous to attempt such treatments in the wild. The disease
and causative agent were unknown until less than 10 years ago and there is
still much to learn about the biology, ecology and distribution of the patho-
gen. Currently, our only defense is to bring vulnerable species into captivity
until alternative approaches are devised. These are massive challenges to
the zoo and conservation research communities. If we are to avoid losing a
substantial proportion (the majority?) of the world’s amphibians in the face
of this daunting threat, these communities need to raise their game to a new
level of cooperation along the lines of a Manhattan project-style response
for amphibian conservation.
Andrew A. Cunningham and Peter Daszak
References
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amphibian chytrid Batrachochytrium dendrobatidis. Diseases of Aquatic Organisms
68:51-63.
Berger, L., Speare, R., Daszak, P., Green, D.E., Cunningham, A.A., Goggin, C.L., Slocombe,
R., Ragan, M.A., Hyatt, A.D., McDonald, K.R., Hines, H.B., Lips, K.R., Marantelli, G.
and Parkes, H. 1998. Chytridiomycosis causes amphibian mortality associated with
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Berger, L., Speare, R. and Hyatt, A.D. 1999. Chytrid fungi and amphibian declines: overview,
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Daszak, P., Berger, L., Cunningham, A.A., Hyatt, A.D., Green, D.E. and Speare, R. 1999.
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Daszak, P., Strieby, A., Cunningham, A.A., Longcore, J.E., Brown, C.C. and Porter, D.
2004. Experimental evidence that the bullfrog (Rana catesbeiana) is a potential car-
rier of chytridiomycosis, an emerging fungal disease of amphibians. Herpetological
Journal 14:201-207.
Garner, T.W.J., Perkins, M.W., Govindarajulu, P., Seglie, D., Walker, S.F., Cunningham,
A.A. and Fisher, M.C. 2006. The emerging amphibian pathogen Batrachochytrium
dendrobatidis globally infects introduced populations of the North American bullfrog,
Rana catesbeiana. Biology Letters 2:455-459.
Green, D.E., Converse, K.A. and Schrader, A.K. 2002. Epizootiology of sixty-four amphib-
ian morbidity and mortality events in the USA, 1996-2001. Annals of the New York
Academy of Science 969:323-339.
Green, D.E. and Sherman, C.K. 2001. Diagnostic histological findings in Yosemite toads
(Bufo canorus) from a die-off in the 1970s. Journal of Herpetology 35:92-103.
Kusrini, M.D. and Alford, R.A. 2006. Indonesia’s Exports of Frogs’ Legs. TRAFFIC Bulletin
21:13-24.
Mazzoni, R., Cunningham, A.C., Daszak, P., Apolo, A., Perdomo, E. and Speranza, G.
2003. Emerging pathogen of wild amphibians in frogs (Rana catesbiana) farmed for
international trade. Emerging Infectious Diseases 9:995-998.
Muths, E., Corn, P.S., Pessier, A.P. and Green, D. E. 2003. Evidence for disease-related
amphibian decline in Colorado. Biological Conservation 110:357-365.
Weldon, C., du Preez, L.H., Hyatt, A.D., Muller R. and Speare, R. 2004. Origin of the amphib-
ian chytrid fungus. Emerging Infectious Diseases 10:2100-2105.

ESSAY 4.6. CLIMATE CHANGE AND AMPHIBIAN DECLINES

Evidence that climate change is currently affecting life on earth has become
overwhelming in the last few years. Although some continue to question the
link between global warming and human activity, the rate of current trends in
temperature shifts is vastly different from past global climate change events.
For example, in lower elevation cloud forests in the Andes, the warming trend
experienced from the 1970s to the present is 18 times faster than at the end
of the last ice age (Bush et al. 2004). Species of plants and animals in many
different taxonomic groups and regions of the world are already responding
to warming, by shifting their ranges uphill or towards the poles and tracking
cooler environments (Walther et al. 2002). The timing of seasonal cycles, such
as flowering in plants and migration in birds, has also been shifting with the
changing climate (Root et al. 2003). Spawning dates and breeding migra-
tions in a number of amphibian species have already become significantly
earlier (Root et al. 2003). Although these changes might appear trivial, they
do alter the composition and dynamics of communities, with complex and
unpredictable outcomes. For example, in Britain, newts are now arriving
much earlier at breeding ponds, allowing their predatory larvae to be much
larger by the time frogs arrive. This may be detrimental for frog species that
now deposit their eggs and tadpoles into pools full of hungry newt larvae
(Beebee 1995). Along with changes in species’ ranges and phenology, current
models predict a great loss of species in the future due to a warmer and more
extreme climate (Thomas et al. 2004). In fact, climate change may already
be causing extinctions.
Declines and disappearances of amphibian populations have become
increasingly apparent in the past 20 years. Some declines appear to be
explained by clear anthropogenic effects such as habitat loss, while others,
with less apparent causes, have been classified as “enigmatic” (Stuart et
al. 2004). Research activity has increasingly focused on determining the
causes of these declines. High-elevation species seem to be most affected,
and based on the fate of known species it appears that many have probably
disappeared prior to being described. In the Neotropics, harlequin frogs of the
genus Atelopus appear to have been most affected by “enigmatic declines”.
A number of undescribed Atelopus species collected by South American
herpetologists are currently awaiting description. In the short time since these
species were collected, scientists have returned to their original collection
sites only to find that most of these undescribed species have vanished
from their apparently pristine environments. Many more species are likely
to have become extinct before even being collected. For instance, in many
countries, most historical collection sites are directly adjacent to a highway
or rural road, while vast tracts of land remain almost entirely unsurveyed. This
makes it likely that many species unknown to science have already been lost.
The situation is even worse when we take molecular evidence into account. Atelopus zeteki (Critically Endangered) from El Cope, Panama, taken about nine months before the population crash happened at that site. © Ross A. Alford
Cryptic species that looked extremely similar to each other were probably
hidden among some of the widespread species. We may never know how
many species we have truly lost.
Could amphibian disappearances be attributed to global warming pat- 2005). As with humans, a synergistic interaction between temperature change 1). The probability of this correlation occurring by chance is less than one in
terns? Skeptics have voiced critical opinions on this idea. Amphibians have and infectious disease may explain many of the recent disappearances of a thousand. In the laboratory, B. dendrobatidis grows best at temperatures
been on earth for at least 280 million years, persisting through at least 60 amphibians. One disease for which this synergism may be important is the between 17 and 25°C, but grows more slowly outside this range, and may
glacial periods in the last 12 million years alone. So why would the recent chytrid fungus Batrachochytrium dendrobatidis, which infects the skin of not cause disease at temperatures over 30°C (Piotrowski et al. 2004). Even
temperature shifts be more threatening to amphibians than the ever-changing amphibians, causing the disease chytridiomycosis. Outbreaks of this disease short periods at body temperatures of 37°C can clear frogs of infections by
environment they evolved in? The answer may be found in the interaction have occurred in apparently naïve populations, and are clearly linked to the the pathogen (Woodhams et al. 2003). Such temperatures may be experienced
between climate change and another variable – disease. extirpation of frog communities in the immediate range of harlequin frogs by frogs after a short period in direct sunlight, but cannot be attained under
Climate change is already impacting human health by facilitating the (Lips et al. 2006). Approximately 80% of missing harlequin frog species heavy cloud cover. Although it has been suggested that the probability of
spread of disease, causing an estimated 150,000 deaths a year (Patz et al. vanished directly after a relatively warm year (Pounds et al. 2006; see Figure population declines increases with altitude, this does not seem entirely
 Chapter 4. The State of the World’s Amphibians 51

true for harlequin frogs. Pounds et al. (2006) demonstrate that patterns of
extinction in harlequin frogs vary at different altitudes, and suggest that they
are caused by interactions among temperature, atmospheric moisture, cloud
cover, and host-pathogen dynamics. This hypothesis could be tested in other
regions of the world, such as Australia, where the patterns of declines in
amphibian populations show many parallels with those in the Neotropics.
In eastern Queensland, Australia, at least 14 species have experienced
Tyler, M.J. 2004. Effect of season and temperature on mortality in amphibians due to
0,8 – – 12
chytridiomycosis. Australian Veterinary Journal 82:434-439.
Bush, M.B., Silman, M.R. and Urrego, D.H. 2004. 48,000 years of climate and forest change
in a biodiversity hot spot. Science 303:827-829. – 10
Lips, K.R., Brem, F., Brenes, R., Reeve, J.D., Alford, R.A., Voyles, J., Carey, C., Livo, L.,
0,6 –
Pessier, A.P. and Collins, J.P. 2006. Emerging infectious disease and the loss of biodiver-
sity in a Neotropical amphibian community. Proc Natl Acad Sci USA 103:3165-3170. –8

declines in recent decades, with six species disappearing altogether, and McDonald, K.R., Méndez, D., Müller, R., Freeman, A.B., and Speare, R. 2005. Decline in

others becoming locally extinct at high-elevation sites. Batrachochytrium
dendrobatidis has been implicated in most of these declines. In these areas,
where B. dendrobatidis now appears to be endemic, patterns of infection
and mortality are consistent with the climate-linked epidemic hypothesis,
with infection levels and mortality highest during cooler months (Berger et
al. 2004; Retallick et al. 2004; McDonald et al. 2005; Woodhams and Alford
2005). Future predictions for amphibians in the region are grim, with 13 to
68% of frog species endemic to Queensland predicted to become extinct under
future climate scenarios (Thomas et al. 2004). This prediction is based on the
effects of changing climatic conditions alone. The true impacts of climate
change may be far more complex and potentially detrimental when acting
in synergism with disease.
What seems to be clear is that amphibian extinctions are strongly cor-
related with the changing climate. The disappearance of frogs may be an
early warning of a new episode of mass extinction, and this time, there is
no cataclysm to blame, only ourselves.
Robert Puschendorf, Ross A. Alford, and Jodi J.L. Rowley
References
Beebee, T.J.C. 1995. Amphibian breeding and climate change. Nature 374:219-220.
Berger, L., Speare, R., Hines, H.B., Marantelli, G., Hyatt, A.D., McDonald, K.R., Skerratt,
L.F., Olsen, V., Clarke, J.M., Gillespie, G., Mahony, M., Sheppard, N., Williams, C. and
Number of species
Tropical AT (ºC)
the prevalence of chytridiomycosis in frog populations in North Queensland, Australia.
0,4 – –6
Pacific Conservation Biology 11:114-120.
Patz, J.A., Campbell-Lendrum, D., Holloway, T. and Foley, J.A. 2005. Impact of regional
climate change on human health. Nature 438:310-317. –4
Piotrowski, J., Annis, S. and Longcore, J.E. 2004. Physiology of Batrachochytrium dendro- 0,2 –
batidis, a chytrid pathogen of amphibians. Mycologia 96:9-15.
–2
Pounds, J.A., Bustamante, M.R., Coloma, L.A., Consuegra, J.A., Fogden, M.P., Foster, P.N.,
La Marca, E., Masters, K.L., Merino-Viteri, A., Puschendorf, R., Ron, S.R., Sanchez-
Azofeifa, G.A., Still, C.J. and Young, B.E. 2006. Widespread amphibian extinctions 0– –0
from epidemic disease driven by global warming. Nature 439:161-167. 1982 1985 1988 1991 1994
Retallick, R.W.R., McCallum, H. and Speare, R. 2004. Endemic infection of the amphibian
chytrid fungus in a frog community post-decline. PLoS Biology 2:1965-1971. Year
Root, T.L., Price, J.T., Hall, K.R., Schneider, S.H., Rosenzweig, C. and Pounds, J.A. 2003.
Fingerprints of global warming on wild animals and plants. Nature 421:57-60.
Stuart, S.N., Chanson, J.S., Cox, N.A., Young, B.E., Rodrigues, A.S.L., Fischman, D.L. Figure 1. Last year of observation in harlequin frog species for which there
and Waller, R.W. 2004. Status and Trends of Amphibian Declines and Extinctions is sufficient population data (red line) and air temperature for the tropics (ºC)
Worldwide. Science 306:1783-1786. of the preceding year (blue line). Reproduced, with permission of authors,
Thomas, C.D., Cameron, A., Green, R.E., Bakkenes, M., Beaumont, L.J., Collingham, Y.C., from Pounds et al. 2006 (Nature 439:161-167).
Erasmus, B. F. N., de Siquera, M.F., Grainger, A., Hannah, L., Hughes, L.S., Huntley,
B., van Jaarsveld, A.S., Midgley, G.F., Miles, L., Ortega-Huerta, M.A., Peterson, A.T.,
Phillips, O.L. and Williams, S.E. 2004. Extinction risk from climate change. Nature Woodhams, D.C., Alford, R.A. and Marantelli, G. 2003. Emerging disease of amphibians
427:145-148. cured by elevated body temperature. Diseases of Aquatic Organisms 55:65-67.
Walther, G.-R., Post, E., Convey, P., Menzel, A., Parmesank, C., Beebee, T.J.C., Fromentin, Woodhams, D.C. and Alford, R.A. 2005. The Ecology of Chytridiomycosis in Rainforest
J.-M., Hoegh-Guldberg, O. and Bairlein, F. 2002. Ecological responses to recent climate Stream Frog Assemblages of Tropical Queensland. Conservation Biology 19:1449-
change. Nature 416:389-395. 1459.

ESSAY 4.7. THE STATUS OF THE CHINESE GIANT SALAMANDER ANDRIAS DAVIDIANUS

The Chinese Giant Salamander Andrias davidianus, is the largest living amphib- Reserve. Three canoes with over 150 bow-hooks were seen in the main river The Chinese Giant Salamander Andrias davidianus (Critically Endangered) is
ian, with adults reaching a total length of more than 200cm and a weight of of Yongshun Chinese Giant Salamander reserve of Hunan Province. the largest living amphibian, with adults reaching a total length of more than
50kg (Peng et al. 1998). Although very similar to its congener, the Japanese Giant As the Chinese Giant Salamander has a great economic value mostly as 200cm and a weight of 50kg. © Kejia Zhang
Salamander Andrias japonicus, the Chinese Giant Salamander differs in the ar- luxury food, this species has long been excessively exploited by local people.
rangement of tubercles on the head and throat, which are mostly in pairs, and Because of the high demand in some cities such as Guangzhou and other
much smaller and fewer than those of its Japanese cousin. The snout is also less provincial capitals, the value of salamander meat is increasing steadily in
rounded and the tail a little longer in the Chinese species. The species is referred local black markets. The price in China has risen from around US$5 per kg
to locally as wawayu (baby fish), because its call resembles a baby’s cry. in the early 1980s to over US$350 in recent years. High prices are strong
Formerly widespread in China, most local wild populations of the Chinese Gi- incentives for harvesting salamanders from the wild, and combating illegal
ant Salamander have declined greatly in both range and number since the 1950s, harvesting and trade has become extremely difficult.
largely due to habitat fragmentation or loss, and as a result of overharvesting for In order to protect the wild populations of Chinese Giant Salamander, 21
human consumption. Recent surveys show that the Chinese Giant Salamander nature reserves have been established in Henan, Jiangxi, Chongqing, Shaanxi,
is distributed in a few mountainous areas in the middle and lower tributaries of Guizhou, Guangdong, and Hunan Provinces since the 1980s (and see Table 1),
the Yangtze, Yellow and Pearl Rivers, across the 17 provinces of Henan, Shaanxi, but many of these reserves have very limited funds available to effectively
Shanxi, Gansu, Qinghai, Sichuan, Chongqing, Guizhou, Hubei, Anhui, Zhejiang, conserve the salamanders. Sadly, because of funding difficulties four of these
Jiangxi, Hunan, Fujian, Yunnan, Guangdong and Guangxi (Wang et al. 2004). reserves have now been degazetted.
Many populations are now on the verge of extinction, and some are already To date, about 100 legal Chinese Giant Salamander farms have been
extinct (Zhao 1998; Zhang and Wang 2000). Today, because of this continuing, established throughout the species’ range. Most of them were set up during
drastic, population decline the Chinese Giant Salamander is listed as Critically the 1990s, directly aimed at supplying the ever-growing food market, and more
Endangered on the IUCN Red List of Threatened Species. than 20 farms have bred the giant salamanders and successfully reared young
Although habitat loss is certainly a threat, uncontrolled harvesting is the primary to maturity. During 2005, over 32,000 larvae were bred from more than 6,000
reason for the steep decline in numbers of Chinese Giant Salamander. Before parents in seven farms. But, at present, even sustainable breeding programmes
the wildlife protection act was declared by the Chinese government in 1988, the are insufficient to meet market demands. To date, only a few restaurants in
Chinese Giant Salamander was a legal fishery product in China. Indeed, there was Beijing, Hunan and Guangdong have been approved by the government to sell
already a marked decline in the number of salamanders being collected from the Chinese Giant Salamander, with the result that most trade of this animal has
wild by the 1970s. For example, a traditional company in Taibai county of Shaanxi been driven to the black market (Zhang, pers. obs.). In addition, it certainly seems
province reported that, while demand remained high, the quantity of Chinese possible that a number of farms have obtained adult salamanders illegally from
Giant Salamander purchased declined from 3,813kg in 1973 to 1,300kg in 1979, the wild, and this in itself may pose a threat to remaining wild populations.
presumably in response to a reduction in wild stocks. Likewise, from the 1950s to
the 1980s, the amount purchased of this species had declined more than 80% in
some areas in Hunan and Anhui Provinces (Wang et al. 2004). In addition, whereas Table 1. Chinese Giant Salamander reserves in China with location, grade, area and year established (updated from Wang et al. 2004).
the salamander was found in 28 counties of north-eastern Sichuan Province in the
1970s, it appears to have disappeared from 11 of them by the middle of the 1980s.
Western Hunan Province had 10 counties with salamanders in the 1960s, but the Reserve Location Grade Area (ha) Established year
species could only be found in five of them in the 1980s (Liu 1989). Lushi Giant Salamander Reserve Lushi, Henan Provincial 184,350 1982
Despite protection by the Chinese Government since 1988 (the Chinese Xixia Giant Salamander Reserve Xixia, Henan Provincial 131,040 1982
Government has declared the giant salamander a Class II Protected Species), Qingyaoshan Giant Salamander Reserve Xinan, Henan Provincial 9,000 1988
illegal collection of Chinese Giant Salamander from the wild is still a lead- Songxian Giant Salamander Reserve Songxian, Henan Provincial 600 1998
ing threat widely contributing to population declines, and in many counties Luanchuan Giant Salamander Reserve Luanchuan, Henan Provincial 800 1996
the poaching has actually increased during recent years. Hunters now use Youyang Giant Salamander Reserve Youyang, Chongqing County 4,000 1989
pesticides, electrical shock tools, explosives, and other modern methods, Taibai Xushuihe Giant Salamander Reserve Taibai, Shannxi Provincial 5,740 1986
instead of traditional bow-hooks to capture salamanders. The increasing Lingkou Giant Salamander Reserve Luonan, Shannxi Provincial 5,715 1998
use of such unsustainable methods has resulted in very serious declines in Zhuxi Giant Salamander Reserve Zhuxi, Hubei Provincial 800 1986
salamander populations. Furthermore, the species is in danger of losing its Zhongjianhe Giant Salamander Reserve Xianfeng, Hubei Provincial 264 1994
title of “world’s largest amphibian”, as the weights of giant salamanders being Dachenshan Giant Salamander Reserve Loudi, Hunan County 100 1987
captured today appears to be much lower than formerly, presumably due to Sangzhi Quanhe Giant Salamander Reserve Sanzhi, Hunan County 4,810 1983
the increased hunting pressure. In Sangzhi County in Hunan Province most Yongshun Liangcha Giant Salamander Reserve Yongshun, Hunan - 24,400 1988
individuals weighed c. 35-40kg; all individuals captured in the same locality Chenxi Longmen Giant Salamander Reserve Chengxi, Hunan County 1,700 1984
during the 1970s were <5kg (Liu 1989). According to our field surveys, and Zhangjiajie Giant Salamander Reserve Zhangjiajie, Hunan National level 14,285 1998
questionnaires to local people in the main distributions of the salamander in Qianxi Giant Salamander Reserve Qianxi, Guizhou County 1,000 1986
six provinces from 2000 to 2001, illegal poaching is still prevalent across the Jingan Liaohe Giant Salamander Reserve Jinan, Jiangxi County 100 1980
range, and is even prevalent within protected areas. For example, more than Guishan Giant Salamander Reserve Heyuan, Guangdong Municipal 1,199 2001
100kg of salamanders were collected per year in Dabie mountain area of Anhui Nuoshuihe Giant Salamander Reserve Tongjiang, Sichuan Provincial 9,480 1998
Province (given weights of 0.5-1.0kg, this represents around 7-100 animals), Chuanxihe Giant Salamander Reserve Dayi, Sichuan County - 2004
and the poachers could have captured about 50kg (30-50 animals) worth of Wenxian Giant Salamander Reserve Wenxian, Gansu Provincial 21,160 2004
giant salamander per year in recent years in Hupingshan National Natural
52
The many problems outlined above suggest that wild populations of the
Chinese Giant Salamander are on the verge of extinction, particularly if no
effective conservation measures are taken in the immediate future. The
development and enforcement of a comprehensive conservation strategy
and action plan is now urgently needed to prevent the extinction of this
extraordinary amphibian.
Zhang Ke-jia
and Wang Xiaoming
ESSAY 4.8. AMPHIBIANS AS INVASIVE ALIEN SPECIES
While only a small proportion of the species that are moved around the
world with human movements and global trade actually cause harm, those
that do – termed invasive alien species – are devastating. Such “biological
invasions” are now considered one of the biggest factors in biodiversity
loss and extinctions (Baillie et al. 2004). Many threatened amphibians are
heavily impacted by invasive alien species. However, amphibians themselves
can also be invasive. The list of “100 of the world’s worst invasive alien
species” includes three amphibians, as well as a fungal disease agent, for
which invasive amphibians are vectors of spread (Lowe et al. 2000; and see
www.gisd.org).
Judas’ kiss: the role of invasive amphibians in
the spread of frog chytrid fungus
Amphibian species have been reported as declining since the 1970s,
and in many of them the fungal disease chytridiomycosis is implicated.
Moreover, of the recorded nine recent extinctions of amphibians since
1980, eight were sudden disappearances in suitable habitats and they
are thought to be the result of this disease, probably operating in synergy
with climate change. At first, this may seem to be unrelated to the issue
of invasive species, but in fact, it perfectly illustrates the complexity of
biological invasion issues: not only is the pathogen itself an invasive
alien species, but its spread is facilitated by invasive alien species that
are amphibians!
The origin of the virulent pathogen Batrachochytrium dendrobatidis
is not known with absolute certainty, but a recent study has led to
the conclusion that in all likelihood it originated in Africa. Hence, the
impacts on amphibian populations in the rest of the world result from
the unintentional introductions of a pathogen as an invasive alien spe-
cies. In an ironic twist of fate, several invasive frog species play a role
in the exportation of the pathogen out of Africa, its ‘escape” into the
wild and its ongoing spread and impact on threatened amphibians. It
is thought to have started with the export, out of Africa, of the African
Clawed Frog (Xenopus laevis) for use in laboratories. This frog was a
carrier of chytridiomycosis, and the establishment of feral populations
of this species in countries where it had been moved to, allowed the
pathogen access to other species, such as the American Bullfrog Rana
catesbeiana which now is considered to be one of the main vectors of
the disease (Weldon et al. 2004). The bullfrog, native in parts of the USA
and Canada, is itself a highly invasive species, and has been introduced
to Hawaii, parts of western USA and south-western Canada, Mexico
and the Caribbean, South America, Europe and Asia (GISD 2006a). It has
been traded internationally as a food item, and within countries it can
spread when populations held for culinary use escape and establish.
Frog species in the pet trade are also potential agents of spread of the
disease and infected amphibians have been identified in European and
US zoos. In Australia, chytridiomycosis has been found in Cane Toads
(Bufo marinus), a recently introduced invasive alien species. Chytrid can
persist in contaminated water or mud and it can hence also be spread
by wet or muddy boots, vehicles, cattle and other animals moving among
aquatic sites. Nevertheless, invasive alien frogs and toads have acted in
an (involuntary) ‘Judas’ role, exposing many native species of amphibian
around the world to the pathogen, with disastrous results.
An African Clawed Toad Xenopus laevis (Least Concern) with a fish in its
mouth. Chytridiomycosis was detected in museum specimens of this species
dating back to 1938, and one hypothesis posits that the international trade
in this species may have introduced this fungal disease to other regions of
the world. © Carlos Garin Lobos
Threatened Amphibians of the World
References
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Technology Press, Shanghai, China. [In Chinese]
Wang, X.-M., Zhang, K.-J., Wu, W., Wang, Z.-H., Ding, Y.-Z. and Huang, S. 2004. The decline
of the Chinese giant salamander and applications for its conservation. Conservation
status and distribution of Chinese giant salamander. Oryx 38:197-202.
Peng, K.-M., Fen, R.-P. and Cheng, X.-P. 1998. The Chinese giant salamander. Journal of
Chinese Wildlife 19:11. [In Chinese]
Frog eats frog
The American Bullfrog is not only a vector for disease, but is also a highly
invasive alien species in its own right, affecting native species, amphibian
and otherwise, through competition or direct predation. In British Columbia
(Canada), for example, it has been recorded in competition with, and predat-
ing upon, the Red-legged Frog (Rana aurora), and in Puerto Rico there have
been reports of bullfrogs preying on several bird species, including White-
cheeked Pintail (Anas bahamensis) (GISD 2006a). In the United States, its
impacts include: competition with the Relict Leopard Frog (Rana onca, EN),
which was once thought to be extinct, in Arizona and Nevada (Bradford et
al. 2004); predation on both larval and juvenile Red-legged Frogs in California
(Doubledee et al. 2003); and competition with and predation on the Oregon
Spotted Frog (Rana pretiosa, VU) in Oregon (Pearl et al. 2004).
There goes the neighbourhood
The Caribbean Treefrog, also called the Common Coqui (Eleutherodactylus
coqui) is a relatively small treefrog native to Puerto Rico. While there are
ecological concerns about its introduction into Hawai’i, it also creates a
nuisance to people: it is their loud call that is the main reason that many
Hawai’ians consider the species as a pest. It is feared that the high-pitched
call of the frog may affect the tourism industry and there are also concerns
that property value may be affected due to the high biomass of frogs on
infested sites (Kraus and Campbell 2002; GISD 2006b).
Cane toad conquest
The Cane Toad is native in parts of South and Central America and parts of the
Caribbean. It has been introduced widely throughout the world as a biological
control agent for various insect pests of sugarcane and other crops. Its alien
range now includes many countries, including parts of the Caribbean where
it is not native, and where, for example, it has been implicated in the decline
of the endemic Bermuda Skink (Eumeces longirostris) (Davenport et al. 2001).
It has also been introduced in parts of Asia, Australia and the Pacific. The
Cane Toad has become a major invasive species in most of its introduced
range. It competes with native amphibians and other animals for food and
breeding habitat, and also directly preys on them. In fact, it has been quoted
as eating “almost any terrestrial animal” (Hinkley 1962). In addition, the
toad’s toxicity means that many native predators are killed. For example, in
Australia native snakes have been found dead with the Cane Toad in their
mouths or guts, and in Japan’s remote Iriomote Island, it is feared that the
recent arrival of Cane Toads may result in the poisoning of the threatened
Iriomote Cat (Prionailurus iriomotensis) (GISD 2006c).
In Australia, Cane Toads were introduced to Queensland in 1935 and they
have since expanded their range to encompass more than a million square
kilometres of tropical and subtropical Australia. When researchers radio-tracked
toads at the “invasion front” they found astonishing locomotor performance,
with animals moving up to 1.8km per night during the rainy months. As is the
case with many anurans, toads with longer legs can move faster than those
with shorter legs. This turned out to be happening at a grand scale in Australia:
longer-legged toads were disproportionately common among the first wave of
arrivals at any new site. In general, toads at the invasion front were longer-
The Cane Toad Bufo marinus (Least Concern), a native of Central and South
America, has been widely introduced in the Caribbean, the Philippines, Japan,
parts of Melanesia and the Pacific Islands and Australia. It was introduced
to Australia in 1935 to control sugar cane pests (which it failed to do) from
whence its name derives. © Craig Morley
Ermi, Z. 1998. China Red Data book of Endangered Animal: Amphibia and Reptilia. Science
Press, Beijing, Hong Kong, China, and New York, USA.
Zhang, K.-J. and Wang, X.-M. 2000. Status of conservation biology of Chinese giant
salamander. In: Proceedings of the fourth Asian Herpetological Conference (ed.),
pp. 172. Chengdu, China.
Liu, G.-J. 1989. The Rare and Precious Animal in China---giant salamander. Chinese Journal
of Zoology 24:43-45. [In Chinese.]
legged than toads from older populations, confirming that the invasion process
has been assisted by the evolution of improved dispersal ability among toads
at the front. This means that it would be expected that over many generations,
rates of invasion will be accelerated owing to rapid adaptive change in the
invaders. The annual rate of progress of the toad’s invasion through tropical
Australia has increased about five-fold since the toads first arrived. In other
words, an already strong invader is actually in the process of getting even better
at it (Philips et al. 2006). This should be a salutary reminder that, as with all
alien invasive species, prevention of introduction is the best option, and any
management should be undertaken as soon as possible, before the invader has
had time to evolve into a more dangerous adversary.
Maj de Poorter
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Bradford, D.F., Jaeger, J.R. and Jennings, R.D. 2004. Population status and distribu-
tion of a decimated amphibian, the relict leopard frog (Rana onca). Southwestern
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Davenport, J., Hills, J., Glasspool, A. and Ward. J. 2001. Threats to the Critically Endan-
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and the persistence of California red-legged frogs. Journal of Wildlife Management.
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6th September 2006]
Global Invasive Species Database, 2006b. Bufo marinus. Available from: http://www.
issg.org/database/species/ecology.asp?si=113&fr=1&sts=sss [Accessed 6th
September 2006]
Hinkley, A.D. 1962. Diet of the giant toad, Bufo marinus (L.), in Fiji. Herpetologica
18:253-259.
Kraus, F. and Campbell, E. 2002. Human-mediated escalation of a formerly eradicable
problem: The invasion of Caribbean frogs in the Hawaiian Islands. Biological Inva-
sions 4:327-332.
Lowe, S., Browne, M., Boudjelas, S., and De Poorter, M. 2000. 100 of the World’s Worst
Invasive Alien Species. A selection from the Global Invasive Species Database;
Published by ISSG as special lift-out in Aliens 12, December 2000. 12 pp. http://www.
issg.org/booklet.pdf (English) http://www.issg.org/spanish.pdf (Spanish)
Pearl, C.A., Adams, M.J., Bury, R.B., and McCreary, B. 2004. Asymmetrical effects of
introduced Bullfrogs (Rana catesbeiana) on native ranid frogs in Oregon. Copeia
2004:11-20
Phillips, B.L., Brown, G.P. Webb, J.K., and Shine, R. 2006. Invasion and the evolution of
speed in toads. Nature 439:803.
Weldon C., du Preez, L.H., Hyatt, A.D., Muller, R. and Speare, R. 2004. Origin of the Amphib-
ian Chytrid Fungus. Emerging Infectious Diseases 10:2100-2105.
In recent years, Eleutherodactylus coqui (Least Concern), a species native to Puerto
Rico, has become infamous in the Hawaiian islands, where it probably arrived in
plant pots and bromeliads. There, it has spread widely (the Hawaiian archipelago
has no native amphibians) where they reportedly annoy the local people when they
sing loudly by windows and in gardens, at night. © Alejandro Sanchez
 CHAPTER 5. AMPHIBIANS OF THE AFROTROPICAL REALM

Franco Andreone, Alan Channing, Robert Drewes, Justin Gerlach, are likely to be rare, and have small ranges) are discovered, and as the taxonomic status of Figure 1. Summary of Red List categories
Frank Glaw, Kim Howell, Malcolm Largen, Simon Loader, Stefan Lötters, many species complexes is resolved. for amphibians in the Afrotropical Realm.
Leslie Minter, Martin Pickersgill, Christopher Raxworthy, Mark-Oliver Rödel, The Afrotropical realm currently accounts for about 13% (240) of all globally threatened The percentage of species in each category
Arne Schiøtz, Denis Vallan and Miguel Vences amphibian species. When looking at the Red List Categories, the Afrotropics account for is also given.
only 7% of CR species, but 14% of the EN species, and 15% of the VU species. Hence, on
the basis of current knowledge, threatened Afrotropical amphibians are more likely to be
THE GEOGRAPHIC AND HUMAN CONTEXT in a lower category of threat, when compared with the global distribution of threatened Red List Category Number of species
species amongst categories. This might partly be explained by the fact that the amphibian Extinct (EX) 0
The Afrotropical Realm includes all of mainland Sub-Saharan Africa and the southern Arabian fauna is very poorly known in many parts of the continent (for example the Albertine Rift, Extinct in the Wild (EW) 0
Peninsula1, as well as several large offshore islands: Zanzibar and Pemba; Madagascar; the Congo Basin, the Ethiopian plateau and the Upper Guinea forests of West Africa), Critically Endangered (CR) 33
the western Indian Ocean islands of the Seychelles (including Aldabra), Mauritius (includ- and these are places that could have many threatened species. However, the lower than Endangered (EN) 107
ing Rodrigues), Réunion, Mayotte, and the Comoros; the Gulf of Guinea islands (Bioko, average level of threat is also likely to be genuine, in part because the fungal disease chy- Vulnerable (VU) 100
Príncipe, São Tomé and Pagalu); Socotra; and the Cape Verde Islands. Although Africa tridiomycosis, although present in Africa (Weldon et al. 2004), is not so far believed to be Near Threatened (NT) 53
finally separated from the rest of the southern land-mass of Gondwanaland some 100 a serious threat in the region (although there are few monitoring programmes, and further Least Concern (LC) 467
Ma, the African, Arabian and Eurasian plates abutted at least at the end of the Oligocene, information could change this perception). Species that are threatened by this disease are Data Deficient (DD) 209
and it is only from about 15-10 Ma that Africa has been an isolated land mass (when the more likely to experience sudden and dramatic declines, pushing them very quickly into the Total Number of Species 969
African and Arabian land masses rifted apart; see Kingdon 1989; Goudie 2005). The island higher categories of threat.
3%
of Madagascar, on the other hand, separated from the African mainland between 165-140 Surprisingly, especially for such a poorly known region, the percentage of DD species 11%
Ma and has been isolated (as an island) from all other land masses for 87-91 million years (22%) is slightly lower than the global average (23%). As more African amphibian species 22%
(Storey et al. 1995; Torsvik et al. 2000). are discovered and named, the percentage of DD species might increase for a period, at
An extensive process of rifting which began about 30 Ma has left mainland Africa very least until the conservation status of these species can be adequately assessed. 10%
diverse topographically, with many high mountain ranges, especially on the east of the There have been no recorded recent extinctions of amphibians in the Afrotropical Realm.
continent, and with rift valleys that include some of the deepest lakes in the world. Much However, of the 33 CR species, three are flagged as Possibly Extinct: Arthroleptides dutoiti
of Africa is still volcanically active, including Mount Kilimanjaro, its highest peak, soaring from Mount Elgon in Kenya, which has not been recorded since 1962, despite extensive 6%
some 5,895m above sea-level. A combination of intra-plate hotspots, extensive Cenozoic searches (Lötters et al. 2003); Nectophrynoides asperginis from the Udzungwa Mountains
doming (45 Ma), vulcanism and coastal upwarp has created the world’s largest plateau. in the Eastern Arc of Tanzania (which might have become extinct in the wild in 2004, though
This is characterized by a “basin and swell” topography unique to the interior of the Afri- it still survives in captivity); and Conraua derooi, from the Togo-Volta Highlands of eastern
can continent, and stretching from South Africa to East Africa, with high points including Ghana and western Togo (previously not recorded since the 1960s3). 48%
Mount Mulanje (3,002m) in southern Malawi and Mount Rungwe (2,691m) in southern
Tanzania (Beentje et al. 1994; Goudie 2005). East and North-east Africa are bisected by the
6,000-km-long Great Rift (or Gregory Rift), the “passive” margin between the African and SPECIES RICHNESS AND ENDEMISM
East African Plates. This massive feature is visible from space and runs from the Luangwa
Valley in Zambia, northe-ast through Ethiopia, where it meets the Saudi Arabian Plate and Species Richness and Endemism Across Taxa
continues north-west to the Caucasus Mountains.
Not surprisingly, the region is ecologically diverse, with an enormous range of vegetation The 969 native amphibian species in the Afrotropical Realm represent 16% of the currently
types from deserts to woodlands and grasslands to rainforests. In West Africa, the agricul- known global total of 5,915 species. Of these 969 species, 954 (or 98%) are endemic to the
turally productive Guinea and Sudanese savannah regions slowly merge into the lowland Afrotropics (Table 1). The overwhelming majority of African amphibians are frogs and toads
forests stretching across West Africa. A little further to the east, the highly fragmented, lower (Anura), 98% of which are endemic. All 28 species of Afrotropical caecilian (Gymnophiona)
Guinea rainforests of Nigeria and western Cameroon open into the vast, lush, relatively intact occurring in the region are endemic. Members of the order Caudata (salamanders) are
tropical rainforests of the Congo Basin. Apart from the chain of volcanic mountains centred completely absent. A total of 618 species (64%) are members of families that are endemic
on western Cameroon and extending into the sea as the Gulf of Guinea islands, West Africa to the region.
has very few uplands (with the exception of Mount Nimba, and the Loma and other nearby The amphibian fauna of the Afrotropical Realm has been relatively isolated from that of
mountains in Sierra Leone, Guinea and Liberia). The eastern edge of the Congo Basin is the rest of the world for much of its evolutionary history, and remains so today (the Saharan
flanked by the Albertine Rift, a series of high, block-faulted mountain chains that separates and Arabian deserts forming an effective, though fairly recent, barrier to most species).
the Congo Basin rainforest of Central Africa from the forest/savannah mosaic habitats of This isolation accounts for the high level of family and genus-level endemism within the
East Africa, most famous for their teeming herds of wildebeest and zebra. region. For example, of the 16 families of amphibians found in the region, nine are also
To the north the Great Rift cuts through the Ethiopian Highlands, home to more than endemic. From the perspective of amphibian biogeography, the region is almost defined by
two-thirds of Africa’s unique Afro-alpine habitat, while to the south-east of the Great Rift the distribution of the African treefrogs and reed frogs (family Hyperoliidae) (Poynton 1999),
are the Eastern Arc Mountains that continue into the Southern Highlands of Tanzania south which are present through nearly all of Sub-Saharan Africa, Madagascar, the Seychelles
to Mount Mulanje in Malawi and the Chimanimani Highlands of Zimbabwe. Patches of for- Islands, and the Gulf of Guinea islands (though absent from southern Arabia, the Comoros4,
est (now highly fragmented) fringe the coast of much of eastern Africa, from the Juba and the Mascarenes, and the Cape Verde Islands). The amphibian fauna comprises elements
Shabelle River Valleys in Somalia south to south-eastern South Africa. In the south-western with both tropical and southern temperate origins (Poynton 1999).
part of South Africa, the unique Mediterranean fynbos vegetation is dominant, with its There are 112 amphibian genera in the Afrotropical Realm, of which 105 (94%) are
remarkable floristic endemism, and further north the Succulent Karoo becomes dominant endemic. These endemic genera represent nearly one-quarter (23%) of the 460 amphibian
along the west coast of South Africa. genera worldwide. The Afrotropics, therefore, account for a larger proportion of the overall
Africa has a low human population density (approximately 30 people per square kilometer diversity of amphibians at the generic level than at the species level. The most species-rich
in 2005), of which over 60% live in rural areas, but a high population growth rate (over 2.1% endemic genus in the region is Hyperolius (125 species). At the opposite end of the spectrum,
per annum, though this is decreasing). Historically, the region has been subject to relatively there are 42 monotypic genera endemic to the Afrotropical Realm, which equates to exactly Table 1. The number of Afrotropical amphib-
low levels of anthropogenic disturbance, related not only to low human population densities, ians in each taxonomic Family present in
but also to widespread poverty (gross income per capita was around US$600 in Sub-Saharan the region.
African in 2004). However, 35% of the continent’s Gross Domestic Product is concentrated in
South Africa, where the human impacts on natural ecosystems and biodiversity have been
more severe than in most of the region. Economic growth rates in Africa have been amongst the Family Native species Percentage of Percentage of Native genera Percentage of Percentage of
lowest in the world and still are, though there has been a recent tendency for somewhat faster (endemics to species in region species in family (endemics genera in region genera in family
region) that are endemic that are endemic to region) that are endemic that are endemic
growth (4.5% in 2004). Although many of Africa’s natural habitats are still intact, some parts of to region to region
the continent have been affected much more than others. Deforestation has been particularly
severe in the Upper Guinea forests of West Africa, in the mountains and lowlands of Nigeria and Anura
western Cameroon, in many of the mountainous regions of central and East Africa, and in the Arthroleptidae 51 (51) 100 100 3 (3) 100 100
East African coastal lowlands. This forest loss has been driven largely by expanding subsistence Astylosternidae 29 (29) 100 100 5 (5) 100 100
agriculture to support the burgeoning human populations, but also by commercial agriculture Bufonidae 105 (97) 92 20 15 (14) 93 41
and logging. As mentioned above, habitat loss and fragmentation has been particularly severe Heleophrynidae 6 (6) 100 100 1 (1) 100 100
in South Africa, and above all in the fynbos vegetation of the extreme south and south-west Hemisotidae 9 (9) 100 100 1 (1) 100 100
(which has been heavily impacted by invasive species and fire, and well as by general urban Hylidae 1 (0) 0 0 1 (0) 0 0
and agricultural development). There has also been extensive habitat degradation in the semi- Hyperoliidae 253 (253) 100 100 18 (18) 100 100
arid Sahel region, much of it driven by over-exploitation of resources for basic subsistence by Mantellidae 158 (158) 100 100 5 (5) 100 100
growing human populations. However, in central and south-central Africa, extensive tracts of Microhylidae 87 (87) 100 20 18 (18) 100 26
lowland forest, savannah and grassland remain largely intact. On Madagascar, habitat loss Petropedetidae 102 (102) 100 100 13 (13) 100 100
has been especially severe, with almost no natural habitat surviving on the central plateau, Pipidae 23 (23) 100 77 4 (4) 100 80
and much of the country characterised by very high levels of soil erosion. Ranidae 109 (103) 94 15 14 (9) 64 23
Rhacophoridae 4 (4) 100 1 1 (1) 100 11
Sooglossidae 4 (4) 100 100 2 (2) 100 100
GLOBAL CONSERVATION STATUS TOTAL ANURA 941 (926) 98 18 101 (94) 93 26
Gymnophiona
A total of 969 amphibian species are recorded from the Afrotropical Realm, of which 240 Caeciliidae 22 (22) 100 19 9 (9) 100 100
(25%) are considered threatened (see Figure 1 for details). Although this is significantly Scolecomorphidae 6 (6) 100 100 2 (2) 100 100
less than the global average of 33%2, it is still almost one-quarter of the overall amphibian TOTAL GYMNOPHIONA 28 (28) 100 16 11 (11) 100 100
fauna of the Afrotropics. As is the case globally, the percentage of threatened species is TOTAL ALL AMPHIBIANS 969 (954) 98 16 112 (105) 94 23
expected to increase as the status of DD species is clarified, as new species (many of which
54 Threatened Amphibians of the World

Boophis luteus (Least Concern) is a treefrog in
the Family Mantellidae, which is endemic to
Madagascar. This species is locally abundant
along streams in both pristine and degraded
rainforest. © Piotr Naskrecki
Table 2. The number of species within each
IUCN Red List Category in each Family and
Order in the Afrotropical Realm. Introduced
species are not included.
one-third (33%) of the 126 monotypic genera of amphibians worldwide. This unexpectedly
high percentage is probably a reflection of the poor state of knowledge of the Afrotropical
amphibian fauna; it is likely that many of these genera will prove not to be monotypic as
more species are discovered. For example, recent descriptions include a second species of
the previously monotypic genus Callulina (De Sá et al. 2004), a second species of Cophyla
(Vences et al. 2005), a second species of Acanthixalus (Rödel et al. 2003), and a second and
third species of Alexteroon (Amiet 2000). The seven non-endemic genera in the Afrotropics
include five ranid genera (Euphlyctis5, Hoplobatrachus, Ptychadena, Rana6, Tomopterna7)
and the widespread genera Bufo8 and Hyla5.
As noted already, 16 of the world’s 48 amphibian families (33%) occur in the Afrotropics.
The nine endemic families to the region (Arthroleptidae, Astylosternidae, Heleophrynidae,
Hemisotidae, Hyperoliidae, Mantellidae, Petropedetidae, Sooglossidae and Scolecomorphi-
dae) represent 19% of the global level of diversity of amphibians at the family level9. The
characteristics of these families are provided in Chapter 1.
Among the non-endemic families, the majority of Afrotropical species are in the Bufoni-
dae, the Microhylidae and the Ranidae. Of the Afrotropical Bufonidae, 68 species (65% of
those occurring in the region) are within the widespread genus Bufo8. The remaining 14
genera have mostly small numbers of species, with the exception of Nectophrynoides (11
species). The Afrotropical toad genera of Nectophrynoides (endemic to eastern Tanzania)
and Nimbaphrynoides (endemic to Mount Nimba in Côte d’Ivoire, Guinea and Liberia) are
remarkable in that they include 13 of the 14 known live-bearing anurans in the world10 (with
the possibility that the poorly known, monotypic Afrotropical bufonid genera, Didynamipus
and Laurentophryne, might also be live-bearing).
The Afrotropical Microhylidae species are very unevenly distributed within the region,
with 56 species endemic to Madagascar (10 genera, including Plethodontohyla – 15 spe-
cies, Platypelis – 11 species, Scaphiophryne – 10 species, and Stumpffia – 8 species). On
the African mainland, there are 29 species in eight genera. The largest concentrations of
microhylids are in South Africa (15 species, eight of which are endemic) and Tanzania (12
species, nine of which are endemic, with four endemic genera). Microhylid diversity is low
in the rest of Africa, with only one species in West Africa, and none at all in the equatorial
rainforest belt. The largest genera on the African mainland are Breviceps (15 species) and
Phrynomantis (5 species). The Afrotropical microhylids exhibit a wide diversity of reproductive
modes, including both larval- and direct-developers.11
The Afrotropical Ranidae species are all larval-developers, and occur throughout the
African mainland, with one species naturally occurring on Madagascar. There are 14 genera,
the dominant genus being Ptychadena (47 species), with other notable genera including
Amnirana (11 species), Strongylopus (11 species), Afrana (10 species), and Tomopterna (9
species). Among the Afrotropical ranids is the largest frog in the world, the Goliath Frog
Conraua goliath, with some individuals recorded as weighing more than 3kg.12
Of the remaining families, the highly aquatic Pipidae, although small in number of species
(but with more than 75% of the species in the family occurring in the Afrotropics), is a very
visible and abundant component of the amphibian fauna, being dominated by the generally
widespread and resilient genus Xenopus (16 species).
The caecilian family Caeciliidae is very poorly known in the Afrotropics, as in other parts
of the world. Only 21 species (in nine genera) are known, of which six species are endemic
to the granitic islands of the Seychelles, though strangely none at all occur in Madagascar.
The Seychelles is the only country in the world in which the order Gymnophiona forms
a majority of the amphibian fauna (and see Essay 5.1). One species in the Caeciliidae is
endemic to the island of São Tomé. On the African mainland, members of this family appear
to occur mainly in the West African forest belt, east to Cameroon and western Democratic
Republic of Congo, and in the coastal areas of East Africa and the Eastern Arc Mountains,
south to southern Malawi. There is a single species present in south-western Ethiopia, and
a single species (known from just one specimen) is known from the Albertine Rift highlands.
Caecilians are unknown from the Congo Basin, but this is probably due to lack of sampling,
and they also appear to be absent from the Kenyan Highlands (except for the northern
outliers of the Eastern Arc Mountains).
Not surprisingly, the larger families – Bufonidae, Hyperoliidae, Mantellidae, Microhylidae,
Petropedetidae – have the largest absolute numbers of globally threatened species (Table
2). The percentage of threatened species ranges greatly between the families, from 0% for
the Hylidae, Rhacophoridae and Scolecomorphidae to 100% for the endemic Sooglossidae
of the Seychelles (all four of the species have tiny ranges). The percentage of threatened
species is also very high in the Astylosternidae (73%), reflecting the poor state of forest
conservation in the mountainous regions of Cameroon. The percentages are also high
(>30%) in the Bufonidae, Heleophrynidae, and Microhylidae. In the Bufonidae, only 11 of
the 68 species of Afrotropical Bufo (16%) are globally threatened, but 27 of the remaining
37 species in other genera (80%) are threatened. The Bufonidae are also noteworthy in that
30 of the 38 threatened species fall in the Critically Endangered and Endangered categories,
underlining how urgent the conservation needs are in this family.
Certain families have lower levels of threat, notably the Pipidae, Hemisotidae and
Ranidae13, all of which have a high proportion of adaptable species in the Afrotropics.
The results for the Arthroleptidae are hard to interpret because of the major taxonomic
uncertainties in this family (31% of the species are DD). There is very little information on
the threat levels to Afrotropical caecilians (39% DD); two of the three threatened species
occur in Seychelles, and the other in Kenya, but almost nothing is known of the conservation
status of most of the West African species.
Geographic Patterns of Species Richness and
Endemism
A map of overall species richness of amphibians in the Afrotropical Realm (Figure 2) shows
that species richness is lowest in arid regions, such as the Sahel, the Horn of Africa, and
south-west Africa. However, the apparent region of species paucity on the southern Congo
Basin is almost certainly an artefact reflecting the very limited herpetological work in that
part of Africa.
Although the regions with the highest known species richness, such as south-western
Cameroon and eastern Madagascar are genuine reflections of amphibian diversity, these are
also the rainforest regions that have received the most research attention from herpetolo-
gists. Other regions, such as the high mountains of the Albertine Rift of eastern Democratic
Republic of Congo, Rwanda, Burundi and south-western Uganda, are likely to be richer in
species than the current data indicate. Some regions showing higher species richness, such
as the Eastern Arc Mountains of Tanzania, and Mount Nimba in Liberia, Guinea and Côte
d’Ivoire, are likely to represent genuine patterns, whereas others (for example, in the Upemba
region of southern Democratic Republic of Congo and the Taï National Park in south-western
Côte d’Ivoire) are probably a reflection of locally intensive survey efforts.
There are seven major concentrations of threatened species (Figure 3a): the Upper Guinea
forests from Sierra Leone to Togo; south-eastern Nigeria and south-western Cameroon;
the mountains of the Albertine Rift; the Ethiopian Highlands; the Eastern Arc Mountains;
southern South Africa (especially in the south-western Cape); and eastern Madagascar
(especially in the northern and southern extremities of the eastern rainforest zone). These
concentrations of threatened species correlate with those for other taxa (Stuart and Collar
1988; Baillie et al. 2004). Smaller concentrations of threatened amphibians are found in: São
Tomé, the Kenyan Highlands; southern Malawi (around Mount Mulanje); eastern Zimbabwe
(the Chimanimani mountains, in particular) and adjacent Mozambique; and the Seychelles
Islands. These geographic concentrations reflect the parts of the region where amphibians
have naturally small ranges, and where habitat destruction is ongoing. However, at least
some of the threatened species in the Upper Guinea forests were once more widespread,
but have declined due to extremely severe habitat destruction. Because of the relatively
small number of CR species, there are few significant concentrations of CR species, and
the overall pattern is similar to that for threatened species (Figure 3b).
Species Richness and Endemism within Countries
Amphibians occur naturally in every mainland country in Sub-Saharan Africa and southern
Arabia (Figure 4). They are also present on Madagascar, and are indigenous to the following

Family CR EN VU NT LC DD Total number Number threatened % threatened

of species or Extinct or Extinct
Anura
Arthroleptidae 3 9 2 3 18 16 51 14 27
Astylosternidae 2 11 8 2 5 1 29 21 72
Bufonidae 9 21 8 2 49 16 105 38 36
Heleophrynidae 2 0 0 0 4 0 6 2 33
Hemisotidae 0 0 1 0 4 4 9 1 11
Hylidae 0 0 0 0 1 0 1 0 0
Hyperoliidae 1 19 29 17 133 54 253 49 19
Mantellidae 7 12 16 12 77 34 158 35 22
Microhylidae 3 13 16 2 36 17 87 32 37
Petropedetidae 3 13 8 10 39 29 102 24 24
Pipidae 1 1 0 1 15 5 23 2 9
Ranidae 1 7 7 4 68 22 109 15 14
Rhacophoridae 0 0 0 0 4 0 4 0 0
Sooglossidae 0 0 4 0 0 0 4 4 100
TOTAL ANURA 32 106 99 53 453 198 941 237 25
Gymnophiona
Caeciliidae 1 1 1 0 11 8 22 3 14
Scolecomorphidae 0 0 0 0 3 3 6 0 0 Grandisonia sechellensis (Least Concern) is known only from the islands of Mahé, Praslin
TOTAL GYMNOPHIONA 1 1 1 0 14 11 28 3 11 and Silhouette in the Seychelles. The Seychelles is the only place in the world where
TOTAL ALL AMPHIBIANS 33 107 100 53 467 209 969 240 25 caecilians form the majority of the amphibian fauna, and this species is sympatric with five
other caecilian species. © Renaud Boistel
 Chapter 5. Amphibians of the Afrotropical Realm 55

Bufo togoensis (Near Threatened) is a toad

from the Upper Guinea forests of West
Africa, ranging from eastern Sierra Leone to
western Togo. It lives only in primary forest,
a. usually in close association with the streams
in which it breeds, and is affected by ongoing
deforestation throughout its range. © Piotr
Figure 2. The species richness of amphibians in the Afrotropical Realm, with darker colours Naskrecki
corresponding to regions of higher richness. Colour scale based on 10 quantile classes;
maximum richness equals 123 species.
Figure 4. The number of amphibians present
in and endemic to each Afrotropical country.
islands: Seychelles (on most of the granitic islands, but not on the coral atolls), Zanzibar, *denotes countries not entirely within the
Pemba, Mafia, Bioko, Príncipe and São Tomé (but apparently not on Pagalu, Socotra, the Afrotropical Realm, hence only the species
Comoros4, the Mascarenes or the Cape Verde Islands). whose ranges fall within the region are
Madagascar has been intensively studied (Blommers-Schlösser and Blanc 1991; Glaw and included.
Vences 1994; Andreone et al. 2005) and has the largest number of species of any country in
the Afrotropical Realm (226 species), and 70% of these are in the endemic family Mantellidae
(see also Essay 5.2). The only other country with more than 200 species is the Democratic Figure 5. Percentage of species endemic to
Republic of Congo, and only four other countries have more than 100 species (Cameroon, each Afrotropical country. Countries with no
Tanzania, South Africa, and Nigeria), although, if including species awaiting description, endemic species are not included. *denotes
Cameroon has more than 200 species (J.-L. Amiet pers. comm.; see Essay 5.3). countries not entirely within the Afrotropical
However, these figures need to be treated with considerable caution. The rate of new Realm, hence only the species whose ranges
species’ descriptions in Madagascar has been very high over the last decade, with the number fall within the region are included.
of described species from the country doubling since 1991 (compare Blommers-Schlösser
and Blanc [1991] with Figure 4). However, the rate of species descriptions in recent years
from the African mainland has been much slower than from Madagascar, and this is largely Figure 6. The number of threatened amphib-
a reflection of the very limited amount of herpetological work that has been conducted on ians present in and endemic to each Afrotropi-
the continent in recent decades. Only the work of Schiøtz (1967, 1975, 1999) focusing on cal country. Countries with no threatened spe-
the Hyperoliidae, and that of Tandy and Keith (1972) focusing on African Bufo, has taken cies are not included in the diagram. *denotes
a continental approach to the amphibian fauna. In particular, there has been very limited countries not entirely within the Afrotropical
work carried out in the Congo Basin since the late 1950s in what might be expected to be b. Realm, hence only the species whose ranges
one of the most diverse parts of the region. Similarly, with the exception of a small amount fall within the region are included.
of work in Uganda (Drewes and Vindum 1994; Vonesh 2001), the Albertine Rift has received
no attention for a similar period of time, despite the fact that this area has higher species Figure 3. a) The richness of threatened amphibians in the Afrotropical Realm, with darker
richness and endemism among vertebrates than any other part of mainland Africa (Stuart colours corresponding to regions of higher richness. Colour scale based on 10 quantile Figure 7. Percentage of native species that are
and Collar 1988; Brooks et al. 2001). There has been much greater focus on the amphibian classes; maximum richness equals 31 species. b) The richness of CR amphibians in the Af- threatened. Countries with no threatened spe-
fauna of Cameroon (Perret 1966; Amiet 1983, 1989), but serious exploration of the fauna of rotropical Realm, with darker colours corresponding to regions of higher richness. Maximum cies are not included in the diagram. *denotes
West Africa only resumed in the mid 1990s (Rödel 2000; see Essay 5.4). Apart from some richness equals three species. countries not entirely within the Afrotropical
Realm, hence only the species whose ranges
fall within the region are included.

Madagascar Seychelles Madagascar Seychelles

Congo D.R.
Cameroon São Tomé and Principe
Cameroon São Tomé and Príncipe
Tanzania Madagascar
South Africa Tanzania Cameroon
Nigeria South Africa
Angola Tanzania South Africa Tanzania
Kenya
Côte d’Ivoire Ethiopia Congo D.R. Madagascar
Gabon Cameroon
Zambia Côte d’Ivoire Burundi
Ghana Congo, D.R.
Nigeria Rwanda
Malawi Chad*
Guinea Ghana South Africa
Equatorial Guinea Angola
Mozambique Yemen* Ethiopia Yemen*
Ethiopia
Zimbabwe Kenya Rwanda Côte d’Ivoire
Central African Republic Somalia Burundi Ethiopia
Uganda
Congo Togo
Kenya Togo
Liberia Mali*
Sierra Leone Seychelles Ghana
Namibia Namibia
Swaziland Zimbabwe Uganda Nigeria
Botswana
Rwanda Burundi Zimbabwe Uganda
Senegal Guinea
Somalia Guinea Zimbabwe
Burundi
Côte d’Ivoire
Mali* Eritrea Malawi Liberia
Gambia Ghana Equatorial Guinea Guinea
Togo
Burkina Faso Lesotho Liberia Malawi
Sudan* Central African Republic
Benin Gabon Kenya
Lesotho Uganda
Chad* Mozambique Congo D.R.
Gabon
Eritrea
Guinea Bissau Malawi São Tomé and Príncipe Equatorial Guinea
Seychelles Sierra Leone Togo Mozambique
Niger*
Saudi Arabia* Liberia Sierra Leone Sierra Leone
Djibouti Rwanda
São Tomé and Principe Namibia Gabon
Yemen* Zambia
Mauritania* Equatorial Guinea Yemen* Namibia
Oman*
United Arab Emirates* Nigeria Zambia Zambia

0 50 100 150 200 250 0 20 40 60 80 100 0 10 20 30 40 50 60 0 10 20 30 40 50 60

Number of species Percentage of endemic species Number of threatened species Percentage of threatened species
Endemics Non-endemics Endemics Non-endemics
Figure 4 Figure 5 Figure 6 Figure 7
56
Kassina cochranae (Near Threatened) is in the
endemic Afrotropical Family Hyperoliidae. It is
an arboreal, forest-dwelling species known
from the forest zone of Sierra Leone, Liberia
and Guinea. Certain populations are probably
suffering from severe deforestation as a result
of agricultural expansion, logging and growing
human settlements. © Piotr Naskrecki
Table 3. The habitat preferences of amphib-
ians in the Afrotropical Realm.
Figure 8. The habitat preferences of Afrotropi-
cal amphibians. The plot on the left-hand side
shows the number of species in the region in
each habitat type. On the right-hand side,
the percentage of these species which are
threatened is given.
Hyperolius puncticulatus (Least Concern),
one of at least 125 treefrogs in the genus Hy-
perolius, ranges from coastal Kenya, through
eastern and southern Tanzania (including
the island of Zanzibar) to the highlands of
Malawi. The males of this species call from
vegetation around pools, where they breed.
© Alan Channing
Threatened Amphibians of the World
preliminary work in Ethiopia (Largen 2001) and Somalia (Lanza 1990), work in eastern Africa one of the native species, the Mascarene Ridged Frog Ptychadena mascareniensis (LC),
has been very limited, although there has been some recent attention on Tanzania (e.g., being found nowhere else15. In both Seychelles and São Tomé and Príncipe, the level of
Poynton 2003), and a field guide to the amphibians of East Africa, covering Kenya, Tanzania endemism is 100%.
and Uganda (Channing and Howell 2006). On the African mainland, Tanzania has more endemic species than any other country (see
The frogs of the southern third of the African continent are probably somewhat better Essay 5.5). More than 50 endemic species are also known from Cameroon, the Democratic
known, thanks to a number of detailed studies, for example: Poynton (1964) and Channing Republic of Congo, and South Africa. In terms of percentage of the fauna being endemic, the
(2001) covering the whole region; Wager (1986), Lambiris (1989a), Passmore and Carruthers highest endemism on the African mainland is found in South Africa, Tanzania and Ethiopia
(1995), and Minter et al. (2004) in South Africa, Lesotho and Swaziland; Channing and Griffin (all have around 40% of species endemic). Although the actual numbers and percentages
(1993) in Namibia; Lambiris (1989b) in Zimbabwe; Broadley (1971) in Zambia; Stewart (1967) will change as new information becomes available, the overall patterns are concordant with
in Malawi; Poynton (1966) in Mozambique; and the classic Amphibia Zambesiaca works of those of other taxonomic groups, and are almost certainly real.
Poynton and Broadley (1985ab, 1987, 1988, 1991). However, even in South Africa, which Threatened species occur in 27 of the 49 countries in which there are native amphibians
is by far the most studied country in the region, new species continue to be described (for (Figure 6). In fact, threatened species are concentrated in relatively few countries. Only four
example, Turner et al. 2004). countries, Madagascar, Cameroon, Tanzania and South Africa, have more than 20 globally
In reality, our understanding of the country-level species richness of amphibians in the threatened species. The top three countries, Madagascar, Cameroon and Tanzania, are
Afrotropics is still very incomplete. Future investigations will doubtless result in increases undoubtedly genuine centres of threatened species, but the number is almost certainly
in the numbers of species in every country; however, based on the known species richness grossly under-estimated for the Democratic Republic of the Congo. Here there are likely
of other groups such as birds, these increases are likely to be greatest for the Democratic to be many threatened species, probably many of them still undescribed, in the mountains
Republic of Congo, Nigeria, Angola, Côte d’Ivoire, Gabon, Ghana, Guinea, Equatorial Guinea, around the Albertine Rift (where there is also severe forest loss due to expanding subsis-
Mozambique, Ethiopia, Central African Republic, Uganda, Congo, Liberia, Sierra Leone, tence agriculture). The percentage of threatened amphibian species is highest in the island
Rwanda, Burundi, Togo, Sudan, Benin14, and Chad. Meanwhile, Madagascar, Cameroon and nations of Seychelles and São Tomé and Príncipe (Figure 7). The highest percentage on the
Tanzania, which have received the most herpetological attention (after South Africa) in recent African mainland is 27% for Cameroon, a reflection of the poor state of forest conservation
years, show no signs of slowing in their rates of new species’ descriptions. in the mountainous regions of the country. In many countries, the percentage of threatened
Not surprisingly, endemism (in terms of relative proportions; Figure 5) is much higher species can be expected to rise as new species discoveries are made or crucial habitats
in island nations. Madagascar has far more endemics than any other country, with all but are destroyed.
Assessments of the conservation status of Afrotropical amphibians at national level have
been carried out only for South Africa, Lesotho and Swaziland (Branch 1988; Monadjem et
al. 2003; Minter et al. 2004).
There are only 33 Critically Endangered Afrotropical species, nine of which occur in
Habitat type Number % of all Threatened % of species Madagascar, seven in Cameroon and five in Tanzania. Outside these three countries, there
of species in species and Extinct occurring are three Critically Endangered species in South Africa, two each in Côte d’Ivoire and Kenya,
each habitat occurring in species in habitat that
the habitat are Threatened
and one each in Gabon, Ghana, Guinea, Liberia, Togo and Zimbabwe. However, the numbers
or Extinct in West Africa are very likely to increase during the coming years, both because of newly
discovered species and because of destruction of forest and mountain habitats.
Forest 648 67 208 32
All tropical forest 624 64 201 32
Lowland tropical forest 495 51 114 23 HABITAT AND ECOLOGY
Montane tropical forest 322 33 137 43
Savannah 227 23 7 3 Habitat Preferences
Grassland 267 28 52 19
Shrubland 213 22 29 14 Two-thirds of Afrotropical amphibians occur in forests, and over 40% are believed to be able
Secondary terrestrial habitats 415 43 74 18 to survive in secondary terrestrial habitats (Table 3; Figure 8). Similar percentages make use
Flowing freshwater 380 39 112 29 of flowing water and of standing, open water habitats, but marshes and swamps appear
Marsh/swamp 223 23 38 17 to be used by fewer species. Forest-dwelling amphibians are more likely to be threatened
Still open freshwater 375 39 53 14 than those occurring in any other habitats, with almost one-third of them being globally
Arid and semi-arid habitats 41 4 1 2 threatened. Almost 30% of amphibians associated with flowing water (generally streams)
are threatened. Consequently, forest-associated amphibians that live along streams are
particularly likely to be threatened, a combination that has also been associated with rapid
declines worldwide (Stuart et al. 2004).
For tropical forests overall, 32% of Afrotropical species are globally threatened. However,
Number of species % of species in habitat that are threatened in montane tropical forest, 43% of known species are threatened, compared with 23%
Forest
in lowland tropical forest. These figures probably reflect smaller range sizes of montane
species, and the lack of effective habitat conservation measures in many mountainous
Savannah parts of the region.
Grassland Amphibians occurring in savannah and arid habitat are particularly unlikely to be threat-
Shrubland ened. Africa has 47% of the world’s 484 savannah-associated amphibians, and 44% of the
world’s 94 arid-habitat species, but only 14% of the global total of 4,712 forest-dwelling
Secondary terrestrial habitats
species. Many of these “species” are in fact complexes of several species, and when these
Flowing freshwater are disaggregated, the number of savannah amphibians will increase. In addition, the
Marsh/swamp number of threatened savannah species can also be expected to increase as their habitats
shrink and dry out (these habitats are especially vulnerable to climate change) and their
Still open freshwater
ranges become smaller.
Arid and semi-arid habitats
0 200 400 600 0 10 20 30 Reproductive Modes
Larval development is by far the most common reproductive mode in the Afrotropics (85%
of species), compared with 10% for direct development and 2% live-bearing (this compares
with the global picture of 68% larval development, 30% direct development, and 1%
live-bearing) (Table 4). Although live-bearing is uncommon, the Afrotropics account for
36% of the world’s known live-bearing amphibians, and all but one of the world’s known
live-bearing frogs and toads. However, it should be noted that the reproductive mode of
many species is unknown, and more terrestrial breeders are likely to be identified (Rödel
and Ernst 2002; Rödel et al. 2002).
In the Afrotropics, the percentage of globally threatened direct-developing amphibians
is only slightly higher than for larval-developing species (at a global level, direct-developing
species are much more likely to be threatened). The high percentage of threatened live-
bearing species in the Afrotropics is a reflection of the high levels of threat to species in
the genera Nectophrynoides and Nimbaphrynoides, many of which have very small ranges
in fragile, primarily montane, and poorly protected, habitats.
MAJOR THREATS
As in other parts of the world, habitat loss is overwhelmingly the major threat to amphibians
in the Afrotropics (Table 5; Figure 9), affecting over 90% of the threatened species. Other
commonly recorded threats include invasive species, fire and pollution. Over-utilization ap-
pears to be a minor threat in the region (at least based on current knowledge), and disease,
which is a very important factor in other parts of the world, is cited as a threat to only 2%
of threatened species in the Afrotropics (although the amount of information on the chytrid
fungus, and its pathogenicity, is very limited in the region).
In terms of the types of habitat loss that are impacting threatened amphibians in the
Afrotropics, expanding croplands, vegetation removal (mainly logging), and urbanization /
industrial development are approximately equivalent, and each is affecting more than 70% of
threatened species. Livestock grazing has less impact (probably because it is more prevalent
in regions of lower amphibian species richness, and in any case much of Africa has long
been grazed by large wild mammals), and tree plantations also appear not to be a significant
threat in most places (although this is a serious threat in South Africa and Swaziland). In
 Chapter 5. Amphibians of the Afrotropical Realm 57

Reproductive mode All Species Threatened or % Threatened

Extinct Species or Extinct
Direct development 100 26 26
Larval development 819 190 23
Live-bearing 21 10 48
Not known 29 14 48

Table 4. Afrotropical amphibians categorized by reproductive mode.

Threat type Threatened species % Threatened

species
Habitat loss 221 92
Agriculture – Crops 191 80
Agriculture – Tree plantations 18 8
Agriculture – Livestock 79 33
Timber and other vegetation removal 179 75
Urbanization and industrial development 177 74
Invasive species 52 22
Utilization 15 6
Accidental mortality 1 0.4
Pollution 30 13
Natural disasters 3 1
Disease 4 2
Human disturbance 1 0.4
Changes in native species dynamics 2 1
(excluding disease)
Fire 50 21

Leptopelis parkeri (Vulnerable), a treefrog in

Table 5. The major threats to globally threatened amphibians in the Afrotropical Realm. the Family Hyperoliidae, is endemic to several
Only present threats to species are tallied. mountain blocks in the Eastern Arc chain of
All habitat loss
Tanzania. It is a species of closed, intact
rainforest, not surviving in seriously disturbed
parts of West Africa, planned mining activities might become increasingly serious as threats Invasive species habitats outside forest, and is threatened by
to important sites for amphibians (for example, in Guinea and Ghana). Utilization habitat loss as a result of expanding human
A total of 84 species (18 of which are threatened) are recorded as being used for some Accidental mortality settlements and agriculture, and the har-
or other purpose in the region (Table 6). The most commonly recorded reason for harvest- vesting of wood. © David Moyer / Wildlife
Persecution
ing Afrotropical amphibians is for the international pet trade (especially in Madagascar), Conservation Society
followed by human consumption (although the number of species used as human food is Pollution
probably greatly under-recorded). Much of the harvesting of amphibians in the region is not Natural disasters
considered to constitute a major threat to the species. Of the 84 species being harvested,
Disease
utilization is considered to be a threat for 32 (of which only 15 are threatened species for
which harvesting is believed to be contributing to deterioration in their status). Human disturbance

Changes in native sp. dynamics

Fire
POPULATION STATUS AND TRENDS
0 20 40 60 80 100
Estimates of Population Trends Percentage of threatened species affected

A summary of the inferred population trends of Afrotropical amphibians is presented in

Table 7. For nearly all species, these trends are inferred from trends in the state of the Figure 9. The major threats impacting threatened amphibians in the Afrotropical Realm.
habitats on which the species depend. Species with decreasing populations are typical
forest-dependent species that can tolerate little disturbance to their habitats. The percent-
age of decreasing and increasing species in the Afrotropics is very similar to the global
results. However, the percentage of species where the situation is stable is slightly lower, Purpose Subsistence Sub-national/ Regional/ Number
and the percentage for which it is unknown is slightly higher than the global averages (27% National International of species
and 30%, respectively). Food – human 24 (4) 5 (2) 1 25 (4)
Medicine – human and veterinary 3 1 1 3 (0)
“Rapidly Declining” Species Pets, display animals 0 1 61 (15) 61 (15)
Research 0 1 1 2 (0)
The Afrotropics appears to have been shielded to some extent from the amphibian declines
that are taking place in some other regions, with only 30 (6%) of the 470 globally “rapidly
declining” species occurring within the region (a full list of all “rapidly declining” species is Table 6. The purposes for which amphibians are used in the Afrotropical Realm. The numbers
provided in Appendix IV and includes their occurrence within each of the realms). Most of in brackets are the number of species within the total that are threatened species.
the “rapidly declining” species are threatened primarily by the reduction of suitable habitat.
Twelve of these are forest-obligate species from the Upper Guinea region of West Africa
(see Essay 5.4), where habitat loss has been particularly severe. Another six are endemic
to South Africa, where urbanization and agricultural intensification in key habitats has been Population Trend Number of species % of species
relatively more severe than on the rest of the continent (see Essay 5.6). The remaining eight Decreasing 425 44 The aptly named Tomato Frog Dyscophus
species are scattered widely across the continent in Cameroon, Gabon, Kenya, Madagascar Stable 221 23 antongilii (Near Threatened) of the Fam-
(two species), São Tomé and Príncipe, Tanzania, and Zimbabwe. Increasing 8 1 ily Microhylidae is endemic to north-eastern
The Afrotropics have three “rapidly declining” species, affected by severe over-harvesting. Unknown 315 33 Madagascar. Although it is an adaptable
Two of these, Mantella cowanii (CR) and Scaphiophryne gottlebei (CR), both from Madagas- species, being found in a variety of habitats
car, appear to have declined due to over-collection for the international pet trade (Andreone from primary rainforest to disturbed urban
et al. 2006). In the case of Mantella cowanii, it declined initially because of the loss of most Table 7. The population trends for all extant Afrotropical amphibians. areas, the pollution of waterbodies is a
of its habitat, but subsequently the remnant populations were hit by over-harvesting. The potential threat to this species. © Russell
third species, the Goliath Frog Conraua goliath (EN) from Cameroon and Equatorial Guinea, A. Mittermeier
has declined mainly due to over-harvesting for human consumption, though there is also
some international trade. Family Number of species Percentage of
So far, only one “rapidly declining” species undergoing an enigmatic decline has been in “rapid decline” species in family
in “rapid decline”
recorded in the Afrotropics, namely the Kihansi Spray Toad Nectophrynoides asperginis (CR).
This species, endemic to the spray zone of one waterfall in Tanzania, decreased initially Arthroleptidae 3 6
because of the damming of an upstream river. However, its final catastrophic decline to Bufonidae 5 5
probable extinction is believed to have been caused by the fungal disease chytridiomycosis Hemisotidae 1 11
(Krajick 2006), which has been implicated or suspected in most of the enigmatic declines Hyperoliidae 6 2
worldwide. Globally, enigmatic declines account for 48% of the “rapid declines” worldwide, Mantellidae 1 6
but for only 3% so far in the Afrotropics, though it is possible that additional enigmatic Microhylidae 2 2
declines have not yet been detected. One suspected case concerns Arthroleptides dutoiti Petropedetidae 7 7
(CR) in Kenya. This species has the ecological characteristics of other species that have Ranidae 5 5
succumbed to chytridiomycosis elsewhere in the world, and now appears to be absent from
former sites, even though suitable habitat remains. It is not listed as an enigmatic decline
species here, as its decline might have taken place prior to 1980. Table 8. The number of species in “rapid decline” in the Afrotropical Realm by Family .
58
Endnotes
1 We follow the WWF biogeographic realms in this
chapter, but recognize that this is not ideal for
amphibians, especially with regard to the inclusion
of the Arabian Peninsula within the Afrotropical
Realm. The small Arabian amphibian fauna is
generally of Palaearctic origin.
2 P<0.01 (binomial test)
3 At the time of writing, this species had been redis-
covered in the Togo Hills (Leache et al. 2006), and
been found outside its former range in the Atewa
mountains (M.-O. Rödel, pers. comm.).
4 Although no amphibians have been found on the
islands that make up the country Comoro, two
undescribed species are present on the French
island of Mayotte, which is part of the Comoro
Archipelago (Vences et al. 2003). Because they
are undescribed, they are not included in this
analysis.
5 The genera Euphlyctis and Hyla occur only in the
southern Arabian peninsula within the Afrotropical
Realm, and not on the African continent south of
the Sahara.
6 The only evidence for the occurrence of the
genus Rana in Africa is the doubtfully valid Rana
demarchii of unknown provenance in Ethiopia
(Largen 2001).
7 The genus Tomopterna occurs marginally in
the Palaearctic, as it is present in parts of the
Sahara region; the genus is, however, essentially
Afrotropical.
8 Frost et al. (2006) split the genus Bufo in Africa.
9 Frost et al.’s (2006) rearrangement results in
19 families in the Afrotropics, of which nine
are endemic: Arthroleptidae; Brevicipitidae;
Heleophrynidae; Hemisotidae; Hyperoliidae;
Mantellidae; Ptychadenidae; Phrynobatrachidae;
and Pyxicephalidae. However, in this section
we follow the former taxonomic arrangement of
families based on Frost (2004).
10 The only other species is the apparently extinct
Eleutherodactylus jasperi from the Neotropics.
11 Frost et al. (2006) separate the genera Breviceps,
Balebreviceps, Callulina and Probreviceps into a
new family endemic to the Afrotropics: Brevicipi-
tidae. Loader et al. (2004) consider that the genus
Spelaeophryne also belongs with the brevicipitine
group. Under this arrangement, all the African
mainland species remaining in Microhylidae are
larval developers.
12 Under the new arrangement by Frost et al.
(2006), the genus Amnirana is the only remaining
member of the Ranidae in the Afrotropics. The
genus Conraua is moved to Petropedetidae, and
Euphyctis and Hoplobatrachus are transferred to
the predominantly Indomalayan Dicroglossidae.
Ptychadena, Hildebrandtia and Lanzarana are
transferred to a new endemic Afrotropical family,
Ptychadenidae, and the remaining Afrotropical
genera previously considered to be in Ranidae
are transferred to the endemic new family Pyxi-
cephalidae.
13 It should be noted that the species of Ranidae
that have been separated by Frost et al. (2006) as
Pyxicephalidae have a clear center of endemism
and species richness in southern Africa; these
species appear to be on average more strongly
threatened than other frogs in South Africa (Van
der Meijden et al. 2006).
14 Indeed, since we concluded this analysis, Nago
et al. (2006) added 17 new country records of
amphibians from Benin.
15 However, Vences et al. (2004) show that even
the Ptychadena mascareniensis on Madagascar
represent a different species from the animals on
the African mainland, but as the formal taxonomic
amendments have not yet been made, we have
not included this change in our analysis. When
the formal taxonomic change is made, 100% of
the amphibian species occurring on Madagascar
will be considered to be endemic.
Threatened Amphibians of the World
Unlike most other regions, the Afrotropical “rapidly declining” species show no distinct
taxonomic pattern (see Table 8), with small percentages (less than 7% of the species) in
all of the larger families.
KEY FINDINGS
• A total of 969 species are recorded from the Afrotropical Realm, of which 240 (nearly
25%) are considered threatened.
• At the species level, 954 amphibians (99% of those present) are endemic to the Afro-
tropics; of the 16 families found in the region, nine are endemic, and of 112 amphibian
genera occurring, 105 are endemic.
• The percentage of threatened species is very high in the family Astylosternidae
(72%), reflecting the poor state of forest conservation in the mountainous regions of
Cameroon. On the other hand, certain families have lower levels of threat (Pipidae,
Hemisotidae and Ranidae), all of which have a high proportion of adaptable species
in the Afrotropics.
• Geographic concentrations of threatened species occur in the Upper Guinea forests from
Sierra Leone to Togo; south-eastern Nigeria and south-western Cameroon; the mountains
of the Albertine Rift; the Ethiopian Highlands; the Eastern Arc Mountains; eastern and
southern South Africa (especially in the south-western Cape); and eastern Madagascar
(especially at the northern and southern tips).
• Madagascar has the largest number of species of any country in the Afrotropical Realm
(226 species). Only five other countries have more than 100 species (Democratic Republic
of Congo, Cameroon, Tanzania, South Africa, and Nigeria), with all except Nigeria having
more than 50 endemics. Madagascar, Cameroon, Tanzania and South Africa each have
more than 20 globally threatened species. Madagascar has many more endemics than
any other country.
• Among species occurring in tropical forests, 43% of species in montane tropical forest
are threatened, compared with 23% in lowland tropical forest, probably reflecting smaller
range sizes of montane species, and the lack of effective habitat conservation in montane
regions. Africa has 47% of the world’s 484 savannah-associated amphibians (yet only
3% of these species are threatened) and 44% of the world’s 94 arid-habitat species (2%
threatened).
• Habitat loss, primarily due to expanding croplands, vegetation removal (mainly logging),
and urbanization/industrial development, is affecting over 90% of the threatened spe-
cies in the region. Other commonly recorded threats include invasive species, fire and
pollution; disease is cited as a threat to only 2% of threatened species.
• Only 6% of the 470 globally “rapidly declining” species occur within the region; 12 of
these species are forest-obligate species from the Upper Guinea region of West Africa,
where habitat loss has been especially severe.
• No amphibian extinctions have yet been recorded from the Afrotropics, but three species
are possibly extinct.
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ESSAY 5.1. SEYCHELLES AMPHIBIANS
Due to their permeable skin few amphibians are able to tolerate dry or salty
conditions, a characteristic that has prevented most species from colonising
oceanic islands. The most diverse oceanic island amphibian fauna is found
in the Seychelles, an archipelago of about 115 granitic and coralline islands
in the Indian Ocean, some 1,600km east of mainland Africa, and northeast of
the island of Madagascar (Figure 1). Eleven native amphibian species have
been recorded from the Seychelles islands, comprising both recent colonists
and ancient endemics. Recent colonization by one species, the Mascarene
Grass Frog Ptychadena mascariensis (LC), is probably the result of human
introduction (Vences et al. 2004) in the 1800s, whilst the endemic Seychelles
Treefrog Tachycnemis seychellensis (LC) is believed to be descended from a
natural colonist from Madagascar (Vences et al. 2003). The remaining spe-
cies are all ancient endemics that have probably existed on the island since
their isolation from the Indian landmass some 65 million years ago. These
species comprise the endemic frog family Sooglossidae and seven species
of burrowing caecilians (Order Gymnophiona).
Sooglossidae are an exceptional family in several respects. Four species
are currently recognized in two genera, although recent morphological and mo-
lecular data suggests that a further three species remain to be described and
that the genera need to be redefined. They are all small frogs, with Gardiner’s
Frog Sooglossus gardineri (VU) among several species up for contention as
the world’s smallest frog (with adults being as small as 9mm in length). All
sooglossids are found in the damp forests of the two highest islands, Mahé
and Silhouette. These habitats have only seasonal or fast-flowing stems, and
the sooglossids have abandoned the normal frog life-cycle in favour of ter-
restrial development. The Seychelles Frog Sooglossus sechellensis (VU) lays
its eggs on land and the female carries the tadpoles until they develop into
frogs. Sooglossus gardineri is even more specialized, with the male guarding
terrestrial eggs that hatch into 3-mm long froglets; the entire tadpole stage is
passed within the egg. Thomasset’s Seychelles Frog Nesomantis thomasseti
(VU) has recently been discovered to have a very similar breeding strategy,
but nothing is known of the reproductive behaviour of the Seychelles Palm
Frog S. pipilodryas (VU). This latter species is the most recently described of
the Seychelles amphibian species, having been discovered in 2000 (Gerlach
and Willi 2002). It lives in the axils of endemic palms where it lay protected
from discovery by the dense spines of the palm leaves. It is the only arboreal
sooglossid, the other species all being associated with crevices in boulder
fields or the leaf-litter and root-mat of the forest floor.
The caecilian fauna of the Seychelles islands comprises six species in
three endemic genera. Not surprisingly, and as with caecilians in general
(see Essay 1.3), these are the least well known of all Seychelles amphib-
ians. Their burrowing habits make them difficult to locate and study. For
most species, their distributions are reasonably well defined and there are
some observations of breeding habits. Different species appear to show the
full range of reproductive strategies, form aquatic larvae, thorough to ter-
restrial larvae and direct development. The limited ecological data available
indicate that Grandisonia alternans (LC) is a widespread species occurring in
all habitats and Hypogeophis rostratus (LC) is a similar generalist, although
more associated with lowland habitats. In contrast, Praslina cooperi (VU) and
some of the small Grandisonia species appear to be specialists of the high
forest. Current research into this group is attempting to develop monitoring
methods and to identify aspects of their ecology that are of importance to
their conservation.
ESSAY 5.2. THE ENDEMIC AND THREATENED AMPHIBIANS OF MADAGASCAR
Separated from mainland Africa by a sea channel of about 300km, the biodiver-
sity of Madagascar has experienced a distinct evolutionary trajectory that has
resulted in a very high degree of endemism in both its fauna and flora. Among
the vertebrates, the amphibians of this large island (around 580,000km², the
fourth in the world for size) are currently represented by more than 230 frog
species1, a number that is still preliminary and tentative, since many more
remain to be discovered or are awaiting description (see Essay 1.1) (Figure 1).
Chapter 5. Amphibians of the Afrotropical Realm
Half of Seychelles amphibians are threatened due to their naturally
restricted ranges and deteriorating habitats (five species are categorised
as Least Concern, 5 Vulnerable and one Endangered). The sooglossid frogs
and the caecilian Praslina cooperi have particularly restricted ranges, being
associated only with the damper rain-forests. Habitat deterioration is a
significant threat to the caecilian Grandisonia brevis (EN), which has a re-
stricted range and occurs in habitats that are suffering from ongoing invasion
by alien plant species. In addition, there is some suggestion that changes
in rainfall patterns may have impacts on some species, which may result
from mid-year declines in rainfall restricting the activity and distribution of
the rain-forest specialists (Gerlach 2000). Losses of several areas of marsh
habitats have probably caused population declines in some species, such as
another caecilian Grandisonia alternans (Gerlach 2000).
There have been a number of successful conservation programmes in
Seychelles, concentrating mainly on birds. Critically Endangered species have
been rescued from extinction by limited habitat management, but mostly by
inter-island transfers. These programmes have been to the benefit of spe-
cies such as the Seychelles Magpie-robin Copsychus seychellarum and the
Seychelles White-eye Zosterops modesta. Such options are not available
for the amphibians due to their close dependence on their high forest or
marsh-land habitats. Consequently, preservation of their habitat is essential;
most species are present in protected areas, although habitat management
is still required to control the spread of invasive plant species. The island
with the highest diversity of amphibians (Silhouette) is currently unprotected
Nesomantis thomasseti (Vulner-
able) is restricted to Mahé and
Silhouette Islands in the Seychelles,
and has been recorded from the
Morne Seychellois National Park.
© Naomi Doak
Remarkably, out of the entire amphibian fauna, only two species, the adaptable
and widely distributed Ptychadena mascareniensis, which is present also in
mainland Africa, and Hoplobatrachus tigerinus, introduced to Madagascar from
southern Asia, are not considered to be endemic. Interestingly, recent molecular
studies have shown that the P. mascareniensis populations from Madagascar
are already sufficiently differentiated from those from mainland Africa, and
likely represent a different species (Vences et al. 2003, 2004).
59
Figure 1. Map of the Seychelles Islands.
and designation of this island as a national park would significantly enhance
amphibian conservation in Seychelles.
To date, there has been no evidence of any sudden amphibian decline in
Seychelles, although the potential impact of an invasion by chytridiomycosis
would be considerable. Monitoring programmes are in place for the frogs, and
are being developed for the caecilians. Updates on research and conservation
progress can be found at http://members.aol.com/jstgerlach/herps.htm
Justin Gerlach
References
Gerlach, J. 2000. Seychelles Amphibia – a mixture of secure and declining species.
Froglog 40(5).
Gerlach, J. and Willi, J. 2002. A new species of Sooglossus frog (Sooglossidae). Am-
phibia-Reptilia 23:445-498.
Vences, M., Kosuch, J., Glaw, F., Bohme W. and Veith, M. 2003. Molecular phylogeny
of hyperoliid treefrogs: biogeographic origin of Malagasy and Seychellean taxa and
re-analysis of familial paraphyly. Journal of Zoology and Systematic Evolutionary
Research 41:205-215.
Vences, M., Kosuch, J., Rodel, M.-O., Channing, A., Glaw, F. and Bohme, W. 2004.
Phylogeography of Ptychadena mascareniensis suggests transoceanic dispersal in a
widespread African-Malagasy frog lineage. Journal of Biogeography 31:593-601.
Malagasy amphibians are represented by four families of frogs (Gym-
nophiones and Urodeles being absent): Mantellidae, Microhylidae, Hyperolii-
dae, and Ranidae2, with ranids being represented only by the aforementioned
non-endemic Ptychadena mascareniensis and Hoplobatrachus tigerinus (Glaw
and Vences 1994). Mantellidae is the most speciose group and is endemic to
Madagascar and the Comoro Islands. This family includes the highly diverse
genus Mantidactylus (with nearly 90 species), the well-known genus Mantella
60
(15 species), Boophis (53 species), Aglyptodactylus (3 species), and the mono-
specific genus Laliostoma. At the time of writing, Mantidactylus has been
split into several genera (Glaw and Vences 2006), which differ significantly
in aspects of morphology, life history and distribution: Wakea, Blommersia,
Guibemantis, Spinomantis, Gephyromantis, Boehmantis, and Mantidactylus.
As a general trait, Mantidactylus (sensu lato) and the closely related Mantella
show peculiar reproductive features, such as the absence of amplexus and
of nuptial pads in males, with eggs laid outside water, and the presence
(in most species) of femoral glands, which are glandular structures on the
inside of the thighs, and related to reproductive behaviour. They also exhibit
a variety of life history traits, with species adapted to terrestrial, aquatic,
and arboreal habitats (Andreone and Luiselli 2003). The diurnal Mantella
species are characterized mainly by their bright aposematic colouration, small
size, and accumulations of alkaloids in the skin. The Mantella species are,
therefore, toxic, and are apparently rarely predated upon by other species,
a situation similar in many respects to that of the Neotropical dendrobatid
frogs (Clark et al. 2005).
In contrast to these frogs, amphibians in the genus Boophis are mainly
arboreal species, breed in water, and have a typical larval development.
Egg-laying usually occurs in streams, except for some species that reproduce
in lentic waters (Aprea et al. 2004; Glaw and Vences 2006). Two further
genera, Aglyptodactylus and Laliostoma, are mainly terrestrial and breed
in temporary ponds, often breeding explosively during which time they form
large aggregations. The genus Aglyptodactylus is also peculiar in having the
males that assume a somehow bright yellow colouration during the breeding
season (Glaw and Vences 1994, 2006).
Microhylidae are represented by 10 genera and more than 50 species
with a diverse life history. The cophyline microhylids (belonging to the gen-
era Cophyla, Platypelis, Anodonthyla, Plethodontohyla, Madecassophryne,
Rhombophryne, and Stumpffia) are closely tied to rainforest habitats and
have a reproduction that is characterized by the presence of parental care (for
example, both the parents, or at least one of them, remain with the tadpoles
during their development) and non-feeding tadpoles (Andreone et al. 2004).
The genera Dyscophus, Paradoxophyla, and Scaphiophryne are different in
this habitat preference, living mainly in open areas, but sometimes in arid
and sub-arid conditions (Andreone et al. 2006a). Most of them are mainly
terrestrial, although some species, such as S. gottlebei are partly rupicolous
and able to climb vertical walls within the narrow canyons of the Isalo
Massif (Andreone et al. 2005a). In particular, Scaphiophryne species have
tadpoles that are largely filter feeding, with some peculiar specializations
(see Mercurio and Andreone 2005).
The Malagasy Hyperoliidae includes the single endemic genus Heterix-
alus comprising eleven species, which inhabit grasslands and forest edges.
They are quite similar in habitat preferences and general behaviour to other
hyperoliids from mainland Africa (Glaw and Vences 1994).
Nine species of Malagasy frogs have been categorized as Critically
Endangered according to the IUCN Red List Categories and Criteria, namely:
Mantella aurantiaca, M. cowani, M. expectata, M. milotympanum, M. viridis,
Scaphiophryne gottlebei, Mantidactylus pauliani, Boophis williamsi, and
Stumpffia helenae (Andreone et al. 2005b). A further 21 species were classed
as Endangered and 25 as Vulnerable. In general, the main threat affecting
the local amphibian fauna is the high rate of forest loss (just less than 1%
per annum; Harper et al. 2007) (Figure 2). The different life history traits of
the amphibians are consequently mirrored by their differential ecological
sensitivity and conservation needs (Andreone and Luiselli 2003). In fact,
most of the Malagasy frogs inhabit the eastern rainforest, an ecosystem
that allowed the rapid diversification of some groups, such as Boophis,
Mantidactylus (sensu lato), and cophyline microhylids. The original eastern
rainforest block is now severely fragmented due to deforestation, and this
fragmentation has often resulted in high levels of threats among the native
amphibian fauna (Figure 3). However, in some cases this loss in amphibian
species richness is not immediately evident, because species have differing
sensitivities to habitat alterations (Andreone et al. 2005b).
Figure 1. Richness map of amphibian species in Madagascar, with dark red
colours corresponding to areas of higher richness. Colour scale based on 10
quantile classes. Maximum richness equals 91 species.
Threatened Amphibians of the World
In addition to the threat of habitat loss, some species (e.g. those belong-
ing to Mantella, Scaphiophryne, and Dyscophus) have been highly sought
after for the international pet trade due to their biological peculiarities and
remarkable colouration (Raxworthy and Nussbaum 2001; Andreone et al.
2006b; Mattioli et al. 2006). There is still a paucity of reliable data on the
impact of trade on wild populations, although in some cases it is evident
that collecting represents a confounding threat when the habitat is already
compromised and the populations are small. This is the case, for example, for
the rare harlequin mantella, Mantella cowani, which was collected in high
numbers and survived in a very degraded environment on the high plateau
of Madagascar (Andreone and Randrianarina 2003; Chiari et al. 2005). Fortu-
nately, the collecting and exportation of this species, classified as Critically
Endangered, are now banned, a measure that should reinforce its protection
(Andreone et al. 2006b). Due to concerns about overharvesting for commercial
trade, all frogs in the genus Mantella as well as the species Scaphiophryne
gottlebei are now included on CITES Appendix II. Other species are also of
conservation concern, such as the Tomato Frog, Dyscophus antongili, the only
native species included on CITES Appendix I. Although its distribution area is
wider than formerly believed (Andreone et al. 2006b), the habitat degradation
around the town of Maroantsetra, where most known populations occur, is
high. There is also evidence that the populations have apparently declined
in numbers, and the species appears to have vanished from sites at which it
was formerly known to occur (Chiari et al. 2006).
Fortunately, field surveys conducted during the last 15 years have revealed
no known extinctions of Malagasy amphibians resulting from habitat loss,
disease or other agents (Andreone et al. 2005b), as has been reported
elsewhere. On the other hand, 12 highly threatened species now have their
last remaining populations confined to a single site (Ricketts et al. 2005),
and several of these sites, such as the Ankaratra Massif and Fierenana,
remain unprotected. In general, the two areas with the majority of threatened
species are the northern Tsaratanana-Marojejy-Masoala highlands and the
southeastern Anosy Mountains (Andreone et al. 2005b).
Following the remarkable declaration by Malagasy president Marc
Ravalomanana to triple the existing coverage of the island’s protected areas
network (see Figure 2), amphibians are now beginning to be considered in
the identification of globally important sites for conservation (see Essay
11.3). This is all the more important since, as already noted, several highly
threatened species experience no protection whatsoever. For example, of
the nine Critically Endangered amphibians, six are not recorded from any
protected area (Andreone et al. 2005b, 2006b). Amphibians also represent
an excellent candidate to become a symbol for the conservation strategies in
Madagascar. Indeed, it is clear that, as with lemurs, the frogs of Madagascar
have the potential to become an important tool for the conservation of wildlife
in Madagascar. This aspect, as well as long-term conservation planning, was
the focus of a workshop (A Conservation Strategy for the Amphibians of
Madagascar) held in Antananarivo in September 2006 specifically dedicated
to the amphibians of Madagascar, and attended by more than 80 scientists
and conservation practitioners.
Franco Andreone and Herilala Randriamahazo
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the genus Paradoxophyla (Microhylidae: Scaphiophryninae) from Masoala rainforest,
northeastern Madagascar. Acta Herpetologica 1:15-27.
Andreone F., Mattioli F. and Mercurio V. 2005a. The call of Scaphiophryne gottlebei, a
microhylid frog from the Isalo Massif, south-central Madagascar. Current Herpetol-
ogy 24:33-35.
Andreone, F. and Luiselli, L.M. 2003. Conservation priorities and potential threats
influencing the hyper-diverse amphibians of Madagascar. Italian Journal of Zoology
70:53-63.
Figure 2. Forest cover map for Madagascar, and existing protected areas in
black. Red corresponds to forest clearance between 1990 and 2000, brown to
clearance between 1975 and 1990, and green to forest cover in 2000.
Andreone F. and Randrianirina J.E. 2003. It’s not carnival for the harlequin mantella!
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Vallan, D. and Vences, M. 2005b. Species Review of Amphibian Extinction Risks in
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adaptations revealed through a molecular analysis of the secretive cophyline frogs of
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phibians and reptiles in Madagascar. Amphibian and Reptile Conservation 2:15-23.
Ricketts, T.H., Dinerstein, E., Boucher, T., Brooks, T.M., Butchart, S.H.M., Hoffmann, M.,
Lamoreux, J.F., Morrison, J., Parr, M., Pilgrim, J.D., Rodrigues, A.S.L., Sechrest, W.,
Wallace, G.E., Berlin, K., Bielby, J., Burgess, N.D., Church, D.R., Cox, N., Knox, D.,
Loucks, C., Luck, G.W., Master, L.L., Moore, R., Naidoo, R., Ridgely, R., Schatz, G.E.,
Shire, G., Strand, H., Wettengel, W. and Wikramanayake, E. 2005. Pinpointing and
preventing imminent extinctions. Proceedings of the National Academy of Sciences
USA 102:18497-18501.
Vences, M., Vieites, D.R., Glaw, F., Brinkmann, H., Kosuch, J., Veith, M. and Meyer, A.
2003. Multiple overseas dispersal in amphibians. Proceedings of the Royal Society
B 270:2435-2442.
Vences, M., Kosuch, J., Rödel, M.-O., Lötters, S., Channing, A., Glaw, F. and Böhme,
W. 2004. Phylogeography of Ptychadena mascareniensis suggests transoceanic
dispersal in a widespread African-Malagasy frog lineage. Journal of Biogeography
31:593-601.
1 Although only some 226 species are currently classified through the Global Amphibian
Assessment, several new species were recently described, including Paradoxaphyla
tiarano from Masoalo Forest in north-eastern Madagascar (Andreone et al. 2006a).
2 The Ranidae were disaggregated into several families by Frost et al. (2006). In
Madagascar, Pthychadena is ascribed to Ptychadenidae and Hoplobatrachus to
Dicroglossidae. Thus, the number of families becomes five.
Figure 3. Richness map of threatened amphibian species in Madagascar, with
dark red colours corresponding to areas of higher richness. Colour scale based
on five quantile classes. Maximum richness equals nine species.

ESSAY 5.3. FROG BIODIVERSITY IN CAMEROON
Around 200 species of frogs occur in Cameroon (Amiet 1989), including sev-
eral unnamed taxa, but the final total could be as high as 210 due to limited
knowledge of the extreme north and south-east of the country. Comparisons
with the less well prospected neighbouring countries in central Africa are
difficult, but the frog fauna of Cameroon is almost certainly the second most
diverse in continental Africa, after the Democratic Republic of the Congo.
Components of Cameroon’s frog fauna
Cameroon’s frog fauna largely consists of four ecological components. The
savannah component (c. 35 species) is associated with grassy and herbaceous
habitats over a broad latitudinal extent, from the small savannahs near
Yaoundé, north to the steppes around Lake Chad. Very few species occupy
this entire area, most being distributed in latitudinal bands reflecting their
different tolerances of drought. All have wide distributions outside Cameroon,
and their tadpoles develop in still water.
The montane component, represented by about 50 species, many endemic,
is restricted not only to mountains but also to high plateaus in western
Cameroon. The boundary between this component and lowland faunas is
between 800 and 1,200m. These species prefer forests or are indifferent to
vegetation type, and for many the tadpoles develop in streams.
The forest component includes some 80 species that are generally confined
to closed-canopy forest. These species depend mainly on ecological condi-
tions created by a closed canopy, rather than on the exploitation history of
the forest (primary or secondary), or on floristic composition. Amiet’s (1989)
study of the Yaoundé forest frog fauna showed that it consisted overwhelm-
ingly of species living in leaf-litter (43%), or on vegetation (39%), with half
of the water-dependent species associated with streams. This component
includes many species and some genera that are, more or less, restricted to
Cameroon, especially in the west.
The “parasylvicolous” component includes some 30 species confined to
the forest zone and gallery forests in the savannah zone. These species do
not live in intact forest, but in habitats that have been disturbed, either as
a result of natural processes, or because of degradation by humans (Amiet
1989). Most of these species occur widely outside Cameroon, their tadpoles
developing in still or slow-flowing waters.
There is also a small component of about six species that occur widely in
the savannahs, and in degraded habitats in the forest zone, often in villages
and large urban areas.
These components are not homogeneous, and can be subdivided into
elevational zones (montane component) or latitudinal zones (savannah
component). However, the limits of these zones are less clear than those
that separate the components themselves.
Causes of the diversity of Cameroon’s frog
fauna
Several factors contribute to the high level of frog diversity in Cameroon.
The first is the wide variety of natural conditions in the country. An old
tourist advertisement boasts that Cameroon is “the whole of Africa in just
one country”, which is not an exaggeration. Diversity is boosted by interplay
between climatic and geomorphological factors. The country covers over 12°
of latitude, including most of the climatic gradient of western and central
Africa north of the equator, from a very rainy equatorial to tropical climate
(with rainfall decreasing as one goes northwards).
This pattern is modified by two major axes of relief. The first is the
Cameroon mountain ridge, a series of massifs, often of volcanic origin with
peaks over 2,000m, the highest being Mount Cameroon (4,095m). The ridge
runs SSW-NNE, with numerous southern and western slopes exposed to the
monsoon rains. The second is the Adamawa Plateau, which, at an altitude
of 1,000-1,200m, crosses the entire country between 6° and 8°N, and has
a relatively cool and humid climate. Thus, there is a complex patchwork of
rainfall patterns in Cameroon, analysed in depth by Suchel (1972, 1988). In
Astylosternus ranoides (Endangered) is known only from western Cameroon, where it has been recorded from the
Bamboutos Mountains, Lake Oku, and Mount Neshele, at altitudes of 2,000-2,600m asl. © Jean-Louis Amiet
Chapter 5. Amphibians of the Afrotropical Realm
addition to affecting the overall climate, these high plateaus and mountain
massifs also display vertical climatic zoning, resulting in the diversification
of ecological conditions over very short distances.
While the diversity of current climatic conditions undoubtedly contributes
to the diversity of Cameroon’s frog fauna, climatic changes during the Qua-
ternary are the main cause. These changes (see Maley 1996, 2001) resulted
in repeated modifications to the distribution of bioclimatic zones, and have
been a major cause of frog speciation, affecting in particular lowland forest
and montane species. Lowland tropical and montane forests both experienced
phases of expansion and regression. When habitats became fragmented,
allopatric speciation took place in isolated areas, causing remarkable diver-
sification in genera such as Cardioglossa, Astylosternus or Leptodactylodon,
which include pairs and trios of closely related species. Some of these close
relatives have remained allopatric, while others have become sympatric or
even syntopic (Amiet 1980, 1987). However, Quaternary climatic changes are
too recent to explain the presence of 15 genera, endemic, or largely endemic,
to Cameroon (Amiet 1989). Most of these have tadpoles that develop in well-
oxygenated, running water, a habitat that is plentiful in the hilly highlands of
the west and south-west. These genera presumably evolved in situ during
the formation of the Cameroon mountain ridge.
More recently, human impact on the environment has had favourable
consequences on savannah and “parasylvicolous” species. Maintenance
of savannahs by fire since the last episode of forest regression has aided
the expansion of savannah species (for example, in the northern part of the
southern Cameroon Plateau, which under current climatic conditions should
be forest, but which is in fact dominated by savannah). In the forest zone,
agricultural clearings have provided habitats favourable to “parasylvicolous”
species, especially with the increasing destruction of forests since the start
of the colonial period. Some “parasylvicolous” species of western origin,
now quite common, were not found by the first collectors, and some of
these might have reached Cameroon in the last 150 years, and continue to
expand towards the east or south-east. Judging by the speed at which certain
“parasylvicolous” species settle in newly opened forest sites, this hypothesis
does not seem outlandish.
Geographic patterns of diversity
Figure 1 shows how the frog fauna is distributed in relation to the main biocli-
matic units of the country. In regions Ic, II, Va and Vb, the numbers of species
have probably been underestimated, but regions Ia, Ib and III will remain the
centres of Cameroon’s frog diversity. The high diversity in regions Ia and III
is due to the large number of endemic and near-endemic species. Region III
includes virtually all of Cameroon’s montane species, none of which occur in
other mountainous regions in Africa. Regions Ia and Ib include several “para-
sylvicolous” species (those in Ia and Ib being of “western” and “Congolese”
origin, respectively). However, the higher diversity in Ia is due to the presence
of several endemic species on the coastal plain, especially between the lower
Sanaga and the Mungo rivers, and also south of the lower Nyong River (both
probably being forest refugia during past climatic fluctuations).
Threats to Cameroon’s frog fauna
The ecological conditions and the overall diversity and endemism of frogs vary
across the regions of Cameroon, and this affects the threats facing the frog
fauna. Moreover, future environmental changes in the context of the country’s
socio-economical and political situation need to be considered.
The frog faunas in regions IV, Va and Vb are not at great risk, despite
these regions being greatly affected by human activity, as most of the species
are adaptable. Conversely, in region II, the savannah species are, in theory
at least, doomed to eventual extinction, as the forest should be expanding
rapidly under current climatic conditions. However, the species in question
also occur in regions IV and V, and forest species would presumably spread
in region II.
Figure 1. Map of Cameroon show-
ing the major bioclimatic regions,
with the number of species recorded
from each region. Key for bioclimatic
regions: Ia: coastal plain tropical
lowland forest (92 species); Ib:
southern Cameroonian plateau trop-
ical lowland forest (83); Ic: Congo
basin tropical lowland forest (57);
II: southern Cameroonian plateau
gallery forests and Guinea savan-
nah (51); III: Cameroon mountain
ridge mountains and high plateaus
(78); IV: Adamawa Plateau (40); Va:
northern Cameroon Sudan savannah
(23); and Vb: northern Cameroon Su-
dano-Sahelian savannah and Sahel
(29). For methodology in defining
these regions and estimating spe-
cies totals, see (Amiet 1983).
61
The situation is much worse for the forest species in regions Ia, Ib and Ic,
with the increasingly serious loss of forest, which seems inevitable due to lack
of political commitment to conservation. However, frogs are less seriously
affected than large mammals because, thanks to their small size, they are
often able to survive in what little forest remains. They can also survive in
areas of shifting cultivation where rotations allow for rapid recovery of the
forest cover. Logging has limited impact as there is very little clear-cutting in
Cameroon, but commercial tree plantations (such as oil palm) do have a very
negative impact on forest frogs, and are especially prevalent in region Ia.
Region III has lost much forest, especially on the Bamileke Plateau.
However, on the humid southern and western mountain slopes cleared
forest can be replaced quickly by dense secondary vegetation suitable for
many montane frogs (few species are completely dependent on intact forest).
However, the tendency of the Bamileke and Banso peoples to clear large
areas of habitat is alarming. For frogs living at very high elevations in open
habitats, the threat level is low.
The Goliath Frog Conraua goliath (EN) requires particular mention as
the only directly threatened frog in Cameroon. In the past it was sold in
large numbers to at least one American importer for the ridiculous practice
of jumping competitions. Now, these frogs, which used to be eaten only
occasionally, have become prized game actively sought after, at least in the
area of Nkongsamba (G. Renson, pers. comm.). Special traps have even been
invented for catching this species, a victim of the current fad for bushmeat.
With lack of law enforcement and endemic corruption, conservation measures
are ineffective, and the future of the Goliath Frog depends on a change in
food habits among local people.
The most effective measure of conservation for forest and montane frogs
in Cameroon would be total preservation or restoration of a 30m-wide band
of forest on both sides of water courses, whatever their size and location
(including in commercial plantations). This solution, based on this author’s
29 years of field research in Cameroon, would safeguard a huge portion of
biodiversity in all taxonomic groups, and would contribute to quantitative
and qualitative improvements in water reserves. Such a measure, if it were
adopted, would require the means, and above all the political will, to imple-
ment in a resolute manner.
Jean-Louis Amiet
References
Amiet, J.-L. 1980. Révision du genre Leptodactylodon (Amphibia Anura, Astylosterninae).
Annales de la Faculté des Sciences de Yaoundé 27:69-224.
Amiet, J.-L. 1987. Aires disjointes et taxons vicariants chez les Anoures du Cameroun:
implications paléocliamtiques. Alytes 6:99-115.
Amiet, J.-L. 1989. Quelques aspects de la biologie des Amphibiens Anoures du Cameroun.
Année biologique 28:73-136.
Maley, J. 1996. The African rain forest - main characteristics of changes in vegetation
and climate from the upper Cretaceous to the Quaternary. Proceedings of the Royal
Society of Edinburgh 104B:31-73.
Maley, J. 2001. The impact of arid phases on the African rain forest through geological
history. In: W. Weber, L.J.T. White, A. Vedder and L. Naughton-Treves (eds.), African
Rain Forest Ecology and Conservation, pp. 68-87. Yale University Press, New Haven,
Connecticut, USA.
Schiøtz, A. 1999. Treefrogs of Africa. Chimaira, Frankfurt am Main, Germany.
Suchel, J.-B. 1972. La répartition des pluies et les régimes pluviométriques au Cameroun.
Travaux et documents de géographie tropicale, 5. Université fédérale du Cameroun
et C.N.R.S.
Suchel, J.-B. 1988. “Les climats du Cameroun”. Thèse de Doctorat d’Etat, Université de
Saint-Etienne, France. t. I-III: 1188 pp., et t. IV, atlas.
62 Threatened Amphibians of the World

ESSAY 5.4. THREATENED ISLANDS OF AMPHIBIAN DIVERSITY IN WEST AFRICA

160
140
120
# of described species

100
80
60
40
20
0
1830 1850 1870 1890 1910 1930 1950 1970 1990 2010

Figure 1. Number of described West African amphibian species per decade (including known but not yet described new species for the last decade). Black
arrows indicate the start of periods of more intensive survey and taxonomic work. These significant increases were mainly due to (arrows from left to right)
P. Chabanaud; J. Guibé, M. Lamotte and co-workers; A. Schiøtz and J.-L. Perret and M.-O. Rödel and co-workers. With the exception of the 1960s (A. Schiøtz)
new amphibian species have been described at a near constant rate of about 0.7 species per annum. Since 2000 the description rate increased to more than 1.8
species per annum and is not yet reaching any visible plateau, hence illustrating that the West African amphibian fauna still is far from being well known.

West Africa, here defined as the region from Senegal in the west to Nigeria expect that several species have already gone extinct without having been
in the east and extending north to the southern border of the Sahara Desert, discovered and described scientifically.
covers almost all larger African biomes, ranging from rainforests along the But even selective logging might be a severe danger to the endemic
Atlantic Ocean’s coast, over various types of savannahs to semi-deserts. In West African forest frogs. We have shown that habitat degradation mark-
the border region of Guinea, Côte d’Ivoire, and Liberia, as well as within edly alters the composition of forest frog communities, and many endemic
Sierra Leone, there are also several higher mountain ranges representing rare and range-restricted forest frogs do not prevail in logged or fragmented
examples, at least in the West African context, of montane grassland. forests (Ernst and Rödel 2005). Moreover, not only does forest degradation
Amphibian research in West Africa dates back to the middle of the 19th negatively affect particular species, but it severely and non-randomly reduces
century. In general, the diversity of amphibians in West Africa is high, with the functional diversity of forest species assemblages, i.e., specific life his-
around 175 amphibian species having been recorded from this region, includ- tory traits disappear (Ernst et al. 2006). The latter is particularly important
ing four caecilians and 171 anurans. While some forests may support as many for ecosystem renewal and reorganization following change and it provides
as 40 different amphibian species in an area as small as 2 ha, amphibian adaptive capacity in a world of complex systems, and human-dominated
richness is also high in some savannah formations (sometimes surpassing 30 environments. The loss especially concerns species with tadpoles that are
frog species in only a few square kilometres). However, while new species dependant on larger stagnant or slow-flowing waters. In conclusion, West
have been described on a continual basis, species description curves have Africa still supports an extraordinary rich amphibian fauna, albeit restricted
never attained any sort of saturation (Figure 1) suggesting that many new to steadily declining islands of diversity.
species remain undiscovered.
Around one-fifth of the region’s amphibian fauna is considered to be Mark-Oliver Rödel, P. Joël Adeba, Raffael Ernst, Annika Hillers,
threatened (34 species), including 14 species that are listed as Vulnerable, S. Gilles A. Nago, Johannes Penner and Martin Wegmann
16 Endangered, and four Critically Endangered. A further 19 species are clas-
sified as Near Threatened. The four Critically Endangered species include two
species, Nimbaphrynoides occidentalis and N. liberiensis, known only from References
a few square kilometres on Mount Nimba’s grassland mountain ridge and a
very limited area in Liberia’s rainforest on Mount Nimba’s southern slopes, Bakarr, M., Oates, J., Fahr, J., Parren, M., Rödel, M-O. and Demey, R. 2004. Guinean Forests
respectively. A third species, Bufo taiensis, is only known from four specimens of West Africa. In: R.A. Mittermeier, P. Robles-Gil, M. Hoffmann, J.D. Pilgrim, T.M.
collected from the region of Taï National Park in Côte d’Ivoire, while the fourth Brooks, C.G. Mittermeier, J.L. Lamoreux and G. Fonseca (eds.), Hotspots: Revisited,
species, Conraua derooi (currently considered Possibly Extinct on the IUCN Red pp. 123-130. CEMEX, Mexico City, Mexico.
list), was described from, and only very recently rediscovered, in the southern Chatelain, C., Gautier, L. and Spichiger, R. 1996. A recent history of forest fragmentation
part of the Ghanaian-Togolese mountainous borderline. in southwestern Ivory Coast. Biodiversity and Conservation 5:783-791.
Many West African forest amphibians have very small ranges, possibly due Ernst, R. and Rödel, M.-O. 2005. Anthropogenically induced changes of predictability in
to fluctuations in the region’s forest cover (Wieringa and Poorter 2004). These
fluctuations presumably also served as a catalyst for amphibian speciation and
are a reason for today’s high diversity. In the Pleistocene and pre-Pleistocene,
rainforests were either increasing towards the north in humid periods or were Figure 2. Endemism of West African amphibian species. Shown are 42
shrinking to comparatively small forest refugia during drier periods (Falk et well-known amphibian communities and the respective proportion of endemic
al. 2003). The recent distributions of West Africa’s endemic frogs matches species. The level of endemism ranges from local (red, e.g. Nimbaphrynoides
well with the rough location of these postulated forest refugia, namely south- occidentalis, estimated range: 111km²) and regional (orange, e.g. Kassina la-
western Ghana (Rödel et al. 2005), south-western Côte d’Ivoire (Rödel and mottei, 13,002km²) endemics, to species that occur in the Upper Guinea forest
Branch 2002), and the Mount Nimba area (Rödel et al. 2004; Figure 2). zone (yellow), West Africa (clear green), Central and West Africa (green, e.g.
However, new evidence lends support for a much finer grained picture, Phlyctimantis boulengeri, 294,445km²) to species with an almost sub-Saharan
adding a few more distinct Pleistocene or Pre-Pleistocene forest remnants to distribution (dark green, e.g. Kassina senegalensis, 12,263,903km²).
the Upper Guinea highlands (Rödel et al. unpubl.). The Upper Guinea highlands
region stretches from Sierra Leone and Guinea through Liberia to western Côte
d’Ivoire, and forms part of the Guinean Forests of West Africa biodiversity
hotspot (Bakarr et al. 2004). In order to understand West Africa’s forest history
and to reveal potentially overlooked centres of endemism, an accurate knowl-
edge of the location of historical forest refugia is needed. Current research
aims to locate these former forest refugia by investigating the relationships
between various leaf litter frog species and populations from West African
forests. The recent distribution patterns of these amphibians and the genetic
divergences of populations of these frogs will help us to reconstruct the history
of West Africa’s forest cover (Hillers et al. unpubl.).
On this regional scale we have also started to analyse landscape charac-
teristics such as climate, vegetation, altitude, and topo-diversity with respect
to their potential influence on amphibian species richness. Our analyses have
revealed positive correlations of species richness with rainfall, forest cover,
and habitat diversity. Based on these three parameters, we have been able to
model areas believed to harbour a high richness of amphibian species. These
areas coincide with the aforementioned areas of former forest refugia, which
hence would be not only islands of high endemism but also of corresponding
high amphibian diversity (Penner et al. unpubl.).
Unfortunately, these areas are also among the most threatened regions in
West African. South-western Côte d’Ivoire, the area with the most extensive
tracts of remaining forest in the country, lost about 80% of its forests within
the last 30 years (Chatelain et al. 1996). Guinea has little more than about
5% of its former rainforests left (UICN 1996), and many of its remaining
forested mountain ranges are threatened by open-cast mining operations.
Given the distributional limitations of many forest amphibians and the vast
tracts of forest areas that have already been lost, it is not unreasonable to
 Chapter 5. Amphibians of the Afrotropical Realm 63

tropical anuran assemblages. Ecology 86:3111-3118.
Ernst, R., Linsenmair, K.E., and Rödel, M.-O. 2006. Diversity erosion beyond the species
level: Dramatic loss of functional diversity after selective logging in two tropical
amphibian communities. Biological Conservation 133:143-155.
Falk, T.M., Teugels, G.G., Abban, E.K., Villwock, W. and Renwartz, L. 2003. Phylogeographic
patterns in populations of the blackchinned Tilapia complex (Teleostei, Cichlidae)
from coastal areas in West Africa: support for the refuge zone theory. Molecular
Phylogenetics and Evolution 27:81-92.
Rödel, M.-O., Bangoura, M.A. and Böhme, W. 2004. The amphibians of south-eastern
Republic of Guinea (Amphibia: Gymnophiona, Anura). Herpetozoa 17:99-118.
Rödel, M.-O. and Branch, W.R. 2002. Herpetological survey of the Haute Dodo and Cavally
forests, western Ivory Coast, Part I: Amphibians. Salamandra 38:245-268
Rödel, M.-O., Gil, M., Agyei, A.C., Leaché, A.D., Diaz, R.E., Fujita, M.K. and Ernst, R.
2005. The amphibians of the forested parts of south-western Ghana. Salamandra
41:107-127.
UICN 1996. L’atlas pour la conservation des forêts tropicales d’Afrique. Editions Jean-
Pierre de Monza, Paris, France.
Wieringa, J.J. and Poorter, L. 2004. Biodiversity hotspots in West Africa; patterns and
causes. In: L.F. Poorter, F. Bongers, F. Kouamé and W.D. Hawthorne (eds.), Biodiversity
of West African forests. An ecological atlas of woody plant species, pp. 61-72. CABI
Publishing, Cambridge, Massachusetts, USA.

ESSAY 5.5. THE AMPHIBIAN FAUNA OF THE EASTERN ARC MOUNTAINS OF KENYA AND TANZANIA

In 1983 three botanists – Jon Lovett, Roger Polhill and John Hall – were sitting Spray Toad Nectophrynoides asperginis (CR) occurring only in the Kihansi Rovero, F., Stanley, W.T., Stuart, S.N. 2007. The biological importance of the Eastern
together at the foot of the Uluguru Mountains discussing recent discoveries gorge, and now possibly extinct in the wild. This species is adapted to the Arc Mountains of Tanzania and Kenya. Biological Conservation 134:209-231.
of rare plants in the Udzungwa Mountains, previously thought to be endemic peculiar habitat influenced by the constant spray provided by waterfalls. In Davenport, T.R.B., Machaga, S.J., Mwamtobe, A., Mwaipungu, O. and Hayes, B. 2003.
to the Usambaras and Ulugurus (Lovett 1998). From the analysis they were this case, the small distribution is explained by the extremely peculiar local Some notes on the amphibians of Mt Rungwe. Unpublished Report. Southern Highlands
making, it was clear that these rare forest plants were found only on the conditions in the lower Kinhasi gorge. The other three species inhabit sites Conservation Programme / Wildlife Conservation Society (SHCP/WCS). 4 pp.
crystalline peaks of a series of isolated mountain ranges in south-eastern with no apparent special conditions; they have been sought unsuccessfully Loader, S.P., Gower, D.J., Howell, K.M., Doggart, N., Rödel, M.-O., Clarke, B.T., De Sá,
Kenya and Tanzania. These mountains were showing remarkable similarities in other suitable sites within the same forest patch. Differences thus exist R.O., Cohen, B.L. and Wilkinson, M. 2004. Phylogenetic relationships of African
in terms of species richness and composition and since they are set in a half- between assemblages at sites at similar altitudes in adjoining mountain microhylid frogs inferred from DNA sequences of mitochondrial 12S and 16S rRNA
moon ‘arc’ shape, they decided to name them collectively as the Eastern Arc fragments or within single fragments, indicating a fine-scale geographic genes. Organisms, Diversity and Evolution 4:227-235.
Mountains (Lovett 1998). The name first appeared in an article written by Jon turnover in the herpetofauna of these areas. Lovett, J.C. 1985. Moist forests of Tanzania. Swara 8:8-9.
Lovett for the Kenyan magazine Swara, in 1985. The distinct nature of the herpetofauna species assemblages at high Lovett, J.C. 1998. Naming the Arc. The Arc Journal 7.
The Taita Hills of Kenya and the Pare, Usambara, Nguru, Nguu, Ukaguru, altitude, and the high elevational turnover of species, clearly demonstrates Lovett, J.C., Marchant, R., Taplin, J., Küper, W., 2004. The oldest rainforests in Africa: stabil-
Uluguru, Rubeho, Udzungwa, Malundwe and Mahenge Mountains in Tan- the importance of conserving forest at all altitudes. Areas of forest or ity or resilience for survival and diversity? In: A. Purvis, J.L. Gittleman, and T.M. Brooks
zania comprise the Eastern Arcs. They lie within the intertropical montane marginal habitats, at all elevations, might also be vital in generating high (eds.), Phylogeny and Conservation. Cambridge University Press, Cambridge, UK.
region and were formed by heavily metamorphosed Pre-Cambrian basement species diversity, and need to be considered in the development of a holistic Menegon, M., Salvidio, S., 2005. Amphibian and Reptile diversity in the southern Uzungwa
rocks, rising to 2,635 m in elevation (Kimhandu peak in the Ulugurus). They conservation strategy for the area. This suggests that not only is it important Scarp Forest Reserve, South-Eastern Tanzania. In: B.A. Huber, B.J. Sinclair and K.H.
are part of one of the Earth’s richest biodiversity hotpots and amongst the to conserve the forest along an elevational gradient, but it is also important Lampe (eds.), African Biodiversity: Molecules, Organisms, Ecosystems, pp. 205-212.
most important regions in Africa for concentrations of endemic animals to conserve fragments in geographically complex terrains, where isolated Proceedings of the 5th International Symposium on Tropical Biology, Museum Koenig,
and plants, with at least 93 species of endemic vertebrate (Burgess et populations and therefore potentially new species may still exist, and are Bonn, Germany.
al. 2007). awaiting discovery. Moyer, D.C. 1993. Foraging ecology, habitat selection, and community structure of
The Danish batrachologist Arne Schiøtz published a short paper highlight- afromontane forest birds in Tanzania. M.Sc. thesis, Louisiana State University and
ing the importance of the basement hills of Tanzania (the name Eastern Arc Michele Menegon and Tim Davenport Agricultural and Mechanical College, Department of Zoology and Physiology. Baton
was still to be coined) as a regional centre of amphibian endemism (Schiøtz Rouge, Louisiana, USA.
1981). The Eastern Arcs are home to about 94 named amphibian species, Schiøtz, A. 1981. The Amphibia of the forested basement hills of Tanzania - a biological.
of which 57 are endemic or near-endemic. A further 17 species have been References indicator group. African Journal of Ecology 19:205-208.
recently discovered and are awaiting formal description (all of which are Wilkinson, M., Loader, S.P., Gower, D.G., Sheps, J.A. and Cohen, B.L., 2002. Phylogenetic
probably endemic). A rough estimate, therefore, of the total number of Burgess, N.D., Butynski, T.M., Cordeiro, N.J., Doggart, N., Fjeldså, J., Howell, K.M., Kila- Relationships of African Caecilians (Amphibia: Gymnophiona): Insights from Mitochon-
amphibians (described and undescribed) in the Eastern Arc Mountains is hama, F., Loader, S.P, Lovett, J.C., Menegon, M., Moyer, D.C., Nashanda, E., Perkin, A., drial rRNA Gene Sequences. African Journal of Herpetology 52:83-92.
121 species, of which 74 occur are endemic or nearly so. The remarkable
species richness and the high level of endemism is due to the great age of
the Eastern Arc Mountains (they uplifted at least 30 million years ago), to
their archipelago-like arrangement, and to the climatic influence of the Indian
Ocean that kept the mountains relatively wetter and warmer than the sur-
rounding areas during past climatic fluctuations. The high number of endemic
species in small areas, the co-occurrence of recent and old lineages, and the
consequent biogeographical implications make these mountains of extreme
biological and conservation interest.
Historically, the Eastern Arc Mountains were probably covered by a
mosaic of rainforests. These rainforests were concentrated mostly on the
eastern slopes and on the upland plateaux and were interspersed with
open grassland areas and with dry, semi-deciduous forests on the western
slopes. The majority of endemic species occur in the wet forest that covered
parts of the eastern slopes of the mountains. Although the forest environ-
ment has been well-studied, the grasslands and ecotones could represent
a further frontier of batrachological exploration. At the southern tip of the
Udzungwas, the so called ‘Makambako gap’ has long been considered the
southernmost limit. However, recent cross-taxon studies are revealing that
the Southern Highlands, especially the forests of Mt Rungwe and Livingstone,
also contain many species previously assumed to be endemic to the Eastern
Arcs. For example, amphibian species such as Nectophrynoides viviparus (VU),
Nectophrynoides tornieri (VU), and Leptopelis barbouri (VU) demonstrate Mt
Rungwe’s Eastern Arc affinities (Davenport et al. 2003).
Recently, molecular analysis has shown that a number of genera and
species in the Eastern Arc Mountains are genetically ancient. For example,
DNA sequence data of caecilians, including Eastern Arc species, suggest
that the origin of the caecilian fauna of Africa may ‘predate the break-up of
Gondwana’ (Wilkinson et al. 2003). Similar suggestions are made concerning
the microhylids and brevicipitids fauna of Africa based on DNA sequence
data, which have exposed the extremely ancient age of these lineages
(Loader et al. 2004). The present pattern of distribution and the presence
of recent and old lineages could be interpreted as a consequence of stable
local conditions, and to some degree the pattern can be attributed to local
speciation or to local low extinction rates – in other words, long species
persistence (Lovett et al. 2004).
Among the taxa occurring in the Arc, forest bufonids are of particular
interest with a number of endemic genera and species like the extraordinary
Churamiti maridadi (CR), with its shining skin resembling wet lichen, or the
bicoloured Nectophrynoides viviparus. Ongoing molecular studies on forest
bufonids are revealing the presence of lineages of East African origin as well
as others derived from Guineo-Congolian taxa and ancient African-Asian
linkages. Forest bufonids in the Arc are revealing a much more abundant
species radiation than expected with several new species and genera
awaiting description.
Many of the endemic species are confined to high-elevation sites and their
distribution pattern may be due to relictualization. Other endemic species
demonstrate intriguing and highly restricted ranges, deserving of special
conservation strategies. These species display a ‘single site’ distribution,
since they are confined to a single valley or parts of it, at certain elevations,
and yet are surrounded by apparently suitable habitat. In just the Uzungwa
Scarp Forest Reserve there are three strictly endemic species showing
such distribution patterns: two dwarf forest bufonids, Nectophrynoides
wendyae (CR) and N. poyntoni (CR), and the treefrog Hyperolius kihangensis
(EN)(Menegon and Salvidio 2005). Another ‘single site’ species is the Kihansi Figure 1. Map showing the 13 blocks of the Eastern Arc Mountains of Tanzania and Kenya (reproduced, with permission, from Moyer 1993).
64
ESSAY 5.6. THREATENED AMPHIBIANS OF SOUTHERN AFRICA
Although many species of amphibians have been described from Africa in
recent years, amphibians as a group remain poorly known relative to other
vertebrate taxa. An exception is South Africa, which has perhaps the best-
studied fauna in the region, culminating in the Atlas and Red Data Book of the
Frogs of South Africa, Lesotho and Swaziland (Minter et al. 2004). The larger
southern African region, including Botswana, Namibia, Mozambique, Malawi
and Zimbabwe, is home to 185 species of amphibians, of which 32 species
(17%) have been assigned a status of conservation concern, including five as
Critically Endangered, 16 as Endangered, and 11 as Vulnerable.
These threatened species have several things in common: they mostly
occupy small areas, either on coastal plains, or in highland or montane
grasslands or forests. The coastal areas are preferred localities for housing
developments, and for farming wheat and other crops. The highlands and
mountains are areas with high rainfall, and are selected for planting huge
tracts of pine plantations. These threatened species evolved on highlands
where there was sufficient moisture, often separated from other highlands by
dry plains. The moist conditions that promoted speciation, thereby assisting
the formation of a rich diversity of amphibians (see, for example, Channing
et al. 2002, 2005; Clarke & Poynton 2005) are the same environmental condi-
tions that are suitable for growing trees and other crops. Agriculture leads
to severe habitat modification and fragmentation. All the well-known areas
planted with exotic pines, such as the Amatola Mountains, the Elandsberg
Mountains, and even Table Mountain in Cape Town, are home to endemic
species of amphibians that are now threatened with extinction due to habitat
loss and conversion. Furthermore, many of these species may be at risk of
infection with the fast-spreading chytrid fungus, which is responsible for
the extinction of amphibian species in many parts of the world (Daszak et
al. 1999; Mendelson et al. 2006).
One-quarter of threatened species do not have normal life-cycles with
free-swimming tadpoles. Instead, their tadpoles remain within the egg, and
develop directly into small frogs. This negates the need for nearby pools of
water, enabling these species to survive in areas where there is little chance
of pools forming, such as in very sandy areas, forests and mountain tops.
However, the species with tadpoles display some very unusual breeding
strategies: some deposit eggs out of water, and the tadpoles must either
move to water, or wait until the nest is flooded. The tadpoles of these spe-
cies also display a range of adaptations. Most develop in streams or quiet
pools, but the tadpoles of Broadley’s Ridged Frog Ptychadena broadleyi (EN)
live on wet rocks in forests, and both species of ghost frogs Heleophryne sp.
have tadpoles that develop in fast-flowing rocky streams. The tadpoles hang
onto rocks in these turbulent habitats by using a large sucker-like mouth. The
Chirinda Toad Stephopaedes anotis (EN) breeds in small pockets of water
trapped in tree-holes.
Protected areas have proved to be the most important conservation tactic
in southern Africa, provided that the conservation area is large enough to
include a viable population size of the threatened species. Protected areas
Afrana johnstoni (Endangered) is known only from montane grassland and forest habitats on Mount Mulanje in
southern Malawi. © Alan Channing
Threatened Amphibians of the World
that provide a haven for globally threatened amphibians include: Inyanga
National Park, Chimanimani National Park and Chirinda Forest on the eastern
highlands of Zimbabwe (Afrana inyangae, EN; Bufo inyangae, EN; Arthro-
leptis troglodytes, CR; Probreviceps rhodesianus, EN; Stephopaedes anotis,
EN; Strongylopus rhodesianus, VU); Mt Mulanje Forest Reserve in Malawi
(Afrana johnstoni, EN; Nothophryne broadleyi, EN; Arthroleptis francei, EN;
Ptychadena broadleyi); Nyika National Park in Malawi (Bufo nyikae, VU), and
Table Mountain National Park (Capensibufo rosei, VU; Heleophryne rosei, CR)
on the Cape Peninsula of South Africa. Smaller reserves occur along the south
and east coasts of South Africa, though these may not be adequate to sustain
viable populations of threatened species in the region, such as Knysna Spiny
Reed Frog Afrixalus knysnae (EN), Natal Spiny Reed Frog Afrixalus spinifrons
(VU), and Pickersgill’s Reed Frog Hyperolius pickersgilli (EN).
Several threatened species occur almost entirely within strictly man-
aged, albeit different, areas. For example, the Cape Platanna Xenopus gilli
(EN) occurs within the well-established Table Mountain National Park, with
other populations in the newly proclaimed and still developing Agulhas
National Park. However, even within these national parks, its security cannot
be guaranteed, principally due to hybridization with the Common Platanna
Xenopus laevis (LC) (Kobel 1981; Picker et al. 1996) although the extent of this
problem may be limited (Evans et al. 1997). The Desert Rain Frog Breviceps
macrops (VU) occurs in a narrow coastal zone along the west coast of South
Africa and southern Namibia. This is a diamond mining area where access
is restricted, and no development is permitted. However, the diamonds are
mined by removing all the old beach sand above the bedrock. This effectively
destroys the habitat where this species is found. Of course, even within many
protected areas, a lack of management means that natural forest is still being
removed, and agricultural activities (both subsistence and large-scale) have
been reported from within these. Most (81%) threatened species occur largely
(72%) or entirely (9%) outside of any conservation area.
Active protection of threatened species does occur in some cases, as in
the Western Cape province of South Africa, where CapeNature was able to
both thwart construction plans for a road that would have had devastating
impacts on the breeding habitat of the Critically Endangered Micro Frog
Microbatrachella capensis, and establish a new breeding site. This species
is also being actively monitored by CapeNature, although continual active
intervention by local authorities is required to halt the damaging effects of
alien invasive plants on the breeding sites. As far as can be determined,
the only threatened species that are subject to official long-term monitoring
activities are Rose’s Ghost Frog Heleophryne rosei (CR), the Micro Frog, and
Cape Platanna. Among local conservation authorities, only CapeNature seems
to be paying sufficient attention to the problem of threatened amphibians. In
many other protected areas, including those to the north of southern Africa, it
is not unusual for staff of reserves to be unaware of the importance of their
reserve as a haven for a population of a globally threatened species.
Conservationists are able to determine effectively which species are
Table Mountain National Park on the Cape Peninsula of South Africa is home to the only known population of the
eponymous Table Mountain Ghost Frog Heleophryne rosei (Critically Endangered), a cryptic species found from 240-
1,060m asl. © Richard Boycott
threatened. We are also able to determine which species require the most
appropriate conservation response, be it the establishment of a protected
area or controlling invasive species. However, it seems that we are unable to
initiate long-term programmes involving active management. Will we sit on
the sidelines and tick off the species as they become extinct? Where there
was once a rallying call from African conservationists, ‘Save the Rhino!”, we
now need to encourage a ‘Save the Frog!” campaign.
Alan Channing and Andrew Turner
References
Channing, A., Moyer, D.C. and Howell, K.M. 2002. Description of a new torrent frog in the
genus Arthroleptides from Tanzania (Amphibia, Anura, Ranidae). Alytes 20:13-27.
Channing, A., Menegon, M., Salvidio, S. and Akker, S. 2005. A new forest toad from
the Ukaguru Mountains, Tanzania (Bufonidae: Nectophrynoides). African Journal of
Herpetology 54:149-157.
Clarke, B.T. and Poynton, J.C. 2005. A new species of Stream Frog, genus Strongylopus (An-
ura:Ranidae) from Mount Kilimanjaro, Tanzania, with comments on a ‘northern volcanic
mountains group’ within the genus. African Journal of Herpetology 54:53-60.
Daszak P., Berger L., Cunningham, A.A., Hyatt, A.D., Green, D.E. and Speare, R. 1999.
Emerging infectious diseases and amphibian population declines. Emerging Infectec-
tious Diseases 5:735-48.
Evans, B.J., Morales, J.C., Picker, M.D., Kelley, D.B. and Melnick, D.J. 1997. Comparative
molecular phylogeography of two Xenopus species, X. gilli and X. laevis, in the south-
western Cape Province, South Africa. Molecular Ecology 6:333-343.
Kobel, H.R., du Pasquier, M. and Tinsley, R.C. 1981. Natural hybridization and gene intro-
gression between Xenopus gilli and Xenopus laevis laevis (Anura:Pipidae). Journal
of Zoology 194:317-322.
Minter, L.R., Burger, M., Harrison, J.A., Braack, H.H., Bishop, P.J. and Kloepfer, D. (eds.).
2004. Atlas and Red Data Book of the Frogs of South Africa, Lesotho and Swaziland.
Smithsonian Institution, Washington D.C., USA.
Mendelson, J.R., Lips, K.R., Gagliardo, R.W., Rabb, G.B., Collins, J.P., Diffendorfer, J.E.,
Daszak, P., Ibanez, R., Zippel, K.C., Lawson, D.P., Wright, K.M., Stuart, S.N., Gascon,
C., da Silva, H.R., Burrowes, P.A., Joglar, R.L., La Marca, E., Lotters, S., du Preez, L.H.,
Weldon, C., Hyatt, A., Rodriguez-Mahecha, J.V., Hunt, S., Robertson, H., Lock, B.,
Raxworthy, C.J., Frost, D.R., Lacy, R.C., Alford, R.A., Campbell, J.A., Parra-Olea, G.,
Bolanos, F., Domingo, J.J., Halliday, T., Murphy, J.B., Wake, M.H., Coloma, L.A., Kuzmin,
S.L., Price, M.S., Howell, K.M., Lau, M., Pethiyagoda, R., Boone, M., Lannoo, M.J.,
Blaustein, A.R., Dobson, A., Griffiths, R.A., Crump, M.L., Wake, D.B. and Broody, E.D.
2006. Confronting Amphibian Declines and Extinctions. Science 313(5783):48.
Picker, M.D., Harrison, J.A. and Wallace, D. 1996. Natural hybridisation between Xenopus
laevis laevis and X. gilli in the south-western Cape Province, South Africa. In: R.C.
Tinsley and H.R. Kobel (eds.), The Biology of Xenopus, pp. 61-71. Zoological Society
of London, Oxford, UK.
 CHAPTER 6. AMPHIBIANS OF THE AUSTRALASIAN REALM
Jean-Marc Hero, Stephen Richards, Ross Alford, Allen Allison, Philip Bishop,
Rainer Günther, Djoko Iskandar, Fred Kraus, Frank Lemckert, James Menzies,
Dale Roberts and Michael Tyler
THE GEOGRAPHIC AND HUMAN CONTEXT
The Australasian Realm as defined here includes Australia and New Zealand, and subsumes
the Papuan Sub-region incorporating the Solomon Islands, Papua New Guinea and parts
of Indonesia (Papua Province on the island of New Guinea, and the Malukus, including the
islands of Halmahera, Ceram, Obi, Misool, Aru, Ambon, Buru and Kai). The Oceanic Region,
which includes islands in the northern and eastern Pacific Basin, is usually treated as a
separate biogeographic region (Olsen et al. 2001), but as it has only three native amphibians
occur in the region (two in Fiji and one in Palau) and their evolutionary affinities are with
Australasian taxa so it is combined with Australasia for the purpose of this analysis.
The Australasian continental plate includes mainland Australia, the island of Tasmania
and the southern half of the island of New Guinea. It was part of the southern landmass
of Gondwana until about 96 Ma, though connections with Antarctica and South America
continued until more recently. The Indo-Australian Plate started drifting northwards (with
the Indian and Australasian portions separating), colliding with the Eurasian Plate to the
north approximately 15 Ma, and creating the uplift that formed New Guinea’s spectacular
mountainous spine. Much of New Guinea, particularly the northern portions, is a complex
mosaic of ‘terrains’, fragments of the earth that have embedded onto the mainland at
different times, each with different origins and ages of impact. This incredibly complex
geological history and one of the most active orogenies on earth has resulted in a spec-
tacularly diverse topography and fauna in a relatively small area. The Torres Strait, a
shallow water body that now separates Australia from New Guinea, has formed a bridge
between the two land-masses on several occasions in the past during glacial periods
when global sea levels were lower. The most recent land connection between Australia
and New Guinea lasted until about 7,000 years ago. Tasmania was similarly connected to
the Australian mainland during times of glacial maxima when Bass Strait was exposed.
The islands of Wallacea form the western boundary of the region. These islands enabled
cross colonization of plants between Southeast Asia and Australia-New Guinea but the
deep ocean straits between islands appear to have hindered the exchange of many groups
of amphibians (though some dispersal appears to have taken place due to tectonic ac-
cretion, for example allowing Oreophryne to reach Sulawesi from the east, and Rana to
reach New Guinea from the west).
Around 85 Ma, a land mass known as Tasmantis broke away from Australia. All that now
remains of this largely submerged landmass is New Zealand at its southern end and New
Caledonia at the northern end. Located on the south-eastern boundary of the Australian plate
where it converges with the Pacific plate, New Zealand consists of two main islands, called
the North and South Islands, and a number of smaller islands. Its position on the boundary of
two converging plates is reflected in the many active volcanoes on the North Island.
Australia is one of the flattest countries in the world. Because it lies in the middle of
the Australian tectonic plate there has been very little geological activity since the plate
separated from Gondwana. Instead the landscape has been moulded by constant weather-
ing and erosion over millions of years. The highest peak is Mount Kosciuszko (2,228m) in
the Snowy Mountains of New South Wales; these form part of the Great Dividing Range,
which runs north-south along nearly the entire east coast of the country. This mountain
range promotes significant rainfall along the east coast, but substantially prevents rainfall
from the Pacific Ocean reaching the interior. Hence, the east coast includes a wide range
of wet forest habitats (especially in the more mountainous areas), but the vast interior of
Australia, stretching all the way to the west coast, is predominantly desert, semi-desert
or savannah. The north of the country is tropical, and is dominated by savannah, eucalypt
woodland, grassland, and desert habitats, with rainforests being found along parts of the
east coast (mainly in mountainous regions) and in pockets scattered further west. There are
significant areas of rocky escarpment country in the Kimberley and Arnhem Land. Temper-
ate climates are found only in the south-east and south-west corners of the country. Inland
droughts lasting several seasons are not uncommon as rainfall can be highly variable and
patchily distributed.
The island of New Guinea is divided politically, with the eastern half of the island along
with many islands off the north and east coast (including Admiralty and Bismarck archi-
pelagos, and the islands of Bougainville and Buka in the Solomons Archipelago) forming the
country of Papua New Guinea, and the western half of the island belonging to Indonesia. A
wide, east-west range of mountains dominates the interior of the island, with the highest
peak being snow-covered Puncak Jaya (5,030m) in Papua, Indonesia. To the north and south
of the central mountains are vast lowland rainforests and complex wetland habitats. These
are most conspicuously developed in the Trans-Fly region of south-central New Guinea
where seasonally flooded savannah grasslands form a habitat unique in New Guinea (but
typical of northern Australia).
New Zealand is geographically very isolated, the closest countries being Australia some
2,000km to the west, and New Caledonia, Fiji and Tonga, between 1,400 and 2,000km to the
north. Mountains cover about two-thirds of the South Island and 20% of the North Island,
the highest peak being Aoraki/Mount Cook (3,754m) on the South Island. On either side of
the central mountain ridges are rolling fertile plains, and due to its southerly latitudes the
country has a temperate climate. Rainfall is generally moderate to high in New Zealand.
Human population density is very low throughout the Australasian Realm. The Solomon
Islands have the highest population density (17 people/km2), followed by New Zealand
(15), Papua New Guinea (11), Indonesia (Papua Province only: 6) and Australia (less than
3). Indonesia is the fourth most populous country in the world, but only 1% of Indonesians
live in Papua Province, even though the landmass of Papua constitutes around 22% of the
total landmass of the country. Population growth is high (over 3% per annum) in Papua
Province, partly due to a high birth rate but also due to immigration from other provinces
of Indonesia. The Solomon Islands and Papua New Guinea also have growth rates above
2% (approaching 4% in many coastal areas), but Australia and New Zealand have growth
rates that are less than 1% per annum.
New Guinea has been subject to relatively low anthropogenic disturbance and it has been
stated that around 70% of the natural environment on the island remains intact (Mittermeier
Ceratobatrachus guentheri (Least Concern)
is an abundant direct-developing ranid frog
from the Solomon Islands and Bougainville
and Buka Islands in Papua New Guinea. It
inhabits the forest floor in tropical rainforest,
and persists in secondary forest, rural gardens
and other degraded areas. © Piotr Naskrecki
et al. 2002), although some parts of the island have experienced serious habitat loss (and the
level of loss is likely to be higher than has been realised). The extensive tropical rainforest
wilderness on the island is the world’s third largest rainforest block after the Amazon and
Congolese forests. However, logging concessions now cover much of the lowland forest in
New Guinea and nearly all of New Britain; unsustainable logging is the most immediate
threat to the biodiversity of these regions. In Papua New Guinea the majority of people still
practice subsistence agriculture and live in traditional societies, and the GDP per capita is
low ($2,400 in 2005). The Solomon Islands has a similarly low GDP per capita ($1,700 in
2002), and Indonesia ($3,700 in 2005 averaged across the whole country, not just Papua
Province) is only slightly higher.
Australia has a very low human population density, but with the majority of the people
living along the east and south-east coast, coupled with the high GDP per capita, habitat
loss and degradation has been particularly serious in these regions. There has also been
extensive habitat loss in south-western Australia (which is a Mediterranean-type system).
Likewise, New Zealand has experienced significant habitat loss and degradation with natural
habitats covering only 22% of the country. Invasive species have had a significant impact
on native species both in Australia and New Zealand.
GLOBAL CONSERVATION STATUS
There are 544 amphibians currently recorded from the Australasian Realm, 69 (13%) of Figure 1. Summary of Red List categories
which are considered globally threatened or Extinct. This is much lower than the global for amphibians in the Australasian Realm.
average of 33%1. However, the number of DD species, 178 (33%), is much higher than The percentage of species in each category
the global average of 23%. As sufficient information becomes available to assess these is also given.
species, it is expected that some of them will contribute to an increase in the percentage
of threatened species in the region.
The Australasian Realm accounts for only 3% of globally threatened amphibians, which Red List Category Number of species
is relatively low considering that 9% of the world’s amphibian species occur here. The region Extinct (EX) 3
contains 4% of the CR, 3% of the EN and 4% of the VU species in the world. Extinct in the Wild (EW) 0
Of the world’s 34 known amphibian extinctions, three occurred in the Australasian Realm Critically Endangered (CR) 16
and all of them were species endemic to Australia. The only two species within the family Endangered (EN) 21
Rheobatrachidae are now both Extinct: the Southern Gastric-brooding Frog Rheobatrachus Vulnerable (VU) 29
silus was last seen in the wild in 1981 (Richards et al. 1993), while the Northern Gastric- Near Threatened (NT) 10
brooding Frog Rheobatrachus vitellinus was last recorded in the wild in 1985 (Ingram and Least Concern (LC) 287
McDonald 1993; McDonald and Alford 1999) (see Essay 6.1). The third Extinct species, Data Deficient (DD) 178
Taudactylus diurnus, has not been recorded in the wild since 1979 (Czechura and Ingram Total Number of Species 544
1990). The reason(s) for the decline of all three species has still not been confirmed despite
1% 3%
extensive research (Tyler and Davies 1985), but the disease chytridiomycosis (which was 4%
unknown at the time of these extinctions) must now be suspected. In addition, there are five 5%
2%
Critically Endangered species that are considered Possibly Extinct (Litoria castanea, Litoria 33%
lorica, Litoria nyakalensis, Litoria piperata and Taudactylus acutirostris). This represents 4%
of the global total of 130 Possibly Extinct species.
SPECIES RICHNESS AND ENDEMISM
Species Richness and Endemism Across Taxa
53%
There are 544 amphibian species occurring in the Australasian Realm (9% of the world’s
total). Only one of the three orders of amphibians, the Anura, is found here but of the seven
anuran families native to the region, four are endemic. The level of endemism increases at
the generic level, with 90% (45 of 50) of genera endemic, and again at the species level,
with 99% (538 of 544) of species endemic.
The high level of endemism is largely due to the insular nature of the region. The boundary
between Australasia and Indomalaya is also defined here in a manner that minimizes the
number of amphibians occurring across both regions. Whereas the boundary between the
Australasian and Indomalayan realms is often taken as Wallace’s line (an imaginary line
running between Borneo and Sulawesi and Bali and Lombok in Indonesia; e.g. Newton 2003),
the boundary is here taken to lie further to the east between Sulawesi and the Maluku islands
(approximating Tyler 1999a), as this reflects amphibian distribution patterns better.
Seven of the world’s 48 amphibian families are found in the region, and four of these
are endemic: Leiopelmatidae, Limnodynastidae, Myobatrachidae, and Rheobatrachidae.
The characteristics of these families are provided in Chapter 12. There have been various
taxonomic arrangements of the families defined here as Limnodynastidae, Myobatrachidae
66 Threatened Amphibians of the World

The Australian Lace-lid Nyctimystes dayi (En- and Rheobatrachidae. Limnodynastidae and Rheobatrachidae are sometimes included as
dangered) is a treefrog (Hylidae) that is endem- subfamilies of Myobatrachidae, with the species defined here as members of the Myobatra-
ic to the Wet Tropics of northern Queensland, chidae included in the subfamily Myobatrachinae (Heyer and Liem 1976). Another taxonomic
Australia. In the 1980s and early 1990s it arrangement recognizes two families, separating the Limnodynastidae as a separate family
disappeared from upland sites throughout its from the Myobatrachidae (Zug et al. 2001; Davies 2003a, b; Iskandar and Colijn 2000). The
range, but still persists in suitable habitat at Rheobatrachidae were placed in a separate family by Laurent (1980, 1986), and these three
low elevations. © Jodi Rowley separate families are recognized here.3
The three non-endemic families occurring in the region are Microhylidae, Hylidae, and
Ranidae. Together, these three non-endemic families contain 77% of the region’s species.
The family with the largest number of species in the region is Microhylidae with 197 species
(all endemic), and representing 46% of the family’s global species diversity. Most of these
species are found on New Guinea, and comprise almost half of the total amphibian fauna
on the island. There are 19 genera of the Microhylidae in the region, which represents 28%
of the family’s generic diversity. All of these are endemic to the region except for the genus
Table 1. The number of Australasian am- Oreophryne which occurs as far west as the Philippines. The three most speciose genera
phibians in each taxonomic Family present are Oreophryne (34 species in the region) that within Australasia is restricted to the Papuan
in the region. sub-region, and Cophixalus (35 species) and Austrochaperina (24 species) both of which
occur in Australia and in the Papuan sub-region. All microhylids occurring in the region have
a reproductive strategy in which embryonic and larval development occurs entirely within
Family Native species Percentage of Percentage of Native genera Percentage of Percentage of the egg capsule and they do not rely on free-standing water for breeding.
(endemics species in region species in family (endemics genera in region genera in family The second largest family in the region is the Hylidae4 with 161 species, of which 159
to region) that are endemic that are endemic to region) that are endemic that are endemic
are endemic to the region (representing 20% of the family’s global species diversity). Three
to region to region
hylid genera, representing only 6% of the global diversity at the genus level, are represented
Anura in the region. The largest genus, and the only one not endemic to the region, is Litoria (124
Hylidae 161 (159) 99 20 3 (2) 67 4 species), which ranges across Australia and the Papuan sub-region. Two of the species in
Leiopelmatidae 4 (4) 100 100 1 (1) 100 100 the genus occur in both the Australasian and Indomalayan regions (Litoria infrafrenata and L.
Limnodynastidae 50 (50) 100 100 8 (8) 100 100 rubella) and one is endemic to the Indomalayan region (Litoria everetti). The second largest
Microhylidae 197 (197) 100 46 19 (18) 95 26 genus, Nyctimystes (24 species), also occurs in Australia and the Papuan sub-region while
Myobatrachidae 71 (71) 100 100 11 (11) 100 100 the smallest genus, Cyclorana (13 species), is endemic to Australia. All hylids occurring in
Ranidae 59 (55) 91 8 7 (4) 57 10 the region lay eggs in water where free-swimming larvae develop until metamorphosis.
Rheobatrachidae 2 (2) 100 100 1 (1) 100 100 The remaining non-endemic family occurring in the region is the globally widespread
TOTAL ANURA 544 (538) 99 10 50 (45) 90 12 Ranidae, represented in the region by 59 species (55 endemic) comprising 9% of the global
TOTAL ALL AMPHIBIANS 544 (538) 99 9 50 (45) 90 10 species diversity of the family. Within Australasia ranids are primarily found in the Papuan
sub-region with only one species, Rana daemeli, extending its range into Australia. The three
native amphibians of the Oceanic Region are members of the region’s most speciose ranid
genus, Platymantis (29 species). In total there are seven native ranid genera in the region,
which is 10% of the global diversity of the family at the genus level. Four of these genera
Family EX CR EN VU NT LC DD Total Number threatened % Threatened are endemic: Batrachylodes (8 species), Ceratobatrachus (1 species), Discodeles (5 species)
of Order or Extinct or Extinct and Palmatorappia (1 species). The species in these genera have a ‘direct development’
Anura reproductive strategy like the microhylids, but species within the genera Limnonectes and
Hylidae 0 6 6 12 2 93 42 161 24 15 Rana have aquatic larvae.5
Leiopelmatidae 0 1 1 2 0 0 0 4 4 100 There are 69 threatened or extinct species in the Australasian Realm spread across all
Limnodynastidae 0 1 7 2 1 37 2 50 10 20 seven families (Table 2). The family with the highest number of threatened species (24) is
Microhylidae 0 2 4 6 3 71 111 197 12 6 the Hylidae. The most diverse family in the region, the Microhylidae, has relatively few
Myobatrachidae 1 6 2 4 3 49 6 71 13 18 threatened species (only 12 of 197 species being threatened). Comparing the percentage
Ranidae 0 0 1 3 1 37 17 59 4 7 of species that are threatened or extinct reveals large variation between families. Two
Rheobatrachidae 2 0 0 0 0 0 0 2 2 100 families have 100% of their species threatened or Extinct. Both species within the family
TOTAL ANURA 3 16 21 29 10 287 178 544 69 13 Rheobatrachidae are Extinct, and all four species within the New Zealand endemic family
Leiopelmatidae are threatened (and see Essay 6.2).
The least threatened families are the Microhylidae and Ranidae, with only 6% and 7%
Table 2. The number of species within each of species in threatened categories, respectively. Both of these families have in common
IUCN Red List Category in each Family and that they are primarily found in the Papuan sub-region, and all microhylids and the majority
Order in the Australasian Realm. Introduced of ranids in the region are direct-developing species. Compared with the other families,
species are not included. they also have a much higher percentage of species in the Data Deficient category: 56%
for Microhylidae and 29% for Ranidae. This is partly a reflection of how poorly surveyed the
Papuan sub-region is for amphibians. As more research is done in the Papuan sub-region it
is to be expected that the number of threatened species in these two families will increase,
at least to some extent, although direct-developing species in this region do not appear to
be at risk from chytridiomycosis. Microhylids in northern Australia have not declined due
to this disease, even in areas where other species have disappeared.

Geographic Patterns of Species Richness

and Endemism
A map of overall amphibian species richness within the Australasian Realm is shown in
Figure 2. The highest species richness of amphibians is found in the tropics, where the
number of species is highest in the central mountains of New Guinea and the Wet Tropics
rainforests of north-eastern Queensland in Australia. In Australia amphibian species rich-
ness is concentrated in the areas of highest rainfall including the east (in particular in the
temperate forest systems in the New South Wales / Queensland border area) and north
coastal regions, and the south-western tip of Western Australia. Areas of lowest species
Figure 2. The species richness of amphibians richness coincide with Australia’s arid interior and central southern coast. In the Papuan
in the Australasian Realm, with darker colours sub-region, Papua New Guinea has a higher documented species richness than Papua
corresponding to regions of higher richness. Province (Indonesia), but this is most likely an artefact of sampling because there has been
Colour scale based on 10 quantile classes; substantially greater survey effort in Papua New Guinea.
maximum richness equals 51 species. The pattern of species richness illustrated in Figure 2 will require modification as survey
effort is increased in poorly documented areas. Australia and New Zealand are both well
surveyed for amphibians and the rate of new species descriptions is low. In comparison, the
Papuan sub-region is one of the least studied regions in the world, and it is estimated that
more than half of the amphibian species in New Guinea remain to be described (Günther
2006a). Many species in the Papuan sub-region are still known only from their type localities
and perhaps one other locality, whereas the known range of species in Australia and New
Zealand is much more likely to be representative of their actual ranges. There has been
relatively little herpetological research conducted in the Solomon Islands and on the offshore
islands of Papua New Guinea, and it is expected that documented amphibian diversity in
these regions will increase in the future. However, there has recently been considerable
survey work on the islands offshore of south-eastern Papua New Guinea and the known
amphibian diversity there has increased markedly. Although most of these new additions
have yet to be described, a number of new range records have been published (Kraus and
Austrochaperina parkeri (Data Deficient) is only Allison 2004; Kraus and Shea 2005).
known from two localities near Lae in Papua The vast majority of threatened species in the Australasian Realm are endemic to
New Guinea. It is in the Family Microhylidae, Australia (see Figure 3) (and see Essay 6.3). The highest concentrations are found along
which comprises over half the frog species of the east and south-east coast, largely congruent with the areas of highest species species
the island of New Guinea, all of which breed richness. However, the north coast, a relatively species-rich region, has no threatened
by direct development, and many of which are species. Two factors appear to be contributing to this pattern: firstly, the fertile plains of
very poorly known. © Fred Kraus the east and south-east coast of Australia are where the vast majority of people live, and
 Chapter 6. Amphibians of the Australasian Realm 67

Figure 3. a) The richness of threatened

amphibians in the Australasian Realm, with
darker colours corresponding to regions of
higher richness. Colour scale based on five
quantile classes; maximum richness equals
11 species. b) The richness of CR amphibians
in the Australasian Realm. Maximum richness
equals five species.

a. b.

hence where ongoing habitat loss is most severe; and, secondly, the current distribution of
the amphibian chytrid fungus excludes the north coast west of north-eastern Queensland.
The chytrid fungus was first detected near Brisbane on the east coast of Australia in 1978
(Speare and Berger 2005). Since then it has been found in the north and south, and it now
appears to be established in coastal regions from Far North Queensland on the east coast,
southwards through New South Wales and Victoria, and along the southern coast to Adelaide
in South Australia. On the west coast it has been found in Perth and across most of the high
rainfall zone in south-western Western Australia (Speare and Berger 2005). The spread of
recorded occurrences of the disease is continuing with amphibians infected with chytrid
being recorded in Tasmania for the first time in late 2004 (Obendorf 2005). It is possible
that, if chytrid spreads into areas where it is not already present, more species will become
threatened. However, chytrid fungus has relatively limited climatic tolerances with respect
to both humidity and temperature (Woodhams et al. 2003). It might be less likely to have
major impacts in the wet dry-tropical eucalypt woodland/grassland communities across
much of northern Australia where dry seasons are too extreme and summer temperature
too high for persistence, or in south-western Australia for similar reasons: extreme summer
drought and associated high temperatures.
In New Zealand the map of amphibian species richness and threatened species richness
are identical because all four native amphibians are threatened. In the Oceanic region, only
one species is threatened, the Fijian Ground Frog Platymantis vitiana (EN), which is now
found only on the small islands of Ovalau, Gau, Taveuni, and Viwa in Fiji (its tiny range is
difficult to see in Figure 3a).
The Papuan sub-region (see Essay 6.4) has very few threatened species, particularly when
considering its high species richness. The documented distribution of threatened species
is incomplete due to severe lack of knowledge, but may also partly reflect a lack of certain
threatening processes. For example, the chytrid fungus, which has had such a devastating
impact on amphibians in Australia, has not been found in the region (though attempts to
document this are limited to non-existent). There has also been relatively less habitat loss
and degradation compared with Australia. However, widespread environmentally destructive
logging throughout the Papuan sub-region (which is ongoing especially in parts of Papua
New Guinea) may threaten numerous species in the future, particularly on smaller island
archipelagos such as the Solomon Islands and the Admiralty and Bismarck Archipelagos.
The distribution of Critically Endangered species is highly congruent with that of
threatened species overall (Figure 3b). New Zealand and the east coast of Australia remain
focal points, with particular concentrations of Critically Endangered species in the Wet
Tropics region of far northern Queensland, and in the central mountains from Fraser Island
to Sydney. Tasmania is no longer highlighted, possibly because the chytrid fungus has only
very recently been confirmed on the island and its impact on local amphibian populations
is not yet known. In the Papuan sub-region one Critically Endangered species, Albericus
siegfriedi, is known6. It has been recorded from only one location, Mount Elimbari, in Papua
New Guinea (barely visible in Figure 3b).
Species Richness and Endemism within Countries
Amphibian species richness in the Australasian Realm is greatest on the two largest land
masses of Australia and New Guinea (see Figure 2). Only three native amphibians occur in
the Oceanic region (0.05% of the global total), compared with 187 birds (2% of the global
total) (BirdLife International 2004). The only two countries in the Oceanic region that have
native amphibians are Fiji (Platymantis vitiana and P. vitiensis) and Palau (Platymantis
pelewensis). New Caledonia has one introduced species (Litoria aurea) as do the Northern
Mariana Islands, American Samoa, the Bonin Islands, the Federated States of Micronesia,
Guam and Tuvalu (Bufo marinus in each case).
Papua New Guinea has the most native amphibians in the Australasian Realm (245
species) and this number is predicted to almost double as species descriptions continue.
Australia is the second-most diverse country (214 species), although as species descrip-
tions continue it is likely that Indonesian New Guinea (Papua) will eventually overtake
Australia’s total in the region.7 The number of species in the Solomon Islands is relatively
low (19 species) and there will be a moderate increase in the known fauna with future
herpetofaunal surveys. Documentation of the New Zealand native frog fauna is almost
certainly complete, although recent genetic work on isolated populations of Leiopelma
hochstetteri (Gemmel et al. 2003) suggests that there might be more than six distinct cryptic
species. New Zealand also has three species introduced from Australia, two of which are
threatened in their native ranges.
The only overview of the amphibians of the whole region is that of Tyler (1999a). There
is an extensive overview literature, and guide books, on the amphibians of Australia (e.g.,
Barker et al. 1995; Glasby et al. 1993; Tyler 1999b; Cogger 2000), and recent assessments of
amphibian declines throughout Australia include Hero and Shoo (2003), Hero and Morrison
(2004), Hero et al. (2005) and Hero et al. (2006). Slatyer et al. (2007) made an assessment
of endemism and species-richness patterns of Australian frogs. The most recent overview
of the small frog fauna of New Zealand is by Gill and Whitaker (1996). Menzies (2006)
provides the first review of the amphibians of the Papuan sub-region. Figure 4. The number of amphibians present
The percentage of species that are endemic to each country in the region varies from in and endemic to each Australasian country.
100% to 5% (Figure 5). Three countries in the region – New Zealand, Fiji and Palau – have *denotes countries not entirely within the Aus-
endemism of 100%, although all three of these countries have only very few species (four, tralasian Realm, hence only the species whose
two and one, respectively). Of the three countries with high species richness, Australia ranges fall within the region are included.
has the highest level of endemism at 93%, followed by Papua New Guinea with 67% and
Indonesia with 59% (the lower levels of endemism in the latter two countries are due to the
biogeographically arbitrary border between them, and the percentage of endemics on the Figure 5. Percentage of species endemic to
island of New Guinea is over 95%). The Solomon Islands has the lowest level of endemism each Australasian country. *denotes countries
with only one of its nineteen native species, Discodeles malukuna (DD), being endemic. not entirely within the Australasian Realm,
However, in a biogeographical context these levels of endemism are meaningless. Like the hence only the species whose ranges fall
island of New Guinea, the level of endemism in the Solomon Islands Archipelago would within the region are included.
be substantially greater if this natural and discrete biogeographic ‘unit’ had not been split
by political boundaries.
All of the countries in the Australasian Realm that have native amphibians also have Figure 6. The number of threatened am-
threatened species with the exception of Palau (see Figure 6). Australia contains almost phibians present in and endemic to each
three-quarters (71%) of the region’s 66 threatened species and all but two of these species Australasian country. Countries with no
are endemic to the country. Predicting the impact generated by the possible spread of the threatened species are not included in the
pathogenic chytrid fungus throughout Australia is difficult as its spread into semi-arid and arid diagram. *denotes countries not entirely
areas across much of southern and northern Australia might have less impact than expected, within the Australasian Realm, hence only the
as described above. In contrast, the most diverse country, Papua New Guinea, has only 10 species whose ranges fall within the region
threatened species, of which eight are endemic. Indonesia and New Zealand, with 148 and are included.
four native species, respectively, amazingly have the same number of threatened species.
As more survey work is completed in the Papuan sub-region, the number of threatened
species in Indonesia, Papua New Guinea and the Solomon Islands probably will increase. Figure 7. Percentage of native species that are
This increase is likely to come from documenting newly described species with severely threatened. Countries with no threatened spe-
constrained ranges and threatened habitats, as well as from improved data that will permit cies are not included in the diagram. *denotes
a more realistic assessment of species currently in the Data Deficient category. countries not entirely within the Australasian
Realm, hence only the species whose ranges
fall within the region are included.

Papua New Guinea New Zealand Australia New Zealand

Australia Fiji
Papua New Guinea Fiji
Indonesia* Palau
New Zealand Australia
Solomon Islands Australia
Indonesia* Solomon Islands
New Zealand Papua New Guinea
Solomon Islands Papua New Guinea
Fiji Indonesia*

Palau Solomon Islands Fiji Indonesia*

0 50 100 150 200 250 0 20 40 60 80 100 0 10 20 30 40 50 0 20 40 60 80 100

Number of species Percentage of endemic species Number of threatened species Percentage of species threatened
Endemics Non-endemics Endemics Non-endemics
Figure 4 Figure 5 Figure 6 Figure 7
68 Threatened Amphibians of the World

Left: Litoria prora (Least Concern) is a hylid

treefrog from New Guinea. It is an arboreal
species and is found near small streams and
forest pools in lowland and foothill rainforests.
It breeds in pools and slow-flowing portions of
streams. © Stephen Richards

Right: The Sunset Frog Spicospina flammocae-

rulea (Vulnerable) in the Family Myobatrachi-
dae is known only from a small area of south-
western Australia, where it was discovered
in 1994. It occurs only in a few isolated peat
swamps. © Ross Knowles

There is extreme variation in the percentage of species that are threatened in each country
in the region (see Figure 7). With 100% of its native species threatened, New Zealand is
top of the list, followed by Fiji with 50%, but these countries only have four and two native
species, respectively. When considering countries with high species richness, Australia has
by far the greatest percentage of threatened species, 22% of extant species, compared with
only 4% in Papua New Guinea and 3% in Indonesia. These figures reflect the impact that chy-
tridiomycosis, habitat loss and other threatening processes have had in Australia compared
with the relatively low impact of threats in New Guinea, and perhaps also reflect the lack
of data from Indonesia and Papua New Guinea. Australia, however, still has a significantly
lower percentage of its species threatened than the global average of 33%.
Assessments of the conservation status of Australasian amphibians at national level
have been carried out in Australia (Tyler 1997; Campbell 1999; Hero and Morrison 2004;
Hero et al. 2006; Clayton et al. 2006; and see Essay 6.3), and in New Zealand (Newman
1996; Hitchmough 2002).
There are only 16 Critically Endangered species in the Australasian Realm, but 14 of
these occur in Australia (where the impacts of chytridiomycosis and habitat loss have been
most severe) and one each in New Zealand and Papua New Guinea.
HABITAT AND ECOLOGY
Habitat Preferences
Most amphibians in the region are found in forests (73%), and in particular in tropical forest
(68%) (Table 3; Figure 8). There are also large numbers of species in grassland (19%) and
shrubland (13%). However, the percentage of species in each of these habitats that are
threatened is notably different: 14% of forest-dwelling species are threatened, but only
6% of grassland species and 10% of shrubland species are threatened. Similar numbers of
amphibians are associated with flowing freshwater and still open freshwater; however, more
than twice the total number of species associated with flowing freshwater is threatened
compared with species in still open freshwater. This reflects a global trend for amphibians
associated with forests and flowing freshwater to be more likely to decline rapidly (Stuart
et al. 2004). Hero and Morrison (2004), Hero et al. (2005) and Hero et al. (2006) previously
noted that montane wetland and streamside species are the most likely ones to experience
enigmatic declines in Australia.
Significantly more lowland tropical forest species below 1,000m (52%) than montane
tropical forest species above 1,000m (33%) occur in the region, and a higher percentage of
lowland species is threatened (16%) compared with montane species (11%). This is most
likely because the mountain ranges in Australia, where most threatened species in the region
are found, are relatively low in elevation, and because the higher elevation mountain ranges
of New Guinea are very poorly surveyed (so that a large proportion of montane species in
this region is considered Data Deficient).
No amphibians occurring in savannah are considered threatened and this is largely a
reflection of the wide ranges of these species. Interestingly, 25% of species in the region
are found in secondary terrestrial habitats, and a significant proportion (7%) of these spe-
cies is threatened.
Breeding Strategies
The most common reproductive mode amongst Australasian amphibians involves a free-
swimming larval stage or ‘larval development’ (53%), followed closely by ‘direct development’
(47%) in which embryonic and larval development is completed within the egg capsule
(Table 4). There are no known live-bearing species in the region, but the reproductive
Reproductive mode All Species Threatened or % Threatened
Extinct species or Extinct
Direct development 255 24 9
Larval development 288 45 16
Live-bearing 0 0 -
Not known 1 0 0
Table 4. Australasian amphibians categorized by reproductive mode.
strategy of many species is inferred. When compared with global trends in reproductive
mode, the Australasian Realm has a significantly higher proportion of direct developers
(30% of species globally are direct developers), and a significantly lower proportion of
larval developers (68% of species globally are larval developers). This is largely caused
by nearly 200 species of the family Microhylidae occurring mainly in New Guinea, all of
which are direct developers.
The percentage of larval-developing species that are threatened is almost double that of
direct developers (Table 4). This contradicts the global trend, where 45% of direct developers
and 28% of larval developers are threatened. This apparent contradiction can be explained
by considering the geographical distribution of species exhibiting different breeding strat-
egies. In Australia, where 47 of the region’s 66 threatened species are found, 188 of the
217 native species are larval developers. In Papua New Guinea, where only 10 species are
threatened, 160 of the 244 native species are direct developers. The situation is similar in
Indonesia, where only four species are threatened, and 83 of the 148 native species are
direct developers. Australia is the only country in the region in which the proportion of larval
developers is higher than that of direct developers. Compared with the other two megadiverse
countries in the region, Papua New Guinea and Indonesia, Australia is also the country most
impacted by threats such as chytridiomycosis, introduced species, and habitat loss (Hero
and Morrison 2004; Hero et al. 2005, 2006). Chytridiomycosis is more often implicated in
declines of stream-associated, usually larval-developing, species (Lips et al. 2003), although
some direct developers that are associated with streams have also been impacted by the
disease. In New Zealand, contrary to the general pattern, the fully terrestrial Leioipelma
archeyi (CR) has experienced a decline due to chytridiomycosiss (Bell et al. 2004), whereas
no declines have been detected in the sympatric stream-dwelling frog (L. hochstetteri, VU).
In Fiji and Palau, all of the native species are fully terrestrial direct developers.
MAJOR THREATS
The most significant threat to species in the region is habitat loss, impacting three-quarters
of threatened species (Table 5; Figure 9). This trend is global and habitat loss has been
identified as a major threat for birds and mammals as well as amphibians (Baillie et al.
2004). Of particular interest is the second-most significant threat, invasive species8 (primarily
salmonids and Gambusia; Gillespie and Hero 1999), which are affecting two-thirds of all
threatened species in the region, compared with only 11% of threatened species globally.
This highlights the particularly severe impact that invasive species are having on native
wildlife in Australia, New Zealand, and oceanic islands such as Fiji.
Pollution is a significant threat to almost half of the threatened species in the region,
which is a higher proportion than the global figure of 29%. Disease is also having a major
impact, and is believed to be affecting 24 threatened species in the region. The chytrid
fungus has now been identified in wild populations of 48 native Australian frogs, and also

Habitat type Number of % of all Threatened % of species

Number of species % of species in habitat that are threatened
species in each species and Extinct occurring
habitat occurring in species in habitat that Forest
the habitat are Threatened Savannah
or Extinct
Grassland
Forest 398 73 55 14
Shrubland
All tropical forest 372 68 47 13
Lowland tropical forest 281 52 44 16 Secondary terrestrial habitats

Table 3. The habitat preferences of amphib- Montane tropical forest 177 33 19 11 Flowing freshwater
ians in the Australasian Realm. Savannah 41 7 0 0 Marsh/swamp
Grassland 106 19 6 6
Still open freshwater
Shrubland 70 13 7 10
Figure 8. The habitat preferences of amphib- Secondary terrestrial habitats 136 25 10 7 Arid and semi-arid habitats
ians in the Australasian Realm. The plot on the Flowing freshwater 177 32 35 20
0 100 200 300 400 0 5 10 15 20
left-hand side shows the number of species in Marsh/swamp 95 17 14 15
the region in each habitat type. On the right- Still open freshwater 171 31 15 9
hand side, the percentage of these species Arid and semi-arid habitats 5 1 0 0
which are threatened is given.
 Chapter 6. Amphibians of the Australasian Realm 69

Table 6. The purposes for which amphibians

Threat type Threatened species % Threatened species Purpose Subsistence Sub-national/ Regional/ Number of are used in the Australasian Realm. The
Habitat loss 50 76 National International species numbers in brackets are the number of species
Agriculture – Crops 4 6 Food - human 15 (0) 1 (0) 0 15 (0) within the total that are threatened species.
Agriculture – Tree plantations 5 8 Food - animal 1 (0) 0 0 1 (0) The two species listed as being used in “Re-
Agriculture – Livestock 24 36 Handicrafts, curios, etc. 0 1 (0) 0 1 (0) search” (Rheobatrachus silus and R. vitellinus)
Timber and other vegetation removal 17 26 Pets, display animals 3 (2) 6 (3) 11 (2) 12 (3) are now Extinct.
Urbanization and industrial development 29 44 Research 0 2 (2*) 0 2 (2*)
Invasive species 44 67
Utilization 1 2
Accidental mortality 2 3
Pollution 31 47 Table 7. The population trends for all extant
Natural disasters 12 18 Population Trend Number of species % of extant species Australasian amphibians.
Disease 24 36 Decreasing 53 10
Human disturbance 16 24 Stable 286 53
Fire 16 24 Increasing 2 0.4
Unknown 200 37

Table 5. The major threats to globally threatened amphibians in the Australasian Realm.
Only present threats to species are tallied.
Table 8. The number of species in “rapid
Family Number of Percentage Number of Percentage decline” and “enigmatic decline” in the
species in of species in species in of species Australasia Realm by Family.
“rapid decline” family in “enigmatic in family in
“rapid decline” decline” “enigmatic
All habitat loss decline”
Invasive species Hylidae 13 8 10 6
Utilization Leiopelmatidae 1 25 1 25
Limnodynastidae 7 14 3 6
Accidental mortality
Microhylidae 2 1 0 0
Persecution Myobatrachidae 9 13 7 10
Pollution Ranidae 1 2 0 0
Rheobatrachidae 2 100 2 100
Natural disasters

Disease

Human disturbance

Changes in native sp. dynamics
Fire
0 20 40 60
Percentage of threatened species affected
Figure 9. The major threats impacting threatened amphibians in the Australasian Realm.
in the introduced cane toad, Bufo marinus (Speare and Berger 2005). Captive populations of
two additional species have been infected, although there is no evidence of infection within
their wild populations (Speare and Berger 2005). Not all species infected have declined, and
there are some species, for example Taudactylus eungellensis (CR), which have experienced
declines in the past that might have been due to chytridiomycosis and are now showing signs
of recovery. Why some species appear to be more resilient to the disease than others is still
not known, although there is evidence that both innate immune defenses and environmental
factors play a role (Woodhams and Alford 2005; Woodhams et al. 2006).
A total of 30 species (three of which are threatened and two considered Extinct) are
recorded as being utilized by humans. The most common reason for harvesting Australasian
amphibians is for local human consumption, and this occurs only in the Papuan sub-region.
Some amphibians are harvested for the international pet trade; the most common species
being treefrogs of the genus Litoria, which originate from both Australia and the Papuan
sub-region. Much of the harvesting of amphibians in the region is not considered to constitute
a major threat to the species; of the 30 species being harvested, this activity is considered
to be a threat to just six species.
not well understood, while the other 12 have experienced declines due to reduced habitat.
Australia has the highest number of “rapidly declining” species in the region, with 31 species,
followed by Papua New Guinea with three species, and New Zealand with one species. All
80 100 three species in Papua New Guinea are declining as a result of reduced habitat, and the
reason for the decline of the one New Zealand endemic species has been identified as most
likely a result of chytridiomycosis (Bell et al. 2004). Most of the “enigmatic declines” in the
region are concentrated in eastern Australia, where chytrid is widespread.
Of the 544 species occurring in the region, 6% are considered “rapidly declining” which
is slightly less than the global average of 8% of all amphibians. Considering that the
region has a significantly lower percentage of threatened species than the global average
(12% compared with 33%), the number of “rapidly declining” species is relatively high.
This highlights the severity of the threats impacting certain species in the region, and in
particular how quickly these threats can push species to the brink of extinction. Of the 66
species that are currently considered threatened in the region, 29 are also “rapidly declining”.
Consequently, although the region does not have a large proportion of threatened species,
many species that are threatened, are also in serious decline.
The “rapidly declining” species show a distinct taxonomic pattern (Table 8). Among the
larger families, the Limnodynastidae and the Myobatrachidae show the highest percent-
ages of species in serious decline, and in the Myobatrachidae almost 10% of species are The Silver-eyed Barred Frog Mixophyes balbus
in “enigmatic decline”. The largest number of species in serious decline is found in the (Vulnerable) in the Family Limnodynastidae
Hylidae, but percentage wise this family appears to be less seriously affected than the Lim- occurs mainly in eastern New South Wales,
nodynastidae and the Myobatrachidae. Some less speciose families have high percentages Australia. The species has declined and dis-
of species in serious decline and “enigmatic decline”, most notably the Rheobatrachidae appeared from a number of locations where
and the Leiopelmatidae, the former family now being extinct (according to the traditional it was once common. It is typically found in
family-level amphibian classification). association with permanent streams in tem-
perate and sub-tropical rainforest and in wet
sclerophyll forest. © Frank Lemckert

POPULATION STATUS AND TRENDS

Estimates of Population Trends
A summary of the inferred population trends of Australasian amphibians is presented in Table
7. For many species, particularly those in the Papuan sub-region, these trends are inferred
from broad trends in the state of the habitats on which the species depend. In Australia
and New Zealand, where most of the declining species are found, many of the population
trends are based on field observations (for example, Campbell (1999) for Australia, and
Bishop (2006) for New Zealand). Compared with global trends, the population trends in the
Australasian Realm is quite different, with many more species considered to have stable,
rather than decreasing, populations. More than half of the amphibians in the region are
considered to have stable populations, compared with only 27% globally, and only 10% of
species in the region are considered to be in decline (vs. 42% globally). This would suggest
that although there have been dramatic declines, and in some cases extinctions, amongst
some populations of species in Australia and New Zealand, amphibian populations overall
are relatively stable. In Australia, this stability might be because the major effects of feral
introductions, large-scale habitat clearance and chytridiomycosis have generally now played
out. The main feral predators (cats and foxes) and habitat-changers (rabbits and domestic
livestock) reached all parts of Australia some time ago. The large-scale clearance of important
natural habitats was generally completed by 1950, and legislation to minimize such habitat
clearance is now generally in place. The chytrid fungus had reached much of Australia by
the late 1990s. As a result, there is some relative stability compared with the situation 20,
50 and 100 years ago. For a substantial proportion of species (37%), there is still insufficient
information to determine a trend in population size.

“Rapidly Declining” Species

There are 35 “rapidly declining” species in the Australasian Realm, which is 7% of the global
total of 470 species (a full list of all “rapidly declining” species is provided in Appendix IV
and includes their occurrence within each of the realms). Twenty-three of these species are
experiencing, or have experienced, “enigmatic declines”, in which the causal factors are
70
The Fiji Treefrog Platymantis vitiensis (Near
Threatened) is widespread in Fiji on the islands
of Viti Levu, Vanua Levu, Ovalau and Taveuni. It
is naturally a forest species, living especially
along streams in giant bird’s nest ferns, but
occurs at much lower densities in secondary
habitats. It is in the Family Ranidae and breeds
by direct development, the eggs being laid in
leaf axils. © Paddy Ryan
Endnotes
1 P<0.01 (binomial test)
2 Frost et al.’s (2006) taxonomic rearrangement
results in eight families in the Australasian Realm,
of which only two are endemic: Limnodynastidae
and Myobatrachidae. Frost et al. include the
Nearctic tailed frogs (Ascaphidae) within the New
Zealand frogs (Leiopelmatidae), and so under this
arrangement, the latter family is not endemic to
the region. However, in this section we follow the
former taxonomic arrangement of families based
on Frost (2004).
3 Frost et al. (2006) present another arrangement,
separating the genus Mixophyes (6 species),
from the Limnodynastidae and placing it within
the family Myobatrachidae, and also including
the Rheobatrachidae within the family Myobatra-
chidae.
4 Savage (1973) recognized the Australo-Papuan
hylids as a separate family, the Pelodyradidae.
This has more recently been treated as a sub-
family, Pelodryadinae (see discussion in Frost et
al. 2006).
5 Frost et al. (2006) separate the ranids found in this
region into three families. Those species within
the genus Rana remain within the family Ranidae,
while the genus Limnonectes becomes part of a
new family Dicroglossidae, and the remaining gen-
era (Batrachylodes, Ceratobatrachus, Discodeles,
Palmatorappia and Platymantis) become part of
the new family Ceratobatrachidae.
6 One species recently described for Biak Island in
Indonesia (Litoria biakensis) appears to be seri-
ously threatened (Günther 2006b) and might be
Critically Endangered. However, the species was
described after the completion of data collection
for this book, and is not included in this analysis.
7 The totals listed here for Indonesia only include
species occurring on the islands within the Aus-
tralasian Realm. For the overall country totals for
Indonesia see Appendix V)
8 In this analysis, we have not considered the
chytrid fungus Batrachytrium dendrobatidis to be
an invasive, pending further information on its
geographic origin.
Threatened Amphibians of the World
KEY FINDINGS
• A total of 544 species occur in the Australasian Realm, of which 66 are considered
threatened and three are Extinct (13%). Five species are also considered Critically
Endangered (Possibly Extinct).
• Of the 544 species in the region, 99% are endemic; of 50 amphibian genera, 45 are
endemic; and of the 7 families found in the region, four are endemic. Only one of the
three Orders of amphibians, the anurans, occurs in the region.
• There are five families that have percentages of threatened species higher than the
regional average: Leiopelmatidae (100%), Rheobatrachidae (100%), Limnodynastidae
(20%), Myobatrachidae (18%), and Hylidae (14%). Two families have lower than the
regional average of threatened species: Ranidae (7%) and Microhylidae (6%).
• The vast majority of threatened species in the Australasian Realm are endemic to
Australia, and are concentrated along the east and south-east coast.
• Papua New Guinea has the highest number of native species in the region, followed by
Australia and Indonesia. Of these three countries, Australia has the highest percentage
of endemic species with 93%.
• Australia has the largest number of threatened species (47), and New Zealand has the
highest percentage of threatened species (100%) in the region.
• Most amphibians in the Australasian Realm are found in forests (73%), and in particular
tropical forest (68%). There are more species occurring in lowland tropical forest below
1,000m asl (52%) than in montane tropical forest (33%), and a higher percentage of
lowland species are threatened (16%) compared with montane species (11%).
• Habitat loss is the most common threat, affecting 76% of species, followed by invasive
species (67%), pollution (47%), and disease (36%).
• There are 35 “rapidly declining” species occurring in the region, 31 of these species
are endemic to Australia. Most of these species (66%) have experienced “enigmatic”
declines, probably due to chytridiomycosis, while the remainder are suffering from
reduced habitat.
• Chytridiomycosis has been linked to the enigmatic declines in the Wet tropics region
of north Queensland, and is probably implicated elsewhere in eastern Australia. It has
also been linked to declines in New Zealand.
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ESSAY 6.1. GASTRIC-BROODING FROGS
The discovery in 1973 of a new species of frog, living in rainforest, near
Brisbane in eastern Australia, did not at first attract much attention. The
person who discovered it, David Liem, erected a new genus, Rheobatrachus,
for it and named the species Rheobatrachus silus (Liem 1973). Interest in the
species initially hinged upon its position in the amphibian tree of life, but in
the following year a female was observed to give birth to baby frogs through
the mouth (Corben et al. 1974). This remarkable breeding biology led to it
being anointed the common name of the “Gastric-brooding Frog”.
Gastric brooding represents a unique form of parental care among
amphibians (Corben et al. 1974). Following fertilization, the female ingests
the eggs. The jelly surrounding the eggs contains a compound that switches
off the cells in the lining of the stomach wall that secrete hydrochloric acid
to aid digestion. The compound also paralyses the muscles in the stomach
wall and increases the number of small blood vessels. In effect, collectively
the changes convert the female’s stomach into a womb. The eggs change
into cream-coloured tadpoles and finally metamorphose into young frogs
after a period of six to seven weeks. The young are believed to live entirely
upon their yolk for this period. Similarly, the mother does not actively feed
during this time, living instead upon her body fat and glycogen from the
liver. Young eventually emerge as fully formed froglets, and after four
days the female’s digestive tract returns to normal and she recommences
Birth of a gastric brooding frog Rheobatrachus silus. © Michael J. Tyler
ESSAY 6.2. LEIOPELMATID FROGS: THE WORLD’S MOST ARCHAIC FROGS
The Leiopelmatidae represent a unique evolutionary lineage among am-
phibians, and are thought to be the most archaic frogs in the world. These
frogs are found only in New Zealand, and all members belong to the genus
Leiopelma. Three species are now extinct (L. auroraensis, L markhami and
L.waitomoensis), and only four species are extant: L. archeyi, L. hamiltoni, L.
hochstetetteri and L. pakeka.1
Members of the Leiopelmatidae exhibit a number of primitive traits that
separate them from most other species, including: vestigial tail-wagging
muscles, cartilaginous inscriptional ribs, the presence of amphicoelous
vertebrae, and nine presachral vertebrae (most frogs have eight). Ascaphus
truei, the tailed frog of the North-western United States, is the only other
extant frog known to possess these features (Stephenson 1961). In addition,
there is some evidence to suggest that Leiopelma evolved from an ancestor
with ‘normal’ tadpole development and the degree of cranial remodeling
that occurs in Leiopelma larvae at metamorphosis is intermediate between
salamanders and Ascaphus (Bell and Wassersug 2003).
In addition to these primitive traits, Leiopelma differ from most other frogs
because they lack external eardrums and produce only limited vocalizations.
There are no mating calls and the only noises that any of the four species
are known to make are chirps or yelps when harassed. It is currently thought
that these frogs communicate more like salamanders than other frogs, using
chemosignals to recoginize size and individuality of conspecifics (Lee and
Waldman 2002; Waldman and Bishop 2004).
Although members of the genus Leiopelma have many similarities, each
species varies in its physical characteristics, habitat preferences, distribution
range, and threats faced. Hochstetter’s Frog Leiopelma hochstetteri (VU) is the
more aquatic frog (albeit semi-aquatic) of the four extant species. It is widely
distributed in at least 10 fragmented and isolated populations in the northern
half of the North Island. They are nocturnal and shelter by day in wet crevices
Chapter 6. Amphibians of the Australasian Realm
feeding (Tyler and Davies 1983). Since the stomachs of all frog species are
extremely similar, much of the initial research focusing on stomach acid
secretion in amphibians was done using an imported pest species, the
Cane Toad Bufo marinus.
In 1979, the Gastric-brooding Frog disappeared and, following numerous
fruitless searches, is now regarded as Extinct. Because of the unique nature
of its breeding habits, the species had attracted worldwide interest, and
so attention was focussed upon the many potential causes for its sudden
disappearance. The first was habitat degradation due to timber harvesting,
as although much of the habitat was pristine rainforest, other parts were
greatly disturbed. Stream siltation could have played a significant role,
because it covered the large stones beneath which the frogs hid during the
day. Excessive collecting was also suggested as a reason for the decline of
the species, but this has since been disproved; a second species (the Mount
Glorious Day Frog, Taudactylus diurnus) disappeared simultaneously from
the same site, and is now also considered Extinct.
In 1984, a second species of Rheobatrachus was found approximately
800km north of the range of Rheobatrachus silus, in vine forest near Mackay
on the Queensland coast. It differed from its sibling species in its larger
size and brilliant yellow markings on the ventral surfaces. The species
was named the Northern or Eungella Gastric-brooding Frog R. vitellinus
(Mahony et al. 1984) and evidence of gastric brooding and oral birth was
reported later that year (McDonald and Tyler 1984). Compared with Rheo-
batrachus silus, which gives birth to 18-25 young measuring 13mm, the
female Northern Gastric-brooding Frog gives birth to 22 young measuring
up to 16mm in length. Oddly, upon examination, the stomach of a female
carrying young did not exhibit any of the physical changes observed in R.
silus. Instead, it retained a normal muscular structure and cells secreting
hydrochloric acid. It is presumed that the mechanism employed by R.
vitellinus to avoid digesting its young was the same as that used by all
vertebrate animals to avoid digesting the stomach wall, namely by coating
the young with mucus.
Sadly, some 18 months after its discovery, the Eungella Gastric-brooding
Frog disappeared without trace and has not been seen since. It had inhabited
dense vegetation in the remote Eungella National Park and there were no
Prematurely born tadpole aged approximately two weeks.
© Michael J. Tyler
or under stones or logs close to the water’s edge in shaded streams. Males of
this species exhibit sexual dimorphism in the form of more muscular, robust
forelimbs than females. This is another distinctive trait, as the other three
Leiopelma species are not sexually dimorphic, apart from females reaching
greater body size. One of the main threats to this species is the destruction
and modification of its habitat, which is still occurring either directly (e.g.
afforestation, gold mining, storm water discharge) or indirectly (e.g. feral
goats and pigs causing erosion leading to stream siltation). The New Zealand
Department of Conservation has purchased considerable amounts of suitable
land to prevent further degradation of some of their habitat, and attempts
are being made to better monitor their populations.
Archey’s Frog Leiopelma archeyi (CR) is the smallest of the indigenous
species (<38mm). They are restricted to two regions on the North Island of
New Zealand, occurring on the Coromandel Peninsula and the Whareorino
Forest, west of Te Kuiti. In both of these areas, it occurs sympatrically with
Leiopelma hochstetteri. They prefer to live at relatively high altitudes from
400-1000m in moist native forest; they are terrestrial and nocturnal, spending
most of the day hidden under stones or logs away from streams or creeks.
Leiopelma archeyi is a terrestrial breeder, laying a small clutch of eggs in a
moist site under stones or logs. They exhibit parental care with the tailed
froglets remaining on their father’s back for several weeks until metamor-
phosis is nearly complete. Populations of this species have crashed in recent
years with monitored populations decreasing by 88% over the 1996-2001
period (Figure 1). Several factors, including the severity and rapidity of the
population crash, the geographic spread of the decline (from south to north),
and the discovery of frogs with chytridiomycosis (caused by Batrachochytrium
dendrobatidis), all point to disease being the major cause of the decline. A
breeding facility has recently been opened at Auckland Zoo with the intention
of producing a self-sustaining captive population. A top priority of the New
71
signs of habitat disturbance or any event linked to the species’ disappear-
ance. The causes for the disappearance of this species, and along with it
a unique mode of reproduction among amphibians, is unknown. Flannery
(2006) asserts that the demise of the Southern Gastric-brooding Frog is
attributable to climate change, but the rate of loss was simply too fast for
this to be the case. In any event, there was no drought and no detectable
environmental change. However, there is some circumstantial evidence that
an environmental factor must have been involved because a small colony
remained alive in a laboratory in Adelaide for three years following their
disappearance from the wild.
At the Second World Congress of Herpetology held in Australia at the end
of 1993, Australian researchers learned that their experience of disappearing
frogs was mirrored by other species on other continents. The search for a
common cause or causes has been ongoing ever since.
Michael J. Tyler
References
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Brooding Frog, pp. 44-57. Croom Helm, London, UK.
An x-ray of a female gastric-brooding frog that has given birth. The dark space
in the abdomen is the stomach which held the young. © Michael J. Tyler
Zealand Native Frog Recovery Group is to investigate ways of preventing
further declines of this species.
Hamilton’s Frog Leiopelma hamiltoni (EN) and the Maud Island Frog
L. pakeka (VU) are the largest living indigenous frogs (<50mm) in New Zealand.
These two species were once considered conspecific, but they have since been
described as separate species based on allozyme variation (Bell et al. 1998).
There is some debate as to whether there should be a distinction between
the two, with further analysis using 12S ribosomal RNA and cytochrome b
gene sequences finding little variation (Holyoake et al. 2001). However, as
both these species are limited to offshore islands with no chance for genetic
exchange, they need to continue to be managed as separate evolutionary
significant units if not kept as separate species.
Both L. hamiltoni and L. pakeka live amongst the boulders and moss
covered rocks of the remnants of coastal forest. They are terrestrial and
nocturnal, spending most of the day hidden under stones or logs away from
streams or creeks. They are terrestrial breeders, laying a small clutch of
eggs in a moist site under suitable substrates. The only naturally occurring
L. hamilitoni population consists of less than 300 adult frogs on Stephens
Island, and after five years of intensive monitoring this population is thought
to be stable. They are limited to a single rock-tumble (ca. 300 m2) that has
been fenced off to stop the predation of these frogs by Tuatara (Sphenodon
punctatus). New habitat was created for the frogs by the construction of a
second rocky tumble on the same island and, in 1992, 12 adult frogs were
transferred to the new site (Brown 1994). The results of the translocation were
mixed, with three frogs remaining at the new site and two homing back to
the original frog site (Brown 1994; Tocher and Brown 2004). In 2004, a fence
was erected to join both of these sites thereby significantly enlarging the
amount of tuatara-free habitat. Also in 2004, a translocation of 40 individuals
to Nukuwiata, another predator-free island in the Marlborough Sounds, took
72
place. Initial reports are that this appears to have been a success with frogs
remaining in good condition through the first year.
Leiopelma pakeka, like L. hamiltoni, was only found naturally on one
small predator-free island, Maud Island, in the Marlborough Sounds. The
population has been estimated at between 27,500 and 39,500 individuals
found mainly in a 16-ha patch of native bush. One hundred of these frogs
were translocated to restored forest habitat on the same island in 1984-1985
and, 20 years later, this new population appears to be thriving with new
recruitment and higher adult weights than the source population (Bell et al.
2004b). Two inter-island translocations to predator-free islands have occurred
since then in order to increase the distribution of L. pakeka and therefore
lower their risk of extinction. The first of these involved moving 300 frogs to
Motuara Island in 1997 where the population survival has been high follow-
ing a 3-month settling in period and evidence of breeding success has been
recorded. A second inter-island translocation to Long Island occurred in 2005
and appears to be successful as all the recaptured frogs have gained weight
and some appear to be gravid.
The continued survival of these ancient species in New Zealand will largely
depend upon conservation measures and research carried out by the Native
Frog Recovery Group and the Department of Conservation.
Phil J. Bishop, Jen M. Germano and Ben D. Bell
References
Bell, B.D. and Wassersug, R.J. 2003. Anatomical Features of Leiopelma Embryos and
Larvae: Implications for Anuran Evolution. Journal of Morphology 256:160-170.
Bell, B.D., Daugherty, C.H. and Hay, J.M. 1998. Leiopelma pakeka, n. sp. (Anura: Leiopelma-
tidae), a cryptic species of frog from Maud Island, New Zealand, and a reassessment
of the conservation status of L. hamiltoni from Stephens Island. Journal of the Royal
Society of New Zealand 28:39-54.
Bell, B.D., Carver, S., Mitchell, N.J. and Pledger, S. 2004a. The recent decline of a New
Zealand endemic: how and why did populations of Archey’s frog Leiopelma archeyi
crash over 1996-2001. Biological Conservation 120:189-199.
Bell, B.D., Pledger, S. and Dewhurst, P.L. 2004b. The fate of a population of the endemic
ESSAY 6.3. AMPHIBIAN DECLINES IN AUSTRALIA
Concern about the status of Australian frogs first arose in the 1980s with
the disappearance of the famous gastric-brooding frogs: Rheobatrachus
silus (described in 1973, and last seen in the wild in 1981), and R. vitellinus
(described in 1984, and last seen in the wild in 1985) (Ingram and McDonald
1993; and see Essay 6.1). Intensive searches for both of these species were
conducted in the late 1980s, with no individuals located then, or since, and
they are both now listed as Extinct on the IUCN Red List. This represents the
loss of an entire family, as well as a unique reproductive strategy.
Around the same time that the gastric-brooding frogs disappeared, concern
was raised over the status of another iconic myobatrachid – the Corroboree
Frog Pseudophryne corroboree (CR). Declines of the Corroboree Frog were
first reported in the late 1980s (Osborne 1989). Although these declines and
disappearances raised concerns at the time, they were regarded as isolated
events. However, increasing awareness that amphibian declines might be a
global problem, and a review of the status of Australian frogs by Tyler (1991),
revealed that the phenomenon of declines was more widespread. Tyler
identified 23 species that were thought to have suffered recent population
declines. The critical nature of the phenomenon was reinforced by Richards
et al. (1993), whose survey of 47 sites in the Australian Wet Tropics showed
that declines were occurring on a regional scale; six species had disappeared
from almost all sites above 400m elevation. These species subsequently
disappeared from the few higher-elevation sites at which they were present
in the initial surveys (McDonald and Alford 1999).
Fortunately, Australia has well-defined procedures in place at a national
level for dealing with threats to biodiversity, and these procedures were quickly
mobilized. In the early 1990s, regional species recovery teams were organized,
Threatened Amphibians of the World
700
600
500
Population estimate (Nhat)
400
300
200
100
0
Aug-83 Jan-84 Feb-85 Feb-86 Dec-86 Feb-88 Feb-89 Feb-90 Jun-91 Jan-93 Feb-94 Dec-94 Dec-96 Feb-97 Feb-98 Feb-99 Feb-00 Dec-00 Dec-01 Dec-02
Figure 1. Jolly-Seber population estimates of population size of Leiopelma archeyi (Critically Endangered) on the Tapu Ridge study plot, 1983-2002. Error
bars = 1SE. (Source: Bell et al. 2004a).
frog Leiopelma pakeka (Anura: Leiopelmatidae) translocated to restored habitat on
Maud Island, New Zealand. New Zealand Journal of Zoology 31:123-131.
Brown, D. 1994. Transfer of Hamilton’s frog, Leiopelma hamiltoni, to a newly cre-
ated habitat on Stephens Island, New Zealand. New Zealand Journal of Zoology
21:425-430.
Holyoake, A., Waldman, B. and Gemmell, N.J. 2001. Determining the species status of one
of the world’s rarest frogs: a conservation dilemma. Animal Conservation 4:29-35.
Lee, J.S.F. and Waldman, B. 2003. Communication by fecal chemosignals in an archaic
frog. Copeia 3:679-686.
Stephenson, E.M. 1961. New Zealand native frogs. Tuatara 8:99-106.
and regional and national meetings and workshops were held. Each regional
recovery team was responsible for developing and implementing a recovery
plan for the species under its jurisdiction. These plans went through formal
review processes at the State and Federal levels. Initially, most plans called
primarily for additional research, since the nature and causes of the declines
were not well understood. At a national level, the federal department of the
environment sponsored the production of the first version of the Action Plan
for Australian Frogs (Tyler 1997). Knowledge of Australian frog populations
and their status increased dramatically during the early to mid-1990s, largely
as a result of the work carried out under the species recovery process. Of the
23 species identified as being of concern by Tyler (1991), sufficient data had
been collected to show that the apparent decline of 12 species was instead
just a manifestation of local fluctuations. However, an additional 16 species
were identified as suffering declines, such that there were now 27 species of
concern. Observations confirmed that many of them were experiencing serious
declines. In late 1997, a meeting was held to review the action plan and discuss
what was known about the status of frogs across the country, and to attempt to
devise some standard methods for investigating and mitigating the effects of
frog declines. The outcomes of this meeting were published in the book Declines
and Disappearances of Australian Frogs (Campbell 1999).
At the time of the 1997 meeting, declines of nine species were linked to
specific causes, including habitat loss and degradation, and the introduction
of exotic predators. However, 18 species fell into a category since termed
“enigmatic declines” by Stuart et al. (2004). Comparative analyses have
shown that these species share a variety of characteristics; for example,
McDonald and Alford (1999) found that species closely associated with stream
Tocher, M.D. and Brown, D. 2004. Leiopelma hamiltoni homing. Herpetological Review
35:259-261.
Waldman, B. and Bishop, P.J. 2004. Chemical communication in an archaic anuran amphib-
ian. Behavioral Ecology 15:88-93.
1 As noted in the Introductory chapter, Frost et al. (2006) include the Ascaphidae (Tailed
Frogs) in this family, with the result that this primitive family is expanded to contain
six species, and its distribution is in New Zealand and the Pacific Northwest of North
America, with the New Zealand species breeding by direct development, and the
American ones by larval development.
habitats have a higher probability of declining, and many authors pointed out
that declines at high elevations had occurred in protected areas.
In 1998, the pathogenic chytrid fungus, Batrachochytrium dendrobatidis,
was described (Berger et al. 1998). Histological examination of dead
and dying individuals collected at the time of some of the declines in
Queensland confirmed that the affected frogs had been heavily infected
with chytrid (Berger et al. 1999). It is now generally accepted that
chytridiomycosis was the proximate cause of mortality in the widespread
die-offs of frogs in the Australian Wet Tropics in the late 1980s and early
1990s, and was probably to blame for the earlier disappearances of
the gastric-brooding frogs and day frogs (Taudactylus spp.) in southern
Queensland. It also appears to be a factor in the decline of Corroboree
Frogs in southern Australia, and may be the cause of local die-offs of
many other species. Since its discovery, chytrid has been detected in 48
native frog species in the wild (Speare and Berger 2005), as well as in the
introduced and now widespread Cane Toad, Bufo marinus. The nomadic
behaviour of this introduced species means that it is one possible vector of
the disease, at least on a local scale. The wide distribution of this disease
and the serious threat that it constitutes to Australian amphibians, and
indeed globally, was explored in detail at a conference in August 2000,
attended by national and international experts (Speare et al. 2001).
Nyctimystes dayi (Endangered) and Litoria nannotis (Endangered) are
rainforest specialists from Australia’s Wet Tropics; both species vanished
from upland sites throughout the region, most likely due to the disease
chytridiomycosis. © Jodi Rowley

One of the outcomes of this conference was the formal nomination of
chytridiomycosis, to the Australian federal Department of Environment
and Heritage, as a threatening process. This nomination went through
several drafts and commenting stages, and was formally adopted in June
2005. Even before its final adoption, the threatening process nomination
prompted state and federal governments to increase levels of funding
of research and management aimed at reducing the threat. A threat
abatement plan was prepared and accepted in 2006 (AGDEH 2006).
Conservation actions targeting threatened amphibians in Australia have
included habitat creation and modification, introduced species control, and
captive breeding. These actions have had some success at individual sites.
Several frog species that survived at low elevations in the Wet Tropics, when
dramatic population declines associated with chytridiomycosis were taking place
at high elevations, appear to be recolonizing some high-elevation sites without
any management intervention. However, 47 species (more than 20%) of extant
Australian amphibians are still listed as threatened in the Global Amphibian
Assessment, and remain susceptible to chytrid (as it spreads to new areas),
continued habitat modification and destruction, and introduced species. A great
deal of work remains before the status of Australian frogs is fully understood and
measures to preserve their remaining diversity can be implemented.
Ross A. Alford and Jodi J.L. Rowley
ESSAY 6.4. AMPHIBIANS OF THE PAPUAN REGION
The Papuan region extends from Indonesia’s Maluku Islands, in the west,
to the Solomon Islands and Fiji, in the east. It is dominated by the world’s
largest and highest tropical island, New Guinea, and includes thousands of
islands, ranging from tiny coral atolls to forest-covered ranges that rise several
kilometres above the ocean. The amphibian fauna here consists entirely of
frogs; salamanders and caecilians do not extend east of Wallace’s Line, which
extends between Bali and Lombok, and Borneo and Sulawesi.
The Papuan frog fauna has been poorly documented compared with
most other regions of the globe, despite the fact that many new species
are being discovered and described each year. More than 320 species are
known from the region, although it has been estimated that the total will
exceed 600 given the current rates of discovery (Günther 2006). Three
families dominate the frog fauna: Microhylidae, Hylidae, and Ranidae.
The Microhylidae is the most species-rich group of frogs on New Guinea.
Chapter 6. Amphibians of the Australasian Realm
References
Berger, L., Speare, R., Daszak, P., Green, D.E., Cunningham, A.A., Goggin, C.L., Slocombe,
R., Ragan, M.A., Hyatt, A.D., McDonald, K.R., Hines, H.B., Lips, K.R., Marantelli, G.
and Parkes, H. 1998. Chytridiomycosis causes amphibian mortality associated with
population declines in the rainforests of Australia and Central America. Proceedings
of the National Academy of Science USA 95:9031-9036.
Berger, L., Speare, R. and Hyatt, A. 1999. Chytrid fungi and amphibian declines: Overview,
implications and future directions. Declines and disappearances of Australian Frogs. In:
A. Campbell (ed.), pp. 23-33. Environment Australia, Canberra, Australia.
Campbell, A. 1999 Declines and disappearances of Australian Frogs. Environment
Australia, Canberra, Australia. Avaliable as pdf at http://www.deh.gov.au/biodiversity/
threatened/publications/pubs/frogs.pdf
Australian Government Department of the Environment and Heritage. 2006. Threat Abatement
Plan: Infection of Amphibians with Chytrid Fungus Resulting in Chytridiomycosis. Department
of the Environment and Heritage, Commonwealth of Australia, Canberra, Australia.
Ingram, G.J. and McDonald, K.R. 1993. An update on the decline of Queensland’s frogs.
In Herpetology in Australia: A Diverse Discipline. In: D. Lunney and D. Ayers (eds.),
pp.297-303. Surrey Beatty and Sons, Sydney, Australia.
McDonald, K. and Alford, R.A. 1999. A review of declining frogs in northern Queensland.
In Declines and disappearances of Australian Frogs. In: A. Campbell (ed.), pp. 14-22.
Environment Australia, Canberra, Australia.
They occupy a diverse range of environments, from sea level to around
4,000m asl, and can be found in semi-aquatic, terrestrial, subterranean,
and arboreal (canopy) habitats. The New Guinea microhylids include some
of the smallest frogs on earth (such as Oreophryne minuta, DD) that is adult
at just 9-11.5mm; Richards and Iskandar 2000), but also include a number of
very large species, including one of the few species (Asterophrys turpicola,
LC) known to attack and bite as a means of self-defence (Richards et al.
1994). All New Guinean microhylids share the habit of laying their eggs
out of water. Embryos develop directly into small froglets, by-passing a
free-swimming tadpole stage. It appears that the father is always respon-
sible for guarding the embryos and, in the case of the sharp-snouted frog
Sphenophryne cornuta (LC), for carrying the freshly hatched youngsters
around on their back (Bickford 2004).
The Hylidae, or treefrogs, are generally found in bushes or trees. However,
in the savannah habitats of southern New Guinea, there are two species
(Litoria nasuta, LC, and L. nigrofrenata, LC) that live exclusively on the ground.
Their long legs and slender build allow them to jump enormous distances to
escape predators. Treefrogs reach their greatest diversity in the mountains
and forests of New Guinea’s interior. Here, one can find bizarre species, such
as Litoria prora (LC), with its elongated nose spike, and its habit of hanging
its eggs from leaves over small forest pools. New Guinea is dominated by
a mountainous spine running across its centre, including the highest peaks
between the Himalaya and Andes. These precipitous mountains are drained
by a myriad of torrential streams, providing a challenging environment for
frogs. Many New Guinean treefrogs in the genera Litoria and Nyctimystes
have taken advantage of these extreme habitats, laying large eggs, glued
under stones, in steep streams. The eggs hatch into tadpoles with enormous
sucker-mouths that cling to rocks, in even the fastest of torrents (e.g.,
Günther 2006).
The Ranidae, or ‘true-frogs’, a family dominating much of the temperate
northern hemisphere, is poorly represented on New Guinea. However, one
group of ranid frogs, the platymantines, dominates the smaller archipelagos
to the west, north, and north-east of New Guinea. Like the microhylids,
these frogs have a ‘direct-development’ reproductive strategy, and it has
been suggested that this has assisted their colonization of far-flung islands
in the South Pacific. Frogs of the genus Platymantis occur on nearly all of
the smaller archipelagos north of New Guinea, and have occupied the
wide range of habitats used by microhylids on mainland New Guinea.
The platymantine radiation is most evident on the Solomon Islands, with
Platymantis and several endemic genera (Batrachylodes, Ceratobatrachus
and Palmatorrapia) dominating the fauna. Many platymantine species are
endemic to single islands or small island groups, and so may be susceptible
to the massive habitat destruction occurring on most archipelagos from
major logging operations.
Three other frog families are poorly represented in the Papuan region:
Limnodynastidae, Myobatrachidae and Bufonidae. Limnodynastidae and
Myobatrachidae dominate the nearby Australian fauna, but only seven
species occur here and they all occur on New Guinea. Perhaps the most
interesting of these are the three species of Lechriodus (Limnodynastidae)
– large, broad-headed frogs that construct floating foam ‘nests’ into which
they place their small eggs during reproduction. Bufonidae, or Toads, do not
occur naturally in New Guinea and are represented by two exotic species,
Bufo marinus and Bufo melanostictus. Bufo marinus, from the Neotropics, is
widespread in eastern New Guinea and in many islands of Melanesia (Lever
2001). Bufo melanostictus is native to western Indonesia and has recently
been introduced to Papua Province (New Guinea) (Iskandar and Colijn 2000;
Menzies and Tapilatu 2000). The impacts of these species (if any) on native
Papuan frogs are poorly understood, and require urgent assessment.
Male Sphenophryne cornuta (Least Concern) are responsible for carrying the
freshly hatched youngsters around on their back. © Stephen Richards
Litoria prora (Least Concern) hangs its eggs from leaves over small forest
pools. © Stephen Richards
73
Osborne, W.S. 1989. Distribution, relative abundance and conservation status of Corro-
boree Frogs, Pseudophryne corroboree (Anura: Myobatrachidae). Australian Wildlife
Research 16:537-547.
Richards, S.J., McDonald, K.R. and Alford, R.A. 1993. Declines in populations of Australia’s
endemic tropical rainforest frogs. Pacific Conservation Biology 1:66-77.
Speare, R, and Berger, L. 2005. Chytridiomycosis in amphibians in Australia. http://www.
jcu.edu.au/school/phtm/PHTM/frogs/chyspec.htm
Speare, R., Alford, R.A., Aplin, K., Berger, L., Bishop, P., Cullen, B., Cunningham, A., Daszak,
P., Dovey, L., Halliday, T. Hines, H., Lynch, M., Marantelli, G., McDonald, K., Orchard,
S. and Owens, L. 2001. Developing management strategies to control amphibian
diseases: Decreasing risks due to communicable diseases. School of Public Health and
Tropical Medicine, James Cook University, Townsville, Australia. Available as a pdf at
http://www.jcu.edu.au/school/phtm/PHTM/frogs/adms/AmpDisStrategies.pdf
Stuart, S.N., Chanson, J.S., Cox, N.A., Young, B.E., Rodrigues, A.S.L., Fischman, D.L.
and Waller, R.W. 2004. Status and Trends of Amphibian Declines and Extinctions
Worldwide. Science 306:1783-1786.
Tyler, M. 1997. The action plan for Australian Frogs. Wildlife Australia, Canberra,
Australia.
Tyler, M.J. 1991. Declining amphibian populations – a global phenomenon? An Australian
perspective. Alytes 9:43-50.
Albericus siegfriedi (Critically Endangered) is known only from high elevations
on Mt Elimbari in Papua New Guinea. © Stephen Richards
Globally, the Papuan region has the highest proportion of Data Deficient
frog species. Many species are known from only one or two localities, and
rugged terrain and logistical and bureaucratic hurdles have hindered attempts
to relocate and assess the conservation status of these frogs. Although few
species appear to be at immediate risk of extinction, the severely restricted
distributions of some species, coupled with increasing rates of forest de-
struction, have raised concerns that a number of species may be susceptible
to extinction in the future. One such example is the small, shrub-dwelling
microhylid Albericus siegfriedi (CR), which is found only at high elevations
on Mt Elimbari in Papua New Guinea, where the only known population is
isolated in a patch of forest that is being gradually consumed by an expanding
local human population.
Further research is required to better understand the distribution and
conservation status of Data Deficient frogs in this poorly documented region.
Given a land-tenure system throughout the region that largely precludes the
designation of protected areas by the central government, conservation of
the region’s unique frog fauna will depend on developing education programs
for, and conservation initiatives with, local landowners. Fortunately, the
Papuan region retains one of the largest blocks of tropical forest on earth,
and appropriate action now will help to ensure the long-term survival of its
spectacular frog fauna.
Stephen Richards
References
Bickford, D.P. 2004. Male parenting of New Guinea froglets. Nature 418:601-602.
Günther, R. 2006. Derived reproductive modes in New Guinean anuran amphibians and
description of a new species with paternal care in the genus Callulops (Microhylidae).
Journal of Zoology 268:153-170.
Iskandar, D.T. and Colijn, E. 2000. Preliminary Checklist of Southeast Asian and New
Guinean Herpetofauna I. Amphibians. Treubia 31(Suppl.):1-134.
Lever, C. 2001. The Cane Toad. The history and ecology of a successful colonist. Westbury
Academic and Scientific, Yorkshire, UK.
Menzies, J.I. and R.F. Tapilatu. 2000. The introduction of a second species of toad (Am-
phibia: Bufonidae) into New Guinea. Science in New Guinea 25:70-73.
Richards, S.J. and Iskandar, D.T. 2000. A new minute Oreophryne (Anura: Microhylidae) from
the mountains of Irian Jaya, Indonesia. Raffles Bulletin of Zoology 48:257-262.
Richards, S.J., Johnston, G.R. and Burton, T.C. 1994. A remarkable new asterophryine
microhylid frog from the mountains of New Guinea. Memoirs of the Queensland
Museum 37:281-286.

Figure 1. Summary of Red List categories
for amphibians in the Indomalayan Realm.
The percentage of species in each category
is also given.
Red List Category Number of species
Extinct (EX)
Extinct in the Wild (EW)
Critically Endangered (CR)
Endangered (EN)
Vulnerable (VU)
Near Threatened (NT)
Least Concern (LC)
Data Deficient (DD)
Total Number of Species
2% 3%
26%
31%
Table 1. The Extinct amphibians of the Indo-
malayan Realm.
Species
Adenomus kandianus
Nannophrys guentheri
Philautus adspersus
Philautus dimbullae
Philautus eximius
Philautus extirpo
Philautus halyi
Philautus hypomelas
Philautus leucorhinus
Philautus malcolmsmithi
Philautus nanus
Philautus nasutus
Philautus oxyrhynchus
Philautus rugatus
Philautus stellatus
Philautus temporalis
Philautus travancoricus1
Philautus variabilis
Philautus zal
Philautus zimmeri
Theloderma gordoni (Least Concern) is an Asian treefrog in the family Rhacophoridae. This species is known from monsoon forests in Thailand
and Vietnam, and is believed to breed in cavities in trees. © Nikolai L. Orlov
CHAPTER 7. AMPHIBIANS OF THE INDOMALAYAN REALM
Raoul Bain, S.D. Biju, Rafe Brown, Indraneil Das, Arvin Diesmos, Sushil Dutta,
David Gower, Robert Inger, Djoko Iskandar, Yoshio Kaneko, Michael Wai Neng
Lau, Madhava Meegaskumbura, Annemarie Ohler, Theodore Papenfuss,
Rohan Pethiyagoda, Bryan Stuart, Mark Wilkinson and Feng Xie
THE GEOGRAPHIC AND HUMAN CONTEXT
20
0 The Indomalayan Realm (sometimes termed the Oriental region) encompasses all of South
32 and Southeast Asia, including the Indonesian and Philippine archipelagos, and incorporating
134 the major offshore islands of Sri Lanka, Hainan, and Taiwan, as well as Japan’s Ryukyu
143 archipelago. The western and northern boundaries follow that of Olson et al. (2001), reaching
104 Pakistan, the Himalaya, and southern subtropical China, although the boundary between
311 the Palaearctic and Indomalayan Realm is somewhat unclear in south-east China. However,
255 as here defined, the eastern boundary between Indomalaya and Australasia, which is usu-
999 ally taken as Wallace’s line (an imaginary line named for Alfred Russell Wallace running
between Borneo and Sulawesi, and between Bali and Lombok in Indonesia), is here taken
to lie further to the east, such that the region includes all of Nusa Tenggara and a number
13% of islands in Maluku (but excluding Seram, Amboin, Buru, Obi, Halmahera, Tanimbar and a
few other smaller islands) (see Tyler 1999).
The geological, evolutionary, and climatic history of this region is complex and is reflected
by the evolutionary history and diversity of its fauna. Peninsular India, which includes Sri
Lanka, consists of a single tectonic plate (the Deccan or Indian Plate) that separated from
14%
Gondwanaland about 130 Ma and, after breaking away from Madagascar and the Seychelles
around 90 Ma, rafted across the Tethys Sea eventually colliding with Eurasia at about 65-
40 Ma (Beck et al. 2005). This massive collision resulted in the uplift of the Himalaya and
10%
the Tibetan plateau, which caused dramatic climatic changes over vast expanses across
South Asia. Although the climate of almost the whole of Peninsular India is monsoonal,
the region is varied both in terms of topography and vegetation, including, for example,
rainforests (e.g., in the Western Ghats, south-western Sri Lanka, and Myanmar), arid areas
(such as the Thar Desert in north-western India), low-lying swamps and mangroves (in the
Sundarbans), and island systems (Andamans and Nicobars). Peninsular India is relatively
flat, and is highest in the south-west, with the western flank of the plateau being formed
by the Western Ghats.
Similarly, the coming together of the Indian plate with the Asian continental landmass
has influenced much of the topography in mainland Southeast Asia, including the general
Country north-south orientation of the mountains and main rivers. Much of this region (often referred
Sri Lanka to more generally as Indo-Burma) is characterized by distinct seasonal weather patterns (for
Sri Lanka example, in northern Vietnam and southern coastal China, the dominant weather pattern is
Sri Lanka the north or north-easterly monsoon during the northern winter and east or south-easterly
Sri Lanka monsoon in the summer). Originally, most of the region was dominated by broadleaf forests;
Sri Lanka the most diverse forests are the lowland mixed wet evergreen forests, which occur in
Sri Lanka climates with one to four dry months.
Sri Lanka Another geological highlight of the region is the islands forming part of the Malay Archi-
Sri Lanka pelago, comprising the Greater Sundas – including Borneo and Sumatra (the third and sixth
Sri Lanka largest islands on earth, respectively), Java, and Sulawesi – the Lesser Sundas, the Philip-
Sri Lanka pines, and several islands of the Moluccas). This is one of the most active seismic regions
Sri Lanka in the world, and the site of some of the most dramatic seismic events known, including
Sri Lanka the eruption of Krakatau in 1883 and the earthquake that caused a massive tsunami in the
Sri Lanka Indian Ocean, just off the coast of Aceh, Sumatra, on December 26th, 2004. The highest point
Sri Lanka in the region is Gunung Kinabalu in northern Borneo at 4,101m. The islands of the Sunda
Sri Lanka shelf were connected to mainland Southeast Asia through most, if not all, of the Tertiary,
Sri Lanka and were also periodically connected during episodes of northern glaciation during the
India Quaternary, which is why the fauna and flora of these two regions have much in common.
Sri Lanka At the same time, oscillations in sea levels caused periodic severing of these ephemeral
Sri Lanka land bridges, isolating nearby continental islands, and presumably allowing for the evolution
Sri Lanka and accumulation of endemic species. The climate is tropical, and the vegetation, at least
up until a few decades ago, comprised mainly lowland evergreen rainforest.
Human population density is very high across this region (averaging 124 people per
square kilometre across Southeast Asia), including, as it does, several of the most populous
countries on earth, such as India (with an estimated 1.1 billion people) and Indonesia (220
million). Population density ranges from a whopping 336 people per square kilometre in
India, to 277 per square kilometre in the Philippines, 117 people per square kilometre in
Indonesia, to 25 people per square kilometre in Lao P.D.R. The percentage of the population
concentrated in urban areas also varies, with nearly 20% of people in Cambodia concentrated
in urban areas, 30% in India, around 48% in Indonesia, and nearly two-thirds of people in
the Philippines and Malaysia. With the exception of Singapore (gross national income per
capita of US$24,000), all countries have a GNI per capita of less than US$5,000.
Given the high human population densities in the region, the impact of society on eco-
systems has been severe. Mainland Southeast Asia was probably one of the first regions
where agriculture developed (Diamond 1997), and there has been a long history of shifting
or permanent small-scale agriculture. More recently, though, the exploitation of Southeast
Asia’s valuable timber for commercial trade, and the demand for land to grow cash-crops
and trees, have led to widespread and rampant forest loss, particularly of lowland evergreen
forest, dominated mainly by the giant dipterocarps. Several estimates of forest loss across
the region are available; one recent study estimates that Kalimantan’s protected lowland
forests declined by 56% between 1985 and 2001 primarily from logging (Curran et al. 2004),
and that less than 33% of lowland forest and peat swamp remains across all of Indonesian
Borneo (Whitten et al. 2005). Unfortunately, even where rainforest habitat remains relatively
intact, the unmitigated harvest and trade of some of the larger species of animals has been
so intensive that the term “empty forest syndrome” was coined (Redford 1992); this “empty
forest” phenomenon is particular apparent in China, Vietnam, Laos and Cambodia.
GLOBAL CONSERVATION STATUS
A total of 329 (33%) of the amphibian species in the Indomalayan Realm are considered to
be globally threatened or Extinct (Figure 1). This is very similar to the global average. The
Indomalayan Realm contains 17% of all globally threatened amphibians. When looking at
the Red List Categories, Indomalaya accounts for only 7% of the world’s CR species, but 17%
of the EN species, and 21% of the VU species. Hence, on the basis of current knowledge,
threatened Indomalayan amphibians are more likely to be in a lower category of threat,
when compared with the global distribution of threatened species amongst categories. The
percentage of DD species, 26% (255 species), is also similar to, though slightly higher than,
the global average of 23%. This high percentage is not surprising, given that much of the
region is still very poorly surveyed for amphibians.
Twenty of the world’s 34 known amphibian extinctions (59%) have occurred in this region
(Table 1), 19 of these in Sri Lanka and one in southern India. Eighteen of these species are
frogs from the genus Philautus, and most of these probably had tiny ranges and died out
as a result of extensive forest loss, perhaps as long ago as the late 1800s or early 1900s
(Manamendra-Arachchi and Pethiyagoda 2005). In addition, one Critically Endangered
species in the Indomalayan Realm is considered to be possibly extinct, Philautus jacobsoni
from central Java, Indonesia. It is suspected that the apparent concentration of extinctions
in Sri Lanka, as opposed to other parts of the region, is a result of better knowledge of this
country due to recent herpetological work (e.g., Manamendra-Arachchi and Pethiyagoda
2005; and see Essay 4.1), in which the extant fauna has been extensively surveyed and
compared with the historical baseline provided by museum specimens.
SPECIES RICHNESS AND ENDEMISM
Species Richness and Endemism Across Taxa
The 999 native amphibian species in the Indomalayan Realm represent 17% of the currently
known global total of 5,915 species. Of these, 800 (or 80%) are endemic to the region (Table
2). All three amphibian orders, are represented in the Indomalayan Realm, but the frogs
account for 92% of the species. Contrary to the situation in some other regions, endemism
is much lower in the salamanders (46%) as compared with the frogs and toads (80%). This is
because most of the Indomalayan salamanders occur in central China, on the poorly defined
boundary (which is in fact a broad overlap zone) with the Palaearctic, thus occurring in both
regions. Caecilian endemism is 100%. Although Indomalaya has the second highest number
of species of any realm (though well behind the Neotropics), it has only 14 families, which
is fewer than any realm except Australasia. Three of these families are endemic. Only 45
species (5% of the species in the region) are members of these endemic families, although
the treefrog family Rhacophoridae occurs only marginally in the Palaearctic and Afrotropical
Regions and is predominantly Indomalayan.
Under current climatic conditions, there is essentially no isolation between the Palaearctic
and Indomalayan Realms, especially in China, and the boundary between these two faunas is
somewhat arbitrary. The effect of this indistinct boundary is to reduce the level of endemism
of both regions. Summaries of the amphibian fauna of the Indomalayan Realm are provided
by Bourret (1942), Inger (1999), Iskandar and Colijn (2000), and Zhao (1999).
There are 81 genera (18% of the global total) occurring in the region, of which 37 (46%)
are also endemic. Endemism at the generic level is much lower among the salamanders (with
no endemic genera) than it is among the frogs and toads (46%), contrary to the situation in
the Palaearctic. Generic level endemism is 100% among the caecilians. The most speciose
endemic genera in the region are Ichthyophis (34 species), Ansonia (22 species), Kalophrynus
(15 species), Nyctibatrachus (12 species) and Micrixalus (11 species). At the opposite end
of the spectrum, there are 11 monotypic genera endemic to the Indomalayan Realm, all
of which are frogs. The 44 non-endemic genera in Indomalaya include 37 frog genera (13
genera from the Ranidae, eight from the Megophryidae, seven from the Rhacophoridae,
five from the Microhylidae, two from the Hylidae, and one each from Bombinatoridae and
Bufonidae) and seven salamander genera (five from the Salamandridae, and one each from
the Cryptobranchidae and the Hynobiidae). These non-endemics include the widespread
genera Bufo, Rana and Litoria. It should be noted that future taxonomic changes are likely
to have a major impact on the patterns outlined above, with a tendency for the number of
genera (including monotypic genera) to increase.
 Chapter 7. Amphibians of the Indomalayan Realm 75

As noted already, 29% (14/48) of the world’s amphibian families occur in the Indo-
malayan Realm, and three of these are endemic: Nasikabatrachidae, Ichthyophiidae, and Family Native species Percentage of Percentage of Native genera Percentage of Percentage of
Uraeotyphlidae.2 The characteristics of these families are provided in Chapter 1. Among (endemics to species in region species in family (endemics to genera in region genera in family
region) that are endemic that are endemic region) that are endemic that are endemic
the non-endemic families, the majority of Indomalayan species are in the Bufonidae (true
to region to region
toads), Megophryidae (Asian spadefoots), Microhylidae (narrow-mouthed toads), Ranidae
(true frogs), Rhacophoridae (Asian treefrogs), and Salamandridae (newts and relatives). The Anura
Bufonidae occur widely in the Indomalayan Realm as far south and east as Sulawesi and Bombinatoridae 4 (3) 75 30 2 (1) 50 50
the southern parts of the Philippines, with 84 species in eight genera.3 Most species in the Bufonidae 84 (72) 86 15 8 (7) 88 21
region are endemic, but 12 species are shared with the Palaearctic. All Indomalayan species Hylidae 12 (4) 33 0.5 2 (0) 0 0
breed by larval development, and occur in many different habitats. Megophryidae 90 (56) 62 44 10 (2) 20 20
There are 90 species across 10 genera in the Megophryidae in the Indomalayan Realm. Microhylidae 88 (77) 89 18 14 (9) 64 13
This family is predominantly Indomalayan, with over 70% of its species occurring in the Nasikabatrachidae 1 (1) 100 100 1 (1) 100 100
region, and 44% of them globally endemic (all other species occurring in the Palaearctic Ranidae 375 (290) 77 44 24 (11) 46 28
Region). Thirty-four species in China cross the Indomalayan-Palaearctic boundary. The family Rhacophoridae 263 (230) 87 84 8 (1) 13 11
ranges from Nepal, Bangladesh, and north-eastern India, through central and southern China, TOTAL ANURA 917 (733) 80 14 69 (32) 46 9
and Southeast Asia as far as Java, Borneo and the Philippines. Caudata
The Microhylidae range very widely through the region, with 88 species, 77 of which Cryptobranchidae 1 (0) 0 0 1 (0) 0 0
are endemic. They occur in a wide variety of habitats, and all Indomalayan species breed by Hynobiidae 6 (3) 50 7 1 (0) 0 0
larval development, except for eight species of the genus Oreophryne in the eastern parts Salamandridae 21 (10) 48 14 5 (0) 0 0
of Indonesia and the Philippines which are direct developers. Most of the non-endemic TOTAL CAUDATA 28 (13) 46 2 7 (0) 0 0
species are shared with the Palaearctic. Gymnophiona
The Ranidae constitute the largest family in the Indomalayan Realm, accounting for Caeciliidae 10 (10) 100 9 2 (2) 100 8
over one-third of the total amphibian fauna of the region. One-third of the ranids are in Ichthyophiidae 39 (39) 100 100 2 (2) 100 100
the genus Rana4. The family is found throughout the region, occurring in most habitats, Uraeotyphlidae 5 (5) 100 100 1 (1) 100 100
and all species breed by larval development, except in the genera Ingerana (5 species) and TOTAL GYMNOPHIONA 54 (54) 100 31 5 (5) 100 15
Platymantis (27 species).5 TOTAL ALL AMPHIBIANS 999 (800) 80 14 81 (37) 46 8
The Rhacophoridae are a predominantly Indomalayan family, with 263 species (and
eight genera) occurring in the region, of which 230 are endemic. The family occurs widely
through the region, east to the Philippines and Sulawesi. Many of the species are arboreal, Table 2. The number of Indomalayan am-
and this family includes the flying frogs. The family is split approximately evenly between phibians in each taxonomic Family present
direct developers (many species in the genus Philautus) and larval developers (some of in the region.
which use foam nests).6
The Salamandridae are predominantly a Palaearctic family, but 21 species occur in the
Indomalayan Realm, 10 of which are endemic. Most of the Indomalayan species occur in
southern China. All Indomalayan species breed by larval development.
Among the smaller non-endemic families, the Bombinatoridae (fire-bellied toads) have
an unusual distribution. This family occurs mainly in the Palaearctic, but two species also
occur in southern China (one extending into northern Vietnam), and another two (in the
genus Barbourula) are highly isolated from the rest of the family in Kalimantan (southern
Borneo) and in the Palawan island group (south-western Philippines). Some of these are
highly aquatic species, with the genus Bombina breeding by larval development (the breeding
remaining unknown in Barbourula).
The Hylidae are absent from much of the region, but eight species in the genus Hyla The Hole-in-the-Head Frog Huia cavitympanum
occur in the northern parts of the region (mainly in China) and four species in the genus (Least Concern) is in the Family Ranidae and
Litoria occur on islands in the extreme east of the region in Indonesia. is endemic to central and northern Borneo,
The giant salamanders (Cryptobranchidae) are represented in the region by a single non- where it inhabits rainforests in hilly terrain.
endemic species (the Chinese Giant Salamander Andrias davidianus), which until recently The tadpoles cling to rocks in strong rapids in
occurred widely in southern China. These animals are aquatic and are associated with clear clear streams. © Nikolai L. Orlov
streams where they breed by larval development.
The Asian salamanders (Hynobiidae) are mainly a Palaearctic family having their distribu-
tion centred on Japan and China, with six species occurring in the Indomalayan Realm. All This close-up view of the head of Ichthyophis
species have larval development. tricolor (Least Concern) shows the tentacle
The caecilian family Caeciliidae occurs predominantly in the Neotropics and Afrotropics, which is characteristic of caecilians. This
but 10 species occur in India, mainly in the Western Ghats in the south of the country, though subterranean species from the Western Ghats
one species is present in the north-east. All Indomalayan species are assumed to have in India lives in soil in wet semi-evergreen
terrestrial eggs and breed by direct development, although direct evidence is very scarce tropical forest, but also occurs in farmland and
(only known for one species, Gegeneophis ramaswamii). rubber plantations. Like other members of the
There are high percentages of threatened and extinct species in most families in the Family Ichthyophiidae, it has aquatic larvae in
Indomalayan Realm (Table 3). The three small families Bombinatoridae, Nasikabatrachidae, streams. © Photo by John Measey, courtesy of
and Cryptobranchidae are entirely composed of threatened species, and all but one spe- The Natural History Museum, London
cies of Hynobiidae is threatened. The threat level among the salamanders is much higher
than that of the frogs, with over half of the species at risk. The salamander faunas of the
Palaearctic, Nearctic and Neotropics also face high threat levels. In the Indomalayan Realm,
over-harvesting for medicine and food, coupled with habitat loss and/or restricted range,
are probably the most significant threats facing these species (see later). Conversely, threat
levels among caecilians appear to be very low, but this is probably an artefact of over 85%
of the species in the region being Data Deficient (see Gower and Wilkinson 2005). Table 3. The number of species within each
IUCN Red List Category in each Family and
Order in the Indomalayan Realm. Introduced
species are not included.

Family EX CR EN VU NT LC DD Total number Number threatened % Threatened

of species or Extinct or Extinct
Anura
Bombinatoridae 0 0 1 3 0 0 0 4 4 100
Bufonidae 1 3 17 13 11 27 12 84 34 40
Hylidae 0 0 0 0 0 9 3 12 0 0
Megophryidae 0 1 6 20 10 34 19 90 27 30
Microhylidae 0 1 8 12 11 31 25 88 21 24
Nasikabatrachidae 0 0 1 0 0 0 0 1 1 100
Ranidae 1 8 43 55 43 141 84 375 107 29
Rhacophoridae 18 17 50 33 25 57 63 263 118 45
TOTAL ANURA 20 30 126 136 100 298 206 917 312 34
Caudata
Cryptobranchidae 0 1 0 0 0 0 0 1 1 100
Hynobiidae 0 0 3 2 0 0 1 6 5 83
Salamandridae 0 1 5 3 4 6 2 21 9 43
TOTAL CAUDATA 0 2 8 5 4 6 3 28 15 54
Gymnophiona
Caeciliidae 0 0 0 0 0 1 9 10 0 0
Ichthyophiidae 0 0 0 2 0 5 32 39 2 5
Uraeotyphlidae 0 0 0 0 0 0 5 5 0 0
Figure 2. The species richness of amphibians in the Indomalayan Realm, with darker colours TOTAL GYMNOPHIONA 0 0 0 2 0 6 46 54 2 4
corresponding to regions of higher richness. Colour scale based on 10 quantile classes; TOTAL ALL AMPHIBIANS 20 32 134 143 104 310 255 999 329 33
maximum richness equals 84 species.
76 Threatened Amphibians of the World

Figure 3. a) The richness of threatened

amphibians in the Indomalayan Realm, with
darker colours corresponding to regions of
higher richness. Colour scale based on 10
quantile classes; maximum richness equals
29 species. b) The richness of CR amphibians
in the Indomalayan Realm, with darker colours
corresponding to regions of higher richness.
Colour scale based on four quantile classes;
maximum richness equals eight species.

a. b.

Figure 4. The number of extant amphibians
present in and endemic to each Indomalayan
country. *denotes countries not entirely within
the Indomalayan Realm, hence only the spe-
cies whose ranges fall within the region are
included. Note that 102 described species
are known from Sri Lanka, but because 19
of these are extinct, only 83 are included in
this figure.
Figure 5. Percentage of species endemic to
each Indomalayan country. Countries with no
endemic species are not included. *denotes
countries not entirely within the Indomalayan
Realm, hence only the species whose ranges
fall within the region are included.
Figure 6. The number of threatened am-
phibians present in and endemic to each
Indomalayan country. Countries with no
threatened species are not included in the
diagram. *denotes countries not entirely
within the Indomalayan Realm, hence only the
species whose ranges fall within the region
are included.
Figure 7. Percentage of native species that are
threatened. Countries with no threatened spe-
cies are not included in the diagram. *denotes
countries not entirely within the Indomalayan
Realm, hence only the species whose ranges
fall within the region are included.
In general, the largest families in the region house the highest number of threatened
frog species: Rhacophoridae, Ranidae, Bufonidae, Megophryidae and Microhylidae (Table
3). The Rhacophoridae has a particularly high percentage (45%) of threatened species, much
of this reflecting high threat levels (54% of species) in the genus Philautus, in which many
species have tiny ranges and can be seriously affected, even by the loss of small patches
of habitat. In the Bufonidae, over 40% of the species are threatened, which is similar to
confamilial levels in the Neotropics and Afrotropics (but not in the Palaearctic). Most
threatened bufonid species (75%) are dependent on clear mountain streams in forests for
breeding, a very threatened habitat (their larvae can be adversely affected by even modest
levels of silt in the stream, which is a common affect of logging and other forms of forest
clearance). The Megophryidae are dependent on the same habitats, are similarly impacted
by siltation, and also face a high threat level (30%). The Indomalayan species of the family
Ranidae are also facing extensive threats from both over-harvesting for human food and
from habitat loss. The Microhylidae have the lowest level of threat among the larger frog
families, but over 27% of the species are Data Deficient, higher than any other frog family,
so this may be an underestimate. There are no threatened Hylidae in the region.
The great majority (90%) of the threatened amphibians in the Indomalayan Realm are
either Endangered or Vulnerable. Furthermore, 17 of the 32 Critically Endangered species
are rhacophorids, and 15 are in the genus Philautus (seven of these in India, seven in Sri
Lanka, and one from Indonesia).
Geographic Patterns of Species Richness and
Endemism
A map of overall species richness of amphibians in the Indomalayan Realm (Figure 2) shows great
variation across the region. However, more than perhaps any other major biogeographic region,
this map is somewhat biased by sampling intensity, and probably represents a misleading picture
of amphibian species richness in this part of the world. Some of the overall patterns are probably
accurate; for example, the large areas of low species richness in the drier parts of northern,
central and eastern India, and in the lower Mekong Delta, and the peaks of highest species
richness in the Western Ghats, south-western Sri Lanka, the Malaysian Peninsula, and northern
Borneo. The overall patterns of species richness in southern China are also likely to be reasonably
accurate. Through most of mainland Southeast Asia (excluding Malaysia), north-eastern India,
Nepal, Bhutan, Sumatra, Kalimantan (Indonesian Borneo), Sulawesi, the Lesser Sunda Islands,
and the Philippines, the patterns on Figure 2 are likely to reflect sampling intensity. Areas that
are particularly poorly surveyed, include Bhutan, Myanmar, Thailand, Laos, Cambodiag, Vietnam
and most of Indonesia (the stark contrast in recorded species richness between Malaysian and
Indonesian Borneo emphasizes this point) (see Essay 7.1). As the results of future surveys and
taxonomic work are incorporated into the Global Amphibian Assessment, our understanding of
the patterns of Indomalayan amphibian species richness will change considerably.
The same caveats apply to the interpretation of the distribution of threatened species
(Figure 3a) in the Indomalayan Realm. The concentrations of threatened species in the
Western Ghats, Sri Lanka, southern China, northern Borneo, Java, and the Philippines
probably reflect reality because these areas have been relatively heavily surveyed. How-
ever, there are likely to be important concentrations in places such as Myanmar, Thailand,
Laos, Cambodia, Sumatra, Kalimantan, and Sulawesi that remain undetected due to lower
sampling effort. Not surprisingly, given the small number of species involved, there are few
noteworthy concentrations of Critically Endangered species in the region (Figure 3b), the
most important being in Sri Lanka and southern India around the Western Ghats (see Essay
7.2; Essay 1.2). Much of the apparent concentration in China reflects the originally wide
distribution of one species, the Chinese Giant Salamander.
Species Richness and Endemism within Countries
Amphibians occur naturally in 20 countries in the Indomalayan Realm (all except the Mal-
dives). India has the largest number of species (236 extant) in the region (Figure 4), followed
quite closely by China, Indonesia, and Malaysia (all have over 200 extant species). Vietnam
and Thailand have over 100 extant species (Sri Lanka has 102 described species, but 19 are
now considered extinct), but many countries have very low totals that almost certainly reflect
inadequate survey effort (most notably Myanmar, Laos, Nepal, Cambodia, and Bhutan). India
has by far the largest number of endemics (151 species), and China, Philippines, Indonesia,
Sri Lanka, and Malaysia each have more than 50 endemics.
The amphibian fauna of parts of the Indomalayan Realm has been summarized in numer-
ous national-level publications, including: India (Tiwari 1992; Dutta 1997; Das 1999, 2002;
Daniel 2002; Daniels 2005); Pakistan (Khan 2006); Sri Lanka (Dutta and Manamendra-Arachchi
1996); Nepal (Schleich and Kästle 2002); China (Ye et al. 1993; Zhao and Adler 1993; Fei et
al. 1999, 2005; Zhao et al. 2000); Japan (Maeda and Matsui 1999; Uchiyama et al. 2002;
Goris and Maeda 2004); Peninsular Malaysia (Berry 1975); Thailand (Nabhitabhata 1989;
Chan-ard 2003); Vietnam (Bain and Nguyen 2004; Bourret 1942; Inger et al. 1999; Ohler
et al. 2000; Orlov et al. 2001, 2002; Ziegler 2002); Laos (Bourret 1942; Stuart 1999, 2005;
Teynie et al. 2004); Cambodia (Ohler et al. 2002; Stuart et al. 2006; Stuart and Emmett in
press); Thailand and Peninsular Malaysia (Chan-ard et al. 1999); Singapore (Lim and Lim
2002); Borneo (Inger 1966; Inger and Stuebing 1997; Malkmus et al. 2002); Java and Bali
(Iskandar 1998; McKay 2006); and Philippines (Alcala and Brown 1998).
Although India has many more endemics than any other country in the region, Japanh, Sri
Lanka and the Philippines (see Essay 7.3) have higher percentages of endemic species (all
above or around 80%; Figure 5). Endemism is over 60% in India, over 50% in China, over 40%
in Taiwan, and over 30% in Indonesia. The percentage endemism in a number of countries,
such as Indonesia, can be expected to rise as the fauna becomes better known.
India has more threatened species (63) than any other country in the Indomalayan Realm
(Figure 6). Countries with over 40 threatened species are Sri Lanka, Philippines, Malaysia
and China. Indonesia has 29 threatened species; this surprisingly small number is probably
a reflection of how poorly the amphibian fauna is known in this country (Essay 7.1). Vietnam
has 15 threatened species, also probably a significant under-estimate (30% of the species
in this country are Data Deficient). In much of mainland Southeast Asia, the numbers of
threatened species are likely to be seriously under-estimated (partly because much of the
amphibian fauna remains to be discovered, for example in Cambodia, Laos, Myanmar and
Thailand; and see Essay 7.4).
The percentage of threatened amphibian species is highest in island nations (Figure 7),
notably Sri Lanka (63%), the Philippines (49%), and Japan (46%). All other Indomalayan

India* Japan* India* Sri Lanka

China* Sri Lanka Sri Lanka Philippines
Indonesia* Philippines
Philippines Japan*
Malaysia
India* Taiwan, Province
Viet Nam Malaysia of China
China*
Thailand China* India*
Taiwan, Province
Philippines of China
Indonesia* Malaysia
Sri Lanka Indonesia*
Malaysia Viet Nam China*
Myanmar
Lao P.D.R. Viet Nam Japan* Bhutan
Brunei Darussalam Nepal Taiwan, Province Indonesia*
of China
Nepal Myanmar Lao P.D.R. Viet Nam
Cambodia Bhutan
Taiwan, Province Cambodia Cambodia
of China Lao P.D.R.
Bangladesh Nepal Nepal
Thailand
Singapore Brunei Darussalam Brunei Darussalam
Cambodia
Japan*
Pakistan* Thailand Lao P.D.R.
Pakistan*
Singapore Bhutan Bangladesh
Bhutan
Timor-Leste Bangladesh Bangladesh Thailand

0 50 100 150 200 250 300 0 20 40 60 80 100 0 20 40 60 80 0 20 40 60 80

Number of species Percentage of endemic species Number of threatened species Percentage of species threatened
Figure 4 Endemics Non-endemics Figure 5 Figure 6 Endemics Non-endemics Figure 7
 Chapter 7. Amphibians of the Indomalayan Realm 77

countries have levels of threat that are much lower than global average of 33%, though as
mentioned above, this is likely to be under-estimated in several countries.
Assessments of the conservation status of Indomalayan amphibians have been carried out
in only a few countries, for example: India (Molur and Walker 1998); Japan (Japan Agency
of Environment 2000; Ota 2000); China (Zhao 1998; Xie and Wang 2004); and Philippines
(Wildlife Conservation Society of the Philippines 1997). A regional overview of the threatened
status of amphibians and reptiles in South Asia was published by Bamabaradeniya and
Samarasekara (2001), and Pawar et al. (2007) carried out an assessment and prioritization
of areas for amphibian conservation in north-eastern India.
There are only 32 Critically Endangered Indomalayan species, but 13 of these occur in
India and 11 in Sri Lanka. Outside these two countries, there are three Critically Endangered
species each in China and Indonesia, and one each in Malaysia and the Philippines.

HABITAT AND ECOLOGY

Habitat Preferences
Most Indomalayan amphibians (82%) occur in forests, including 66% in lowland tropical
forest, and 47% in montane tropical forest (Table 4). As in other regions, forest species are
more threatened than those occurring in other terrestrial habitats, and montane forest species
are more threatened than those in lowland forest. However, the level of threat to lowland
tropical forest species in the Indomalayan Realm (33%) is higher than that in the Afrotropi-
cal Region (23%), though very similar to that in the Neotropical Region (30%). Among the
aquatic habitats, the level of threat is highest in flowing freshwater. So, as in other regions,
forest-dwelling and stream-associated amphibians are more likely to be threatened than
those occurring in any other habitats. This is the combination of habitat preferences that has
been associated with rapid declines in amphibian populations worldwide (Stuart et al. 2004).
Almost one-quarter of the fauna (23%) can survive in secondary terrestrial habitats (Table 4;
Figure 8). This latter figure is higher than in either the Afrotropics or the Neotropics. Table 4
and Figure 8 show that amphibians occurring in savannahs, shrubland, and arid and semi-arid
habitats are less likely to be threatened than those occurring in other habitats.

Reproductive modes
Larval development is by far the most common reproductive mode in the Indomalayan Realm
(81% of species), compared with 18% for direct development (Table 5). There are no live-
bearing species in the region. These figures compare with the global picture of 68% larval
development, 30% direct development, and 1% live-bearing. The presumed direct-developing
Indomalayan amphibians are dominated by rhacophorid treefrogs in the genus Philautus,
and also include the ranid frog genera Platymantis and Ingerana, and the microhylid genus
Oreophryne, as well as perhaps the caecilian genera Gegeneophis and Indotyphlus.
In the Indomalayan Realm, the percentage of globally threatened or Extinct direct-de-
veloping species is much higher than in the larval-developing species (Table 5), a pattern
repeated in several other regions.
MAJOR THREATS
As is the case in all other regions, habitat loss is overwhelmingly the major threat to amphib-
ians in the Indomalayan Realm (Table 6; Figure 9), affecting nearly 90% of the threatened
species. Pollution is the next most serious threat, impacting nearly one-third of threatened
species. All other threats have much lower impacts, although utilization is implicated in
the rapid decline of over 20 species (see below). Many of those species being utilized are
listed as Near Threatened, so they do not show in this analysis. Chytridiomycosis has not
been recorded in the region.
The impacts of vegetation removal (mainly via logging) (affecting 64% of the threatened
species) and expanding croplands (61%) are the most severe types of habitat loss impacting
amphibians, followed by urbanization and industrial development (46%) and tree plantations
(18%). Livestock constitutes a less important threat in most cases.
This unidentified, and possibly undescribed,
A total of 143 species (26 of which are threatened) are recorded as being harvested by species of Leptobrachium from the Annamite
people in the region. The most common reasons for harvesting are for human consumption Mountains in Cambodia is from the Asian
(112 species, mostly at local and national levels), pet trade (31 species, mostly at international spadefoot Family Megophryidae. Like most
and national levels), and medicine (27 species, mostly at local and national levels) (Table 7). other members of the Family, it is associ-
Not all of the amphibian harvesting in the region is considered to constitute a major threat ated with streams in hilly forested areas. ©
to these species. Of the 143 species being harvested, utilization is considered to be a threat David Emmett
for 87 (of which 26 are threatened species for which harvesting is believed to be contributing
to a deterioration in their status). Twenty of these 26 species seriously threatened by over-
harvesting occur in China, where many species of amphibians are extensively harvested for
human food and medicines. Examples include 16 species of ranid frog (eight in the genus
Paa), and six species of salamander (including the Chinese Giant Salamander). Threatened
species outside China that are heavily harvested include three species of ranid frog in the
Philippines and two in Indonesia.
POPULATION STATUS AND TRENDS
Estimates of Population Trends
A summary of the inferred population trends of Indomalayan amphibians is presented in
Table 8, inferred from trends in the state of the habitats on which the species depend (though
in some cases, population declines have been noted, especially for species that are being
over-harvested). The overall population trends of Indomalayan amphibians are worse than
the global trends (where 42% are decreasing and only 27% are stable). In both cases, the
percentage of increasing species is very small.

“Rapidly Declining” Species

Habitat type Number of % of all Threatened % of species Of the 470 globally “rapidly declining” species, 58 (12%) occur within the Indomalayan
species in species or Extinct occurring Realm. Twenty of these 58 species are in decline due to over-exploitation, 37 due to reduced
each habitat occurring in species in habitat that
habitat, and one due to so-called “enigmatic declines”. Not surprisingly for this region, more
the habitat are Threatened
or Extinct declines are attributed to reduced habitat and over-exploitation than to enigmatic declines.
The Indomalayan Realm accounts for 53% of the world’s rapid declines due to over-exploita-
Forest 823 82 298 36 tion, but only 18% of the reduced habitat declines, and 0.4% of the enigmatic declines.
All tropical forest 800 80 291 36 Although one species in the region, Leptophryne cruentata (CR) from Java, has been recorded
Lowland tropical forest 655 66 218 33 as undergoing an enigmatic decline, the causes of this decline are not known, and have not
Montane tropical forest 474 47 195 41 so far been linked to either chytridiomycosis or climate change (although these two threats
Savannah 17 2 0 0 have now been associated with many such declines that have taken place elsewhere in the
Grassland 95 10 21 22
Shrubland 116 12 14 12
Secondary terrestrial habitats 229 23 50 22
Flowing freshwater 505 51 155 31
Marsh/swamp 113 11 19 17
Still open freshwater 300 30 55 18
Number of species % of species in habitat that are threatened
Arid and semi-arid habitats 3 0.4 0 0
Forest

Savannah

Table 4. The habitat preferences of amphibians in the Indomalayan Realm. Grassland

Shrubland

Secondary terrestrial habitats

Reproductive mode All Species Threatened or % Threatened Flowing freshwater
Extinct species or Extinct
Marsh/swamp
Direct development 181 121 67
Still open freshwater
Larval development 807 204 25 Figure 8. The habitat preferences of Indoma-
Live-bearing 0 0 - Arid and semi-arid habitats layan amphibians. The plot on the left-hand
Not known 11 4 36 side shows the number of species in the region
0 200 400 600 800 0 10 20 30
in each habitat type. On the right-hand side,
the percentage of these species which are
Table 5. Indomalayan amphibians categorized by reproductive mode. threatened is given.
78 Threatened Amphibians of the World

Table 6. The major threats to globally threat-

ened amphibians in the Indomalayan Realm. Threat type Threatened species % Threatened Species
Only present threats to species are tallied. Habitat loss 272 88
Agriculture – Crops 188 61
Agriculture – Tree plantations 57 18
Agriculture – Livestock 15 5
Timber and other vegetation removal 198 64
Urbanization and industrial development 142 46
Invasive species 10 3
Utilization 26 8
Accidental mortality 10 3
Pollution 100 32
Natural disasters 25 8
Disease 1 0.3
Human disturbance 18 6
Fire 17 6

Figure 9. The major threats impacting threat- The Shanjing Emperor Newt Tylototriton shanjing (Near Threatened) from the Family Sala-
ened amphibians in the Indomalayan Realm. mandridae is known only from Yunnan in southern China where it inhabits hill forests and
All habitat loss
secondary forest. It is subject to over-collection for traditional Chinese medicine, and small
numbers are also exported for the international pet trade. © Henk Wallays
Invasive species

Utilization

Accidental mortality
Persecution
Pollution
Natural disasters
Disease
Human disturbance
Changes in native sp. dynamics
Table 7. The purposes for which amphibians
are used in the Indomalayan Realm. The Purpose
numbers in brackets are the number of species
within the total that are threatened species. Food – human
Food – animal
Medicine – human and veterinary
Pets, display animals
Research
Specimen collecting
Table 8. The population trends for all extant
Indomalayan amphibians. Population Trend
Decreasing
Stable
Increasing
Unknown
Table 9. The number of species in “rapid
decline” and “over exploited decline” in the Family
Indomalayan Realm by Family.
Bufonidae
Megophryidae
Microhylidae
Ranidae
Rhacophoridae
Cryptobranchidae
Salamandridae
Breeding and larval development in the
microhylid frog Metaphrynella sundana
(Least Concern) takes place in water-filled
tree holes from where the males call. It lives
in lowland primary rainforest, and is widely
distributed in Borneo, with a single specimen
having been collected from northern Sumatra.
© Björn Lardner
world (Lips et al. 2006; Pounds et al. 2006). A full list of all “rapidly declining” species is
provided in Appendix IV and includes their occurrence within each of the realms.
The “rapidly declining” species in the Indomalayan Realm show a distinct taxonomic
pattern (Table 9), as over half of them, and 75% of “over-exploited” species are ranids.
Among the larger families, the Bufonidae, Salamandridae, Ranidae and Microhylidae show a
higher tendency to serious decline than the Megophryidae and Rhacophoridae. There are no
Indomalayan species in serious decline in the Bombinatoridae, Hylidae, Nasikabatrachidae,
Hynobiidae, Caeciliidae, Ichthyophiidae and Uraeotyphlidae (though these last three are very
poorly known caecilian families in which most species are Data Deficient). In one small family,
Fire
Cryptobranchidae, the only species in the region is in “rapid decline” and “over-exploited
decline”. Among the larger families, “over-exploited declines” are concentrated in the
0 20 40 60 80 100 Ranidae and the Salamandridae.
Percentage of threatened species affected Species in “rapid decline” in the Indomalayan Realm show a clear geographic pattern
as well, since the major concentration of declines is in Malaysia (32 species) and Indonesia
(31 species), followed by China (15), Vietnam (seven), and Thailand and the Philippines (both
with five). For Malaysia and Indonesia, most of the declining species are on Borneo, where
there has been very severe loss of lowland rainforest (see Essay 7.5). Of the “over-exploited
Subsistence Sub-national/ Regional/ Number declines”, 14 are in China, and seven in Vietnam.
National International of species
109 (22) 32 (8) 8 (1) 112 (22)
1 (0) 0 0 1 (0) KEY FINDINGS
27 (2) 13 (2) 3 (0) 27 (2)
5 (2) 21 (6) 30 (6) 31 (7) • A total of 999 species are recorded from the Indomalayan Realm, of which 329 (33%)
1 (1) 6 (1) 1 (0) 7 (1) are considered threatened or Extinct.
1 (1) 0 0 1 (1) • At the species level, 800 amphibians (80%) are endemic to the Indomalayan Realm;
of the 14 families found in the region, three are endemic, and of 81 amphibian genera
occurring, 37 are endemic. Endemism would be higher, were it not for the unclear and
somewhat arbitrary boundary with the Palaearctic Region, especially in China.
• The percentage of threatened and/or extinct species is higher than in many other parts of
Number of species % of extant species the world, and highest in the families Bombinatoridae (100%), Nasikabatrachidae (100%),
509 52 Cryptobranchidae (100%), Hynobiidae (83%), Rhacophoridae (45%), Salamandridae
161 16 (43%), and Bufonidae (40%).
2 0.2 • Overall, the threat levels are much higher among salamander species (54%) than frogs
307 31 (34%). Caecilians are very poorly known in the region, with over 85% of the species
being Data Deficient.
• Geographic concentrations of threatened species in the Indomalayan Realm occur in the
Western Ghats (southern India), Sri Lanka, southern China, northern Borneo, Java and the
Philippines; there are likely to be important concentrations in places such as Myanmar,
Number of Percentage Number of Percentage Thailand, Laos, Cambodia, Sumatra, Kalimantan, and Sulawesi that remain undetected.
species in of species species in of species • India has the largest number of species (236) in the region, followed quite closely by
“rapid decline” in family in “over-exploited in family in
“rapid decline” decline” “over-exploited
China, Indonesia and Malaysia (all have over 200 species). Many countries have very
decline”
low totals that almost certainly reflect inadequate survey efforts.
• India has by far the largest number of endemics (151 species) within the region,
8 10 0 0 and China, Philippines, Indonesia, Sri Lanka, and Malaysia each have more than 50
2 2 0 0 endemics.
6 7 2 2 • India has more threatened species (63) than any other country in the Indomalayan Realm.
30 8 15 4 Countries with over 40 threatened species are Sri Lanka, Philippines, China, and Malaysia.
9 3 0 0 • The percentage of threatened amphibian species is highest in island nations, notably
1 100 1 100 Sri Lanka (63%), the Philippines (49%), and Japan (46%).
2 10 2 10 • Threatened species tend to show distinct habitat preferences, with forest-dwelling and
stream-associated species being the most threatened (36% and 31%, respectively). This
mirrors patterns seen elsewhere in the world.
• Habitat loss, primarily due to the impacts of vegetation removal (mainly logging),
expanding croplands, and urbanization and industrial development is affecting nearly
90% of the threatened species in the region. Pollution impacts nearly one-third of the
threatened species. Chytridiomycosis, the emerging amphibian fungal disease, has not
been recorded as a significant threat in the region so far.
• Of the 470 globally “rapidly declining” species, 12% occur within the region. Most of
these rapid declines (69%) are caused by severe habitat loss, and 34% are due to over-
exploitation. Over-exploitation is a more serious threat in the Indomalayan Realm than
in any other part of the world, except the Palaearctic.
• Twenty amphibian extinctions have been recorded from the Indomalayan Realm, 19 in
Sri Lanka and one in India. One species (from Indonesia) is possibly extinct.
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80
ESSAY 7.1. AN OVERVIEW OF OUR KNOWLEDGE ON INDONESIAN AMPHIBIANS
Oreophryne minuta (Data Deficient) from the Derewo River Basin in the moun-
tains of western Papua, Indonesia, at 2,000m asl. © Djoko Iskandar
Straddling the tropics with more than 17,000 islands, Indonesia has the
longest coastline of any country in the world, and also has among the high-
est mountains in the tropics. Indonesia consists of three bioregions, namely
Sundaland, Wallacea, and the Papuan realm. Sundaland comprises the three
large islands of Borneo, Sumatra and Java, although politically Borneo is
divided among three nations. Wallacea includes the large island of Sulawesi,
and the smaller islands of two extensive archipelagos: the Malukus and the
Lesser Sundas (Nusa Tenggara). The Papuan realm comprises Indonesia Papua
(on New Guinea), plus a number of offshore islands.
That Indonesia is ranked as the second richest country in terms of known
biodiversity in the world (Mittermeier et al. 1997) is significant when one
considers that much of the region remains unexplored or poorly surveyed.
Nonetheless, the remarkable diversity and endemism of its flora and fauna
is underscored in the reports of numerous undescribed species discovered in
practically every new expedition undertaken in the country. According to the
results of the Global Amphibian Assessment, there are nearly 350 species of
amphibians documented from Indonesia (ranking Indonesia as the sixth most
important country for amphibian diversity), of which nearly half (46%) are
endemic. However, as an indication of just how poorly known the fauna is,
one-third of amphibian species in the country are classed as Data Deficient
on the IUCN Red List (significantly higher than the global average).
The amphibian fauna on Indonesia has been for the most part overlooked
since the end of World War II. The islands of Sumatra and Sulawesi are
particularly poorly known, and very few publications deal with these two
large islands. The faunas of Kalimantan (Indonesia Borneo) and Indonesian
Papua, on the other hand, are relatively better documented, but only as a direct
result of the influence of herpetological surveys undertaken in neighbouring
Sabah and Sarawak (Malaysia) in Borneo, and Papua New Guinea in New
Guinea. Whereas the number of described amphibians on Sumatra stood at
68 in 1923, the total now stands at around 100 species mainly due to new
discoveries, although this number is likely to increase exponentially given
our current knowledge of undescribed forms. There is a particular paucity of
information on amphibian species in montane or even or at medium elevations
on Sumatra (Inger and Voris 2001; Inger and Iskandar 2005). This is also the
case for Borneo. For example, at the time of the publication of the first edition
of their guide to the frogs of Borneo, Inger and Stuebing (1997) recorded 15
Philautus species from Borneo, though not a single species was observed
in the Indonesian part (Kalimantan) even though this area occupies roughly
two-thirds of the land mass. More than 80% of the amphibian species of
Borneo have been described from either Sabah or Sarawak.
Sulawesi has very few species in common with other islands. At present,
the island is considered to be species poor, although there are indications
that numerous species await formal description, especially in the genera
Limnonectes and Rhacophorus (Iskandar and Tjan 1994; Evans et al. 2002).
Iskandar and Tjan (1994) reported at least 13 undescribed amphibian species
ESSAY 7.2. DIVERSITY AND CONSERVATION STATUS OF THE WESTERN GHATS AMPHIBIANS
The Western Ghats are a chain of mountains in western India running parallel
to the coast for over 1,600km. The mountains in the northern portion begin
as low-lying hills close to the Tapti River in Gujarat, increase in height as
they pass southwards through the States of Maharashtra, Goa, Karnataka
and Kerala, and end abruptly in the Mahendragiri Hills of Tamil Nadu State
- the southernmost tip of Peninsular India. Along their entire length, there is
only one major discontinuity, the biogeographically important ‘Palghat Gap’
of Kerala, which is approximately 30km wide and has an elevation of less
than 100m above sea level (Figure 2).
Perhaps surprisingly for such a populous country as India, about one-third
of the Western Ghats is still covered by natural vegetation, including about
20,000km2 of rapidly diminishing tropical moist forest (Collins 1990). Although
these areas may constitute only around 5% of the total land area of India, they
contain at least 30% of India’s native species (Rodgers and Panwar 1988).
The results of the Global Amphibian Assessment indicated that 237 am-
phibian species are present within India1. This impressive diversity includes
212 species of frogs and toads in seven families and 37 genera, at least 25
Threatened Amphibians of the World
and several other new records and doubled the number of species hitherto
known from the island. However, the low number of amphibian genera
represented is an indication that this island may truly have an impoverished
fauna. The Lesser Sunda Islands are essentially arid and consequently low
in species diversity, as is the case for the Malukus. When a workshop on
the Biodiversity of New Guinea was held in Biak1, a study revealed that the
ratio of publications on Indonesian Papua compared with Papua New Guinea
was roughly 1 to 14, suggesting that very few works have been done in the
Indonesian part. This is also evident in the number of amphibian species in
Indonesian Papua, which currently has at least 100 fewer described species
than Papua New Guinea, even though Indonesia represents roughly half the
land mass (see also Essay 6.4).
Of the three greater regions, Sundaland is richest in terms of species,
although amphibian composition differs greatly among the three main islands,
particularly at the generic level. The island of Borneo, the largest and the
most ecologically diverse of three, has an extremely high number of endemic
species (see Essay 7.5), though bearing in mind that most Bornean species
have been described from Sabah or Sarawak. By contrast, the Wallacean
region is depauperate in species (especially in the Lesser Sundas), with a
little over 30 species recorded.
There is much variation in body-size among Indonesia amphibians, rang-
ing from about 10mm in Oreophryne minuta (DD) from Papua (Richards and
Iskandar 2000) – one of the smallest amphibian species in the world – to
about 300mm in Limnonectes blythii (NT) of Sumatra, one of the largest frog
species. Reproductive strategies include parental care (Inger 1966; Inger and
Voris 1988; Brown and Iskandar 2002; Günther 2006) to tadpole laying in an as
yet unnamed Sulawesian species (Iskandar and Tjan 1994; and see Chapter
1). Most eggs are laid in a single gelatinous mass, but many Limnonectes
and Platymantis and all the New Guinean microhylids have a derived mode
of reproduction (parental care).
Unfortunately, Indonesia is a country experiencing an exceptionally high
rate of forest loss due to a combination of land conversion and forest fires,
and while only 10% of the country’s amphibians are listed as threatened,
this probably will be shown to be considerably higher with further survey
work. Most forest loss has occurred in the last three decades, a result of
commercial logging and major agricultural projects (including oil palm plan-
Nyctixalus margaritifer (Vulnerable) occurs on the island of Java, Indonesia,
at elevations above 700m asl. It was rediscovered in 1997 after a long period
without any records. © Djoko Iskandar
species of caecilians, and a single species of salamander (Tylototriton ver-
rucosus) that lives in the mountains of the north-east. The amphibian fauna
of India as a whole has been discussed in a number of publications, including
Inger and Dutta (1986), Inger et al. (1987), Daniels (1992), Dutta (1997), Pillai
and Ravichandran (1999), and Das (2000)2.
The amphibian diversity of the Western Ghats is distinctive both in its
diversity and endemism (Biju 2001; Biju and Bossuyt 2003). These mountains
currently hold 131 recognized amphibian species in 25 genera, with 114 of
these species (87%) being entirely restricted to this biodiversity hotspot.
Higher-taxonomic-level endemism clearly makes this region important in
the Asiatic region (Roelants et al. 2004; Bossuyt et al. 2004), with two
families (Nasikabatrachidae and Uraeotyphlidae), and eight genera (Indirana,
Indotyphlus, Melanobatrachus, Micrixalus, Minervarya, Nasikabatrachus,
Nyctibatrachus, Uraeotyphlus) being endemic3.
According to the results of the GAA, 53 amphibian species of the Western
Ghats, or 40% of the amphibian fauna, are threatened with extinction4 (Figure
1). In addition to the high number of threatened species, it is also worrying
tations) in combination with government policies. In Sumatra, for example,
illegal and unsustainable logging and non-timber forest product extraction
are widespread, and fueled by high demand for hardwood timber from China,
North America, Europe, and Japan. Fires have become a major threat in recent
years, and may often be linked to logging operations that create flammable
conditions by both leaving fuelwood on the forest floor, and through expos-
ing the understory to drying (Whitten et al. 2004). It may be suspected that
global climate change is resulting in the drying and dessication of a number
of large aquatic areas, such as the Fly River Basin and lowland areas, but
further detailed study is needed.
In conclusion, our knowledge of Indonesia’s amphibians is scattered and
largely based on what is known from faunistic surveys in a few areas only; no
single area, even on Java, has reliable data on the ecology and distribution
of amphibians. Survey work is hampered by the fact that many areas in the
region are too remote or inaccessible for quick study and assessment, and
the facilities and resources available for study are generally lacking. However,
notwithstanding, there are several regions within Indonesia that represent
urgent priorities for further survey work, particularly since they are likely
to be characterized by high levels of endemism, including: Mounts Leuser
and Kerinci in Sumatra, the Muller and Meratus ranges in Kalimantan, the
Mengkoka Mountains in Sulawesi, and most parts of Indonesian Papua. Fur-
thermore, we have virtually no amphibian records for small- to medium-sized
islands such as Karimata in the west and most of the Malukus, especially as
many of them have relatively unexplored high mountains.
Djoko T. Iskandar
References
Brown, R.M. and Iskandar, D.T. 2000. Nest Site Selection, Larval Hatching and Advertise-
ment Calls of Limnonectes arathooni from southwestern Sulawesi (Celebes) Island,
Indonesia. Journal of Herpetology 34.3:404-413.
Evans, B.J., Brown, R.M., McGuire, J.A., Supriatna, J., Andayani, N., Diesmos, A., Iskan-
dar, D., Melnick, D.J., Canatella, D.C. 2003. Phylogenetics of Fanged Frogs: Testing
Biogeographical Hypotheses at the Interface of the Asian and Australian Faunal Zone.
Systemic Biology 52:794-819.
Günther, R. 2006. Derived reproductive modes in New Guinean anuran amphibians and
description of a new species with paternal care in the genus Callulops (Microhylidae).
Journal of Zoology 268:153-170
Inger, R.F. 1966. The systematics and zoogeography of the Amphibia of Borneo. Fieldiana
Zoology 52:1-402.
Inger, R.F. and Iskandar, D.T. 2005. A collection of Amphibians from West Sumatra, with
description of a new species of Megophrys (amphibia: Anura) Raffles Bulletin of
Zoology 53:133-142.
Inger, R.F. and Voris, H.K. 1988. Taxonomic status and reproductive biology of Bornean
tadpole-carrying frogs. Copeia 1988:1060-1062.
Inger, R.F. and Voris, H.K. 2001. The biogeographical relations of the frogs and snakes of
Sundaland. Journal of Biogeography 28:863-891.
Inger, R.F. and Stuebing, R.B. A Field Guide to the Frogs of Borneo. Natural History Publica-
tions, Kota Kinabalu, Borneo.
Iskandar, D.T. and Tjan, K.N. 1996. The Amphibians and Reptiles of Sulawesi, with notes
on the distribution and chromosomal number of frogs. In: D.J. Kitchener and A. Suyanto
(eds.), Proc. of the First International Conference on Eastern Indonesian-Australian
Vertebrates, pp.39-46. Western Australian Museum for Lembaga Ilmu Pengetahuan
Indonesia, Perth, Western Australia, Australia.
Mittermeier, R.A., Gil, P.R. and Mittermeier, C.G. 1997. Megadiversity. Earth’s Biologically
Wealthiest Nations. CEMEX, Mexico City, Mexico.
Richards, S.R. and Iskandar, D.T. 2000. A new tiny species of frog of the genus Oreophryne
(Anura, Microhylidae) from the mountains of Irian Jaya, Indonesia Raffles Bulletin of
Zoology 48:257-262.
Whitten, T., van Dijk, P.P., Curran, L., Meijaard, E., Wood, P., Supriatna, J. and Ellis, S.
2004. Sundaland. In: R.A. Mittermeier, P. Robles-Gil, M. Hoffmann, J.D. Pilgrim, T.M.
Brooks, C.G. Mittermeier, J.L. Lamoreux and G. Fonseca (eds.), Hotspots Revisited:
Earth’s Biologically Richest and Most Endangered Terrestrial Ecoregions, pp. 164-172.
CEMEX, Mexico City, Mexico.
1 The Irian Jaya Biodiversity Conservation Priority-Setting Workshop. Conservation
International. Biak 7-11 January 1997.
that many of the once locally common species (e.g., Nyctibatrachus aliciae,
N. minor, Micrixalus fuscus, Rhacophorus lateralis and several of the more
widespread Philautus species) appear to have visibly declined in recent years
(S.D. Biju pers. obs.). Several species have not been recorded since their
original descriptions (e.g., Philautus flaviventris and P. chalazodes), and the
possibility exists that they are extinct.
It is very likely that the leading threat to the amphibian species of the
Western Ghats is the continuing conversion or modification of natural
habitats. This loss of habitat is largely driven by the continuing growth of
the human population in this area, and the basic needs of these people for
both agricultural and urban land. Significant threats to the remaining natural
forests also come from the ongoing expansion of plantations (including both
non-native timber plantations and tea and coffee estates); commercial logging
operations; the extensive extraction of forest products such as firewood; and,
perhaps more localized, but nonetheless highly damaging, mining for metal
ores and gemstones. While there are no records to date in the Western Ghats
of the disease chytridiomycosis, which has been implicated in the consider-
 Chapter 7. Amphibians of the Indomalayan Realm 81

Ansonia ‘Bufo’ Pedostibes Kaloula Melanobatrachus

2 species 10 species 1 species 1 species 1 species

Microhyla Ramanella Uperodon Nasikabatrachus Euphlyctis

3 species 6 species 1 species 1 species 2 species

Fejervarya Hoplobatrachus Sphaerotheca Minervarya ‘Rana’

10 species 2 species 4 species 1 species 4 species

Indirana Micrixalus Nyctibatrachus ‘Philautus’ Polypedates

10 species 11 species 12 species 21 species 2 species

Rhacophorus Gegeneophis Indotyphlus Ichthyophis Uraeotyphlus

5 species 7 species 2 species 7 species 5 species

able declines of amphibian communities in Latin America and Australia, there known only from the original, historical description, which can often be brief or
0.7%
is a need for field surveys to confirm the current absence of this pathogen. incomplete in a contemporary context, and in many instances does not include 7.6%
The primary means of protecting the amphibians of the Western Ghats is enough specific details about the initial collection locality. Additionally, the Red List Category Number
31.3% 20.6%
through the region’s extensive system of protected areas. There are a total type-series has sometimes been lost or misplaced, meaning that there is very of species
of nine National Parks and 45 Wildlife Sanctuaries in the mountain range little information available to guide contemporary workers. Extinct (EX) 1
(Kothari et al. 1989; recent updating), covering a total area of 16,935km2 or However, the greatest impediment to conservation and management Extinct in the Wild (EW) 0
11% of the Ghats (Collins 1990). Many of the threatened amphibian species of this rich amphibian fauna may be hidden in the vast number of species Critically Endangered (CR) 10
have at least some part of their range within these parks and reserves, but that remain to be described. Intense fieldwork (Biju 2001) has revealed that Endangered (EN) 27
it will be important to rapidly characterize how many Critically Endangered many clearly morphologically distinct species have not yet been scientifi- Vulnerable (VU) 16
and Endangered species are not present within these protected areas. Urgent cally documented. In addition, the intra-population structure of several of Near Threatened (NT) 6
steps are needed to also protect the remaining habitat of these species to the currently well-recognized species remains very poorly known. It is Least Concern (LC) 30
prevent imminent extinctions. becoming increasingly clearer that some of the taxa that are considered to Data Deficient (DD) 41 12.3%
22.9%
One of the largest problems for conserving the amphibian fauna of the be common and widespread in the Western Ghats may actually represent Total taxa 131
4.6%
Western Ghats is the lack of detailed systematic and other biological infor- cryptic ‘species complexes’ – groups of similar looking taxa that form distinct
mation for much of the region’s amphibian species. Some 41 species from evolutionary lineages (Bickford et al. 2007). In view of the ongoing threats
the Western Ghats are categorized as Data Deficient – these are species for to the remaining natural habitat of the Western Ghats, and considering that
which insufficient details are available on the taxonomic identity, distribution, a number of species currently hidden within these complexes will have
or threats to determine whether these animals are of global conservation restricted ranges, these species may, in turn, be of significant conservation Figure 1. Summary of the Red List status for amphibians in the Western Ghats,
concern. A number of the Data Deficient species from the Western Ghats are concern. Molecular approaches, such as DNA barcoding (see Essay 11.11), based on the results of the Global Amphibian Assessment.
82
Figure 2. Map of the Western Ghats showing a generalized boundary of the
region, elevation, and protected areas in white.
ESSAY 7.3. PHILIPPINE AMPHIBIAN SPECIES BIODIVERSITY IS INCREASING BY LEAPS AND BOUNDS
The discovery and description of the diversity of Philippine amphibians began
with early European and American professional naturalist collectors who
made ancillary collections of amphibian specimens and returned these to
museums in their native countries. Descriptions of these specimens were later
prepared by early herpetologists such as Duméril, Bibron, Peters, Boettger,
Boulenger, Günther, Mertens, Wiegmann, and Stejneger, among others. The
first published descriptions of endemic Philippine species were soon followed
by discoveries of strange and unique species that captured the attention and
curiosity of biologists around the world.
Looking back as students of the history of herpetology in the Philippines,
it is convenient now for us to think of five separate chapters in the study of
Philippine herpetological diversity (Brown et al. 2002). These include the initial
period of exploration described above, followed by the career of Edward Taylor
(1913-1975). We think of Taylor as the “father” of Philippine herpetology because
his work, involving multiple detailed monographs, resulted in descriptions of so
many of the truly spectacular Philippine endemics and a first true appreciation
of the staggering magnitude of herpetological diversity in the archipelago. The
third phase was marked by the work of Robert Inger and the publication of his
monograph “Systematics and zoogeography of Philippine Amphibia (Inger 1954).
Inger’s comprehensive review of Philippine amphibians marked a turning point
in the history of herpetology in the country because of his systematic application
of an explicitly stated species concept, statistical treatment of natural variation,
and other advances. The fourth stage of Philippine herpetology includes the
lengthy and productive collaboration of Angel Alcala and the late Walter Brown
(1958-2000). This body of work included numerous comprehensive taxonomic
reviews, new species descriptions, and a variety of the first ecological and
developmental studies in Philippine herpetology. Finally, we consider the present
day, on-going effort to review the amphibians and reptiles of the Philippines a
fifth phase in the development of the study of the herpetofauna of the country.
An examination of species accumulation over these years (Figure 1) provides
us with an appreciation of the magnitude of taxonomic contributions from each
of these five distinct periods in Philippine herpetology.
In 1993, the discovery of a new species of forest frog in the genus Platy-
mantis from the mossy forests of Panay Island in central Philippines triggered
a major reconsideration of species boundaries within this group. The new
species (P. panayensis, EN) differed from an adjacent population (P. hazelae,
EN) on nearby Negros Island by subtle differences in morphological propor-
tions, slight differences in coloration, and by its distinct male advertisement
call (Brown et al. 1997). The realization that closely related species may
differ primarily by advertisement call unleashed a plethora of active fieldwork
and taxonomic studies, resulting in a doubling of the number of species of
Platymantis from 12 to 24 species, between 1997 and 2001 (summarized in
Alcala and Brown 1999). With the appreciation that advertisement calls may
provide us with insight into truly biologically meaningful suite of characters,
we undertook a comprehensive assessment of the acoustic diversity of Philip-
pine forest frogs along with a thorough re-evaluation of the species diversity
in the Philippine members of the genus Platymantis. The other major advance
Threatened Amphibians of the World
will be extremely useful to rapidly map many aspects of amphibian diversity
in the Western Ghats.
Urgent work is now needed to describe, document, and protect these ex-
ceptional biota. To resolve the existing confusion, greater emphasis is needed
on serious coordinated research activities for the Western Ghats amphibians.
Perhaps most urgently, detailed collaborative studies by scientists working
in different geographical areas of this long mountain chain are needed to
finally determine the correct taxonomic identification of many of the region’s
amphibians. The conservation management of the exceptional amphibian
fauna of the Western Ghats, and the biological diversity of this mountain
range as a whole, can only benefit from such an investment.
S D Biju, Rachunliu G Kamei, Bhatta, G., Varad Giri, Neil Cox,
Indraneil Das and Franky Bossuyt
References
Bhatta, G., Dinesh, K.P., Prashanth, P. and Kulkarni, N.U. 2007. A new species of Ge-
geneophis Peters (Amphibia: Gymnophiona: Caeciliidae) from Goa, India. Zootaxa
1409:51-59.
Bickford, D., Lohman, D.J., Sodhi, N.S., Ng, P.K.L., Meier, R., Winker, K., Ingram, K. and
Das, I. 2007. Cryptic species as a window on diversity and conservation. Trends in
Ecology and Evolution 22(3):148-155.
Biju, S.D. 2001. A Synopsis to the Frog Fauna of the Western Ghats, India. Occasional
Papers of the Indian Society for Conservation Biology 1:1-24.
Biju, S.D and Bossuyt, F. 2003. New frog family from India reveals an ancient biogeographi-
cal link with the Seychelles. Nature 425:283-284.
Bossuyt, F., Meegaskumbara, M., Beenaerts, N., Gower, D., Pethiyagoda, R., Roelants,
K., Mannaert, A., Wilkinson, M., Bahir, M.M., Manamendra-Arachchi, K., Ng, P.K.L.,
Schneider, C.J., Oommen, O.V. and Milinkovitch, M.C. 2004. Local endemism within
the Western Ghats-Sri Lanka biodiversity hotspot. Science 306:479-481.
Collins, D. 1990. The Last Rain Forests. Mitchell Beazley and IUCN, London.
Daniels, R.J. 1992. Geographical distribution patterns of amphibians in the Western Ghats,
India. Journal of Biogeography 19:521-529.
Das, I. 2000. Updates in amphibian systematics and nomenclature for the Indian region.
Biodiversity India, ISCB Newsletter 8-12:3-5.
Das, I. and Dutta, S.K. 2006. New species of Polypedates (Anura: Rhacophoridae) from the
Western Ghats, southwest India. Journal of Herpetology. 40(2):214-220.
Dutta, S.K. 1997. Amphibians of India and Sri Lanka (Checklist and bibliography). Odessy
Publishing House, Bhubaneswar. 342 pp.
Frost, D.R. 2007. Amphibian Species of the World: an online reference. Version 5.0 (12
in improving our understanding of species diversity in the Philippines has been
the application of molecular phylogenetic approaches (Brown and Guttman
2002; Evans et al. 2003; Brown 2004) to new collections of genetic samples
of amphibian species from a robust geographic coverage throughout the major
island groups of the country. The combination of these new tools have pro-
vided new insights into species boundaries and helped uncover the presence
of numerous cryptic species that had gone unnoticed for so many decades.
Initial results of this ongoing work fortified our understanding of the degree
to which biodiversity in Philippine Amphibia has been grossly underestimated
by traditional, primarily morphology-based taxonomic practices (Figure 1).
The result is a new appreciation of diversity that provides a fuller, more bal-
anced, and biologically meaningful appreciation of the complex interactions
of characteristics that have surfaced as most meaningful for the process of
lineage diversification in Philippine amphibians (Figure 2).
Our current understanding of amphibian species diversity in the Philippines
stands at 97 indigenous species (Brown et al. 2002; Diesmos et al. 2002).
However, species descriptions of new frogs of the genus Platymantis currently
in progress will soon increase that number to around 130 taxa. And, if work on
other undescribed species of frogs of other genera that we are aware of were
to be completed, the total number would eventually reach at least 165 species.
If current trends in rates of species discoveries hold as biologists explore the
still many biologically unexplored regions of the country, we expect a possible
doubling of the richness of Philippine Amphibia within the next two decades.
200
180
160
140
Cumulative Total
120
Taylor
100
Günther, Peters, Boulenger,
80 Boettger, Stejneger, etc.
60
Linnaeus
40
20
0
1840
1850
1860
1870
1880
1890
1900
1910
1920
Year of description
Figure 1. Species accumulation curve for Philippine amphibians, including endemic (circles) and non-endemic (squares) species. Estimates of numbers of new
species awaiting description are based on a combination of morphological, behavioural, and ecological character differences, with species’ distinctiveness
confirmed by bioacoustic and molecular data.
July 2007). Electronic Database accessible at http://research.amnh.org/herpetology/
amphibia/index.html. American Museum of Natural History, New York, USA.
Frost, D.R., Grant, T., Faivovich, J.N., Bain, R.H., Haas, A., Haddad, C.F.B., de Sá, R.O.,
Channing, A., Wilkinson, M., Donnellan, S.C., Raxworthy, C.J., Campbell, J.A., Blotto,
B.L., Moler, P., Drewes, R.C., Nussbaum, R.A., Lynch, J.D., Green, D.M. and Wheeler,
W.C. 2006. The amphibian tree of life. Bulletin of the American Museum of Natural
History 297:1-370.
Inger, R.F. and Dutta, S.K. 1986. An overview of the amphibian fauna of India. Journal of
the Bombay Natural History Society 83:135-146.
Inger, R.F., Shaffer, H.B., Koshy, M. and Bakde, R. 1987. Ecological structure of a herpeto-
logical assemblage in South India. Amphibia-Reptilia 8:189-202.
Kothari, A., Pratibha, P., Shekhar, S. and Dilnavaz, V. 1989. Management of National
Parks and Sanctuaries in India: A Status Report. Ministry of Environment and Forests,
Government of Kerala, New Delhi.
Pillai, R.S. and Ravichandran, M.S. 1999. Gymnophiona (Amphibia) of India, a taxonomic
study. Records of the Zoological Survey of India, Occasional Paper No. 172. 117 pp.
Rodgers, W.A. and Panwar, H.S. 1988. Planning a Wildlife Protected Area, Network India.
Wildlife Institute of India, Dehra Dun.
Roelants, K., Jiang, J. and Bossuyt, F. 2004. Endemic ranid (Amphibia: Anura) genera
in southern mountain ranges of the Indian subcontinent represent ancient frog
lineages: evidence from molecular data. Molecular Phylogenetics and Evolution
31:730-740.
Roelants, K., Gower, D.J., Wilkinson, M., Loader, S.P., Biju, S.D., Guillaume, K., Moriau, L.
and Bossuyt, F. 2007. Global patterns of diversification in the history of modern amphib-
ians. Proceedings of the National Academy of Sciences, USA 104:887-892.
Van Bocxlaer, I., Roelants, K., Biju, S.D., Nagaraju, J. and Bossuyt, F. 2006. Late Cretaceous
Vicariance in Gondwanan Amphibians. PLoS ONE 1(1):e74.
1 Frost (2007) currently recognizes 265 amphibian species present in India. This total
includes a number of recently described species, such as the tree frog Polypedates
occidentalis Das and Dutta (2006), not included in the 2005 results of the GAA.
2 Frost (2007) recognizes a diversity of 237 species of frogs and toads in 12 families
(including the Western Ghats endemic Micrixalidae) and 50 genera, 27 species of
caecilians, and a single species of salamander. Since this last version, a few additional
new species of amphibian, such as the caecilian Gegeneophis goaensis (Bhatta et al.
2007), have been described from the Western Ghats.
3 Also, see Frost et al. (2006), Van Bocxlaer et al. (2006), Frost (2007) and Roelants et
al. (2007), for emerging information on the high biological diversity of the Western
Ghats.
4 In addition to the 52 threatened species (Critically Endangered, Endangered or
Vulnerable), there are six Near Threatened species, and a single species regarded as
Extinct.
The irony of the astonishing rates of species discovery, even as so many
species are declining (Hanken 1999; Stuart et al. 2004; Köhler et al. 2005),
coupled with the devastating loss of forested habitat in the Philippines,
convinces us that no higher conservation urgency in the world exists than
that of the megadiverse Philippine global biodiversity hotspot (Myers et
al. 2000; Catibog-Sinha and Heaney 2006). With so few trained workers
actively working to discover and describe new species of Philippine amphib-
ians, we are left with doubt as to whether we can survey and characterize
Philippine amphibian biodiversity within the timeframe of our own careers.
Consequently, there can be no greater priority than training new students in
amphibian field studies and doing everything possible to overcome logistical
and bureaucratic obstacles to field work while at the same time investing in
collections and related repositories. It is through the building of natural history
collection resources (including digital photographic archives, sound libraries,
and genetic collection resources) that will enable tomorrow’s generations
of biologists the opportunity to re-assess our work in light of technological
advances of the future. Coupled with constant and regular conservation
status assessments, we are convinced that these efforts provide the best
chances of preventing impending catastrophic amphibian extinctions that
loom on the horizon if we fail to take action now (Lips et al. 2003; Sodhi et
al. 2004; Stuart et al. 2004).
Rafe Brown, Arvin Diesmos, and Angel Alcala
Anticipated in 4-5 years given current rate of
descriptions and number of currently known
new amphibian species
Today
Inger Brown and Alcala
1930
1940
1950
1960
1970
1980
1990
2000
2010
 Chapter 7. Amphibians of the Indomalayan Realm 83

References Sodhi, N.S., Koh, L.P., Brook, B.W. and Ng, P.K.L. 2004. Southeast Asian biodiversity: an Stuart, S.N., Chanson, J.S., Cox, N.A., Young, B.E., Rodrigues, A.S.L., Fischman, D.L. and
impending disaster. Trends in Ecology and Evolution 19:654-660. Waller, R.W. 2004. Status and trends of amphibian declines and extinctions worldwide.
Alcala, A.C. and Brown, W.C. 1999. Philippine frogs of the genus Platymantis (Amphibia: Science 306:1783-1786.
Ranidae). Philippine Journal of Science 128:281-287.
Brown, R.M. 2004. “Evolution of ecomorphological variation and acoustic diversity in Figure 2. Representatives of the species complexes defined by Brown et al. (1997), in part corresponding to the ecomorph classes defined by Brown (2004):
mate-recognition signals of Southeast Asian forest frogs (subfamily Platymantinae)”. P. indeprensus of the Platymantis dorsalis Species Group (ground frogs; A); P. isarog of the Platymantis hazelae Species Group (shrub frogs; B); P. banahao
Ph.D. Dissertation, University of Texas, Austin, Texas, USA. of the Platymantis guentheri Species Group (treefrogs; C); and an example of an enigmatic intermediate: Platymantis insulata (a terrestrial species, nested
Brown, R.M., Diesmos, A.C. and Alcala, A.C. 2002 (“2001”). The state of Philip- within the treefrog clade; D). All photos © Rafe Brown, 2006. Courtesy of HerpWatch Philippines
pine herpetology and the challenges for the next decade. Silliman Journal
42:18-87.
Brown, R.M. and Guttman, S.I. 2002. Phylogenetic systematic of the Rana signata complex
of Philippine and Bornean stream frogs; reconsideration of Huxley’s modification of
Wallace’s Line at the Oriental-Australian faunal zone interface. Biological Journal of
the Linnean Society 76:393-461.
Brown W.C., Brown, R.M. and Alcala, A.C. 1997. Species of the hazelae group of Platyman-
tis (Amphibia: Ranidae) from the Philippines, with descriptions of two new species.
Proceedings of the California Academy of Sciences 49:405-421.
Catibog-Sinha, C.S. and Heaney, L.R. 2006. Philippine biodiversity: Principles and
Practice. Haribon Foundation for Conservation of Natural Resources, Quezon City,
Philippines.
Diesmos, A.C., Brown, R.M., Alcala, A.C., Sison, R.V., Afuang, L.E. and Gee, G.V.A. 2002.
Philippine amphibians and reptiles. In: P.S. Ong, L.E. Afuang and R.G. Rosell-Ambal
(eds.), Philippine Biodiversity Conservation Priorities: a Second Iteration of the National
Biodiversity Strategy and Action Plan, pp.26-44. Department of the Environment and a. c.
Natural Resources–Protected Areas and Wildlife Bureau, Conservation International
Philippines, Biodiversity Conservation Program–University of the Philippines Center
for Integrative and Developmental Studies, and Foundation for the Philippine Environ-
ment. Quezon City, Philippines.
Evans, B.J., Brown, R.M., McGuire, J.A., Supriatna, J., Andayani, N., Diesmos, A.C.,
Iskandar, D., Melnick, D.J. and Cannatella, D.C. 2003. Phylogenetics of fanged frogs:
testing biogeographical hypotheses at the interface of the Asian and Australian faunal
zones. Systematic Biology 52:794-819.
Hanken, J. 1999. Why are there so many new amphibian species when amphibians are
declining? Trends in Ecology and Evolution 14:7-8.
Inger, R.F. 1954. Systematics and zoogeography of Philippine Amphibia. Fieldiana
33:181-531.
Köhler, J., Vieites, D.R., Bonett, R.M., García, F.H., Glaw, F. Skeinke, D. and Vences, M.
2005. New amphibians and global conservation: a boost in species discoveries in a
highly endangered vertebrate group. BioScience 55:693-696.
Lips, K.R., Reeve, J. and Witters, L.R., 2003. Ecological factors predicting amphibian
population declines in Central America. Conservation Biology 17:1078-1088.
b. d.

ESSAY 7.4. AMPHIBIAN SPECIES DISCOVERY IN MAINLAND SOUTHEAST ASIA

The global number of recognized amphibian species has risen dramatically over
the past two decades (Hanken 1999; Köhler et al. 2005), making amphibians
one of the animal groups with the highest proportional rate of description of
new species (Hanken 1999). This increase is primarily due to the discovery of
truly ‘novel’ forms by intensified scientific collecting in previously unexplored
parts of the world. Molecular genetic and bioacoustic tools in systematic
studies have also aided the discovery of ‘cryptic’ species that were previously
overlooked because they morphologically resemble other species.
Mainland Southeast Asia (defined in Figure 1) is no exception to this global
trend of rapid, recent discovery of new amphibian species. Beginning in 1834
with the description of Hoplobatrachus rugulosus from Hong Kong, a total of
299 currently recognized species of limbed amphibians (excluding caecilians)
were discovered and described from the region (Figure 1). Most strikingly, the
years 2005 and 2006 (through the time of writing in early November) each
yielded 16 new amphibian species, the highest number of annual descriptions
from the region since the advent of Linnean classification. The descriptions of
2005-2006 came from every country in the region, suggesting that the very re-
cent boost is not explained by scientific collecting in a localized ‘hotspot’. Prior
to the last few years, there have been three peaks of discovery in mainland
Southeast Asia, with 10-11 currently recognized species described in each
of the years 1937, 1962, and 1983 (Figure 1). Two of these peaks reflect the
significant contributions of 11 new species from Vietnam by Bourret (1937)
and seven new species from Guangxi Province, China, by Liu and Hu (1962),
while the third peak is primarily a coincidence of species descriptions from
southern China by a number of Chinese authors.
Every amphibian species described to date from mainland Southeast Asia
has been distinguished from its closest relatives on the basis of morphological
differences. However, a number of studies have used bioacoustic data (e.g.,
Kuramoto and Wang 1987; Wogan et al. 2003) or molecular genetic data (e.g.,
Li et al. 2001; Bain et al. 2003) to either discover a new species, or corroborate
The sociological nature of species discovery and description in the range species, each having much smaller geographic ranges. Conversely, newly
has changed over time. Historically, scientists working on amphibians in the discovered species are often prematurely labeled as endemic to a limited
region tended to work and publish alone or with very few colleagues. Today, geographic area only because scientists have just become aware of them. Al-
the process of discovering and describing species in the region involves col- though international cooperation among amphibian taxonomists is increasing,
laboration among scientists from within and outside of range countries. For many studies have been restricted by political boundaries. As a result, some
example, the average number of authors on species descriptions increased species have been described on opposite sides of borders under different
over time in peak years, with 1.0 in 1937, 1.73 in 1962, 2.0 in 1983, 2.38 names, and their known geographic distributions are limited by these sampling
in 2005, and 2.94 in 2006. The 2005-2006 boom in species descriptions of restrictions. These challenges indicate that much remains to be learned, and
amphibians from mainland Southeast Asia represented papers authored discovery efforts in the field and laboratory promise to rapidly improve our
by (alphabetically) American, Burmese, Cambodian, Canadian, Chinese, understanding of amphibian species diversity in the region.
French, German, Indian, Japanese, Russian, Thai, and Vietnamese authors.
The exchange of expertise and division of labour inherent in these growing Bryan L. Stuart and Raoul H. Bain
collaborations may partly explain the increasing productivity of amphibian
taxonomists working in the region.
Our current understanding of amphibian species diversity in the Philippines References
stands at 102 indigenous species and five or six introduced species (Brown et
al. 2002; Diesmos et al. 2002; Brown 2007). However, species descriptions of Bain, R.H., Lathrop, A., Murphy, R.W., Orlov, N.L. and Ho, C.T. 2003. Cryptic species of a
new frogs of the genus Platymantis currently in progress will soon increase cascade frog from Southeast Asia: taxonomic revisions and descriptions of six new
that number to around 140 taxa. Novel findings await discovery in the vast species. American Museum Novitates 3417:1-60.
areas that are still under-surveyed. A number of geographically widespread, Bourret, R. 1937. Notes herpétologiques sur l’Indochine française. XIV. Les batraciens
single species that are suspected to represent species complexes have not de la collection du Laboratoire des Sciences Naturelles de l’Université. Descriptions
yet been studied with molecular genetic or bioacoustic tools. Some of these de quinze espèces ou variétés nouvelles. Annexe au Bulletin Général de l’Instruction
widespread species may be found to contain multiple, morphologically cryptic Publique 4:5-56.
20
18
16
Number of species

14
the morphological distinctiveness of a new species. Every molecular genetic
12
study to date that has broadly sampled populations across the range of a
widespread frog species in mainland Southeast Asia has uncovered genetic 10

diversity that has been interpreted as unrecognized species diversity (Stuart 8

et al. 2006). These findings suggest that species diversity in the region re- 6
mains significantly underestimated. Molecular genetic and bioacoustic tools
4
are likely to play increasingly important roles in the process of discovering
amphibian species diversity in the region. 2

One of the most striking examples of recent Southeast Asian amphibian 0

species discoveries is found in the cascade frogs of the genus Odorrana. Of the
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1840
1845
1850
1855
1860
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30 species of Odorrana described from the region, 19 have been described since
2001. Many of these new Odorrana are morphologically very similar and have
been confused with other species for over a century (Bain et al. 2003; Stuart et al.
2006). Molecular genetic tools have shown that several morphologically similar,
but genetically distinct, lineages of Odorrana coexist in the same streams. The
ecological and behavioural mechanisms that these coexisting species use to
maintain their genetic distinctiveness are currently unknown.
Year of description
Figure 1. Annual amphibian species descriptions (n = 299 descriptions) from mainland Southeast Asia, defined as Vietnam, Laos, Cambodia, Thailand (north of
the Isthmus of Kra), Myanmar (north of the Isthmus of Kra), and southern China (Yunnan, Guangxi, Guangdong, Fujian, Hainan, Hong Kong and Taiwan). Data were
obtained from Frost (2006) and subsequently published literature, except that Hoplobatrachus rugulosus was not treated as a junior synonym of H. chinensis. Other
synonyms, species having the vague type locality of “China,” and caecilians (owing to poorly defined species boundaries) were not included.
84
Brown, R.M. 2007. Introduction, Robert F. Inger’s Systematics and Zoogeography of
Philippine Amphibia (Foreword to the 2007 reprint of Inger’s 1954 monograph). In:
Systematics and Zoogeography of Philippine Amphibia, pp. 1-17. Natural History
Publications, Kota Kinabalu.
Frost, D.R. 2006. Amphibian Species of the World: an Online Reference. Version 4 (17
August 2006). Electronic database accessible at http://research.amnh.org/herpetol-
ogy/amphibia/index.php. American Museum of Natural History, New York, USA.
Hanken, J. 1999. Why are there so many new amphibian species when amphibians are
ESSAY 7.5. AMPHIBIANS OF BORNEO
Borneo, the world’s third largest island, straddles the equator and until rela-
tively recently was entirely covered in tropical forests. Given this favorable
environment, it is not surprising that Borneo is one of the global hot spots
for frogs. No fewer than 148 species are now known from Borneo (Inger and
Stuebing 2005) and new species continue to be discovered. Most explorations
of this rich fauna have concentrated on the Malaysian portions of the island,
Sarawak and Sabah. Relatively little work has been carried out in Kalimantan,
which occupies roughly two-thirds of the land mass.
This rich fauna is diverse taxonomically and consists of species in six
families, most of which are also very rich in species in the adjacent land
masses of Sumatra and the Malay Peninsula. These frogs vary in size from
the tiny Microhyla perparva (NT) at 15mm to the giant Bufo juxtasper (LC) at
215mm in body length. The fauna is also ecologically diverse, demonstrat-
ing almost all the modes of life that frogs are capable of, from species that
burrow (e.g., Calluella smithi DD), to those that live mostly high in trees (e.g.,
Rhacophorus pardalis LC).
The breeding habits of the Bornean species also cover almost the entire
range of variation known for frogs (Inger and Tan 1996). Most species deposit
their eggs in the water of ponds or streams, but a few lay their eggs in water-
containing tree holes (e.g., Metaphrynella sundana LC). Some even lay their
eggs under leaf litter on the forest floor (e.g., Limnonectes palavanensis LC),
where they are guarded by the male who later transports the small tadpoles
on his back to a small rain pool on the forest floor. Tadpoles vary also in shape
and size (Inger 2005). Those that develop in ponds, such as the tadpoles of
Polypedates otilophus (LC), have fat, almost spherical bodies and are large
(up to about 65mm), while those that develop in streams tend to have more
slender bodies. The tadpoles of the Slender Litter Frogs (genus Leptolalax)
have very lithe bodies and wriggle into the crevices between the rocks lin-
ing the bottoms of the swift streams in which they live. Some tadpoles, for
example, those of the Torrent Frogs (genus Meristogenys), live in rapids and
have a large sucker on the underside that enables them to cling to rocks in the
strongest currents. There are also a few kinds of small tadpoles, like those of
the Sticky Frogs (genus Kalophrynus) that do not feed, but subsist through their
short developmental periods on the store of yolk in the eggs. These tadpoles
are usually found in very small, shallow pools of rain water, sometimes those
that form in rotting logs. There is also a group of species, the Bush Frogs
(genus Philautus) that do not have free-swimming tadpoles. These frogs have
very large (relative to their body size) eggs, rich in yolk, which are usually laid
under the moist leaf litter of the forest floor. The embryo develops within the
gelatinous envelope of the egg and hatches out as a tiny froglet.
From many points of view this is a rich, very diverse frog fauna. But,
because it is a tropical forest fauna, it is at risk because of forest conversion
and clearing, primarily due to logging, agriculture and mining. The entire forest
A calling male tree hole frog Metaphrynella sundana (Least Concern), with
the vocal sac inflated. Some tree holes may be as much as 10 metres or more
above the ground (©) Björn Lardner
Threatened Amphibians of the World
declining? Trends in Ecology & Evolution 14:7-8.
Köhler, J., Vieites, D.R., Bonett, R.M., Garcia, F.H., Glaw, F., Steinke, D. and Vences, M.
2005. New Amphibians and Global Conservation: A Boost in Species Discoveries in
a Highly Endangered Vertebrate Group. BioScience 55:693-696.
Kuramoto, M. and Wang, C.-S. 1987. A new rhacophorid treefrog from Taiwan, with
comparisons to Chirixalus eiffingeri (Anura, Rhacophoridae). Copeia 1987:931-942.
Li, C., Ye, C.-Y. and Fei, L. 2001. Taxonomic studies of Odorrana andersonii in China (Anura:
Ranidae). Acta Zootaxonomica Sinica 26:234-238.
A Bornean endemic, Ansonia spinulifer (Near Threatened), is a characteristic
species reported from lowland localities throughout Borneo from 150-750 m
asl. © Djoko Iskandar
fauna depends on the very high humidities and moderate temperatures cre-
ated by the closed canopy of the forest. Nearly one-third of the Bornean frog
species (29%) – all but one of them known only from Borneo – are listed as
threatened on the IUCN Red List, and all because of serious habitat modifica-
tion. Although economic development poses a hazard to all of these rainforest
frogs, it is clear that certain ecological or behavioral characteristics put some
species at particular risk. The frog fauna of Borneo is roughly equally divided
between those species that breed in streams (61 of 148 species) and those
that breed in small bodies of standing water (64 species). However, when one
considers only the threatened species, then those that breed in streams (19 of
42 species) outnumber those that that breed in standing water (10 species).
The majority (14 species) of these threatened stream-breeding amphibians
have tadpoles specialized for living in strong, clear currents.
One of the immediate consequences of selective logging, which harvests
only the largest trees, is stream siltation, which results in the accumulation
of a fine layer of silt covering the stream bottom. Forest clearing results in an
even deeper layer of silt. This silt clogs the bottom crevices in which some
kinds of tadpoles live (e.g., tadpoles of the genus Leptolalax) and prevents
the growth of rock-clinging algae on which other kinds of tadpoles feed
(e.g., tadpoles of the Torrent Frogs, Meristogenys). The result is a sharp,
Limnonectes rhacoda (Near Threatened) is a lowland rainforest species currently known only from Indonesian Borneo. © Djoko Iskandar
Liu, C.-C. and Hu, S.-Q. 1962. A herpetological report of Kwangsi. Acta Zoologica Sinica
14(Suppl.):73-104.
Stuart, B.L., Inger, R.F. and Voris, H.K. 2006. High level of cryptic species diversity revealed
by sympatric lineages of Southeast Asian forest frogs. Biology Letters 2:470-474.
Wogan, G.O.U., Htun Win, Thin Thin, Kyi Soe Lwin, Awan Khwi Shein, Sai Wunna Kyi,
and Hla Tun. 2003. A new species of Bufo (Anura: Bufonidae) from Myanmar (Burma),
and redescription of the little-known species Bufo stuarti Smith 1929. Proceedings of
the California Academy of Sciences 54:141-153.
rapid decline in the populations of these two groups of species that breed in
streams having clear water and gravel or rock bottoms.
Opening of the forest exposes the leaf litter in which many frog species live
to much higher temperatures and lower humidity. The Bush Frogs (Philautus)
place their fertilized eggs under dead leaves, which in undisturbed forests,
remain continually moist. As soon as the forest is opened, more sunlight
reaches the floor raising the temperature and drying out the floor litter. Ten
of the 16 Bornean species of Philautus are considered to be threatened.
While there has been no direct study of the impact of high temperature and
low humidity on litter-dwelling frogs, the failure to find these species in
open areas, such as surrounding agricultural fields, suggests a direct relation
between forest clearance and disappearance of these species.
Yet some species manage to survive in secondary forests, those forests
from which some trees have been removed. Large areas cleared of forest in
Borneo have been converted to non-native tree plantations, mostly oil palm,
but also Acacia mangium. These tree plantations have some of the physical
characteristics of natural forest, such as a closed canopy, reduction in sunlight
reaching the floor, and high humidity. If these plantations are adjacent to for-
est, even secondary forest, it is possible that some rain forest frogs may move
into and survive in these environments. In fact, an as yet incomplete survey
of the frogs living in Acacia plantings in Sarawak has discovered species of
frogs characteristic of rain forests in these exotic environments. At least one
threatened floor-dwelling species, Kalophrynus intermedius (VU), has been
recorded within an Acacia planting. These particular plantations are adjacent
to secondary forests, and this secondary forest is probably the source of the
Acacia-dwelling frogs. It may also be significant that pesticides have not
been broadcast in these plantings, and more investigation of the fauna of
tree plantations is needed to determine what portion of the fauna can adjust
to living in that kind of exotic environment, and for how long populations can
persist. Until such study is completed, we may not know how grim the future
is for this interesting, unique amphibian fauna.
Robert F. Inger
References
Inger, R.F. 2005. The systematics and zoogeography of the Amphibia of Borneo. Natural
History Publications, Kota Kinabalu (reprint), Malaysia.
Inger, R.F. and Stuebing, R.B. 2005. A field guide to the frogs of Borneo. 2nd edn. Natural
History Publications, Kota Kinabalu, Malaysia.
Inger, R.F. and Tan, F.-L. 1996. The natural history of amphibians and reptiles in Sabah.
Natural History Publications, Kota Kinabalu, Malaysia.
 CHAPTER 8. AMPHIBIANS OF THE NEARCTIC REALM

Don Church, David M. Green, Geoffrey Hammerson, Joseph Mitchell, contains 4% of all globally threatened amphibians. It accounts for only 2% of CR species Figure 1. Summary of Red List categories
Gabriela Parra Olea and Georgina Santos Barrera and 3% of the EN species, but 7% of the VU species. Hence, threatened Nearctic amphib- for amphibians in the Nearctic Realm. The
ians are more likely to be in a lower category of threat, when compared with the global percentage of species in each category is
distribution of threatened species amongst categories. The percentage of DD species, 7% also given.
THE GEOGRAPHIC AND HUMAN CONTEXT (25 species), is also less than the global average of 23%. This result is not surprising for
such a well-surveyed region.
The Nearctic Realm includes most of mainland North America, south to central Mexico Two of the world’s 34 known amphibian extinctions have occurred in this region, namely Red List Category Number of species
(excluding the Caribbean coast of Mexico, and extreme southern Florida in the United the Vegas Valley Leopard Frog Rana fisheri and the poorly known Catahoula Salamander Extinct (EX) 2
States). North America currently has a land connection to South America that was formed Plethodon ainsworthi in Mississippi. In addition, the only amphibian in the world currently Extinct in the Wild (EW) 1
relatively recently, beginning in the late Pliocene, some 3.5 Ma. Even earlier in the Perm- listed as Extinct in the Wild with only captive populations in existence, the Wyoming Toad Critically Endangered (CR) 10
ian to early Jurassic Periods, some 285-190 Ma, North America had land connections with Bufo baxteri, is also found here. Three Critically Endangered species in the Nearctic Realm Endangered (EN) 24
Eurasia and Africa. Biogeographic relationships between these regions are seen today, for are also considered possibly extinct: Tlaloc’s Leopard Frog Rana tlaloci from the Mexican Vulnerable (VU) 46
example, in the presence of giant salamanders in the Appalachians and in China and Japan. Plateau, the Bigfoot Splayfoot Salamander Chiropterotriton magnipes2 from San Luis Potosi Near Threatened (NT) 40
The Appalachian Mountains in the eastern United States began to uplift some 680 Ma, and Queretaro, Mexico, and the Cave Splayfoot Salamander Chiropterotriton mosaueri from Least Concern (LC) 189
when the European and African plates pressed against the east side of the North American northern Hidalgo, Mexico (the first two of these also occur in the Neotropical Region). These Data Deficient (DD) 25
plate, long before the time when the first amphibians crawled or hopped. These mountains taxa represent 2.3% of the 130 possibly extinct species in the world. There is little in the Total Number of Species 337
had eroded almost entirely by the Mesozoic Era, but further uplift during the Cenozoic Era way of overall discernible pattern in Nearctic amphibian extinctions thus far. 1% 3%
7%
and subsequent erosion by newly formed rivers and streams shaped the Appalachians as 7%
we know them today. The moist temperate forests that now cover these mountains grow
on severely eroded remnants of a formerly towering cordillera that was putatively higher SPECIES RICHNESS AND ENDEMISM 14%
than the Himalaya.
The western part of the region is much more mountainous and includes the Rocky Species Richness and Endemism Across Taxa
Mountains, the Cascade Range, the Alaskan Range, and many others. The topographical
and geological histories of this region are complex. Three major mountain-building episodes The 337 native amphibian species in the Nearctic Realm represent 6% of the currently
restructured the topography of the west during the period of about 170 to 40 Ma (Jurassic known global total of 5,915 species. Of these 337 species, 230 (or 68%) are endemic to the
12%
to Cenozoic Periods). Orogenous uplift continues today in the Cascades and other western- Nearctic (Table 1). Both salamanders and frogs occur in the region, but no caecilians. Unlike
most ranges as seen in modern eruptions of mountain peaks like Mount Saint Helens. The the situations in all other major biogeographic realms, there are more species of salamanders 56%
Rockies and the Sierra Madre Mountains in Mexico date from uplifts that occurred during (Caudata) than frogs and toads (Anura), the former accounting for 58% of the species in
the Laramide orogeny some 70-40 Ma in the Palaeocene and Eocene Epochs. The highest the Nearctic. Endemism is also much higher in the salamanders (89%), as compared with
point in the Nearctic Realm is Mount McKinley in Alaska at 6,194m. frogs and toads (39%). This presumably reflects a more fine-grained pattern of isolation
During the more recent geological past, as in the Pleistocene, sea levels rose and fell and speciation, and accordingly generally much smaller range sizes, among salamanders
during several glacial-interglacial episodes. The eastern coastline of North America during in the Appalachian Mountains in the east and Sierra Nevada Mountains in the west. Of the
the last major glacial event (that ended about 10,000 years ago) was as far as 100km east of 17 families that are native to the region, only three of them are endemic (Table 1). Only 10
its current location and -130m to -180m below current sea level. The Coastal Plain terraces species (3%) are members of these endemic families, but this low percentage is a reflection
are the result of previous sea level rise and fall events. Glacial and interglacial episodes in of the fact that several families occur only marginally in other regions.
the Pleistocene had dramatic effects on the flora and fauna in most of North America with Under current climatic conditions, there is less isolation between the Nearctic and
major latitudinal and elevational shifts in ranges of entire communities and ecosystems (Flint Neotropical Realms than there is between the Afrotropical and Palaearctic Realms, and
1971). Many of the current distributions of amphibians in the Nearctic are the result of these there are points of contact between the two faunas along the Caribbean coast of Mexico
historical events. Many amphibians that now range northward to New York or nearly so, for and in Florida (although the transvolcanic belt in central Mexico does form a barrier to faunal
example, expanded their ranges up the Coastal Plain following glacial retreat. dispersal). The result of this indistinct boundary is to reduce the level of endemism of both
Boreal coniferous forests dominate the northern part of the region through Alaska and regions. The families Scaphiopodidae, Ambystomatidae, Amphiumidae, and Sirenidae are
Canada, although high-elevation mountains such as in the southern Appalachians and the nearly endemic to the Nearctic, and Leptodactylidae and Rhinophrynidae are almost endemic
Rocky Mountains also support such forests. The eastern part of the region was originally to the Neotropics. Proteidae is also overwhelmingly Nearctic with just one species in the
covered with mixed and broad-leaved forests after the Pleistocene glaciations, giving way Palearctic. Duellman and Sweet (1999), Green (1997) and Lannoo (2005) provide summaries
to vast grasslands in the central part of the continent east of the Rockies. The south-eastern of the amphibian fauna of the Nearctic.
portion of the Nearctic Realm was dominated by pine forests after the last glaciation some Of the 51 genera (11% of the global total) that occur in the region, 22 (33%) are also The Lesser Siren Siren intermedia (Least
10,000 years ago. In keeping with the topography, habitats are much more complex in the endemic. Endemism at the generic level is much higher among the salamanders (67%) than Concern) is one of the four species of siren in
west, ranging from hot deserts, to sagebrush, Mediterranean-type chaparral, montane it is among frogs and toads (4.2%), a pattern mirrored in the Palaearctic. The most speciose the family Sirenidae, a group of highly aquatic
coniferous forests, alpine tundra, and extremely wet temperate rainforests in the Pacific genera in the region are Plethodon (48 species), Rana (38 species), Bufo (28 species), Eurycea salamanders with small front legs and no back
Northwest. The Mexican portion of the Nearctic Realm supports desert and desert grassland (24 species), and Ambystoma (21 species). At the opposite end of the spectrum, there are legs. It ranges from the south-eastern United
between the Sierra Madre Oriental and the Sierra Madre Occidental. High mountains in nine monotypic genera endemic to the Nearctic Realm, eight of which are salamanders States to north-eastern Mexico, and lives
these regions support xeric oak and pine vegetation. Descriptions of biotic communities, and seven of which are in the family Plethodontidae. The 32 non-endemic genera in the in shallow water with abundant vegetation
vegetation, and ecoregions in the Nearctic Realm are in Scott (1995), Brown et al. (1998), Nearctic include 23 frog genera (13 genera in Hylidae, three in Leptodactylidae, two each in (burrowing into the bottom mud if the water
Ricketts et al. (1999), and Barbour and Billings (2000), among others. Microhylidae and Scaphiopodidae, and one each in Bufonidae, Ranidae and Rhinophrynidae) dries up). © Michael Graziano
Compared with other parts of the world, the Nearctic Realm has a low human population and nine salamander genera (four in Plethodontidae, two in Sirenidae, and one each in
density (approximately 15 people per square km in 2005), only 23% of which lives in rural Ambystomatidae, Amphiumidae, and Salamandridae). These non-endemics include the
areas, and a low population growth rate (1.2% per annum), which is gradually decreasing. widespread genera Bufo and Rana.
The human population is distributed very unevenly, with high concentrations in southern Table 1. The number of Nearctic amphibians in
Canada, on the Coastal Plain and Piedmont in the eastern and north-eastern United States, each taxonomic Family present in the region.
and in California and Florida. Gross income per capita in the United States and Canada was
over US$38,000 in 2004, but the southern part of the region in northern Mexico is poorer,
with a gross income per capita of less than US$10,000 in 2004. The high wealth of much Family Native species Percentage of Percentage of Native genera Percentage of Percentage of
of the region helps to explain why damage to natural ecosystems has been extensive in (endemics to species in region species in family (endemics genera in region genera in family
region) that are endemic that are endemic to region) that are endemic that are endemic
relation to the low human population density; due to large-scale agriculture, urbanization, to region to region
roads, wetland loss. Economic growth rates in the region have been relatively high (3.5%
in 2005), especially for a wealthy region. Anura
The human impact on Nearctic ecosystems varies greatly through the region. There Ascaphidae 2 (2) 100 100 1 (1) 100 100
have been high levels of anthropogenic disturbance over the last 300 years, with very Bufonidae 28 (12) 43 3 1 (0) 0 0
extensive forest loss in the eastern and southern United States and conversion of the Hylidae 39 (14) 36 2 13 (0) 0 0
formerly extensive grasslands to commercial agriculture in the centre of the country. Over Leptodactylidae 23 (3) 13 0.2 3 (0) 0 0
the last century, there has been some forest regeneration in parts of the east, although this Microhylidae 4 (0) 0 0 2 (0) 0 0
regrowth has recently been targeted for much timber harvesting. Damage to ecosystems Ranidae 38 (21) 55 3 1 (0) 0 0
has been as severe in the drier west due to timbering and cattle grazing. The coniferous Rhinophrynidae 1 (0) 0 0 1 (0) 0 0
stands in the northern boreal zone were relatively untouched until recently. Urbanization and Scaphiopodidae 7 (4) 57 57 2 (0) 0 0
suburbanization are expanding rapidly in all cities and towns and contributes substantially TOTAL ANURA 142 (56) 39 1 24 (1) 4 0.3
to habitat loss. Pollution of rivers, streams and lakes is a problem in many areas of the Caudata
Nearctic, and approximately 40% of these in the United States are classified by the United Ambystomatidae 21 (16) 76 53 1 (0) 0 0
States Environmental Protection agency as polluted. Freshwater wetland loss has been high Amphiumidae 3 (2) 67 67 1 (0) 0 0
(Fretwell et al. 1996) because of land conversion to urbanization and agriculture. Overviews Cryptobranchidae 1 (1) 100 33 1 (1) 100 50
of threats to and losses of animals, habitats, and ecosytems in North America are in LaRoe Dicamptodontidae 4 (4) 100 100 1 (1) 100 100
et al. (1995) and Noss et al. (1995). Plethodontidae 147 (136) 93 37 17 (13) 76 45
Proteidae 5 (5) 100 83 1 (1) 100 50
Rhyacotritonidae 4 (4) 100 100 1 (1) 100 100
GLOBAL CONSERVATION STATUS Salamandridae 6 (4) 67 6 2 (1) 50 6
Sirenidae 4 (2) 50 50 2 (0) 0 0
A total of 83 species (24%) of the 337 amphibian species in the Nearctic Realm is consid- TOTAL CAUDATA 195 (174) 89 33 27 (18) 67 29
ered to be globally threatened or Extinct (Figure 1). This is significantly less than the global TOTAL ALL AMPHIBIANS 337 (230) 68 4 51 (19) 37 4
average of 33%1, but very similar to the situation in the Palaearctic. The Nearctic Realm
86 Threatened Amphibians of the World

The Marbled Salamander Ambystoma opacum Figure 2. The species richness of amphibians in the Nearctic Realm, with darker colours
(Least Concern) occurs in the eastern United As noted already, 17 of the world’s 48 amphibian families (35%) occur in the Nearctic, with corresponding to regions of higher richness. Colour scale based on 10 quantile classes;
States in various wooded habitats, in the vicin- three endemic: Ascaphidae, Dicamptodontidae and Rhyacotritonidae3. The characteristics maximum richness equals 52 species.
ity of swamps and vernal pools. For most of of these families are provided in Chapter 1.
the year the adults are subterranean in rodent Among the non-endemic families, the majority of Nearctic species are in the Bufonidae,
tunnels, but in the autumn they congregate, Hylidae, Leptodactylidae, Ranidae, Ambystomatidae and Plethodontidae. Of the Nearctic The members of all four families are mainly large salamanders that are completely aquatic
sometimes in large numbers as shown here, to Bufonidae, all 28 species are in the widespread genus Bufo4. This family occurs through most as both larvae and adults.
lay eggs in forest depressions such as vernal of the region, and 16 species (57%) are shared with the Neotropics. All Nearctic species The Rhyacotritonidae (torrent salamanders) and Dicamptodontidae (Pacific giant
pool basins. © Don Church have free-living aquatic larvae. salamanders) are wholly restricted to the Pacific north-west. Members of both families are
There are 39 species of Hylidae in the Nearctic currently placed in 13 genera (Faivovich inhabitants of coldwater mountain streams.
et al. 2005). This family is also very widespread within the region; 25 species (64%) range The small frog family Scaphiopodidae (American spadefoots), with just seven species,
into the Neotropics. Within this region, these species include treefrogs, chorus frogs and occurs mainly in the Nearctic, but three of the species also range into the Neotropics. The
cricket frogs, and are associated with many different habitats. All Nearctic species have remaining families include the Microhylidae (globally widespread, with just four species
free-living aquatic larvae. in the region, none of which are endemic), the primitive Rhinophrynidae (which marginally
Although the family Leptodactylidae is almost endemic to the Neotropics, 23 species ranges into the region), and the Salamandridae (mainly a Palaearctic family, with six spe-
occur in the Nearctic in Mexico, and some range into the southern United States. These cies in the region).
include representatives of the genera Eleutherodactylus and Craugastor, which breed by The threatened and extinct species in the Nearctic are overwhelmingly in the Plethod-
direct development on land, and Leptodactylus, which have free-living aquatic larvae.5 ontidae (46, 84% of the total in the region; Table 2). This is a reflection of the many species
All of the Nearctic Ranidae are in the genus Rana6, The family is very widespread in with small ranges that occur in mountainous regions (especially in the Appalachians and in
the region, occurring in most aquatic and wetland habitats, and all members in the region Mexico), where they are threatened in many places by forest loss. Nearly one-third (31%) of
have free-living aquatic larvae. the lungless salamanders in the region are globally threatened or extinct. There is also a high
The mole salamanders (Ambystomatidae) are mainly a Nearctic family, with 21 species percentage of threatened species (29%) among the mole salamanders (Ambystomatidae),
(70% of the global total) occurring in the region, 16 of which are endemic. The species are which are also threatened mainly by habitat loss.
mainly subterranean as adults and juveniles, with aquatic larvae that develop mainly in Most threatened frog species occur in the larger families, namely Bufonidae, Lepto-
temporary pools and occasionally in streams. dactylidae, and Ranidae (Table 2). The threatened toads (Bufonidae) occur mainly in the
The Plethodontidae (lungless salamanders) is by far the largest family in the Nearctic, western United States, where they have been impacted by habitat degradation, and in
with 147 species (40% of the global total), of which 136 are endemic. The family includes some cases by introduced bullfrogs (Rana catesbeiana) and the disease chytridiomycosis.
The Wood Frog Rana sylvatica (Least Con- both direct-developing species, which breed terrestrially with no association with water The eight threatened species of Leptodactylidae are all Mexican endemics threatened by
cern) ranges farther north than any other (for example, Plethodon, Aneides, and Batrachoseps), and aquatic larval developing species, habitat loss (and perhaps, in a few cases, by chytridiomycosis). The threatened Nearctic
North American amphibian. It is abundant typically associated with streams (for example, Desmognathus and Eurycea). Although the species in the Ranidae range widely in Mexico and the western and southern United
and widespread, and occurs in various kinds family is widespread through much of the region (but avoiding highest latitudes), it reaches States. As with the other families, they are impacted by habitat loss and chytridiomycosis,
of forest and woodland habitats, and also extraordinary diversity in the southern Appalachians (see Essay 8.1). Most species in this but invasive species and water pollution have also been detrimental. It is noteworthy that
willow thickets and grass/willow/aspen family are strongly tied to forest habitats (though some, especially in the genus Eurycea, the percentage of threatened treefrogs (Hylidae) in the region is much lower than for the
associations. The eggs are laid, and larvae live in caves). other large families.
develop, usually in small fish-free ponds As home to four small salamander families that occur mainly in the eastern United States, The great majority (88%) of the threatened amphibians in the Nearctic are in the Endan-
in wooded or, sometimes, open areas. the Nearctic contributes uniquely to global amphibian diversity. Both the Amphiumidae gered and Vulnerable categories. The generally low number of Critically Endangered species
© Don Church (amphiumas) and Sirenidae (sirens) occur largely on the Coastal Plain of the eastern and masks some important family-level differences, with 14% of the Nearctic Ambystomatidae,
southern United States (ranging only marginally into the Neotropics). The Cryptobranchidae and 11% of the Nearctic Ranidae falling into this category.
(giant salamanders) has one species in the Nearctic (the Hellbender Cryptobranchus alle-
Table 2. The number of species within each ganiensis; NT); two others are in East Asia. The Proteidae (mudpuppies and waterdogs) has Geographic Patterns of Species Richness
IUCN Red List Category in each Family and five species in the Nearctic (all in the eastern United States), with one species in Europe. and Endemism
Order in the Nearctic Realm. Introduced spe-
cies are not included. Overall species richness of amphibians in the Nearctic Realm (Figure 2) is highest in the
southeastern United States, especially on the Coastal Plain and in the Appalachian Moun-
tains. Species richness is low in the drier parts of northern Mexico, through much of the
Family EX EW CR EN VU NT LC DD Total number Number Threatened % Threatened central and western United States (although there are more species in moist areas along
of species or Extinct or Extinct the northwest Pacific coast), and in Canada. The Nearctic Realm is well studied, and Figure
Anura 2 probably reflects genuine patterns of amphibian species richness.
Ascaphidae 0 0 0 0 0 0 2 0 2 0 0 Concentrations of threatened species (Figure 3a) in the Nearctic are less marked than in
Bufonidae 0 1 0 4 1 2 20 0 28 6 21 many tropical regions. Nevertheless, there are some notable concentrations in the southern
Hylidae 0 0 0 2 1 2 34 0 39 3 8 Appalachians (centred on the Great Smoky Mountains), the mountains of northern Mexico,
Leptodactylidae 0 0 1 2 5 1 9 5 23 8 35 along the Edwards Plateau in Texas, and in the mountains of California. These geographic
Microhylidae 0 0 0 0 0 0 4 0 4 0 0 concentrations generally reflect the topographically diverse montane parts of the region where
Ranidae 1 0 4 1 5 5 20 2 38 11 29 amphibians have naturally small ranges and where habitat destruction is ongoing. The Texan
Rhinophrynidae 0 0 0 0 0 0 1 0 1 0 0 concentration largely reflects subterranean salamanders in the genus Eurycea and blind spe-
Scaphiopodidae 0 0 0 0 0 1 6 0 7 0 0 cies in the genus Typhlomolge. Not surprisingly, given the small number of species involved,
TOTAL ANURA 1 1 5 9 12 11 96 7 142 28 20 there are no noteworthy concentrations of Critically Endangered species in the region, with
Caudata the exception perhaps of the California floristic region (see Essay 8.2) (Figure 3b).
Ambystomatidae 0 0 3 1 2 1 13 1 21 6 29
Amphiumidae 0 0 0 0 0 1 2 0 3 0 0 Species Richness and Endemism within Countries
Cryptobranchidae 0 0 0 0 0 1 0 0 1 0 0
Dicamptodontidae 0 0 0 0 0 1 3 0 4 0 0 Although Mexico as a whole has more amphibian species than the United States, within
Plethodontidae 1 0 2 12 31 21 63 17 147 46 31 the Nearctic Realm the United States has more than double the number of species (261
Proteidae 0 0 0 1 0 1 3 0 5 1 20 species vs. 116) (Figure 4). Species richness in Canada is low, with just 45 species.
Rhyacotritonidae 0 0 0 0 1 2 1 0 4 1 25 The amphibian fauna in Mexico is less well known than that of either the United
Salamandridae 0 0 0 1 0 1 4 0 6 1 17 States or Canada, although the Nearctic parts of Mexico are probably better surveyed
Sirenidae 0 0 0 0 0 0 4 0 4 0 0 than the more diverse Neotropical areas to the south. The Mexican amphibian fauna
TOTAL CAUDATA 1 0 5 15 34 29 93 18 195 55 28 is not well summarized in review literature other than in Flores-Villela (1993) and
TOTAL ALL AMPHIBIANS 2 1 10 24 46 40 189 25 337 83 24 Flores-Villela et al. (1995). However, there are some recent regional publications from
northern Mexico (Lemos-Espinal et al. 2004; Vazquez-Díaz and Quintero-Díaz 2005). The
 Chapter 8. Amphibians of the Nearctic Realm 87

Figure 3. a) The richness of threatened am-

phibians in the Nearctic Realm, with darker
colours corresponding to regions of higher
richness. Maximum richness equals four spe-
cies. b) The richness of CR amphibians in the
Nearctic Realm. Maximum richness equals
one species.

a. b.

United States United States United States Mexico*

of America* of America* of America*

United States
Mexico* Mexico* Mexico* of America*

Canada Canada Canada Canada

0 50 100 150 200 250 300 0 20 40 60 80 0 10 20 30 40 50 60 0 5 10 15 20 25 30

Number of species Percentage of endemic species Number of threatened species Percentage of species threatened

Figure 4 Endemics Non-endemics Figure 5 Figure 6 Endemics Non-endemics Figure 7

Figure 4. The number of amphibians present

amphibians of the United States and Canada are amongst the best known in the world, HABITAT AND ECOLOGY in and endemic to each Nearctic country.
with a very extensive review literature and many books, including Green (1997), Harding *denotes countries not entirely within the
(1997), Conant and Collins (1998), Petranka (1998), Stebbins (2003), and Lannoo (1998, Habitat Preferences Nearctic Realm, hence only the species whose
2005). Summaries of the amphibian fauna of the Nearctic are provided by Duellman ranges fall within the region are included.
and Sweet (1999) and Lannoo (2005). There are numerous State and Province level Most Nearctic amphibians (79%) occur in forests; only 17% of which can survive in secondary
summaries of the amphibian fauna, such as Stebbins (1972), Mount (1975), Martof terrestrial habitats (Table 3; Figure 8). As in other regions, forest-dwelling and stream-as-
et al. (1980), Nussbaum et al. (1983), Green and Pauley (1987), Dundee and Rossman sociated amphibians are more likely to be threatened than are those occurring in any other Figure 5. Percentage of species endemic to
(1989), Klemens (1993), Leonard et al. (1993), Russell and Bauer (1993), Degenhardt et habitats, with nearly 20% of species being globally threatened. This combination of habitat each Nearctic country. *denotes countries
al. (1996), Weller and Green (1997), Bartlett and Bartlett (1999), Hammerson (1999), preferences is associated with rapid declines in amphibian populations worldwide (Stuart not entirely within the Nearctic Realm, hence
Hunter et al. (1999), Dixon (2000), Johnson (2000), Hulse et al. (2001), Minton (2001), et al. 2004). Amphibians occurring in savannahs, marshes and swamps, arid and semi-arid only the species whose ranges fall within the
MacCulloch (2002), Dodd (2004), Trauth et al. (2004), Werner et al. (2004), Corkran and habitats, and secondary terrestrial habitats are less likely to be threatened than those region are included.
Thoms (2006) and Matsuda et al. (2006). occurring in these other habitats (Table 3; Figure 8).
The United States has far more endemic Nearctic species (182) than the other two
countries in the Nearctic (Figure 4). Mexico has 68 endemic Nearctic species, but there Reproductive Modes Figure 6. The number of threatened amphib-
are no endemic amphibians in Canada. In the United States, 70% of the Nearctic species ians present in and endemic to each Nearctic
are endemic, compared with 59% in Mexico (Figure 5). Larval development is the most common reproductive mode in the Nearctic (65% of species), country. *denotes countries not entirely
Only one globally threatened amphibian species occurs in Canada, compared with 31 compared with 35% for direct development and no live-bearing species (Table 4; this compares within the Nearctic Realm, hence only the
in Mexico and 52 in the United States (Figure 6). The percentage of threatened amphibian with the global picture of 68% larval development, 30% direct development, and 1% live-bearing). species whose ranges fall within the region
species is highest in Mexico (27%) compared with the United States (20%) (Figure 7). In The Nearctic amphibians clearly have a larger proportion of direct-developing species than the are included.
both cases, this level of threat is lower than the global average of 33%. global average. Most of the direct-developing Nearctic amphibians are plethodontid salamanders,
Assessments of the conservation status of Nearctic amphibians at the national level especially in the genera Plethodon (48 species), Batrachoseps (19), Chiropterotriton (10), and Pseu-
have been carried out by NatureServe (e.g., Stein et al. 2000), since updated as part of the doeurycea and Aneides (both 6). The other concentration of direct-developing frogs in the region is Figure 7. Percentage of native species
Global Amphibian Assessment (Young et al. 2004). There are also official governmental among the leptodactylid frogs in the genus Eleutherodactylus (12 species) and Craugastor (9). that are threatened. *denotes countries not
threatened species listing processes in Canada (http://www.cosewic.gc.ca/index.htm) and In the Nearctic, the percentage of globally threatened or Extinct direct-developing entirely within the Nearctic Realm, hence
in the United States (http://www.fws.gov/Endangered/esa.html). An official Mexican red amphibians is much higher than for larval-developing species (33% as apposed to 20%). only the species whose ranges fall within the
list of amphibians is in preparation (the draft is on http://www.semarnat.gob.mx/leyesyn- As in other regions, this is probably because direct-developing species have smaller ranges region are included.
ormas/Normas%20Oficiales%20Mexicanas%20vigentes/NOM-ECOL-059-2001.pdf). on average, and are therefore more seriously affected by habitat loss.
The proportion of Critically Endangered Nearctic species is highest in Mexico (6%),
being 1% in the United States, and zero in Canada. This reflects the generally poor
conservation situation for amphibians in Mexico (see Neotropical section). However, MAJOR THREATS
the only Extinct and Extinct in the Wild species in the region are in the United States,
although, as mentioned above, two of the Critically Endangered species in Mexico are As in other parts of the world, habitat loss is overwhelmingly the major threat to amphibians
possibly extinct. in the Nearctic Realm (Table 5; Figure 9), affecting nearly 80% of the threatened species.

Habitat type Number % of all Threatened % of species

Number of species % of species in habitat that are threatened
of species in species and Extinct occurring
each habitat occurring species in habitat that Forest
in the habitat are Threatened Savannah
or Extinct
Grassland
Forest 266 79 51 19
Shrubland Table 3. The habitat preferences of amphib-
Savannah 10 3 0 0 ians in the Nearctic Realm.
Grassland 70 21 10 14 Secondary terrestrial habitats
Shrubland 77 23 8 10 Flowing freshwater
Secondary terrestrial habitats 56 17 5 9
Marsh/swamp Figure 8. The habitat preferences of Nearctic
Flowing freshwater 169 50 32 19 amphibians. The plot on the left-hand side
Still open freshwater
Marsh/swamp 87 26 8 9 shows the number of species in the region
Still open freshwater 141 42 16 11 Arid and semi-arid habitats in each habitat type. On the right-hand side,
Arid and semi-arid habitats 21 6 2 10 the percentage of these species which are
0 50 100 150 200 250 0 5 10 15 20
threatened is given.
88
The Columbia Torrent Salamander Rhyacotri-
ton kezeri (Near Threatened) is restricted to a
small area in the Coastal Ranges of Oregon
and Washington in the United States. It is
one of only four torrent salamander species in
the Family Rhyacotritonidae, which is endemic
to the Pacific Northwest. All the species are
strictly associated with streams in forest.
© Henk Wallays
Couch’s Spadefoot Scaphiopus couchii (Least
Concern) occurs in northern Mexico and the
south-western United States in arid and semi-
arid shrublands, short grass plains, mesquite
savannah, creosote bush desert and cultivated
areas. It burrows underground or occupies
rodent burrows when inactive, but emerges to
breed explosively when temporary pools are
formed by the rain. © Twan Leenders
Threatened Amphibians of the World
Pollution is the next most serious threat, affecting one-third of the threatened species.
Reproductive mode All species Threatened or % Threatened Invasive and introduced species, disease, human disturbance, and fire also have significant
Extinct species or Extinct impacts on certain species. Most other threats are of relatively minor importance. Although
Direct development 118 39 35 chytridiomycosis has been recorded widely in the region (e.g., Fellers et al. 2001; Bradley et
Larval development 219 44 65 al. 2002), and is associated with some serious declines, its overall impacts still seem to be
Live-bearing 0 0 - less than in the Neotropics. However, there have been some reports of crashes in amphibian
Not known 0 0 - populations in North America (e.g., Kagarise Sherman and Morton 1993; Drost and Fellers
1996) that could well be attributable to chytridiomycosis. Ongoing research may show that
this fungal disease is a greater threat than current documentation indicates.
Table 4. Nearctic amphibians categorized by reproductive mode. Of the types of habitat loss impacting amphibians in the Nearctic, the impacts of veg-
etation removal, mainly via logging (which affects nearly half of the threatened species),
and urbanization and industrial development (nearly 40%) are the most severe, followed
by expanding croplands (nearly 30%) (Table 5). Tree plantations and livestock are less
Threat type Threatened species % Threatened species important threats in most cases.
All habitat loss 63 79 A total of 31 species (only nine of which are threatened) are recorded as being used
Agriculture – Crops 23 29 for some or other purpose in the region (Table 6). The most common reasons for harvesting
Agriculture – Tree plantations 12 15 Nearctic amphibians are for the pet trade and human consumption. Much of the harvesting
Agriculture – Livestock 11 14 of amphibians in the region is not considered to constitute a major threat to the species.
Timber and other vegetation removal 39 49 Of the 31 species being harvested, utilization is considered to be a threat for just 14 (of
Urbanization and industrial development 31 39 which only six are threatened species for which harvesting is believed to be contribut-
Invasive species 17 21 ing to deterioration in their status, most prominently the Critically Endangered Axolotl
Utilization 6 8 Ambystoma mexicanum).
Accidental mortality 6 8
Persecution 1 1
Pollution 27 34 POPULATION STATUS AND TRENDS
Natural disasters 9 11
Disease 14 18 Estimates of Population Trends
Human disturbance 12 15
Changes in native species dynamics 1 1 Roughly half of Nearctic amphibian species are believed to have stable populations whereas
(excluding disease) nearly one-third are thought to be decreasing (Table 7). The population trend is unknown for
Fire 13 16 a substantial percentage of species. For the majority of species, these trends are inferred
from trends in the state of the habitats on which the species depend (although in a few cases
population declines and increases have been noted). Species with decreasing populations
Table 5. The major threats to globally threatened amphibians in the Nearctic Realm. Only are typically forest-dependent species that can tolerate little disturbance to their habitats.
present threats to species are tallied. Overall trends for Nearctic amphibians reflect a healthier situation than is the case globally
where 43% are decreasing and only 27% are stable. In both cases, however, the percentage
of increasing species is very small.
“Rapidly Declining” Species
All habitat loss
Of the 470 globally “rapidly declining” species, only 24 (5%) occur within the Nearctic
Invasive species Realm (a full list of all “rapidly declining” species is provided in Appendix IV and includes
Utilization their occurrence within each of the realms). One of these 24 species, Rana megapoda, is
Accidental mortality in decline due to over-exploitation, 14 due to reduced habitat, and nine due to so-called
“enigmatic declines” (now generally attributed to the synergistic effects of chytridiomycosis
Persecution
and climate change (Lips et al. 2006; Pounds et al. 2006)).
Pollution The “rapidly declining” species show a distinct taxonomic pattern (Table 8). Among the
Natural disasters larger families, the Ranidae and Bufonidae show by far the highest percentages of species
in serious decline and in “enigmatic decline”. There are no Nearctic species in serious
Disease
decline in the Hylidae, and the percentages are also low in the Leptodactylidae and the
Human disturbance Plethodontidae. Some small families have high percentages of species in serious decline,
Changes in native sp. dynamics most notably the Proteidae, Salamandridae and Scaphiopodidae. The relatively small
Fire
number of “rapid declines” in the Nearctic are widely scattered across the region without
any clear geographic pattern.
0 20 40 60 80 100
Percentage of threatened species affected
KEY FINDINGS
Figure 9. The major threats impacting threatened amphibians in the Nearctic Realm. • A total of 337 species are recorded from the Nearctic Realm, of which 80 (24%) are
considered threatened.
• The Nearctic is the only region of the world in which salamanders outnumber frogs (195
as opposed to 142 species).
• At the species level, 230 amphibians (68% of those present) are endemic to the Nearctic;
of the 17 families found in the region, three are endemic, and of 51 amphibian genera
occurring, 19 are endemic. Endemism is lower in the Nearctic than in some other regions,
especially due to the unclear and somewhat arbitrary boundary with the Neotropics in
several places.
• The percentage of threatened or Extinct species is lower than in many other parts of
the world, but highest in the families Leptodactylidae (35%), Plethodontidae (31%),
Ranidae (29%), Ambystomatidae (29%), Rhyacotritonidae (25%), Bufonidae (21%), and
Salamandridae (17%).
• Geographic concentrations of threatened species in the Nearctic are less marked than
in many tropical regions, but there are some notable concentrations in the southern Ap-
palachians (centred on the Great Smoky Mountains), the mountains of northern Mexico,
the Edwards Plateau in Texas, and the mountains of California.
• The United States has the largest number of species in the Nearctic Realm (261 species),
and also has more endemics (182). Mexico has 68 species endemic to the Nearctic,
whereas Canada has none.
• The United States has the largest number of threatened species (52), followed by Mexico
(31). However, the percentage of threatened species is higher in Mexico (27% versus
20%).
• Threatened species tend to show distinct habitat preferences, with forest-dwelling and
stream-associated species being the most threatened (19% and 19%, respectively). This
pattern is mirrored elsewhere in the world.
• Habitat loss, primarily due to vegetation removal (mainly logging), urbanization/indus-
trial development, and expanding croplands is affecting almost 80% of the threatened
species in the region. Pollution impacts over one-third, and invasive species over 20%,
of the threatened species. Disease (usually chytridiomycosis) is negatively affecting
18% of globally threatened species in the region, but is implicated in 38% of the rapid
declines.
• Only 5% of the 470 globally “rapidly declining” species occur within the region. Most
of these rapid declines (58%) are caused by severe habitat loss, but chytridiomycosis is
a growing concern.
• Two amphibian extinctions have been recorded from the Nearctic, and one species is
Extinct in the Wild, all of these being from the United States. A further two species are
possibly extinct (both from Mexico).
 Chapter 8. Amphibians of the Nearctic Realm 89

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Dundee, H.A. and Rossman, D.A. 1989. The Amphibians and Reptiles of Louisiana. Louisiana State University Research 0 2 (1) 3 (1) 4 (1)
Press. Baton Rouge, Louisiana, USA. Specimen collecting 1 (1) 8 (4) 1 (1) 8 (4)
Faivovich, J., Haddad, C.F.B., Garcia, P.C.O., Frost, D.R., Campbell, J.A. and Wheeler, W.C. 2005. Systematic
review of the frog family Hylidae, with special reference to Hylinae: phylogenetic analysis and taxonomic
revision. Bulletin of the American Museum of Natural History 294:1-240.
Fellers, G.M., Green, D.E. and Longcore, J.E. 2001. Oral chytridiomycosis in the mountain yellow-legged Table 7. The population trends for all extant
frog (Rana muscosa). Copeia 2001:945-953. Population Trend Number of species % of species Nearctic amphibians.
Flint, R.F. 1971. Glacial and Quaternary Geology. John Wiley and Sons, New York, New York, USA. Decreasing 103 31
Flores-Villela, O. 1993. Herpetofauna Mexicana. Carnegie Museum of Natural History. Special Publica- Stable 162 48
tions 17:1-73. Increasing 2 0.6
Flores-Villela, O.A., Mendoza-Quijano, F. and Gonzalez-Porter, G. 1995. Recopilacion de claves para la Unknown 68 20
determinacion de anfibios y reptiles de Mexico. Publicaciones Especiales del Museo de Zoologia
Universidad Nacional de Autonoma de Mexico 10:1-285.
Fretwell, J.D., Williams, J.S. and Redman, P.J. (compilers). 1996. National Water Summary on Wetland
Resources. United States Geological Survey. Water-Supply Paper 2425, Reston, Virginia, USA. Table 8. The number of species in “rapid de-
Frost, D.R. 2004. Amphibian Species of the World: an Online Reference. Version 3.0 (22 August, 2004). Family Number of Percentage Number of Percentage cline” and “enigmatic decline” in the Nearctic
Electronic Database accessible at http://research.amnh.org/herpetology/amphibia/index.html. American species in of species species in of species Realm by Family.
“rapid decline” in family in “enigmatic in family in
Museum of Natural History, New York, USA.
“rapid decline” decline” “enigmatic
Frost, D.R., Grant, T., Faivovich, J.N., Bain, R.H., Haas, A., Haddad, C.F.B., de Sá, R.O., Channing, A., decline”
Wilkinson, M., Donnelly, S.C., Raxworthy, C.J., Campbell, J.A., Blotto, B.L., Moler, P., Drewes, R.C.,
Nussbaum, R.A., Lynch, J.D., Green, D.M. and Wheeler, W.C. 2006. The amphibian tree of life. Bulletin Bufonidae 5 18 3 11
of the American Museum of Natural History 297:1-370. Leptodactylidae 1 4 1 4
Gibbs, J.P., Breisch, A.R., Ducey, A.R., Johnson, G., Behler, J.L. and Bothner, R.C. In press. Amphibians Ranidae 8 21 4 11 The Coastal Tailed Frog Ascaphus truei (Least
and Reptiles of New York State: Identification, Natural History, and Conservation. Oxford University Scaphiopodidae 1 14 0 0 Concern) ranges from southern British Co-
Press, New York, New York, USA. Ambystomatidae 1 5 0 0 lumbia and northern California, and is one of
Green, D.M. (ed.) 1997. Amphibians in Decline: Canadian Studies of a Global Problem. Herpetological Plethodontidae 6 4 3 2 only two species in the Family Ascaphidae.
Conservation, Number 1, Society for the Study of Amphibians and Reptiles, St. Louis, Missouri, USA. Proteidae 1 20 0 0 Both species are associated with clear, cold,
Green, N.B. and Pauley, T.K. 1987. Amphibians and Reptiles in West Virginia. University of Pittsburgh Press, Salamandridae 1 17 0 0 swift-moving mountain streams in forests and
Pittsburgh, Pennsylvania, USA. lay eggs in long strings under stones in water.
Hammerson, G.A. 1999. Amphibians and Reptiles in Colorado. Second edition. University Press of Colorado, © Suzanne L. Collins, CNAH
Boulder, Colorado, USA.
Harding, J.H. 1997. Amphibian and Reptiles of the Great Lakes Region. University of Michigan Press, Ann Noss, R.F., LaRoe, E.T., III and Scott, J.M. 1995. Endangered Ecosystems of the United States: A Preliminary
Arbor, Michigan, USA. Assessment of Loss and Degradation. United States Department of the Interior, National Biological
Hulse, A.C., McCoy, C.J. and Censky, E.J. 2001. Amphibians and Reptiles of Pennsylvania and the Northeast. Service, Washington, D.C., USA.
Cornell University Press, Ithaca, New York, USA. Nussbaum, R.A., Brodie, E.D., Jr. and Storm, R.M. 1983. Amphibians and Reptiles of the Pacific Northwest.
Hunter, M.L., Jr., Calhoun, A.J.K. and McCollough, M.1999. Maine Amphibians and Reptiles. University of University Press of Idaho, Moscow, Idaho, USA.
Maine Press, Orono, Maine, USA. Petranka, J.W. 1998. Salamanders of the United States and Canada. Smithsonian Institution Press,
Johnson, T.R. 2000. The Amphibians and Reptiles of Missouri. Second edition. Missouri Department of Washington D.C., USA.
Conservation, Jefferson City, Missouri, USA. Pounds, J.A., Bustamante, M.R., Coloma, L.A., Consuegra, J.A., Fogden, M.P.L., Foster, P.N., La Marca,
Kagarise Sherman, C. and Morton, M.L. 1993. Population declines of Yosemite toads in the eastern Sierra E., Masters, K.L., Merino-Viteri, A., Puschendorf, R., Ron, S.R., Sánchez-Azofeifa, G.A., Still, C.J. and
Nevada of California. Journal of Herpetology 27:186-198. Young, B.E. 2006. Widespread amphibian extinctions from epidemic disease driven by global warming.
Klemens, M.W. 1993. Amphibians and Reptiles of Connecticut and Adjacent Regions. State Geological and Nature 439:161-167.
Natural History Survey of Connecticut, Bulletin 112, Hartford, Connecticut, USA. Ricketts, T.H., Dinerstein, E., Olson, D.M., Loucks, C.J., Eichbaum, W., Dellasaia, D., Kavanagh, K., Hedao,
Lannoo, M. (ed.). 1998. Status and Conservation of Midwestern Amphibians. University of Iowa Press, P., Hurley, P.T., Carney, K.M., Abell, R. and Walters, S. 1999. Terrestrial Ecosystems of North America,
Iowa City, Iowa, USA. A Conservation Assessment. Island Press, Washington, D.C., USA.
Lannoo, M. (ed.). 2005. Amphibian Declines: The Conservation Status of United States Species. University Russell, A.P. and Bauer. A.M. 1993. The Amphibians and Reptiles of Alberta. University of Calgary Press,
of California Press, Berkeley and Los Angeles, California, USA. Calgary, Alberta, and University of Alberta Press, Edmonton, Alberta.
LaRoe, W.T., Farris, G.S., Puckett, C.E., Doran, P.D. and Mac, M.J. (eds.). 1995. Our Living Resources: A Report Scott, G.A.J. 1995. Canada’s Vegetation, A World Perspective. McGill-Queen’s University Press, Montreal
to the Nation on the Distribution, Abundance, and Health of U.S. Plants, Animals, and Ecosystems. United and Kingston, Canada.
States Department of the Interior, National Biological Service, Washington, D.C., USA. Stebbins, R.C. 1972. California Amphibians and Reptiles. University of California Press. Berkeley, Cali- Endnotes
Lemos-Espinal, J., Smith, H.M. and Chiszar, D. 2004. Introducción a los Anfibios y Reptiles del Estado de fornia, USA. 1 P<0.01 (binomial test).
Chihuahua. UNAM-CONABIO, México, D.F. Stebbins, R.C. 2003. A field guide to western reptiles and amphibians. Third edition. Houghton Mifflin 2 At the time of writing, Chiropterotriton magnipes
Leonard, W.P., Brown, H.A., Jones, L.L.C., McCallister, K.R. and Storm, R.M. 1993. Amphibians of Washington Company, Boston, USA. was discovered in two new localities, away from
and Oregon. Seattle Audubon Society, Seattle, Washington, USA. Stein, B.A., Kutner, L.S. and Adams, J.S. (eds.). 2000. Precious Heritage: The Status of Biodiversity in the the type locality (G. Parra Olea pers. comm.).
Lips, K.R., Brem, F., Brenes, R., Reeve, J.D., Alford, R.A., Voyles, J., Carey, C., Livo, L., Pessier, A.P. and United States. Oxford University Press, New York, New York, USA. 3 By merging the Dicamptodontidae into the Ambys-
Collins, J.P. 2006. Emerging infectious disease and the loss of biodiversity in a Neotropical amphibian Stuart, S.N., Chanson, J.S., Cox, N.A., Young, B.E., Rodrigues, A.S.L., Fischman, D.L. and Waller, R.W. 2004. tomatidae, Frost et al. (2006) reduce the number
community. Proceedings of the National Academy of Sciences 103:3165-3170. Status and trends of amphibian declines and extinctions worldwide. Science 306:1783-1786. of families in the region to 16, and the number of
MacCulloch, R.D. 2002. The ROM Field Guide to Amphibians and Reptiles of Ontario. Royal Ontario Museum Trauth, S.E., Robinson, H.W. and Plummer, M.V. 2004. The Amphibians and Reptiles of Arkansas. University endemic families to two. Frost et al. (2006) also
and McClelland and Stewart Ltd., Toronto, Ontario. of Arkansas Press, Fayetteville, Arkansas, USA. merged Ascaphidae into Leiopelmatidae. However,
Martof, B.S., Palmer, W.M., Bailey, J.R. and Harrison III, J.R. 1980. Amphibians and Reptiles of the Carolinas Vazquez-Díaz, J. and Quintero-Díaz, G.E. 2005. Anfibios y Reptiles de Aguascalientes. CONABIO-CIEMA- in this section we follow the former taxonomic
and Virginia. University of North Carolina Press, Chapel Hill, North Carolina, USA. Internacional de Relojes, México. arrangement of families based on Frost (2004).
Matsuda, B., Green, D.M. and Gregory, P.T. 2006. The Amphibians and Reptiles of British Columbia. Royal Weller, W.F. and Green, D.M. 1997. Checklist and current status of Canadian amphibians. Herpetological 4 Frost et al. (2006) transfer these species to other
British Columbia Museum, Victoria, Canada. Conservation 1:309-328. genera.
Minton, S.A. Jr. 2001. Amphibians and Reptiles of Indiana. Indiana Academy of Science, Indianapolis, Werner, J.K., Maxell, B.A., Hendricks, P. and Flath, D.L. 2004. Amphibians and Reptiles of Montana. Mountain 5 Frost et al. (2006) transfer Eleutherodactylus and
Indiana, USA. Press Publishing Company, Missoula, Montana, USA. Craugastor to the family Brachycephalidae.
Mount, R.H. 1975. The Reptiles and Amphibians of Alabama. Auburn University Agricultural Experiment Young, B.E., Stuart, S.N., Chanson, J.S., Cox, N.A. and Boucher, T.M. 2004. Disappearing Jewels: The Status 6 Frost et al. (2006) transfer most of these to the
Station. Auburn, Alabama, USA. of New World Amphibians. NatureServe, Arlington, Virginia, USA. genus Lithobates
90
ESSAY 8.1. APPALACHIAN AMPHIBIANS
Although widely distributed throughout much of the United States, southern
Canada south to northern Mexico in the Sierra Madre Occidental, the Tiger
Salamander Ambystoma tigrinum (Least Concern) is absent from most of the
Appalachian Mountains and occurs only in the Shenandoah Valley at the base
of the Blue Ridge Mountains in west-central Virginia. © Don Church
The Appalachian Mountains lie in a 2,414km north-east to south-west
stretch of eastern North America from Newfoundland and Labrador in
Canada to north-central Alabama in the United States (with their foothills in
north-eastern Mississippi). The topographic system is divided into a series
of ranges, with the highest of the group being Mt. Mitchell in North Carolina
(2,040m). This summit is also the highest point on the continent east of the
Mississippi River. Because North America and Africa were once connected,
the Appalachians were once part of the same mountain chain as the Atlas
Mountains in Morocco. Many floristic and some animal species (e.g., giant
salamanders) are also related to those in the mountains of western China
based on ancient continental connections.
The Appalachian region supports a startling diversity of amphibians, all
the more so since more than one-third of the species (37 of the 98 present)
are endemic. Salamander genetic diversity in the Appalachians is the highest
in the world, although Central America supports a higher species richness
(see Essay 9.3). In total, 72 salamander species inhabit the region, with 60
representatives of the family Plethodontidae (the lungless salamanders). All
of the region’s endemic amphibians are plethodontids. Although the area’s
salamander fauna has been intensively studied for over 150 years, new spe-
cies are still being discovered. Several species with restricted distributions
remained hidden to herpetologists among mountains and caves until the
middle of the 20th century. Dunn (1926) recognized 22 species in his seminal
revision of the Plethodontidae. By 1967, the plethodontid count was at 26
species (Brame 1967). However, the advent of molecular genetic techniques
in the mid-1960s sparked another scientific inquisition into the region’s
salamander fauna. The number of described plethodontid species more
than doubled by the beginning of the 21st century. For example, Conant and
Collins (1998) recognized 56 Appalachian species in their field guide. Genetic
analyses continue to discover cryptic species that are difficult to resolve with
only morphological differences.
Colour, pattern, and morphological variation abound geographically and
among species in the Appalachians. Green Salamanders, Aneides aeneus (NT),
have, for example, a bright green moss-like pattern, splayed limbs, and large
feet used for climbing and living in rock crevices. Seepage and streamside
salamanders include the bright yellow, sleek Two-lined Salamanders, mem-
bers of the genus Eurycea, the robust, brilliant Red Salamander, Pseudotriton
ruber (LC), and the large Spring Salamander, Gyrinophilus porphyriticus (LC),
a predator of all of the rest. The many streamside salamanders in the genus
Desmognathus have dull brown to black colours and varying dull reticulate
and spot patterns, but are amazingly abundant in mountain streams. Some
species are fully aquatic, like the Shovel-nosed Salamander, Desmognathus
marmoratus (LC), whereas others spend as much time on land as in the water,
like the Allegheny Mountain Dusky Salamander, Desmognathus ochrophaeus
(LC). The many species in the completely terrestrial salamander group, genus
Plethodon, lay eggs in moist microhabitats in forests where all embryonic
development occurs within the egg capsule. Hatchlings are adult forms and
fully terrestrial. The Red-backed Salamander, Plethodon cinereus (LC), may be
the most abundant vertebrate in the Appalachians. Their biomass per square
meter can be higher than all the mammals and equal to that of all the birds
in the area (Burton and Likens 1975).
The lungless salamanders within the Appalachian region have been very
valuable to our understanding of speciation and other evolutionary processes
(Frost 2000). The diversity of the Appalachian amphibian fauna is enhanced
by a few species from other families that have small relict populations. For
example, the Tiger Salamander Ambystoma tigrinum (LC) is found patchily
along the Coastal Plain, but is also known to occur in only a single location
within the Appalachian region. This disjunct population is dependent upon
a unique complex of ancient sinkhole ponds in the Shenandoah Valley
at the base of the Big Levels massif of the Blue Ridge Mountain chain in
west-central Virginia (Buhlmann and Hoffman 1990; Mitchell and Buhlmann
1999). Although the next nearest Tiger Salamander population occurs on
Virginia’s Coastal Plain, mitochondrial DNA evidence reveals that this relict
mountain population is most closely related to a population in west-central
South Carolina, but became isolated 100,000-300,000 years ago (Church et
al. 2002). Genetic and fossil data strongly suggest that the Tiger Salamander
once had a more continuous distribution throughout the Appalachian region
but became isolated in the Big Levels area before or during the last Pleisto-
cene glaciation event. Interestingly, these Shenandoah Valley sinkhole ponds
also support disjunct populations of plant, dragonfly, and damselfly species
Threatened Amphibians of the World
(Fleming and Van Alstine 1999; Roble 1999) suggesting that this region of Brame, A.H., Jr. 1967. A list of the world’s recent and fossil salamanders. Herpeton
Appalachia may have provided Pleistocene refugia for entire communities. 2:1-26.
Future studies of these relict populations will reveal how species ranges Buhlmann, K.A. and Hoffman, R.L. 1990. Geographic distribution: Ambystoma tigrinum
have contracted and expanded and, hence, what the consequences may be tigrinum. Herpetological Review 21:36.
from future climate change. Burton, T.M. and Likens, G.E. 1975. Salamander populations and biomass in the Hubbard
Several of the endemic Appalachian plethodontid salamander species have Brook Experimental Forest, New Hampshire. Copeia 1975:541-546.
extremely restricted distributions. These species are susceptible to habitat Bury, R.B., Dodd, C.K., Jr., and Fellers, G.M. 1980. Conservation of the amphibia of the
loss and climate change and are, therefore, important flagship species for United States: a review. U.S. Department of the Interior, Fish and Wildlife Service.
the region. The Shenandoah Salamander, Plethodon shenandoah (VU), for Resource Publication 134:1-34.
example, is restricted to talus slopes between 914 and 1,143m asl on the Church, S.A., Kraus, J.M., Mitchell, J.C., Church, D.R., and Taylor, D.R. 2002. Phylogeog-
north and north-west slopes of Hawksbill Mountain, Stony Man Mountain, raphy of the eastern tiger salamander: Effects of the Pleistocene glaciation on current
and The Pinnacles in the Blue Ridge of north-central Virginia (Petranka genetic relationships. Evolution 57:372-383.
1998; Mitchell and Reay 1999). The conservation status of this species is Conant, R., and Collins, J.T. 1998. A Field Guide to Reptiles and Amphibians, Eastern and
apparently secure for the immediate future given that its entire range lies Central North America. Third, expanded edition. Houghton Mifflin Company, Boston,
within Shenandoah National Park. The Peaks of Otter Salamander, P. hubrichti Massachusetts, USA.
(VU), is another small, entirely terrestrial species restricted to a 19-km high Dodd, C.K., Jr. 2004. The Amphibians of Great Smoky Mountains National Park. University
elevation length of the Blue Ridge Mountains in west-central Virginia. of Tennessee Press, Knoxville, Tennessee. USA.
Although abundant within parts of its range, this salamander is susceptible Dunn, E.R. 1926. The Salamanders of the Family Plethodontidae. Smith College, Northamp-
to timbering and habitat fragmentation due to its very low dispersal rate ton, Massachusetts, USA.
and restricted distribution (Kramer et al. 1993). The Junaluska Salamander, Fleming, G.P. and Van Alstine, N.F. 1999. Plant communities and floristic features of
Eurycea junaluska (VU), is another rare Appalachian salamander found patchily sinkhole ponds and seepage wetlands in southeastern Augusta County, Virginia.
and in low abundance in the Cheoah River drainage of North Carolina and Banisteria 13:67-94.
along Fighting Creek and Tellico River in Tennessee. This very rare species Frost, D. 2000. Species, descriptive efficiency, and progress in systematics. In: R.C. Bruce,
has a biphasic life history, with stream-dwelling larvae and adults that enter R.G. Jaeger and L.D. Houck (eds.), The Biology of Plethodontid Salamanders, pp. 7-29.
forests. Thus, the Junaluska Salamander is vulnerable to degradation of both Kluwer Academic/Plenum Publishers, New York, USA.
stream and forest habitats. Kramer, P., Reichenbach, N., Hayslett, M. and Sattler, P. 1993. Population dynamics
Although not endemic to Appalachia, the Hellbender, Cryptobranchus and conservation of the Peaks of Otter salamander, Plethodon hubrichti. Journal of
alleganiensis (NT), is another important flagship amphibian for the region. Herpetology 27:431-435.
This species is the sole representative of its genus, and one of only three McCoy, C.J. 1982. Amphibians and reptiles in Pennsylvania. Special Publications of the
surviving species within the family of giant salamanders, Cryptobranchidae. Carnegie Museum of Natural History 6:1-91.
Hellbenders are bizarre animals with fleshy folds of skin undulating along Mitchell, J.C., Breisch, A.R. and Buhlmann, K.A. 2006. Habitat Management Guidelines
their sides and are capable of reaching 74 cm in total length. Living their for Amphibians and Reptiles of the Northeastern United States, pp. 106. Partners in
entire lives in large, fast-flowing streams, they have laterally compressed Amphibian and Reptile Conservation, Technical Publication HMG-3, Montgomery,
tails and are capable of swimming quickly underwater. Their large, dorsally Alabama, USA.
flattened heads support only diminutive eyes, but have huge mouths capable Mitchell, J.C. and Buhlmann, K.A. 1999. Amphibians and reptiles of the Shenandoah Valley
of engulfing crayfish and small fish, the mainstays of their diet. A mature male Sinkhole Pond System in Virginia. Banisteria 13:129-142.
constructs a nesting cavity under a large flat rock during the breeding season Mitchell, J.C., Pauley, T.K., Withers, D.I., Cupp, P.V., Braswell, A.L., Miller, B., Roble, S.M.
and aggressively guides gravid females to enter, whereupon he detains her and Hobson, C.S. 1999. Conservation status of the southern Appalachian herpetofauna.
until she oviposits her clutch of up to 750 eggs. The male then fertilizes the Virginia Journal of Science 50:13-36.
eggs, evicts the mother, and subsequently guards the nest from intruders Mitchell, J.C. and Reay, K.K. 1999. Atlas of Amphibians and Reptiles of Virginia. Virginia
for a period of the incubation (Bishop 1943; Dodd 2004). Hellbenders were Department of Game and Inland Fisheries, Special Publication Number 1, Richmond,
traditionally eaten by Native Americans, who probably skinned them first as Virginia, USA.
their slimy skin secretions are toxic to many predators (McCoy 1982). The Petranka, J.W. 1998. Salamanders of the United States and Canada. Smithsonian Institu-
majority of the Hellbender’s range occurs within the Appalachian region, tion Press, Washington D.C., USA.
where it was undoubtedly abundant in streams prior to European colonization Roble, S.R. 1999. Dragonflies and damselflies (Odonata) of the Shenandoah Valley Sinkhole
(Petranka 1998). Populations have since declined drastically across much of Pond System and vicinity, Augusta County, Virginia. Banisteria 13:101-127.
their range and have gone locally extinct due to their susceptibility to stream
impoundment, siltation from timber harvest operations and development, and
pollution from industry (Bury et al. 1980; Mitchell et al. 1999). Although many
river systems within the Hellbender’s range are protected, this species, like The Blue Ridge Red Salamander Pseudotriton ruber (Least Concern) ranges
many amphibians, is highly susceptible to anthropogenic activities outside from the Hudson River in New York southwestward to Indiana and southward
the primary habitat. to Louisiana and the Gulf Coast; adults are both aquatic and terrestrial.
Despite the high species richness © Don Church
and abundance of the salamanders in
the Appalachian Mountains of North
America, many of them are threat-
ened. Population extinction and range
contraction is occurring for several
of these unique vertebrates. None
appears to be in imminent danger
of complete extinction, but urban
sprawl, pollution, climate change, and
some types of agriculture and timber
harvesting are causing declines in
many areas. Fortunately, large parts
of the Appalachian Mountain chain,
especially in the southern Appala-
chians, are in U.S. National Park and
National Forest Service ownership.
Conservation and management of
these spectacular amphibians is
ongoing at many levels, such as with
the Habitat Management Guidelines
produced by Partners in Amphibian
and Reptile Conservation (Bailey et al.
2006; Mitchell et al. 2006).
Don R. Church, Thomas S.B.
Akre, and Joseph C. Mitchell
References
Bailey, M.A., Holmes, J.N., Buhlmann, K.A.
and Mitchell, J.C. 2006. Habitat Manage-
ment Guidelines for Amphibians and Rep-
tiles of the Southeastern United States,
pp. 88. Partners in Amphibian and Reptile
Conservation, Technical Publication HMG-
2, Montgomery, Alabama, USA.
Bishop, S.C. 1943. Handbook of Salamanders
of the United States and Canada. Cornell
University Press, Ithaca, New York, USA.

ESSAY 8.2. AMPHIBIAN DECLINES IN CALIFORNIA
California, along with Central America and Australia, has been a focal area
for the study of amphibian population declines, because of the severe de-
clines of many of its species. The region is recognized as one of the world’s
biodiversity hotspots (the “California Floristic Province”) and contains a
heterogeneous landscape that sustains a wide variety of ecosystems, such
as Sonoran deserts, marshes and wetlands, oak woodlands, high-elevation
alpine systems, temperate rain forests, and many others. The amphibian fauna
is diverse and, at the time of writing, includes 64 recognized native species
(Figure 1a), including 40 species of salamanders from five families and nine
genera, and 24 species of frogs and toads from five families and six genera
(plus two introduced species; AmphibiaWeb 2006). Amphibians in California
can be found in nearly all habitat types ranging from near Mount Whitney (at
3,657 m, the highest peak in the contiguous United States) to Death Valley
(85 m below sea level). Despite the fact that California contains some of the
largest contiguous protected habitats in the continental United States, nearly
one-quarter of amphibians in California are threatened (Figure 1b).
Many potential causes for the widespread declines of amphibians have
been proposed. In general these can be grouped into two major categories:
1) factors general to the overall biodiversity crisis, including habitat destruc-
tion, alteration and fragmentation, introduced species and over-exploitation,
and 2) factors associated with amphibians that might account for declines
in relatively undisturbed habitats. The first category includes relatively well
understood direct ecological phenomena, whereas the second includes com-
plex and elusive mechanisms, such as climate change, increased ultraviolet
(UV-B) radiation, chemical contaminants, infectious diseases, and the causes
of deformities (or malformations). The underlying mechanisms behind these
factors are complex and may be working synergistically with more evident
factors, such as habitat destruction and introduced species, to exacerbate
declines. Many biologists believe that there are some dominant causes, such
as new infectious diseases, whereas others are not convinced that there is
a single overarching cause for global declines, but that many factors are
threatening amphibian populations to a greater or lesser extent.
In California, amphibian declines are associated with many of the various
hypotheses. Habitat destruction, alteration, and fragmentation have affected
a large number of species including the Foothill Yellow-legged Frog R. boylii
(NT), the Arroyo Toad Bufo californicus (EN), and the California Tiger Sala-
mander Ambystoma californiense (VU), to name a few (Lannoo 2005). Some
amphibians suffered declines long ago. In the 19th century, the California
Gold Rush brought waves of new settlers who quickly over-exploited some
frog species for food. They also altered the environment in ways that have
had much more substantial effects on amphibians. Cities were built, rivers
dammed and diverted, forests were cleared, and the waterways of Great
Central Valley were completely altered for agriculture and to provide water
for cities and industrial growth. The effect on California’s ecosystems has been
profound. As elsewhere, habitat conservation has become a central theme
in efforts to preserve the region’s biodiversity. However, recent amphibian
declines in California have occurred in remote habitats well protected from
development.
Beginning in the 1980s, scientists began to document alarming amphib-
ian disappearances in protected habitats in California, Central America, and
Australia. Some of the best documented examples of declines can be found
in the Sierra Nevada, which not only contains large sections of roadless,
undisturbed habitat, but also has a rich history of biological surveys going
back to the turn of the 20th century (Grinnell and Storer 1924). These data
are useful for comparisons with present day distributions of amphibians.
For example, historical surveys noted abundant amphibians throughout the
Sierra Nevada, and at higher elevations (>1500 m) the Mountain Yellow-
legged Frog Rana muscosa (CR) (Figure 2) was termed the most abundant of
all vertebrates! But, in the last three decades, nearly the entire amphibian
fauna of the Sierra Nevada has collapsed. Air pollution, increases in harmful
ultraviolet radiation (UV-B), introduced predators, and emerging diseases have
The Mountain Yellow-legged Frog Rana muscosa (Critically Endangered)
from California and Nevada, USA, has declined by the loss of over 93% of
historic populations. This frog is extinct in the state of Nevada, but scattered
populations remain in the Sierra Nevada along eastern California and in three
mountain tops surrounding Los Angeles (fewer than 200 individuals survive in
only eight populations in the southern mountains). Major threats to remaining
populations include air pollution, disease and introduced predators (trout).
© Vance Vredenburg
Chapter 8. Amphibians of the Nearctic Realm
all been proposed as key factors that may explain the enigmatic declines in
this protected landscape.
The first reports of amphibian collapse in the Sierra Nevada came from
Yosemite Toads Bufo canorus (EN) (Stebbins and Cohen 1995). Hundreds
of animals were found dead and dying, and many populations simply
disappeared. A similar pattern was found with Mountain Yellow-legged
Frogs (Stebbins and Cohen 1995), while Western Toads Bufo boreas (NT),
Foothill Yellow-legged Frogs, and even Long-toed Salamanders Ambystoma
macrodactylum (LC) also suddenly declined throughout their ranges. On the
other hand, Pacific Chorus Frogs Pseudacris regilla (LC) remained abundant
(Stebbins and Cohen 1995).
The Sierra Nevada became a testing ground for hypotheses that could
account for disappearances in remote, seemingly pristine, habitats. One
example is the UV-B hypothesis, which states that human-induced climate
modification results in increased levels of harmful UV-B that can kill am-
phibian eggs exposed to direct sunlight. A correlational analysis showed
that there was no negative pattern between expected high UV-B dose and
disappearance of frogs, but did reveal that frog populations occurring closer
to pesticide and fertilizer sources were more likely to have gone extinct
than populations in more remote areas (Davidson et al. 2001). Disease, in
particular chytridiomycosis, is also a factor and responsible for the collapse
of amphibian populations in Central and South America (Berger et al. 1998;
Lips et al. 2006), and Australia (Berger et al. 1998). This disease was found
in collapsing populations of Mountain Yellow-legged Frogs in the Sierra
Nevada (Rachowicz et al. 2006), although it also appears to be endemic in
other amphibian species that have stable populations.
Amphibian declines may be the window into the future of what we can
expect as humans continue to alter their environment on a global scale. As we
learn more about the mechanisms responsible for declines, we may be able
to reverse some of them, though there remain few examples. In the Sierra
Nevada, one study has shown that, in the absence of disease, population
recovery of threatened frogs is possible (Vredenburg 2004). Trout have been
extensively introduced throughout the Sierra Nevada for sport fishing, and
more than 90% of the naturally fishless lakes now contain non-native trout.
The Mountain Yellow-legged Frog, which is adapted to living in environments
without any fish, has declined dramatically (more than 93% of historical
populations are now extinct; Vredenburg et. al. 2007), and while there are
several potential causes, removal of introduced trout from entire lakes leads
to recovery of local frog populations.
Vance T. Vredenburg, Michelle S. Koo, and David B. Wake
91
References
AmphibiaWeb. 2006. AmphibiWeb: Information on amphibian biology and conservation
[web application]. in. Available: http://amphibiaweb.org/, Berkeley, California:
AmphibiaWeb. (Accessed 2006).
Berger, L., Speare, L., Daszak, P., Green, D.E., Cunningham, A.A., Goggin, C.L., Slocombe,
R., Ragan, M.A., Hyatt, M.A., McDonald, K.R., Hines, H.B., Lips, K.R., Marantelli, G.,
and Parkes, H. 1998. Chytridiomycosis causes amphibian mortality associated with
population declines in the rain forests of Australia and Central America. Proceedings
of the National Academy of Sciences USA 95:9031-9036.
Davidson, C., Shaffer, H.B. and Jennings, M.R. 2001. Declines of the California red-legged
frog: Climate, UV-B, habitat, and pesticides hypotheses. Ecological Applications
11:464-479.
Gastner, M.T. and Newman, M.E.J. 2004. Diffusion-based method for producing
density-equalizing maps. Proceedings of the National Academy of Sciences USA
101:7499-7504.
Grinnell, J. and Storer, T. 1924. Animal life in the Yosemite. University of California
Press, Berkeley, USA.
Lannoo, M.J. (eds.). 2005. Amphibian declines: the conservation status of the United
States species. University of California Press, Berkeley, USA.
Lips, K.R., Brem, F., Brenes, R., Reeve, J.D., Alford, R.A., Voyles, J., Carey, C., Livo, L.,
Pessier, A.P.,and Collins, J.P. 2006. Emerging infectious disease and the loss of biodi-
versity in a Neotropical amphibian community. Proceedings of the National Academy
of Sciences USA 103:3165-3170.
Rachowicz, L.J., Knapp, R.A., Morgan, J.A.T., Stice, M.J., Vredenburg, V.T., Parker, J.M.
and Briggs, C.J. 2006. Emerging infectious disease as a proximate cause of amphibian
mass mortality in Rana muscosa populations. Ecology 87:1671-1683.
Stebbins, R.C. and Cohen, N.W. 1995. Declining Amphibians. A Natural History of Amphib-
ians. Princeton University Press, Princeton, NJ, USA.
Stuart, S., J.S. Chanson, N.A. Cox, B.E. Young, A.S.L. Rodrigues, D.L. Fishman, and R.W.
Waller. 2004. Status and trends of amphibian declines and extinctions worldwide.
Science 306:1783-1786.
Vredenburg, V.T. 2004. Reversing introduced species effects: Experimental removal of
introduced fish leads to rapid recovery of a declining frog. Proceedings of the National
Academy of Sciences USA 101:7646-7650.
Vredenburg, V.T., Bingham, R., Knapp, R., Morgan, J.A.T., Moritz, C. and Wake, D. 2007.
Concordant molecular and phenotypic data delineate new taxonomy and conservation
priorities for the endangered mountain yellow-legged frog (Ranidae: Rana muscosa).
Journal of Zoology, London 271:361-374.
Figure 1. Amphibian species
diversity by U.S. state (A),
and percent of threatened
amphibian species by state
(b) visualized using den-
sity equalizing cartograms
(technique after Gastner and
Newman 2004). State size
and shape are purposefully
distorted in proportion to
(in A) the total number of
amphibian species, and (B)
the percentage of threatened
species by state (data sourc-
es; A: AmphibiaWeb 2006; B:
Stuart et al. 2004).
 CHAPTER 9. AMPHIBIANS OF THE NEOTROPICAL REALM

Cochranella vozmedianoi (Data Deficient) is Federico Bolaños, Fernando Castro, Claudia Cortez, Ignacio De la Riva, Taran
a poorly known species endemic to Cerro El Grant, Blair Hedges, Ronald Heyer, Roberto Ibáñez, Enrique La Marca, Esteban
Humo, in the Península de Paria, in northern Lavilla, Debora Leite Silvano, Stefan Lötters, Gabriela Parra Olea, Steffen
Venezuela. It is a glass frog from the Family Reichle, Robert Reynolds, Lily Rodríguez, Georgina Santos Barrera, Norman
Centrolenidae that inhabits tropical humid Scott, Carmen Ubeda, Alberto Veloso, Mark Wilkinson and Bruce Young
forests, along streams. It lays its eggs on the
upper side of leaves overhanging streams. The
larvae fall into the stream below after hatch- THE GEOGRAPHIC AND HUMAN CONTEXT
ing. © Juan Manuel Guayasamin
The Neotropical Realm includes all of mainland South America, much of Mesoamerica
(except parts of northern Mexico), all of the Caribbean islands, and extreme southern Texas
and Florida in the United States.
South America has a long history of geographic isolation that began when this continent
separated from other Southern Hemisphere land masses 40-30 Ma. The Andes, one of the
largest mountain ranges on earth and reaching 6,962m at Acongagua in Argentina, began to
uplift 80-65Ma as South America drifted west from Africa. The other prominent mountainous
areas on the continent are the Tepuis of the Guianan Shield, and the highlands of south-
eastern Brazil. The complex patterns of wet and dry habitats on the continent are the result
of an array of factors, including the climatic effects of cold ocean currents interacting with
these mountain ranges, orographic barriers to winds carrying humidity within the continent,
and the constant shifting of the intertropical convergence zone, among others.
Figure 1. Summary of Red List categories for The geological history of Mesoamerica is very complex and still not completely under-
amphibians in the Neotropical Realm. stood. The land north of the Isthmus of Tehuantepec in southern Mexico is historically part ANURA Plectrohyla hazelae
of the North American continent, with the highest point at Pico de Orizaba at 5,610m asl. Bufonidae Plectrohyla siopela
The land south of the Isthmus to the southern Nicaragua lowlands is a mosaic of plates that Andinophryne colomai Plectrohyla thorectes
Red List Category Number of species have rearranged themselves and alternately been submerged and exposed by the sea several Atelopus arthuri Scinax heyeri
Extinct (EX) 7 times during the last 65 million years. The region encompassed by Panama, Costa Rica, and Atelopus balios Leptodactylidae
Extinct in the Wild (EW) 0 southern Nicaragua formed over the last three to ten million years through a combination of Atelopus carbonerensis Craugastor anciano
Critically Endangered (CR) 358 volcanic activity and uplift. The result is a jumble of mountain ranges interrupted by valleys Atelopus chiriquiensis Craugastor andi
Endangered (EN) 456 and lowlands, with the highest point being Volcán Tajumulco at 4,220m asl in Guatemala. Atelopus chrysocorallus Craugastor angelicus
Vulnerable (VU) 324 The closing of these different blocks of land in present-day Nicaragua during the Pliocene Atelopus coynei Craugastor coffeus
Near Threatened (NT) 140 (5.3 - 1.8 Ma) has had a marked impact on the distribution of amphibians today. Atelopus famelicus Craugastor cruzi
Least Concern (LC) 956 The geological history of the Caribbean also remains under intense study, but most Atelopus guanujo Craugastor escoces
Data Deficient (DD) 675 geologists now agree that the Greater Antilles (Cuba, Jamaica, Hispaniola, and Puerto Rico) Atelopus halihelos Craugastor fecundus
Total Number of Species 2,916 are geological cousins of the plates that make up northern Central America. Some 100 Ma, Atelopus lozanoi Craugastor fleischmanni
these islands were lined up more or less between North and South America in approximately Atelopus lynchi Craugastor guerreroensis
0.2%
12% the location of present day Central America. Over the last 70 million years, these islands Atelopus mindoensis Craugastor merendonensis
23% have drifted east to their current positions. The trailing edge of this parade of islands has Atelopus muisca Craugastor omoaensis
fused to North America and now makes up northern Central America. Some of the Greater Atelopus nanay Craugastor polymniae
16% Antilles may have had temporary land connections with North and/or South America as Atelopus oxyrhynchus Craugastor saltuarius
they drifted eastward. The Lesser Antilles formed in a completely different manner. As the Atelopus pachydermus Craugastor stadelmani
Caribbean Plate carrying the Greater Antilles moved eastward, the America Plate subducted Atelopus peruensis Craugastor trachydermus
beneath it, creating an island arc known today as the Lesser Antilles. Atelopus pinangoi Crossodactylus trachystomus
The region is enormously varied ecologically. Equatorial South America is dominated by the Atelopus planispina Cryptobatrachus nicefori
11% lowland rainforest of the Amazon Basin. South of this the habitats become progressively drier Atelopus sorianoi Cycloramphus ohausi
33% and less suitable for amphibians, with the exception of the now extensively cleared Atlantic Atelopus senex Eleutherodactylus bernali
Forest of southern and eastern Brazil, eastern Paraguay, and north-eastern Argentina, and the Atelopus sernai Eleutherodactylus emleni
5% Bufo fastidiosus Eleutherodactylus eneidae
temperate forests of Chile. There are important wetland areas in the Llanos of Venezuela and
Colombia, and the Pantanal of Brazil, Paraguay and Bolivia. The topographically varied Andean Bufo fluviaticus Eleutherodactylus glanduliferoides
region includes all varieties of habitats, from some of the wettest lowland rainforest in the Pacific Bufo holdridgei Eleutherodactylus jasperi
lowlands of Colombia to the Atacama Desert in northern Chile (the driest place on earth), and Melanophryniscus macrogranulosus2 Eleutherodactylus karlschmidti
from cold temperate habitats in the extreme south of the continent to high mountain paramos in Rhamphophryne rostrata Eleutherodactylus olanchano
the tropics. In Mesoamerica there is a very complex patchwork of natural habitats, with humid Centrolenidae Eleutherodactylus orcutti
mountain slopes rising above both dry (generally on the Pacific side) and wet (Caribbean side) Centrolene ballux Eleutherodactylus schmidti
lowland habitats. The Caribbean islands are also a complex mosaic of habitats, with low-lying Centrolene heloderma Eleutherodactylus semipalmatus
islands tending to be semi-arid, and wetter environments occurring where trade winds encounter Hyalinobatrachium crybetes Eleutherodactylus zongoensis
the higher Caribbean mountains giving rise to a variety of moist tropical forest types. Dendrobatidae Gastrotheca lauzuricae
On average, the Neotropics have a relatively low human population density (approxi- Aromobates nocturnes Holoaden bradei
mately 27 people per square kilometre in 2005), only 22% of which lives in rural areas, and Colostethus dunni Odontophrynus moratoi
a population growth rate (1.4% per annum) that is decreasing. However, some Caribbean Colostethus edwardsi Paratelmatobius lutzii
islands are among the most densely populated places on Earth. Historically, South America Colostethus jacobuspetersi Paratelmatobius mantiqueira
has been subject to relatively low levels of anthropogenic disturbance, but human impact Colostethus ruizi Phrynopus spectabilis
has been higher in northern Mesoamerica and on the Caribbean islands. The low impact Colostethus vertebralis Telmatobius cirrhacelis
in much of the region is related to low human population densities. The gross income per Dendrobates abditus Telmatobius niger
capita was around US$3,500 in the region in 2004. However, 33% of the region’s Gross Mannophryne neblina Telmatobius vellardi
Domestic Product is concentrated in Mexico and a further 30% in Brazil, where the human Hylidae Ranidae
impacts on natural ecosystems and biodiversity have been much more severe. Economic Aplastodiscus flumineus Rana omiltemana
growth rates in the region have been modest (2.7% in 2003). Bromeliohyla dendroscarta Rana pueblae
The human impact on ecosystems is very variable through the region. Natural habitats Bokermannohyla claresignata Rana tlaloci*
have been particularly severely damaged on many of the Caribbean islands (and especially Bokermannohyla izecksohni Rhinodermatidae
in Haiti), and in the northern part of Mesoamerica (from central Mexico, south to Guatemala, Charadrahyla altipotens Rhinoderma rufum
Honduras and El Salvador). There has also been extensive habitat loss through much of the Charadrahyla trux
Andes, and especially in the Atlantic Forests and Cerrado of central, southern and eastern Ecnomiohyla echinata CAUDATA
Brazil, and in the native southern temperate forests of Chile. The forests of the Amazon Hyla bocourti Plethodontidae
Basin and Guianan Shield are still largely intact, although there is much clearance currently Hyla chlorostea Bolitoglossa jacksoni
taking place along the southern edge of the forest zone in Brazil. Habitat loss in the region Hypsiboas cymbalum Bradytriton silus
has been driven largely by expanding subsistence agriculture to support growing human Isthmohyla calypsa Chiropterotriton magnipes*
populations, and also by commercial agriculture and logging. Isthmohyla debilis Ixalotriton parva
Isthmohyla graceae Oedipina paucidentata
Isthmohyla rivularis Pseudoeurycea aquatica
GLOBAL CONSERVATION STATUS Isthmohyla tica Pseudoeurycea naucampatepetl
Megastomatohyla pellita Pseudoeurycea nigromaculata
A total of 2,916 amphibian species (49% of the world’s total) are recorded from the Neotropi- Plectrohyla calvicollina Pseudoeurycea praecellens
Table 1. The Critically Endangered (Possibly cal Realm, of which 1,145 (39%) are considered to be globally threatened (Figure 1). This is Plectrohyla celata Thorius infernalis
Extinct) amphibian species in the Neotropical significantly more than the global average of 33%1. As is the case globally, the percentage Plectrohyla cembra Thorius magnipes
Realm (*denotes species that also occur in the of threatened species is expected to increase as the status of DD species is clarified, as Plectrohyla cyanomma Thorius narismagnus
Nearctic Region). A full list of CR(PE) species new species (some of which are likely to be rare, and/or have small ranges) are discovered, Plectrohyla ephemera Thorius narisovalis
can be found in Appendix IX. and as the taxonomic status of many species complexes is resolved.
 Chapter 9. Amphibians of the Neotropical Realm 93

The Neotropical Realm contains 60% (1,145) of all globally threatened amphibians. The
region accounts for a massive 79% of CR species, 59% of the EN species, and 48% of the
VU species in the world. In other words, unlike the case elsewhere, threatened Neotropical
amphibians are more likely to be in a higher category of threat (CR or EN), when compared
with the global distribution of threatened species amongst categories. This tendency for
threatened species to be in CR and EN is probably explained in part by the effects of habitat
loss on species with very small ranges, in particular in the Andes, Mesoamerica, and the
Caribbean islands, and also the very severe impact of enigmatic declines that are probably
due to the synergistic effects of the pathogenic chytrid fungus and climate change, especially
through the higher elevations of the region.
There have been seven recorded recent extinctions of amphibians in the Neotropical
Realm (21% of the global total): Atelopus ignescens (the Jambato Toad from Ecuador); Ate-
lopus longirostris (Ecuador); Atelopus vogli (Venezuela); Bufo periglenes (the famous Golden
Toad from Monteverde, Costa Rica); Phrynomedusa fimbriata (southern Brazil); Craugastor
chrysozetetes (Honduras); and Craugastor milesi (Honduras). With the exception of the
Golden Toad, all of these were stream-associated species that occurred at middle to high
elevations (above 700m asl) – the typical ecological profile of species that have experienced
rapid declines (Lips et al. 2003; Ron et al. 2003; Burrowes et al. 2004; Stuart et al. 2004; La
Marca et al. 2005). Some additional undescribed species are possibly extinct, especially in
the genus Atelopus (see Pounds et al. [2006] for details).
In addition, 121 Critically Endangered species in the Neotropics are considered possibly
extinct. This represents 93% of the 130 possibly extinct species in the world, thus dramati-
cally highlighting the extinction crisis that has unfolded with the Neotropical amphibians.
Most of the Critically Endangered (Possibly Extinct) species (listed in Table 1) share the
same ecological characteristics as those that have gone extinct. Of the 121 possibly extinct
species, 22 are harlequin toads in the genus Atelopus (representing 29% of the described
species in the genus), which have experienced catastrophic declines, especially in southern
Mesoamerica, and in the Andes south at least to Peru (La Marca et al. 2005; Pounds et al.
2006; and see Essay 9.1). Four genera concentrated in Mesoamerica also have large propor-
tions of possibly extinct species: Isthmohyla (36%), Plectrohyla (20%), Craugastor (14%)
and Thorius (17%) (see, for example, Lips et al. [2004, 2006] and Mendelson et al. [2004]).
In each of these cases, further work is likely to show that the percentage of possibly extinct
species has been underestimated. Another genus for which the percentage of possibly extinct
species might have been underestimated is Cycloramphus from southern Brazil (see Eterovick
et al. [2005] and Heyer et al. [1988] for more details). The genus Telmatobius, which occurs
in the Andes from Ecuador southwards, is also subject to extensive disappearances, but
much of this information is only just now becoming available, and for the most part is not
yet included in these results (De la Riva 2005; and see Essay 9.2). Possibly extinct species
range very widely in the Neotropics, generally in mountainous regions from southern Mexico
(for example, Plectrohyla cyanomma) south to Chile (for example, Rhinoderma rufum). A
number of recent declines and possible extinctions in Colombia have come to light since
the GAA data were collected (F. Castro pers. obs.).
The percentage of DD species is very similar to the global average of 23%. As mentioned
above, many of these DD species are likely to be threatened, but many others could be
LC, especially those that occur in poorly surveyed low-lying areas, such as in parts of the
Amazon basin, and the Cerrado of Brazil.
SPECIES RICHNESS AND ENDEMISM
Species Richness and Endemism Across Taxa
Of the 2,916 native amphibian species in the Neotropical Realm, 2,808 (or 96%) are endemic
to the Neotropics (Table 2). All three orders of amphibians are represented in the Neotropical
Realm. The overwhelming majority of Neotropical amphibians (89%) are frogs and toads
(Anura), 97% of which are endemic. All species of Neotropical caecilians (Gymnophiona),
and 91% of Neotropical salamanders (Caudata), are endemic. Only 392 species (13%) are
members of families that are endemic to the region, but this low percentage is really a
reflection of the fact that the very large family Leptodactylidae (accounting for 42% of
Neotropical amphibian species) marginally occurs in the Nearctic Region.
The Strawberry Poison Frog Dendrobates
pumilio (Least Concern) is a poison frog from
the Family Dendrobatidae, a Family famous
for the stunning coloration of its species. This
species lives on the floor of lowland rainforest
in Nicaragua, Costa Rica and Panama. The
females lay egg clutches on the forest floor,
and carry the larvae, after hatching, to water-
filled bromeliads, where they complete their
development. © Piotr Naskrecki
The Burrowing Toad Rhinophrynus dorsalis
(Least Concern) is the only member of the
Family Rhinophrynidae. It is a lowland species
ranging from southern Texas to Costa Rica. It
can be found in forest, thorn scrub, savan-
nah, and cultivated areas with friable soils.
It is usually subterranean, except after heavy
rains, when it emerges to breed explosively in
temporary pools. © Paddy Ryan
Under current climatic conditions, there is less isolation between the Neotropical and Ne-
arctic Regions than there is between the Afrotropical and Palaearctic Regions, and there are
points of contact between the two faunas along the Caribbean coast of Mexico, and Florida
(although the transvolcanic belt in central Mexico does form a barrier to faunal dispersal).
The result of this indistinct boundary is to reduce the level of endemism of each region.
The families Leptodactylidae and Rhinophrynidae are nearly endemic to the Neotropics, and
Scaphiopodidae, Ambystomatidae, Amphiumidae and Sirenidae are almost endemic to the
Nearctic. Salamandridae is also a northern element that is only marginally present in the
Neotropics. Of the 20 families that are native to the region, six are endemic. Amphibian
family-level diversity is higher than in any other biogeographic realm, but endemism is lower
than in the Afrotropics (where there are nine endemic families) because of the relative lack
of isolation. From the perspective of amphibian biogeography, the region is almost defined
by the distribution of the Neotropical frogs (family Leptodactylidae), which are present
through nearly all of Mexico, Central America, South America, and the Caribbean islands.
Summaries of the amphibian fauna of the Neotropics are provided by Campbell (1999a),
Duellman (1999) and Hedges (1999).
There are 189 genera occurring in the region (41% of the global total), of which 157 are
also endemic. These endemic genera represent over one-third (34%) of the 460 amphibian
genera worldwide. The Neotropics, therefore, account for a larger proportion of the overall
diversity of amphibians at the species level than at the generic level. The most species-rich
genus in the region is Eleutherodactylus (607 species, and 715 if the genus Craugastor is
included within it, contra Crawford and Smith [2005]). At the opposite end of the spectrum,
there are 46 monotypic genera endemic to the Neotropical Realm, which equates to just
over one-third (33%) of the 126 monotypic genera of amphibians worldwide. Interestingly,
nine of these monotypic genera are in the family Microhylidae, which is not particularly
diverse in the region. The 32 non-endemic genera in the Neotropics include 13 genera from
Hylidae, four from Plethodontidae, three from Leptodactylidae, two each from Microhylidae,
Scaphiopodidae and Sirenidae, and one each from Bufonidae, Ranidae, Rhinophrynidae,
Ambystomatidae, Amphiumidae and Salamandridae. These non-endemics include the
widespread genera Bufo and Rana.
Of the 20 amphibian families that occur in the Neotropics (42% of the global total), six
are endemic to the region: Allophrynidae, Brachycephalidae, Centrolenidae, Dendrobatidae,
Rhinodermatidae, and Rhinatrematidae.3 The characteristics of these families are provided
in Chapter 1.
Among the non-endemic families, the majority of Neotropical species are in Bufonidae,
Hylidae, Leptodactylidae and Microhylidae. Of the Neotropical Bufonidae, 121 species
(47% of those occurring in the region) are within the widespread genus Bufo. There are 77 Table 2. The number of Neotropical amphib-
ians in each taxonomic Family present in
the region.

Family Native species Percentage of Percentage of Native genera Percentage of Percentage of

(endemics to species in region species in family (endemics to genera in region genera in family
region) that are endemic that are endemic region) that are endemic that are endemic
to region to region
Anura
Allophrynidae 1 (1) 100 100 1 (1) 100 100
Brachycephalidae 8 (8) 100 100 1 (1) 100 100
Bufonidae 256 (240) 94 50 13 (12) 92 35
Centrolenidae 138 (138) 100 100 3 (3) 100 100
Dendrobatidae 234 (234) 100 100 9 (9) 100 100
Hylidae 610 (585) 96 73 46 (33) 72 67
Leptodactylidae 1,235 (1,215) 98 98 55 (52) 95 95
Microhylidae 56 (52) 93 12 19 (17) 89 25
Pipidae 7 (7) 100 23 1 (1) 100 14
Ranidae 34 (17) 50 3 1 (0) 0 0
Rhinodermatidae 2 (2) 100 100 1 (1) 100 100
Rhinophrynidae 1 (0) 0 0 1 (0) 0 0
Scaphiopodidae 3 (0) 0 0 2 (0) 0 0
TOTAL ANURA 2,585 (2,499) 97 48 153 (130) 85 36
Caudata
Ambystomatidae 15 (9) 60 30 1 (0) 0 0
Amphiumidae 1 (0) 0 0 1 (0) 0 0
Plethodontidae 221 (210) 95 58 14 (10) 71 34
Salamandridae 2 (0) 0 0 1 (0) 0 0
Sirenidae 2 (0) 0 0 2 (0) 0 0
TOTAL CAUDATA 241 (219) 91 41 19 (10) 53 16
Gymnophiona
Caeciliidae 81 (81) 100 72 15 (15) 100 58
Rhinatrematidae 9 (9) 100 100 2 (2) 100 100
Figure 2. The species richness of amphibians in the Neotropical Realm, with darker colours TOTAL GYMNOPHIONA 90 (90) 100 52 17 (17) 100 52
corresponding to regions of higher richness. Colour scale based on 10 quantile classes; TOTAL ALL AMPHIBIANS 2,916 (2,808) 96 47 189 (157) 83 34
maximum richness equals 144 species.
94 Threatened Amphibians of the World

species in Atelopus, 19 in Melanophryniscus, and 10 in Rhamphopryne, but the remaining

Family EX CR EN VU NT LC DD Total number Number % Threatened nine genera have small numbers of species. Most of the Neotropical species breed by larval
of species Threatened or Extinct development, but breeding (where it is known) is by direct development in Metaphryniscus,
or Extinct Oreophrynella and Osornophryne and unknown in Andinophryne, Crepidophryne, Rhampho-
Anura pryne and Truebella. The family is widely distributed through most of the Neotropics, with
Allophrynidae 0 0 0 0 0 1 0 1 0 0 highest species richness in the equatorial regions.
Brachycephalidae 0 0 0 1 1 1 5 8 1 13 Hylidae is overwhelmingly a Neotropical family, the main radiation outside the region
Bufonidae 4 73 29 30 10 77 33 256 136 53 occurring in Australia and New Guinea. At the species level, 73% of the family (585
Centrolenidae 0 6 16 29 10 28 49 138 51 37 species) is endemic to the Neotropics, where it occurs widely through most of the region
Dendrobatidae 0 20 29 16 14 58 97 234 65 28 (excluding Chile), with especially high species richness in Brazil (over 300 species, with
Hylidae 1 65 58 35 24 312 115 610 159 26 very high diversity in the Atlantic Forest). There are over 160 species in Mesoamerica. Two
Leptodactylidae 2 145 246 172 61 350 259 1,235 565 46 subfamilies, Phyllomedusinae and Hylinae, occur in the Neotropics (and see Essay 1.5).
Microhylidae 0 0 2 5 2 37 10 56 7 13 The genera have recently been extensively revised (Faivovich et al. 2005), and under this
Pipidae 0 0 1 0 0 6 0 7 1 14 new arrangement, the genera Dendropsophus, Scinax, Hypsiboas, Plectrohyla, Hyloscirtus,
Ranidae 0 5 2 7 4 14 2 34 14 41 Phyllomedusa, Hyla and Bokermannohyla all have more than 20 species. The Neotropical
Rhinodermatidae 0 1 0 1 0 0 0 2 2 100 hylids are associated with many different habitats, but species richness is highest in forests,
Rhinophrynidae 0 0 0 0 0 1 0 1 0 0 and all known breeding is by larval development.
Scaphiopodidae 0 0 0 0 0 3 0 3 0 0 The family Leptodactylidae, which is almost endemic to the Neotropics, is by far the
TOTAL ANURA 7 315 383 296 126 888 570 2,585 1,001 39 largest family of amphibians worldwide. It ranges widely throughout the region, with the
Caudata highest species richness in the tropical Andes from Venezuela and Colombia south to Bolivia
Ambystomatidae 0 8 2 0 0 3 2 15 10 67 (over 550 species), with significant diversity in Brazil (nearly 300 species), Mesoamerica (c.
Amphiumidae 0 0 0 0 0 1 0 1 0 0 160 species), the Caribbean islands (c. 160 species) and the Southern Cone (c. 130 species).
Plethodontidae 0 35 70 28 14 28 46 221 133 60 The family includes the largest genus of vertebrates, Eleutherodactylus (607 species) with the
Salamandridae 0 0 1 0 0 1 0 2 1 50 following genera including more than 20 species: Craugastor, Leptodactylus, Gastrotheca,
Sirenidae 0 0 0 0 0 2 0 2 0 0 Telmatobius, Physalaemus, Phrynopus, Cycloramphus, and Hylodes. The family includes
TOTAL CAUDATA 0 43 73 28 14 35 48 240 144 60 species that breed by direct development and larval development. Although these frogs
Gymnophiona occur in many habitats, species richness is highest in forests.4
Caeciliidae 0 0 0 0 0 29 52 81 0 0 The Neotropical Microhylidae species are widely distributed within the region, from
Rhinatrematidae 0 0 0 0 0 4 5 9 0 0 Mexico south to central Argentina, but not on the Caribbean islands (except Trinidad). With
TOTAL GYMNOPHIONA 0 0 0 0 0 33 57 90 0 0 the exception of Chiasmocleis (19 species), all genera are small. Most species occur at low
TOTAL ALL AMPHIBIANS 7 358 456 324 140 956 675 2,915 1,145 39 elevations, with highest species richness in the equatorial regions. In the Neotropics, all
species breed by larval development, and many are subterranean when not breeding.
The Neotropical Ranidae species (all of which are in the widespread genus Rana) are all
Table 3. The number of species within each larval developers, and occur predominantly in Mesoamerica, with three species reaching South
IUCN Red List Category in each Family and America, and none on the Caribbean islands (except Trinidad). Of the remaining families, the
Order in the Neotropical Realm. Introduced highly aquatic Pipidae, with a single genus in the region, Pipa, ranges from Panama south
species are not included. to Bolivia. Scaphiopodidae, Amphiumidae, Salamandridae and Sirenidae are Nearctic taxa
that only marginally occur in the northern Neotropics. Ambystomatidae (mole salamanders)
is also a Nearctic element, but there is an important radiation of species in central Mexico,
including the famous Axolotl Ambystoma mexicanum (CR). Ambystomatidae breed by larval
development, and some retain their aquatic larval features throughout their life cycles.
Most Neotropical salamanders are in the large family Plethodontidae (lungless salaman-
ders). This family has its highest species richness in the Nearctic, but there are c. 110 species
in Mexico, c. 40 in Guatemala, c. 25 in Honduras, c. 40 in Costa Rica, c. 25 in Panama, but
only 28 in the whole of South America (12 of these being endemic to Colombia). The largest
Neotropical genus is Bolitoglossa (91 species), with Pseudoeurycea, Oedipina and Thorius
each having more than 20 species. Lungless salamanders occur as far south as Bolivia. With
the possible (but even then unlikely) exception of a single species (Pseudoeurycea aquatica),
all Neotropical species breed by direct development, and almost all are associated with
forest habitats (and see Essay 9.3).
The caecilian family Caeciliidae is very poorly known in the Neotropics, as in other parts
of the world. A total of 81 species (in 15 genera) is recorded from the region, comprising 72%
of the family at the species level. Species richness is highest in the Amazon Basin, with only
16 species in Mesoamerica (as far north as southern Mexico), and four species reaching
northern Argentina. The species exhibit a wide variety of reproductive modes, from larval
and direct development, to live-bearing. The majority are subterranean species in the forest
floor, but certain species (in the genera Atretochoana, Potomotyphlus and Typhlonectes) are
aquatic, sometimes referred to as “rubber eels”.
Not surprisingly, the larger families – Bufonidae, Centrolenidae, Dendrobatidae, Hylidae,
Leptodactylidae and Plethodontidae – have the largest absolute numbers of globally threat-
ened species (Table 3). The percentage of threatened species ranges greatly between the
families, from zero for the Allophrynidae, Rhinophrynidae, Scaphiopodidae, Amphiumidae,
a) Sirenidae, Caeciliidae, and Rhinatrematidae, to 100% for the endemic Rhinodermatidae of
Chile and Argentina. The zero percentages of threatened species in the caecilian families is
probably because these species are so poorly known (with >50% of species Data Deficient
in both the Caeciliidae and Rhinatrematidae). The percentages of threatened species are
also very high in the salamander families Ambystomatidae (67% - reflecting the serious
conservation problems on the Mexican plateau) and Plethodontidae (60% - reflecting the poor
state of forest conservation in the pine-oak regions of Mexico, Guatemala, and Honduras).
The percentage levels of threat are also high (>30%) in the Bufonidae (53% - reflecting, in
part, the devastating declines in the genus Atelopus), Leptodactylidae (46% - with several
genera, including Craugastor, Eleutherodactylus and Telmatobius showing high levels of
threat), Ranidae (41%), and Centrolenidae (37%).
Some of the larger families have more species in the Endangered category than in
Critically Endangered or Vulnerable (e.g., Dendrobatidae, Leptodactylidae, Plethodontidae).
However, Centrolenidae show a similar pattern to birds and mammals (Baillie et al. 2004),
with least in Critically Endangered and most in Vulnerable. Conversely, Bufonidae and Hylidae
have most in Critically Endangered and least in Vulnerable, showing how severely impacted
these families have been by recent dramatic declines (indeed, 55% of threatened Neotropi-
cal Bufonidae are Critically Endangered). Among the smaller families, there is also a high
percentage of Critically Endangered species in Rhinodermatidae and Ambystomatidae.

Geographic Patterns of Species Richness and

Figure 3. a) The richness of threatened
amphibians in the Neotropical Realm, with
darker colours corresponding to regions of
higher richness. Colour scale based on 10
quantile classes; maximum richness equals
42 species. b) The richness of CR amphibians
in the Neotropical Realm. Colour scale based
on five quantile classes; maximum richness
equals 18 species.
Endemism
A map of overall species richness of amphibians in the Neotropical Realm (Figure 2), shows
that species richness is highest in the tropical regions, notably Costa Rica and Panama,
the Pacific lowlands of western Colombia and north-western Ecuador, the Guianan Shield,
the Atlantic Forest of southern Brazil, and in particular in the Amazon Basin (especially in
the west). Species richness is lowest in more temperate regions (Mexico, Argentina and
Chile), on the Caribbean islands, and especially in arid regions, such as northern Mexico
and northern Chile (there being no amphibians at all in most of the Atacama Desert region).
The higher taxon diversity is particularly low in the Caribbean islands, where there are just
four families and five genera (compared with 15 families and 67 genera in Mesoamerica).
b) The vast majority (88%) of the Caribbean’s amphibian species belong to just one genus,
Eleutherodactylus. No salamanders or caecilians occur on these islands.
 Chapter 9. Amphibians of the Neotropical Realm 95

Brazil Jamaica Colombia Haiti

Colombia Cuba Mexico* Dominican Republic
Ecuador
Peru Puerto Rico Ecuador Jamaica
Mexico* Mexico* Brazil Cuba
Venezuela Chile Peru Puerto Rico
Bolivia Guatemala Mexico*
Panama Brazil
Costa Rica Venezuela Venezuela Guatemala
Argentina Haiti Costa Rica Dominica
Guatemala Honduras Guadeloupe
Guyana Guadeloupe
Colombia Panama Montserrat
Honduras
French Guiana Peru Cuba Honduras
Suriname United States of America* Haiti Virgin Islands (British)
Paraguay Dominican Republic Ecuador
Nicaragua Ecuador
Argentina Chile
Cuba Honduras
Chile Bolivia Costa Rica
Dominican Republic
Haiti Chile Grenada
United States of America* Grenada
Jamaica Martinique
Uruguay Martinique
Belize Puerto Rico St Vincent and the Grenadines
St Vincent and the Grenadines
Dominican Republic Nicaragua Colombia
Argentina
Trinidad and Tobago El Salvador Panama
El Salvador Guatemala
Trinidad and Tobago El Salvador
Jamaica Bolivia
Puerto Rico Belize Trinidad and Tobago
Dominica Guyana Virgin Islands (US)
Virgin Islands (British)
Dominica Virgin Islands (US) Uruguay Venezuela
Guadeloupe Costa Rica French Guiana Peru
Virgin Islands (US) Virgin Islands (British)
Grenada Dominica Argentina
Martinique Trinidad and Tobago Guadeloupe Belize
St Vincent and the Grenadines Panama Suriname Brazil
Antigua and Barbuda Guyana United States of America* Nicaragua
Bahamas Suriname
Cayman Islands Virgin Islands (British) Bolivia
Montserrat French Guiana Grenada Uruguay
Anguilla Uruguay Martinique Guyana
Barbados Nicaragua Montserrat United States of America*
Netherlands Antilles
St Kitts and Nevis Paraguay St Vincent and the Grenadines French Guiana
St Lucia Belize Virgin Islands (US) Suriname
0 200 400 600 800 0 20 40 60 80 100 0 50 100 150 200 250 0 20 40 60 80 100
Number of species Percentage of endemic species Number of threatened species Percentage of species threatened
Endemics Non-endemics Endemics Non-endemics
Figure 4 Figure 5 Figure 6 Figure 7

Figure 4. The number of amphibians present

As with other parts of the tropics, Figure 2 probably does not reflect genuine patterns
of amphibian species richness everywhere in the region, due to uneven survey effort.
In particular, species richness is probably under-sampled in the Guianan Shield, in the
Venezuelan, Colombian and Bolivian Amazon, in the Peruvian and Bolivian Andes, and in
the Cerrado of Brazil. However, sampling appears to be less uneven than in the Old World
tropics (similar maps for the Afrotropical, Indomalayan and Australasian Realms make
much less overall biogeographic sense). New species continue to be discovered at a rapid
rate almost everywhere in the Neotropics, but nevertheless the overall patterns of species
richness are probably reasonably clear.
Over 85% of the threatened amphibian species in the Neotropics occur in the region
from southern Mexico to Ecuador and northern Venezuela, and on the Greater Antilles
(Cuba, Hispaniola, Jamaica and Puerto Rico) (see Figure 3a). This region represents by far
the greatest concentration of threatened amphibian species anywhere in the world. Within
this region there are peaks of threatened species in montane areas in southern Mexico,
Guatemala, Honduras, Costa Rica, western Panama, Cuba, Hispaniola, Jamaica, Puerto
Rico, and the Andes of Venezuela, Colombia and Ecuador. Outside this region, the largest
concentration of threatened amphibian species in the Neotropics is in the Atlantic Forests
of southern Brazil. There are lesser concentrations in the Peruvian and Bolivian Andes, and
in the austral forest zone of Chile. It is possible that, due to the poor state of knowledge,
the levels of threat have been under-estimated in the Peruvian and Bolivian Andes; further,
with very recently recorded declines in Telmatobius species (De la Riva 2005), and probably
in other species too, not yet included in our data, it is likely that a new concentration of
threatened species will soon be identified in the Andes running from Peru south to Bolivia,
Chile, and Argentina. These concentrations of threatened species correlate with those for
other taxa (Baillie et al. 2004). These geographic concentrations reflect the topographi-
cally diverse (usually montane) parts of the region where amphibians have naturally small
ranges, and where habitat destruction is ongoing (and in central and southern Peru is in
part a reflection of the over-harvesting of some frogs (e.g., in the genus Telmatobius) for
human consumption). However, these are also the places where rapid population declines
and disappearances have been noted due to chytridiomycosis and climate change (Heyer
et al. 1988; La Marca and Reinthaler 1991; Young et al. 2001; Lips et al. 2003, 2004, 2006;
Ron et al. 2003; Burrowes et al. 2004; Mendelson et al. 2004; De la Riva 2005; Eterovick et
al. 2005; La Marca et al. 2005; Pounds et al. 2006).
The concentrations of Critically Endangered species (Figure 3b) broadly match those
of threatened species as a whole. The greatest concentrations of these most severely
threatened species are in southern Mexico (in particular Veracruz and Oaxaca), Guatemala,
Honduras, Costa Rica, western Panama, the Ecuadorian Andes, and Haiti (especially the
Massif de la Hotte and Massif de la Selle). Lesser concentrations of Critically Endangered
species are found in the Andes of Venezuela, Colombia, Peru, southern Brazil, central Chile,
Puerto Rico, Dominican Republic, Jamaica, and eastern Cuba (although recent data from the
Colombian Andes suggest that the next update of the GAA might reveal this region to be a
major concentration of Critically Endangered species (F. Castro pers. obs.).
Mexico (Flores-Villela 1993; Flores-Villela et al. 1995; Calderon Mandujano et al. 2005); in and endemic to each Neotropical country.
Yucatán (Campbell 1999b; Lee 1996, 2000); Guatemala (Lee 2000, Campbell 1999b, 2001); *denotes countries not entirely within the
Belize (Campbell 1999b, Lee 2000); Honduras (McCranie and Wilson 2002; McCranie et al. Neotropical Realm, hence only the species
2006); El Salvador (Köhler et al. 2005); Nicaragua (Köhler 2001); Costa Rica (Savage 2002; whose ranges fall within the region are
Guyer and Donnelly 2005); Panama (Ibáñez et al. 1999, 2000); Colombia (Ruiz-Carranza et al. included.
1996); Venezuela (La Marca 1992, 1997; Barrio Amorós 2004); Ecuador (Coloma 2005); Peru
(Lehr 2002); Bolivia (De la Riva et al. 2000, Köhler 2000); Argentina (Cei 1980, 1987; Lavilla et
al. 2000; Lavilla and Cei 2001); Chile (Veloso and Navarro 1988; Formas 1995); Brazil (Sociedade Figure 5. Percentage of species endemic to
Brasileira de Herpetologia 2004); the Guianas (Hoogmoed 1979); the Guianas (Señaris and each Neotropical country. *denotes countries
MacCulloch 2005); French Guiana (Lescure and Marty 2000); the Caribbean islands (Crother not entirely within the Neotropical Realm,
1999; Schwartz and Henderson 1988; Powell and Henderson 1999); the Lesser Antilles hence only the species whose ranges fall
(Malhotra and Thorpe 1999); Guadeloupe (Breuil 2002); Netherlands Antilles (Powell et al. within the region are included.
2005; van Buurt 2005); and Trinidad and Tobago (Murphy 1997). There have also been some
important reviews of particular taxonomic groups, for example on the Hylidae of Mesoamerica
(Duellman 2001), on the western Ecuadorian Eleutherodactylus (Lynch and Duellman 1997), Figure 6. The number of threatened amphib-
and Neotropical plethodontid salamanders (Wake and Lynch 1976; Wake 2003). ians present in and endemic to each Neotropi-
Brazil has more endemic species (489) than any other country in the Neotropics (Figure cal country. Countries with no threatened spe-
4), or in the world, followed by Colombia (337), Mexico (234), Peru (181), Ecuador (159), and cies are not included in the diagram. *denotes
Venezuela (155). More than 50 endemic species are also known from Cuba and Bolivia. In countries not entirely within the Neotropical
terms of percentage of the fauna being endemic, the highest endemism is on the Greater Realm, hence only the species whose ranges
Antilles (Figure 5), with Jamaica at 100%, Cuba at 97% and Puerto Rico at 78%. Although fall within the region are included.
the percentage endemism in Haiti and the Dominican Republic is lower, for the island of
Hispaniola as a whole it is 100%. On the mainland the highest percentage endemism is
found in Mexico (70%), Chile (69%), and Brazil (65%), with levels over 40% in Venezuela, Figure 7. Percentage of native species that are
Haiti, Guadeloupe, Colombia, and Peru (Figure 5). threatened. Countries with no threatened spe-
Threatened species occur in 35 of the 44 countries in which there are native amphibians cies are not included in the diagram. *denotes
(Figure 6). In fact, threatened species are concentrated in relatively few countries. Colombia countries not entirely within the Neotropical
has more threatened amphibian species than any other country in the Neotropics (209), Realm, hence only the species whose ranges
followed by Mexico (190), Ecuador (163) and Brazil (110). A further 12 countries have 20 fall within the region are included.
or more threatened species: Peru, Guatemala, Venezuela, Costa Rica, Panama, Honduras,
Cuba, Haiti, Dominican Republic, Argentina, Bolivia and Chile. The percentage of threatened
amphibian species is highest in the Greater Antilles (Figure 7), with Haiti at a staggering
92%, the Dominican Republic at 86%, Jamaica at 81%, Cuba at 80% and Puerto Rico at
72%. Overall, the percentage threat levels for amphibians on the Caribbean islands are
worse than anywhere else in the world, and is a reflection of the very poor state of habitat
conservation, coupled with chytridiomycosis in some places (see Essay 9.6) (of course, in
these relatively species-poor countries, even a limited number of threatened species can
result in a high percentage of species at risk of extinction). The highest percentage of
threatened species on the mainland is in Mexico (57%), closely followed by Guatemala
(55%), with a further 12 countries having levels greater than 30%: Guadeloupe, Dominica,

Species Richness and Endemism within Countries

Amphibians occur naturally in every mainland country in Mesoamerica and South America,
and on all but the smallest Caribbean islands (Figure 4). However, only one extant species
occurs naturally in St. Lucia, St. Kitts and Nevis, the Netherlands Antilles, Barbados and
Anguilla, and only two on Antigua and Barbuda, the Bahamas, the Cayman Islands and
Montserrat. There are no indigenous amphibians on the Galapagos Islands.
The two countries with the largest number of species in the Neotropical Realm are Brazil
(751 species; and see Essay 9.4) and Colombia (697 species; see Essay 9.5). There is also
very high species richness in Ecuador (447 species), Peru (411), Mexico (336), Venezuela (298)
and Bolivia (209). Brazil, Colombia, Ecuador, Peru, and Mexico are the top five countries in Rhinatrema bivittatum (Least Concern) is a
the world in terms of amphibian species richness. Another eight Neotropical countries have caecilian from the small Family Rhinatremati-
more than 100 species (Panama – 195, Costa Rica – 179, Argentina – 157, Guatemala – 138, dae, occurring in Brazil, Guyana, Suriname and
Guyana – 118, Honduras – 116, French Guiana – 104, Suriname – 103). French Guiana. It is a subterranean species in
These figures are, of course, a reflection of current knowledge, and as mentioned earlier, lowland rainforest, and is presumed to breed
certain regions and countries have been better studied than others. In certain places, the in streams by larval development, like other
existing knowledge has been well summarized in review literature, and in books, including: members of its Family. © Peter Stafford
96 Threatened Amphibians of the World

Habitat type Number of % of all Threatened % of species

species in species or Extinct occurring
each habitat occurring in species in habitat that
the habitat are Threatened
or Extinct
Forest 2,478 85 1,029 42
All tropical forest 2,407 83 1,007 42
Lowland tropical forest 1,405 48 427 30
Montane tropical forest 1,494 51 773 52
Savannah 200 7 6 3
Grassland 429 15 120 28
Shrubland 343 12 73 21
Secondary terrestrial habitats 621 21 110 18
Flowing freshwater 1,133 39 505 45
Marsh/swamp 168 6 23 14
Still open freshwater 746 26 116 16
Arid and semi-arid habitats 25 1 5 21

Table 4. The habitat preferences of amphibians in the Neotropical Realm.

Number of species % of species in habitat that are threatened

Forest

Savannah

Grassland

Shrubland

Secondary terrestrial habitats

Flowing freshwater

The Upland Coqui Eleutherodactylus portori-
censis (Endangered) is one of more than 600
species in the genus Eleutherodactylus in the
Family Leptodactylidae. It occurs in mesic,
upland broadleaf forests, and calls from
bushes and tree trunks, and has not been
recorded outside forest habitat. The eggs
are laid in bromeliads, and these develop
directly without a free-living larval stage.
© Alejandro Sanchez
Montserrat, Honduras, British Virgin Islands, Ecuador, Chile, Costa Rica, St Vincent and the
Grenadines, Martinique, Grenada, and Colombia (Figure 7). Overall, percentage threat levels
are high in the Neotropics compared with the rest of the world, but the overall levels of
threat do appear to be lower in the Guianan Shield and Amazonian Brazil.
Assessments of the conservation status of Neotropical amphibians at national level are
still at an early stage, but there have been assessments in El Salvador (Greenbaum and
Komar 2005), Panama (Young et al. 1999), Venezuela (Rodríguez and Rojas-Suárez 1995),
Brazil (Ministério do Meio Ambiente 2003), Argentina (Lavilla et al. 2000), Chile (Glade 1993)
and Bolivia (Reichle 2006). An official Mexican red list of amphibians is in preparation (the
draft is on http://www.semarnat.gob.mx/leyesynormas/Normas%20Oficiales%20Mexic
anas%20vigentes/NOM-ECOL-059-2001.pdf. Hedges (2006) provided an overview of the
conservation of Caribbean amphibians.
Some countries have particularly high proportions of Critically Endangered species. The
most extreme example is Haiti, where 31 species are CR, 10 are EN and five are VU (out of a
total amphibian fauna of 50 species). In the neighbouring Dominican Republic, the situation
is marginally less severe (10 CR, 16 EN, 5 VU out of 36 species), though still very serious.
Puerto Rico has 7 CR, 5 EN and 1 VU out of a fauna of 18 species. On the mainland, things are
particularly bad in Honduras (30 CR, 24 EN, 19 VU out of 116 species), and also very disturbing
in Chile (9 CR, 4 EN, 7 VU out of 55), Mexico (69 CR, 80 EN, 41 VU out of 335), Guatemala (27
CR, 30 EN, 19 VU out of 140), and Costa Rica (19 CR, 22 EN, 20 VU out of 179).
In general, the levels of threat are worse in Mesoamerica than South America, because
habitat loss has in general been more severe in the former, and also chytridiomycosis has
been especially severe in this region (and see Essay 9.7). The situation is also serious and
deteriorating in the Andean countries, where there is also significant habitat loss, and
chytridiomycosis is currently spreading (De la Riva 2005; La Marca et al. 2005). However,
because most of the Andean countries also have large, intact Amazonian amphibian faunas,
the percentage of threatened species is not usually as high as in some of the Mesoamerican
countries. The percentage of threatened species in Peru and Bolivia is almost certainly
underestimated due to paucity in knowledge.
HABITAT AND ECOLOGY
Habitat Preferences
Most Neotropical amphibians (85%) occur in forests, and only just over 20% can survive
in secondary terrestrial habitats (Table 4; Figure 8). Compared with Afrotropical species,
for example, Neotropical amphibians appear to be less able to survive in disturbed areas.
They also appear to make more use of flowing water habitats than still, open freshwater
habitats, or marshes and swamps. This is presumably a reflection of the great diversity of
stream-associated species in the Andes and Mesoamerica. Forest-dwelling amphibians
are more likely to be threatened than those occurring in any other terrestrial habitats, with
over 40% of them being globally threatened. A similar percentage of amphibians associated
with flowing water (generally streams) is threatened. Forest-associated amphibians that
live along streams are particularly likely to be threatened, a combination that has also been
associated with rapid declines worldwide (Stuart et al. 2004).
The percentage of threatened species varies considerably between different types of
forest. In montane tropical forest, over 50% of known species are threatened, compared with
just over 30% in lowland tropical forest. These figures probably reflect smaller range sizes of
montane species, the lack of effective habitat conservation measures in many mountainous
parts of the region, and the high incidence of chytridiomycosis in montane areas (Lips et
al. 2003; Burrowes et al. 2004). Amphibians occurring in savannahs, marshes and swamps,
still open freshwater habitats, and secondary terrestrial habitats are much less likely to be
threatened than those occurring in other habitats (Table 4; Figure 8).
Reproductive Modes
Of those species where reproduction is known or reasonably inferred, larval development is
the most common reproductive mode in the Neotropics (59% of species), compared with 38%
for direct development and 1% live-bearing (this compares with the global picture of 68%
larval development, 30% direct development, and 1% live-bearing) (Table 5). The Neotropical
Marsh/swamp
Still open freshwater
Arid and semi-arid habitats
0 800 1,600 2,400 0 15 30 45
Figure 8. The habitat preferences of Neotropical amphibians. The plot on the left-hand side
shows the number of species in the region in each habitat type. On the right-hand side, the
percentage of these species which are threatened is given.
Reproductive mode All species Threatened or % Threatened
Extinct species or Extinct
Direct development 1,105 584 53
Larval development 1,719 555 32
Live-bearing 24 1 4
Not known 68 5 7
Table 5. Neotropical amphibians categorized by reproductive mode.
amphibians clearly have a larger proportion of direct-developing species than the global
average, and this is largely because of the enormous genus Eleutherodactylus (607 species),
all but one of which (the possibly extinct Eleutherodactylus jasperi from Puerto Rico) are
believed to be direct developers, but also because of other large genera such as Craugastor
and Phrynopus. In addition, all but one of the 221 Neotropical plethodontid salamanders are
believed to be direct developers. Although live-bearing is uncommon, the Neotropics account
for 39% of the world’s known live-bearing amphibians (all but one of these live-bearing
species are caecilians, with the exception of the aforementioned E. jasperi).
In the Neotropics, the percentage of globally threatened direct-developing amphibians
is much higher than for larval-developing species. This is probably because direct-develop-
ing species have smaller ranges on average, and are therefore more seriously impacted
by habitat loss. This result is interesting because chytridiomycosis appears to have its
greatest impact on stream-associated, usually larval-developing species (Lips et al. 2003)
(though it should be noted that some stream-associated species, such as the species in
the Craugastor rugulosus group, are direct-developers, and have been severely impacted by
chytridiomycosis). The low percentage of threatened live-bearing species in the Neotropics
could be a reflection of the high number of Data Deficient caecilians.
MAJOR THREATS
Habitat loss is overwhelmingly the major threat to amphibians in the Neotropics (Table 6;
Figure 9), affecting nearly 90% of the threatened species. The two other most commonly
recorded threats are pollution and disease (both affecting nearly 30% of threatened species).
With the exception of fire (17%), all other threats are of minor importance. Over-utilization
appears to be a minor threat in the region as a whole (at least, based on current knowledge),
but it can have a serious impact on some species (e.g., on the genus Telmatobius in Peru,
and probably elsewhere).
In terms of the types of habitat loss that are impacting amphibians in the Neotropics,
the impacts of expanding croplands (affecting just over 70% of threatened species) and
vegetation removal (mainly logging) (64%) are the most severe, but urbanization / industrial
development and livestock grazing are each affecting more than 40% of threatened species.
However, the importance of different types of habitat loss varies within the region. For
example, removal of vegetation for charcoal production is a major mechanism of habitat
loss in the Greater Antilles, especially in Haiti (Hedges 2006).
The distribution of chytridiomycosis in the Neotropics is only gradually becoming
clear. Ron (2005) documented confirmed records of the disease widely in Mesoamerica
(southern Mexico, Guatemala, Costa Rica, Panama), the Caribbean (Dominican Republic,
Puerto Rico), and north-western South America (Ecuador, Venezuela). More recently the
 Chapter 9. Amphibians of the Neotropical Realm 97

Threat type Threatened species % Threatened Species

Habitat loss 1,007 89
Agriculture – Crops 806 71
Agriculture – Tree plantations 143 13
Agriculture – Livestock 478 42
Timber and other vegetation removal 728 64
Urbanization and industrial development 538 47
Invasive species 81 7
Utilization 34 3
Accidental mortality 3 0.3
Pollution 336 29
Natural disasters 73 6
Disease 324 28
Human disturbance 101 9
Changes in native species dynamics 1 0.1
(excluding disease)
Fire 197 17

Table 6. The major threats to globally threatened amphibians in the Neotropical Realm.
Only present threats to species are tallied.

All habitat loss

Invasive species

Utilization

Accidental mortality

Persecution

Pollution

Natural disasters

Disease
The Titicaca Water Frog Telmatobius culeus
Human disturbance (Critically Endangered) in the Family Lepto-
Changes in native sp. dynamics Population Trend Number of species % of species dactylidae is endemic to Lake Titicaca in Peru
Decreasing 1313 45 and Bolivia. It is a wholly aquatic species,
Fire
Stable 725 25 breeding in shallow waters close to the
0 20 40 60 80 100 Increasing 15 0.5 shoreline. It was previously common, but has
Percentage of threatened species affected Unknown 856 29 declined massively due to the over-harvesting
of adults, the presumed predation of larvae
by introduced trout, water extraction from
Figure 9. The major threats impacting threatened amphibians in the Neotropical Realm. Table 8. The population trends for all extant Neotropical amphibians. the lake, and domestic and agricultural water
pollution. © Mikael Lundberg

Purpose Subsistence Sub-national/ Regional/ Number Family Number of Percentage Number of Percentage
National International of species species in of species species in of species
Food – human 41 (20) 16 (7) 5 (1) 44 (21) “rapid decline” in family in “enigmatic in family in
Food – animal 1 (0) 1 (1) 0 2 (1) “rapid decline” decline” “enigmatic
decline”
Medicine – human and 15 (12*) 5 (2) 4 (1) 19 (13*)
veterinary Bufonidae 89 35 76 30
Poisons 5 (0) 0 2 (0) 7 (0) Centrolenidae 4 3 4 3
Wearing apparel, accessories 1 (0) 0 0 1 (0) Dendrobatidae 17 7 11 5
Handicrafts, curios, etc. 2 (0) 1 (0) 1 (0) 2 (0) Hylidae 53 9 39 6
Pets, display animals 10 (2) 29 (8) 125 (38*) 132 (38*) Leptodactylidae 110 9 53 4
Research 0 6 (1) 6 (1) 11 (1) Ranidae 6 18 4 12
Specimen collecting 0 0 1 (1) 1 (1) Rhinodermatidae 1 50 1 50
Ambystomatidae 5 36 0 0 Table 9. The number of species in “rapid
Plethodontidae 20 9 6 3 decline” and “enigmatic decline” in the
Table 7. The purposes for which amphibians are used in the Neotropical Realm. The numbers Neotropical Realm by Family.
in brackets are the number of species within the total that are threatened species. *One of
the species in the brackets is actually now listed as Extinct.

“Rapidly declining” species

disease has been recorded more widely in the continent, for example in southern Peru
(Seimon et al. 2005), southern Brazil (Carnaval et al. 2005) and the Pampas region of
Argentina (Herrera et al. 2005). The earliest records of the fungus in the region date from
the early 1980s, and coincide roughly with the onset of amphibian declines (Carnaval et
al. 2006; Lips et al. 2006).
A total of 181 species are recorded as being used for some or other purpose in the region
(61 of which are threatened (though not necessarily by use) and one now considered Extinct).
The most common reason for harvesting Neotropical amphibians is for the international pet
trade, followed by local human consumption (Table 7). Well-known examples of utilization
in the region include the pet trade in colourful, poisonous frogs in the genera Dendrobates,
Epipedobates and Phyllobates, and the horned frogs (in the genus Ceratophrys), and
harvesting Telmatobius frogs for local human consumption in parts of the Andes. Much of
the harvesting of amphibians in the region is not considered to constitute a major threat
to the species, but there are exceptions (for example, in the case of several species of
Telmatobius). Of the 180 extant species being harvested, utilization is considered to be a
major threat for 71 (of which only 34 are threatened species for which harvesting is believed
to be contributing to deterioration in their status).
POPULATION STATUS AND TRENDS
Estimates of Population Trends
A summary of the inferred population trends of Neotropical amphibians is presented in Table
8. In the absence of more rigorous population monitoring studies, these trends are largely
inferred from trends in the state of the habitats on which the species depend (though in some
cases, dramatic population declines have been noted). Species with decreasing populations
are typically forest-dependent species that can tolerate little disturbance to their habitats.
The overall trends of Neotropical amphibians are very similar to the global results.
The Neotropics are home not only to nearly 50% of the world’s amphibian species, but to
nearly two-thirds (65%; 305 species) of the world’s “rapidly declining” species (Stuart et
al. 2004) (a full list of all “rapidly declining” species is provided in Appendix IV and includes
their occurrence within each of the regions). The Neotropics are the global epicentre for Flectonotus pygmaeus (Least Concern), in the
amphibians in catastrophic decline. Twelve of these 305 species are in decline due to over- Family Leptodactylidae, occurs in Venezuela
exploitation, 99 due to reduced habitat, and 194 due to so-called “enigmatic declines”, and Colombia. It is a species of pre-montane
which are currently attributed to chytridiomycosis and climate change (Lips et al. 2006; humid forests, and is particularly associated
Pounds et al. 2006). with bromeliads. The eggs are carried on the
The “rapidly declining” species show a distinct taxonomic pattern (Table 9). Among back of the female in a pouch, and the larvae
the larger families, Bufonidae show by far the highest percentage of species in serious are deposited in bromeliad axils. © Francisco
decline, and in particular in “enigmatic decline”. Most of this very serious situation can José López-López
be accounted for by the genus Atelopus (73 species in “rapid decline”, 72 in “enigmatic
decline”). There are many species in serious decline in Hylidae and Leptodactylidae, but
percentage wise these families are much less seriously affected than Bufonidae. However,
some genera seem to be particularly affected, notably Isthmohyla (6 out of 14 species in
serious decline), Plectrohyla (21 out of 41), Craugastor (22 out of 108), Telmatobius (14
out of 52), and Thoropa (3 out of 6). The 43 species of Eleutherodactylus (especially from
the Caribbean islands) in rapid decline should be seen in the context of a genus of 610
species. Some small families have high percentages of species in serious decline, most
notably Rhinodermatidae and Ambystomatidae. The two species in the Rhinodermatidae,
in particular, require comment. One of them, Darwin’s Frog Rhinoderma darwinii (VU) is
currently in “enigmatic decline”. The other, Rhinoderma rufum (CR), also has declined
enigmatically and was last seen in 1978; however, it is not recorded as a “rapidly declining”
species, as its population crashed prior to 1980, the year from which “rapid declines” have
been measured (Stuart et al. 2004).
The “rapid declines” in the Neotropics are concentrated in particular regions, most
especially in Mesoamerica (from central Mexico south to Panama, as typified by the
genera Isthmohyla, Plectrohyla and Craugastor) and the Andes (as typified by Atelopus and
98 Threatened Amphibians of the World
Bolitoglossa pesrubra (Endangered) is a lungless salamander from the Family Plethodonitidae, and is restricted to the Cordillera de Talamanca
in Costa Rica. It still occurs in many places within its range, but has drastically declined in some sites where it was formerly abundant, while
appearing to be stable in others. © Twan Leenders
Telmatobius). There have also been many declines in the Greater Antilles, for the most part
in the genus Eleutherodactylus. In addition, there are some smaller foci of “rapid declines”
in Chile (especially in Alsodes and Rhinoderma) and the Brazilian Atlantic Forest (where
declines are still poorly documented, but involve several genera, including Cycloramphus
and Thoropa (Heyer et al. 1988; Eterovick et al. 2005). The growing evidence suggests that
the Neotropics is in the process of losing most of its montane, stream-associated amphibian
fauna in the space of just a few decades.
KEY FINDINGS
• A total of 2,916 species are recorded from the Neotropical Realm, of which 1,145 (39%)
are considered threatened.
• At the species level, 2,808 amphibians (96% of those present) are endemic to the
Neotropics - roughly half of all recognized amphibians worldwide; of the 20 families
found in the region, six are endemic, and of 189 amphibian genera occurring, 157 are
endemic.
• The percentage of threatened species is very high in the families Rhinodermatidae (100%),
Ambystomatidae (67%), Plethodontidae (60%), Bufonidae (53%), Salamandridae (50%),
Leptodactylidae (46%) and Ranidae (41%), reflecting both habitat loss and declines most
likely related to chytridiomycosis and climate change.
• Geographic concentrations of threatened species occur in the Greater Antilles (Cuba,
Hispaniola, Jamaica and Puerto Rico); Mesoamerica (Central America south to Panama);
the tropical Andes (especially in Colombia and Ecuador, but also increasingly in Peru and
Bolivia); the Venezuelan highlands; central Chile; and the Atlantic Forests of southern
Brazil.
• Brazil has the largest number of species in the Neotropical Realm (751 species), and
has more endemics than any other country (489). Thirteen other countries have more
than 100 species (Colombia, Ecuador, Peru, Venezuela, Bolivia, Panama, Costa Rica,
Argentina, Guatemala, Guyana, Honduras, French Guiana, and Suriname), with the first
five of these countries having more than 50 endemics.
• Colombia has the largest number of threatened species (209), followed by Mexico
(190), Ecuador (163), and Brazil (110). Peru, Guatemala, Venezuela, Costa Rica, Panama,
Honduras, Cuba, Haiti, Dominican Republic, Argentina, Bolivia, and Chile each have 20
or more globally threatened species.
• Among species occurring in tropical forests, 52% of species in montane tropical
forest are threatened, compared with 30% in lowland tropical forest, probably
reflecting smaller range sizes of montane species, the lack of effective habitat
conservation in many montane regions, higher human population densities in
mountainous areas, the widespread incidence of chytridiomycosis, and the
increased vulnerability of montane species to the impacts of climate change.
Further, 45% of Neotropical amphibians associated with flowing water (most of
which are montane) are threatened.
• Habitat loss, primarily due to expanding croplands, vegetation removal (mainly logging),
urbanization/industrial development, and livestock grazing, is affecting almost 90% of
the threatened species in the region. Disease (usually chytridiomycosis) and pollution
are both impacting nearly 30% of globally threatened species.
• A massive 65% of the 470 globally “rapidly declining” species occur within the region;
these are concentrated in Mesoamerica, the Andes and the Greater Antilles where
habitat loss and chytridiomycosis have been especially severe. A total of 63% of the
“rapid declines” in the Neotropics are classified as “enigmatic declines” (probably due
to chytridiomycosis and climate change).
• Seven amphibian extinctions have been recorded from the Neotropics, and a further 121
species are possibly extinct (again concentrated in Mesoamerica, the Andes, and the
Greater Antilles).
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The Suriname Toad Pipa pipa (Least Concern) occurs widely in the Amazon Basin and the
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stage. © Manfred Beier

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Reichle, S. 2006. Distribution, diversity and conservation status of Bolivian amphibians. PhD Mathematisch salamanders in tropical America. Natural History Museum of Los Angeles County Science Bulletin species) from Panama to the upper Amazon
Naturwissenschaftliche Fakultaet, Rheinische Friedrichs-Wilhem Universitaet Bonn, Germany. 25:1-65. Basin forms a new family Hemiphractidae. The
Rodríguez, J.P. and Rojas-Suárez, F. 1995. Libro Rojo de la Fauna Venezolana. Provita, Fundación Polar, Young, B.E., Lips, K.R., Reaser, J.K., Ibáñez, R., Salas, A.W., Cedeño, J.R., Coloma, L.A., Ron, S., La Marca, genera Adelophryne, Atopophrynus, Barycholos,
Caracas, Venezuela. E., Meyer, J.R., Muñoz, A., Bolaños, F., Chaves, G. and Romo, D. 2001. Population declines and priori- Dischidodactylus, Craugastor, Eleutherodactylus,
Ron, S.R. 2005. Predicting the distribution of the amphibian pathogen Batrachochytrium dendrobatidis in ties for amphibian conservation in Latin America. Conservation Biology 15:1213-1223. Euparkerella, Geobatrachus, Holoaden, Ischnoc-
the New World. Biotropica 37:209-221. Young, B.E., Sedaghatkish, G., Roca, E. and Fuenmayor, Q. 1999. El Estatus de la Conservación de la nema, Phrynopus, Phyllonastes and Phyzelaphryne
Ron, S.R., Duellman, W.E., Coloma, L.A. and Bustamante, M.R. 2003. Population decline of the Jambato Toad Herpetofauna de Panamá: Resumen del Primer Taller Internacional sobre la Herpetofauna de Panamá. are transferred to the existing family Brachyc-
Atelopus ignescens (Anura: Bufonidae) in the Andes of Ecuador. Journal of Herpetology 37: 116-126. The Nature Conservancy, Arlington, Virginia, USA. ephalidae, creating a new grouping of nearly 800
species covering almost the same geographic
range as the former Leptodactylidae. The genera
The Pumpkin Toadlet Brachycephalus ephippium (Least Concern) is a member of the small Family Brachycephalidae from the Atlantic Forests of southern and eastern Brazil. It is a com- Cryptobatrachus and Stefania are transferred to
mon species in leaf-litter on the floor of primary and secondary forest. The egg clutches are deposited on the forest floor, and these develop directly without a free-living larval stage. © the new family Cryptobatrachidae (21 species)
Juarez Silva endemic to northern South America. The genera
Flectonotus and Gastrotheca are transferred to
the new family Amphignathodontidae (nearly
60 species), ranging from Costa Rica south to
Argentina. The genera Atelognathus, Batrachyla,
Ceratophrys, Insuetophrynus, Lepidobatrachus,
and Telmatobius are transferred to the new
family Ceratophryidae (c. 80 species) ranging
from Colombia south to Chile and Argentina. The
genera Alsodes, Crossodactylodes, Crossodacty-
lus, Cycloramphus, Eupsophus, Hylodes, Hylorina,
Limnomedusa, Macrogenioglottus, Megaelosia,
Odontophrynus, Proceratophrys, Thoropa, and
Zachaenus are transferred to the new fam-
ily Cycloramphidae (together with Rhinoderma
from Rhinodermatidae), with c. 130 species, in
southern tropical and temperate South America.
The genera Edalorhina, Engystomops, Eupemphix,
Physalaemus, Pleurodema, Pseudopaludicola, and
Sumuncuria are transferred to Leiuperidae (75
species) ranging from southern Mexico throughout
Central and South America south to central Chile
and central Argentina. With these changes, the
family Leptodactylidae is reduced to including the
genera Hydrolaetare, Leptodactylus (including
the subgenus Lithodytes for the former genera
Adenomera and Lithodytes), Paratelmatobius and
Scythrophrys, comprising c. 90 species through
much of the Neotropics and southern Nearctic,
including on some Caribbean islands.
100
ESSAY 9.1. THE EXTRAORDINARY CASE OF THE NEOTROPICAL HARLEQUIN FROGS (ATELOPUS): MASS EXTINCTION WITHIN A GENUS
Atelopus sorianoi (Critically Endangered) has the most restricted geographic range of any Venezuelan Atelopus species, being known from a single stream in
an isolated cloud forest in the Cordillera de Mérida. The last record of the species was in 1990. © Pascual Soriani
Figure 1. Richness map of species in the genus Atelopus, with dark red
colours corresponding to regions of higher richness. Maximum richness
equals four species.
Threatened Amphibians of the World
The harlequin frogs (genus Atelopus) are small, colourful “jewels” dis-
tributed in the humid forests and paramos of Central and South America.
The genus is the largest in the family Bufonidae, with about 80 described
species distributed from Costa Rica south to Bolivia and eastward through
the Amazon basin into the Guianas (Figure 1). Despite interest by scientists
in these species, their conservative morphology and variable coloration
have often obscured their taxonomy. Many species have highly variable
colour patterns, and different species frequently have similar colour pat-
terns. Recent genetic studies reveal both unappreciated genetic diversity
among populations, but also great variation within a given taxon. More
than 30 previously unrecognized species are currently in the process of
being described or being elevated to the species level.
Most Atelopus species are associated with streams, although many
occur part time of the year in terrestrial habitats (Lötters 1996). They
range from sea level to approximately 4,800m elevation, but the majority
live in highlands at 1,500-3,000m. Some species, such as A. varius (CR), A.
chiriquiensis (CR), A. carbonerensis (CR), and the now Extinct A. ignescens
and A. vogli, have been characterized as locally abundant, with hundreds of
animals seen in a few hundred meters, often during annual breeding events
(La Marca and Reinthaler 1991; Manzanilla and La Marca 2004; Pounds and
Crump 1994; Ron et al. 2003). Local endemism is common in the genus,
making species particularly vulnerable to extinction. At least 26 species
are known from only one site (per Ricketts et al. 2005).
Sadly, these beautiful and once common diurnal amphibians are now
vanishing. A recent study based on 113 Atelopus species (i.e., including
also undescribed forms and a few just recently named), revealed that 37%
of these species have undergone significant declines, and only 10 species
have what are believed to be stable populations (La Marca et al. 2005).
The majority of the declining species have disappeared in the last two
decades only, and many, such as Atelopus sorianoi (CR) are feared extinct;
at least 30 species have been missing from all known localities for at
least eight years. All species restricted to elevations of above 1000m have
declined and 75 percent have disappeared. At least three Atelopus species
are considered as Extinct according to the IUCN Red List Categories and
Criteria, namely Atelopus ignescens and A. longirostris from Ecuador, and
A. vogli from Venezuela (Lötters et al. 2004). To put things in perspective,
harlequin frogs represent about 15% of the 442 Critically Endangered (CR)
amphibian species on the IUCN Red List.
The first red flag that something was amiss came many years ago (La
Marca and Reinthaler 1991). Several potential causes were then discussed
trying to explain the observed declines, but today the most commonly cited
cause is the pathogenic chytrid fungus Batrachochytrium dendrobatidis,
which strikes even in undisturbed montane habitats. Habitat loss may
explain a few of the disappearances. However, it is not considered to
be a major cause in the case of the Atelopus species, since almost 20%
of the harlequin frogs have disappeared from protected areas. Other
potential causes of declines, such as introduced species, trade, and
pollution may partly explain a few cases, too, but are unlikely to have
affected the majority of the species. Synergistic combinations of factors
are expected to affect some harlequin frogs, as illustrated by the case of
Atelopus zeteki (CR), believed to be nearly extinct in the wild due to the
combined effects of habitat change, illegal collecting, and fungal disease
(Mendelson et al. 2006).
A recent finding suggests that large-scale warming of our planet is cor-
related with mass extinction in Atelopus (Pounds et al. 2006). The warming
trend, estimated at about 0.18ºC per decade, has been to the benefit of the
fungal pathogen. The rise in temperatures has most probably increased
the amount of evaporation in the tropical montane environments inhabited
by harlequin frogs, which in turn has been translated into increased cloud
formation. Increased cloud cover in turn leads to a decrease in incoming
solar radiation, thus reducing daytime temperatures, and by night may
result in a green-house effect that impedes natural heat loss from the
ground, with resulting warmer night-time temperatures. These cooler days
and warmer nights brings the pathogenic fungus to near optimum thermal
conditions, believed to be between17 and 25ºC, thereby encouraging its
growth, reproduction and propagation. The theory gains support since
most of the species have disappeared in the altitudinal band between
1,000 and 2,400m elevation, while recent “re-appearances” have occurred
either in low-elevation (humid lowland forests) or high-elevation (paramo)
habitats (Lötters et al. 2005). Both extremes in temperature conditions for
the fungal pathogen may act as “thermal refuges” for the few surviving
Atelopus populations.
Unfortunately, the problem of declines and extinctions in Atelopus is not
likely to diminish in the foreseeable future. Under a scenario of double CO2
concentrations within the next century (Malcolm et al. 2006), the rate of
amphibian extinctions is expected to increase in many regions, including the
Tropical Andes, where most harlequin frogs are known to occur. Atelopus may
actually be a good indicator of what is happening to other less conspicuous
species that could experience similar declines. The loss of these important
links in the trophic web has unforeseeable consequences (Ranvestel et
al. 2004). Furthermore, the resulting impoverished biodiversity may also
represent a loss in potential advances in biomedicine and biotechnology
(Mendelson et al. 2006).
Currently, there is no known effective protection against Batrachochytrium
dendrobatidis in the wild. Therefore, one of the most tempting alternatives to
cope with the Atelopus declines is to initiate captive-breeding programmes
(see Essay 11.5). The success with the Panamanian Golden Frog, Atelopus
zeteki (Zippel 2002), which is now available in breeding colonies in numerous
zoos, is promising in this regard. Nevertheless, comprehensive captive-breed-
ing programs for all Atelopus species threatened with extinction appear to
be impossible due to the many species involved.
Enrique La Marca and Stefan Lötters
References
La Marca, E. and Reinthaler, H.P. 1991. Population changes in Atelopus species of the
Cordillera de Mérida, Venezuela. Herpetological Review 22:125-128.
La Marca, E., Lips, K.R., Lötters, S., Puschendorf, R., Ibáñez, R., Rueda-Almonacid, J.V.,
Schulte, R., Marty, C., Castro, F., Manzanilla-Puppo, J., García-Pérez, J.E., Bolaños,
F., Chaves, G., Pounds, J.A., Toral, E. and Young, B.E. 2005. Catastrophic popula-
tion declines and extinctions in neotropical harlequin frogs (Bufonidae: Atelopus).
Biotropica 37:190-201.
Lötters, S. 1996. The Neotropical toad genus Atelopus. Checklist—biology—distribution.
M. Vences and F. Glaw, Verlags GbR, Cologne, Germany.
Lötters, S., La Marca, E., Stuart, S., Gagliardo, R. and Veith, M. 2004. A new dimension
of current biodiversity loss? Herpetotropicos 1:29-31.
Lötters, S., La Marca, E., Gagliardo, R.W., Señaris, C.J. and Veith, M. 2005. Harlequin
frogs back? Some thoughts and speculations. Froglog 70:1-3.
Malcolm, J.R., Canran, L., Neilson, R.P., Hansen, L. and Hannah, L. 2006. Global warming
and extinctions of endemic species from biodiversity hotpots. Conservation Biology
20:358-548.
Manzanilla, J. and La Marca, E. 2004. Population status of the Rancho Grande harlequin
frog (Atelopus cruciger Lichtenstein and Martens 1856), a proposed critically endan-
gered species from the Venezuelan Coastal Range. Memoria de la Fundación La Salle
de Ciencias Naturales, Caracas, 62(157):5-29.
Mendelson III, J.R., Lips, K.R., Gagliardo, R.W., Rabb, G.B., Collins, J.P., Diffendorfer, J.E.,
Daszak, P., Ibáñez, R., Zippel, K.C., Lawson, D.P., Wright, K.M., Stuart, S.N., Gascon, C., da
Silva, H.R., Burrowes, P.A., Joglar, R.L., La Marca, E., Lötters, S., du Preez, L.H., Weldon,
C., Hyatt, A., Rodriguez-Mahecha, J.V., Hunt, S., Robertson, H., Lock, B., Raxworthy, C.J.,
Frost, D.R., Lacy, R.C., Alford, R.A., Campbell, J.A., Parra-Olea, G., Bolaños, F., Calvo
Domingo, J.J., Halliday, T., Murphy, J.B., Wake, M.H., Coloma, L.A., Kuzmin, S.L., Stanley
Price, M., Howell, K.M., Lau, M., Pethiyagoda, R., Boone, M., Lannoo, M.J., Blaustein,
A.R., Dobson, A., Griffiths, R.A., Crump, M.L., Wake, D.B. and Brodie, E.D., Jr. 2006.
Confronting Amphibian Declines and Extinctions. Science 313:48.
Pounds, J.A. and Crump, M.L. 1994. Amphibian declines and climate disturbance: The case
of the golden toad and the harlequin frog. Conservation Biology 8:72-85.
Pounds, J.A., Bustamante, M.R., Coloma, L.A., Consuegra, J.A., Fogden, M.P.L., Foster, P.N.,
La Marca, E., Masters, K.L., Merino-Viteri, A., Puschendorf, R., Ron, S.R., Sanchez-
Azofeifa, G.A., Still, C.J. and Young, B.E. 2006. Widespread amphibian extinctions
from epidemic disease driven by global warming. Nature 439:161-167.
Ranvestel, T.W., Lips, K.R., Pringle, C.M., Whiles, M.R. and Bixby, R.J. 2004. Neotropical
tadpoles influence stream benthos: evidence for ecological consequences of amphibian
declines. Freshwater Biology 49:274-285.
Ricketts, T.H., Dinerstein, E., Boucher, T., Brooks, T.M., Butchart, S.H.M., Hoffmann, M.,
Lamoreux, J.F., Morrison, J., Parr, M., Pilgrim, J.D., Rodrigues, A.S.L., Sechrest, W.,
Wallace, G.E., Berlin, K., Bielby, J., Burgess, N.D., Church, D.R., Cox, N., Knox, D.,
Loucks, C., Luck, G.W., Master, L.L., Moore, R., Naidoo, R., Ridgely, R., Schatz, G.E.,
Shire, G., Strand, H., Wettengel, W. and Wikramanayake, E. 2005. Pinpointing and
preventing imminent extinctions. Proceedings of the National Academy of Sciences
USA 102:18497-18501.
Ron, S.R., Duellman, W.E., Coloma, L.A. and Bustamante, M.R. 2003. Population decline
of the jambato toad Atelopus ignescens (Anura: Bufonidae) in the Andes of Ecuador.
Journal of Herpetology 37:116-126.
Stuart, S.N., Chanson, J.S., Cox, N.A., Young, B.E., Rodrigues, A.S.L., Fischman, D.L. and
Waller, R.W. 2004. Status and trends of amphibian declines and extinctions worldwide.
Science 306:1783-1786.
Zippel, K. 2002. Conserving the Panamanian golden frog: Proyecto Rana Dorada. Herpe-
tological Review 33:11-12.

ESSAY 9.2. CONSERVATION STATUS OF THE ANDEAN FROGS OF THE GENERA TELMATOBIUS AND BATRACHOPHRYNUS
The frogs of the genera Telmatobius and Batrachophrynus constitute a
remarkable group of endemic Andean anurans that occur from central
Ecuador in the north, to northern Chile and Argentina in the south (Figure
1). The most recent classification of Telmatobius (Lavilla 2005) includes 56
species, with Peru harbouring the highest diversity (23 species), followed by
Bolivia (15 species), Argentina (14 species), Chile (9 species; the description
of an additional species was in press at the time of writing this report) and
Ecuador (3 species). Batrachophrynus includes two species (endemic to
central Peru), and while we treat it as distinct in the current essay, recent
studies have indicated that Batrachophynus is not a valid genus and should
rather be included in Telmatobius (Aguilar and Pacheco 2005; Córdova and
Descailleaux 2005; Sinsch et al. 2005).
The genus Telmatobius is mostly aquatic, occupying a wide, albeit
montane, altitudinal range (1,300-5,000m asl), and inhabiting habitats as
diverse as cloud forests to humid paramos and dry puna. Many species have
co-existed with humans for centuries, and, undoubtedly, habitat destruc-
tion, mining, agricultural practices and livestock (especially camelids) have
influenced the distribution and local abundance of some taxa (De la Riva
2005). Several species are captured either for food or because of certain
putative medicinal or magical properties; other potential threats include
water pollution and the introduction of trout for fishing. Particularly concern-
ing is the case of the giant Lake Titicaca Frog (Telmatobius culeus, CR) and
the even larger Lake Junin Frog (B. macrostomus, EN), both of which have
been affected by over-fishing and other problems. Although some protection
measures have been implemented (such as captive breeding), they have
proved to be mostly unsuccessful (Pérez 2005).
Since Telmatobius and Batrachophrynus frogs inhabit montane areas
and are stream- or lake-breeders, their biology and ecology render them
particularly susceptible to chytrid infection. Indeed, there is growing
evidence that chytridiomycosis is having a direct impact on populations
of these frogs. Reports of serious population declines in Telmatobius first
came from Ecuador, and a recent summary of the conservation status
of Ecuadorian Telmatobius yields conclusive evidence of a catastrophic
decline of the three species endemic to that country (Merino-Viteri et
al. 2005). Specimens found in the 1980s and 1990s had malformations
and symptoms of diseases, including chytridiomycosis. Despite intensive
surveys for living animals in recent years (including 2005), the last living
Telmatobius specimen seen in Ecuador was a tadpole of T. niger (CR) with
severe epidermal damages, collected on 1 December 1994; T. vellardi (CR)
was last seen in 1987, and T. cirrhacelis (CR) in 1981. The three Ecuadorian
Telmatobius are now likely extinct.
While the situation in Ecuador has been thoroughly investigated, there
is almost no published information for other range countries. In Perú, Lehr
(2005) stated that these frogs are threatened due to agricultural practices,
water pollution, and commercial utilization. Seimon et al. (2005) reported
a case of chytridiomycosis affecting T. marmoratus (VU), collected in July
2002 in the department of Cusco, although without reference to population
declines. This species has the broadest distribution of any Telmatobius,
and occurs in the Altiplano and Puna highlands above the tree line. Healthy
specimens were found in the department of Puno in southern Peru during
recent fieldwork (February 2006; De la Riva, unpubl.), and although it is
plausible that some populations of this widespread species are extinct
or have declined due to chytridiomycosis, the presence of larvae and
Telmatobius marmoratus (Vulnerable) has the widest range of any species in
the genus, being known from the Andean region of southern Peru, northern
and central Bolivia, and northern Chile. © Ignacio de la Riva
Figure 1. Richness map of species in the genera Telmatobius and Batracho-
phrynus, with dark red colours corresponding to areas of higher richness.
Maximum richness equals four species.
Chapter 9. Amphibians of the Neotropical Realm
adults in many sites indicates that, overall, the species is not severely
threatened. However, this situation could change in the future if climate
change facilitates a shift to favorable conditions for chytrid in previously
unsuitable zones.
Unfortunately, more alarming data, albeit preliminary, concern spe-
cies from the humid paramos and upper cloud forest regions of Peru. In
1999-2001, Telmatobius were largely extirpated from the department of
Cajamarca, where farmers frequently encountered dead animals (R. Schulte,
in. litt.). Recent fieldwork (February 2006) by a team of five herpetologists
surveying nine Andean valleys in the department of Puno and southern
Cusco did not yield a single specimen of Telmatobius, despite thorough
searches in appropriate sites. In several places, local people explained that
all “kaylas” frogs (=Telmatobius) vanished two years ago.
In Bolivia, the situation seems to be similar. De la Riva (2005) docu-
mented a severe decline of a newly described upper cloud forest species,
T. espadai. Tadpoles of this species were extremely abundant in Río Apaza
(Cochabamba) in 1990, but no tadpoles were found in 1994 and 1999, and
only a single one in 1998. A recent examination of the oral structures of this
individual showed an almost complete destruction of keratinized structures,
which is consistent with chytridiomycosis. Another paramo/upper cloud
forest species, T. sanborni, occurred at least between Pelechuco (La Paz,
Bolivia), where it was abundant at least in 2001, and Ollachea (Puno, Peru)
(De la Riva 2005). No trace of this species was found during the recent
surveys in Peru in February 2006, although it has yet to be searched for in
Bolivia. The last records of two other threatened species, T. sibiricus (EN)
and T. verrucosus (VU), are from 2004 (De la Riva 2005).
Chilean species of Telmatobius occur in mostly desertic conditions
and dry puna, and are subject to the same general threats as other
highland species (Formas et al. 2005). At present, there is no direct or
indirect evidence of chytrid infection in Chilean Telmatobius. The avail-
able information in Argentina is not promising (Lavilla and Barrionuevo
2005). Early in 2006, S. Barrionuevo (pers. comm.) found evidence of
chytrid fungus in individuals of Telmatobius atacamensis (CR; a species
already threatened by mining activities) in the environs of San Antonio
de los Cobres (Salta) and in a population of Telmatobius pisanoi (EN)
near El Pichao (Tucumán). Another species, T. laticeps (EN), fairly com-
mon in the past in the area of Tafí del Valle (Tucumán) disappeared from
its range in the last few years due to unknown causes; the same might
have happened with the forest-dwelling T. ceiorum (EN; S. Barrionuevo
and M.L. Ponssa, pers. comm.).
In summary, frogs of the genera Telmatobius (and Batrachophynus) are
severely threatened, and at a scale comparable only to the bufonid genus
Atelopus (see Essay 9.1). In Telmatobius, as with Atelopus, many species
are still to be named, many of those already described are known only from
the type locality or nearby, and several extinctions have already taken place.
However, there is one important difference: Telmatobius has no lowland
species, and thus the entire genus faces the very real danger of extinction
in the very near future. Unfortunately, the case of Telmatobius frogs seems
to strongly support Lips et al.’s (2006) statement: “…it is no longer correct
to speak of global amphibian declines but, more appropriately, of global
amphibian extinctions.”
Ignacio De la Riva and Esteban O. Lavilla
101
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Monog. Herpetol. 7, AHE, Valencia, Spain.
102
ESSAY 9.3. NEOTROPICAL SALAMANDERS
Living salamanders comprise about 550 species, representing approximately
one-tenth of living amphibians. The order Caudata hosts the largest amphib-
ian, the Chinese Giant Salamander (Andrias davidianus, CR), in which adults
measure 180cm from nose to tip of tail (and see Essay 4.7), as well as one
of the smallest, Thorius arboreus (EN), one of several species of the genus
that achieve sexual maturity at about 15mm in length. Although the number
of salamander species is small compared with that of frogs, the diversity
of species and life histories, coupled with late 20th century declines and
disappearances worldwide, make salamanders an important model for
understanding the causes of global change (i.e., climate change, pollution,
habitat loss, etc) and their effect on biodiversity.
Salamanders are more commonly representatives of the northern
temperate regions. Only a few groups have colonized tropical regions: the
Salamandridae in south-eastern Asia and the Plethodontidae in tropical
America. The magnitude and extent of these tropical invasions differs greatly.
While tropical Asia has been colonized by only a few species, the Neotropics
have been the stage for a large-scale radiation encompassing almost 40%
of all salamander species.
The main Neotropical salamander radiation is restricted to a single clade,
the supergenus Bolitoglossa (Parra-Olea et al. 2004), which is represented
by more than 180 species and 12 genera, and ranges from northern México
to Brazil (Figure 1). Bolitoglossine salamanders share fully terrestrial life
histories, internal fertilization, direct development within encapsulated eggs,
and a highly specialized feeding mechanism. These derived traits have played
a major role in the success of bolitoglossines in the tropics (Wake 1987). A
second radiation in the Neotropics can be found in the genus Ambystoma in
the Transvolcanic Axis of central Mexico. Although this radiation has produced
relatively few species, it includes several independently evolved paedomor-
phic lineages. A prime example is the Axolotl (Ambystoma mexicanum, CR),
which is widely used to illustrate paedomorphosis in vertebrate evolution.
“Cryptic speciation” is common in salamanders. Their morphological evolu-
tion is conservative, and different populations often share traits that have
arisen through convergence or that have evolved in parallel. Genetic studies
are thus often necessary to identify new species. Such studies, coupled with
fieldwork, have shown that salamanders in the tropics often exhibit a pattern
of local isolation, with extreme genetic differentiation occurring over short
distances (“tropicality” syndrome) (Garcia-Paris et al. 2000). Units that have
been previously treated as single species often comprise multiple and geneti-
cally distinct lineages. Accordingly, the total number of salamander species
remains unknown, and new forms are steadily being described. In Mexico,
for example, the number of recognized species has risen from 93 to 128 in the
past 10 years — a 39% increase (Flores-Villela and Canseco 2004).
Amphibian populations are declining worldwide and Neotropical salaman-
ders are no exception. In several localities where salamanders were seen or
collected by the hundreds in the 1970s and 1980s, it is now difficult to find a
single individual. Some declines have occurred in seemingly pristine areas,
such as Cerro San Felipe in Oaxaca, Mexico, and Reserva de Monte Verde
in Costa Rica (Parra-Olea et al. 1999). The results of the Global Amphibian
Assessment found that out of a total of 226 species in the Neotropics, 36 are
Critically Endangered, 74 Endangered, and 28 Vulnerable. An additional 42
were listed as Data Deficient. Habitat loss and water pollution is the major
threat to most species of salamanders, and, for some species, over-collecting
(e.g., for food), the introduction of exotic species, and urban development
are significant threats. Other factors such as climate change, increased
UV-B radiation, chemical contamination, and emerging infectious disease
are currently being evaluated.
ESSAY 9.4. BRAZIL: THE WORLD LEADER IN AMPHIBIAN DIVERSITY
More than 8.5 million km2 in size, Brazil is the fifth largest country in the world
and the largest among those in the tropics. It has the largest continental biota
on Earth, and inspired the concept of Megadiversity countries (Mittermeier et
al. 1997). Conservative estimates indicate that Brazil is home to 13% of global
biodiversity, and has the richest flora with more than 56,000 species.
Not surprisingly, then, Brazil is also the world leader in amphibian diversity
and endemism. The Global Amphibian Assessment records 751 native spe-
cies, of which around 65% are endemic. However, the rate of description of
new species is very high, and according to an updated list of the Brazilian
Society of Herpetology (SBH) there are now 794 amphibian species in Brazil.
Only Colombia can come close to rivaling Brazil in terms of absolute numbers
of amphibians present (see Essay 9.5). Further, even the current number of
recognized amphibians is an underestimate, as evidenced by the number of
species that have been discovered and described just in recent years. Several
areas have never been inventoried and likely would reveal many new species
waiting to be described, including large areas in the Cerrado and Amazon,
such as southern Pará and Maranhão states, western Bahia state, northern
Mato Grosso, and almost all of Tocantins state.
Amphibian diversity is not evenly distributed across the country, and
there is a noticeable concentration of species in some regions, for example
in the Atlantic Forest (Figure 1). Unfortunately, the Atlantic Forest is also
the center of origin for reports of amphibian declines in Brazil (Eterovick
et al. 2005) (Figure 2). The Atlantic Forest (or Mata Atlântica) stretches
along Brazil’s Atlantic coast from the northern state of Rio Grande do Norte
south to Rio Grande do Sul, extending from 4º to 32ºS. Long isolated from
other major rainforest blocks in South America, the Atlantic Forest has an
extremely diverse and unique mix of vegetation and forest types, with eleva-
tion varying from the sea level to about 2,900m. Unfortunately, this biome
has been largely destroyed, and only about 7% of its original native forest
cover remains intact, and it is regarded as a recognized global biodiversity
Threatened Amphibians of the World
To date, the chytrid fungus Batrachochytrium dendrobatidis has been
found in four species of plethodontid salamanders (Lips et al. 2006) and in
four ambystomatid species (G. Parra-Olea, unpubl.), but massive die offs of
salamanders have, as yet, not been linked to this pathogen. Climate change
and forest fragmentation could have important impacts on salamanders,
especially considering the limited dispersal abilities of these animals.
Bioclimatic envelope modelling suggests that the terrestrial salamander
Pseudoeurycea leprosa (VU) in Mexico could lose almost 75% of its range
area over the next 50 years because of climate change (Parra-Olea et al.
2005). This will be true for all terrestrial salamanders that inhabit pine and/or
pine-oak forests, and will be exacerbated for the majority of species with
small distributional ranges.
Traditionally, aquatic salamanders of the genus Ambystoma, such as the
Axolotl and the Achoque, A. mexicanum, have played an important role in local
communities. The Aztecs considered the Axolotl as the transfiguration of the
deity Xolotl, and both species are exploited by local communities as a food
source and as a remedy for respiratory infections (see Essay 2.3). The main
threats to most Ambystoma species include contamination and drying out of
their aquatic habitats, the introduction of exotic species, and over-exploitation.
For example, the local conditions of Lake Pátzcuaro, to which the Achoque
is endemic, have changed following an increase in water temperature, a
decrease in the mean depth of the lake, and the introduction of exotic fish
(Centrarchidae, Cyprinidae and Cyclidae) and their accompanying parasites
(i.e. Bothriocephalus acheilognathi) (García et al. 1993). Furthermore, between
1987 and 2000 the harvest of the Achoque was approximately 27,592 kg
(Huacuz 2002). A management plan has been proposed for the Achoque and a
captive-breeding program was started by a group of nuns from the Pátzcuaro
convent, with the main objective of sustainably harvesting the species from
the wild for the production of cough syrup for the community. However, we
still know little about the size of the population, its genetic structure, or its
dynamics, so evaluating its chances of survival is difficult.
Given the alarming declines and disappearances that have been witnessed
among Neotropical salamanders, examining the various potential threats is
urgent, particularly where these threats act in synergy (Pounds et al. 2006).
Most importantly, we cannot begin to propose adequate management plans
for species, unless we know and understand the taxonomic and phylogenetic
identity of the species of concern. Resolving taxonomic uncertainties is
thus fundamentally important. Studies involving the use of phylogenetics
for uncovering cryptic species diversity will help reveal the true diversity of
the group, and in turn also help identify unique lineages and hot spots of
diversity. Finally, they will identify the affinities of individual populations,
thus facilitating appropriate decisions about the translocation or reintroduc-
tion of salamanders.
Gabriela Parra-Olea
References
Flores-Villela, O. and Canseco-Marquez, L. 2004. Nuevas especies y cambios taxonomicos
para la herpetofauna de México. Acta Zoologica Mexicana 2:115-144.
Huacuz, E.D. 2002. Programa de Conservación y Manejo de Ambystoma dumerili. El
Achoque del Lago de Patzcuaro. Universidad Michoacana de San Nicolas de Hidalgo,
Fondo Mexicano para la Conservación de la Natraleza, A. C. y Secretaria de Medio
Ambiente y Reccursos Naturales. Michoacán, México.
García, A.I., Pérez-Ponce de León, G. and García, P.L. 1993. Contribución al conocimiento de
hotspot (Mittermeier et al. 2004).
In a recent review of amphibian declines in Brazil, Eterovick et al. (2005)
found that most species experiencing declines in the Atlantic Forest are
recorded at elevations up to 1,000m, some of them associated with streams
(Colostethus olfersioides VU, Bokermannohyla langei DD, Crossodactylus dis-
par DD, C. gaudichaudii LC, Hylodes babax DD, Cycloramphus boraceiensis LC,
C. mirandaribeiroi DD, C. semipalmatus NT, Hyalinobatrachium uranoscopum
LC), others with cliffs dripping with water for tadpole development (Cycloram-
phus duseni DD, Thoropa lutzi EN, T. petropolitana VU). The species are either
direct developers (Adelophryne baturitensis VU), or their mode of reproduction
is unknown (Colostethus carioca DD, Cycloramphus eleutherodactylus DD). A
few species are recorded only above 1,500m (Eleutherodactylus paranaensis
DD, Cycloramphus granulosus DD, Paratelmatobius lutzii DD).
But declines are not only reported from the Atlantic Forest. The Brazil-
ian Cerrado, another global biodiversity hotspot, makes up one-fifth of the
country, and is the most extensive woodland-savannah in South America.
The Cerrado receives abundant rainfall between October and April, while the
rest of the year is characterized by a pronounced dry season. The Cerrado
also contains a rich montane meadow vegetation found at higher portions
of some of the mountain ranges in south-eastern Brazil, such as the Serra do
Espinhaço. Eterovick et al. (2005) reported the first declines of amphibians
in the Cerrado, among the species Crossodactylus bokermanni (DD) and
Epipedobates flavopictus (LC). They recorded only a few adult individuals of
Crossodactylus bokermanni, from the Serra do Cipó in 2001 in the Parque
Nacional da Serra do Cipó, and yet the species was known to be common in
the same study area in 1971-1974.
The main cause for declines in amphibians in Brazil is undoubtedly
habitat destruction, largely as a consequence of deforestation, agricultural
expansion, mining, fire, and infrastructure development and urbanization.
However, other factors, such as severe winters, pollution and acid rain, and
Figure 1. Richness map of Neotropical salamander species (n=242) in the gen-
era Batrachoseps, Bolitoglossa, Chiropterotriton, Cryptotriton, Dendrotriton,
Ixalotriton, Nototriton, Nyctanolis, Oedipina, Parvimolge, Pseudoeurycea and
Thorius. Dark red colours correspond to higher richness. Colour scale based
on five quantile classes. Maximum richness equals 17 species.
las comunidades de helmintos de dos especies de anfibios endémicos del Lago de Pátz-
cuaro, Michoacán: Rana dunni y Ambystoma dumerilii. Cuad. Mex. Zool. 1:73-80.
García-Paris, M., Good, D.A., Parra-Olea, G. and Wake D.B. 2000. Biodiversity of Costa
Rican salamanders: Implications of high levels of genetic differentiation and phylo-
geographic structure for species formation. Proceedings of the National Academy of
Sciences, USA 97:1640-1647.
Lips, K.R., Brem, F., Brenes, R., Reeve, J.D., Alford, R.A., Voyles, J., Carey, C., Livo, L.,
Pessier, A.P.,and Collins, J.P. 2006. Emerging infectious disease and the loss of biodi-
versity in a Neotropical amphibian community. Proceedings of the National Academy
of Sciences, USA 103:3165-3170.
Parra-Olea, G., García-Paris, M., and Wake, D.B. 1999. Status of some populations of
Mexican salamanders (Amphibia: Plethodontidae). Revista de Biologia Tropical
47:217-223.
Parra-Olea, G., García-Paris, M. and Wake, D.B. 2004. Molecular diversification of the genus
Bolitoglossa and its evolutionary and biogeographic consequences for the invasion of
American tropics. Biological Journal of the Linnean Society 81:325-346.
Parra-Olea, G., Martínez-Meyer, E. and Pérez Ponce de León, G. 2005 Forecasting climate
change effects on salamander distribution in the highlands of Central Mexico.
Biotropica 37:202-208.
Pounds, J.A., Bustamante, M.R., Coloma, L.A., Consuegra, J.A., Fodgen, M.P., Foster, P.N.,
La Marca, E., Masters, K.L., Merino-Viteri, A., Puschendorf, R., Ron, S.R., Sanchez-
Azofeifa, A., Still, C.J. and Young, B.E. 2006. Widespread amphibian extinctions from
epidemic disease driven by global warming. Nature 439:160-167.
Wake, D.B. 1987. Adaptive radiation of salamanders in Middle American cloud forests.
Annals of the Missouri Botanical Garden 74:242-264.
extended dry periods, are all possible causative factors (Eterovick et al. 2005).
Furthermore, disease may also play an important role. Carnaval et al. (2006)
conducted histological screenings of 96 preserved specimens of anurans
collected at 10 sites in the Atlantic Forest and found chytrid fungus to be
widely distributed, having recorded the disease in specimens of Colostethus
olfersioides, Bokermannohyla gouveai DD and Hypsiboas freicanecae DD,
as well as Thoropa miliaris LC and Crossodactylus caramaschii LC. More
concerningly, the altitudinal range is broad, spanning from less than 100m
to about 2,400m (in the Parque Nacional do Itatiaia). The widespread occur-
rence of chytrid in the Atlantic Forest adds to the challenge of conserving
an already threatened biome. More recently, Toledo et al. (2006) extended
the distribution of the fungus in Brazil ca. 630km southward from the previ-
ous southernmost record of Carnaval et al. (2006), reaching São Francisco
de Paula in the state of Rio Grande do Sul (the southernmost limits of the
Atlantic rainforest), and speculated on its distribution in the Cerrado and
Pantanal (and see Ron 2005).
The Brazilian politics of conservation include important legal instruments,
such as lists of threatened species and the selection of priority areas for con-
servation in all Brazilian biomes. On the other hand, the country still has much
to do to improve its protected areas network, particularly given the noticeable
gaps in coverage in the Atlantic Forest. The Cerrado biome deserves special
attention because of the scarce knowledge about amphibians and its high rate
of habitat loss due to the rapid advance of the agricultural frontier. Brazilian
herpetologists have clearly made dramatic strides forward in the last two
decades in improving our knowledge on natural history, ecology, and basic
life-history of many species, but a great deal remains to be done to better
understand the causes of the declines being witnessed among the country’s
amphibians, and the most appropriate means to mitigate these threats.
Débora Leite Silvano

References
Carnaval, A.C.O.Q.C., Puschendorf, R., Peixoto, O.L., Verdade, V.K., and Rodrigues, M.T.
2006. Amphibian Chytrid Fungus Broadly Distributed in the Brazilian Atlantic Rain
Forest. EcoHealth 3:41-48.
Eterovick, P.C., Carnaval, A.C.O.Q.C., Borges-Nojosa, D.M., Silvano, D.L., Segalla,
M.V. and Sazima, I. 2005. Amphibian Declines in Brazil: An Overview. Biotropica
37:166-179.
Mittermeier, R.A., Robles Gil, P. and Mittermeier, C.G. (eds.). 1997. Megadiversity: Earth’s
Biologically Wealthiest Nations. CEMEX, Mexico City, Mexico.
Mittermeier, R.A., Robles-Gil, P., Hoffmann, M., Pilgrim, J.D., Brooks, T.M., Mittermeier,
C.G., Lamoreux, J.L. and Fonseca, G. 2004. Hotspots Revisited: Earth’s Biologically
Richest and Most Endangered Ecoregions. Second Edition. CEMEX, Mexico City,
Mexico.
Ron, S.R. 2005. Predicting the distribution of the amphibian pathogen Batrachochytrium
dendrobatidis in the New World. Biotropica 37(2):209-221.
Toledo, L.F., Britto, F.B., Araujo, O.G.S., Giasson, L.M.O. and Haddad, C.F.B. The occurrence
of Batrachochytrium dendrobatidis in Brazil and the inclusion of 17 new cases of
infection. South American Journal of Herpetology 1(3):185-191.
ESSAY 9.5. A BRIEF OVERVIEW OF THE AMPHIBIANS OF COLOMBIA
The amphibian fauna of Colombia is among the largest and most diverse on
the planet. According to the results of the Global Amphibian Assessment,
nearly 700 recognized species of amphibians are known from, or expected to
occur in, Colombia, and our current estimate stands at 732. The diversity of
amphibians in Colombia is, to a certain degree, the fortuitous consequence
of human politics – it is as if Colombia’s borders were drawn with the specific
intent of maximizing its amphibian diversity. That is, Colombia’s amphibian
diversity is a function not only of the area of this tropical country, but also its
specific location. For example, the two countries with the greatest number of
amphibian species are Brazil and Colombia (Ecuador is a distant third, with
“only” 449 species). With 752 recognized species listed in the GAA, Brazil has
a slightly larger amphibian fauna, but its area is over eight times greater than
that of Colombia. Consequently, Brazil has 8.8X10-5 species per km2, whereas
Colombia has 6.1X10-4 species per km2 – a full order of magnitude more.
In occupying the north-western-most portion of South America, Colombia
includes the rich amphibian fauna of the rain-soaked Pacific lowlands and
adjacent Andean foothills, and this is augmented by capturing many species
(e.g., the dendrobatid Colostethus panamensis, LC) and lineages (e.g., the
brachycephalid genus Craugastor) that extend into Colombia from Central
America. The eastern borders reach far into Amazonia, and further north the
Llanos secure fauna associated with the Orinoco river drainage. Predomi-
nantly Venezuelan lineages, such as the aromobatid Aromobates, extend into
Colombia in the Serranía de Perijá, and the isolated Sierra Nevada de Santa
Chapter 9. Amphibians of the Neotropical Realm
Figure 1. Richness map of all amphibian species in Brazil, with dark red
colours corresponding to regions of higher richness. Colour scale based on
10 quantile classes. Maximum richness equals 139 species.
Marta harbors an endemic fauna that includes such enigmatic species as the
brachycephalid Geobatrachus walkeri (EN).
Nevertheless, although Colombia’s regional span contributes greatly to
the diversity of amphibians, it is the Andean backbone that is most significant
(Lynch et al. 1997). Whereas to the south and north-east the Andes form
comparatively simple systems, in Colombia they divide into three isolated
ranges that radiate from the Nudo de Pasto, and these ranges harbor about
two-thirds of Colombian amphibians. Among the Andean species, most occur
in the cool, moist cloud forests between 1,200 and 2,500m asl, and many are
confined to extremely small areas. Although experimental data are lacking, it
is assumed that this isolation is due to the adaptation of species to specific
environments and their inability to survive under even mildly different condi-
tions. For example, although two adjacent mountains may share identical
environmental conditions, the different environment (e.g., higher temperature
and lower precipitation) of the intervening valley would serve as a barrier to
dispersion and gene flow (Lynch and Duellman 1997).
The limited geographic distribution of most Colombian amphibians makes
them extremely susceptible to habitat alteration and destruction. This poses
a special challenge for Colombian policy makers because humans have
targeted precisely the same elevations of the Andes for their development
activities. For many Andean species, the removal of a single remaining patch
of forest may mean the extinction of the species. For example, Atopophrynus
syntomopus (CR) is the only species of its genus and is known from a single
103
Figure 2. Richness map of all threatened amphibian species in Brazil, with
dark red colours corresponding to regions of higher richness. Colour scale
based on five quantile classes. Maximum richness equals 24 species.
locality in the Cordillera Central of Antioquia in an area that has been
subjected to extreme deforestation. In addition to the habitat alteration
that accompanies human development, other actual or potential threats to
Colombian amphibians include the introduction of exotic species (including
the illicit introduction of the North American Bull frog, Lithobates catesbeiana,
for commercial purposes), global climate change and increased exposure to
ultraviolet radiation, and infectious disease – especially chytridiomycosis,
caused by the chytrid fungus Batrachochytrium dendrobatidis (Rueda-Almo-
nacid et al. 2004).
According to the IUCN Red List, just less than half (47%) of all Colombian
amphibian species are classified as Least Concern. Of the remainder, 18% are
Endangered or Critically Endangered, and another 18% are Near threatened
or Vulnerable. Quantitative data derived from rigorous monitoring studies are
lacking for all of those species, but data are so scant for an additional 17%
of Colombian species that not even a rough estimate of their status could
be made, and they are designated as Data Deficient.
Given the state of knowledge of Colombian amphibians, two areas of
research are in urgent need of increased attention. First, taxonomic research
– including the exploration of under-sampled areas and the production of
revisionary, monographic studies of groups and regions – must be expanded to
complete the identification of Colombian amphibians. Although few localities
can be considered thoroughly sampled, Acosta-Galvis (2000) highlighted a
number of high-priority regions, including mid- to high-elevations of the central
Geobatrachus walkeri (Endangered) is a frog known only from the north-
western and western slopes of the Sierra Nevada de Santa Marta in northern
Colombia. © Taran Grant
Figure 1. Priority regions for further inventory and survey work in Colombia.
104
and northern Cordillera Central and Cordillera Occidental, higher elevations
along the length of the Serranía de Perijá, the páramos of the Cordillera
Oriental in southern Cundinamarca and Tolima departments, the Serranía
del Darién along the Colombo-Panamanian border, the southern Cordillera
Occidental and adjacent lowlands in Cauca and Nariño departments, and
the Cordillera Oriental and rainforests of Putumayo, Amazonas, Caquetá,
Guaviare, Vaupés, and Gauinía departments (Figure 1).
The discovery and identification of previously unknown species in all
major groups of Colombian amphibians shows no sign of slowing in the
foreseeable future. Indeed, as the expansion of institutional (e.g., natural
history collections, molecular laboratories, GIS databases, parallel comput-
ing facilities) and human resources (e.g., active scientists and hyperactive
students trained in amphibian systematics) continues, we anticipate that the
current rate of discovery will continue or increase in the coming years. For
example, although it once seemed that the Amazonian fauna was spatially
quite uniform (albeit highly diverse), denser sampling, exploration of previ-
ously unstudied localities, and analysis of non-traditional data – especially
ESSAY 9.6. THREATENED AMPHIBIANS OF THE WEST INDIES
Eleutherodactylus orientalis (Critically Endangered) from a small flat top
mountain in eastern Cuba called El Yunque de Baracoa. The very small
range and declining quality of habitat in the region are causes for concern.
Adults are only 11mm long, only one millimetre longer than the smallest
frog in the world, E. iberia, which also occurs in eastern Cuba. © S. Blair
Hedges, Penn State
The West Indies is a complex assortment of islands and countries located
between North and South America. In total land area (224,000km²) it is similar
in size to Great Britain and includes three major island groups: a chain of four
large and old islands (the Greater Antilles), a relatively flat limestone bank to
the north (Bahamas and Turks and Caicos Islands), and a classical volcanic
island-arc in the east (the Lesser Antilles). The Greater and Lesser Antilles are
located within, or at the border of the Caribbean Sea, but the Bahamas Bank
is within the Atlantic Ocean. From the standpoint of biodiversity, the West
Indies – sometimes called the “Caribbean Islands” – includes the Cayman
and Swan Islands, but usually excludes islands neighboring Central and South
America (e.g., Aruba, Curaçao, Trinidad and Tobago, etc.), which have faunas
that more closely resemble those on the continents (Figure 1).
The origin of the West Indian fauna has been debated for more than a
century, and continues to be an area of active research. Because the Greater
Threatened Amphibians of the World
DNA sequences – are revealing much greater complexity, and what were
believed to be widespread species are frequently found to involve numer-
ous, even distantly related species of more modest distributions. As our
appreciation of the diversity of Colombian species increases, so too does
our understanding of their basic biology so crucial to implementing effective
conservation strategies to ensure their survival.
A second critical area of research in need of attention is the establishment
of long-term, reliably funded studies that monitor natural populations at key
localities in both pristine and fragmented, or otherwise developing, areas.
Such monitoring programs would allow researchers to track the spread of
infectious diseases, understand the response of individual species to differing
pressures, and distinguish natural and normal population fluctuations from
extreme and abnormal declines, all of which is necessary to design and
implement an efficient conservation strategy to ensure the survival of one
of the world’s most diverse and fascinating amphibian faunas.
Taran Grant, Andrés R. Acosta-Galvis and John D. Lynch
Antilles were once connected, as a geological unit, with North and South
America in the late Cretaceous (~60-70 million years ago), it has been sug-
gested that the present fauna arose by “vicariance” – in other words, trav-
eled with the islands as they broke away from the continents. But fossil and
genetic research has failed to identify more than a few West Indian groups
that fit this model, if any. Most, or all, groups probably arrived to the West
Indies by flying, swimming, or floating on flotsam (mats of vegetation). The
east to west direction of ocean currents means that the source for almost
all flotsam in the West Indies is South America (or, more rarely, Africa), and
this agrees with the evolutionary affinities of much of the non-flying land
fauna (Hedges 2001, 2006).
With its 172 native species, the amphibian fauna of the West Indies is
remarkably diverse for such a small land area. Yet it is also peculiar in that
all the native amphibians are frogs – there are no salamanders or caecilians
– and most (147 species) belong to a single genus of direct-developing lep-
todactylid frogs, Eleutherodactylus (Schwartz and Henderson 1991; Hedges
1999). West Indian amphibians range in adult size, from the smallest frog
in the world, Eleutherodactylus iberia of eastern Cuba, at 10mm, to the gi-
ant ditch frog Leptodactylus fallax (the “Mountain Chicken”) of the Lesser
Antilles, reaching 210mm.
Frogs of the genus Eleutherodactylus (see Essay 1.4) lay their eggs on
land, bypassing the aquatic tadpole stage, which eventually hatch into min-
iatures of the adults. One species of Eleutherodactylus in Puerto Rico even
gives birth to living froglets. Parental care is common among the species,
and many guard their eggs during development. The sex of the egg-guarder
follows evolutionary lines, with the father having this job in the Puerto Rican
group of species, whereas in Jamaica the mother is usually the guarder
(Townsend 1996). Individual species are adapted to a great many terrestrial
niches, including underground burrows, rock caves, cliffs, salt marshes,
waterfalls, bromeliads, tree holes, leaf litter, and vegetation of all types.
The term “ecomorph” has been used with West Indian Eleutherodactylus
to recognize the morphological and ecological convergence in species from
different islands and their adaptations to these niches.
Other native amphibians include a modest radiation of toads (11 species,
family Bufonidae) and an assortment of treefrogs (nine species, family Hyli-
dae). With eight species, Cuba is the center of diversity for toads, whereas
Jamaica and Hispaniola are hot spots of hylid frog diversity, with four species
each. There are several species of aquatic ditch frogs (Leptodactylus) as well,
including the Mountain Chicken that occurs on Montserrat and Dominica. A
single species of dendrobatid frog occurs on Martinique. All but a few species
of West Indian amphibians are endemic to a single island.
Many species occur in the lowlands, but a peak in species density occurs
between 550 and 1,150m elevation, usually corresponding to cloud forest
habitat. On average, species body size decreases by about one mm per
100m of increasing elevation, so a lowland species is typically twice as long
Figure 1. Map of the West Indies
(Caribbean Islands).
Eleutherodactylus glanduliferoides
(Critically Endangered) from the
Massif de La Selle of Haiti. This
species is only known from a few
places on the northern slope of these
mountains, just above the densely
populated capital city of Port-au-
Prince. The region is completely
deforested and this species is
possibly extinct. © S. Blair Hedges,
Penn State
References
Acosta-Galvis, A.R. 2000. Ranas, salamandras y caecilias (Tetrapoda: Amphibia) de
Colombia. Biota Colombiana 1:289-319.
Lynch, J.D. and Duellman, W.E. 1997. Frogs of the genus Eleutherodactylus in western
Ecuador. Systematics, ecology, and biogeography. Special Publication. Natural History
Museum, University of Kansas 23:1-236.
Lynch, J.D., Ruiz-Carranza, P.M. and Ardila-Robayo, M.C. 1997. Biogoegraphic patterns of
Colombian frogs and toads. Revista de la Academia Colombiana de Ciencias Exactas,
Físicas y Naturales 21:237-248.
Rueda-Almonacid, J.V., Lynch, J.D. and Amézquita, A. 2004. Libro Rojo de Anfibios de Co-
lombia. Conservación Internacional Colombia, Instituto de Ciencias Naturales-Universi-
dad Nacional de Colombia, Ministerio del Medio Ambiente, Bogotá, Colombia.
(~56mm) as one on a mountaintop at 2,500m (Hedges 1999). The number of
sympatric (co-occurring) species varies among and within islands, with the
highest number recorded being 24 species near the Haitian village of Castillon,
in the Massif de la Hotte of Hispaniola.
From a conservation standpoint, the West Indies is one of the hottest
biodiversity hotspots (Smith et al. 2005). On average humans have destroyed
more than 90% of the original native habitat in the West Indies and it is no
surprise that these forest-dwelling species have been decimated. Clearing
of land is often for subsistence farming, but trees are also sold for building
materials or made into charcoal for cooking fuel. Charcoaling is practiced in
Jamaica, Cuba, and Hispaniola and is one of the major sources of income
in Haiti where the human population has soared to over eight million and
where essentially no original forests remain (Hedges and Woods 1993;
Young et al. 2004).
A recent assessment of the status of West Indian amphibians found that
84% of the species are threatened (Young et al. 2004), with 37% listed as
Critically Endangered, 36% as Endangered, and 11% as Vulnerable. There is
no other region of the world that has such a high proportion of threatened
species. Among those 63 species listed as Critically Endangered, eight are
considered to be “possibly extinct” because they have not been seen in
many years (Hedges 1993, 1999; Young et al. 2004). These include the live-
bearing species Eleutherodactylus jasperi of Puerto Rico, as well as several
stream-dwelling species.
Besides the major threat from deforestation, a few species have disap-
peared from forested areas and the reason for this is unclear. Certainly, no
forests in the West Indies are pristine because of introduced flora and fauna
that impact the native biota. For example, Old World rats and mongooses
can be encountered throughout forested areas high on mountains in the
Greater Antilles and these species are known to prey on amphibians. Still,
it remains to be established whether introduced predators, climate change,
a chytrid fungus, another threat, or rather a combination of these factors is
the primary cause for the presumed extinctions of these amphibians (Bur-
rowes et al. 2004).
Most countries have made some efforts to control deforestation, such as
the designation of national parks and protected areas. These efforts are to be
applauded, but unfortunately most have had limited or no success in slowing
the destruction of habitat. This is especially true in the countries such as Haiti
and the Dominican Republic, where clear-cutting and charcoaling continue
within protected areas, mostly because budgets allocated to environmental
protection are insufficient. Essentially no original forests remain in Haiti
(Hedges and Woods 1993), and therefore many endemic species – including
those not found in the Dominican Republic – will likely become extinct in
the near future unless something is done soon. Species in other countries
may not be far behind.
It is imperative that international agencies, both conservation-based and

otherwise, step in to help protect the environment in Haiti and elsewhere in
the West Indies before it disappears. Salaries for park guards and logistical
support (e.g., vehicles, communication equipment, and supplies) for existing
parks would provide the most immediate benefit. An accurate knowledge
of the amphibian fauna is also a pre-requisite for conservation efforts, as
is basic systematic work, which needs to be supported through increased
training of local scientists, better education of government officials charged
with protecting the environment (e.g., scientists do not pose a threat!), and
greater facilitation of work by foreign scientists. Together with prompt
and vigorous support directed towards existing protected areas, the real
threats facing the amphibian fauna of the West Indies may be reduced
– if we are lucky.
S. Blair Hedges
ESSAY 9.7. SPREAD OF DISEASE IN LATIN AMERICAN AMPHIBIAN POPULATIONS
In 2005-2006, the world watched as avian flu spread across Asia and into
Europe. As we write this, Americans are expecting the disease to cross the
Atlantic and infect birds in the United States. Fortunately, at least in the case
of bird flu, we have an example of a global monitoring system that can track
the spread of disease across geopolitical boundaries, as well as a network
of health-care providers that are prepared to treat patients infected by the
virus. Although many people will be saved, many will still suffer from this
emerging infectious disease despite the benefit of advanced warning systems,
treatment facilities, and trained personnel. But let us consider a different
hypothetical situation. What would happen if:
• Entire populations of humans were dying from an unknown disease in
remote regions of the world?
• Few scientists or doctors were aware of the situation?
• Nobody contacted the news media or policy makers?
• We realized that this was an epidemic, but we had no monitoring network
or treatment facilities?
• The few personnel trained to detect and potentially treat the disease
were distantly located and had neither financial resources nor sufficient
infrastructure to offer any help?
• We could prevent deaths in hospitals, but nowhere else?
Where and how could we even begin to offer help? With few data, no
dedicated sources of funding, and no infrastructure, the situation would seem
impossible. Yet, to do nothing would clearly be unacceptable. In all respects,
this is the situation for Latin American amphibians, as population after popula-
tion succumbs to the frog-killing fungal disease, chytridiomycosis.
The current crisis of global amphibian extinctions is the result of multiple
causal factors (Collins and Storfer 2003), but none is more insidious than
chytridiomycosis. Considered an emerging infectious disease (Daszak et al.
2000), caused by a recently discovered fungal pathogen, Batrachochytrium
dendrobatidis (Bd), chytridiomycosis has been found infecting more than 80
species of amphibians in Latin America and over 150 species worldwide,
and more areas and species are predicted to be affected in the future (Ron
2005). The disease has been directly implicated in numerous population
declines and extinctions, and is acknowledged to have caused many other
such catastrophes that were not observed. Yet, we still lack a complete
understanding of Bd’s basic biology, pathology, epidemiology, and taxo-
nomic/geographic distribution.
A case study from El Copé, Panama (Lips et al. 2006) illustrates the
devastation that Bd can inflict upon diverse tropical amphibian communities
that are typical of upland areas in Central and South America. Following a
period of eight years of monitoring both amphibian abundance and disease
prevalence, field crews obtained the first Bd-positive sample on 23 September
2004 (Figure 1). Within one week, several dead frogs had been found, and
shortly thereafter as many as 19 dead frogs were found in a single day. After
Figure 1. Timelines of sequential catastrophic declines of amphibians, as
a result of chytridiomycosis, in Costa Rica and Panama. Modified from Lips
et al. (2006).
An undescribed species of Eleutherodactylus found dead floating in a
stream during a die off near El Cope, Panama in 2004. © Forrest Brem &
Roberto Brenes
Chapter 9. Amphibians of the Neotropical Realm
References
Burrowes, P.A., Joglar, R.L., and Green, D.E. 2004. Potential causes for amphibian declines
in Puerto Rico. Herpetologica 60:141-154.
Hedges, S.B. 1993. Global amphibian declines: a perspective from the Caribbean. Biodi-
versity and Conservation 2:290-303.
Hedges, S.B. 1999. Distribution patterns of amphibians in the West Indies. In: W.E. Du-
ellman (ed.), Patterns of distribution of amphibians: A global perspective, pp. 211-254.
The Johns Hopkins University Press, Baltimore, Maryland, USA.
Hedges, S.B. 2001. Caribbean biogeography: an outline. In: C.A. Woods and F.E. Sergile
(eds.), Biogeography of the West Indies: Patterns and Perspectives, pp. 15-33. CRC
Press, Boca Raton, Florida, USA.
Hedges, S.B. 2006. Paleogeography of the Antilles and the origin of West Indian terrestrial
vertebrates. Annals of the Missouri Botanical Garden 93:231-244.
four months, researchers had found 350 dead frogs, representing 40 species
among eight families of anurans and salamanders, all of which were heavily
infected with Bd. This report was the first definitive link between catastrophic
population declines of amphibians and the sudden appearance of Bd at a site.
Within six months, amphibian abundance had been reduced by more than
75%, with 50% of species missing and almost all species affected to some
degree. Nocturnal surveys prior to October 2004 often produced as many as
170 captures, representing as many as 23 species, but those same transects
now produce no more than five or six frogs, representing only two or three
species. Within a remarkably short space of time, the amphibian community
of El Copé has been devastated by this disease, and is not anticipated to
recover, because populations have been greatly reduced and because Bd
persists at this site.
Although many diseases can impact host populations by causing
temporary or permanent declines in abundance, only recently has disease
been considered a major cause of species extinctions (de Castro and Bolker
2005). Theoretical work on disease ecology predicts that, as an epidemic
infectious disease reduces the abundance of its hosts, there is an increase
in the relative abundance of immune individuals; thus, disease transmission
is reduced to zero, such that the pathogen becomes extinct before the host.
Bd is unusual in that it affects a broad taxonomic array of species, and is
highly virulent killing most infected individuals. These are classic traits of
invasive or novel pathogens, and the arrival of Bd in the Neotropics, and its
subsequent extermination of the native fauna, is similar to the effect smallpox
had when it was introduced from Europe to the Americas by Christopher
Columbus and his crew.
A large number of amphibian species have already been identified as hosts
of Bd, and it is likely that both the environment and the frogs are potential
disease reservoirs (Lips et al. 2006). When populations begin to decline,
they can be wiped out in a short period of time with little chance of recovery
or replenishment from other populations. It is this lethal combination of an
exotic virulent pathogen (with a broad host range) invading a highly endemic
amphibian fauna with small geographic distributions that produces high levels
of species extinction in very short periods of time. In conclusion, we can
expect to see many more losses of amphibian species from the Neotropics,
as this disease continues to expand its range. Worse yet, bioclimatic modeling
(e.g., Ron 2005; and see Essay 11.4) suggests that Bd can survive in many
other parts of the globe, and is likely to infect hundreds, if not thousands, of
additional species in Africa and Asia.
Many questions remain regarding how the relationships among Bd,
amphibian hosts, and environmental conditions produce such a wide range
of amphibian population responses. We are beginning to appreciate the
complex challenges involved in mitigating or even reversing impacts of
emerging infectious diseases. We also know that traditional conservation
efforts that are tied to habitat protection are inadequate by themselves in
such instances, so effective solutions remain elusive. Trade and transport
105
Hedges, S.B. and Woods, C.A. 1993. Caribbean hot spot. Nature 364:375.
Schwartz, A. and Henderson, R.W. 1991. Amphibians and reptiles of the West Indies: de-
scriptions, distributions, and natural history. University of Florida Press, Gainesville.
Smith, M.L., Hedges, S.B., Buck, W., Hemphill, A., Inchaustegui, S., Ivie, M., Martina, D.,
Maunder, M., and Ortega, J.F. 2005. Caribbean islands. In: R.A. Mittermeier, P. Robles-
Gil, M. Hoffmann, J.D. Pilgrim, T.M. Brooks, C.G. Mittermeier, J.L. Lamoreux and G.
Fonseca (eds.), Hotspots revisited: Earth’s biologically richest and most endangered
terrestrial ecoregions, pp. 112-118. CEMEX, Mexico City, Mexico.
Townsend, D.S. 1996. Patterns of parental care in frogs of the genus Eleutherodactylus.
Contributions to West Indian herpetology: a tribute to Albert Schwartz. R. Powell
and R. W. Henderson. Society for the Study of Amphibians and Reptiles, Ithaca,
New York, USA.
Young, B.E., Stuart, S.N., Chanson, J.S., Cox, N.A., and Boucher, T.M. 2004. Disappearing jew-
els: the status of New World amphibians. NatureServe, Arlington, Virginia, USA.
of amphibians and other wildlife may spread Bd (Hanselmann et al. 2004),
but the actual frequency and impact of these factors is not known. Regional
and local climatic conditions may influence the growth and survival of Bd
(Pounds et al. 2006), but until a mechanism is identified that can link global
or regional climate changes to individual mortality or reduced population
growth, we cannot hope to design effective conservation measures that
would mitigate these impacts. At this time, while we can effectively treat
animals in captivity (Nichols et al. 2001), we cannot control the spread of Bd
in the wild, or treat animals or ecosystems in situ. Another problem is that Bd
is difficult to detect because it is microscopic and may persist in ecosystems
for undetermined periods of time.
Chytridiomycosis is an alarming model system for disease-driven extinc-
tions of an entire fauna. For one emerging infectious disease to appear
so suddenly and have such uncontrollable effects on global biodiversity,
demands an immediate response. We are facing the synchronous extinction
of a significant proportion of an entire group of vertebrates, and we propose
that it is no longer correct to speak of global amphibian declines but, more
appropriately, of global amphibian extinctions.
Karen R. Lips and Joseph R. Mendelson III
References
Collins, J.P. and Storfer, A. 2003. Global amphibian declines: sorting the hypotheses.
Diversity and Distributions 9:89-98.
Daszak, P, Cunningham, A.A, Hyatt, A.D. 2000. Emerging infectious diseases of wildlife
– threats to human health and biodiversity. Science 287:443-449.
de Castro, F. and Bolker, B. 2005. Mechanisms of disease-induced extinction. Ecol Let-
ters 8:117-126.
Hanselmann, R., Rodriguez, A., Lampo, M., Fajardo-Ramos, L., Aguirre, A.A, Kilpatrick,
A.M., Rodriguez, J.P. and Daszak, P. 2004. Presence of an emerging pathogen of
amphibians in introduced bullfrogs Rana catesbeiana in Venezuela. Biological
Conservation 120:115-119.
Lips, K.R., Brem, F., Brenes, R., Reeve, J.D., Alford, R.A., Voyles, J., Carey, C., and
Collins, J.P. 2006. Infectious disease and global biodiversity loss: pathogens and
enigmatic amphibian extinctions. Proceedings of the National Academy of Sciences,
USA 103:3165-3170.
Nichols, D.K., Lamirande, E.W., Pessier, A.P. and Longcore, J.E. 2001. Experimental trans-
mission of cutaneous chytridiomycosis in dendrobatid frogs. J. Wildl. Dis. 37:1-11.
Pounds, J.A., Bustamante, M., Coloma, L., Consuegra, J., Fogden, M., Foster, P., LaMarca,
E., Masters, K.L., Merino-Viteri, L.A, Puschendorf, R., Ron, S., Sanchez-Azofeifa, G.,
Still, C., and Young, B. 2006. Widespread amphibian extinctions from epidemic disease
driven by global warming. Nature 439:161-167.
Ron, S. 2005. Predicting the distribution of the amphibian pathogen Batrachochytrium
dendrobatidis in the New World. Biotropica 37:209-221.
 CHAPTER 10. AMPHIBIANS OF THE PALAEARCTIC REALM

Figure 1. Summary of Red List categories Brandon Anthony, J.W. Arntzen, Sherif Baha El Din, Wolfgang Böhme, Dan Palaearctic Realm contains 6% of all globally threatened amphibians. The Palaearctic accounts
for amphibians in the Palaearctic Realm. CogĄlniceanu, Jelka Crnobrnja-Isailovic, Pierre-André Crochet, Claudia Corti, for only 3% of CR species and 5% of the EN species, but 9% of the VU species. Hence, on the
The percentage of species in each category Richard Griffiths, Yoshio Kaneko, Sergei Kuzmin, Michael Wai Neng Lau, basis of current knowledge, threatened Palaearctic amphibians are more likely to be in a lower
is also given. Pipeng Li, Petros Lymberakis, Rafael Marquez, Theodore Papenfuss, Juan category of threat, when compared with the global distribution of threatened species amongst
Manuel Pleguezuelos, Nasrullah Rastegar, Benedikt Schmidt, Tahar Slimani, categories. The percentage of DD species, 13% (62 species), is also much less than the global
Max Sparreboom, ùsmail Uøurtaû, Yehudah Werner and Feng Xie average of 23%, which is not surprising given that parts of the region have been well surveyed.
Red List Category Number of species Nevertheless, the percentage of DD species is much higher than in the Nearctic.
Extinct (EX) 2 Two of the world’s 34 documented amphibian extinctions have occurred in this region: the
Extinct in the Wild (EW) 0 THE GEOGRAPHIC AND HUMAN CONTEXT Hula Painted Frog Discoglossus nigriventer from Israel and the Yunnan Lake Newt Cynops
Critically Endangered (CR) 13 wolterstorffi from around Kunming Lake in Yunnan Province, China. In addition, one Critically
Endangered (EN) 40 The Palaearctic Realm includes northern Africa, all of Europe, and much of Asia, excluding Endangered species in the Palaearctic Realm is considered possibly extinct, Scutiger macu-
Vulnerable (VU) 58 the southern extremities of the Arabian Peninsula, the Indian Subcontinent (south of the latus from central China. This represents 1% of the 130 possibly extinct species in the world.
Near Threatened (NT) 48 crest of the Himalaya), Southeast Asia, and the southern parts of China. It is by far the Clearly, as with the Nearctic, there is little in the way of overall discernible pattern so far in
Least Concern (LC) 245 largest of the six biogeographic realms. Palaearctic amphibian extinctions.
Data Deficient (DD) 62 ThePalaearctic Realm isdominated bytheEurasian tectonic plate, which comprises allofEurope Despite a lower overall threat to the amphibian fauna in the Palaearctic compared with
Total Number of Species 468 and most of Asia. Over the past 60 million years, the African, Arabian and Indian plates have collided some other regions, many widespread Least Concern species are seriously threatened in many
0.4% 3%
with and compressed the Eurasian plate, forming an extensive array of mountain ranges, generally parts of their ranges. Many countries in Europe, report high levels of threat to their amphibian
13%
9% running in an east-west direction. These ranges include: the Pyrenees; the Alps; the Carpathians; the faunas. For example, Switzerland lists 70% of its species as nationally threatened (Schmidt and
Balkan mountains; the Caucasus; the mountains of Turkey, Iran, Afghanistan, Pakistan and central Zumbach 2005), even though only one of these species is globally threatened.
Asia; and the Himalaya. The northern part of the African Plate and most of the Arabian plate are
12% now dominated by a Palaearctic fauna, whereas the fauna on the Indian Plate is overwhelmingly
Indomalayan (as is the fauna on the south-eastern part of the Eurasian Plate). Although the Palaearc- SPECIES RICHNESS AND ENDEMISM
tic has an extensive land connection with the Afrotropical Region, there is relatively little mixing of
the faunas, due to the barrier to dispersal formed by the Sahara and Arabian Deserts, particularly Species Richness and Endemism Across Taxa
10% for organisms such as amphibians. However, there is much more overlap with the Indomalayan
fauna, and the boundary between the two regions is hard to delineate in China. The 468 native amphibian species in the Palaearctic Realm represent 8% of the currently
52% For the reasons given above, the southern part of the region is much more mountain- known global total of 5,915 species. Of these 468 species, 260 (or 56%) are endemic to the
ous than the northern part, and includes the highest mountains in the world, peaking at Palaearctic (Table 1). Salamanders, newts, frogs and toads are present in the region, but there
Sagarmatha (Mount Everest) at 8,850m. All of the world’s mountains higher than 8,000m are no caecilians. Anurans account for over three-quarters (77%) of the species. Endemism is
occur in the Himalayan and Karakorum ranges. much higher in the salamanders and newts (86%) compared with the frogs and toads (47%).
Boreal coniferous forests dominate the northern part of the region, through Scandinavia This is presumably a reflection of the generally much smaller range sizes among salamanders,
and Russia. Further to the south, both the western and eastern parts of the region were and that the families Salamandridae and Hynobiidae radiated mainly in the Palaearctic
originally covered in mixed and broad-leaved temperate forests, giving way to steppe lands (see, for example, Weisrock et al. 2006). Of the 15 amphibian families that are native to the
in the central part of the region. However, very little remains of the original temperate forest region, only three are endemic (Table 1). Only 19 species (4% of the species in the region)
habitat through most of the region. In keeping with the topography, habitat diversity is greater are members of these endemic families, although the salamander family Hynobiidae occurs
in the southern part of the region, ranging from deserts, to grassland, to Mediterranean-type only marginally in the Indomalayan Region and is predominantly Palaearctic.
scrubland, and to coniferous forests. Under current climatic conditions, there is essentially no isolation between the Pa-
The Palaearctic Realm has a very uneven human population density (approximately 32 laearctic and Indomalayan Regions, especially in China, and the boundary between these
people per square kilometre in Europe in 2005, compared with 137 in China, 339 in Japan, two faunas is somewhat arbitrary. The effect of this indistinct boundary is to reduce the
but only 8 in Russia and 11 in Saudi Arabia). The percentage of the population living in rural level of endemism of both regions. Summaries of the amphibian fauna of the Palaearctic
areas also varies greatly, ranging from 11.5% in Saudi Arabia, to 26.7% in Europe and Rus- are provided by Borkin (1999) and Zhao (1999).
sia, 34.3% in Japan, and 59.5% in China. In general, most of the people in the region are There are 66 genera (14% of the global total) occurring in the region, of which 22 (33%)
concentrated in eastern Asia and western Europe. The gross income per capita also varies are also endemic. Endemism at the generic level is much higher among the salamanders and
hugely across the region, from US$1,740 in China in 2005, to US$4,460 in Russia, US$11,770 newts (72%) than it is among the frogs and toads (10%), a pattern mirrored in the Nearctic.
in Saudi Arabia, US$27,900 in western Europe, and US$38,390 in Japan. Economic growth The most speciose genera in the region are Rana2 (83 species), Bufo2 (39 species), Paa (24
rates are very high in China (reaching 10% annually), and now averaging 6% in eastern species), Hynobius (24 species), Xenophrys (23 species) and Amolops (20 species). There
Europe, Russia, and central Asia, but much lower (<3%) in western Europe and Japan. are eight monotypic genera endemic to the Palaearctic Realm, all of which are salamanders.
The high economic growth rate in China, and high overall wealth in Europe, as well as high The 44 non-endemic genera in the Palaearctic include 37 frog genera (14 genera from
Pachyhynobius shangchengensis (Vulner- human population densities, explain why damage to natural ecosystems has been so extensive in the Ranidae, nine from the Megophryidae, seven from the Rhacophoridae, four from the
able) is a member of the Asian salamander the eastern and western parts of the region. In China, but in Europe and the Middle East to an even Microhylidae, and one each from the Bombinatoridae, Bufonidae and Hylidae) and seven
Family Hynobiidae. It is known only from the greater extent, humans modified extensive areas of natural habitat over 1,000 years ago. High levels salamander genera (five from the Salamandridae, and one each from the Cryptobranchidae
Dabieshan area in central China, and it lives of habitat modification are still ongoing in China and many other parts of the region. Conversely, in and the Hynobiidae).
in slow-flowing hill streams in forested areas, the central and northern parts of the region, many ecosystems are still largely intact. As noted, 15 of the world’s 48 amphibian families (31%) occur in the Palaearctic, of
where its larvae also develop. Over-harvesting which three are endemic: Discoglossidae, Pelobatidae, and Pelodytidae. The characteristics
for human consumption is a major threat to of these families are provided in Chapter 1.
this species. © Arnaud Jamin GLOBAL CONSERVATION STATUS Among the non-endemic families, the majority of Palaearctic species are in the Bufonidae,
Megophryidae, Ranidae, Rhacophoridae, Hynobiidae and Salamandridae. Of the Palaearctic
A total of 113 species (24%) of amphibian in the Palaearctic Realm is considered to be globally Bufonidae, all 39 species are in the widespread genus Bufo3. This family occurs through most
Table 1. The number of Palaearctic amphib- threatened or Extinct (see Figure 1). This is significantly less than the global average of 33%1. The of the region, and 20 species (51%) are shared with other regions (mainly Indomalayan, but
ians in each taxonomic Family present in also Afrotropical). All Palaearctic bufonids breed by larval development, and some of these
the region. occur in a wide variety of different habitats.
There are 72 species of Megophryidae in the Palaearctic in nine genera. This family is
predominantly Indomalayan, but 30% of the species are endemic to the Palaearctic, occur-
Family Native species Percentage of Percentage of Native genera Percentage of Percentage of ring mainly in central China. They occur especially in mountainous regions, and the genus
(endemics to species in region species in family (endemics genera in region genera in family Scutiger occurs as high as 4,000m in eastern Tibet.
region) that are endemic that are endemic to region) that are endemic that are endemic
The Ranidae (true frogs) constitute the largest family in the Palaearctic, accounting for
to region to region
just over one-third of the total amphibian fauna of the region. Just over half of the species
Anura are in the genus Rana4. The family is found in almost all parts of the region, occurring in
Bombinatoridae 7 (6) 86 60 1 (0) 0 0 most habitats, and breeds by larval development (except the genus Ingerana (four species
Bufonidae 39 (19) 49 4 1 (0) 0 0 in the Palaearctic) which are believed to breed by direct development).5
Discoglossidae 12 (12) 100 100 2 (2) 100 100 The Rhacophoridae (Asian treefrogs) is another predominantly Indomalayan family, with
Hylidae 13 (6) 46 14 1 (0) 0 0 39 species occurring in the Palaearctic, only six of which are endemic. All the Palaearctic
Megophryidae 72 (38) 53 30 9 (0) 0 0 species are in the east of the region (mainly in China). In this region, most of the species
Microhylidae 13 (2) 15 0.5 4 (0) 0 0 breed by larval development, some using foam nests, but a few in the genus Philautus are
Pelobatidae 4 (4) 100 100 1 (1) 100 100 probably direct developers.
Pelodytidae 3 (3) 100 100 1 (1) 100 100 The Asian salamanders (Hynobiidae) are mainly a Palaearctic family centred on Japan
Ranidae 158 (71) 45 11 14 (0) 0 0 and China, with a few species ranging further west as far as Iran, Afghanistan, ex-Soviet
Rhacophoridae 39 (6) 15 2 7 (0) 0 0 Central Asia and north of European Russia. There are 43 species (87% of the global total
TOTAL ANURA 360 (168) 47 3 41 (4) 10 1 for the family) in the region, 40 of which are endemic (and see Essay 10.1).
Caudata The Salamandridae are more diverse in the Palaearctic than elsewhere, with 61% of the
Cryptobranchidae 2 (1) 50 33 1 (0) 0 0 species occurring in the region. The family also occurs in the northern part of the Indomalayan
Hynobiidae 43 (40) 93 87 7 (6) 86 86 region, and there are also six species in the Nearctic. They are widespread in the region, but
Plethodontidae 8 (8) 100 2 2 (2) 100 7 are especially diverse in Europe and China. Most species lay eggs and have free-living aquatic
Proteidae 1 (1) 100 17 1 (1) 100 50 larvae, but 12 species (in the genera Salamandra and Lyciasalamandra) are live-bearers that
Salamandridae 54 (43) 80 61 14 (9) 64 56 do not lay eggs. In some of these species, the young are nourished inside the mother from
TOTAL CAUDATA 108 (93) 86 17 25 (18) 72 29 the embryos of their siblings that die before birth, a unique process in amphibians.
TOTAL ALL AMPHIBIANS 468 (260) 56 4 66 (22) 33 5 The Palaearctic is also important for three other smaller families. The fire-bellied toads
(Bombinatoridae), 60% of which occur in the Palaearctic, are widespread in the region,
 Chapter 10. Amphibians of the Palaearctic Realm 107

and also extend into Indomalaya. Two of the three species of Cryptobranchidae (giant
salamanders) occur in the Palaearctic, one in China, the other in Japan. These are the
largest amphibians in the world, and are associated with clear streams where they breed
by larval development. Finally, one of the six species of Proteidae occurs in the Palaearctic
(the other five being in the Nearctic). The Palaearctic species is the Olm Proteus anguinus
(VU) which occurs in underground streams in karstic landscapes, in north-eastern Italy,
Slovenia, Croatia and a few locations in Bosnia-Herzegovina. It lays eggs, and the adults
retain their larval form.
The remaining families include the Hylidae (treefrogs: a large family occurring mainly in
the Americas, New Guinea and Australia, with just 13 species in the region, all in the genus
Hyla), Microhylidae (narrow-mouthed frogs: globally widespread, with just 13 species in
the region, none of which are endemic), and the Plethodontidae (lungless salamanders: a
large mainly Nearctic and Neotropical family, with just eight species in the region, seven
centred on Italy and one in Korea).
The threatened and extinct species in the Palaearctic show very distinct taxonomic
patterns (Table 2). Salamanders, which account for less than one-quarter of the species in Figure 2. The species richness of amphibians in the Palaearctic Realm, with darker colours The Alpine Newt Triturus alpestris (Least Con-
the region, constitute 45% of the threatened or extinct species, with high levels of threat in corresponding to regions of higher richness. Colour scale based on 10 quantile classes; cern), in the Family Salamandridae, is widely
all families. Among the larger salamander families, 56% of the Hynobiidae are threatened, maximum richness equals 40 species. distributed in Europe, occurring in both alpine
and over 40% of the Salamandridae. In both cases, habitat loss and the impact of excessive and lowland habitats including forests, mead-
utilization are the most common threats. This tendency for salamanders to show very high ows and pastureland. The species breeds,
levels of threat is found also in the Indomalayan, Nearctic and the Neotropical Realms. The and the larvae develop, in stagnant waters,
high level of threat in the Hynobiidae is probably related to the small geographic ranges of including shallow ponds, temporary pools, lakes,
many of the species, and their narrow ecological niches in mountainous regions ditches, drinking troughs, ruts and sometimes
Most of the threatened frog species occur in the larger families, Megophryidae and slow-moving streams. © Henk Wallays
Ranidae (Table 2). The Megophryidae (Asian spadefoots) has a high percentage (one-third)
of threatened species, largely due to habitat loss in China, and often very small geographic
ranges. These species are usually dependent on clear mountain streams for breeding, a
very threatened habitat. The threatened Palaearctic Ranidae (which account for half of the
threatened frogs in the region) are overwhelmingly concentrated in China, where they are
generally threatened both by over-harvesting for human food, and by habitat loss.
Remarkably, the toads (Bufonidae), which in most other regions show high levels
of threat, have no threatened species at all in the Palaearctic. In this region, the family
is dominated by widespread, generally adaptable species. There are also low levels of a.
threat in the Rhacophoridae. There are no globally threatened Palaearctic species in
the Hylidae and the Microhylidae. Among the small endemic families, threat levels are
25% in the Discoglossidae (painted frogs and midwife toads) and Pelobatidae (European
spadefoots), but, at least on the basis of current knowledge, negligible in the Pelodytidae
(parsley frogs). It is perhaps noteworthy that 31% of Palaearctic species in the primitive
suborder Archaeobatrachia (Bombinatoridae, Discoglossidae, Megophryidae, Pelobatidae
and Pelodytidae) are globally threatened, compared with an average of 17% for anurans
as a whole in the Palaearctic.
The great majority (87%) of the threatened amphibians in the Palaearctic are in the Endan-
gered and Vulnerable categories. Again, the generally low number of Critically Endangered
species masks some important family-level differences, with 12% of the Palaearctic Hyno-
biidae, and 50% of the Palaearctic Cryptobranchidae falling into this category. Salamanders
account for 69% of the Critically Endangered species in the region.

Geographic Patterns of Species Richness and b.

Endemism
A map of overall species richness of amphibians in the Palaearctic Realm (Figure 2) shows
that species richness is low through most of the region, especially in most of Russia and
central Asia. In dry areas, such as in much of the Arabian Peninsula and on the Tibetan
Plateau, and in very cold areas such as the northernmost belt of the region, there are no
amphibians at all. Species richness is highest in China, and moderate in parts of Europe,
Japan, and Korea. The high richness in China is partly a reflection of certain species of
Indomalayan origin occurring there, as well as the absence of glaciations in much of central
and southern China during the Pleistocene ice ages. The Palaearctic Realm is generally
well studied, and Figure 2 probably reflects genuine overall patterns of amphibian species
richness, though more species are still being discovered regularly, especially in the eastern
parts of the region, and sometimes even in western Europe.
Threatened species (Figure 3a) in the Palaearctic are overwhelmingly concentrated in
central China (especially in Sichuan and northern Yunnan). Elsewhere, there is little discern-
ible pattern, but with some threatened species in Mediterranean Europe, north-west Africa,
Asia Minor, central Asia, and Japan. Not surprisingly, given the small number of species
involved, there are no noteworthy concentrations of Critically Endangered species in the
region (Figure 3b) (much of the apparent concentration in China reflects the wide distribu-
tion of one species, the Chinese Giant Salamander Andrias davidianus, which is subject to
over-exploitation as a food source).
Figure 3. a) The richness of threatened amphibians in the Palaearctic Realm, with darker
colours corresponding to regions of higher richness. Colour scale based on five quantile
classes; maximum richness equals 11 species. b) The richness of CR amphibians in the
Palaearctic Realm. Maximum richness equals one species.
Species Richness and Endemism within Countries
Amphibians are native in 81 countries and territories in the Palaearctic Realm (Figure 4). China
has by far the largest number of species and of endemics in the region (265 species, of which
137 are endemic). More than half (57%) of Palaearctic species occur in China, and 29% are
endemic to this country (see Essay 10.2). India, although second on the list, extends only
marginally into the Palaearctic, and consequently has no endemics in the region. Only four
other countries (Japan, Italy, France and Spain) have more than 30 species; remarkably, the
Russian Federation, despite its huge size, has only 28 species and no endemics, due to the
severe continental climate over a large part of its territory. The low number of species and the
low level of endemism in most countries are caused in part by low overall species richness, Table 2. The number of species within each
but also by the small size of many of the countries in the region. After China, only Japan and IUCN Red List Category in each Family and
Order in the Palaearctic Realm. Introduced
species are not included.

The Oriental Fire-bellied Toad Bombina orientalis (Least Concern) is one of ten species in
the Family Bombinatoridae. It is widespread in the Korean peninsula, north-eastern China, Family EX CR EN VU NT LC DD Total number Number Threatened % Threatened
and parts of the Russian Far East. Although generally common, there is concern about of species or Extinct or Extinct
the level of harvest for traditional Chinese medicine, and for the international pet trade. Anura
© Twan Leenders Bombinatoridae 0 0 0 2 0 5 0 7 2 29
Bufonidae 0 0 0 0 3 31 5 39 0 0
Discoglossidae 1 0 0 2 4 5 0 12 3 25
Hylidae 0 0 0 0 0 12 1 13 0 0
Megophryidae 0 2 11 11 5 27 16 72 24 33
Microhylidae 0 0 0 0 0 13 0 13 0 0
Pelobatidae 0 0 1 0 1 2 0 4 1 25
Pelodytidae 0 0 0 0 0 3 0 3 0 0
Ranidae 0 2 11 18 14 89 24 158 31 20
Rhacophoridae 0 0 0 1 5 23 10 39 1 3
TOTAL ANURA 1 4 23 34 32 210 56 360 62 17
Caudata
Cryptobranchidae 0 1 0 0 1 0 0 2 1 50
Hynobiidae 0 5 8 11 2 11 6 43 24 56
Plethodontidae 0 0 1 2 4 1 0 8 3 38
Proteidae 0 0 0 1 0 0 0 1 1 100
Salamandridae 1 3 8 10 9 23 0 54 22 41
TOTAL CAUDATA 1 9 17 24 16 35 6 108 51 47
TOTAL ALL AMPHIBIANS 2 13 40 58 48 245 62 468 113 24
108 Threatened Amphibians of the World

China* China* Turkey

India* Japan*
Italy Turkey Algeria
Japan*
France China* Japan* China*
Spain
Russian Federation Italy Greece
Turkey Tajkistan
Greece Greece Japan*
Czech Republic Iran Spain Iran
Germany
Serbia and Montenegro Iran Italy
Austria Italy
Croatia Algeria Afghanistan
Iran
Slovenia Turkey India* Iraq
Portugal
Romania Albania Morocco
Slovakia Korea, Republic
Croatia Spain
Ukraine
Bosnia and Herzegovina France Tunisia
Hungary Afghanistan
Switzerland Morocco Albania
Bulgaria Morocco
Pakistan* Slovenia Croatia
Korea, Republic
Poland Afghanistan Slovenia
Greece
Albania Bosnia and Kazakhstan
Belgium Herzegovina
Netherlands Egypt Georgia Georgia
Denmark
Korea, D.P.R. Iraq Korea, D.P.R.
Luxemburg Sudan*
Macedonia, F.Y.R. Kazakhastan Korea, Republic
Belarus
Algeria Spain Korea, D.P.R. France
Georgia Korea, Republic Bosnia and
Latvia Herzegovina
France
Moldova Portugal India*
Morocco
Sweden Serbia and Switzerland
Algeria Montenegro
Kazakhstan
Lithuania Switzerland Portugal
Azerbaijan Pakistan* Serbia and
Liechtenstein Tunisia Montenegro

0 50 100 150 200 250 300 0 20 40 60 0 20 40 60 80 0 10 20 30 40

Number of species Percentage of endemic species Number of threatened species Percentage of species threatened
Endemics Non-endemics Endemics Non-endemics
Figure 4 Figure 5 Figure 6 Figure 7

Figure 4. The number of amphibians present

in and endemic to each Palaearctic country.
*denotes countries not entirely within the
Palaearctic Realm, hence only the species
whose ranges fall within the region are in-
cluded. Only countries with at least 10 native
species are included.
Figure 5. Percentage of species endemic to
each Palaearctic country. *denotes countries
not entirely within the Palaearctic Realm,
hence only the species whose ranges fall
within the region are included. Only coun-
tries with at least one endemic species are
included.
Figure 6. The number of globally threatened
amphibians present in and endemic to each
Palaearctic country. Countries with no glob-
ally threatened species are not included in
the diagram. *denotes countries not entirely
within the Palaearctic Realm, hence only the
species whose ranges fall within the region
are included.
Figure 7. Percentage of native species that
are globally threatened. Countries with no
globally threatened species are not included
in the diagram. *denotes countries not entirely
within the Palaearctic Realm, hence only the
species whose ranges fall within the region
are included.
Italy have more than 10 endemic species (25 and 11, respectively). Thirty-six countries in the
region have fewer than 10 native amphibian species (not shown on Figure 4), and only four
of these (Afghanistan, Egypt, Sudan and Tajikistan) have endemics (one each).
The amphibian fauna of the Palaearctic has been well summarized in extensive review
literature. Important regional works include: Ananjeva and Borkin (1981), Arnold (2002, 2003),
Balletto et al. (1985), Borkin (1984, 1999), Cog lniceanu et al. (2000), Corti et al. (1999),
Duguet and Melki (2003), Gasc et al. (1997), Griffiths (1996), Grossenbacher and Thiesmeier
(1999, 2003, 2004), Kuzmin (1996, 1999), Kuzmin and Semenov (2006), Leviton et al. (1992),
Nöllert and Nöllert (1992), Pleguezuelos (1997), Salvador (1996), Schleich et al. (1996),
Tarkhnishvili and Gokhelashvili (1999), Thorn (1968), Thorn and Raffaëlli (2001) and Zhao
(1999). There are numerous national-level publications, including, but by no means limited
to: Arnold (1995), Baran and Atatür (1998), Bauwens and Claus (1996), Bons and Geniez
(1996), Cabela and Tiedemann (1985), Castanet and Guyetant (1989), Crespo and Oliveira
(1989), Disi (2002), Fei et al. (1999, 2005), García-París (1985), García-París et al. (2004),
Geniez et al. (2005), Goris and Maeda (2004), Groenveld (1997), Grossenbacher (1988), Khan
(2006), Kuzmin et al. (1998), Llorente et al. 1995, Maeda and Matsui (1999), Malkmus (2004),
Moravec (1994), Parent (1979), Pleguezuelos et al. (2002), Puky et al. (2003), Saleh (1997),
Salvador and García-Paris (2001), Schleich and Kästle (2002), Scoccianti (2002), Sindaco et
al. (2006), Societas Herpetologica Italica (1996), Uchiyama et al. (2002), Werner (1988), Ye
et al. (1993), Zhao and Adler (1993), and Zhao et al. (2000).
Although China has many more endemics than any other country in the region, Japan has
the highest percentage of endemic species at almost 70% (Figure 5), because of the insular
nature of the country. In the Palaearctic part of China, over 50% of the amphibians are endemic.
In addition to species-poor Tajikistan (already mentioned), high levels of endemism (>20%)
also exist in Iran, Italy, and Turkey. In both Japan and China, the levels of endemism are high
in frogs and salamanders. In Iran, Italy and Turkey, the endemics are mainly salamanders.
China has many more threatened species (64) than any other country in the Palaearctic
(Figure 6). Well over half of the threatened species in the region (57%) occur in China, and
50% are endemic to China. Endemic species in China are much more threatened (45%)
than non-endemics (2%). The number of globally threatened species is less than 10 in all
other countries in the region (Figure 6). The percentage of threatened amphibian species
is highest in Turkey (35%), and is greater than 20% in Algeria, China, Greece, Japan, and
Iran (Figure 7). With the exception of Turkey, in all Palaearctic countries, the level of threat
is much lower than the global average of 33%. The high percentage of threatened species
in Turkey is a reflection of the presence of nine species of salamanders with small ranges,
all threatened by habitat loss.
These figures all relate to the number of globally threatened amphibian species in each
country. Many countries, especially in Europe, list many globally Least Concern species as
nationally threatened. Assessments of the conservation status of Palaearctic amphibians
have been carried out in several countries, including, for example: Switzerland (Grossen-
bacher 1994; Schmidt and Zumbach 2005); Japan (Japan Agency of Environment 2000; Ota
2000); Spain (Pleguezuelos et al. 2002); and China (Zhao 1998; Xie and Wang, 2004). Corbett
(1989) published an overview of amphibian conservation in Europe, while Cox et al. (2006)
presented a review of the status and distributions in the Meditteranean (and see Essay 10.3).
Xie et al. (2007) proposed a conservation plan for the amphibians of China.
There are only 13 Critically Endangered Palaearctic species, but seven of these occur in
China, two each in Iran and Japan, and one each in Afghanistan and Turkey. Outside China,
all Critically Endangered species in the Palaearctic are salamanders.
HABITAT AND ECOLOGY
Habitat Preferences
Most Palaearctic amphibians (78%) occur in forests, but almost one-third of the fauna (31%)
can survive in secondary terrestrial habitats (Table 3; Figure 8). This latter figure is almost
double that of the Nearctic Realm, perhaps suggesting a higher percentage of adaptable
species in the Palaearctic. As natural habitats have been completely lost in many parts of
the Palaearctic, some amphibian species are now dependent of artificial habitats. As in other
regions, forest-dwelling and stream-associated amphibians are more likely to be threatened
than those occurring in any other habitats, with over 20% of these species being globally
threatened. This is the combination of habitat preferences that has been associated with
rapid declines in amphibian populations worldwide (Stuart et al. 2004). Amphibians occur-
ring in savannahs, arid and semi-arid habitats, and secondary terrestrial habitats are less
likely to be threatened than those occurring in other habitats (Table 3; Figure 8). At least
one highly threatened species, Ranodon sibiricus (EN), is mainly associated with forests,
but is surviving at much reduced densities in secondary habitats due to the loss of forest
cover (Kuzmin and Thiesmeier 2001).
Reproductive Modes
Larval development is by far the most common reproductive mode in the Palaearctic (94%
of species), compared with 3% for direct development and 3% live-bearing (Table 4; this
compares with the global picture of 68% larval development, 30% direct development,
and 1% live-bearing). The Palaearctic has a smaller proportion of direct-developing species
than in any other region of the world. The direct-developing Palaearctic amphibians include
plethodontid salamanders in the genera Speleomantes and Karsenia, and frogs in the genera
Philautus and perhaps Ingerana. The live-bearing species are all salamandrid salamanders
in the genera Salamandra and Lyciasalamandra. All of the live-bearing salamanders of the
world are found only in the Palaearctic Realm.

Habitat type Number of % of all Globally % of all

Number of species % of species in habitat that are threatened
species in species Threatened species in
each habitat occurring in and Extinct habitat that Forest
the habitat species are globally Savannah
Threatened
or Extinct Grassland

Shrubland
Table 3. The habitat preferences of amphib- Forest 365 78 81 22
Savannah 16 3 0 0 Secondary terrestrial habitats
ians in the Palaearctic Realm.
Grassland 114 24 17 15 Flowing freshwater
Shrubland 150 32 23 15
Marsh/swamp
Figure 8. The habitat preferences of Palaearc- Secondary terrestrial habitats 143 31 15 10
Flowing freshwater 346 74 87 25 Still open freshwater
tic amphibians. The plot on the left-hand side
shows the number of species in the region Marsh/swamp 127 27 17 13 Arid and semi-arid habitats
in each habitat type. On the right-hand side, Still open freshwater 254 54 36 14
0 100 200 300 0 8 16 24
the percentage of these species which are Arid and semi-arid habitats 26 6 0 0
threatened is given.
 Chapter 10. Amphibians of the Palaearctic Realm 109

The Marsh Frog Rana ridibunda (Least Con-

cern) occurs widely from western Europe to
western China, with invasive populations
spreading in several places in the Palaearctic
outside its natural range. It is a highly oppor-
tunistic semi-aquatic ranid frog, living in most
habitats, especially in open, well-warmed
areas with abundant herbaceous vegetation.
It may also be found in slightly saline water.
© Miroslav Samardži

In the Palaearctic, the percentage of globally threatened or Extinct live-bearing am-

Reproductive mode All Species Threatened or % Threatened phibians is much higher than in direct-developing and larval-developing species (Table 4),
Extinct species or Extinct though, in view of the small number of species involved, the significance of this finding
Direct development 14 3 21 is not clear.
Larval development 439 101 23
Live-bearing 13 9 69
Not known 2 0 0 MAJOR THREATS
Habitat loss is overwhelmingly the major threat to amphibians in the Palaearctic (Table 5;
Table 4. Palaearctic amphibians categorized by reproductive mode. Figure 9), affecting over 80% of the threatened species. Pollution and utilization are the
next most serious threats, each impacting over one-third of the threatened species. Invasive
species, disease, human disturbance and natural disasters also have significant impacts on
certain species. Most other threats are of relatively minor importance. Chytridiomycosis has
Threat type Threatened species % Threatened species been recorded in many parts of Europe, with records from seven countries and 20 out of
All habitat loss 90 81 28 species examined (Garner et al. 2005, 2006). It has been associated with some serious
Agriculture – Crops 41 37 declines in Spain, but so far only of non-threatened species (Bosch et al. 2001). However, it
Agriculture – Tree plantations 4 4 does not seem to kill amphibians in most areas where it has been recorded in Europe, and
Agriculture – Livestock 16 14 so its overall impact currently seems to be less severe than in some other regions.
Timber and other vegetation removal 48 43 In terms of the types of habitat loss that are impacting amphibians in the Palaearctic, the
Urbanization and industrial development 59 53 impacts of urbanization and industrial development (affecting over half of the threatened
Invasive species 9 8 species), vegetation removal (mainly logging in Asia) (>40%) and expanding croplands
Utilization 41 37 and agricultural intensification (over one-third) are the most severe. Tree plantations and
Accidental mortality 5 4 livestock are less important threats in most cases. The heavy impact of urbanization and
Pollution 41 37 industrialization reflects the high human population density in China, Japan, and Europe
Natural disasters 11 10 in the areas where amphibian species richness is highest. These factors, as well as the
Disease 2 2 extensive development of agriculture, are also important threats in some parts of the former
Human disturbance 11 10 Soviet Union. The coding of major threats used in the Global Amphibian Assessment does
Fire 5 4 not sufficiently depict habitat loss resulting from the over-exploitation of water resources,
which is increasingly affecting amphibians in the south-western part of the Palaearctic
(especially in the Iberian Peninsula, Greece and northern Africa), resulting in the widespread
Table 5. The major threats to globally threatened amphibians in the Palaearctic Realm. Only loss of breeding sites.
present threats to species are tallied.

All habitat loss

Invasive species

Utilization

Accidental mortality

Persecution

Pollution

Natural disasters

Disease

Human disturbance

Changes in native sp. dynamics

Fire
The Green Toad Bufo viridis (Least Concern)
from the Family Bufonidae has an extremely
0 20 40 60 80 100 wide range in North Africa, Europe, and West
Percentage of threatened species affected and Central Asia. It is one of the most adapt-
able amphibians of the Palaearctic, living in
forests, steppes, dry grasslands, alpine areas,
arid habitats (usually close to oases), and
Figure 9. The major threats impacting threatened amphibians in the Palaearctic Realm. urban areas. © Maik Dobiey
110 Threatened Amphibians of the World

The Common Spadefoot Pelobates fuscus riorating, given the high percentage of decreasing species. One reason for the large number
(Least Concern), one of four species in the of decreasing species is that the majority of the region’s amphibians occur in China, where
Family Pelobatidae, ranges from western habitat loss and over-harvesting are both serious threats. Many species in Europe are also
Europe to central Asia. It occurs mostly in in decline, especially due to agricultural intensification.
open areas, and burrows in loose soil when
not breeding. It breeds in small still water “Rapidly Declining” Species
bodies, including ditches, ponds and lakes.
© Miroslav Samardži Only 42 (9%) of the 470 globally “rapidly declining” species occur within the Palaearctic
Realm (a full list of all “rapidly declining” species is provided in Appendix IV and includes
their occurrence within each of the realms). Eighteen of these species are in decline due to
over-exploitation, 23 due to reduced habitat, and one due to so-called “enigmatic declines”. Not
surprisingly for this region, more declines are attributed to reduced habitat and over-exploitation
than to enigmatic declines. The Palaearctic accounts for 47% of the world’s rapid declines due
to over-exploitation, but only 11% of the reduced habitat declines, and 0.4% of the enigmatic
declines. One species in the region, Nanorana pleskei (NT) from Qinghai, Gansu and Sichuan
Provinces in China, has been recorded as undergoing an enigmatic decline, the causes of this
decline are not known, and have not so far been linked to either chytridiomycosis or climate
change (although these two threats have now been associated with many such declines that
have taken place elsewhere in the world (Lips et al. 2006; Pounds et al. 2006)).
The “rapidly declining” species show a distinct taxonomic pattern (Table 8). Among the
larger families, the Ranidae and Salamandridae show a higher tendency to serious decline
Table 6. The purposes for which amphibians than the Bufonidae, Megophryidae and Hynobiidae. There are no Palaearctic species in rapid
are used in the Palaearctic Realm. The num- Purpose Subsistence Sub-national/ Regional/ Number decline in the two treefrog families, Hylidae and Rhacophoridae, nor in the Microhylidae.
bers in brackets are the number of species National International of species Some small families have high percentages of species in serious decline, most notably the
within the total that are threatened species. Food – human 83 (23) 33 (9) 10 (1) 88 (24) Discoglossidae and Pelobatidae (both families with large tadpoles) and Cryptobranchidae.
Food – animal 3 (0) 0 0 3 (0) Among the larger families, “over-exploited declines” are concentrated in the Ranidae and
Medicine – human and 42 (10) 24 (4) 7 (0) 45 (10) the Salamandridae.
veterinary The “rapid declines” in the Palaearctic also show a clear geographic pattern. The major
Pets, display animals 7 (2) 39 (9) 47 (9) 54 (15) concentration of declines is in China (23 species), followed by Iberia (8), north-western
Research 1 (1) 8 (1) 3 (0) 8 (1) Africa (three), and Iran (two). Of the “over-exploited declines”, 16 are in China, and two
Specimen collecting 1 (1) 7 (7) 8 (8) 9 (9) in Iran (Neurergus newts in the pet trade). The rapid declines in Iberia and north-western
Africa have been linked to loss of habitat due to a long-term drying trend in the climate
(see Pleguezuelos et al. 2002), linked with the over-exploitation of water resources, which
results in the widespread loss of breeding sites.
Table 7. The population trends for all extant
Palaearctic amphibians. Population Trend Number of species % of extant species
Decreasing 246 53 KEY FINDINGS
Stable 126 27
Increasing 5 1 • A total of 468 species are recorded from the Palaearctic Realm, of which 113 (24%) are
Unknown 89 19 considered globally threatened or Extinct.
• At the species level, 260 amphibians (56% of those present) are endemic to the Palaearc-
tic; of the 15 families found in the region, three are endemic, and of 66 amphibian genera
occurring, 22 are endemic. Endemism is lower in the Palaearctic than some other regions,
Table 8. The number of species in “rapid especially due to the unclear and somewhat arbitrary boundary with the Indomalayan
decline” and “over exploited decline” in the Family Number of Percentage Number of Percentage Region, especially in China.
Palaearctic Realm by Family. species in of species species in of species • The percentage of threatened or Extinct species is lower than in many other parts of the
“rapid decline” in family in “over-exploited in family in
“rapid decline” decline” “over-exploited
world, but highest in the families Proteidae (100%), Hynobiidae (56%), Cryptobranchidae
decline” (50%), Salamandridae (41%), Plethodontidae (38%), Megophryidae (33%), Bombinatori-
dae (29%), Discoglossidae (25%) and Pelobatidae (25%).
Bufonidae 1 3 0 0 • Overall, the threat levels are much higher among salamanders and newts (47%) than
Discoglossidae 3 25 0 0 frogs and toads (17%).
Megophryidae 2 3 0 0 • Geographic concentrations of threatened species in the Palaearctic are overwhelmingly
Pelobatidae 2 50 0 0 centred on China; elsewhere there is little discernible pattern, though there are small
Ranidae 20 13 12 8 concentrations in Mediterranean Europe, north-west Africa, Asia Minor, Central Asia
Cryptobranchidae 1 50 1 50 and Japan.
Hynobiidae 2 5 1 2 • China has the largest number of species in the Palaearctic Realm (265 species), and also
Plethodontidae 1 13 0 0 has more endemics (137). Another five countries (India, Japan, Italy, France, and Spain)
Salamandridae 10 19 4 7 have more than 30 species, but only two of these (Japan and Italy) have more than 10
endemics.
• China also has by far the largest number of threatened species (64), all other countries
having fewer than 10 threatened species. However, the percentage of threatened species
A total of 153 species (50 of which are threatened) are recorded as being used in the region is higher in Turkey (35% cf. 24% in China).
(Table 6). The most common reasons for harvesting Palaearctic amphibians are for food (88 • Threatened species tend to show distinct habitat preferences, with forest-dwelling
species, mostly at local and national levels), pet trade (54 species, mostly at national and and stream-associated species being the most frequently threatened (22% and 25%,
international levels), and medicinal use (45 species, mostly at local and national levels). Not respectively). This mirrors patterns seen elsewhere in the world.
all of the harvesting of amphibians in the region is considered to constitute a major threat to • Habitat loss, primarily due to the impacts of urbanization and industrial development,
the species. Of the 153 species being harvested, utilization is considered to be a threat for vegetation removal (mainly logging), expanding croplands and agricultural intensifica-
96 (of which 46 are threatened species for which harvesting is believed to be contributing tion, is affecting over 80% of the threatened species in the region. Over-harvesting and
to a deterioration in their status). Over 60% of these species seriously threatened by over- pollution each impact over one-third of the threatened species. Disease has not been
harvesting occur in China, where many species of amphibians are extensively harvested recorded as a significant threat in the region so far, although chytridiomycosis is spreading
for human food and medicines. Examples include the Chinese Giant Salamander (which has in Europe.
undergone catastrophic declines), 12 species of ranid frogs (eight in the genus Paa), and • The overall trends of Palaearctic amphibians are worse than the global situation, with
six species of hynobiid salamanders. Threatened species outside China that are harvested 53% of the species in decline.
include six species of hynobiid salamanders (in the genus Hynobius) in Japan (mainly pet • Of the 470 globally “rapidly declining” species, 9% occur within the region. Most of
trade), nine species of salamandrid salamanders (in the genera Lyciasalamandra and Neur- these rapid declines (55%) are caused by severe habitat loss, and 43% are due to over-
ergus) in Iran and Turkey (pet trade), and the hynobiid Ranodon sibiricus in Kazakhstan (mainly exploitation. Over-exploitation is a much more serious threat, especially in China, than
medicinal use). A much higher percentage of Palaearctic threatened species is negatively in any other part of the world.
impacted by utilization (37%) than in any other region (Indomalaya is the next highest at • Two amphibian extinctions have been recorded from the Palaearctic, one in China, the
8%). Harvesting of non-threatened amphibians in Russia as food and medicine exports to other in Israel. A third species (from China) is possibly extinct.
China (mainly Rana dybowskii, but also R. amurensis, R. asiatica, R. nigromaculata, Bufo
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7:1-207. Vollzug Umwelt, 48 S.
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4: Schwanzlurche (Urodela) I. Aula-Verlag. Wiesbaden, Germany. Atlante Degli Anfibi E Dei Rettili D’Italia. Polistampa.
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4: Schwanzlurche (Urodela) IIa. Aula-Verlag. Wiesbaden, Germany. del Museo Civico di Storia Naturale “G. Doria”, Genova 91:95-178.
Grossenbacher, K. and Thiesmeier, B. (eds.) 2004. Handbuch der Reptilien und Amphibien Europas, Volume Stuart, S.N., Chanson, J.S., Cox, N.A., Young, B.E., Rodrigues, A.S.L., Fischman, D.L. and Waller, R.W. 2004.
4: Schwanzlurche (Urodela) IIb. Aula-Verlag. Wiesbaden, Germany. Status and trends of amphibian declines and extinctions worldwide. Science 306:1783-1786.
Japan Agency of Environment. 2000. Threatened Wildlife of Japan – Red Data Book. 2nd ed. Reptilia/Am- Tarkhnishvili, D.N. and Gokhelashvili, R.K. 1999. The Amphibians of the Caucasus. Advances in Amphibian
phibia (in Japanese with English summary). Japan Wildlife Research Center, Tokyo, Japan. Research in the Former Soviet Union, 4. Pensoft, Sofia, Moscow, Russia.
Khan, M.S. 2006. Amphibians and Reptiles of Pakistan. Krieger Publishing Company, Malabar, Florida, USA. Thorn, R. 1968. Les Salamandres d’Europe, d’Asia, et d’Afrique du Nord. Éditions Paul Lechevalier, Paris,
Kuzmin, S.L. 1996 Threatened amphibians in the former Soviet Union: the current situation and the main France.
threats. Oryx 30:24-30. Thorn, R. and Raffaëlli, J. 2001. Les Salamanders de L’Ancien Monde. Societé Nouvelle des Editions Endnotes
Kuzmin, S.L. 1999. The Amphibians of the Former Soviet Union. Pensoft, Sofia and Moscow, Russia. Boubée, Paris, France. 1 P<0.001 (binomial test)
Kuzmin, S.L., Borkin, L.J., Vorobyeva, E.I., Darevsky, I.S., Munkhbayar, Kh. and Semenov, D.V. 1998. Uchiyama, R., Maeda, N., Numata, K. and Seki, S. 2002. A Photographic Guide: Amphibians and Reptiles 2 Frost et al. (2006) transfer many of the species to
Amphibians and Reptiles of Mongolian People’s Republic: General Problems. Amphibians. Nauka, in Japan. Heibonsha, Tokyo, Japan. other genera.
Moscow, Russia. Weisrock, D.W., Papenfuss, T.J., Macey, J.R., Litvinchuk, S.N., Polumeni, R., U÷urtaú, I.H., Zhao, E., 3 Frost et al. (2006) transfer many of these species
Kuzmin, S.L. and Thiesmeier, B. 2001. Mountain Salamanders of the Genus Ranodon. Advances in Amphibian Jowkar, H. and Larson, A. 2006. A molecular assessment of phylogenetic relationships and lineage to other genera.
Research in the Former Soviet Union, volume 6. Pensoft Publishers, Sofia-Moscow. accumulation rates within the family Salamandridae (Amphibia, Caudata). Molecular Phylogenetics 4 Frost et al. (2006) transfer many of these species
Kuzmin, S.L. and Semenov, D.V. 2006. Conspect of the fauna of amphibians and reptiles of Russia. KMK, and Evolution 18:434-448. to other genera.
Moscow, Russia. Werner, Y.L. 1988. Herpetofaunal survey of Israel (1950-85), with comments on Sinai and Jordan and on 5 Under Frost et al’s. (2006) arrangement, the
Leviton, A.E., Anderson, S.C., Adler, K. and Minton, S.A. 1992. Handbook to Middle East Amphibians and zoogeographical heterogeneity. In: Y. Yom-Tov and E. Tchernov, (eds.), Zoogeography of Israel, pp. Palaearctic species in the genera Chaparana, Eu-
Reptiles. Society for the Study of Amphibians and Reptiles, Lawrence, Kansas, USA. 355-388. Monographiae Biologicae 62, W. Junk, Dordrecht, The Nerthlands. phlyctis, Fejervarya, Hoplobatrachus, Limnonectes,
Lips, K.R., Brem, F., Brenes, R., Reeve, J.D., Alford, R.A., Voyles, J., Carey, C., Livo, L., Pessier, A.P. and Xie, F., Lau, M.W.N., Stuart, S.N., Chanson, J., Cox, N.A. and Fischman, D.L. 2007. Conservation needs of Nanorana, Occidozyga, Paa, and Sphaerotheca are
Collins, J.P. 2006. Emerging infectious disease and the loss of biodiversity in a Neotropical amphibian amphibians in China: A review. Science in China, Series C, Life Sciences 50:265-276. transferred to the predominantly Indomalayan
community. Proceedings of the National Academy of Sciences, USA 103:3165-3170. Xie, Y. and Wang, S. 2004. China Species Red List, Vol. 1 Red List, Higher Education Press, Beijing, family Dicroglossidae. Most of these species are
Llorente, G. A., Montori, A., Santos, X. and Carretero, M. A. 1995. Atlas dels Amfibis I Rèptils de Catalunya China. in China, but others occur further west in western
i Andorra. Edicions El Brau, Figueres, Spain. Ye, C.-Y, Fei, L. and Hu, S.Q. 1993. Rare and Economic Amphibians of China. Sichuan Publishing House of Asian and even north-western Africa. Some
Maeda, N. and Matsui, M. 1999. Frogs and Toads of Japan. Bun-ichi Sogo, Shuppan, Japan. Science and Technology, Chengdu, China. other Palaearctic “ranids” are transferred to other
Malkmus, R. 2004. The Amphibians and Reptiles of Portugal, Madeira and the Azores-Archipelago. A.R.G. Zhao, E.M. 1998. China Red Data Book of Endangered Animals – Amphibia and Reptilia. Science Press, families under Frost et al.’s proposed classification:
Gantner Verlag K.G. Ruggel, Germany. Beijing, China. one species to the Afrotropical family Pyxicephali-
Maslova, L.V. and Lyapustin, S.N. 2005. Illegal export of amphibians and reptiles from the Russian Far East to Zhao, E.M. 1999. Distribution patterns of amphibians in temperate East Asia. In: Duellman, W.E. (ed.), dae (Tomopterna cryptotis); four species in the
countries of the Asian region: the situation in 2003. In: N.B. Anayeva and O. Tsinenko (eds.), Herpetologia Patterns of Distribution of Amphibians: A Global Perspective, pp. 421-443, Johns Hopkins University genus Ingerana to the Oceanian and Indomalayan
Petropolitana, pp. 191-192, Societas Europaea Herpetologica, St. Petersburg, Russia. Press, Baltimore, Maryland, USA. Ceratobatrachidae; and two species in the genus
Moravec, J. (ed.) 1994. Atlas Rozši eni Obojživlniku v eské Republice. Národní Museum, Prague, Czech Zhao, E.M. and Adler, K. 1993. Herpetology of China. Society for the Study of Amphibians and Reptiles, Ptychadena to the Afrotropical Ptychadenidae.
Republic. Lawrence, Kansas, USA. Even with these changes, the Ranidae have just
Nöllert, A. and Nöllert, C. 1992. Die Amphibien Europas. Franckh-Kosmos Verlags-GmbH & coco, Stuttgart, Zhao, E.M., Chang, H.W., Zhao, H. and Adler, K. 2000. Revised checklist of Chinese Amphibia and Reptilia. over 100 species in the Palaearctic, and remain
Germany. Sichuan Journal of Zoology 19:196-207. easily the largest family in the region.
112
ESSAY 10.1. CURRENT STATUS OF JAPANESE SMALL SALAMANDERS OF THE GENUS HYNOBIUS
The Japanese archipelago includes more than 3,000 islands. In addition
to the four main islands – Hokkaido, Honshu, Shikoku and Kyushu – the
archipelago includes smaller island groups, such as the Ogasawara-shoto
(the Bonin Islands and Iwo or Volcano Islands), and the Nansei-shoto (Ryukyu
and Satsunan Islands) (Figure 1).
The amphibian fauna of Japan is represented by 56 species. Endemism
is marked, with around 45 species found nowhere else. One-third of the
endemic species belong to the genus Hynobius, a diverse group of 27 small
salamander species in the family Hynobiidae, with its only other representa-
tives in mainland China (six species), the island of Taiwan (three), and the
Korean Peninsula (three). Most species in the genus have allopatric distribu-
tions (for example, the Hokkaido or Ezo Salamander H. retardatus is the only
species found on Hokkaido, the most northerly of the Japanese islands) and
very small ranges. For example, the Critically Endangered Oki Salamander
Hynobius okiensis is confined entirely to the island of Dogo in the Oki Islands
in Shimane Prefecture, while the Tsushima Salamander H. tsuensis (LC) is
found only on the island of Tsushima (Figure 1).
Sixteen species of the genus are recorded from Japan, all of which are
endemic. As a result of studies involving the use of molecular techniques,
three new species have been described since 1984, including Hynobius katoi
(DD) described only in 2004 from montane regions of the Chubu District
in south-central Japan (Matsui et al. 2004). More recent studies are also
revealing the presence of large genetic variation in wide-ranging species,
and the number of species is likely to increase. For example, among the
stream-breeding, montane species, the Odaigahara Salamander H. boulengeri
(VU) has been shown to comprise at least three species, each occurring on
different islands of Japan (Nishikawa et al. 2001). The Blotched Salamander
Hynobius naevius (LC) is also known to contain two lineages that are par-
tially syntopic and surely represent different species (Tominaga et al. 2003).
Similarly, the Hida Salamander H. kimurae (LC) is genetically split into two
distinct lineages that are morphologically quite different as well (Matsui et
al. 2000). Likewise, still-water (lentic) breeding, lowland species are known
to exhibit a great deal of intraspecific variation. It is certain, for example,
that H. nebulosus (LC) from western Japan contains more than two species,
Hokkaido
Oki
hu
ns
Ho
Tsushima
Shikoku
Kyushu
s
nd
la
Bonin Islands
Is
u
ky
u
Ry
Volcano Islands
ESSAY 10.2. CONSERVATION NEEDS OF AMPHIBIANS IN CHINA
China is one of the most geographically varied countries in the world. Its
landscapes include the highest mountains and plateaus, vast river basins,
deserts and wetlands, the most extensive karstic region on earth, and both
temperate and tropical forests. China spans the Indomalayan and Palaearctic
Realms, and much of the country is considered to be a transition area between
them (Zhao 1999). As a result, the country is particularly rich in biodiversity,
with many endemic species, especially in the southern and south-western
parts of the country that were never affected by glaciations.
There are 346 extant native amphibians in China1 comprising 299 species
of frog, 46 salamanders and one caecilian, plus two extinct species: Cynops
wolterstorffi (globally Extinct) (He 1998) and Echinotriton andersoni (extinct
on Taiwan, still present in the Ryukyu Islands, Japan). China’s amphibian
diversity is greater than in any other country in the Old World, and is the fifth
highest in the world. Further, China has the third highest salamander diversity
in the world after the USA and Mexico. Diversity is highest in the centre and
south of the country, and lowest in the north. Three of the regions with the
highest diversity are Hengduan, Wuyi, and Nanling Mountains, where the
faunas comprise Indomalayan and Palaearctic elements. Two other regions
of high diversity are in the southern tropical regions of Xishuangbanna and
Hainan. China is also important in terms of endemic species, with 198 species
occurring nowhere else, putting the country in sixth position globally.
According to the results of the Global Amphibian Assessment, 98 amphib-
ian species in China (28% of the total) are Extinct or globally threatened and
66 species (19%) are categorized as Data Deficient, most of which are likely
to prove to be highly threatened. Threatened species are concentrated mostly
in the central and southern parts of the country, Hainan Island being the
place with the highest concentration of threatened species. Unfortunately,
the current protected area network in China (Xie 2003) does not provide
adequate coverage for Chinese amphibians (Rodrigues et al. 2004). Most of
Threatened Amphibians of the World
although delimiting the geographic boundaries between the two is not easy
due to their complex variation patterns (Matsui et al. 2006). Finally, Tohoku
Salamander Hynobius lichenatus (LC) from eastern Japan is also split into
several distinct genetic lineages that may warrant recognition at the species
level (Aoki and Matsui unpubl.).
Hynobiid salamanders are highly aquatic, occurring in streams and pools,
and breed by means of external fertilization, with the eggs laid in two distinc-
tive egg-sacs that are attached to rocks or submerged vegetation. The male
remains in the vicinity of the egg sacs, possibly for parental care, in some
species. The larvae typically hatch and metamorphose in the same year as the
eggs are laid, but in most species (e.g., Abe’s Salamander Hynobius abei; CR), a
few larvae may overwinter and only metamorphose the following year. In fact,
in the Odaigahara Salamander, the large overwintering larvae sometimes prey
on the next year’s batch of new larvae (Nishikawa and Matsui unpubl.).
Currently, around one-third (20 species) of Japan’s amphibian species are
considered threatened, eight of which are from the genus Hynobius. However,
the impending taxonomic division of several wide-ranging species discussed
above will almost certainly result in an increase in the number of species listed
as threatened in the near future. Abe’s Salamander, currently categorized as
Critically Endangered (which together with the Oki Salamander are the only
two Critically Endangered amphibians in Japan), has recently been found
from several new localities, although these discoveries were made prior to
ongoing land development in the area. Today, H. abei is specially protected
by the Japan Environment Ministry, and the capture of individuals and the
modification of its habitats are strictly regulated.
For many years, the ranges of still-water breeding lowland species have
been much impacted by increasing land development, including road con-
struction and changes in cultivation practices. In an attempt to increase rice
production, cultivation of rice paddies was halted for a season or seasons,
and then bulldozed flat and reconstructed into large rectangles with the
original fertile substrate mud bulldozed back into the paddies. The result of
such development was large-scale devastation of the amphibian fauna, ex-
acerbated by the fact that bulldozing usually took place in autumn and winter,
when populations were hibernating (Matsui 2002). Besides lowland species,
Figure 1. Map of Japan show-
ing the main islands and island
chains, and the distribution of the
genus Hynobius in the Japanese
Archipelago.
Hynobiid salamanders are highly
aquatic, and breed by means of ex-
ternal fertilization. Females deposit
a pair of distinctive egg sacs (here
Hynobius tokyoensis), including a
full clutch of unfertilized eggs, and
males focus on these egg sacs during
breeding, rather than on the female.
© Henk Wallays
the large nature reserves are in areas with few or no amphibians (especially
in the north and west of the country), while many of the most important sites
in the central and southern parts of the country are not properly protected.
With the exception of the Chinese Giant Salamander Andrias davidianus
(CR) (and see Essay 4.7), very few nature reserves have been established for
amphibians in China and managed appropriately, and those that do exist are
usually protected only at the county level.
The eight Critically Endangered species are of very high conservation
priority, and occur in scattered locations across central and southern China.
Seven of these species (two salamanders: Echinotriton chinhaiensis and
Hynobius amjiensis; and five frogs: Oreolalax liangbeiensis, Rana chevronta,
R. minima, R. wuchuanensis and Scutiger maculatus) have very small distribu-
tions, and these can readily be protected in a small number of reserves. Of
these, E. chinhaiensis, O. liangbeiensis, R. minima and R. wuchuanensis are
not present in any nature reserve, and their sites need urgent protection if
they are to be saved from extinction. Unlike the other Critically Endangered
species, the Chinese Giant Salamander still occurs very widely, but is now
extremely rare throughout its range and has disappeared from many areas
(Zhang et al. 2002; Wang et al. 2004). Some other very threatened species
that are very high priorities for conservation attention in China, but that also
occur in other countries, include Leptolalax ventripunctatus, Rana psaltes
and Ranodon sibiricus.
Salamanders (with 59% of species threatened or Extinct) are much more
threatened than frogs (24%), and 63% of the salamander species are used in
some way by humans. At the family level widely divergent patterns are seen.
Over 60% of the threatened species occur in the two largest families, Ranidae
and Megophryidae, both of which also have high percentages of threatened
species (26% and 34%, respectively). However, several smaller families show
a higher propensity to become seriously threatened, especially Bombinatoridae
stream-breeding (lotic) montane species are now also subject to increasing
habitat degradation as a result of the construction of montane roads and
dams, as well as ongoing deforestation at higher elevations. Over-harvesting
of animals for the national pet trade is also an important threat for species
such as Tokyo Salamander H. tokyoensis (VU) and the Oita Salamander H.
dunni (EN). As a result of these threats, many local governments are trying
to implement legislation to restrict the collection of animals and to prevent
land development projects that are detrimental to the salamanders’ survival.
Habitat restoration and captive-breeding programmes have been attempted
for a number of species, but both efforts are currently hampered by an often
limited knowledge of species-specific life histories.
Masafumi Matsui
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Science 20:1467-1476.
(60% of species threatened), Cryptobranchidae (100%), Hynobiidae (74%) and
Salamandridae (40%).
In China, most conservation is implemented at the provincial level. It
is, therefore, important to indicate the relative importance of the different
provinces for amphibian conservation. Sichuan has easily the largest number
of seriously threatened species, followed by Yunnan, and other provinces in
central and southern parts of the country are also important. However, the
percentage of threatened species shows a somewhat different pattern, with
the Provinces of Qinghai, Sichuan, Ningxia and Taiwan scoring the highest,
followed by Hainan, Guizhou, Shaanxi, Shanxi and Henan. Sichuan scores
more highly in terms of both numbers and percentages of threatened species,
reflecting its high level of endemism, with many severely threatened species
with very restricted ranges.
Over 80% of the threatened species are associated with forests, and over
80% occur in or around flowing water, usually streams. Streamside species
appear to be particularly at risk, and in China these species are threatened by
alien invasive species, pollution, dams, over-harvesting, siltation, and general
habitat degradation. Forest species are especially threatened because they
tend not to be very adaptable, and timber harvests in China have increased
18-fold in the past 50 years, while the area of natural forest has declined by
30% (Li and Wilcove 2005).
Habitat loss is by far the most pervasive threat, affecting 94% of threat-
ened species, compared with utilization (34%) and pollution (23%). However,
when utilization is operational, it often drives species to threatened status
more rapidly than is the case with the other threats. A total of 78 species of
amphibian in China is negatively impacted by utilization (Table 1), of which 16
are rapidly declining (as defined by Stuart et al. 2004), out of a total of 22 rapidly
declining species in China. More significantly, utilization is a major factor in 28
out of a total of 34 category deteriorations in China (category deteriorations are
 Chapter 10. Amphibians of the Palaearctic Realm 113

defined in the footnote to Table 1). In short, although utilization impacts many P.A., Pilgrim, J.D., Pressey, R.L., Schipper, J., Sechrest, W., Stuart, S.N., Underhill, L.G.,
Family Number of Number of rapidly Number of fewer species than habitat loss, it is the major threat driving species into rapid Waller, R.W., Watts, M.E.J. and Yan, X. 2004. Effectiveness of the global protected
species adversely declining species category decline in China. Ranid species in particular are threatened by over-harvesting area network in representing species diversity. Nature 428:640-643.
impacted by threatened by deteriorations for
utilization
for food, most notably the genus Paa, which has seven rapidly declining spe- Stuart, S.N., Chanson, J.S., Cox, N.A., Young, B.E., Rodrigues, A.S.L., Fischman, D.L. and
utilization utilized species
cies (accounting for 14 category deteriorations), all as a result of utilization. Waller, R.W. 2004. Status and trends of amphibian declines and extinctions worldwide.
Bombinatoridae 2 0 0 Paa are large, economically valuable frogs, and according to Ye et al. (1993), Science 306(5702):1783-1786.
Bufonidae 4 0 0 75% of the species in China are used as human food at the sub-national or Wang, X.-M., Zhang, K.-J., Wang, Z.-H., et al. 2004. The decline of the Chinese giant sala-
Hylidae 2 0 0 national level. The Chinese Giant Salamander is probably the species that has mander Andrias davidianus and implications for its conservation. Oryx 38:197-202.
Megophryidae 8 0 0 declined most seriously as a result of over-harvesting. Xie, Y. 2003. Problems in the Protected Area Management in China. 2003 World Park
Microhylidae 0 0 0 Xie et al. (2007) have provided the basic outline of a conservation strategy Congress Report.
Ranidae 33 12 21 for China’s amphibians. The key elements are: 1) conservation of important sites Xie, F., Lau, M.W.N., Stuart, S.N., Chanson, J., Cox, N.A. and Fischman, D.L. 2007.
Rhacophoridae 3 0 0 and habitats for amphibians; 2) managing for sustainable harvests; 3) scientific Conservation needs of amphibians in China: A review. Science in China, Series C,
Cryptobranchidae 1 1 3 research (on taxonomy, threatening processes, and basic species biology); 4) Life Sciences 50:265-276.
Hynobiidae 12 1 2 planning for chytridiomycosis (not yet recorded from China); and 5) rationalizing Ye, C.-Y., Fei, L. and Hu, S.-Q. 1993. Rare and Economic Amphibians of China. Sichuan
Salamandridae 13 2 2 governmental responsibility for amphibians (at present, responsibility for frogs Publishing House of Science and Technology, Chengdu, China.
Ichthyophiidae 0 0 0 falls under the State Forestry Administration, whereas that for salamanders is Zhang, K.-J., Wang, X.-M., Wu, W., et al. 2002. Advances in conservation biology
Grand Total 78* 16 28 under the Ministry of Agriculture, thus making a coherent approach to amphibian of Chinese giant salamander. Biodiversity Science 10:291-297. [In Chinese, with
conservation difficult). It is now a high priority for China to develop and imple- English abstract]
ment a nationwide strategy to save its remarkable amphibian fauna. Zhao, E.-M. 1999. Distribution patterns of amphibians in temperate Eastern Asia. Pages
Table 1. The impact of utilization on China’s amphibians. Rapidly declining 421-443. In: W.E. Duellman (ed.), Patterns of Distribution of Amphibians – A Global
species are those that have deteriorated in Red List category since 1980. One Feng Xie and Michael Wai Neng Lau Perspective, pp. 421-443. The John Hopkins University Press, Baltimore, USA.
category deterioration is a movement of a species by one Red List category Zhao, E.-M. 1998. China Red Data Book of Endangered Animals. Amphibia and Reptilia.
since 1980. For example, a movement of one species from Least Concern to Science Press, Beijing, China.
Near Threatened is one category deterioration. A movement of two species References
from Least Concern to Near Threatened is two category deteriorations. A
movement of one species from Near Threatened to Critically Endangered (as He, X.-R. 1988. Cynops wolterstoffi, an analysis of the factors caused its extinction. 1 Since the completion of data collection for the analysis in this book, eleven new
is the case with Andrias davidianus) is three category deteriorations. Sichuan Zoological Journal 17:58-60. amphibian species have been described from China, making a total of 357 species in
Li, Y. and Wilcove, D.S. 2005. Threats to vertebrate species in China and the United the country. However, if the disputed area controlled by India is included, then there
* If the disputed area with India is included, 84 species in China are adversely States. BioScience 55:147-153. are 365 species in China, rising to 376 with the new species descriptions. Note that for
impacted by utilization. Rodrigues, A.S.L., Andelman, S.J., Bakarr, M.I., Boitani, L., Brooks, T.M., Cowling, R.M., the purposes of this essay, Taiwan, Province of China, is included, although statistics
Fishpool, L.D.C., Fonseca, G.A.B., Gaston, K.J., Hoffmann, M., Long, J.S., Marquet, for this island are presented separately elsewhere in the book.

ESSAY 10.3. THE STATUS AND DISTRIBUTION OF AMPHIBIANS IN THE MEDITERRANEAN BASIN

The Mediterranean Basin, Eurasia and Africa lies at the meeting point of
two major landmasses. During the mid-Tertiary, a collision between these
two continental plates – the northward moving African-Arabian continent
and the stationary Eurasian plate – produced an unusual geographical and
topographical diversity, with high mountain ranges (more than 4,500m in eleva-
tion), peninsulas, and one of the largest island archipelagos in the world, with
almost 5,000 islands and islets. Biogeographically, the Mediterranean Basin is
usually taken as that portion of land around the Mediterranean Sea that has a
Mediterranean climate, but for the purposes of this essay we take a relatively
loose definition of the Mediterranean Basin, combining a geographic focus
on states with a pragmatic cut-off point to the north and west in Europe and
Turkey, and in the Sahara Desert to the south. Despite this broader definition,
the region is still characterized first and foremost by its climate, where mild
wet winters alternate with long hot dry summers. In some parts of the Mediter-
ranean (such as the coastal areas of Libya and Egypt), annual rainfall can be
as low as 50mm per year, whereas in the wetter regions, such as the Adriatic
coast of the Balkan countries, annual rainfall can exceed 1,000mm.
As with other Mediterranean-type landscapes across the world, includ-
ing for instance the Cape Fynbos region of South Africa and south-western
Australia, the defining characteristic of the region, at least in terms of
biodiversity, is the richness of plant species. The flora of the Mediterranean
Basin is estimated to include around 25,000 species of vascular plants,
13,000 of which are endemic (Médail and Myers 2004). This remarkable
floristic diversity, combined with a concomitant high level of threat, has led
to the region being designated as a global biodiversity hotspot. However, as
with other Mediterranean-climate regions, diversity and endemism among
vertebrates are much lower than that for plants, particularly for birds and
mammals. The mammal fauna is largely derived from Eurasian and African
elements (even the North African mammal fauna has closer affinities with
tropical Africa than with the Mediterranean Basin), whereas the avifauna
is dominated by Eurasian elements. Diversity among vertebrates is highest
among reptiles, with approximately 355 species present in the region, of
which 170 are endemic (Cox et al. 2006).
Amphibian diversity in the Mediterranean Basin is much lower than
reptile diversity, this being largely a reflection of the extent to which arid
and semi-arid habitats predominate within the region. However, at 64%,
regional endemism among the Mediterranean amphibian species is relatively
very high. The family Discoglossidae (painted frogs and midwife toads) is
almost endemic to the Mediterranean region, and two of the three species
of Pelodytidae (parsley frogs) are fully endemic. All four members of the
Pelobatidae (Eurasian spadefoots) occur in the region, two of them being
wholly restricted to the Mediterranean. Among the newts and salamanders,
49% of the world’s 70 Salamandridae species occur in the region, with five
endemic genera (Chioglossa, Euproctus, Lyciasalamandra, Pleurodeles and
Salamandrina1). The region is also noteworthy for its seven endemic cave
salamander species of the genus Speleomantes (family Plethodontidae).
Until the recent discovery of a species in Korea (Min et al. 2005), these were
thought to be the only Old World members of a family that has around 350
species in the Americas. The single Old World member of the Proteidae, the
Olm Proteus anguinus, is found only in a small part of north-eastern Italy and
otherwise Slovenia, Croatia and a few localities in Bosnia-Herzegovina; the
other five members of the family occur in eastern North America.
The richness of amphibian species in the Mediterranean basin is highest in
Europe (Figure 1), especially in areas of higher rainfall. Diversity is much lower
in the eastern and southern parts of the region. Amphibians clearly avoid arid
areas, and are absent from most of the arid Sahara. According to the results
of the Global Amphibian Assessment, 25.5% of the Mediterranean amphibian
species are considered to be threatened, with 1% Critically Endangered, 12%
Endangered and 12% Vulnerable. The most notable concentration of threatened
species is Sardinia (Italy), with four threatened species present on this island
(Figure 2). One species is listed as Extinct, the painted frog from Israel, Dis-
coglossus nigriventer. Recent surveys conducted in Lebanon following reports
that the species may survive in the Aammiq marsh in south-east Lebanon
unfortunately failed to find any individuals of this species (Tron 2005).
The human population of the Mediterranean-rim countries currently is around
400 million people, with most people concentrated along the coast. A steady
historical and continuing migration towards coastal areas, and specifically in
the south and east of the Mediterranean, has resulted in growing pressure
on the coastal environment and, more importantly, on its biodiversity. After
several millennia of human impact, most of the region’s deciduous forests have
been converted to agricultural lands, evergreen woodlands, and maquis, many
wetlands have been lost through drainage and diversion often to irrigate crops,
and increasing aridity combined with unsustainable farming and range manage-
ment practices have led to continuing desertification and land degradation in
many areas. Interestingly, amphibian families that have a higher percentage
of threatened species are also those considered to be more primitive phyloge-
netically, and coincidentally those that have large tadpoles making them more
dependent on the temporal availability of water, as well as on the presence of
available resources in this water to grow to reach maturity.
More recently, tourism has had a serious impact on the environment, par-
ticularly in western and southern Turkey, and in Cyprus, Tunisia, and Morocco,
as well as the Mediterranean islands of the Balearics (Spain), Sardinia and
Sicily (Italy), and the Cyclades Archipelago and Crete (Greece). Not surprisingly,
then, habitat loss and degradation, mainly due to agriculture and infrastructure
development, has had the largest impact on the region’s amphibian fauna,
currently affecting 19 of the 27 threatened species. However, pollution also
has been a major threat, followed by invasive alien species (e.g., Ficetola et al.
2007) not to mention predation by domestic and commensal mammal species.
Finally, there is also the risk that the fungal amphibian disease chytridiomycosis
could become a more serious threat to amphibians in the Mediterranean basin
in the future. Chytridiomycosis was first recorded in the Mediterranean Basin in
Spain in 1997, and has since been implicated in declines of the Mediterranean
populations of the common midwife toad Alytes obstetricans (Bosch et al. 2001)
as well as the fire salamander Salamandra salamandra and common toad Bufo
bufo (Bosch and Martínez-Solano 2006). If this fungal disease starts to become
as pathogenic to Mediterranean amphibians as it has done to species elsewhere
in the world, then it could rapidly become a much more serious threat in the
region. In particular, other species of midwife toad (i.e., Alytes cisternasii, A.
dickhilleni, A. maurus and A. muletensis) may be susceptible to the disease. If
this is the case, species infected with the disease, and especially those with
small insular ranges, could quickly move into a higher category of threat.
Sherif Baha el Din, Wolfgang Böhme, Claudia Corti, Jelka
Crnobrnja Isailovic, Petros Lymberakis, Rafael Marquez, Claude
Miaud, Tahar Slimani, ùsmail Uøurtaû, and Yehudah L. Werner
References
Bosch, J., Martínez-Solano, I. and García-París, M. 2001. Evidence of a chytrid fungus
infection involved in the decline of the common midwife toad (Alytes obstetricans) in
protected areas of central Spain. Biological Conservation 97:331-337.
Bosch, J. and Martínez-Solano, I. 2006. Chytrid fungus infection related to unusual
mortalities of Salamandra salamandra and Bufo bufo in the Peñalara Natural Park
(Central Spain). Oryx 40:84-89.
Carranza, S. and Amat, F. 2006. Taxonomy, biogeography and evolution of Euproctus
(Amphibia: Salamandriadae), with the resurrection of the genus Calotriton and the
description of a new endemic species from the Iberian peninsula. Zoological Journal
of the Linnean Society 145:555-582.
Cox, N., Chanson, J. and Stuart, S. (Compilers) 2006. The Status and Distribution of Reptiles
and Amphibians of the Mediterranean Basin, pp. v+42. IUCN, Gland, Switzerland
and Cambridge, UK.
Ficetola, G.F., Coïc, C., Detaint, M., Berroneau, M., Lorvelec, O. and Miaud, C. 2007. Pat-
tern of distribution of the American bullfrog Rana catesbeiana in Europe. Biological
Invasions 9:762-772.
Médail, F. and Myers, N. 2004. Mediterranean Basin. In: R.A. Mittermeier, P. Robles-Gil, M.
Hoffmann, J.D. Pilgrim, T.M. Brooks, C.G. Mittermeier, J.L. Lamoreux and G. Fonseca
(eds.), Hotspots: Revisited: Earth’s Biologically Richest and Most Endangered Terrestrial
Ecoregions, pp. 144-147. CEMEX, Mexico City, Mexico.
Min, M.S., Yang, S.Y., Bonett, R.M., Vieites, D.R., Brandon, R.A. and Wake, D.B. 2005.
Discovery of the first Asian plethodontid salamander. Nature 435:78-90.
Myers, N. Mittermeier, R.A, Mittermeier, C.G., Fonseca, G.A.B. de and Kent, J. 2000.
Biodiversity hotspots for conservation priorities. Nature 403:853-858.
Tron, F. 2005. Second Discoglossus nigriventer rediscovery expedition in the central Bekaa
Valley, Lebanon: 17-28 April 2005 Expedition Report. A Rocha International.

Figure 1. Richness map of all amphibians in the Mediterranean basin, with Figure 2. Richness map of all threatened amphibians in the Mediterranean
dark red colours corresponding to regions of higher richness. Colour scale basin, with dark red colours corresponding to regions of higher richness. Colour 1 At the time of writing, the genus Calotriton, represented by two species, had been
based on 10 quantile classes. Maximum richness equals 18 species. scale based on five quantile classes. Maximum richness equals 3 species. removed from the synonymy of Euproctus (Carranza and Amat 2006).
 CHAPTER 11. AMPHIBIAN CONSERVATION – RESPONDING TO THE GLOBAL DECLINE OF AMPHIBIANS
Colostethus ruthveni (Critically Endangered) is
one of seven species of Critically Endangered
or Endangered amphibians that are known to
be confined entirely to the Sierra Nevada de
Santa Marta massif in northern Colombia. The
area is partly protected by the Parque Nacio-
nal Sierra Nevada de Santa Marta and the
recently proclaimed El Dorado Nature Reserve.
© Fundación ProAves / www.proaves.org
Michael Hoffmann, Don Church, James P. Collins, Neil Cox, Claude Gascon,
Joseph H. Mendelson III, Robin D. Moore, Simon N. Stuart and Kevin C. Zippel
Modern amphibians have been on this planet for well over 100 million years;
they are survivors, and their decline is puzzling. Scientific study of amphibians
holds promise for a deeper understanding of the resilience as well as the limits
of environments. This is an era of increased concern about issues in biodiversity
and its maintenance in the face of environmental changes that many believe arise
directly from human activities. Amphibians may serve usefully as bioindicators,
organisms that convey information on the state of health of environments. How
to read the message, and what to do about it, are timely challenges to scientists
and to the public. David Wake, 1991
A foray through the scientific literature of the late 1980s and early 1990s reveals a sense of
alarm, and controversy, about amphibian population declines (e.g., Barinaga 1990; Blaustein
and Wake 1990; Pechmann et al. 1991; Pechmann and Wilbur 1994). Reports of decreasing
amphibian populations, particularly in seemingly undisturbed environments, made it clear
that the cause was not just habitat loss (Pounds and Crump 1994). Care was also called
for in reporting declines, because many observations may reflect only short-term natural
fluctuations (Pechmann et al. 1991; Pechmann and Wilbur 1994). The issue was nonethe-
less of sufficient concern that IUCN’s Species Survival Commission established a Declining
Amphibian Populations Task Force (DAPTF) in 1991, to determine the reality, nature, extent
and causes of declines of amphibians throughout the world (Heyer and Murphy 2005; and
see Essay 11.1).
A little over a decade later, controversy has given way to the widely held conclusion
that the world’s amphibians are indeed declining and a resulting plea for a unified front to
save them (Mendelson et al. 2006; Gascon et al. 2007). Much of the evidence presented
both in the preceding pages and those that follow indicates that global amphibian declines
are not only real, but driven by agents familiar and understood (habitat loss, commercial
use, exotic species) as well as novel and insidious (disease, climate change, toxins, and the
synergistic effects thereof). The remaining controversy is how best to arrest an extinction
episode that is already in progress.
In September of 2005, less than a year after the initial release of the results of the
Global Amphibian Assessment, more than 80 of the world’s leading herpetologists and
conservationists convened at a three-day Amphibian Conservation Summit in Washington,
D.C., to write a consolidated action plan to halt further global losses of amphibian popula-
tions. The assumption was that it is irresponsible to document amphibian declines and
extinctions without also designing and promoting an appropriate response. The resulting
Amphibian Conservation Action Plan (ACAP; see Gascon et al. 2007) outlines the need for
four types of intervention to conserve amphibians: 1) Expanded understanding of the causes
of declines and extinctions; 2) Ongoing documentation of amphibian diversity, and how it is
changing; 3) Development and implementation of long-term conservation programmes; and
4) Emergency responses to immediate crises (see Appendix VI). An initial five-year budget
to implement this Action Plan requires at least US$400 million.
Given the existence of a consensus conservation framework among those present at
the Amphibian Conservation Summit, this chapter is not a new vision for conservation ac-
tion. Nor is it a detailed review of amphibian conservation efforts, a topic covered in detail
elsewhere (e.g., Semlitsch 2003, Heatwole and Wilkinson in press). Rather, this chapter is
intended to summarize the key action items of the ACAP, measured against our knowledge
of threats to amphibians – and biodiversity more broadly – accompanied by conservation
recommendations for immediate uptake by governments, corporations, civil society, biodi-
versity conservation organizations, and the scientific community.
THE CORE RESPONSE: DESIGNING
AND IMPLEMENTING AN IMPROVED PROTECTED
AREAS NETWORK FOR AMPHIBIANS
In the preceding pages, we have already established that for nine out of every 10 amphibian
species classified as threatened, habitat loss and degradation is a major threat. In such
cases, area- or site-based action, which usually takes the form of the establishing protected
areas, is the most effective conservation response for safeguarding biodiversity in the long
term (Bruner et al. 2001). The Programme of Work on Protected Areas of the Convention
on Biological Diversity (CBD) states that protected areas are “…essential components in
national and global biodiversity conservation strategies.” However, habitat protection is not
necessarily sufficient to ensure long-term viability in the face of threats such as emerging
infectious disease (Lips et al. 2006) or climate change (Pounds et al. 2006), and often needs
to be complemented by other actions, such as environmental education and targeted ex-situ
interventions (see later), as well as by actions at the landscape scale that aim to respond
to threats operating at a broader scale (Boyd et al. 2008).
Unfortunately, although protected areas cover 11.5% of the world’s land surface, the
existing network of protected areas is incomplete and does not adequately capture global
biodiversity. Indeed, assessments conducted at differing scales (e.g., Rodrigues et al. 2004a;
Rondinini et al. 2005), consistently reveal large gaps in the network, this being particularly
evident in the tropics (Brooks et al. 2004), leaving many threatened species unprotected.
According to the Global Amphibian Assessment, around 40% of all threatened amphibians
are not represented in any protected area. An evaluation of the protection afforded to threat-
ened amphibians through the current protected areas network suggests that amphibians
are markedly less well covered than other taxa (mammals, threatened birds, and turtles and
tortoises; Rodrigues et al. 2004a,b; Essay 11.2). Many of the amphibians that do not occur
within a protected area are also found only at a single site thereby increasing the risk of
extinction above that of generally more widespread bird and mammal species (Ricketts et
al. 2005). Furthermore, protected areas vary in legal status and management type, resulting
in variable levels of effectiveness, and enforcing area protection and related laws is often
logistically difficult in many countries.
The last two decades saw an explosion of literature on the subject of systematic conserva-
tion planning, a field of conservation biology that aims to identify comprehensive networks
of sites or protected areas that together will contribute to the overall goal of minimizing
biodiversity loss (Pressey et al. 1993; Margules and Pressey 2000). This is necessary because,
as is evident from Chapter 4, neither biodiversity nor threats to it are distributed evenly, with
the result that investments must be made in some places with greater urgency than others.
The significant advances made in the field of systematic conservation planning (Kirkpatrick
1983; Margules and Pressey 2000) have seen the science move beyond theory to on-the-
ground application (e.g., Cowling et al. 2003; BirdLife International 2004).
Strategic decision-making of this order requires information on both the spatial and
temporal options available for inclusion in the conservation planning framework. These
two variables are commonly referred to as irreplaceability and vulnerability, respectively, in
the conservation planning literature (Pressey and Taffs 2001). Irreplaceability estimates the
degree to which spatial options available for conservation of unique biodiversity features
are lost if a particular site is lost. At its most extreme, for example, a site with the entire
population of a species is wholly irreplaceable – there are no other sites available (i.e., spatial
options) for conserving the species (Pressey et al. 1994). Vulnerability is a measure of our
temporal options available. Just as threatened species are more likely to be lost before non-
threatened species, our options for conserving those sites facing high levels of vulnerability
or threat are more limited in time, with places of higher threat likely to lose their biodiversity
value sooner (Rodrigues et al. 2004b). Vulnerability and irreplaceability interact in complex
ways to define conservation priorities. For example, sites of simultaneously high threat and
high irreplaceability are the highest-order priorities as they correspond to places where the
loss of unique biodiversity is imminent. By contrast, conservation in low irreplaceability
regions can afford to be opportunity-driven, as there are plenty of spatial options.
Based on these two concepts, Rodrigues et al. (2004b) expanded their earlier analysis
and provided a first-ever attempt at a global framework for the next step of strategically
expanding the global protected area network to ensure appropriate coverage of mammals,
amphibians, freshwater turtles and tortoises, and globally threatened birds. They identified
several urgent priorities for expanding the network (Figure 1), which fall overwhelmingly in
the tropics, especially in tropical and subtropical moist forests. Not surprisingly, the results
of this study, in terms of geography, show remarkable congruence with areas already known
to represent consensus priorities for reactive conservation action (Brooks et al. 2006).
However, just as any broad-based global priority setting exercise (see Brooks et al. 2006
for review of these) is useful for guiding the allocation of resources on a broad scale, the
regions identified as urgent priorities for the expansion of the global network by Rodrigues
et al. (2004b) are not useful for determining the location of the boundaries of new protected
areas. Consequently, given the coarse scale of their analysis, the key recommendation from
their study is that the highlighted areas become urgent priorities for finer-scale assessments
to identify manageable units of global conservation significance.
The Amphibian Conservation Action Plan adopted the Key Biodiversity Areas approach
(KBAs; Eken et al. 2004; Langhammer et al. 2007) as a tool for identifying globally important
sites for amphibian conservation. Key Biodiversity Areas, which hinge on the concepts of
threat and irreplaceability, build on nearly a quarter century of work by BirdLife International
in relying on a set of criteria for identifying globally significant sites for bird conservation
(Important Bird Areas; see, for example, Fishpool and Evans 2001), but extend this concept
to create a taxon-neutral umbrella to incorporate all taxa (see Essay 11.3). The focus of KBAs
is on species most vulnerable to extinction: globally threatened species and geographically
 Chapter 11. Amphibian Conservation – Responding to the Global Decline of Amphibians
concentrated species. KBA identification for amphibians is already advanced in several
regions of the world, thanks to data collated through the Global Amphibian Assessment
(Langhammer et al. 2007), and will provide a ready basis for directing valuable conservation
resources to sites of known global biodiversity importance.
However, while all KBAs represent global priorities for site-scale conservation, there will
be a subset of sites where conservation action is most urgent. The Amphibian Conservation
Action Plan itself calls for the identification and safeguarding of the 120 KBAs where unique
amphibian biodiversity is most at risk. Fortunately, there has already been considerable effort
aimed at locating such places through the work of the Alliance for Zero Extinction1, a joint
initiative of more than 65 biodiversity conservation organizations around the world that aims
to prevent imminent extinctions by pinpointing and safeguarding sites that represent the
last refuges of highly threatened species. Such sites, termed Alliance for Zero Extinction (or
AZE) sites, are discrete areas containing ~95% or more of the global population of one or
more species listed as Critically Endangered or Endangered on the IUCN Red List (Ricketts
et al. 2005)2. The Alliance has identified 546 sites holding 758 Critically Endangered (CR)
or Endangered (EN) species of amphibians, reptiles (crocodiles, iguanas, and turtles and
tortoises), mammals, birds, and conifers. At least half of these sites (263) have been identified
for 381 amphibians (Figure 2). Of the sites identified so far, 125 have no formal protection
and 27 only partial protection3, yet again emphasizing that a large proportion of the world’s
most threatened biodiversity is not adequately represented in protected areas.
All AZE sites/species are important and top priorities in the sense that they represent
known places where conservation action is needed now. However, establishing protected
areas may not be the most appropriate response in all cases, but will depend on the nature of
the threat. Indeed, in the context of site-scale conservation, one set of immediate priorities
will be sites for which the overwhelming threat driving species declines is loss or degrada-
tion of natural habitat and for which the most obvious conservation action involves habitat
protection, maintenance, or restoration. This will be particularly urgent for sites with no
formal protection, or at least only partial protection. For example, the Massif de la Hotte
in Haiti is the only known site for 13 Critically Endangered or Endangered amphibians, and
the overwhelming threat at this site is habitat loss and degradation. Even though part of the
site is protected as the Parc Natural Macaya, there is little active protection of the area and
habitat loss continues even within park boundaries. Such sites (i.e., those where site-based
threat and habitat loss are high) are priorities for safeguarding, and indeed several, such as
the Parque Nacional Sierra Nevada de Santa Marta in the Colombian Andes, have benefited
from collaborative efforts to safeguard remaining habitat4. At the time of going to press,
efforts were underway to transform a priority cloud forest site in Sri Lanka, Morningside5,
into a Biodiversity Reserve managed by the Forest Department of Sri Lanka in collaboration
with local civil society organizations.
Establishing protected areas is not normally a simple process. There are usually many
competing demands for using particular parcels of land, whatever their size. For a protected
area to be long-lasting, the interests of multiple stakeholders must be considered. A num-
ber of possibilities exist for building support from local stakeholders for protected areas,
including: a) enabling local communities to benefit from nature reserves (such as allowing
sustainable harvest of relevant products, and demonstrating the benefits of securing water
catchments); b) providing local employment (for example in ecotourism, and in reserve protec-
tion, management and monitoring; and c) developing alternative livelihoods in the vicinity of
reserves that enable people to enhance their standards of living in ways that are compatible
with securing a sustainable future for the reserve in question. Most of the new protected
areas that need to be established to conserve amphibians are in mountainous areas of the
humid tropics, where both human population densities and poverty levels tend to be high. The
success of these plans to establish new protected areas depends on forging alliances with
development assistance agencies and community-based organizations to achieve the twin
goals of biodiversity conservation and sustainable economic development. Further, establish-
ment of protected areas is not the only way to secure areas for biodiversity, and other means
that complement protected areas, including land purchase, land ownership/stewardship, and
land concessions and negotiations with private landowners, may be options.
Establishing protected areas that remain resilient to the increasing threat of climate vari-
ability is another major challenge. Climate change, driven by human activity forces species
to track preferred habitat conditions, but this movement may be difficult or impossible in
fragmented landscapes. Further, the rate and magnitude of current climate change is such
that many species may be unable to disperse quickly enough. For instance, species that
inhabit high-elevation montane regions often have nowhere to disperse to as temperatures
rise and other climatic conditions change. Thus, protecting species where they currently
exist is only the beginning – protection is needed where species will be in the future, as
well as connections in the landscape between the two. Biologists are now able to create
115
Figure 1. Global distribution of priority sites
for the expansion of the global protected-area
network, based on data for all mammals,
amphibians, turtles, and threatened birds.
Adapted from Rodrigues et al. (2004b).
computer models of species’ range shifts that can be used to plan corridors, comprising
additional parks, or multiple-use areas such as forest reserves, to limit the damage of
climate change. Landscape conservation of this sort requires a large percentage of remain-
ing natural habitats to be successful. Proactive investment in these areas will help prevent
accelerating extinctions in the near future due to the ‘double whammy’ of climate change
in a fragmented landscape. For example, numerous threatened frog species in Sri Lanka are
hanging on within small fragments of the country’s 5% of remaining rainforest. These species
may already be affected by decreasing precipitation and warmer temperatures – conditions
that increasingly wring small forest fragments dry. In these cases, restored forests around
existing patches will provide water retentive buffers locally, as well as corridors to forest
remnants that respond differently to a changing climate.
It is necessary to emphasize that while there is little doubt that habitat protection
must remain the cornerstone of any conservation action, the ecological requirements of
some species means that conservation action is required at scales larger than that of
protected areas. Among mammals and birds, for example, this is particularly the case for
area-demanding species that have very low population densities or are migratory. Further,
the nature of many threats to biodiversity is such that habitat protection alone, or other
site-based interventions, is not necessarily sufficient to preserve the integrity of the broad-
scale ecological and evolutionary processes necessary to ensure the persistence of species
within individual sites (Lens et al. 2002).
A recent study provides a unique perspective on the appropriate scale for conserva-
tion action for threatened amphibians, using data collated to support IUCN Red List
assessments. Boyd et al. (2008) found that while most species can be conserved through
individual sites, or networks of sites, more than one-fifth (21%, or 353 species) of globally
threatened amphibians cannot be conserved at the site scale alone in the medium-term
(cf. 13% for mammals and 19% for birds). For amphibians, this is entirely due to alterations
in hydrological processes, such as changes in flow regimes (including various aspects of
seasonality and flow rate), which is deemed necessary for 115 species, and water quality
(e.g., temperature, pH, and sediment loads), which is considered an issue for 278 species
(Boyd et al. 2008) (Appendix VIIa and VIIb)6.
Several initiatives and organizations are incorporating broad-scale ecological processes
into conservation planning (e.g., Pressey et al. 2003). However, the task remains challenging
due to problems with setting priorities for ecological processes, and developing targets for
their conservation (Boyd et al. 2008). Encouragingly, the recovery plans for several threatened
species, including the California Red-legged frog Rana aurora7, Corroboree Frog Pseudophryne
corroboree8 and stream-dwelling rainforest frogs of the Wet Tropics region in North-east
Queensland9 incorporate components relating specifically to the development and implementa-
tion of guidelines for maintaining adequate water flow regimes. For example, the recovery plan
for the California Red-legged Frog, the only species in the United States for which a recovery
plan has been developed, includes allocating water budgets to county planners and water
districts, particularly those in the San Simeon Creek and Santa Rosa Creek in San Luis Obispo
County, and managing dams and reservoirs that affect populations of the species. Another
example is the recovery efforts for the Kihansi Spray Toad Nectophrynoides asperginis, which
included the construction of a gravity-fed artificial spray system to create a fine spray in an
attempt to maintain the spray-zone microhabitat in the Kihansi Gorge, following the construction
of a dam upstream on the Kihansi River (Lee et al. 2006; Krajick et al. 2006).
EX-SITU MANAGEMENT AND REINTRODUCTION
PROGRAMMES
While immediate habitat protection must be considered an emergency measure for several
amphibians (see above), even with timely and effective protection from direct anthropogenic
threats, such as habitat destruction and over-collection, there are some species for which
we cannot readily safeguard wild populations from uncontrollable indirect threats, such as
climate change, and direct threats, such as emerging infectious diseases. Reference is often
made to the Golden Toad Bufo periglenes as the model case of an amphibian that disappeared
in pristine and well-protected habitat (Pounds and Crump 1994), but in the current era it
is one of many species that have declined under similar circumstances. In cases involving
threats of such a nature, conservation action in situ must be complemented by immediate
interventions that may be invasive, but unavoidable.
Indeed, with the realization that in the absence of any clear understanding on how to
combat the spread of the chytrid pathogen, the current lack of any management technique
to prevent wild populations succumbing to the disease, and no real understanding on the
116
Figure 2. Map of Alliance for Zero Extinction
sites (n=263) triggered by the presence of a
Critically Endangered or Endangered amphib-
ian (381 species). Yellow sites are either
fully or partially contained within declared
protected areas (n=122), and red sites are
completely unprotected or have unknown
protection status (n=141). In areas of overlap,
unprotected (red) sites are mapped above
protected (green) sites to highlight the more
urgent conservation priorities. Modified from
Ricketts et al. (2005); AZE data version 2.1.
The Massif de la Hotte in Haiti is home to
the last remaining populations of 13 Criti-
cally Endangered and Endangered amphibians,
more than any other single AZE site. © Robin
D. Moore
Threatened Amphibians of the World
disease’s origins or mechanisms of spread, one strategy is establishing emergency ex-situ
survival-assurance colonies where infected animals can be treated10, and as yet unaffected
but susceptible species may be safeguarded. Indeed, this may well be one of the most
important roles for the ex-situ conservation community at this time, as through extensive
field research and predictive modelling, we can, in some cases, anticipate when and where
certain species will be impacted (see Essay 11.4; and Essay 9.7), thereby creating a narrow
window of opportunity for proactive intervention.
Ex-situ survival-assurance colonies may provide the only option for the eventual return
of some species or populations to safeguarded natural habitat. Now, more than ever, there
is a need for the ex-situ conservation community to play an active role in averting spe-
cies extinctions, not just in terms of providing the much-needed facilities and husbandry
expertise, but in supporting disease-related research, education and conservation efforts
on the ground. Ex-situ programmes cannot replace in-situ conservation activities, but they
have the potential to provide the means for species survival while complementary research
and conservation programmes proceed. The role of ex-situ management as a conservation
measure is controversial for several reasons: 1) gaps in our knowledge of the biology and
breeding behaviour of most species; 2) the removal of individuals with potentially higher
fitness from populations perhaps already severely decimated; 3) uncertainty regarding the
number of individuals required to maintain a viable population; 4) absence of resources for
long-term maintenance of captive populations; 5) the low success rates of several previous
ex-situ conservation efforts (see later); 6) the number of species for which such survival-
assurance colonies may be needed, and 7) the degree to which such efforts may avert or
misdirect funding from active conservation on the ground.
Various authors have discussed the candidacy of amphibians for ex-situ management
programmes (Wiese and Hutchins 1994; Bloxam and Tonge 1995) and the role of ex-situ
management in amphibian conservation has been detailed in several works (e.g., Zippel
2005; Griffiths and Kuzmin 2006; Mendelson and Rabb 2006). The valuable role of ex-situ
management, particularly as part of a coordinated recovery programme (see Essay 11.5)
has been recognized as a useful conservation tool by IUCN, which states that it should be
initiated “…to help support the conservation of a threatened taxon, its genetic diversity,
and its habitat”, while noting that “Ex situ programmes should give added value to other
complementary programmes for conservation.”11 Consequently, just as others have made
previous calls for ex-situ conservation action as part of a broader set of conservation priorities
in declining amphibian communities (e.g., Young et al. 2001), the Amphibian Conservation
Action Plan has incorporated ex-situ management as an explicit component of the overall
strategy, with the proviso that dedicated infrastructure of staff members and facilities will
require long-term commitments of reliable financial support.
However, in the face of estimates that the global ex-situ conservation community can
currently manage viable populations of only approximately 50 amphibian species (see
Essay 11.5), we need a reference point for assessing the magnitude of task. In Appendix
VIII, we list 247 species for which ex-situ survival-assurance colonies are proposed as an
urgent conservation measure to ensure the persistence of species in the face of the threat
of chytridiomycosis (either because the chytrid fungus has already been documented in the
species, or because the species has undergone declines and chytridiomycosis is deemed
likely to be the cause12). Mapping the distributions of these species (Figure 3), we see that
most of the species for which this is deemed necessary are concentrated in the Neotropics,
mainly due to the susceptibility of montane, stream-dwelling species in the genera Atelopus,
Craugastor, Telmatobius, and Colostethus to chytridiomycosis, although there are also
concentrations in the Eastern Arc Mountains and in Australia.
An important caveat is that these figures exclude species flagged as Critically Endangered
(Possibly Extinct), since such species represent the most urgent priorities for immediate
survey work to determine whether they persist in the wild (see later). In such cases, any
surviving individuals may need to form the basis for the establishment of an ex-situ sur-
vival-assurance colony, although this depends on the nature of any threat to the species.
In addition, there are other cases where ex-situ survival-assurance colonies are needed
because of the impact of invasive species (for example, Cape Platanna Xenopus gilli, EN),
or to mitigate against severe overexploitation (e.g., several Ambystoma species), or because
habitat loss has been so severe that ex-situ management is deemed necessary pending the
initiation of habitat restoration efforts (for example, Puerto Rican Toad Bufo lemur). Such a
list of priority species is not complete, but serves as a useful starting point for the global
conservation community to initiate targeted management programmes.
While we draw attention to the need for any ex-situ management programme to adhere
to the best available husbandry techniques, we also stress several important considerations.
First, while funding opportunities are more likely to be available for such programmes in
developed countries, there is often reluctance on behalf of developing countries to surrender
their biodiversity to foreign institutions. This reaction is based on historical and commercial/
trade precedents14. One solution involves transferring technology and capacity to developing
nations by means of establishing ex-situ management facilities with sustainable funding
sources, and backed by appropriate guidance in husbandry techniques for the species in
question. One example of such a programme in development (it is neither complete in
construction nor does it have sustainable funding), is the El Valle Amphibian Conservation
Center (EVACC) at the El Nispero Zoo in Panama, an effort spear-headed by the Houston
Zoo in collaboration with several zoo and non-governmental organizations. This project
aims to serve as a biological repository for threatened Panamanian amphibian species and
to use these populations as a possible source for reintroductions, but has a second objec-
tive which is to foster appreciation and raise awareness of amphibians among the public,
through programs of education and research. Indeed, half of the allotted 220-m² space at
EVACC is dedicated to public exhibits and the rest to behind-the-scenes space for managing
survival-assurance populations and research projects. It had been hoped that EVACC would
be operational before the arrival of chytrid in the region, but in April 2006 the first amphibian
deaths due to chytridiomycosis were reported from frogs inhabiting waters of the Rio de Jesús
(collected a few months before). Consequently, the emergency collection of amphibians
was initiated in advance of the facility being completely constructed, and a temporary
quarantine and treatment area was established in a nearby hotel. As of end July 2006,
collecting teams of 50 volunteers representing 26 institutions had collected 10 of the 12
priority species15. The goal is for current project leaders to leave the programmes and turn
over responsibility for conservation initiatives to Panamanian colleagues.
Another possibility involves negotiating ownership rights such that source countries may
retain ownership over frogs and their genetic material sent to foreign zoos to form the basis
for ex-situ survival-assurance colonies. This has already been achieved for the Kihansi Spray
Toad Nectophrynoides asperginis, which is being bred in the Toledo and Bronx Zoos. These
toads are being managed under an agreement with the Government of Tanzania, which
retains ownership of the animals (see Lee et al. 2006; Krajick 2006).
Second, any ex-situ programme must be accompanied by appropriate actions directed
at the long-term conservation of the species in situ. This could include, but may not be
limited to, habitat protection, public education and awareness, research (e.g., breeding
biology), population monitoring, targeted species-specific actions (such as control of
invasives), and so forth.
Since the over-riding conservation role for ex-situ management programmes is the
eventual reintroduction/translocation of species back into the wild, it is worth noting that
 Chapter 11. Amphibian Conservation – Responding to the Global Decline of Amphibians
achieving the first has usually proved much easier than the latter, with much consequent
debate surrounding the value of these techniques (Dodd and Seigel 1991; Seigel and Dodd
2002; Trenham and Marsh 2002; Dodd 2005). Indeed, even considering conservation suc-
cess stories, such as the Mallorcan Midwife Toad Alytes muletensis (see Essay 11.6), a
number of long-running projects have yet to demonstrate similar success, even in developed
countries such as the United States. For example, Wyoming Toads Bufo baxteri, now Extinct
in the Wild, have been part of an active breeding programme since 1993 and yet despite
repeated introductions of toadlets and tadpoles to Mortenson Lake, the population is still
not self-sustaining (Dreitz 2006; and see Essay 11.7). The causes for the failure of successful
reintroduction are not always clear, although disease and other factors are often implicated.
In any case, there are several examples of species that would be extinct at this time if they
had not been included in captive programmes (such as the Wyoming Toad). In such cases,
extinction has been deferred, giving researchers time to solve the in-situ problems.
Fortunately, other long-running conservation breeding programmes are finally showing
some signs of success, such as that for the Puerto Rican Toad Bufo lemur. This species was
believed extinct until its rediscovery in 1967, when a small breeding population of toads was
found in north-west Puerto Rico (near Quebradillas) and subsequently in the Guanica Forest
Preserve in the south-west of the island. No toads have been seen at the north-western site
since 1992, and the Guanica Forest Preserve has become the last locality known to hold a
wild population of this species. An ex-situ programme for the Puerto Rican Toad began in
1980 at the Mayaguez Zoo, with offspring from this programme later sent to zoos in North
America, including Buffalo Zoo in New York and the Toronto Zoo in Canada. A Species
Survival Plan (SSP)16 was formed in 1983, with the goal of assisting the recovery of this
species. To date, the Puerto Rican Crested Toad SSP programme includes some 23 zoos and
aquaria; its efforts include education and habitat restoration, and the SSP members are
supporting high school students who look for toads and study ponds in the northern part of
Puerto Rico and have a special biology club with the toad as their logo. The SSP members
(the zoos and aquaria in the SSP, along with Puerto Rican partners, including teachers,
students, the Guanica Forest Manager, the Puerto Rican DNR and USFWS) construct new
ponds for toads; install gates and signs to protect the breeding beach; print conservation
booklets, bumper stickers and decals for students and interpretive centres; work with the
local Juan Rivero Zoo in Mayaguez to help in their conservation programmes for Puerto
Rican wildlife, including the toad; and support research on toad habitats and diseases by
Puerto Rican graduate students and veterinarians. In zoos, the toads are managed geneti-
cally to ensure a diverse assurance population in the event of the unexpected loss of the
small Guanica population. Tadpoles produced in the ex-situ programme have been sent
back to Puerto Rico for release into the wild, and two artificial release ponds have been
constructed to receive the tadpoles. Releases began in 1983 with 75 toadlets, and about
100,000 tadpoles have been released over the last 10 years. Eventually, in 2003, the first
adult toads released as tadpoles returned to breed in the constructed ponds, evidence of
the first signs of success (Zippel 2005).
While it is not possible to identify here a specific unbiased set of species for which
reintroduction may be an appropriate conservation tool, it is essential that species are
carefully appraised for their suitability for reintroduction, as they should also be appraised
for their suitability for captive breeding, and in accordance with the IUCN/SSC Guidelines
for Reintroductions17. Priorities for reintroduction clearly should focus on globally threatened
species, and since the most successful animal reintroductions have usually involved those
species that have reversible threats (Griffith et al. 1989; Wilson and Stanley Price 1994), it
may well be necessary to give preference to those species for which actions are already
underway to mitigate threats such as direct persecution, pollution, and introductions of
fish or other predators. Reintroducing Axolotls Ambystoma mexicanum to Lake Xochimilco,
for example, is complicated by the fact that existing threats need to be neutralized and
potential disease and genetic risks addressed before animals from ex-situ populations are
reintroduced into the wild (and see Essay 11.8).
This also seems the most appropriate place to mention the occasional need for transloca-
tion of threatened species to new sites as a result of impending habitat loss. An example is
Chirixalus romeri, an endemic to Hong Kong that lost part of it natural habitat when the new
Hong Kong International Airport was constructed. With funding from the Hong Kong Jockey
Club, the University of Hong Kong undertook a study of the ecology and population genetics
of this species to identify suitable sites for relocation of this threatened population. More
than 200 individuals were rescued from Chek Lap Kok in 1992 before full-scale construction
took place. A collaborative partnership was also formed with Melbourne Zoo to breed this
species ex-situ, which was successful and over 2,800 frogs and tadpoles were released into
eight sites in Hong Kong Island and the New Territories where they have been monitored
every year. Breeding populations have established in seven of them.
117
Figure 3. The global distribution of Criti-
cally Endangered and Endangered amphibians
(n=247) requiring ex-situ intervention (in
combination with complementary in-situ and
research-based actions).13 No fewer than 59
of these species have their entire distributions
confined to single sites (i.e., AZE species;
see Figure 2).
Finally, ex-situ rearing of animals can also form part of an important conservation strategy
aimed at promoting reasonable and sustainable use of wildlife, although only if the supply
from farmed animals is such that it replaces that from the wild (see Essay 11.9). In Mexico,
for example, despite the wide availability of Axolotls produced in ex-situ populations, wild
animals are still captured and sold illegally in local markets (McKay 2003; and see Essay
2.3), while in China, even the existence of more than 100 apparently legal Chinese Giant
Salamander farms to supply the ever-growing food market is insufficient to meet market
demands (Essay 4.7). Furthermore, an important complication of some of these commercial
enterprises is that they often farm non-native species, such as the American Bullfrog Rana
catesbeiana, which escape and become invasive (see Essay 4.8). This could have serious
implications for conservation, because the rearing of bullfrogs is a growing industry in parts
of the world such as China and South America, particularly Brazil, Uruguay, and Argentina,
and this species may carry chytridiomycosis (see Mazzoni et al. 2003).
Nonetheless, sustainable community-operated farms and commercial breeding farms can
have benefits, but it is recommended that they use local species instead of exotic ones to
reduce the risk of disease spread and establishment of exotic species. It is also necessary
to monitor commercial breeding farms for highly valuable species to prevent wild-caught
individuals from entering into the trade. Finally, a percentage of the profits generated from
such breeding farms should be channelled back to amphibian conservation whenever pos-
sible (Carpenter et al. 2007).
In conclusion, while there is disagreement about the interplay between in-situ and ex-situ
conservation action, it is preferable not to have one undermining the other. In practice, if
we are to prevent amphibian extinctions, there can be no doubt of the importance of stem-
ming rampant environmental degradation (Pounds et al. 2006). Other amphibian-specific
conservation actions, such as those ex-situ, should continue, but their effectiveness will
likely be reduced or compromised in the face of growing environmental deterioration. Yet,
the two are compatible, complementary actions, insofar as their relative contributions
and limitations are understood, accepted, and explicitly stated: ex-situ action is vital for a
finite set of highly threatened species (but clearly not urgent for all), yet requires parallel
in-situ efforts. Our ability to respond depends largely on the priorities of the global ex-situ
conservation community.
TARGETED IN-SITU ACTIONS
Habitat protection and the establishment of ex-situ survival-assurance colonies represent
the two most urgent in-situ and ex-situ conservation actions required, respectively, if we
are to have any immediate impact on amphibian declines globally. But, as already discussed
above, even though they complement one another, they will not suffice on their own. Indeed,
even Ricketts et al. (2005) note that in the case of sites identified using their methodology to
pinpoint species having their last remaining populations confined to single sites, measures
The El Valle Amphibian Conservation Center
(EVACC) at the El Nispero Zoo in Panama is
an effort spear-headed by the Houston Zoo in
collaboration with several zoo and non-gov-
ernmental organizations. Half of the allotted
220-m² space at EVACC is dedicated to public
exhibits and the rest to behind-the-scenes
space for managing survival-assurance popu-
lations and research projects. © Sara Riger
118
A mascot, highlighting the unique characteris-
tics of the endemic Puerto Rican Crested Toad
Bufo lemur (Critically Endangered), contributes
to island-wide conservation programmes at
schools, fiestas, and zoo education and out-
reach events. © Toronto Zoo
Threatened Amphibians of the World
of habitat protection will need to be complemented by other actions, such as the control of
invasive alien predators/competitors or disease.
Invasive species
The Amphibian Conservation Action Plan does not deal specifically with the issue of the
impact of invasive species, which could have detrimental effects on populations as com-
petitors, predators, and interbreeding to create hybrids. Studies show that alien predators,
particularly introduced fish species (e.g., Trout Oncorhynchus spp., Salmo spp., centrarchids
like bass and sunfish, and Mosquitofish Gambusia spp.), American Bullfrogs, and crayfish
(Procambrus clarkia) have contributed to amphibian population declines (see Kats and Ferrer
2003). Some of these studies documented declines among amphibian communities within
protected areas (e.g., Knapp and Matthews 2000). Although introduced predators may be
ubiquitous in watersheds within the range of the threatened species, areas of high concen-
trations should be identified and the numbers of non-native predators reduced, particularly
within protected areas. For example, the recovery plan for the California Red-legged Frog
Rana aurora draytonii identifies a number of sites as areas that may benefit from non-native
predator removal, including Yosemite National Park.18
While the most important conservation measure is prevention by means of more effective
quarantine measures and enforced legislation (see Policy below), more aggressive action
invariably is required for combating alien predators (e.g., Knapp and Matthews 1998)19.
Control and removal of invasive predators is not always feasible or possible in those cases,
where, for example, these species have commercial or recreational value (e.g., alien game
fish), or where the species exhibit particular traits that characterize successful invaders (e.g.
high reproductive rate), thereby making removal difficult. Control methods generally involve
pond drainage, physical removal, poisoning (e.g., rotenone20), and eliminating breeding
habitat of non-native predators. On the island of Mallorca in the Mediterranean Sea, for
example, considerable effort was expended to control introduced snakes (Natrix maura),
but these labour-intensive efforts mostly involved removing snakes by hand from breeding
sites (Guicking et al. 2006). Although relatively large numbers of snakes were removed from
breeding sites, there have been no studies to assess the effectiveness of this technique in
controlling snake numbers. Most importantly, toads continue to survive and breed in these
pools. One recent study found that large-scale removal of introduced non-native fish could
result in at least partial reversal of the decline of Mountain Yellow-legged Frog Rana mus-
cosa (CR), once a common inhabitant of the Sierra Nevada (California, USA). At three lakes
where fish were removed, amphibian population densities increased significantly following
removal of predaceous fish. Following these population increases, frogs dispersed to adjacent
suitable (but unoccupied) sites, moving between 200 and 900m along streams or across dry
land (Knapp et al. 2007). On the other hand, the afore-mentioned California Red-legged Frog
recovery plan notes that filling in stock ponds known to have non-native species may be more
cost effective and productive over the long-term than removing individuals. In Venezuela,
the transportation of Rana catesbeiana to other regions outside of Mérida has been legally
prohibited, and recommendations for its control and removal were passed (Resolution #64
of 15 July 2002). Furthermore, the national government, via its National Office for Biological
Diversity, Ministry of the Environment, and the State Environmental Authority of Mérida,
actively engage in collection and removal of individuals of all stages of the species (e.g.,
collection of eggs, use of dragnets, nets, air rifles and traps).
New studies involving age-structured population models provide interesting insights into
the control and management of invasive species for the benefit of threatened amphibians.
Doubledee et al. (2003), in discussing the impacts of American Bullfrogs on the California
Red-legged Frog, assessed the effectiveness of different management strategies used to
control bullfrog populations (such as shooting of adults and pond draining). Their results
found that efforts taken to eradicate bullfrog populations by increasing adult mortality
through shooting are likely to be ineffective in promoting coexistence, first because a
large amount of effort – equivalent to a mortality of 65% or greater every two years – is
required to make shooting adults beneficial for Red-legged Frog persistence, and second
because such effort can create fluctuations in bullfrog population dynamics, which in turn
may cause large fluctuations in the Red-legged Frog population. Their results suggest that
a combination of shooting adults and draining livestock grazing ponds is the most success-
ful bullfrog management strategy. Even low shooting efforts significantly decreased adult
bullfrog densities by 80% when coupled with pond draining.
The recent emergence of Batrachochytrium dendrobatidis, the chytrid fungus that causes
chytridiomycosis, may itself be the result of unintentional introductions of a pathogen as an
invasive alien species. One suggestion is that the spread of the pathogen started with the
export of the African Clawed Frog Xenopus laevis from Africa for use in pregnancy testing
during the 1930s onwards. But more research is needed to test this hypothesis. Regardless
of the origin of the chytrid fungus, however, bullfrogs carry it and so establishment of feral
populations of this species in countries is one means by which the pathogen can gain access
to other hosts (Weldon et al. 2004; and see Essay 4.8).
Diseases
To date, there have been few direct intervention efforts to prevent the spread of disease
in wild populations of animals. Relatively successful examples include African Wild Dogs
Lycaon pictus in Tanzania’s Serengeti National Park for rabies (Woodroffe 2001) and
Ethiopian Wolves Canis simensis in the Bale Mountains National Park of Ethiopia (Haydon
et al. 2006). There is still no vaccine for chytridiomycosis, and development would, in any
event, probably take many years. Furthermore, direct treatment of amphibians in the wild
with anti-fungal agents is not without problems: release of anti-fungal agents in a way
that would target the frog population significantly enough to deal with infections would
have unknown effects on the fungal component of the ecosystem (Daszak et al. 2008).
Therefore, population management strategies involving capture of wild individuals and ex
situ treatment, combined with selection for resistant individuals that could ultimately form
the basis of lineages that could be released back into the wild, are the only viable options
at present (see earlier). It may be possible to collect animals for survival-assurance colonies
by making use of predictive modelling and thereby move ahead of the planned spread of the
disease, as was conducted by Project Golden Frog (PGF) (see Essay 11.5) and attempted with
EVACC. Other preventative measures in the interim include the disinfection of vehicles and
footwear with 10% chlorine bleach solution to reduce the risk of pathogen dispersal21, and
distributing signs to make people aware of the disease, the risk of spread, and means to
sterilize potentially affected items. The best way to prevent unwanted diseases is to block
their entry to a region. Australia affords an example of such a strategy. In July 2002, the
Australian Government Minister for the Environment and Heritage decided that a threat
abatement plan (TAP) “was a feasible, effective and efficient way to abate the infection
process, and directed a nationally coordinated threat abatement plan be prepared to guide
management of the impact of the amphibian chytrid fungus on Australian amphibians.” A
TAP was adopted in 2006 and one goal was to prevent amphibian populations or regions
that are currently chytridiomycosis-free from becoming infected by blocking further spread
of the amphibian chytrid within Australia.
Water quality
Earlier we discussed evidence that changes in water quality and water flow regimes are
often a significant threat to amphibian species globally. Changes in water quality may include
changes in sediment load as a result of upstream logging, but often are also due to the
influence of environmental contaminants or pollutants (including pesticides, pharmaceuticals,
and heavy metals). In much of the Neotropics, for example, lowland agricultural regions are
subject to extensive agricultural spraying (e.g., Castillo et al. 1997). Furthermore, abiotic
factors (temperature, salinity, pH, and ultraviolet-B radiation) may also pose a threat when
normal values are exceeded and can, in turn, influence the toxicity of other contaminants.
Most work to date has focused on the detrimental effects of contaminants on the aquatic
life-stage of amphibians, although recent studies have also demonstrated that contaminants
can have negative effects in the terrestrial environment (e.g., Hayes et al. 2002).
As already discussed, actions targeted at contending with changes in hydrological
processes are already incorporated into the recovery plans of several species, and gener-
ally involve actions that may not be directed explicitly at actual sites where declines of
amphibians are occurring (such as protected areas). However, there are instances in which
such intervention may be required. Boone et al. (2007) propose that sites where amphibian
declines are taking place should be evaluated for environmental contamination that may
be present, either as a result of direct application or movement through air or water, based
on which emergency clean-up action may be necessary. Here again, the recovery plan for
the California Red-legged Frog Rana aurora draytonii22 exemplifies a management plan that
identifies several contaminated areas (e.g., Vandenberg Air Force Base in Santa Barbara
County) where the species may be exposed to toxins, and proposes that contaminants should
be removed and other measures that decrease exposure of the frog should be undertaken at
these sites. In such cases, animals should be removed from the site while clean up occurs and
either relocated or allowed to disperse back into ponds once water quality is improved.
Over-harvesting
Finally, species-targeted responses may be necessary to deal with the threat of over-har-
vesting, either for species in use for medicinal or food purposes (e.g., Telmatobius spp. or
Ambystoma spp.; and see Essay 11.8) or for the pet trade (e.g., dendrobatid frogs, Mantella
spp., salamanders, and newts of the genera Tylototriton, Paramesotriton, and Cynops). The
collecting of animals and plants usually is not permitted within protected areas, or at least
not within particular zones. However, the resources available to ensure collecting only in
assigned areas and management are often inadequate, and illegal collecting often takes
place even within nature reserves (as in the case of the Chinese Giant Salamander; see
Wang et al. 2004; Essay 4.7).
In-situ sustainable harvesting has real potential as a management option providing that
the biological requirements of the species concerned are understood and proper management
is in place (see Essay 11.9). Determining sustainable harvest quotas is complex and demands
information on the abundance, behaviour, and life-history characteristics of the species in
question, as well as an understanding of a host of socio-economic factors. In a recent survey
of levels of exploitation of species in the genus Mantella in Madagascar, Rabemananjara
et al. (in press) note that because of the apparent high density of many populations, and
their restricted and patchy distributions, these frogs may be well suited for a controlled
system of sustainable harvesting. Most Mantella species are characterized by a rather high
reproductive output, with up to 150 eggs per clutch, and the capacity to produce several
clutches per season. Despite the high to very high numbers of several Mantella species that
have been continuously exported from Madagascar over the past 15 years, most species
appear not to have been directly impacted as a result of over-harvesting. Rabemananjara
et al. (in press) visited populations of Mantella aurantiaca and M. milotympanum that local
collectors claimed to have heavily exploited during past years (e.g., Vieites et al. 2005), and
found Mantella to be very common at these sites. Rabemananjara et al. (in press) propose
transferring the focus of the trade regulation more to the sites of collection that are usually
in non-protected areas threatened by habitat destruction.
Even where the capture of wild specimens does not yet constitute a threat to a species, the
development of in-situ harvesting practices could provide an incentive motivating local land
owners and entire communities to protect native habitat instead of clearing it, with resultant
conservation benefits. A local, sustainable, production programme for species of relatively
high worth (e.g. Mantella frogs or dendrobatid frogs) could also have the additional benefit
of playing a significant role in poverty reduction strategies in some of the poorest parts of
the world. For example, the Global Environment Facility contributed financial support to a
project in Peru aiming to promote sustainable cultivation of poison dart frogs of the genera
Epipedobates and Dendrobates23 for export. These frogs fetch US$40-120 in hobbyist markets
in the United States, Europe and Japan. The Peru project established a business that exports
the frogs to hobbyists in these countries. This business – with expected annual revenues of
about US$300,000 in Year 3 – in turn provides supplemental livelihoods to more than 250
farmers, an incentive for preserving over 15,000ha of tropical forest, and the sustainable
breeding methods conserve the populations of more than 60 species of frogs.24
THE NEED FOR POLICY-LEVEL INTERVENTION
Policy-based actions are essential for providing the institutional support, human and financial
resources, and legal framework to ensure effective biodiversity conservation. Frequently,
such actions occur by developing and implementing legislation at the national or sub-national
levels, or through international agreements (e.g., Table 1).
Legislation is typically regarded as being directed at the protection of particular species,
such as by regulating the harvesting of individuals (e.g., Convention for the Regulation of
Whaling25), their trade (e.g., Convention on International Trade in Endangered Species of
Wild Fauna and Flora; CITES26), or alterations in their habitat (e.g., the Ramsar Conven-
tion27). In the United States, for example, species listed under the United States Endangered
Species Act (1973)28 are protected from exploitation and disturbance, and their habitats
are subject to legal protection. As of October 2006, more than 30 amphibian species were
listed on the ESA (Table 2) – with a few additional species proposed as candidates for list-
ing (such as Mountain Yellow-legged Frog Rana muscosa, CR) – including several species
not native to the US29.
Legislation can also promote habitat protection, with perhaps the most well known being the
Convention Concerning the Protection of the World Cultural and Natural Heritage, better known
as the World Heritage Convention (1972)30. Legislation also has the potential to protect habitat
 Chapter 11. Amphibian Conservation – Responding to the Global Decline of Amphibians 119

by regulating land use patterns at a broader scale (e.g., Brazil’s Forestry Law). Finally, legislation
can also regulate anthropogenic activities that are frequently the most pervasive causes of National legislation International agreements
species declines (e.g., pollution generated by industry, transport leading to the introduction of Species United States Endangered Species Act (1973): the species listed are Convention on International Trade in Endangered Species
invasive species, consumption of fossil fuels leading to climate change). protected from exploitation and disturbance, and their habitats are of Wild Fauna and Flora (CITES; 1975; global): regulates
There are now more than 500 international treaties that concern the environment31, and subject to legal protection. international trade of the species listed on its appendices.
most countries have ratified key international treaties, although several remain unratified The Bern Convention on the Conservation of European Wildlife
by key governments; for example, several countries in Asia and the Middle East, where and Natural Habitats (1979; regional): ensures the conservation
significant commercial trade in species occurs, have yet to join CITES. Such international of European wildlife and natural habitats via cooperation
treaties give conservation agencies a stronger mandate domestically (Steiner et al. 2003), between member States.
which is important given that most conservation action takes place at the national level. Of Sites and habitats Brazil’s Forestry Law (1965): establishes that each rural property in EC Habitats Directive (EC Council Directive on the Conservation
course, any legislation is only useful if it is adequately implemented, and such implementa- the Amazonian basin must preserve at least 80% of its forest cover. of Natural Habitats of Wild Fauna and Flora; 1992; regional):
tion is lacking in many cases. Thailand’s Wild Animals Reservation and Protection Act (1960) and the natural habitat listed must be maintained at a favourable
National Park Act (1961): legal basis for the creation of conservation status, particularly by creating a network of protected sites
Species-targeted legislation areas or protected areas, including national parks (144 sites), wildlife (Natura 2000 network).
sanctuaries (53 sites), forest parks (42 sites), wildlife non–hunting Ramsar Convention on Wetlands (1975; global): provides the
In terms of international legislation specifically targeting species, the most relevant for amphib- areas (52 sites), biosphere reserves (1 sites), World Heritage Natural framework for national action and international cooperation for
ians is CITES, instituted in 1975 (and see Table 3). Effectively, CITES is an agreement between sites (1 sites), watershed class 1 and conservation of mangroves. the conservation and wise use of wetlands and their resources,
governments designed to ensure that international trade in wild animals and plants does not in particular through the designation of sites under the Ramsar
threaten their survival. Currently, CITES attempts to regulate the trade of some 30,000 different List of Wetlands of International Importance.
species around the world, both plants (25,000) and animals (5,000), whether dead or alive, World Heritage Convention (1972; global): encourages the
whole animals, parts or products (e.g., carved ivory). Membership to the Convention is voluntary, identification, protection and preservation of cultural and
and member countries, known as Parties, act jointly by agreeing to restrict international trade natural heritage (including habitats of threatened species)
in a particular species that might be threatened with extinction. around the world considered to be of outstanding value to
In effect, CITES restricts and regulates trade in wildlife species. All forms of import and humanity. Countries submit places for designation under the
export are regulated by means of appendices, which have varying degrees of flexibility ac- World Heritage List.
cording to the required protection. Appendix I includes species threatened with extinction, Activities China’s Law on Environmental Impact Assessment (2003): requires United Nations Framework Convention on Climate Change
and trade in specimens of these species is permitted only if the purpose of the import is that governmental and non-governmental planning on land use, (1994) and Kyoto Protocol (adopted in 1997, not yet into
non-commercial (e.g., scientific study). Appendix II includes species that are not necessarily urban engineering, communication, and natural resource exploration force; global): caps greenhouse gas emissions in participating
threatened with extinction, but in which trade must be controlled to avoid levels of use that goes through the process of an environmental impact assessment. industrialized nations from 2008 to 2012 and establishes an
would be incompatible with their survival. All trade in Appendix I and Appendix II species is Australian quarantine regulations: strict control measures aimed at international market in emissions credits that will allow these
supposed to be on the basis of a “non-detriment finding” indicating that the trade in question preventing the introduction of pests and diseases (mainly established nations to seek out the most cost-effective means to reduce
will not harm the conservation status of the species. Appendix III contains species that are to protect the agricultural sector, but also human health and the atmospheric concentrations of greenhouse gases.
protected in at least one country, and which has then asked other CITES Parties for assistance native flora and fauna).
in controlling the trade. At the request of any Party, a species can be included on Appendix III. The Federal Constitution of the United States of Mexico
The Secretariat and Parties to the Convention meet every two years to revise the Appendices, (Constitución Federal de Los Estados Unidos Mexicanos) Article
and species may be downgraded or upgraded depending on their status (for an amendment to 27: states that all natural resources are property of the nation,
be made, there must be a two-thirds majority of Parties present), and to debate the benefits of therefore all species of flora and fauna are protected. To use, collect,
opening trade. For this reason, every Party to the Convention designates a Scientific Authority kill, transport or trade any amphibians, either living specimens or
for advice on the benefits or detriments of trade, and as to the current status of species. Par- their parts, requires a collection permit, issued by the Mexican
ties are also required to submit annual reports detailing exports and imports of CITES-listed environmental authority (SEMARNAT).
species (although some countries have never submitted a single report). A Party that does not
agree with a species listing may enter a reservation32 and is then considered a non-Party with
respect to the species in question, until the reservation is withdrawn. Table 1. Examples of national legislation
Since its inception, CITES has proved to be a successful agreement, and remains one of and international agreements relevant to
the largest of its kind in the world. International bans implemented by means of the CITES Scientific Name Where Listed Listing Status IUCN Category amphibian conservation, for the conservation
Appendices have done much to slow the trade in wildlife products across borders, and it is Eleutherodactylus jasperi entire T CR of particular species, for the protection of sites
perhaps a measure of CITES success, that since its inception, no species listed on one of its Rana aurora draytonii subspecies range clarified T NT or habitats, and for the regulation of activities
Appendices has been declared formally Extinct because of international trade. Nonetheless, Rana chiricahuensis entire T VU that can pose threats to biodiversity. Dates
several species known to be threatened by international trade are not currently listed on any Conraua goliath entire T EN correspond to when the agreement entered
CITES Appendix. According to the Global Amphibian Assessment, 47 amphibian species that Discoglossus nigriventer entire E EX into force. For international agreements,
are listed as threatened are adversely impacted by exploitation for the international trade; Rana capito sevosa Wherever found west of Mobile E CR1 there is an indication of whether their scope
currently, only 20 of these species are listed on Appendix I or II of CITES, leaving several and Tombigbee Rivers in AL, MS, is global (any country can ratify it) or regional.
and LA
species as priorities for inclusion on a CITES Appendix (Table 4). Note: many of these laws/agreements could
CITES has some drawbacks, largely because of which trade has been able to continue Rana muscosa southern California DPS E CR be listed under two or more categories (e.g.,
in many parts of the globe unchecked. First, as already noted, not all countries are Parties Atelopus varius zeteki entire E CR2 the EC Habitats Directive simultaneously
to the convention (currently, there are 169 signatories). Since membership is voluntary, any Leiopelma hamiltoni entire E EN provides for the protection of species and
country not Party to the convention need not report to CITES, and so trade between these Eleutherodactylus cooki entire T EN sites/habitats, and for the regulation of
countries goes largely undocumented. These countries are, in effect, loopholes through which Eurycea sosorum entire E VU activities). Many national biodiversity acts,
species can be illegally traded. Secondly, CITES, while legally binding, is an international Ambystoma californiense U.S.A. (CA - Santa Barbara County) E VU such as South Africa’s Biodiversity Act of 2004,
body, and as such governs only international trade. The Convention does not replace domestic Ambystoma californiense U.S.A. (CA - Sonoma County) E VU also fall in multiple categories. Adapted, with
legislation, and its effectiveness is hamstrung by the degree to which countries implement Ambystoma californiense Central CA DPS T VU permission, from Baillie et al. (2004).
and enforce their own domestic wildlife trade policies. In China, for example, the Chinese Plethodon nettingi entire T NT
Giant Salamander Andrias davidianus, several Tylototriton spp. and Hoplobatrachus rugulosa Andrias davidianus entire E CR
(=davidianus d.)
are listed as Class II State Key Protected Animals whose collection, transport, cultivation,
Batrachoseps aridus entire E 3
and sale requires permission from provincial authorities. Regardless, law enforcement is
inadequate and illegal collecting is widespread (Wang et al. 2004). Further, while some Ambystoma cingulatum entire T VU
Parties to CITES have domestic legislation with trade controls stricter than those required Andrias japonicus entire E NT Table 2. Amphibians listed on the United
(=davidianus j.)
by the Convention, the effectiveness with which CITES is implemented varies between States Endangered Species Act as of October
countries. At the time of writing, an Amphibian Identification Guide from Environment Phaeognathus hubrichti entire T EN 2006. E = Endangered; T = Threatened. Endan-
Canada containing all CITES protected species, is in the final stages of preparation. The Eurycea nana entire T VU gered species are those in danger of extinction
guide is designed specifically for customs officers, to aid in the identification of protected Ambystoma macrodactylum entire E LC throughout all or a significant portion of their
croceum
species. It includes a colour key and a descriptive page for each species with full colour range; Threatened species are those that are
illustrations. Texts are in three languages: English, French and Spanish. Also, while CITES Plethodon shenandoah entire E VU likely to become Endangered in the foresee-
is intended to protect species threatened by international trade, it can sometimes hinder Ambystoma tigrinum stebbinsi entire E LC able future. Note that these categories are
conservation, for example, blocking or delaying international movement of entire animals for Typhlomolge rathbuni entire E VU4 independent of the IUCN Categories of Threat
ex-situ management programmes or parts thereof (material for genetic analysis). Bufo californicus entire E EN (final column), but species appearing on the US
(=microscaphus)
Endangered Species list are an under-repre-
Habitat-based legislation Bufo superciliaris entire E LC sentation of species known to be threatened
Bufo houstonensis entire E EN with extinction. 1considered a distinct species;
Several international treaties are concerned with site or habitat protection, including the Bufo periglenes entire E EX 2considered a distinct species; 3considered

Ramsar Convention on Wetlands that has designated 1,429 Wetlands of International Peltophryne lemur entire T CR5 a subspecies of Batrachoseps major (LC);
Importance for conservation and wise use and the World Heritage Convention that has Bufo baxteri (=hemiophrys) entire E EW 4included in the genus Eurycea; 5as Bufo
6
designated 830 properties as part of its World Heritage List. However, the most powerful Nectophrynoides spp. entire E lemur; 611 species recognized (8 threatened,
convention related to protected areas is the Convention on Biological Diversity (CBD)33, signed 2 DD, 1 LC).
by 150 government leaders at the 1992 Rio Earth Summit, and dedicated to: a) conserving
biological diversity; b) sustainable use of its components; and c) fair and equitable sharing
of the benefits arising from the use of genetic resources. The CBD is currently focused on as the central element of their strategy to implement the Convention. Since the 1960s, the
the 2010 Biodiversity Target: to achieve by 2010 a significant reduction of the current rate IUCN World Congress in National Parks and Protected Areas has fundamentally influenced
of biodiversity loss at the global, regional and national level as a contribution to poverty the protected areas agenda. The Congress, which meets every 10 years, provides a forum
alleviation and to the benefit of all life on earth. To achieve this ambitious target, the CBD for discussion on all ecological, social, economic, political, and practical matters related to
has employed a number of strategies, including the development and implementation of protected areas. The establishment of a 10% target for protected areas stemmed from the
national biodiversity strategies and action plans (NBSAPs), thematic programmes (e.g., on Fourth World Parks Congress in Caracas, Venezuela, in 1992, where it was recommended
forests, inland waters, mountains, etc), and cross-cutting issues (e.g., on plants, taxonomy, “that protected areas cover at least 10 percent of each biome by the year 2000” (IUCN
invasive alien species, and protected areas). 1993). Subsequently, the 10% target for protected areas became deeply entrenched in
Protected areas are a central part of the CBD since the Parties themselves have con- the thinking of many conservationists and incorporated into the national legislation of
sistently identified efforts to develop and maintain their national protected area systems many countries for establishing protected areas. It was, and still is, generalized to apply
120 Threatened Amphibians of the World

Table 3. Amphibians listed on CITES Ap-

pendices I and II. No amphibians are listed Appendix I Appendix II Appendix III Order Family Name Appendix I Appendix II Not listed
on Appendix III. Many of the amphibians ANURA Anura Bufonidae Atelopus cruciger y
listed are traded internationally as pets (e.g., Bufonidae Toads Anura Bufonidae Atelopus zeteki y
Phyllobates, Dendrobates, Epipedobates, Altiphrynoides spp. Anura Bufonidae Melanophryniscus y
Mantella, Dyscophus antongilii, Scaphi- Atelopus zeteki cambaraensis
ophryne gottlebei, Ambystoma). Euphlyctis Bufo periglenes Anura Dendrobatidae Dendrobates altobueyensis y
hexadactylus and Hoplobatrachus tigerinus Bufo superciliaris Anura Dendrobatidae Dendrobates bombetes y
are in the frog-leg trade. Andrias davidianus Nectophrynoides spp. Anura Dendrobatidae Dendrobates granuliferus y
is exploited for food, but is not generally in Nimbaphrynoides spp. Anura Dendrobatidae Dendrobates lehmanni y
international trade. Other CITES-listed spe- Spinophrynoides spp. Anura Dendrobatidae Dendrobates opisthomelas y
cies (e.g., some of the African toad genera Dendrobatidae Poison frogs Anura Dendrobatidae Dendrobates speciosus y
such as Nectophrynoides) have never been Allobates femoralis Anura Dendrobatidae Dendrobates steyermarki y
the subject of international trade, and so the Allobates zaparo Anura Dendrobatidae Dendrobates virolensis y
utility of their listing is doubtful. The genus Dendrobates spp. Anura Dendrobatidae Phyllobates terribilis y
Rheobatrachus is Extinct, but not because of Epipedobates spp. Anura Hylidae Agalychnis annae y
international trade. Phyllobates spp. Anura Hylidae Phyllodytes auratus y
Mantellidae Mantellas Anura Leptodactylidae Caudiverbera caudiverbera y
Mantella spp. Anura Mantellidae Mantella aurantiaca y
Microhylidae Red rain frog, tomato frog Anura Mantellidae Mantella bernhardi y
Dyscophus antongilii Anura Mantellidae Mantella cowanii y
Scaphiophryne gottlebei Anura Mantellidae Mantella crocea y
Myobatrachidae Gastric-brooding frogs Anura Mantellidae Mantella expectata y
Rheobatrachus spp. Anura Mantellidae Mantella madagascariensis y
Ranidae Frogs Anura Mantellidae Mantella milotympanum y
Euphlyctis hexadactylus Anura Mantellidae Mantella pulchra y
Hoplobatrachus tigerinus Anura Mantellidae Mantella viridis y
CAUDATA Anura Microhylidae Platypelis milloti y
Ambystomidae Axolotls Anura Microhylidae Scaphiophryne boribory y
Ambystoma dumerilii Anura Microhylidae Scaphiophryne gottlebei y
Ambystoma mexicanum Anura Microhylidae Scaphiophryne marmorata y
Cryptobranchidae Giant salamanders Anura Ranidae Conraua goliath y
Andrias spp. Anura Ranidae Limnonectes macrodon y
Anura Ranidae Rana shqiperica y
Caudata Hynobiidae Hynobius tokyoensis y
Caudata Hynobiidae Ranodon sibiricus y
to individual countries and to the entire planet, despite its proven shortcomings (Soulé and Caudata Plethodontidae Plethodon petraeus y
Sanjayan 1998; Rodrigues et al. 2004a). Caudata Salamandridae Cynops ensicauda y
At the Fifth Congress held in South Africa in 2003, a workshop on “Building Comprehensive Caudata Salamandridae Echinotriton andersoni y
Protected Area Systems” demonstrated that despite substantial gains, global protected area Caudata Salamandridae Lyciasalamandra antalyana y
systems are still far from comprehensive, and reiterated that percentage-based targets are Caudata Salamandridae Lyciasalamandra atifi y
not sufficient for protecting our biodiversity. Results from the Global Amphibian Assessment Caudata Salamandridae Lyciasalamandra billae y
were key to building the case (see Essay 11.2). The Congress report to the CBD stated Caudata Salamandridae Lyciasalamandra fazilae y
that “the global system of protected areas needs to safeguard all globally and nationally Caudata Salamandridae Lyciasalamandra flavimembris y
important areas for biodiversity”, and in the Durban Accord asked the global community for Caudata Salamandridae Lyciasalamandra luschani y
a “commitment to expand and strengthen worldwide systems of protected areas, prioritized Caudata Salamandridae Neurergus kaiseri y
Table 4. Species threatened by international on the basis of imminent threat to biodiversity”. Several countries responded immediately, Caudata Salamandridae Neurergus microspilotus y
trade according to the results of the Global with the President of Madagascar and the Governors of the Brazilian states of Amazonas Caudata Salamandridae Paramesotriton fuzhongensis y
Amphibian Assessment, indicating those spe- and Amapá announcing at the Congress itself that they would significantly expand their Caudata Salamandridae Salamandra algira y
cies already listed on a CITES Appendix, and protected area systems. Caudata Salamandridae Tylototriton kweichowensis y
those for which legislation is required. Building on these commitments, the Seventh Conference of the Parties (the governing body
of the CBD) adopted a Programme of Work on Protected Areas (Decision VII/2834) with “the
objective of the establishment and maintenance by 2010 for terrestrial and by 2012 for marine
areas of comprehensive, effectively managed, and ecologically representative national and • Emissions Trading: the option to buy and sell emissions credits among developed
regional systems of protected areas”. This Programme of Work has four elements (implementa- countries. If one country exceeds its emissions target for one year, while another’s falls
tion, governance and equity, enabling activities, and monitoring) each with several goals. The below its targets, the first is permitted to purchase the second country’s unused emission
first goal of the first element −“to establish and strengthen national and regional systems of credits to meet its emission target.
protected areas integrated into a global network as a contribution to globally agreed goals,” • Joint Implementation: a country can receive emissions credits for a specific project
bears mentioning since it requires the identification of sites of global biodiversity significance undertaken in another country.
in each country to determine which sites are currently not represented in protected area • Clean Development Mechanism: developed countries can receive emissions credit for
systems, and sets priorities for new actions among these sites (and therefore links directly financing projects that reduce emissions in developing countries
back to the need to identify sites requiring urgent habitat protection).
In 2002, the CBD adopted the Global Strategy for Plant Conservation, which includes CO2 emissions from deforestation are another type of emission that can be profitably
16 targets under five general headings: a) understanding and documenting plant diversity; reduced if allowed into a market. At present, avoided deforestation (AD) carbon trading is
b) conserving plant diversity; c) using plant diversity sustainably; d) promoting education restricted to particular voluntary markets in the US. The much larger Kyoto Protocol-based
and awareness about plant diversity; and e) building capacity for the conservation of plant European markets do not currently trade in avoided deforestation, largely due to political
diversity. In view of the amphibian crisis documented in this book, a multifaceted CBD global concerns of the past. Developing countries took the position that developed countries should
strategy on amphibians, building on the ACAP, should now be considered, and at the time of take the first step toward tackling climate change. Now that a European carbon market is
going to press, discussions were ongoing regarding the incorporation of a global strategy in place, developing countries see a financial opportunity to sell emissions reductions from
on amphibians into the CBD. AD. Credits for AD would be related only to the amount of deforestation avoided, not the
total amount of forest. This would be true for either a project-level or country-level scheme.
Environment Canada have produced an Legislation aimed at regulation of activities Imagine a country that has one million hectares of forest, and an annual forest loss baseline
Amphibian Identification Guide containing of ten thousand hectares per year39. If this loss is reduced to 5,000ha per year, the country
all CITES listed species, specifically to aid Among international legislative agreements regulating activities that can pose threats to would be credited for a 5,000ha/year reduction, not their entire million hectare carbon
customs officers in the identification of biodiversity, arguably the most important is the United Nations Framework Convention on stock. A proposal to bring AD into the Kyoto Protocol was introduced in November 2005 by
protected species. Image courtesy of Tamara Climate Change35 and the Kyoto Protocol. The UNFCCC, which entered into force in 1994, is the Coalition for Rainforest Nations (CRN), a group of tropical developing countries led by
Maliepaard. an international treaty that sets an overall framework for intergovernmental efforts to tackle Papua New Guinea and Costa Rica. This concept was originally proposed by Santilli et al.
the challenge posed by climate change. The Convention has near universal membership, with (2005) under the name “compensated reduction.” The CRN proposal has been endorsed to
the ratification of 189 countries36. Recently, a number of nations have approved an addition date by the African Union, European Union, Pacific Island Forum, British Commonwealth, As-
to the treaty: the Kyoto Protocol37, an international and legally binding agreement to reduce sociation of Small Island States, and most recently, Indonesia. Several years of international
greenhouse gas emissions world wide, which came into effect on 16 February 2005. The negotiations will need to take place before an agreement can be finalized.
Kyoto Protocol significantly strengthens the Convention by committing Annex I Parties to In the absence of an international treaty specifically designed to reduce greenhouse gases,
individual, legally binding targets to limit or reduce their greenhouse gas emissions. However, the Intergovernmental Panel on Climate Change40 estimates that global mean temperatures
only Parties to the Convention that have also become Parties to the Kyoto Protocol (i.e., by will rise by between 1.4°C and 5.8°C, with resulting catastrophic consequences for weather
ratifying, accepting, approving, or acceding to it) are bound by the Protocol’s commitments. patterns, biodiversity, and sea levels. However, limiting climate change requires major
165 countries have ratified the Protocol to date, with the most notable exceptions being changes beyond simply signing the Kyoto Protocol; it involves the evolution of a society that
the United States and Australia. becomes carbon neutral on a global scale in this century (Lovejoy and Hannah 2005).
Under the Kyoto Protocol, developed countries (termed Annex 1 countries) have com-
mitted themselves to reducing their collective emissions of six key greenhouse gases by at
least 5.2%, with a reduced emissions target of 2008-2012 (calculated as an average over RESEARCH AS A CONSERVATION ACTION
the five years)38. Although these caps are national-level commitments, most countries will
devolve their emissions targets to individual industrial entities. The complicating factor is that Conservation action is more effective when supported by knowledge, not just on species
actual emission reductions will need to be much larger than 5%, because for some wealthy themselves, but on the threats affecting species and the most effective measures needed for
industrialized countries reduction targets will need to be as much as 15% lower than their addressing those threats. Furthermore, research that is integrated with the implementation of
expected emissions in 2008. To provide countries with the flexibility to meet their emission conservation can allow for adaptive management of projects, thereby building on successes
targets, the Kyoto Protocol has established three market-based mechanisms: and learning from failures in a manner that best serves the species at risk. The results of
 Chapter 11. Amphibian Conservation – Responding to the Global Decline of Amphibians
the Global Amphibian Assessment provided a much-needed context for the well-publicized
phenomenon of amphibian declines, but they also provide us with directions and priorities
for research needs, not only on a taxon basis, but also on a thematic basis.
Among the species that emerge as priorities for research are those flagged as Data
Deficient. The Red List category Data Deficient is assigned to species for which there is
inadequate information available to make a direct, or indirect, assessment of its risk of
extinction based on distribution or population status. As we have already seen, nearly one-
quarter of all amphibians are classified as Data Deficient for one of three reasons: unknown
provenance; taxonomic uncertainty; and/or inadequate data. Data Deficient species tend to
be concentrated in regions with high biodiversity, but are poorly surveyed (Figure 4). There
are also noticeable concentrations in regions where recent taxonomic investigations are
revealing unexpected complexities in terms of species limits, such as in the Western Ghats
of India (see Essay 7.2).
It is necessary to preface a discussion of research priorities and needs with a note that
classification in the Data Deficient category does not imply lack of threat. Indeed, IUCN’s
(2006) user guidelines explicitly note “it may be appropriate …to give them the same degree
of attention as threatened taxa until their status can be assessed.” The pessimistic listing
of actual Data Deficient species as threatened during the Red Listing process can lead to a
muddling of conservation priorities with research priorities. This stems from concerns that
species listed as Data Deficient are seldom beneficiaries of conservation investment, because
they are not listed as threatened (e.g. see Garnett et al. 2003). However, Data Deficient spe-
cies should not be the immediate targets for conservation action per se: by definition, they
are priorities for research action, be it in the form of taxonomic investigation or field-based
survey work, which in turn may reveal the need to conserve these species.41
Taxonomy
Although it is not yet possible to provide a reliable figure on which species are in need of
further taxonomic investigation (i.e., those Data Deficient species so categorized because
of taxonomic uncertainty), there are particular taxa that repeatedly emerge as in need of
taxonomic work, including confused genera such as Arthroleptis (14 of 34 species Data
Deficient), Caecilia (23 of 35), and Ichthyophis (27 of 34).
The taxonomic impediment is by no means restricted to those known species catego-
rized as Data Deficient based on issues surrounding their actual taxonomic status. Just as
species description rates continue to increase (Köhler et al. 2005), new studies that use
complementary diagnostic characters, such as genetic sequences and acoustic features, are
revealing unexpected and cryptic diversity within wide-ranging taxa, with the implication
that single, widespread ‘species’ may actually represent multiple species having smaller
geographic ranges, and, consequently, greater vulnerability to extinction (Stuart et al. 2006a).
In the last mentioned study, the authors found that torrent frogs that have been identified to
date on the basis of morphology as Rana livida and Rana chalconota represent at least 14
different species. Indeed, it is becoming increasingly apparent that morphology alone will
not suffice for the purposes of biodiversity inventory work, and that new and complementary
techniques are required to help identify species that warrant recognition and protection
(see Essay 11.10 and Essay 11.11). In this regard, there is clearly a need to expedite the
application of these techniques to known or suspected species complexes (e.g., Angulo et
al. 2003; Fu et al. 2005).
The implications of such taxonomic investigations for conservation planners are as yet
to be fully revealed, but may be far-reaching. It is to be expected that many Data Deficient
species will really emerge as threatened species in their own right. Likewise, unravelling
the complexities of current wide-ranging species complexes will likely result in a marked
increase in the numbers of species currently considered threatened, and probably will see
many species move from a lower category of threat to a higher category of threat. This is
as a result of the influence of splitting formerly wide-ranging species on overall population
size and geographic range size, which are key factors in the IUCN Red List criteria (Collar
1996). In turn, a surge in threatened species will have implications for those initiatives
incorporating threatened species listings, of which the Red List is the most commonly
used, into priority setting exercises. For example, studies that have investigated how the
influence of differing species concepts will affect the identification of important areas for
conservation have shown conflicting results (e.g., Peterson and Navarro-Sigüenza 1999;
Dillon and Fjeldså 2005).
One possible side-effect of the windfall of species descriptions is that in the race to
describe new species in the face of a biodiversity crisis, the publication of new descriptions
of rare species might inadvertently facilitate their extinctions if the species in question turn
out to have commercial value (Courchamp et al. 2006). Most modern species descriptions
121
Figure 4. Richness map of Data Deficient
amphibian species, with darker reds corre-
sponding to regions of higher richness. Colour
scale based on five quantile classes. Maximum
richness equals 18 species. Note: this map is
only an approximation, since for most Data
Deficient species the extent of occurrence is
poorly known.
provide very detailed information on the locality and habitat where the new species occurs,
turning “…a scientific article into a treasure map for commercial collectors” (Stuart et al.
2006b). The latter authors cite the example of the salamander Paramesotriton laoensis
from northern Laos, which was not known in the international pet trade prior to its recent
description as a new species (Stuart and Papenfuss 2002). Over the past year, Japanese and
German collectors used the published description to find these salamanders, and they are
now being sold to hobbyists in those countries for US$170 to US$250 each. In the face of
recognition that withholding such locality information hampers conservation and science as
much as it does those seeking to profit from it, Stuart et al. (2006b) propose that taxonomists
should work closely with relevant governmental agencies to coordinate publication of the
description with legislation or management plans that thwart overexploitation of the new
species and that while this might lengthen publication time, alternative solutions that allow
taxonomists to continue their work without contributing to species decline are wanting.
Surveys and Monitoring
Clear descriptions of species by taxonomists go hand in hand with survey work. Several
large regions, such as Indonesian Papua, Indonesian Borneo, central Africa and the Albertine
Rift, and parts of the Andean countries, remain very poorly surveyed, and further surveys
very likely will reveal impressive levels of diversity. To cite just one recent study, Stuart
et al. (2006c) undertook the first ever collection of amphibians in hilly eastern Cambodia
since Henri Mouhout’s work in 1859. Their collection comprised 30 species of amphibians,
including two new species Leptobrachium mouhoti and Ophryophryne synoria, not to mention
11 new country records for Cambodia!
While such herpetological collections have great value in terms of improving knowledge
on a country’s national biodiversity assets, and in turn have immediate implications for
national conservation strategies, there is also a need for targeted species surveys. Among
those species urgently in need of survey work – indeed, representing the top priorities
for field-based survey work – are species considered Critically Endangered, and flagged
as Possibly Extinct (PE). This marker was designed specifically to avoid the Romeo Error
(Collar 1998) – scenarios in which we might give up on a species as being extinct before
it truly disappears (Butchart et al. 2006a). The term was first applied to the case of a bird,
the Cebu Flowerpecker Dicaeum quadricolor from the Philippines, which was rediscovered
in 1992 after 86 years without a record (Dutson et al. 1993), having been written off as
extinct at least 40 years earlier on the presumption that no forest remained on the island of
Cebu (Magsalay et al. 1995). Such remarkable rediscoveries are by no means unique, with
several species having been rediscovered, sometimes as long as a century after their date
of last record. Among amphibians, recent rediscoveries include a surviving population of
Atelopus cruciger in Venezuela, and the rediscovery of Atelopus nahumae and A. laetissimus,
both believed to have gone extinct in the Colombian Andes. On the other hand, survey work
conducted in Lebanon to search for Hula Painted Frog Discoglossus nigriventer, a species
known only from the eastern shore of Lake Huleh in Israel and last recorded in 1955, failed
to record the species following tantalizing reports that it might survive in the Aammiq
marsh in south-east Lebanon (although the surveys did result in a new country record for
Pelobates syriacus) (Tron 2005).
Among those amphibians flagged here as CR(PE) (Appendix IX), one recent interesting
rediscovery published at the time of writing is that of Conraua derooi, which has not only
been rediscovered in the Togo Hills (Leaché et al. 2006), but has since been found outside
its former range in the Atewa mountains (M.-O. Rödel, pers. comm.). Habitat loss and deg-
radation is the major threat to this species (including increased sedimentation in breeding
streams), with the result that the most immediate conservation action required is effective
maintenance of natural habitat within the species’ limited range.
Besides survey work, population monitoring is a required conservation action for
most threatened species, especially for those where chytridiomycosis is believed to be a
potential threat but has not yet resulted in population declines. Another set of species for
which monitoring is important are those categorized as Vulnerable under the D2 criterion
(Appendix X). These species usually have stable populations, but typically have very small
distributions. As such, they are especially prone to the effects of human activities or
stochastic events that could result in the species becoming Critically Endangered or even
Extinct in a very short time period.
Threats
As we have already seen, while many declines are due to habitat loss and over-harvesting,
other factors are also operating, some of which are rapidly driving species to extinction.
122 Threatened Amphibians of the World
Some salamander species, such as Neurergus
kaiseri (Critically Endangered) from the south-
ern Zagros Mountains of Iran, are increasingly
threatened by the growing international pet
trade, but have yet to be officially proposed
for listing on a CITES appendix. © Twan
Leenders
Many of the reasons for amphibian declines have been acknowledged for decades (see
Collins and Storfer 2003); others are only now beginning to be better understood (Beebee
and Griffiths 2005). Stuart et al. (2004) coined the term “enigmatic decline species” for those
species that have shown dramatic declines (see Chapter 4), even where suitable habitat
remains, for reasons that are not fully explained. We now know that one of the primary
reasons for these enigmatic declines is likely the emerging infectious disease chytridiomy-
cosis (spread by its causal agent, the pathogenic fungus Batrachochytrium dendrobatidis)
(Lips et al. 2006), perhaps acting in deadly synergy with other factors, particularly climate
change (Pounds et al. 2006).
The most important research needs in terms of threats, include better understanding
of the impacts of climate change, the role of environmental contaminants in declines,
the spread and treatment of disease (specifically chytridiomycosis, but other emerging
infectious diseases, too), and, perhaps most important of all, the context-dependency of
these and other factors (Blaustein and Kiesecker 2002), particularly when compounded
with land use change.
In the past decade, a great deal has been learned about the ecology, biology and impact
of the chytrid pathogen (see Essay 4.5). This knowledge has included the development
of advanced molecular diagnostic tests (Annis et al. 2004; Boyle et al. 2004); analysis
of biochemical defences of amphibians against the pathogen (Rollins-Smith et al. 2002;
and see Essay 2.2); and the use of ecological niche modelling to predict the spread of
the disease (Ron 2005; and see Essay 11.4) and phylogenetic-based predictive models
(Lips et al. 2006). Nonetheless, there remain several unanswered questions concerning
this disease, and these offer important and urgent opportunities for targeted research
(Daszak et al. 2007):
• Studies on the ecology of B. dendrobatidis, including basic and critical aspects of its
natural history, such as how and where it survives, and how long it can persist in the
environment. Initial studies suggest it can survive for up to 8 weeks as a saprobe, an
organism that derives its nutrition from the dead remains of other organisms, in sterile
pond water (Johnson and Speare 2003).
• Understanding how amphibians respond to infection. Do any amphibians become immune
when first infected? We know, for example, that a number of species become susceptible
again even after they have cleared an initial infection. Do amphibians respond to infection
by changing behaviour, e.g., basking, to eliminate infections?
• How does the chytrid fungus spread? And is there any relationship between its spread
and trade in amphibians? Despite the availability of advanced modelling techniques to
predict the spread of disease, one of the highest priorities is to determine the means
by which B. dendrobatidis moves among sites, species, and individuals over varying
scales.
• Why does B. dendrobatidis cause declines in some amphibian populations, while others
are unaffected (e.g., Retallick et al. 2004; Lips et al. 2006). Studies are needed of how
immune responses, antimicrobial peptide responses (Woodhams et al. 2006; Harris et al.
2006), or the genetics of species (or populations) that appear to be tolerant of the disease
(e.g., American Bullfrogs), or completely resistant to infection, differ from susceptible
species.
• How does chytridiomycosis cause death? Does it affect the ability of frogs to respire
or osmoregulate through their skin, or does the pathogen release toxins that ultimately
cause death?
• What caused the emergence of B. dendrobatidis? This includes surveying museum col-
lections (e.g., Puschendorf et al. 2006) and conducting molecular phylogenetic studies
to determine when and where chytrid first emerged, or whether it has always been
widespread and local environmental conditions cause emergence and spread (Rachowicz
et al. 2005).
• Can lineages resistant to chytridiomycosis be selected for in captivity and reintroduced
to native habitats?
Research needs on the effects of environmental contaminants on amphibians are
presented by Boone et al. (2007), who emphasize again that there is a primary need to
examine the interactive effects of either multiple contaminants or contaminants with other
threats (such as disease, pathogens, climate change, and habitat alteration), to help plan
mitigation measures to thwart declines. Other important directions for research into the
threats posed by contaminants include:
• Improved understanding of the impact of contaminants on both aquatic and terrestrial
life stages (currently, there are limited data focusing on the terrestrial life-stage or
interactions between aquatic and terrestrial stages);
• Studies to investigate how contaminants affect populations over long time periods (e.g.,
Semlitsch et al. 1996), and how adaptation to a chemical stressor may influence population
persistence or make individuals vulnerable to other factors (e.g. Semlitsch et al. 2000).
• Investigating the effects of other common pollutants on amphibians (recent studies have
focused heavily on carbaryl, atrazine, and coal combustion wastes), particularly herbicides
and insecticides. For example, the herbicide atrazine disrupts the endocrine system (e.g.,
Hayes et al. 2002), and herbicides can reduce the food base of the community (Diana et
al. 2000; Boone and James 2003). Other groups of contamination resulting from industry
by-products, mining activities and oil and gas prospecting leakages, as well as human
and other animal waste products (pharmaceuticals, sewage by-products), also pose
potential threats that require study (Brooks et al. 2003).
• The types of contaminants and combinations of contaminants that amphibians are
exposed to in nature.
Finally, research related to climate change is needed. Recent climate change is causing
some species to breed earlier (e.g., Beebee 1995; Gibbs and Breisch 2001; Corn 2003), and
recent warming episodes may play some causal role in declines in some areas (Pounds et
al. 1999, 2006). However, there remains a need for further research to understand how
climate change affects amphibians, and given the loss of amphibians from pristine habitat,
the possible synergies between disease dynamics and climate change.
AMPHIBIAN CONSERVATION – CODA
Amphibian declines were open to debate only 15 years ago, but increasing evidence reveals
that many amphibians are undergoing cataclysmic declines, some to extinction, and that a
suite of causes is responsible. How we respond, and in what way, will depend on whether
the conservation community at large can focus attention on known priorities and use the
tools and knowledge available to act.
Some have argued that hope is already lost, the resources do not exist, and that the best
conservation scientists can do is act as scribes documenting the declines of a once diverse
fauna. We contend that to do so, in the face of explicit existing conservation options, would
be not unlike the fire trucks standing by while the Sistine Chapel burns. History has shown
that targeted conservation efforts can help the recovery of even the most threatened species,
including species that have declined to low numbers, such as the Rodrigues Fody Foudia
flavicans (Impey et al. 2002), Black Robin Petroica traverse (Aikman et al. 2001), Mallorcan
mid-wife toad Alytes muletensis (Essay 11.6), Antiguan Racer Alsophis antiguae (Daltry et
al. 2001), and Black-footed Ferret Mustela nigripes (Dobson and Lyles 2000).
In a recent study investigating how many bird extinctions conservationists have prevented,
Butchart et al. (2006b) used data on population sizes, population trends, threatening pro-
 Chapter 11. Amphibian Conservation – Responding to the Global Decline of Amphibians 123

cesses, and conservation actions, to identify at least 26 bird species surviving in the wild Corn, P.S. 2003. Amphibian breeding and climate change: the importance of snow in the mountains. Endnotes
that would have very probably gone extinct without conservation programmes. Of course, Conservation Biology 17:622-625. 1 www.zeroextinction.org
many still require sustained conservation efforts, and a major increase in global conserva- Courchamp, F., Angulo, E., Rivalan, P., Hall, R.J., Signoret, L., Bull, L. and Meinard, Y. 2006. Rarity value and 2 The cornerstones of AZE site identification, as with
tion resources is necessary to extend such efforts to thousands of threatened species. species extinction: the anthropogenic Allee effect. PLOS Biology 4(12):e415. that for KBAs, are the IUCN Red List categories
Nonetheless, such case studies prove that action directed at priority species can be a path Cowling, R.M., Pressey, R.L., Rouget, M. and Lombard, A.T. 2003. A conservation plan for a global biodiversity assigned to taxa that trigger the identification of a
to future recovery (Rodrigues 2006). hotspot – the Cape Floristic Region, South Africa. Biological Conservation 112:191-216. site in the first place. Thus, it is crucial that the Red
We have demonstrated that we have the information at hand to at the very least identify Daltry, J.C., Bloxam, Q., Cooper, G., Day, M.L., Hartley, J., Henry, M., Lindsay, K. and Smith, B.E. 2001. Five Listing process is rigorous, transparent, objective
which species require what type of conservation action, and which species require that action years of conserving the ‘world’s rarest snake’, the Antiguan racer Alsophis antiguae. Oryx 35:119-127. and repeatable, and appropriately supported by
first. This, in the end, is what defines our ability to mitigate biodiversity loss – our ability Daszak, P., Lips, K., Alford, R., Carey, C., Collins, J.P., Cunningham, A., Harris, R. and Ron, S. 2007. Infec- the relevant supporting documentation, given the
to set priorities for where we should invest resources. The resources are limited, but the tious diseases. In: C. Gascon, J.P. Collins, R.D. Moore, D.R. Church, J. McKay, and J.R. Mendelson III utility and value of these in conservation planning
constraints can act as a lens that concentrates options and resources on the most urgent. (eds), Amphibian Conservation Action Plan, pp. 21-25. IUCN/SSC Amphibian Specialist Group. IUCN, (see also Rodrigues et al. 2006).
We cannot expect to ensure habitat protection for the 90% of species for which habitat Gland, Switzerland. 3 Protection status is not known for 16 sites.
loss is a threat, but there are is a finite set of species for which our failure to secure the Diana, S.G., Resetarits, W.J., Schaeffer, D.J., Beckmen, K.B. and Beasley, V.R. 2000. Effects of atrazine 4 In this instance, three conservation NGOs
last remaining tracts of habitat now means, almost certainly, the loss of a country’s, even on amphibian growth and survival in artificial aquatic communities. Environmental Toxicology and - Fundación ProAves Colombia, Conservation
the Earth’s, unique biodiversity assets. Chemistry 19:2961-2967. International, and American Bird Conservancy
So, as we set out to put a face on the amphibian crisis in the pages that follow, we Dillon, S. and Fjeldså, J. 2005. The implications of different species concepts for describing biodiversity - joined by the IUCN/SSC Amphibian Specialist
end with a challenge. A proposed action plan has been developed with many ambitious patterns and assessing conservation needs for African birds. Ecography 28:682-692. Group, teamed up to protect a 1,600-acre site
components, including some not discussed above, such as establishing national networks Dobson, A. and Lyles, A. 2000. Black-Footed Ferret Recovery. Science 288:985-988. (the El Dorado Nature Reserve) on the north-west
in priority countries to monitor trade in amphibians. These actions should form part of a Dodd, C.K., Jr. 2005. Population manipulations. In: M.J. Lannoo (ed.), Amphibian declines: the Conserva- slope of the Sierra Nevada de Santa Marta
unified and coordinated conservation response that must address the top priorities. To tion Status of United States Species, pp. 265-270. University of California Press, Berkeley and Los massif, on the Caribbean coast of Colombia. The
coordinate and facilitate such targeted conservation programmes for amphibians, Mendelson Angeles, USA. site provides what is believed to be a critical last
et al. (2006) called for the formation of an Amphibian Survival Alliance (ASA) – under the Dodd, C.K., Jr. and Seigel, R.A. 1991. Relocation, repatriation, and translocation of amphibians and reptiles: refuge for several highly threatened amphibian
umbrella of the Amphibian Specialist Group (ASG) of the Species Survival Commission of are they conservation strategies that work? Herpetologica 47:336-350. species (including Atelopus laetissimus, A. nahu-
IUCN (see Essay 11.12). The ASA is envisioned as a partnership-based alliance, consisting Doubledee, R.A., Muller, E.B. and Nisbet, R.M. 2003. Bullfrogs, disturbance regimes, and the persistence mae, Colostethus ruthveni, and Cryptobatrachus
of a large and diverse set of stakeholders, and representing a global vehicle for mobilizing of California red-legged frogs. Journal of Wildlife Management 67:424-438. boulengeri) as well as for several threatened bird
and coordinating stakeholders to examine, deliberate, and act on advancing the Amphibian Dreitz, V.J. 2006. Issues in Species Recovery: An Example Based on the Wyoming Toad. BioScience species (Pyrrhura viridicata, Myiotheretes pernix,
Conservation Action Plan. 56:765-771. and Campylopterus phainopeplus).
Our research designed to understand the causes and consequences of amphibian declines Dutson, G.C.L., Magsalay, P.M. and Timmins, R.J. 1993. The rediscovery of the Cebu Flowerpecker Dicaeum 5 Morningside is a 1,000-ha area just to the east of
will require inspired thinking, and must lead to explicit practical recommendations for ac- quadricolor, with notes on other forest birds on Cebu, Philippines. Bird Conserv. Intern. 3:235-243. the Sinharaja World heritage Site, and the only
tions. Our actions, in turn, will need to be methodical and exacting. Our failures will need Eken, G., Bennun, L., Brooks, T.M., Darwall, W., Fishpool, L.D.C., Foster, M., Knox, D., Langhammer, P., known site for five highly threatened amphibians:
to motivate us, while our successes will need to be promoted. The hope is that, in the end, Matiku, P., Radford, E., Salaman, P., Sechrest, W., Smith, M. L., Spector, S. & Tordoff, A. 2004. Key Microhyla karunaratnei (CR), Polypedates fastigo
we can succeed in even the most difficult of cases. Biodiversity Areas as Site Conservation Targets. BioScience 54:110-118. (CR), Philautus simba (CR), Philautus procax (CR),
Fishpool, L.D.C. and Evans, M.I. 2001. Important Bird Areas in Africa and associated islands: Priority sites and Philautus decoris (EN).
for conservation. Pisces Publications and BirdLife International, Newbury and Cambridge, UK. 6 Although disease is a major threat factor, the
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124
17 http://www.iucn.org/themes/ssc/publications/
policy/reinte.htm
18 http://ecos.fws.gov/docs/recovery_plans/2002/
020528.pdf
19 Comprehensive reviews of options for invasive
species control and prevention are provided in
McNeely et al. (2001), Wittenberg and Cock (2001),
and Veitch and Clout (2002). IUCN has also prepared
a set of guidelines to help countries, conservation
agencies and concerned individuals to reduce the
threats posed by invasive alien species to global
biodiversity (http://www.iucn.org/themes/ssc/pub-
lications/policy/invasivesEng.htm).
20 Rotenone is a piscicide that has been used to
remove unwanted fish stocks from a variety of
aquatic habitats, although not without negative
impacts on other aquatic fauna and amphibians.
The impacts of rotenone-containing piscicides on
amphibians have been reviewed by Fontenot et al.
(1994) and McCoid and Bettoli (1996).
21 www.nwhc.usgs.gov/research/amph_dc/sop_
mailing.html
22 http://ecos.fws.gov/docs/recovery_plans/2002/
020528.pdf
23 These species are listed on Appendix II of CITES
(see later), which means that international trade
in these species is permissible if sustainably man-
aged. CITES officials have been supportive of this
project because it will alleviate pressure on poison
dart frog populations.
24 http://www.gefonline.org/projectDetails.
cfm?projID=1485
25 http://www.iwcoffice.org/commission/convention.
htm
26 www.cites.org
27 www.ramsar.org
28 http://www.fws.gov/endangered/esa.html
29 The ESA implements US participation in CITES, and
these were rolled together in the legislation when
the ESA was implemented in 1973 (the same year
as CITES was formed). Consequently, there is a link
between ESA and CITES and some species threatened
by trade have also been listed. In the late 1970s, FWS
had a long list of candidate species (about 200 spe-
cies), including many foreign species (many of which
were unrelated to CITES issues). A Fund for Animals
lawsuit pushed for the listing of most or all of the
candidates, and the resultant mass listing resulted
in the listing of many foreign species. As such, most
of the foreign listings occurred in the first decade of
the ESA and there have been very few since. FWS
currently has little interest in listing new foreign
species. Provisions on take do not apply in foreign
jurisdictions and are unenforceable. Further, FWS
has a stated policy that foreign listed species are the
lowest in priority among listed species, and there are
no recovery plans for listed species and no funding.
30 http://whc.unesco.org/en/conventiontext/
31 www.ecolex.org
32 A Party may enter a Reservation only at the time
that it joins the Convention, or within 90 days of a
change being made in the listing of a species on
the Appendices.
33 www.biodiv.org
34 http://www.biodiv.org/doc/publications/pa-text-
en.pdf
35 http://unfccc.int/2860.php
36 Iraq, Somalia and Timor-Leste are among those yet
to ratify
37 http://unfccc.int/kyoto_protocol/items/2830.php
38 Cuts in the three most important gases – carbon
dioxide (CO2), methane (CH4), and nitrous oxide
(N20) – will be measured against a base year of
1990. Cuts in three long-lived industrial gases
– hydrofluorocarbons (HFCs), perfluorocarbons
(PFCs), and sulphur hexafluoride (SF6) – can be
measured against either a 1990 or 1995 baseline.
39 To know how much deforestation has been reduced, it
is necessary to know how much deforestation would
occurred otherwise. This reference level is referred to
as a baseline. While there are a number of ways this
baseline could be selected (e.g. modeled projections,
present deforestation rate), the best baseline for a
variety of reasons is an average deforestation rate
over a recent historical time period.
40 http://www.ipcc.ch/
41 Conversely, listing species that are known to be
genuinely threatened with extinction as Data
Deficient, either because assessors demand
substantial evidence that a species is threatened
before making such a classification, or to side-step
well intentioned but misguided government policies
that restrict field research on threatened species,
could result in species not receiving conservation
attention before it is too late (see Pimenta et al.
2005; Stuart et al. 2005 for discussion).
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 Chapter 11. Amphibian Conservation – Responding to the Global Decline of Amphibians
ESSAY 11.1. THE DECLINING AMPHIBIAN POPULATIONS TASK FORCE: A SHORT HISTORY
The global amphibian community first became aware of the amphibian de-
cline phenomenon at the first World Congress of Herpetology in Canterbury,
UK, in 1989. Conversation outside the formal sessions was dominated by
anecdotal reports, from all over the world, of once-common species that had
disappeared. In the following year, a workshop set up by the US National
Academy of Sciences lead to the formation of the Declining Amphibian
Populations Task Force (DAPTF) in 1991, under the aegis of IUCN’s Species
Survival Commission. From the start, the DAPTF was formerly linked to the
World Congress of Herpetology (WCH).
The first DAPTF Office was established at Oregon State University, Corval-
lis, under the direction of James L. Vial, assisted by Lorelei Saylor. In 1994,
the DAPTF Office moved to the Open University, UK, with Tim Halliday as
International Director and John Baker as International Coordinator; it remained
there until the closure of the DAPTF in June, 2006. During its existence, the
DAPTF has had five Chairs: David B. Wake, Robert Johnson, W. Ronald Heyer,
James Hanken, and James P. Collins. It has had three International Coordina-
tors: John Baker, John Wilkinson, and Jeanne McKay.
At a meeting in 1992, the DAPTF set itself the following goals (Heyer
and Murphy 2005):
1. Catalyse, catalogue and coordinate efforts to gain an understanding of
amphibian population declines.
2. Identify those target populations, species and regions that merit immediate
attention.
3. Gather and critically examine evidence concerning causal factors contribut-
ing to amphibian declines and identify remedial action.
4. Promote data collection on amphibian populations on a long-term basis.
5. Enlist the support of appropriate scientific disciplines needed to address
the issues.
6. Disseminate information on amphibian declines to the scientific community
and promote public awareness.
7. Advise the IUCN, other conservation organizations, and appropriate
governmental bodies on necessary and immediate action.
An important product of the DAPTF has been its regular, free newsletter,
Froglog. This has been used as a means by which the DAPTF Office can
communicate with the herpetological community, and as a medium through
which herpetologists can report the results of their work and air their views
on amphibian declines. As the DAPTF has developed and grown, Froglog has
increased in size and frequency of publication; in 1997, it went from a quarterly
to bi-monthly publication. As of June 2006, 75 issues of Froglog had been
published. Since 2004, Froglog has been published electronically.
The membership of the DAPTF was informal and it is impossible to say
how many people around the world considered themselves to be members.
The circulation of Froglog reached 3,000 when it was produced only in hard
copy, but many of these copies went to groups rather than individuals. The
membership was organized into 90 Regional Working Groups, covering
specific regions (e.g., Southeast Asia), specific countries, or specific regions
within countries. The work of some of these working groups contributed to
the production of major regional reports on amphibians, such as those for
the Lesser Antilles (Kaiser and Henderson 1994), the former Soviet Union
ESSAY 11.2. AMPHIBIAN GLOBAL GAP ANALYSIS: EVALUATING THE EFFECTIVENESS OF THE WORLD’S PROTECTED AREAS
IN REPRESENTING AMPHIBIANS
The in-situ conservation of viable populations in natural ecosystems is
widely recognized as a fundamental requirement for the maintenance of
biodiversity (e.g., Convention on Biological Diversity). Indeed, the practical
value of protected areas in shielding areas of land from destructive use has
been clearly demonstrated (e.g. Bruner et al. 2001). Protected areas have
therefore received wide recognition as core components of conservation
strategies, and their designation is a requirement of several multilateral
environmental agreements (e.g., the Convention on Biological Diversity and
the Ramsar Convention on Wetlands), as well as national and international
legislation (e.g., European Union Birds and Habitats Directives).
In 1992, the Fourth Congress on National Parks and Protected Areas (Ca-
racas, Venezuela) called for protection of at least 10% of each major biome
by the Year 2000 (IUCN 1993), an ambitious target at a time when only an
estimated 3% of the planet’s land area was under protection. In 2003, the Fifth
World Parks Congress (Durban, South Africa) witnessed the announcement
that 11.5% of the Earth’s land surface is now under some form of protection
(Chape et al. 2003). Most Governments have invested in the creation of
protected areas systems, with more than 100,000 being recognized by the
2003 United Nations List of Protected Areas (Chape et al. 2003).
This extraordinary expansion of the global network of protected areas
should be celebrated as a major conservation achievement, as they are
no doubt contributing very significantly for the protection of the world’s
biodiversity. However, this global network is still largely incomplete. A recent
global gap analysis found that at least 12% of the species of birds, mam-
mals, amphibians and turtles analysed were not covered by any protected
area in any part of their ranges (Rodrigues et al. 2004). Amphibians had
the highest proportion of gap species. Here, we update the results of this
2004 global gap analysis for amphibians, drawing on improved species and
protected area data.
A precise evaluation of how effective the world’s protected areas are
in safeguarding amphibians is not possible with current knowledge. On the
one hand, our data on species’ distributions are very coarse: all we have are
generalized maps of extent of occurrence that for most species are likely to
include vast expanses of unsuitable habitat. On the other hand, our data on
protected areas are very limited: the database is likely to be incomplete,
125
(Kuzmin et al. 1995), Canada (Green 1997), Australia (Campbell 1999), was disbanded as an independent organization, with many of its activities
southern Africa (Minter et al. 2004), and the USA (Lannoo 2005). From the being incorporated into the IUCN/SSC Amphibian Specialist Group (ASG)
beginning, the DAPTF recognized the importance of developing protocols for (see Essay 11.12).
monitoring amphibian populations and for disseminating best practice in
population monitoring among its members. An issue-based DAPTF working Tim Halliday
group addressed this issue and produced an important book on the subject
(Heyer et al. 1994).
Since its inception, the DAPTF has allocated a substantial proportion of References
its budget to an annual programme of Seed Grants. These are small awards
(typically $500 to $2000) given to support projects that further the DAPTF’s Campbell, A. (ed.). 1999. Declines and Disappearances of Australian Frogs. Environment
mission, with the intention that they will not only yield results that further Australia, Canberra, Australia.
our scientific understanding of the amphibian decline phenomenon, but will Green, D.M. (ed.). 1997. Amphibians in Decline: Canadian Studies of a Global Problem.
also lead to recipients setting up new lines of research that will attract SSAR, St. Louis, Missouri, USA.
further, more substantial funding from conventional sources (Halliday 2002). Halliday, T. 2002. Getting started with seed grants – the Declining Amphibian Populations
While the greatest number of proposals has been received from the USA, Task Force. Species 38:18.
the DAPTF steadily increased its global outreach year by year. Up until June Heyer, W.R. and Murphy, J.B. 2005. Declining Amphibian Populations Task Force. In: M.
2006, proposals had been received from over 80 different countries, and the Lannoo (ed.), Amphibian Declines. The Conservation Status of United States Species,
DAPTF had funded projects in 49 countries. While most projects have been pp. 17-21. University of California Press, Berkeley, California, USA.
for research, some Seed Grants were awarded to particular Regional Working Kaizer, H. and Henderson, R.W. 1994. The Conservation Status of Lesser Antillean Frogs.
Groups to fund workshops and meetings. In the last few years, the budget for Herpetological Natural History 2:41-56.
Seed Grants was augmented by large donations from other organizations: an Kuzmin, S.L., Dodd, C.K. and Pikulik, M.M. (eds.). 1995. Amphibian Populations in the
anonymous donor (AD), Conservation International (CI), the Critical Ecosystem Commonwealth of Independent States: Current Status and Declines. Pensoft Publica-
Partnership Fund (CEPF), and the Amphibian and Reptile Monitoring Initia- tions, Moscow, Russia.
tive (ARMI). These donors specified the areas in which their funding can be Lannoo, M. (ed.). 2005. Amphibian Declines. The Conservation Status of United States
used: climate change (AD), specific biodiversity hotspots (CI and CEPF), and Species. University of California Press, Berkeley, California, USA.
activities within the USA (ARMI). Minter, L.R., Burger, M., Harrison, J.A., Braack, H.H., Bishop, P.J. and Kloepfer, D. (eds.).
The decline in the number of applications for DAPTF Seed Grants from 2004. Atlas and Red Data Book of the Frogs of South Africa, Lesotho and Swaziland.
2004 onwards (Table 1) is probably a reflection of the fact that the amphibian Smithsonian Institution, Washington D.C, USA.
decline phenomenon has achieved much higher status in the wider academic
community since the DAPTF was established. This has enabled amphibian
researchers in many parts of the world to obtain substantial grants from con- Table 1. DAPTF Seed Grants submitted and awarded by year.
ventional grant-giving bodies. The changing pattern of applications for DAPTF
Seed Grants reflects the DAPTF’s most important achievement: it stimulated
and sustained interest in amphibian declines until such time as the academic Year No. Submitted No. Awarded Value ($$)
community as a whole came to recognize its full importance. The DAPTF 1992 2 1 1,000
Seed Grant program was very successful in achieving its aim of enabling 1993 31 5 4,975
researchers to initiate innovative lines of research into amphibian declines 1994 1 0 0
and their causes. As of June 2006, the number of papers in refereed journals 1995 38 9 15,517
that acknowledged the DAPTF as a source of funding exceeded 100. 1996 3 2 3,000
In addition to its Seed Grant programme, the DAPTF maintained a small 1997 49 11 22,921
Rapid Response Fund, recognizing that some aspects of amphibian declines, 1998 53 15 27,550
notably mass mortality events, require more immediate action than can be 1999 45 14 23,828
provided by an annual grant program. Between 1998 and 2006, the DAPTF 2000/1 47 21 40,283
received 17 bids to this fund, and funded eight of them. 2002 57 37 58,217
The development of the DAPTF coincided with the development of the 2003 63 19 32,537
internet, enormously facilitating the capacity of organizations like the DAPTF 2004 65 23 43,123
to communicate and disseminate information globally. The DAPTF launched 2005 48 14 25,450
its own web site in 2004 and is now linked to numerous web sites, such 2006 32 12 23,133
as AmphibiaWeb (www.amphibiaweb.org), which cover various aspects Totals 534 183 $321,534
of amphibian biology, status and conservation. In June 2006, the DAPTF
but also likely to include many areas in which amphibian species are poorly species are the dominant Red List category amongst gap species (Figure 1).
protected, or not protected at all. It is therefore impossible to know exactly Less-known species tend to have poorly known distributions, so many of these
which species are adequately conserved in which protected areas. We used species may be subsequently found to occur in protected areas. However, lack
a diversity of approaches to obtain a range of estimate of the degree to which of knowledge is often associated with rarity, and thus many of these species
amphibian species are covered by the global network of protected areas. are likely to have truly small populations and to be highly threatened.
The first three approaches employed (Table 1) investigated the spatial Previous studies found that levels of coverage for amphibians are
overlap between the amphibian distribution maps compiled through the substantially lower than for other vertebrate taxa (Rodrigues et al. 2004).
Global Amphibian Assessment and the 2006 World Database on Protected This is likely a reflection of their much smaller range sizes, as widespread
Areas (WDPA 2006). The WDPA includes about 57,500 protected areas species tend to be covered in protected areas even if by chance alone. Other
mapped as polygons and about 118,000 protected areas mapped as points. explanations include habitat preference – amphibians tend to be associated
Amongst the latter, the 80,500 for which area information was available with freshwater habitats, which are not particularly well addressed by the
were converted to circles, while the remaining protected areas were simply terrestrial network of protected areas – and taxonomic bias – few protected
mapped as points. areas were created with any consideration of amphibian distributions. By
In the most simple of the gap analysis approaches (Gap 1), we looked mapping the gap species found according to each approach it is possible to
for species whose ranges do not overlap any of the protected areas in the obtain a map of the regions where the global protected area network is most
WDPA. This approach considers a species to be ‘covered’ if any part of its incomplete in representing amphibian species (Figure 2).
mapped range overlaps any protected area; otherwise, it is a ‘gap species’. Overall, the areas identified as having higher concentrations of gap species
Given the data limitations explained above, this is very likely to be a crude
overestimate of the overall species coverage. Trying to focus on protected
areas more likely to effectively protect species, in our second approach
(Gap 2) we narrowed down the definition of protected area to include only
those classified by IUCN as ‘strictly protected’, that is, under categories I to Gap analysis approach All species Threatened
VI (IUCN 1994). The third approach (Gap 3) considered only protected areas species
larger than 1,000 ha (thus excluding also all of those mapped as points) which Gap 1 (All protected areas in the WDPA) 815 (14.1%) 356 (19.7%)
are classified under categories I to IV (Table 1). The fourth and final approach Gap 2 (IUCN categories I to IV) 1,390 (24.0%) 599 (33.1%)
(Gap 4) did not use the WDPA data. Instead, it used information provided Gap 3 (IUCN categories I to IV AND 1,473 (25.4%) 635 (35.1%)
by the Global Amphibian Assessment experts on whether or not a species >1,000 ha)
was known to occur in protected areas. A species was considered covered Gap 4 (GAA experts) 1,669 (28.8%) 618 (34.1%)
if protected areas were considered to be a conservation measure in place, Total number of species 5,794 1,810
and a gap species otherwise.
The fraction of species identified as gap species varied between 14.1%
and 28.8% of all amphibians, and between 19.7% and 35.1% of threatened Table 1. Number and percentage of gap species under each approach, for
species (Table 1). The strong bias towards threatened species is again evident all species and for threatened species. The total number of species refers to
from Figure 1, which also demonstrates that species of higher levels of threat those for which distribution maps were compiled in the Global Amphibian
(Critically Endangered) are particularly likely to be gap species. Data Deficient Assessment (i.e., it excludes 124 species of unknown distributions).
126 Threatened Amphibians of the World

fall overwhelmingly in the tropics, especially in tropical and subtropical moist
forests. They are also disproportionately located on islands, and on regions of
high topographic complexity (tropical mountains). In the Western Hemisphere,
the highest concentrations of gap species are found in the tropical Andes
and in the Sierra Madre mountains in Central America. The Atlantic Forest
and the Caribbean are also highlighted. Although the Guayana Shield and
the Amazon are noted centres of endemism (see Chapter 4), the general lack
of gaps there reflects their extensive networks of protected areas and the
relatively wide ranges of most of their species.
For Africa, gap species tend to concentrate almost exclusively in the
mountains, including the Cameroonian highlands, the Eastern Arc Mountains,
the Albertine rift, the Ethiopian Highlands, and eastern Madagascar. Most
gap species found in these regions of high endemism (see Chapter 4) have
very restricted ranges. Concentrations of gap species in Somalia, on the other
100
80
% of species in each Red List Category
hand, reflect this region’s extremely poor protected area coverage.
In Asia, the main concentrations of gap species are found in the Western
Ghats and Sri Lanka, but also in Himalayan slopes, southern China, and Japan,
particularly the Nansei-Shoto (or Ryukyu) Islands. Further east, New Guinea
and the Solomon Islands emerge as major priorities for the establishment
of new protected areas. All of these are areas of very high endemism for
amphibians (see Chapter 4).
Nearly all of the regions identified here as having major gaps in protected
area coverage of amphibian species are located in low-income countries in the
tropics – those that can least afford the costs of establishing and enforcing
protected areas (James et al. 1999). This is the case even if the significant
local benefits of protected areas are factored in (Balmford et al. 2003), because
much of the benefit of the establishment of protected areas is realized at a
global scale (Kremen et al. 2000). Thus, our recommendation for the rapid
Figure 1. Distribution of Red List categories amongst all species of amphib-
ians and amongst those identified as gap species. Red List categories: CR
– Critically Endangered; EN – Endangered; VU – Vulnerable; NT – Near Threat-
ened; LC – Least Concern; DD – Data Deficient; EW – Extinct in the Wild.
establishment of protected areas in these regions goes hand-in-hand with a
recommendation that the costs of this conservation are largely borne by the
global community. Donor country governments, through bilateral and multi-
lateral institutions, as well as NGOs, foundations, and private corporations
and individuals all have an important role to play in financing conservation
(Balmford and Whitten 2003).
Ana S.L. Rodrigues
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terrestrial conservation costs, conservation benefits, and unmet conservation needs.
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P.A., Pilgrim, J.D., Pressey, R.L., Schipper, J., Sechrest, W., Stuart, S.N., Underhill, L.G.,
Figure 2. Number of amphibian gap species per equal-area (3,113km2) Waller, R.W., Watts, M.E.J. and Yan, X. 2004. Effectiveness of the global protected
EW CR EN VU NT LC DD
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ESSAY 11.3. IDENTIFYING AND SAFEGUARDING KEY BIODIVERSITY AREAS FOR AMPHIBIAN CONSERVATION

The overwhelming threat to biodiversity worldwide is the loss of native

habitat (Baillie et al. 2004). Nearly 90% of globally threatened amphibians are
being negatively impacted by habitat loss. Thus, conserving habitat is critical
for the survival of amphibian species. One of the most effective means of
targeting habitat conservation efforts is through the identification of globally
significant sites for biodiversity conservation, termed Key Biodiversity Areas
(KBAs; Eken et al. 2004).
KBAs are identified using a set of globally standard criteria and thresholds
derived from those developed by BirdLife International for the identification
of Important Bird Areas (IBAs; e.g., Fishpool and Evans 2001). The KBA
criteria align with the two principal measures of systematic conservation

Figure 1. Key Biodiversity Areas identified for amphibian species in the

Philippines, indicating the single Alliance for Zero Extinction site (South
Gigante Island. Numbers on the map correspond as follows:
1 - Southwestern Negros; 2 - Northern Negros Natural Park; 3 - Bataan Natural
Park and Subic Bay Forest Reserve; 4 - Buguey Wetlands; 5 - Kalbario-Patapat
National Park; 6 - Pagbilao and Tayabas Bay; 7 - Timpoong and Hibok-hibok
Natural Monument; 8 - Dumaran-Araceli; 9 - Balabac Island; 10 - Puerto
Galera; 11 - Mt. Makiling Forest Reserve; 12 - Balbalasang-Balbalan National
Park; 13 - Apayao Lowland Forest; 14 - North Central Sierra Madre Mountains;
15 - Angat Watershed Forest Reserve; 16 - Mts. Irid-Angilo and Binuang; 17
- Mt. Kulasi; 18 - Malpalon; 19 - Busuanga Island; 20 - Ban-ban; 21 - Biliran
and Maripipi Island; 22 - Anonang-Lobi Range; 23 - Mt. Nacolod; 24 - Mt.
Diwata Range; 25 - Bislig; 26 - Mt. Agtuuganon and Mt. Pasian; 27 - Mt.
Kampalili-Puting Bato; 28 - Mt. Kaluayan-Mt. Kinabalian Complex; 29 - Mt.
Tago Range; 30 - Siburan; 31 - Balogo Watershed; 32 - Mt. Mantalingajan; 33
- South and North Gigante Island; 34 - Mt. Capayas; 35 - Mt. Dajo National
Park; 36 - Victoria and Anepahan Ranges; 37 - Mt. Dingalan; 38 - Aurora
Memorial National Park; 39 - Zambales Mountains; 40 - Mt. Bandila-an; 41
- Mt. Dapiak-Mt. Paraya; 42 - Mt. Busa-Kiamba; 43 - Mt. Sinaka; 44 - Mt.
Hilong-hilong; 45 - Mt. Kambinlio and Mt. Redondo; 46 - Polillo Island; 47
- Marinduque Wildlife Sanctuary; 48 - Central Cebu Protected Landscape;
49 - Rajah Sikatuna Protected Landscape; 50 - San Vicente-Taytay-Roxas
Forests; 51 - Central Panay Mountains; 52 - Mt. Halcon; 53 - Northwest
Panay Peninsula Natural Park; 54 - Mt. Canlaon Natural Park; 55 - Pasonanca
Natural Park; 56 - Basilan Natural Biotic Area; 57 - Mt. Malindang Natural
Park; 58 - Mt. Kitanglad Range Natural Park; 59 - Mt. Matutum Protected
Landscape; 60 - Malampaya Sound Protected Landscape and Seascape; 61
- El Nido Managed Resource Protected Area; 62 - Mt. Guiting-guiting Natural
Park; 63 - Mt. Calavite Wildlife Sanctuary; 64 - Quirino Protected Landscape;
65 - Mt. Kalatungan Range Natural Park; 66 - Mt. Isarog Natural Park; 67
- Casecnan Protected Landscape; 68 - Mt. Apo Natural Park; 69 - Cuernos
de Negros; 70 - Agusan Marsh Wildlife Sanctuary; 71 - Mts. Banahaw-San
Cristobal Protected Landscape; 72 - Samar Island Natural Park; 73 - Mt.
Pulag National Park; 74 - North Eastern Cagayan Protected Landscape
and Seascape; 75 - Northern Sierra Madre Natural Park; 76 - Penablanca
Protected Landscape and Seascape; 77 - Calauit Island; 78 - Culion Island;
79 - Romblon Island
 Chapter 11. Amphibian Conservation – Responding to the Global Decline of Amphibians
planning, vulnerability and irreplaceability (Margules and Pressey 2000),
and can be applied to all taxonomic groups. Sites that hold one or more
globally threatened species or that have globally significant populations of
a geographically concentrated species qualify. Geographically concentrated
species include those that have a limited global range size (provisionally set
at 50,000km²), termed restricted-range species, as well as globally significant
congregations of species.
Boundaries for KBAs are drawn to yield sites that can be managed for
conservation, using data on management units, species’ ecological require-
ments, the extent of remaining habitat, topographical features, and human
settlement patterns. In identifying and delineating KBAs for amphibians,
information from the Global Amphibian Assessment (GAA), including range
polygons, population estimates, and information on threats, conservation
measures, and habitat requirements of species, can be used in combination
with more specific locality data to help identify globally important sites.
The identification of the set of KBAs within a region is typically driven
from the national level. A good example of a country where this process is
far advanced is the Philippines. With more than 20,000 endemic species, the
Philippines is one of the world’s 17 “megadiversity” countries (Mittermeier et
al. 1997), but with less than 7% of its original forest cover remaining intact, it
is also one of 34 global biodiversity hotspots (Mittermeier et al. 2004). As such,
accurately targeting scarce conservation resources is particularly important.
Several priority-setting processes have contributed to this goal, including the
identification of 117 IBAs by BirdLife International and the Haribon Foundation
in 2001 and of 206 conservation priority areas by the Philippine Biodiversity
Conservation Priority-setting Program (PBCPP) in 2002. In 2004, Conservation
International-Philippines, in collaboration with the Haribon Foundation and
the Protected Areas and Wildlife Bureau of the Department of Environment
and Natural Resources, built on this earlier work to identify KBAs for multiple
taxonomic groups, using a systematic, data-driven process. A number of other
partner NGOs, government agencies, research institutions, and individual
experts were involved in contributing data and peer review.
A total of 128 KBAs were identified for threatened and endemic amphib-
ians, mammals, birds, reptiles, and freshwater fish, with 79 sites triggered
by 49 amphibian species (Figure 1). This total includes one Alliance for Zero
ESSAY 11.4. PREDICTING THE DISTRIBUTION AND SPREAD OF PATHOGENS TO AMPHIBIANS
Knowledge of the distribution of species is essential in order to conserve biodi-
versity. Conventional methods for inferring and monitoring changes in species
distributions require labour- and time-intensive efforts and are realistically
practical only for selected taxa and small geographic regions. Nevertheless,
as human-induced pressures on biological diversity and extinctions increase
worldwide, accurate predictions of species distributions are in increasingly
urgent demand by both scientific and conservation communities.
Modelling ecological niches to predict the geographic distribution of
species has become an indispensable tool and has been used to study a
wide variety of processes in ecology, evolution, and conservation (Graham
et al. 2004). This suite of modelling techniques use complex mathematical
algorithms to quantify the environmental space, or envelope, of species by
determining the association between occurrence records and a series of
environmental or biotic niche dimensions (e.g., temperature, precipitation,
vegetation; see Figure 1). This relationship is then used to estimate the
probability of occurrence of the target species in regions where it has not
yet been recorded, and to estimate the geographic extent of suitable habitat.
These areas are “analogous ecologically” to areas where the species has
been reported to occur. As such, they are a function of the species’ ecological
niche or environmental requirements, at least along the niche dimensions
considered (Soberón and Peterson 2005).
More recently, ecological niche modelling has been used in epidemiologi-
cal studies to predict the distributions of pathogens, pests and diseases. For
example, areas of Chagas disease, Ebola haemorrhagic fever, and Sudden Oak
Death outbreaks have been successfully predicted (Peterson et al. 2002, 2004;
Guo et al. 2005). These studies also yielded a detailed understanding of dis-
ease occurrence and risk areas, parasite-host distributional relationships, and
ecological factors contributing to outbreaks. These and other examples reveal
the applicability of ecological niche modelling to better inform programmes
aimed at managing wildlife infectious diseases, particularly in regions where
species survival has been compromised by the arrival of emerging diseases.
Nevertheless, the application of ecological niche modelling on studies of
amphibians’ pathogens has been surprisingly limited.
Emerging infectious diseases have been increasingly identified across
the globe as a threat to wildlife and human welfare (Daszak et al. 2000). An
emerging disease generally has two characteristics: (1) its geographical range
or host range is expanding, or its prevalence has been increasing in recent
years, and (2) these changes are frequently driven by some form of large-
scale anthropogenic environmental change, such as climate anomalies or
deforestation (Daszak et al. 2004). Given the link between disease occurrence
and environmental conditions, ecological niche modelling should prove to be
a valuable tool for examining the dynamics of emerging diseases.
Studies have shown that habitat degradation and climate change can
affect the life cycle of pathogens and can, therefore, influence the patterns
of disease outbreak and transmission. As a consequence, modelling the
distribution of pathogens under different scenarios of future climate change,
habitat loss, and fragmentation can be an important predictive tool to facilitate
disease control. Identifying areas where disease is likely to spread as a result
of climate change and habitat degradation should allow prioritization of
monitoring efforts into high risk and presently uninfected areas. Predictions
of both current and future distributions should also be useful for management
of both pathogens and infected species.
Amphibians may provide a dramatic example of the devastating impact of
emerging diseases on wildlife. Over the past few years, a number of studies
Extinction site (see main text) identified for an amphibian species (the En-
dangered island forest frog, Platymantis insulata, which is entirely restricted
to the tiny island of South Gigante).
The KBA process sometimes focuses on a regional level and involves
multi-country cooperation. For example, in 2002, Nature Kenya and the
Wildlife Conservation Society of Tanzania, with the support of the Critical
Ecosystem Partnership Fund (CEPF), collaborated to identify globally important
sites for conservation within the Eastern Arc Mountains and Coastal Forests
of Tanzania and Kenya biodiversity hotspot (sensu Myers et al. 2000). Building
upon the successful identification of IBAs in the region, a total of 160 KBAs
were identified across five taxonomic groups (plants, gastropods, birds, mam-
mals, and amphibians). A total of 20 KBAs were identified for 44 amphibian
species. Although KBAs continue to be refined as additional species and
habitat data are obtained, the process of communicating these conservation
targets to partners is well underway. In East Africa, in addition to providing
a geographically explicit strategy for US$7 million of CEPF investment in the
hotspot, these sites have also been adopted by a number of conservation
partners in Tanzania and Kenya. Most importantly, they serve as the basis
for priority sites in the $62.2 million Eastern Arc Mountains Endowment
Fund, supported by the Global Environment Facility (GEF). The identification
of KBAs, therefore, clearly serves as a vital tool in focusing and leveraging
funding for conservation.
Once identified, safeguarding KBAs can take many forms, ranging from
formal governmental protection and management, to strengthened manage-
ment at existing sites, to implementing community-based conservation, to
strengthening and monitoring the success of existing projects on the ground.
One recent example from Colombia concerns the proclamation of the El
Dorado Nature Reserve, contiguous with the Parque Nacional Natural Sierra
Nevada de Santa Marta, which is the only known site for seven threatened
amphibians, including the Critically Endangered Santa Marta Harlequin Frog
Atelopus laetissimus and San Lorenzo Harlequin Frog Atelopus nahumae.
Upon learning of the impending sale of plots from the site for the construc-
tion of vacation homes, three conservation organizations, Fundación ProAves,
the American Bird Conservancy, and Conservation International, stepped in
to protect this 1,600-acre site on the north-west slope of the Sierra Nevada
have implicated fungal, viral, bacterial, and parasitic diseases in popula-
tion declines, and even extinctions, of amphibians (Daszak et al. 2003). Of
particular relevance is chytridiomycosis, a disease caused by a pathogenic
fungus that may be related to mass deaths and severe declines in amphibian
populations around the Earth (Longcore et al. 1999; Daszak et al. 2003).
Niche modelling and disease in amphibians:
an illustrative example
The first application of niche modelling as a tool to characterize the environ-
mental envelope and potential distribution of an amphibian pathogen was
INPUT DATA
1. Species occurrence records
ENM
3. Niche model in ecological dimensions
Annual Precipitation
Geographic projection
Annual Mean Temperature
n
tio
va
Ele
127
de Santa Marta massif.
In summary, site conservation is clearly critical in ensuring the long-term
persistence of amphibian species, and in many cases complements and
enhances the effectiveness of other important conservation techniques, such
as disease mitigation, invasive species eradication, and captive breeding.
However, effective conservation action is only possible if we know where such
sites are – and the KBA approach provides the tools necessary to highlight
sites of global conservation significance for amphibians.
Naamal De Silva, Ruth Grace Ambal, Melizar V. Duya,
Matt Foster, Kim Howell, David Knox, Penny Langhammer,
Robinson Mugo and Alex Ngari
References
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IUCN, Gland, Switzerland.
Eken, G., Bennun, L., Brooks, T.M., Darwell, W., Fishpool, L.D.C., Foster, M., Knox, D.,
Langhammer, P., Matiku, P., Radford, E., Salaman, P., Sechrest, W., Smith, M.L.,
Spector, S. and Tordoff, A. 2004. Key Biodiversity Areas as Site Conservation Targets.
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Fishpool, L.D.C. and Evans, M.I. 2001. Important Bird Areas in Africa and associated
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Figure 1. Ecological niche modeling (ENM). Predictions of niche occurrence
are based on: (1) Species geographic distribution in the form of primary point
occurrence data, and (2) Environmental variables. Modeling (3) consists on
finding the association between species occurrence records and environ-
mental or ecological variables. The associations find “resemblances” on the
environmental layers between occurrence records and other regions of the
map. The output is a prediction of the distribution of regions suitable for the
occurrence of the target species (4). The prediction represents the ecological
niche projected back onto geography. Most of the modeling methods yield
maps showing varying probabilities of the presence of suitable conditions
across the region.
2. Ecological variables
Annual Mean Temperature
Vegetation
Annual Precipitation
Minimum Temperature of Coldest Period
Elevation
OUTPUT
4. Distributional prediction
20-40
40-60
20-40
40-60
128 Threatened Amphibians of the World

Figure 2. Predicted distribution of the fundamental niche of (A) the chytrid

fungus Batrachochytrium dendrobatidis, and (B) the Jambato toad, Atelopus A. Chytrid distribution model C. Overlap of distribution areas
ignescens, in Ecuador, South America. Darker yellow and red tones indicate a
higher probability of occurrence. (C) shows areas of overlap between both spe-
cies (gray areas). Red and yellow tones show areas of no overlap. Models were
built from nine chytrid and 76 A. ignescens localities of known occurrence
using MAXENT (Phillips et al. 2004). A 0.5 threshold was used to transform
the probability data to presence/absence data for the chytrid fungus model.
The predicted areas for the pathogen and the host overlap widely.

published by Ron (2005). Niche modelling was used to predict the distribution
of the chytrid fungus Batrachochytrium dendrobatidis, in the New World. The
0-20
predictions indicate that the regions of highest suitability for the chytrid are 20-40
also the areas with the most diverse amphibian faunas worldwide. This study 40-60
60-80
makes recommendations for management based on the fact that regions 80-100
suitable for the establishment of the pathogen are widespread and that the
pathogen has been regularly found in the pet trade. Specifically, Ron (2005)
suggests the implementation of severe trade regulations for live amphibians B. Atelopus ignescens distribution model
to avoid anthropogenic spread of the pathogen.
The predicted occurrence of the chytrid fungus under a wide variety of
environmental regimes and habitat types contrasts with the fact that most
records for the chytrid fungus in the West Indies, Central, and South America
are restricted to a somewhat limited portion of the predicted environmental
niche (e.g., from montane forests and paramos). This asymmetry probably
results from the inclusion of North American records in the process of model
building, and highlights potential biases in the application of environmental
niche modelling when the underlying locality data is not a representative
sample of the pathogen’s environmental niche. Ideally, the input of the model
should consist of positive localities found from uniform sampling throughout
the distribution range. In reality, the distribution of positive records is
biased by accessibility to field sites, availability of museum collections and
0-20
economic resources, and even by scientists’ idiosyncrasies (e.g., a priori 20-40
perceptions of where the pathogen should occur). Other sources of model 40-60
60-80
error are false positive localities and the exclusion of niche dimensions 80-100
that are relevant to limit the pathogen niche. Comprehensible reviews of
these sources of error are provided by Guisan and Zimmerman (2000) and
Guisan and Thuiller (2005).
and spread would also help to develop and test hypotheses on disease of the Jambato Toad Atelopus ignescens (Anura: Bufonidae) in the Andes of Ecuador.
Predicting pathogen and host overlap dynamics that will improve our capacity to design effective conservation Journal of Herpetology 37:116-126.
programmes for amphibians. Soberón, J. and Peterson, A.T. 2005. Interpretation of models of fundamental ecological
To exemplify the applicability of ecological niche modelling in the study of niches and species’ distributional areas. Biodiversity Informatics 2:1-10.
amphibian declines, we modelled the distribution of both the chytrid fungus Pablo A. Menéndez-Guerrero, Santiago R. Ron
and the Jambato Toad, Atelopus ignescens (CR), in Ecuador. The Jambato and Catherine H. Graham
Toad was once widespread and common in the northern Andes and repre-
sents a well-documented case of the disappearance of an amphibian in the
Neotropics during the late 1980s. According to our predictions, the geographic References Taxa
distribution of chytrid in Ecuador overlaps widely with that of the Jambato Chytrid model
Toad (Figure 2). The environmental envelope analysis shows that both have Daszak, P., Cunningham, A.A. and Hyatt, A.D. 2000. Emerging infectious diseases of wildlife Primarily mean diurnal temperature range Ecuadorian amphibian records
2.5 Chytrid records
very similar environmental requirements (i.e., low temperatures during the - threats to biodiversity and human health. Science 287:443-449. Jambato toad records
coldest and driest seasons; Figure 3). These results suggest an ecological Daszak, P., Cunningham, A.A. and Hyatt, A.D. 2003. Infectious disease and amphibian
high

and geographical association between both species that is consistent with population declines. Diversity and Distributions 9:141-150.
PC II

hypotheses implicating the chytrid fungus in the extinction of the Jambato Daszak, P., Tabor, G.M., Kilpatrick, A.M., Epstein, J. and Plowright, R. 2004. Conservation 0.0
Toad and other Andean amphibians (Ron et al. 2003; Pounds et al. 2006). More Medicine and a new agenda for emerging diseases. Annals of the New York Academy
low

detailed studies to explore the overlap between the niche space of threatened
species and the chytrid fungus would be valuable to help clarify the role of
this pathogen in widespread amphibian population declines.
Newly available information
Several new types of information should be used to improve our understand-
ing of the geographic and ecological distribution of the chytrid fungus and
other pathogens of amphibians, as well as to predict how these distributions
will change as a result of anthropogenic activities. The past several years
have witnessed a tremendous increase in the availability of information that
includes extensive databases on species occurrence (hosts and pathogens),
high-resolution spatial data from remote-sensing platforms, comprehensive
treatments of species phylogenies, and new insights on pathogen environ-
mental physiology. Species occurrence records can be used to evaluate the
spatial overlap of amphibian species and pathogens to assess extinction
risks in the context of their respective evolutionary histories. Fine-grained
remotely sensed information and faster computer processors should allow us
to build more precise and refined predictions of amphibian disease outbreaks
using ecological niche modelling. At present, the temporal resolution of
climate maps is 30 years, which is too gross to predict disease driven by
demographic processes. Finally, manipulative experimentation has resulted
in new information about the physiology of chytrid fungus that can also be
used to refine distribution models.
We expect that the combination of these diverse sources of information
in the framework of ecological niche modelling will contribute significantly
to understand disease epidemiology. New insights on disease distribution
of Science 1026:1-11.
Guisan, A. and Thuiller, W. 2005. Predicting species distribution: offering more than simple
habitat models. Ecology Letters 8:993-1009.
Guisan, A. and Zimmermann, N.E. 2000. Predictive habitat distribution models in ecology.
Ecological Modelling 135:147-186.
Graham, C.H., Ferrier, S., Huettman, F., Moritz, C. and Peterson, A.T. 2004. New develop-
ments in museum-based informatics and application in biodiversity analysis. Trends
in Ecology and Evolution 19:497-503.
Guo, Q., Kelly, M. and Graham, C.H. 2005. Support vector machines for predicting distribu-
tion of Sudden Oak Death in California. Ecological Modeling 182:75-90.
Longcore, J.E., Pessier, A.P. and Nichols, D.K. 1999. Batrachochytrium dendrobatidis gen.
et sp. nov., a chytrid pathogenic to amphibians. Mycologia 91:219-227.
Phillips, S.J. Dudík, M., and Schapire R. E. 2004. A maximum entropy approach to species
distribution modeling. Proceedings of the Twenty-First International Conference on
Machine Learning, pp. 655-662.
Pounds, A.J., Bustamante, M.R., Coloma, L.A., Consuegra, J.A., Fogden, M.P.L., Foster, P.N.,
La Marca, E., Masters, K.L., Merino-Viteri, A., Puschendorf, R., Ron, S.R., Sanchez-
Azofeifa, G.A., Still, C.J. and Young, B.E. 2006. Widespread amphibian extinctions
from epidemic disease driven by global warming. Nature 439:161-167.
Peterson, A.T., Bauer, J.T. and Mills, J.N. 2004. Ecologic and geographic distribution of
filovirus disease. Emerging Infectious Diseases 10:40-47.
Peterson, A.T., Sanchez-Cordero, V., Beard, C.B. and Ramsey, J.M. 2002. Ecologic niche
modelling and potential reservoirs for Chagas disease, Mexico. Emerging Infectious
Diseases 8:662-667.
Ron, S.R. 2005. Predicting the distribution of the amphibian pathogen Batrachochytrium
dendrobatidis in the New World. Biotropica 37:209-221.
Ron, S.R., Duellman, W.E., Coloma, L.A. and Bustamante, M.R. 2003. Population decline
-2.5
-2.0 0.0 2.0 4.0
PC I
low high
Primarily temperatures during coldest and driest seasons
Figure 3. Axes I and II from principal components analysis based on 19
environmental variables in Ecuador. Gray circles: 2000 random locations within
the Chytrid fungus predicted niche distribution; green circles: 296 localities of
known occurrence of Ecuadorian endemic species of amphibians in the genera
Colostethus and Atelopus; blue squares: 9 localities of known occurrence of
Chytrid fungus Batrachochytrium dendrobatidis; red circles: 74 localities of
known occurrence of Jambato Toad Atelopus ignescens. So far, all examined
specimens of A. ignescens for chytridiomycosis have tested negative (Ron et
al. 2003). However, the overlap between chytrid fungus model and Jambato
toad’s occurrences in the environmental space is evident. Locality data is
from specimens deposited in Field Museum of Natural History, Museo de
Zoología Pontificia Universidad Católica del Ecuador, Museo de Historia
Natural Gustavo Orcés, Museum of Comparative Zoology Harvard University,
Museum of Zoology University of Michigan, Museum of Vertebrate Zoology
University of California, Natural History Museum University of Kansas, and
Natural History Museum of Los Angeles County.

ESSAY 11.5. ON THE ROLE OF EX-SITU MANAGEMENT IN THE CONSERVATION OF AMPHIBIANS

The best place to conserve wildlife is in the wild. However, in-situ threats
sometimes cannot be mitigated quickly enough to prevent the extinction of
a species in the immediate future, with the result that ex-situ intervention
then becomes the only option available. However, given the magnitude of the
problem versus the currently limited resources of the global ex-situ community,
there simply is no room for most species in need of ex-situ management. For
example, we estimate that the global zoo community can currently manage
viable populations of ~50 amphibian species, which amounts to perhaps
10% of those requiring ex-situ intervention. Much ex-situ space exists in
the private sector, but very little is currently utilized for conservation, with a
few notable exceptions. Other limitations of ex-situ programmes are expense
and risk of disease transmission, inbreeding and artificial selection. Ex-situ
programmes must be placed in the broader context of integrated recovery
efforts: they are one component of the global conservation response, one
that cannot stand alone, and one whose success still ultimately relies upon
mitigation of the in-situ threats.
We see five conservation roles for ex-situ populations, balancing their
potential contributions with their inherent limitations, all of which directly
or indirectly benefit wild populations:
• management for release to augment or restore wild populations
• conservation research
 Chapter 11. Amphibian Conservation – Responding to the Global Decline of Amphibians
• conservation education
• commercial production to relieve in-situ collecting pressure
• fund-raising to support any of the first four activities and/or directly support
in-situ conservation
These roles are not mutually exclusive, and with careful planning, most
ex-situ populations can serve and benefit from multiple roles. Although these
roles have been dealt with in larger monographs (e.g., Griffiths and Kuzmin
2006, Marantelli et al. unpubl.), management for release is a task unique to
the ex-situ community and will therefore be the focus of this essay. Here,
we submit case studies in which programmes implemented by the ex-situ
community have helped prevent the extinction of a threatened species. We
present these case studies as models show-casing a range of options for
future action.
Translocations with time in ex-situ programme
limited to that required for transport
In circumstances where threats have been identified and mitigated or are
spatially limited, and where populations still exist that are demographically
robust enough to sustain harvesting, the simple translocation of animals to
threat-free locations may lead to the re-establishment of populations previ-
ously lost. Such programmes can be achieved cheaply and with minimum risk.
Successful re-establishment of populations has been achieved by this method
for the Natterjack Toad Bufo calamita (LC; Denton et al. 1997). Translocation
of tadpoles from one wild site to another has established new populations
of the Mallorcan Midwife Toad Alytes muletensis (VU; Buley and Gonzalez
2000; and see Essay 11.6) where previously more risky and expensive ex-situ
breeding programmes were employed.
Translocations/reintroductions with time in
ex-situ programme limited to that required for
head-starting
Where threats are mitigated, but extant populations cannot sustain the harvest-
ing levels required to establish new populations through translocation, or where
reduction of mortality can help achieve a positive rate of population increase,
head-starting programmes can be an effective and economical solution. Such
projects utilize naturally produced individuals – usually eggs – and protectively
rear them through periods of otherwise naturally high mortality. This usually
results in the production of large numbers of individuals that would otherwise
ESSAY 11.6. A CONSERVATION SUCCESS STORY: THE MALLORCAN MIDWIFE TOAD ALYTES MULETENSIS
The midwife toad of the Balearic island of Mallorca, Alytes muletensis, gains
its name from the unusual behaviour of the males, which carry fertilized eggs
around their back legs until they hatch; upon hatching the emerging tadpoles
are released into a pool of water to complete their development. The local
Catalan name of ferreret means “little iron-worker” because their ‘chinking’
call sounds like iron being hit with a hammer.
Until the late 1970s, the Mallorcan Midwife Toad was known only from
fossil evidence and was thought to be extinct. However, re-examination of
a specimen originally collected in 1978 revealed that this was not, as first
thought, an introduced common midwife toad (Alytes obstetricans), but an
example of the Mallorcan Midwife Toad (Mayol and Alcover 1981). Subse-
quent surveys revealed the global distribution of the toad to consist of about
13 populations, all contained within the Serra de Tramuntana. Within these
mountains, torrents of water flow through steep-sided karstic gorges during
have perished, without the cost or risk of a captive-breeding programme. A
number of projects have used head-starting to some effect (Sredl et al. 2002;
Gibson and Freeman 1997; Sredl and Healy 1999; Krofta 2003).
Translocations/reintroductions where ex-situ
management is required until in-situ threats
can be mitigated
Often threats are not immediately controllable or even identified and, in
such cases, species may require ex-situ management across generations. In
cases where threats are known to be temporally limited or able to be quickly
mitigated, the ex-situ programme may be short or comprise only a generation
or two. It is always desirable to reduce ex-situ time to minimize risks and
maximize the value of the limited available resources. Captive breeding and
reintroduction of frogs have been used to relocate and re-establish popula-
tions of Romer’s Treefrog (Chirixalus romeri, EN) following the loss of their
habitat to development (Banks 1996; Dudgeon and Lau 1999). Over 100,000
zoo-produced tadpoles of the Puerto Rican Crested Toad (Bufo lemur, CR)
have been released, over the past decade, to augment small wild populations
and restore extirpated populations in areas where breeding habitat has been
restored (Johnson 1999). In some cases the duration of ex-situ management is
indeterminate. Such programmes are resource-intensive and face additional
risks, including genetic loss, artificial selection, and the prospect that the spe-
cies may never be repatriated. For the Kihansi Spray Toad (Nectophrynoides
asperginis, CR), the ex-situ population has circumvented complete extinction,
but there is currently no way to mitigate the in-situ threats (Krajick 2006).
Project Golden Frog (PGF; Zippel 2002) and a collaborative multi-species ef-
fort in Panama by the Atlanta Botanical Garden and Zoo Atlanta (Mendelson
and Rabb 2006) are other examples where the ex-situ component of the
programme will be of indeterminate duration. PGF focused its efforts for the
Panamanian Golden Frog (Atelopus zeteki, CR) along three main objectives:
population and habitat assessments, an intensively managed ex-situ breeding
programme, and range-country education initiatives. Through the breeding
programme, hundreds of offspring have been produced and distributed among
dozens of AZA-accredited zoos. In the US, the frog became a symbol of the
impacts of emerging diseases), with an immediate response from educators
and funding from granting agencies. The Atlanta-based initiative attempted
a pre-emptive extraction of other Panamanian species at an ecosystem
scale. Based on data made available by Lips et al. (2006), the team worked
ahead of and behind the frontline of the progressing chytrid fungus. They
demonstrated that it is possible to collect animals from infected areas, treat
Educational graphics forming part of Panamanian Golden Frog Atelopus zeteki
mixed-species exhibit at the Buffalo Zoo. In addition to A. zeteki, the exhibit
currently houses two other Panamanian species, namely Hylomantis lemur
(Endangered) and Colostethus pratti (Least Concern). © John Kast
Michael Lau returning to
Hong Kong with the first
batch of Romer’s Treefrog
Chirixalus romeri (Endan-
gered) bred at Melbourne
Zoo. © Chris Banks
the winter, carving out semi-permanent plunge pools that provide breeding
grounds for the toad in the spring and summer months when the rest of the
torrent dries up. Although the toad remains restricted to the mountains,
fossil evidence indicates that the species was once widespread across
Mallorca and also inhabited lowland areas (A. Alcover pers. comm.). The
increasingly reduced and fragmented distribution of the species prompted
its listing as Critically Endangered in the 1996 IUCN Red List (Baillie and
Groombridge 1996).
A number of theories have been proposed to explain the decline of the
Mallorcan Midwife Toad. Changes in water demand and population pres-
sure have resulted in increasingly limited availability of breeding sites for
the species and this may have restricted its range to areas where suitable
breeding pools persist. However, most researchers attribute the principal
cause of decline to predation and competition from introduced species such
129
them for chytrid infection regardless of their level of infection, and establish
them in ex-situ populations.
In conclusion, although species persistence must ultimately be achieved in
the wild, it can be facilitated by effective ex-situ conservation programmes.
However, there are several important elements that define a successful
ex-situ programme. Firstly, where ex-situ intervention is warranted to avoid
uncontrollable threats, it is best done in the range country due to disease risks,
political considerations, limited out-of-range capacity, and expense. Ex-situ
facilities worldwide that have adequate resources should partner with zoos
or other institutions in the range country. If capacity does not exist there, it
must be built. Prioritizing efforts within range countries provides renewed
focus, both on capacity building to facilitate in-country accountability and
on government support, for an international response to preserve national
(and global) biodiversity.
Secondly, it is essential that animals removed from the wild and intended
for eventual release be kept in strict isolation (Marantelli et al. in prep) to pre-
vent exposing them to novel pathogens and parasites. The risk is significantly
increased as animals cross more or greater biogeographic barriers from their
natural range. Moreover, specimens must undergo rigorous health screening
prior to return or release to test that isolation has successfully excluded known
organisms. If the ex-situ community is to be responsible, for amphibians or
any organisms they move, they should not gamble on biosecurity issues and
risk becoming the vector for a new disease outbreak in the wildby transfer
of foreign pathogens to the range or host country.
Third, regardless of where the population is maintained, a programme has
a better chance of success if the time spent ex-situ is minimized (thus reducing
the potential for disease exposure, inbreeding, and artificial selection). And,
finally, conservation research, education, and fundraising should be simultane-
ous priorities for ex-situ populations. All of these activities must be linked to,
and for the benefit of, an in-situ conservation programme aimed at reversing
the threats and thereby making the ex-situ programme unnecessary.
Kevin Zippel, Kevin Buley, Richard Gibson,
Graeme R. Gillespie, Robert Johnson, Robert C. Lacy,
Gerry Marantelli, and Joseph R. Mendelson III
References
Banks, C.B. 1996. A conservation program for the threatened Romer’s tree frog (Philautus
romeri). In: P.D. Strimple (ed.), Advances in Herpetoculture, pp. 1-5. International
Herpetological Symposium, Inc, Des Moines, Iowa, USA.
Buley, K.R. and Gonzalez, C. 2000. The Durrell Wildlife Conservation Trust and the Mallorcan Mid-
wife Toad Alytes muletensis - into the 21st century. Herpetological Bulletin 72:17-20.
Denton, J.S., Hitchings, S.P., Beebee, T.J. and Gent, A. 1997. A recovery program for the
natterjack toad (Bufo calamita) in Britain. Conservation Biology 11:1329-1338.
Dudgeon, D. and Lau, M.W.N. 1999. Romer’s Frog reintroduction into a degraded tropical
landscape, Hong Kong, P.R. China. Re-introduction News 17:10-11.
Gibson, R.C. and Freeman, M. 1997. Conservation at home: recovery programme for the
agile frog Rana dalmatina in Jersey. Dodo 33:91-104.
Griffiths, R.A. and Kuzmin, S.L. 2007. Captive breeding of amphibians for conservation.
In: H.H. Heatwole and J.W. Wilkinson (eds.), Conservation and Decline of Amphib-
ians, Amphibian Biology, Volume 8A. Surrey Beatty & Sons, Chipping Norton, New
South Wales, Australia.
Johnson, B. 1999. Recovery of the Puerto Rican crested toad. Endangered Species
Bulletin 24:8-9.
Krajick, K. 2006. The lost world of the Kihansi toad. Science 311:1230-1232.
Krofta, D.M. 2003. California red-legged frog: jumping to survival. Endangered Species
Bulletin 28:18-20.
Lips, K.R., Brem, F., Brenes, R., Reeve, J.D., Alford, R.A., Voyles, J., Carey, C. and Collins, J.P.
2006. Infectious disease and global biodiversity loss: pathogens and enigmatic amphibian
extinctions. Proceedings of the National Academy of Sciences, USA 103:3165-3170.
Mendelson, J.R., III and G.B. Rabb. 2006. Global amphibian extinctions and the role of
living-collections institutions. World Association of Zoos and Aquariums. Proceedings
WAZA Conferences: Proceedings of the 60th Annual Meeting, New York City, USA,
2-6 October 2005: 179-181.
Sredl M.J., Field, K.J. and Peterson, A.M. 2002. Mitigating threats and managing the
Ramsey Canyon leopard frog in Arizona. Nongame and Endangered Wildlife Program
Technical Report 207. Arizona Game and Fish Department, Phoenix, Arizona, USA.
Sredl, M.S. and Healy, B.L. 1999. Conservation and Management Zones: Evaluating an
approach to conserving Arizona populations of the Chiricahua leopard frog (Rana
chiricahuensis). Nongame and Endangered Wildlife Program Technical Report 149.
Arizona Game and Fish Department, Phoenix, Arizona, USA.
Zippel, K.C. 2002. Conserving the Panamanian Golden Frog: Proyecto Rana Dorada.
Herpetological Review 33:11-12.
Tadpoles from a snake-free pool (upper) and a pool subject to predation
from snakes (lower) showing the strikingly different morphologies. © Robin
D. Moore
130
Male Mallorcan Midwife Toad Alytes muletensis (Endangered) carrying
fertilized eggs on its back legs. © Gerardo Garcia
as green frogs, Rana perezi, and, more significantly, the viperine snake,
Natrix maura, a semi-aquatic serpent that preys upon both tadpoles and
adult toads. Archeological and, more recently, molecular evidence (Guicking
et al. 2006) supports the idea that the viperine snake appeared in Mallorca
around 2000 years ago following the arrival of the Romans. Being an island
species that evolved with few natural vertebrate predators, the Mallorcan
Midwife Toad was undoubtedly susceptible to the impacts of predation and
competition from introduced species. The snake may have eliminated the
toad from much of the island, and recent research indicates that predation
from snakes continues to negatively impact some extant populations. In
response to intense predation pressure, tadpoles have evolved a number
of behavioural and morphological anti-predator responses. Upon detecting
chemical cues from snakes in the water, tadpoles reduce activity levels and,
over a matter of weeks, change shape - becoming more streamlined with a
larger tail muscle - to facilitate their escape if they are detected (Moore et al.
2004). Such defences may have allowed the toad to cling onto survival in the
face of intense predation pressure and may have facilitated its co-existence
with the snake in some areas today.
In 1985, at the invitation of the Mallorcan government, the Jersey Wildlife
Preservation Trust (JWPT), now known as the Durrell Wildlife Conservation
Trust (DWCT), initiated a species recovery program for the toad. A captive
population was initially founded at Jersey Zoo, UK, using 20 individuals col-
lected from one site, and the species was bred in captivity for the first time
in 1988. Captive populations stemming from this founder colony are now held
in a number of institutions in both Spain and the UK and these have been
augmented with several more bloodlines that are maintained separately to
ESSAY 11.7. WHEN HABITAT PROTECTION AND SUCCESSFUL CAPTIVE BREEDING ARE NOT ENOUGH: THE CASE OF THE WYOMING TOAD
During the last Ice Age, vast glaciers occupied most of North America’s far
northern latitudes, and the ranges of many northern, cold-climate plants
and animals extended well to the south of their present distributions. With
glacial recession and subsequent warming, the distributions of these species
withdrew northward. One of these, the Canadian Toad Bufo hemiophrys,
disappeared from most of the southern part of its range in the prairie region
of the north-central United States. Until the 1940s, herpetologists believed
that the Canadian Toad lived only as far south as the northern fringe of the
United States, including northern Minnesota and northern North Dakota.
But, in 1946, George Baxter, newly arrived at the University of Wyoming,
discovered an isolated population in the Laramie Basin in south-eastern
Wyoming, about 500 miles away from the nearest known population in
eastern South Dakota. Since then, biologists have scoured the region, but
all known occurrences of this population have been found only within about
50km of the town of Laramie, within an area of about 2,330km², at elevations
between 2,300 and 2,500m asl.
The toad’s habitat in Wyoming consists of wet meadows, marshy wet-
lands, and the moist low-gradient edges of open bodies of water. Baxter’s
initial discoveries were along the floodplains of the Big and Little Laramie
rivers, but recent observations indicate that toads responded to changing
irrigation practices in the basin by switching from floodplain pools and
pooled irrigation water to more isolated ponds and seepage lakes. Toads
survive the Laramie Basin’s long, cold winters by burrowing into the deep
sandy soils of old, vegetated dunes near the breeding areas, or by moving
into rodent burrows.
At the time of its discovery, the toad that Baxter discovered in the Laramie
Basin was regarded as a population of the Canadian Toad. In 1968, Kenneth
Porter of the University of Denver carefully compared the characteristics of the
Wyoming population with those of Canadian Toads in Canada and determined
that the Wyoming toads warranted recognition as a new subspecies, which
he named Bufo hemiophrys baxteri, in recognition of Baxter’s discovery. It was
later considered a distinct species (Smith et al. 1998), and the Laramie Basin
population is now generally known as the Wyoming Toad (Bufo baxteri).
A “wild” Wyoming Toad Bufo baxteri (Extinct in the Wild) from Mortensen
Lake, Albany Country, Wyoming. A single population of Wyoming toads was
discovered at Mortenson Lake in 1987. This location was purchased by The
Nature Conservancy, and the Mortenson Lake National Wildlife Refuge was
established in 1993. © Douglas A. Keinath
Threatened Amphibians of the World
ensure genetic integrity. One of the main aims of the captive-breeding program References
has been to provide toads for reintroduction into their natural habitat.
The first reintroduction of 76 captive toads from JWPT occurred in 1989, Buley, K.R. and Gonzalez-Villavicencio, C. 2000. The Durrell Wildlife Conservation Trust
at the request of the Mallorcan government, into two sites chosen by the and the Mallorcan midwife toad, Alytes muletensis – into the 21st century. Herpe-
Conselleria d’Agricultura Pesca (CDAP). Since then, releases of young toads tological Bulletin 72:17-20.
and tadpoles have been made into numerous unoccupied sites within the Baillie, J. and Groombridge, B. (compilers and editors) 1996. 1996 IUCN Red List of
known historical range of the species. Such sites have been identified after a Threatened Animals. IUCN, Gland, Switzerland and Cambridge, UK.
careful assessment of potential threats, with appropriate habitat and predator Guicking, D., Griffiths, R.A., Moore, R.D., Joger, U. and Wink, M. 2006. Introduced alien or
management carried out in order to maximize the chances of success. Natural persecuted native? Resolving the origin of the viperine snake on Mallorca. Biodiversity
populations are also known to use livestock watering troughs, or cisterns, and Conservation 15:3045-3054.
and construction of such artificial breeding sites in the traditional style has Kraaijeveld-Smit, F.J.L., Griffiths, R.A., Moore, R.D. and Beebee, T.J.C. 2006. Captive
also proved successful in re-establishing populations in areas where water breeding and the fitness of reintroduced species: a test of the responses to predators
is otherwise in short supply. The success of the reintroduction programme is in a threatened amphibian. Journal of Applied Ecology 43:360-365.
illustrated by the fact that about 25% of the current wild population stems Mayol, J. and Alcover, J.A. 1981. Survival of Baleaphryne Sanchiz and Adrover, 1979
from captive-bred stock and that 12 new breeding sites have been established (Amphibia: Anura: Discoglossidae) on Mallorca. Amphibia-Reptilia 3/4:343-345.
since 1989 through reintroductions (Buley and Gonzalez-Villavicencio 2000). At Moore, R.D., Griffiths, R.A., O’Brien, C.M., Murphy, A. and Jay, D. 2004. Induced defences
a recovery group meeting in Jersey in November 2002, a decision was made in an endangered amphibian in response to an introduced snake predator. Oecologia
to suspend reintroductions from ex-situ breeding colonies as a precaution 141:139-147.
against the transmission of diseases, such as the chytrid fungus. It was sug-
gested that conservation efforts rather focus on translocating individuals from
existing populations into new, unoccupied sites and head-starting tadpoles
to increase the chances of success for such translocations. The Sierra Tramuntana of
The recovery program has proven extremely successful in reversing the northern Mallorca, Bale-
decline of the toad. Nineteen breeding populations now exist within the aric Islands, Spain – home
Serra de Tramuntana and six of these have been established from successful to the Mallorcan Midwife
reintroductions of captive individuals. As a result, the species was downlisted Toad Alytes muletensis
by two categories to Vulnerable during the course of the Global Amphibian (Endangered). © Robin
Assessment. The success of this program can be attributed to a number of D. Moore
factors. Firstly, there has been effective collaboration among international
governments, NGOs, academic institutions, and zoos. This collaboration has
meant that captive breeding and reintroduction has been just one component
of a wider recovery program involving threat mitigation and habitat manage-
ment, health screening of wild and captive toads, ongoing monitoring of all
the toad populations, and raising awareness among visitors and stakeholders.
Secondly, these activities have been underpinned by targeted research on
reproductive biology, survey methodologies, the impact of threats, and popula-
tion genetics. Recent research, for instance, has shown that relatively high
levels of heterozygosity and important fitness attributes can be retained for a
few generations of captive breeding in this species, but that both may start to
deteriorate in the long-term (Kraaijeveld-Smit et al. 2006); such findings have
important implications for this, and other, recovery programmes.
Extant populations continue to be monitored and captive populations
maintained as a ‘safety net’ and for conservation research in order to ensure
the continued recovery of the Mallorcan Midwife Toad. This represents a rare
conservation success story and has the potential to serve as a model for other
amphibian recovery programmes throughout the world.
Robin D. Moore and Richard A. Griffiths
Baxter’s field observations over many years indicated that the Wyoming Basin might be a major factor in the decline. Concerted survey efforts in the
Toad was relatively common in the 1950s and 1960s, but then underwent a 1980s fortunately revealed tiny remnant populations but, by 1987, the known
substantial decline in distribution and abundance in the 1970s. In fact, by the range included only a single site. Wild Wyoming Toads have since been found
mid-1980s, the Wyoming population appeared to be extinct. At the time, there only at this site, initially protected by The Nature Conservancy and now
was concern that pesticide applications for mosquito control in the Laramie included in the Mortenson Lake National Wildlife Refuge.
 Chapter 11. Amphibian Conservation – Responding to the Global Decline of Amphibians
In the late 1980s and early 1990s, the Mortenson Lake population persisted
with an adult population in the low 100s, and then declined to just a few adults
by 1993. Breeding output dropped from a few egg masses per year to zero.
Fearing that the toad might disappear completely from Wyoming, biologists
collected the few remaining individuals in 1993 and 1994 and began attempts
to breed them in captivity. This effort, now involving several zoos and other
facilities, generated good numbers of offspring for release into the wild. An
effort in the early 1990s to establish additional populations through releases
of captive-raised toads at Lake George and Rush Lake on the Hutton Lake
National Wildlife Refuge was unsuccessful. At Mortenson Lake, releases of
many thousands of toadlets and tadpoles beginning in the mid-1990s resulted
in at least modest toad survival and even a renewal of reproduction in the wild
in 1998-2000, generating hopes for population recovery. However, subsequent
declines, poor reproduction, and observations of diseased and dead toads in
the early 2000s, put the toad back into an extremely precarious status. The
free-ranging population surely would be completely gone without annual
releases of thousands of captive-reared toadlets. Accordingly, the species
remains functionally extinct in the wild.
Several factors may have contributed to the toad’s decline and current poor
condition. As with most threatened species, the toad has experienced habitat
loss and degradation. Irrigation practices in river floodplains, now used for
hay production, probably made streamside areas less suitable for successful
toad reproduction. For example, de-watering prior to hay-cutting may kill toad
larvae before they metamorphose into toadlets capable of living on land. Also,
some potential breeding sites have dried up as a result of prolonged drought,
while drought-related increases in evaporation have made Mortenson Lake
more saline and perhaps less suitable for toads.
However, recent evidence suggests that the primary threat to the Wyoming
ESSAY 11.8. CONSERVATION ACTION FOR THE MEXICAN AXOLOTL AMBYSTOMA MEXICANUM
The Mexican Axolotl (Ambystoma mexicanum) is one of Latin America’s
most threatened amphibians. The vast wetland upon which Mexico City
was founded – and which once provided a rich and productive habitat for
the Axolotl and other endemic fauna – is now reduced to a handful of small,
isolated patches surrounded by development. Of these, Lake Xochimilco is
the largest, covering just over 2km² – but it is certainly no longer a lake.
The development of the ‘chinampas’ – raised fields of mud and vegetation
reclaimed from the lake – has been going on for centuries and has reduced
the system to a series of canals running between islands of development.
Today, the landscape is often referred to as the ‘floating gardens’ (a misnomer,
as the chinampas are not floating at all). Habitat loss, introduced predators,
pollution, and illegal collection for food and medicines have all taken their
toll on the Axolotl. Consequently, the threats facing this species are complex
and not easily reversible. However, its prominent position within Aztec
mythology (see Essay 2.3) and the ancient lacustrine economy of the region
means that the Axolotl is well known – although poorly understood – among
local people. Some 2000 remeros (local boatmen) earn a living by punting
visitors along the lake’s canal system in gaily decorated trajineras (pleasure
boats), while the chinamperos (local farmers) cultivate the adjacent land,
in much the same way as their ancestors have done for centuries. Fishing
is also important to the local economy, and although non-native carp and
Tilapia may have replaced the Axolotl as the main catch, researchers have
yet to improve upon the highly skilled traditional netting method used by the
fishermen for finding Axolotls.
Over the last five years, a conservation programme on the Axolotl has been
initiated and fostered by a partnership of British and Mexican organizations
(Griffiths et al. 2004). This project was the brainchild of the late Dr Virginia
The Critically Endangered Mexican Axolotl Ambystoma mexicanum is endemic to the canals of Lake Xochimilco in central Mexico, on the southern edge of
Mexico City. © Ian G. Bride
Toad is not habitat degradation, but rather the pathogenic chytrid fungus
(Batrachochytrium dendrobatidis), now associated with amphibian declines
around the world. The fungus has been discovered at Mortenson Lake and
in the captive population. Retrospective analyses indicate that the chytrid
fungus has been present at Mortenson Lake since 1989.
Continued survival of the Wyoming toad depends on intensive manage-
ment. An important immediate concern is maintenance of the toad’s genetic
diversity through careful management of the captive breeding stock. Exces-
sive inbreeding and associated loss of genetic variation potentially could
compromise the toad’s reproductive performance and hinder the population’s
ability to respond to environmental variations.
Even if the Mortenson Lake population were in better condition, the
toad’s long-term survival and recovery would still depend on the identifica-
tion of additional reintroduction sites and the establishment of several
wild populations. This is a basic conservation precaution that minimizes
the probability that localized events do not eliminate the entire species.
Happily, private landowners have begun to step forward and allow toads to
be released into suitable habitat on their property. A recent initiative that
may play an important role in facilitating this effort is known as the Safe
Harbor Agreement (SHA). According to the U.S. Fish and Wildlife Service, a
SHA “encourages landowners to conduct voluntary conservation activities
and assures them that they will not be subjected to increased restrictions
should their beneficial stewardship efforts result in increased endangered
species populations. As long as enrolled landowners allow the agreed upon
habitat improvements to be completed on their property and maintain their
responsibilities, they may make use of the property during the permit term,
even if such use results in the take of individual Wyoming toads or harm to
their habitat. This approach may be critical in establishing additional sites
Graue of the Universita Autonoma Metropolitana at Xochimilco (UAM-X),
who contacted the Durrell Institute of Conservation and Ecology (DICE) in
1999 for assistance with the development of the project. As it was clear
that addressing the many threats that the Axolotl faced would be impos-
sible without the co-operation of local stakeholders, the project focused on
embracing local people within the conservation planning process. This was
done by promoting the Axolotl as a flagship species for nature tourism and
conservation education within the region. Using a field station (run by UAM-X)
on the shores of the lake as a base, and with funding from the IUCN/ SSC
Declining Amphibian Populations Task Force (DAPTF), as well as the British
Government’s Darwin Initiative programme, the project partnership held train-
ing workshops on amphibian biology and conservation for local students and
conservation organizations, nature guiding for local boatmen, and souvenir
production for unemployed artisans.
In addition, the project has been actively engaged in ongoing studies
focusing on the population status of, and threats to, the Axolotl. Despite its
precarious status in the wild, the Axolotl is one of the most familiar amphibians
in laboratories and aquaria throughout the world. Animals were originally col-
lected in 1863 for the Natural History Museum in Paris, and many of today’s
captive animals probably stem from these founders (Smith 1989). As a result of
its well-known reproductive biology, and the availability of captive populations,
there is considerable interest in reintroducing Axolotls to Lake Xochimilco.
However, there are several problems associated with such releases. At the
very least, threats need to be neutralized and potential disease and genetic
problems need to be addressed before captive animals are put back into the
wild (Griffiths et al. 2004). Despite the wide availability of captive-bred Axo-
lotls, wild animals are still captured and sold illegally in local markets (McKay
131
into which captive-reared toadlets might be released with improved chances
of survival and eventual reproduction.”
Recently, a SHA was involved in the release of captive-raised toads on a
private ranch near Mortenson Lake. Well-managed cattle grazing – which is
compatible with and may even facilitate toad recovery – is being continued
on the ranch. Only time will tell if the toads can avoid or overcome the
fungal threat and establish a viable population on this and other potential
reintroduction sites. Meanwhile, the Wyoming Toad remains one of the
world’s most threatened species.
Geoffrey A. Hammerson
References
Anonymous. 2005. Statewide programmatic biological assessment for the Wyoming toad
(Bufo baxteri). Final report submitted to BLM Wyoming State Office, 5353 Yellowstone
Road, Cheyenne, Wyoming 82003-1828.
Odum, R.A. and Corn, P.S. 2005. Bufo baxteri Porter, 1968. Wyoming toad. In: M. Lannoo
(ed.), Amphibian declines: the conservation status of United States species, pp. 390-
392. University of California Press, Berkeley, California, USA.
Smith, H.M., Chiszar, D., Collins, J.T. and van Breukelen, F. 1998. The taxonomic status of
the Wyoming toad, Bufo baxteri Porter. Contemporary Herpetology 1998(1):http//alpha.
selv.edu/ch/1998/1.
Taylor, S.K., Williams, E.S., Thorne, E.T., Mills, K.W., Withers, D.I. and Pier, A.C. 1999.
Causes of mortality of the Wyoming toad. Journal of Wildlife Diseases 35:49-57.
U.S. Fish and Wildlife Service. 1991. Wyoming toad recovery plan. U.S. Fish and Wildlife
Service, Denver, Colorado, USA.
2003). A proposal to upgrade Ambystoma mexicanum from CITES Appendix II
(controlled international trade) to Appendix I (species threatened with extinc-
tion and international trade permitted only in exceptional circumstances) is
currently under discussion by the Mexican authorities.
As a result of a conservation workshop, held at UAM-X in December 2004,
the goals for the conservation of the Axolotl and its habitat were finally dis-
tilled into eight categories: (1) biology of the species; (2) legislation; (3) social
actions; (4) political actions; (5) ecological interactions; (6) local environment;
(7) education; and (8) resource use and harvesting. These goals provided the
framework for the Species and Habitat Action plan for the Axolotl and the
Xochimilco system that was published in 2005 (DICE, UAM-X, 2005). Follow-
ing the workshop, the Senate of the Congress of the Mexican government
passed a motion requesting the President to instruct the Secretariat of the
Commission for the Environment, Natural Resources and Fishing to initiate
an urgent governmental programme to avoid the extinction of the Axolotl in
Lake Xochimilco. What this decision will mean, in practice, remains to be
seen, but it will certainly influence governmental actions that impact the
Axolotl and Xochimilco conservation. The decision, in itself, is an indication
of the leverage that such projects can achieve.
One of the most important products to emerge from the project’s first phase
is the partnership of diverse organizations that all have an interest in the future
of the Axolotl and its habitat: the Grupo por la Investigacione del la Ajolote y
Xochimilco (GIAX), which is co-ordinated by Dr Luis Zambrano of the Institute
of Biology, UNAM. Despite the obvious benefits accruing as a result of raising
awareness and building local capacity, initial data from this first phase of
the project indicate that the status of the Axolotl has deteriorated to such an
extent that the species now warrants classification as ‘Critically Endangered’
according to the IUCN Red List categories and criteria. This reclassification
has provided a sharper focus for the implementation of the action plan. The
new partnerships forged within GIAX will be instrumental in taking the plan
forward during the second phase of the Darwin Initiative project. The training
of remeros as nature guides has resulted in a significant increase in their
income from visitors to Lake Xochimilco. Some of the remeros are currently
being trained as nature guide ‘trainers’, with a view to the training package
eventually being handed over to, and managed by, the guild of remeros. By
the end of the second phase of the project, it is hoped that nature tourism will
be contributing significantly to the sustainable development of the Xochimilco
system, and will be independent of external funding. In parallel, research is
A typical Xochimilco canal scene beyond the busy urban area, showing a
trajinera (pleasure boat) in the foreground. The boats in the distance are
selling ornamental plants grown on the chinampas. © Ian G. Bride
132 Threatened Amphibians of the World

being carried out on the productivity of the Xochimilco system, with a view Some of the 50-plus participants
to balancing the needs of the local fishermen with ameliorating the threats who attended the Axolotl Species/
associated with introduced species and poor water quality. Ultimately, the Habitat Action Plan Workshop, held
future of the Axolotl will depend upon how successfully local stakeholders and at the Universita Autonoma Metro-
their livelihoods engage with the protection of this unique wetland. politana at Xochimilco in December
2004. Participants are holding up
Richard A. Griffiths, Ian G. Bride, and Jeanne E. McKay the name plate of the trajinera that
the Darwin Initiative project bought
for CIBAC. © Ian G. Bride
References
DICE, UAM-X, 2005. The Conservation of the Axolotl (Ambystoma mexicanum) in Xochi-
milco, Mexico City-A Species/Habitat Action Plan. http://www.kent.ac.uk/anthropol-
ogy/dice/research/azaxs/
Griffiths, R.A., Graue, V., Bride, I.G., and McKay, J.E. 2004. Conservation of the axolotl (Am-
bystoma mexicanum) at Lake Xochimilco, Mexico. Herpetological Bulletin 89:4-11.
McKay, J.E. 2003. “An evaluation of captive breeding and sustainable use of the
Mexican axolotl (Ambystoma mexicanum)”. MSc dissertation, University of Kent,
Canterbury, UK.
Smith, H.B. 1989. Discovery of the axolotl and its early history in biological research. In
Developmental Biology of the Axolotl, pp. 3-12. In: J.B. Armstrong and G.M. Malacinski
(eds.). Oxford University Press, New York, USA.

ESSAY 11.9. MANAGING PROBLEMS OF OVER-EXPLOITATION AND TRADE IN AMPHIBIANS

To many cultures, amphibians are an important and commonly available source
of protein. In some places, even tadpoles can be made into a local dish. The
water-holding frogs in Australia store a reservoir of water in the bladder to
last through the extended dry season and are used by the aborigines when no
other source of water is available (Tyler 1976). Certain amphibians are also
commonly used in traditional medicine. For example, in China, nine species
of amphibians are listed in “The Great Pharmacopoeia” published over four
hundred years ago. Up to 32 species are now recognized to be of medicinal
value in traditional Chinese medicine (Ye et al. 1993). Some amphibians are also
exploited to provide useful materials, like the poison from the various Poison
Dart frogs for hunting and the skin of the Marine Toad for the leather industry
in the Americas. Many exotic frogs, such as the colourful Mantellas (genus
Mantella) from Madagascar, and a number of salamander species, are also
collected to supply the pet trade. There is also quite a demand for frogs for use
in the laboratory. And, in some countries such as Mexico, certain amphibians
are used in witch craft and art crafts. Amphibians also play a role in shaping
the local culture. For example, the Mexican Axolotl Ambystomma mexicanum,
which has various mythological connections – the ancient Mexicans considered
it the twin brother of Quetzalcoatl – has played an important role in the local
communities around lakes in the Basin of Mexico (see Essay 2.3). There is even
a frog festival in a branch of the Zhuang Minority in southern China.
These species survived centuries of traditional use and the ‘edible frogs’
remained common and widespread until commercial trade set in leading to a
much larger level of exploitation that is often unsustainable. As early as the
1930s, Bourret (1942) noted the mass commercial trade of large edible frogs
from Viet Nam into China. The collapse of populations of favourite ‘frog legs’
species in Asia shows that even in common, fast-growing and fecund species,
such levels of exploitation are not without limit. Once depleted, the ecological
functions carried out by these frogs are also hampered, which contributes
to the outbreak of insect pests in rice paddies. The demand for amphibians
is unlikely to diminish in the near future due to continual growth in human
population and the increased purchasing power that accompanies growing
economies. The resulting high collecting pressure often acts together with
habitat destruction and degradation and poses a substantial threat to many
amphibians. In China, for example, utilization adversely affects 84 species,
some 30% of the amphibian fauna (Baillie et al. 2004).
The species hardest hit by over-exploitation are those that are rare, re-
stricted or highly valuable. In recent years, a number of frogs and salamanders
such as the Lao Salamander Paramesotriton laoensis (DD) and Kaiser’s Spotted
Newt Neurergus kaiseri (CR) have been subjected to commercial collection
for pet trade at levels that are believed to have removed the majority of
individuals from the wild (Stuart et al. 2006; Leahy 2006). Another example
of a species hard hit by commercial exploitation, but for food, is the Chinese
Giant Salamander Andrias davidianus (see Essay 4.7). This species, the
largest amphibian in the world (adults can weigh more than 40kg) ranges
widely in central and southern China and large populations once existed in
many places. However, it is regarded as a delicacy and fetches a very high
price. The species is particularly susceptible to over-harvesting because it
is nearly fully aquatic and utilizes specific hiding places in clear mountain
streams. Juveniles are also of marketable size and are collected as well as
adults. Highly destructive methods, such as liming or the use of poison, are
sometimes used to collect all the individuals in the stream. Individuals are
also being collected as breeding stock in commercial farms. All these fac-
tors together with the long life-cycle make the Chinese Giant Salamander
especially vulnerable to over-exploitation, and this Critically Endangered
species has now disappeared from many areas, and a conservation action
plan is urgently needed to reverse its decline.
Frog farming has gained momentum in many places and commercial
feed is now available to feed the frogs and their tadpoles. These farms
can make a positive contribution towards conservation if the supply from
breeding farms substitutes for that from the wild, or if part of the income
generated is directed back into biodiversity conservation. However, there
remain challenges, including disease control within farms and the economic
cost/benefit of farming efforts as a business activity (Kusrini and Alford
2006). One negative consequence of amphibian farming is that the American
Bullfrog is one of the favourite species in many parts of the world. These
animals often escape and become invasive, affecting the local ecosystem
and biodiversity. In Mexico, for example, this species has been introduced
into 16 of the country’s 32 states. Most places actually have their own ‘ed-
ible’ frogs that are more suited to the local environment and are much better
candidates for farming than exotic species. However, regardless of species,
frog farming still runs the risk of transmitting diseases from captive frogs to
those in the wild anywhere from the farm surroundings and transport routes
to destination markets.
An alternative to farming and its associated problems is to explore op-
tions for sustainable use of local wild populations. The short life-cycle, high
fecundity and high population size of many large frogs actually renders them
quite resilient to certain levels of harvesting. If the requirements of the species
concerned are known, and proper management is in place, these frogs can be
harvested without affecting their population. There are examples in northern
China where harvest of the Chinese Brown Frog Rana chensinensis (LC), a
species used in traditional Chinese medicine, can be increased by providing
breeding ponds and raising the tadpoles (Ye et al. 1993). Another example
of sustainable use of amphibians is the harvesting of the grass frog Rana
forreri (LC), which is a popular laboratory animal. These animals are harvested
according to an annual quota and then exported to the United States. In
Thailand and Indonesia, frogs (mainly Hoplobatrachus rugulosus, Fejervarya
cancrivora and Fejervarya limnocharis) living and breeding in rice paddies,
not only feed on crop pests but also are harvested for local consumption and
export (Kusrini and Alford 2006). When combined with sustainable land-use
practices, such as organic farming, this increases the economic gains, reduces
or eliminates the need for and use of chemicals, and can bring additional
conservation benefits to many other amphibians and wildlife. It might also
provide an additional incentive to conserve their natural habitats in the case
of non-commensal species.
To prevent unrestricted exploitation and possible extirpation of popula-
tions and species, several countries have implemented legislation that
specifically regulates or prohibits the exploitation of particular amphibian
species. Legislation usually also exists to protect particular natural areas as
parks or sanctuaries, and amphibian populations inside such areas may be
partly or completely protected by the regulations in force for the area. For
species threatened by international trade, the Convention on International
Trade in Endangered Species of Wild Fauna and Flora (CITES) is quite effec-
tive in regulating levels of trade and even in banning the commercial trade
in highly threatened species (see Chapter 11). Adequate local legislations
and enforcement is also needed as many amphibians are consumed locally.
Wildlife trade is dynamic, and monitoring and reporting are important to
provide an early warning system.
Michael Lau, Peter Paul van Dijk and Gracia P. Syed
References
Baillie, J.E.M., Bennun, L.A., Brooks, T.M., Butchart, S.H.M., Chanson, J.S., Cokeliss, Z.,
Hilton-Taylor, C., Hoffmann, M., Mace, G.M., Mainka, S.A., Pollock, C.M., Rodrigues,
A.S.L., Stattersfield, A.J. and Stuart, S.N. 2004. 2004 IUCN Red List of Threatened Spe-
cies. A Global Species Assessment. IUCN, Gland, Switzerland and Cambridge, UK.
Bourret, R. 1942. Les Batraciens de l’Indochine. Institut Océanographique de l’Indochine,
Hanoi. 547 pp.
Kusrini, M.D. and R.A. Alford. 2006. Indonesia’s exports of frogs’ legs. TRAFFIC Bulletin
21(1):13-24.
Leahy, S. 2006. Biodiversity: The insatiable in pursuit of the inedible. http://www.ipsnews.
net/print.asp?idnews=35633
Stuart, B.L., Rhodin, A.G.J., Grismer, L.L. and Hansel, T. 2006. Scientific description can
imperil species. Science 312:1137.
Tyler, M.J. 1976. Frogs. Collins, Sydney, Australia.
Ye, C., Fei, L. and Hu, S. 1993. Rare and Economic Amphibians of China. Sichuan Publishing
House of Science and Technology, Chengdu, China. [In Chinese]

A farmed, market-bought Hoploba-

trachus rugulosus (Least Concern),
the most common species used in
the frog-leg trade in Asia, showing
obvious signs of farm rearing, in-
cluding bulbous toe-tips and trans-
port damage behind the nostrils.
© Peter Paul van Dijk, 1994-2000

An assortment of local language

frog-farming manuals for the rearing
of Hoplobatrachus rugulosus (in Chi-
nese and Thai), Rana catesbeiana
(Thai), Paa spp. (Chinese) and local
ranids (Chinese). © CI/Peter Paul
van Dijk, 2007
 Chapter 11. Amphibian Conservation – Responding to the Global Decline of Amphibians 133

ESSAY 11.10. FILLING THE BLACK HOLE: CHALLENGES IN TAXONOMY TO PROTECT AMPHIBIANS

Less than 10% of species on the planet have been discovered and fewer
than 1% are known beyond brief anatomical descriptions (Wilson 2005).
Without a doubt, our alarmingly inadequate knowledge of the Earth’s diversity
is one of the most significant challenges to effectively protect threatened
species, all the more urgent as biologically rich regions are destroyed at
unprecedented rates.
An excellent example of the significance of this problem is provided by
extinction risk assessments, such as those integral to the Global Amphib-
ian Assessment. However, Red Lists of threatened species only evaluate
formally described species. Although undescribed species may well have
a high risk of extinction, they are ignored by conservation initiatives that
rely on information contained within the IUCN Red List. Therefore, filling
this taxonomic void should be a priority for the implementation of effective
conservation programmes.
Among terrestrial vertebrates, sampling of amphibians appears to be par-
ticularly incomplete. This is indicated by the large number of species described
during recent decades (for example, see Figure 1) at a rate of discovery that
exceeds that of every other vertebrate group (Cannatella and Hillis 2004). Dur-
ing the period 1992-2003, the number of species of amphibians increased by
25% with most of the additions being new discoveries rather than subspecies
elevated to the species rank (Kohler et al. 2005; and see Essay 1.1). Recent
estimates indicate that the number of amphibians awaiting formal descrip-
tion in Southeast Asia and the Neotropics should be well above 1,000 (see
below). These figures suggest that, at a global scale, the taxonomic deficit
could be a serious obstacle for the success of any comprehensive programme
to protect amphibians. Thus, intensive efforts in taxonomy and systematics
are indispensable to secure adequate conservation measures.
Fortunately, the increasing use of genetic markers in systematics could
increase the rate at which amphibian species are discovered. These tools
quantify genetic variation within and among populations, which can expose
genetic clusters that correspond to separate species. These data are usually
complemented with geographic distribution, and morphological, or behav-
ioural data to corroborate species identities. The use of genetic markers
allows the identification of previously unsampled species and enables us to
detect cryptic species that have previously been hiding under a single mor-
phologically identified species. The available techniques include allozymes,
AFLPs, microstatellites, and, most prominently, DNA sequencing (including
DNA bar-coding; see Essay 11.11).
Genetic markers have been used recently to estimate the number of spe-
cies of amphibians that remain to be discovered. In a review of species-level
molecular phylogenies of Neotropical amphibians, Ron et al. (2006) found that
phylogenetic studies have lead to a 28% increase in the number of known
species. The increase was higher (39%) for studies where taxon sampling
has been more intensive (i.e., studies that included more than 50% of the
described species). This indicates that a significant number of species have
either been overlooked by morphology-based taxonomic reviews or have
not been sampled at all.
Given that there are approximately 2,800 described species of amphib-
ians in the Neotropics and assuming, conservatively, that the proportion of
undescribed amphibians lies between 0.28 and 0.39, then the number of
Neotropical species awaiting description should lie between 784 and 1,092.
This estimate is considered conservative because the proportion of species
discovered should increase with taxon sampling, which until now has been
exhaustive only in a few Neotropical clades. Phylogenies with non-exhaus-
tive sampling often include predominantly species of easy access, available
in the pet trade (e.g., poison-arrow frogs of the genus Dendrobates) and/or
distributed in habitats that, because of their relative accessibility and con-
spicuousness, are already described.
Applying the same logic and approach to another amphibian species-rich
faunal region (Indo-China, and the island archipelagos of the Philippines, Malay-
sia, and Indonesia), we can provide estimates of the magnitude of the problem
Advertisement calls provide a
cost-effective tool for species
discovery and identification. Male
Trachycephalus venulosus (Least
Concern) call by inflating large
lateral vocal sacs. They occur in
dry forests in western Ecuador.
© Santiago R. Ron
A potential caveat with molecu-
lar techniques is that some genetic
markers are inadequate to define
species boundaries because they
either have evolved too slowly or to
fast. In addition, the use of genetic
markers is still limited in developing
countries, precisely in the regions
where the taxonomic void is more
extensive. Thus, species identifica-
tion in many circumstances will
continue to rely primarily in morpho-
logical or behavioural traits.
An alternative to discover mor-
phologically cryptic species is through analysis of advertisement calls. References
In anurans, males produce advertisement calls which function to attract
females, defend territories, or confront competing males. Because adver- Alcala, A.C. and Brown, W.C. 1999. Philippine frogs of the genus Platymantis (Amphibia:
tisement calls also have a function in species recognition and discrimination Ranidae). Philippine Journal of Science 128:281-287.
(Gerhardt and Huber 2002), they can be excellent indicators of species Brown, R.M. 2004. Evolution of ecomorphological variation and acoustic diversity in
boundaries, in some cases as reliable as genetic markers. In one increas- mate-recognition signals of Southeast Asian forest frogs (subfamily Platymantinae).
ingly well-studied group, the Ceratobatrachine frogs of Southeast Asia and Ph.D. Dissertation, University of Texas, Austin, Texas, USA.
the South-West Pacific (Alcala and Brown 1999), advertisement calls have Cannatella, D.C. and Hillis, D.M. 2004. Amphibians: living a life of slime. In: J. Cracraft and M.J.
provided a small group of modestly-funded researchers with a powerful Donoghue (eds.), Assembling the tree of life, pp. 430-450. Oxford University Press, Oxford, UK.
means of species delimitation. Combined with analysis of mitochondrial Evans, B.J., Brown, R.M., McGuire, J.A., Supriatna, J., Andayani, N., Diesmos, A.C.,
gene sequences, advertisement calls have served as the primary set of Iskandar, D., Melnick, D.J. and Cannatella, D.C. 2003. Phylogenetics of fanged frogs:
characters for the recognition of more than 45 undescribed species over testing biogeographical hypotheses at the interface of the Asian and Australian faunal
the past 10 years. Of approximately 60 species for which sequence data zones. Systematic Biology 52:794-819.
and call data are both available (Brown 2004, unpubl.), only two cases Gerhardt, H.C. and Huber, F. 2002. Acoustic communication in insects and anurans. The
exist where sequence data and call data lead to differing conclusions University of Chicago Press, Chicago, USA.
with regards to the distinctiveness of a species. Both are cases in which Köhler, J., Vieites, D.R., Bonett, R.M., García, F.H., Glaw, F. Skeinke, D. and Vences, M.
populations of frogs possess very distinctive advertisement calls, but are 2005. New amphibians and global conservation: a boost in species discoveries in a
genetically indistinct from sympatric congeners, and both are suspected highly endangered vertebrate group. BioScience 55:693-696.
cases of hybridization and introgression. Ron, S.R., Santos, J.C. and Cannatella, D.C. 2006. Phylogeny of the túngara frog genus
The overwhelming trend that results from this work is the general conclu- Engystomops (= Physalaemus pustulosus species group; Anura; Leptodactylidae).
sion that genetic and call data are both very illustrative tools for identification Molecular Phylogenetics and Evolution 39:392-403.
of cryptic independent evolutionary lineages, particularly when used in Stuart, B.L., Inger, R.F. and Voris, H.K. 2006. High level of cryptic species diversity revealed
complimentary fashion by field workers. by sympatric lineages of Southeast Asian forest frogs. Biology Letters 2:470-474.
Wilson, E.O. 2005. Systematics and the future of Biology. Proceedings of the National
Santiago Ron and Rafe M. Brown Academy of Sciences of the United States of America 102:6520-6521.
Figure 1. An example of the large number of amphibians that remain to be described is given by the accumulated number of formally described species
of amphibians in Ecuador (period 1758-2006). The number of species has almost doubled since 1970 and additions continue at a fast rate, especially from
montane habitats.
450
400
350
Number of species

300
of underestimated biodiversity using the recent phylogenetic studies of Evans
250
et al. (2003), Brown (2004), and Stuart et al. (2006). Our estimate of undetected
species in this region lies between 271 and 364 undescribed forms. We can 200

further examine the trends in one small oceanic island archipelago country (the 150
Philippines) that has been the subject of recent intensive diversity assessment 100
(see http://www.herpwatch.org) and “ground truth” these estimates with first
50
hand knowledge of new species awaiting description. The estimated numbers
for the Philippines lie somewhere between 32 and 43 undescribed species (see 0
1755 1772 1792 1812 1832 1852 1872 1892 1912 1932 1952 1972 1992
Essay 7.3). We also have personal knowledge of approximately 55 undescribed
species awaiting description in the genera Platymantis and Limnonectes from Year
the Philippines and Sulawesi (R. Brown, A. Diesmos, and A. Alcala, unpubl.), Total (all regions) Montane forests Paramo Amazon rainforest
suggesting again that the estimation process is conservative.

ESSAY 11.11. AMPHIBIAN CONSERVATION: NEW PERSPECTIVES THROUGH DNA BARCODING

Effective conservation requires that conservation practitioners are armed
with the appropriate knowledge, specifically about what to conserve and
where to conserve it. For this reason, conservation efforts on the ground rely
heavily on comprehensive assessments in the field (i.e. species inventories)
supported by appropriate taxonomic studies. Unfortunately, for the majority of
living organisms worldwide, both have been inadequately addressed. Among
amphibians, for example, we know that many tropical regions are still far from
being well sampled (cf. Duellman 1999) and the recent observed increase in
species numbers (over 25 per cent in 11 years; Köhler et al. 2005; and see
Essay 1.1) is largely due to the intensified exploration of tropical areas and
the application of modern techniques such as bioacoustics and molecular
genetics. A major problem in both field surveys and inventory work, as well
as in taxonomy, is that species are often difficult to identify. In amphibians,
this is particularly problematic in species-rich genera, such as Afrotropical tree
frogs (e.g., Hyperolius) or the Neotropical harlequin toads (Atelopus), which
generally display little inter-specific morphological variation.
The amphibian fauna of the Kakamega Forest in western Kenya represents
a case in point. This small forest fragment (240km²), long recognized as a site
of global conservation importance (e.g., Fishpool and Evans 2001), is under
increasing pressure, in particular due to firewood gathering and agricultural
expansion. Despite the fact that it is one of the best sampled regions in
Kenya, five frog species were recorded only recently (Lötters et al. 2006)
resulting in a total of 25 species from this site. However, the taxonomic
status of several species from the Kakamega Forest still remains difficult to
determine due to their cryptic nature. For example, through extensive and
meticulous “detective” work, we found that there are, in fact, two species of
the genus Phrynobatrachus in the Kakamega Forest which are morphologically
indistinguishable (Lötters et al. 2006), and at least three scientific names may
be applicable to them (P. mababiensis from Namibia, P. minutus from Ethiopia
and P. scheffleri from Kenya).
An exciting new tool to help speed the identification of known species
and the discovery of new ones, is termed ‘DNA barcoding’, in which a short
standardized fragment of DNA is sequenced and compared with reference
data (e.g. Herbert et al. 2003). The method is easy to perform and also allows
for more objectivity in species diagnosis, and only small tissue samples are
required from study animals, thereby reducing time and costs. DNA barcoding
is similar to the universal barcodes of products in the retail industry, and it
was with the idea of making the barcode of every species on Earth available,
i.e. “writing the encyclopedia of life”, that the Consortium for the Barcode
134 Threatened Amphibians of the World

of Life was initiated in 2004 (http://www.barcoding.si.edu/). Although still
science fiction, we may be well on our way to one day using handheld ‘tricod-
ers’ to identify any life-form directly in the field, just as Mr. Spock did in the
famous 1960s space drama Star Trek. Apart from the obvious advantages
for conservation-related initiatives, barcoding’s benefits include the ability
to identify fragmentary specimens, and different life-stages (e.g., amphibian
larvae) to the level of species (review in Savolainen et al. 2005).
There has been a lot of discussion about the barcoding movement, includ-
ing the anxiety that classical taxonomy will be replaced by a code-based,
rather than organism-focused, discipline that deals with ‘black box’ species
(i.e., information without knowledge), far removed from actual species and
hence from conservation (e.g. Ebach and Holdrege 2005). DNA barcoding can
only be a supplement to classical taxonomy and consequently conservation
(cf. Gregory 2005). Furthermore, the accuracy of DNA barcoding is not quite
100 per cent. In some cases, although genetic variation is low, data other than
genetic information (such as advertisement calls) can also suggest species
status (e.g. Lötters et al. 2005).
The idea of using a single universal genetic marker for all groups of organ-
isms has been critically discussed. In contrast to a fragment of the cytochrome
oxidase subunit I gene (COI) (Herbert et al. 2003), which has been widely
used in, for example, fish and birds, Vences et al. (2005) advocated that in
amphibians the mitochondrial 16S rRNA gene fulfills the requirements better
then COI due to stronger constancy of priming sites. Currently, a debate has
arisen about the use of mitochondrial versus nuclear DNA, or both combined,
for barcoding (Rubinoff 2006).
But how relevant and useful is DNA barcoding for the purposes of
amphibian conservation? Can we just undertake a “rapid assessment”
into an unexplored region, collect amphibian samples, barcode them, and
immediately obtain results? Certainly, some databases already exist with
catalogued DNA barcode details, such as NCBI/GenBank in the USA, Europe’s
EBI/EMBL Nucleotide Sequence Database, and the DNA Database of Japan
(DDBJ) – all inter-linked. However, the data are unevenly distributed with
respect to taxon and geography. For example, a search of GenBank1, the
largest and most widely used of the databases, for 16S gene sequences of
the Neotropical poison frogs (Dendrobatidae), a family of frogs attractive to
scientists for different reasons, reveals that these are well represented (665
sequences versus ~ 240 described species). In contrast, the abovementioned
Afrotropical frog genus Phrynobatrachus is represented by not more than 67
16S gene sequences, and 17 putative taxa out of about 69 known species.
Apart from problems due to missing sequence data for many species, other
criticisms include the need to rely entirely on the taxonomic identification
of original voucher specimens against which sequence data is checked, the
genetic purity of the sequence itself, and the possible overlap of inter- and
intra-specific variation (Meyer and Paulay 2005).
In summary, DNA barcoding certainly provides new perspectives to
amphibian conservation. However, at the current stage, its applicability
is still limited due to lack of reference data for most species and because
the taxonomy of many species (complexes) is not at the stage that allows
samples from the field to be quickly sequenced and then definitively allocated
to species. Nonetheless, coupled with traditional taxonomic tools, it could
have important uses for disclosing hidden diversity.
Stefan Lötters and Susanne Schick
References
Duellman, W.E. 1999. Patterns of distribution of amphibians: a global perspective. Johns
Hopkins University Press, Baltimore, Maryland, USA.
Ebach, M.C. and Holdrege, C. 2005. DNA barcoding is no substitute to taxonomy. Nature 434:697. Afrixalus osorioi (Least Concern), a banana frog only recently recorded from
Fishpool, L.D.C. and Evans, M.I. (eds.). 2001. Important Bird Areas in Africa and associated the Kakamega Forest, may represent a species complex. However, the DNA
islands: priority sites for conservation. Pisces Publications and BirdLife International, barcode for specimens from the type locality in Angola is not available.
Newbury and Cambridge, UK. © Stefan Lötters
Gregory, T.R. 2005. DNA barcoding does not compete with taxonomy. Nature 434:1067.
Herbert, P.D.N., Cywinska, A., Ball, S.L. and de Waard, J.R. 2003. Biological identifications
through DNA barcodes. Proceedings of the Royal Society of London B 270:313-321.
Köhler, J., Vieites, D.R., Bonett, R.M., Garcia, F.H., Glaw, F., Steinke, D. and Vences, M.
2005. New amphibians and global conservation: a boost in species discoveries in a
highly endangered vertebrate group. BioScience 55:693-696.
Lötters, S., Schmitz, A. and Reichle, S. 2005. A new cryptic species of poison frog from the
Bolivian Yungas (Dendrobatidae: Epipedobates). Herpetozoa 18:115-124.
Lötters, S., Rotich, D., Koester, T.E., Kosuch, J., Muchai, V., Scheelke, K., Schick, S.,
Teege, P., Wasonga, D.V. and Veith, M. 2006. What do we know about the amphib-
ians of the Kenyan central and western highlands? A faunistic and taxonomic review.
Salamandra 42:165-179.
Meyer C.P. and Paulay, G. 2005. DNA barcoding: Error rates based on comprehensive
sampling. PLoS Biology 3:e422.
Rubinoff, D. 2006. Utility of mitochondrial DNA barcodes in species conservation. Con-
servation Biology 20:1026-1033.
Savolainen, V., Cowan, R.S., Vogler, A.P., Roderick, G.K. and Lane, R. 2005. Towards writing
the encyclopedia of life: an introduction to DNA barcoding. Philosophical Transactions
of the Royal Society of London B 360:1805-1811.
Vences, M., Thomas, M., van der Meijden, A., Chiari, Y. and Vieites, D.R. 2005. Compara-
tive performance of the 16S rRNA gene in DNA barcoding of amphibians. Frontiers
in Zoology 2:5. DNA barcoding is a powerful tool for linking different life stages, e.g. frog
tadpoles to adults. In this case, the photograph shows a typical Xenopus
larva collected at Kakamega Forest from where only one Xenopus species
1 http://www.ncbi.nih.gov, accessed 1 May 2006 is known. © Stefan Lötters

ESSAY 11.12. THE AMPHIBIAN SPECIALIST GROUP: PAST, PRESENT AND FUTURE

The Global Amphibian Assessment (GAA) sharpened the scientific com-
munity’s focus on both the global nature and extent of amphibian declines
and extinctions (Stuart et al. 2004). The results of the GAA demonstrated
that the amphibian crisis is complex in nature and requires a comprehen-
sive set of solutions. While traditional approaches to conservation will
continue to form the core of a global strategy, novel approaches will also
be required to address threats such as emerging infectious diseases and
global climate change.
Under the auspices of the World Conservation Union (IUCN) and
Conservation International (CI), an International Amphibian Summit was
convened in September 2005 in Washington, DC, to devise a strategic
global plan of action for amphibian conservation in the form of the Am-
phibian Conservation Action Plan (ACAP). Prior to the Summit, the IUCN
Species Survival Commission (IUCN/SSC) focused on promoting amphibian
conservation through the Global Amphibian Specialist Group (GASG), on
decline-related research through the Declining Amphibian Populations
Task Force (DAPTF; see Essay 11.1), and on monitoring and assessments
through an informal network of researchers contributing data to the GAA.
These programmes have accomplish a great deal, but the magnitude of
the current crisis requires a coordinated, unified approach to amphibian
conservation, research and assessment that is beyond the scope of any
of these individual initiatives (Mendelson et al. 2006). Recognizing this,
a decision was made to merge the GASG, DAPTF and the GAA to form a
single body committed to implementing a global strategy for amphibian
conservation: the Amphibian Specialist Group (ASG).
The logo of the Amphibian Specialist Group
The ASG is a network of conservation professionals working under the
IUCN/SSC framework with the mission of enabling amphibian research and
conservation worldwide. The ASG envisions taking IUCN’s Specialist Group
model to the next level of effectiveness by establishing a Secretariat that
will coordinate regional working groups and a global web of stakeholders
to leverage their intellectual, institutional, and financial capacity. The Sec-
retariat has two Co-Chairs (currently Claude Gascon and James Collins), an
Executive Officer, and Conservation, Research, Assessment, Development
and Communications Divisions, each headed by a director.
The Conservation Division promotes and supports the conservation of
threatened amphibian species globally. By strengthening partnerships among
institutions in developed and developing countries, and by forging links with
groups such as other IUCN/SSC Specialist Groups, the ASG builds regional
capacity to ensure that amphibians are incorporated into conservation plans.
Through a collaborative partnership with Arizona State University, amphib-
ian conservation and research projects in priority regions are supported by
grants through the Critical Ecosystems Partnership Fund (CEPF), based at
Conservation International. Additionally, the ASG also supports conservation
and research through the allocation of Seed Grants, building upon the suc-
cessful model established by DAPTF. An Amphibian Conservation Award will
be presented annually to recognize an individual who has made significant
contributions to amphibian conservation worldwide.
The Research Division works closely with research partners from the
global network of national and regional working groups to apply a strategic
and coordinated approach to research on the causes of global amphibian
declines and to disseminating the results. Many amphibian declines and
extinctions are caused by complex interactions of factors such as disease
and climate change, and understanding these processes is vital for effective
conservation action. The Research division works to narrow knowledge gaps
through targeted research agendas for poorly known regions and groups, and
strives to apply scientific research results to conservation. A central on-line
storehouse of publications and current research results is being developed
to facilitate information exchange amongst researchers and conservation
practitioners. Information will also be presented in multiple languages to
facilitate knowledge-sharing. The ASG will support targeted field research
in poorly known regions of the world and link global expertise with local
expertise in developing countries by conducting training courses for field
practitioners and students. These actions will strengthen regional and
country-based capacity to improve knowledge of the taxonomy, distribution,
abundance, and causes of amphibian declines.
The Assessment Division – which fulfills the ASG’s Red List Authority
responsibilities – will regularly assesses the conservation status of every
amphibian species in the world through the GAA. The ASG aims to make the
GAA a more proactive process based on a “bottom up” flow of data to ensure
the most up-to-date and accurate information on every amphibian species.
The Assessment Division will undertake periodic analysis of the database
and communicate the findings to the global network and beyond. Findings
will be distributed on CDs to those regions where internet access is poor or
unavailable. Information will be disseminated in paper where appropriate via
working groups through workshops and meetings. The Assessment division
will regularly assess the conservation status of every amphibian species
and feed this information into on-the-ground identification of conservation
targets and priorities.
In coordination with the directors of the Conservation, Research and
Assessment divisions, regional working group heads will initiate projects in
their respective regions. To aid in the outreach and fundraising aspects of
the ASG network, an Advisory Board has been established comprising key
stakeholders and advisor-advocates representing policymakers, the media,
business, academia, conservation organizations, and government agencies.
ASG communications will include a new and expanded bi-monthly newsletter
(Froglog) that will be available free of charge on the ASG website (www.
amphibians.org).
The ASG will ensure long-term sustainability of amphibian research and
conservation by building on DAPTF’s worldwide network of expert working
groups and integrating them into IUCN’s global network of Specialist Groups.
The ASG will support the development and dissemination of new tools and
best practices for adoption and application by a network of local, national,
and regional working groups. By mainstreaming amphibian conservation best
practices into the global web of international conservation programmes,
national planning processes, and local practice, the ASG will ensure the
sustainability of amphibian research and conservation.
Robin D. Moore, Claude Gascon and James P. Collins
References
Mendelson, J.R., III, Lips, K.R., Gagliardo, R.W., Rabb, G.B., Collins, J.P., Diffendorfer,
J.E., Daszak, P., Ibanez D, R., Zippel, K.C., Lawson, D.P., Wright, K.M., Stuart, S.N.,
Gascon, C., da Silva, H.R., Burrowes, P.A., Joglar, R.L., La Marca, E., Lotters, S., du
Preez, L.H., Weldon, C., Hyatt, A., Rodriguez-Mahecha, J.V., Hunt, S., Robertson,
H., Lock, B., Raxworthy, C.J., Frost, D.R., Lacy, R.C., Alford, R.A., Campbell, J.A.,
Parra-Olea, G., Bolanos, F., Domingo, J.J.C., Halliday, T., Murphy, J.B., Wake, M.H.,
Coloma, L.A., Kuzmin, S.L., Price, M.S., Howell, K.M., Lau, M., Pethiyagoda, R.,
Boone, M., Lannoo, M.J., Blaustein, A.R., Dobson, A., Griffiths, R.A., Crump, M.L.,
Wake, D.B. and Brodie, E.D., Jr. 2006. Biodiversity: Confronting Amphibian Declines
and Extinctions. Science 313:48.
Stuart, S.N., Chanson, J.S., Cox, N.A., Young, B.E., Rodrigues, A.S.L., Fischman, D.L. and
Waller, R.W. 2004. Status and trends of amphibian declines and extinctions worldwide.
Science 306:1783-1786.
Extinct and Extinct in the Wild
Amphibian Species
136 Threatened Amphibians of the World

EX Adenomus kandianus (Günther, 1872)

Order, Family: Anura, Bufonidae
Country Distribution: Sri Lanka (Extinct)
Geographic Range This species was endemic to Sri Lanka, and is known only from the general type locality of
“Ceylon” (= Sri Lanka). The scientific name suggests that it might have been collected in the vicinity of the city of
Kandy, central Sri Lanka. As the site of collection is unclear, the species has not been mapped.
Population It is known only from the type specimen. There have been no sightings since the original description
and the species is now believed to be extinct (Manamendra-Arachchi and Pethiyagoda 1998). The general area of
Kandy, where this frog is presumed to have occurred, has been well surveyed.
Habitat and Ecology While there is nothing known with certainty about the habitat and ecology of this species, it
presumably bred by larval development in water as do other members of Adenomus.
Major Threats Although the causes of the species extinction not known, if the original collection locality was truly
in the vicinity of Kandy, then it is quite likely that extensive urban development has destroyed any suitable habitat
(Manamendra-Arachchi and Pethiyagoda 1998).
Conservation Measures It was not recorded from any protected areas.
Bibliography: Günther, A. (1872), Manamendra-Arachchi, K. and Pethiyagoda, R. (1998)
Data Providers: Kelum Manamendra-Arachchi, Anslem de Silva

EX Atelopus ignescens (Cornalia, 1849)

Order, Family: Anura, Bufonidae Habitat and Ecology An inhabitant of humid montane forest, humid sub-páramo (high-altitude bushland), and
Country Distribution: Ecuador (Extinct) páramo (high-altitude grassland). This was a stream breeding species. Animals were recorded from disturbed habitats,
including modified grasslands, in the Inter-Andean valleys and residential areas close to Quito.
Major Threats It probably disappeared due to the synergistic effects of the disease chytridiomycosis-confirmed in this
species-and climatic change (local warming and droughts). Habitat loss and the introduction of predatory non-native
trout might also have contributed to some population declines, however these threats are unlikely to have caused
© Suzanne L. Collins / CNAH

the substantial decline of the species throughout its range.

Geographic Range This species ranged from Imbabura to Chimborazo and Bolívar Provinces, in the Inter-Andean
valleys and higher parts of the major Andean Cordilleras of Ecuador, at elevations of 2,800-4,200m asl.
Population There have been no records of this species in Ecuador since 1988, despite extensive searches. It was
formerly abundant along streams, but now appears to be extinct.
Conservation Measures The known range of this species overlapped with several protected areas, including:
Reserva Ecológica Cayambe-Coca; Reserva Ecológica Antisana; Parque Nacional Cotopaxi; Area Nacional de
Recreación el Boliche; Reserva Ecológica Los Illinizas; Parque Nacional Llanganates; and Reserva de Producción
Faunística Chimborazo in Ecuador.
Notes on taxonomy: Coloma, Lötters and Salas (2000) redefined, Atelopus ignescens, and restricted the species to the north-central
Andes of Ecuador. Populations in northern Ecuador (in Cachi Province), and adjacent southern Colombia (Narino Department), previously
assigned to Atelopus ignescens are now considered to belong to an undescribed species.
Bibliography: Coloma, L.A. (2002), Coloma, L.A., Lötters, S. and Salas, A.W. (2000), Cornalia, E. (1849), Gray, P. (1983), La Marca, E. et
al. (2005), Lötters, S. (1996), Pounds, J.A. et al. (2006), Ron, S.R. et al. (2003)
Data Providers: Santiago Ron, Luis A. Coloma, Stefan Lötters, William Duellman, Martín R. Bustamante, Wilmar Bolívar, Enrique La
Marca

EX Atelopus longirostris Cope, 1868

Order, Family: Anura, Bufonidae slope of the Cordillera Occidental, Valle del Cauca Department, in Colombia, refer to an unidentified, probably
Country Distribution: Ecuador (Extinct) un-named species.
Population The species appears to be extinct, with no records in Ecuador since 1989, despite extensive surveys in
suitable habitat.
Habitat and Ecology This was a terrestrial species of lowland and montane tropical rainforests. It was a stream
breeding species.
Major Threats The decline in Ecuador is unexplained, and is possibly due to the disease chytridiomycosis, although
this seems unusual given that the species was also found at lower elevations where chytridiomycosis is often not
considered to be a threat. Other possible factors contributing to the extinction of this species include general climate
change, pollution and habitat loss.
© John D. Lynch

Geographic Range This species was recorded from the Provinces of Esmeraldas, Imbabura, Cotopaxi, and Pichincha,
on the north-western versant of the Ecuadorian Andes, at elevations of 200-2,500m asl. Records from the western
Conservation Measures The range of this species overlapped with the Reserva Ecológica Cotacachi-Cayapas and
possibly reached the northern limit of the Reserva Ecológica Los Illinizas.
Notes on taxonomy: Lötters (1996) discussed controversy over the identity of frogs from Colombia assigned to this species by other authors.
Bibliography: Acosta-Galvis, A.R. (2000), Cannatella, D.C. (1981), Cope, E.D. (1868), La Marca, E. et al. (2005), Lötters, S. (1996),
Merino-Viteri, A. (2001), Pounds, J.A. et al. (2006), Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and Lynch, J.D. (1996)
Data Providers: Martín R. Bustamante, Wilmar Bolívar, Luis A. Coloma, Santiago Ron, Diego Cisneros-Heredia, Fernando Castro, Jose
Vicente Rueda, Stefan Lötters, Andrés Acosta-Galvis

EX Atelopus vogli Müller, 1934

Order, Family: Anura, Bufonidae Population This species is known only from the type series collected by C. Vogel in 1933. Extensive searches in
Country Distribution: Venezuela (Extinct) recent years have failed to find any animals, and the species is now believed to be extinct.
Habitat and Ecology The original habitat at the type locality, semi-deciduous humid forest, has been drastically
modified by repeated clearing and burning, and only a savannah-like environment remains. Details from the collection
series suggest that the species congregated for breeding in small streams.
© Ted Kahn, from Ranas Arleqines, Conservation International

Major Threats The species is believed to have become extinct following the extensive conversion of its habitat
to agricultural use.
Conservation Measures This species has not been recorded from any protected areas.
Notes on taxonomy: This species was recently recognized as distinct from Atelopus cruciger by Lötters, La Marca and Vences
(2004).
Bibliography: Barrio Amorós, C.L. (2004), La Marca, E. et al. (2005), Lötters, S. (1996), Lötters, S., La Marca, E. and Vences, M. (2004),
Pounds, J.A. et al. (2006)
Data Providers: Stefan Lötters, Enrique La Marca, Miguel Vences

Geographic Range This species is known

only from the type locality of “Schluch Las
Peñas (600m), unweit von Maracay”. This
area is now believed to be Pozo del Diablo en
las cabeceras del Río Güey, on the southern
versant of the Cordillera de la Costa, State
of Aragua, Venezuela at 700m asl.
 Extinct and Extinct in the Wild Amphibian Species 137

EW Bufo baxteri Porter, 1964 WYOMING TOAD

Order, Family: Anura, Bufonidae 4 adults (USFWS 2002e). Limited natural reproduction and recruitment of a few metamorphosed juveniles occurred
Country Distribution: United States of America in 2002 (USFWS 2002e). Despite releases of captive-reared individuals, the population at Mortenson Lake NWR
(Extinct in the Wild) appears to be declining (USFWS 2002e).
Current Population Trend: Decreasing Habitat and Ecology Historically, it is associated with floodplain ponds along the Big and Little Laramie Rivers; use
of lakes might have been limited due to saline conditions; irrigation might have flushed out the lakes and made them
more suitable for toads (G. Baxter pers. comm.). Currently, it occurs in the vicinity of lakes and adjacent meadows. It
uses rodent burrows for shelter. Eggs and larvae develop in shallow water.
Major Threats Mortenson Lake, site of the only known extant population, is infected with the amphibian chytrid

© Suzanne L. Collins / CNAH

Geographic Range This species is restricted to the Laramie Basin, Wyoming, USA. The historical range extent was
approximately 2,330km2 (USFWS data). As of 2002, it was extant only at Mortenson Lake National Wildlife Refuge,
which encompasses four impoundments: Mortenson, Garber, Gibbs, and Soda lakes; the refuge is closed to the public
(USFWS 2002e). The extant occurrence probably would be extirpated without recent annual releases of captive-reared
toadlets. Populations at former re-introduction sites (Lake George on the Hutton Lake National Wildlife Refuge, and
Rush Lake) have been lost due to drought (USFWS 2002e).
Population This species was common in the 1950s, but underwent a large decline in the 1960s and 1970s; it was
thought to be extinct in the wild in the mid-1980s, but was found again at Mortenson Lake in 1987 and captive
propagation began in the mid-1990s using toads from Mortenson Lake (USFWS 2002e). At Mortenson Lake, there has
been no natural reproduction by wild toads since 1991; the population is maintained through release of captive-reared
young (Parker, Anderson and Lindzey 2000). In June 2002, a survey at Mortenson Lake NWR yielded 124 yearlings and
fungus (USFWS 2002e). This fungus has been implicated in declines and extinctions of amphibian species worldwide.
Retrospective analysis shows that the fungus has been present at Mortenson Lake since at least 1989. In addition,
chytridiomycosis is the most commonly seen disease in the captive population. Predation, pesticide use, irrigation
practices, and lack of genetic diversity might also limit the abundance of Wyoming Toads in the Laramie Basin. The
cause of the original decline remains unknown but might be associated with the invasion of chytrid into the area (USFWS
2002e). Mortenson Lake recently has become more saline (and less suitable for toads) as a result of drought-related
increases in evaporation (USFWS 2002e). Salt-cedar was found (and removed) at Hutton Lake NWR in 2002. This plant
has the potential to reduce habitat suitability (USFWS 2002e). For the past several years, beginning in late summer,
adults with bacterial and fungal infections have been found moribund or dead (Taylor et al. 1999). Predators, probably
mustelids, killed several radio-tagged individuals in 1998 (Parker, Anderson and Lindzey 2000).
Conservation Measures It occurs in Mortenson Lake National Wildlife Refuge, where a recovery programme,
using captive-bred animals and re-introductions, is being implemented. TNC recently acquired a 1,800-acre tract
at Mortensen Lake and has arranged a conservation easement with an adjacent landowner. Lake George also is
a toad refuge. The success of the recovery programme probably depends on finding some way to combat chytrid-
iomycosis in the wild.
Bibliography: Baxter, G.T. and Stone, M.D. (1980), Blackburn, L., Nanjappa, P. and Lannoo, M.J. (2001), Collins, J.T. (1991), Dickerson,
K. (1999), Dowling, H.G. (1993), Green, D.M. (1983), Lewis, D.L. et al. (1985), Matthews, J.R. and Moseley, C.J. (eds) (1990), Parker, J.,
Anderson, S.H. and Lindzey, F.J. (2000), Smith, H.M. et al. (1998), Spencer, B. (1999), Stebbins, R.C. (1985b), Taylor, S.K. et al. (1999),
U.S. Fish and Wildlife Service (1990a), U.S. Fish and Wildlife Service (2002e)
Data Providers: Geoffrey Hammerson

EX Bufo periglenes Savage, 1967 GOLDEN TOAD

Order, Family: Anura, Bufonidae Geographic Range This species was known only from the Reserva Biológica Monteverde, Costa Rica, at elevations
Country Distribution: Costa Rica (Extinct) of 1,500-1,620m asl.
CITES: Appendix I Population Formerly a common species, no specimen has been seen since 1989. It last bred in normal numbers in
1987, and its breeding sites were well known. In 1988, only eight males and two females could be located. In 1989,
a single male was found, and was the last record of the species. Extensive searches since this time have failed to
produce any more records.
Habitat and Ecology It lived in cloud and elfin forest, and bred in temporary pools at the beginning of the rainy
season.
Major Threats Its restricted range, global warming, chytridiomycosis and airborne pollution probably contributed
to this species’ extinction.
Conservation Measures Its entire range was protected by the Reserva Biológica Monteverde.
© Chris Perrins

Bibliography: Crump, M.L. (1989), Crump, M.L., Hensley, F.R. and Clark, K.L. (1992), Jacobson, S.K. and Vandenberg, J.J. (1991),
Pounds, J.A. et al. (1997), Pounds, J.A. and Crump, M.L. (1994), Pounds, J.A., Fogden, M.P.L. and Campbell, J.H. (1999), Savage, J.M.
(1966), Savage, J.M. (2002)
Data Providers: Alan Pounds, Jay Savage

EX Discoglossus nigriventer Mendelssohn and Steinitz, 1943 HULA PAINTED FROG

Order, Family: Anura, Discoglossidae Geographic Range This species was recorded from only two localities on the eastern shore of Lake Huleh, Israel.
Country Distribution: Israel (Extinct) It seems possible that it could once also be found in adjacent parts of Syrian Arab Republic. The map indicates the
© Heinrich Mendelssohn, Tel-Aviv University

possible former range.

Population It is known only from a few specimens and is now believed to be extinct. The single adult collected in
1955 represents the last confirmed record of this species.
Habitat and Ecology The species was reported to occur in marginal freshwater habitats within the Lake Huleh
wetlands of Israel. It was presumably a larval developing species.
Major Threats The Huleh marshes were drained in the 1950s in an attempt to both eradicate malaria and to make
the land suitable for agricultural use. Of the original 6,000ha of marshland, only 300ha remained after drainage. While
this remaining wetland was set aside as a nature reserve in 1964 (at a considerable distance from the recorded
localities of Discoglossus nigriventer) it seems that this action was too late to save the species.
Conservation Measures The species remains protected by national legislation in Israel. Surveys of potentially
suitable habitat in the Aammiq marsh of nearby south-east Lebanon in April 2004 and April 2005 failed to locate
any animals (Tron 2005). The Aammiq marsh is the only remaining wetland fragment of the Bekaa valley, which was
once a major wetland of the Middle East but has been drastically reduced by conversion to agricultural use and
urban development.
Bibliography: Fromhage, L., Vences, M. and Veith, M. (2004), Honegger, R.E. (1981), Honegger, R.E. (compiler) (1979), Mendelssohn, H.
and Steinitz, H. (1943), Steinitz, H. (1955), Tron, F. (2005)
Data Providers: Theodore Papenfuss, Ahmad Disi, Steven Anderson, Sergius Kuzmin, Avital Gasith, Riyad A. Sadek, Yehudah Werner

EX Phrynomedusa fimbriata Miranda-Ribeiro, 1923

Order, Family: Anura, Hylidae Geographic Range This species has only been recorded from Bibliography: Cruz, C.A.G. (1982), Cruz, C.A.G. (1985), Cruz, C.A.G. (1990), Cruz, C.A.G. (1991)
Country Distribution: Brazil (Extinct) “Alto da Serra”, Paranapiacaba, Santo Andre, in the State of Sao Data Providers: Carlos Alberto Gonçalves da Cruz, Bruno Pimenta
Paulo, south-eastern Brazil. It was collected at an elevation of
around 1,000m asl.
Population It remains known from the holotype only, with no ad-
ditional records for more than 80 years, despite repeated searches.
It is now believed to be extinct.
Habitat and Ecology Although there is no information available
on the biology or ecology of this species, it is possible that it was a
high-altitude stream-breeder.
Major Threats No explanation is currently available for the disap-
pearance of this species.
Conservation Measures There are no protected areas near the
type locality of this species.
138 Threatened Amphibians of the World
EX Craugastor chrysozetetes (McCranie, Savage and Wilson, 1989)
Order, Family: Anura, Leptodactylidae
Country Distribution: Honduras (Extinct)
© James McCranie
EX Craugastor milesi (K. Schmidt, 1933)
Order, Family: Anura, Leptodactylidae
Country Distribution: Honduras (Extinct)
© James McCranie
EX Taudactylus diurnus Straughan and Lee, 1966
Order, Family: Anura, Myobatrachidae
Country Distribution: Australia (Extinct)
© Hal Cogger
Geographic Range This species, an Australian endemic, occurred in disjunctive populations in three sub-coastal
mountain ranges (Blackall, Conondale, and D’Aguilar Ranges) in the south-east Queensland region from Coonoon
Gibber Creek in the north to Mount Glorious in the south (Czechura and Ingram 1990; Hines, Mahony and McDonald
1999). The extent of occurrence of the species was about 1,400km2 (map in Hines, Mahony and McDonald 1999).
Taudactylus diurnus occurred over a relatively narrow altitudinal range of 350-800m asl with most records falling
between 500-800m asl (Czechura and Ingram 1990).
Population In the early 1970s it was considered to be relatively common (McEvoy, McDonald and Searle 1979), but
it has not been sighted in the wild since 1979 despite continued efforts to relocate the species (Hines, Mahony and
McDonald 1999). The disappearance of the species occurred over a period of three to four years, disappearing from
the D’Aguilar Range in late 1975, then from the Blackall Range in late 1978, and finally from the Conondale Range in
early 1979 (Czechura and Ingram 1990). There is no information on population size, structure, genetics or dynamics
(Hines, Mahony and McDonald 1999). This species is now believed to be extinct.
EX Nannophrys guentheri Boulenger, 1882
Order, Family: Anura, Ranidae
Country Distribution: Sri Lanka (Extinct)
Geographic Range This species is known only from the general type locality of “Ceylon” (= Sri Lanka). It is not
possible to produce a meaningful distribution map for the species.
Population Known only from the type series collected more than a 100 years ago. It has not been recorded since it
was originally described. Recent and extensive field surveys of the amphibian fauna of Sri Lanka have been unable
to relocate this frog, and it is now believed to be extinct.
Geographic Range This species was known only from Quebrada de Oro in the Río Viejo, south-east of La Ceiba,
Department of Atlantida, northern Honduras, at 880-1,130m asl.
Population It was always extremely rare, and is now believed to be extinct. Several visits to the only known site in
recent years have failed to locate the species.
Habitat and Ecology Animals were recorded along streams in premontane wet forest. It was presumably a direct
developing species.
Major Threats The species is believed to have disappeared through a combination of threats including deforestation
resulting from agricultural encroachment, human settlement of the region, logging, fires and landslides. The area from
which this species was recorded is subject to extensive landsides; these often severely impact the species habitat.
It should be additionally noted that all streamside Craugastor above 900m asl have disappeared in Honduras in a
manner that is consistent with the disease chytridiomycosis.
Conservation Measures Although the species has not been recorded from a protected area, the Quebrada de Oro
is at the edge of Parque Nacional Pico Bonito.
Notes on taxonomy: This species was previously included in the genus Eleutherodactylus (Crawford and Smith 2005).
Bibliography: Crawford, A.J. and Smith, E.N. (2005), McCranie, J.R. and Wilson, L.D. (2002b), McCranie, J.R., Savage, J.M. and
Wilson, L.D. (1989)
Data Providers: Gustavo Cruz, Larry David Wilson, Randy McCranie
Geographic Range This species was known from west and north-west Honduras. It was recorded from the Cerro
Azul in Copan Departmento, and from Montana del Cusuco (National Park) and Montana del Merendon (west of San
Pedro Sula) in the Departmento of Cortes, within the Sierras of Espiritu Santo and Omoa. Animals were reported
from elevations of 1,050-1,720m asl.
Population It was formerly relatively common, but underwent a precipitous decline. Repeated attempts to relocate
this frog in appropriate habitat and weather conditions in Parque Nacional Cusuco between 1992 and 1996 were
unsuccessful. It is now considered to be extinct.
Habitat and Ecology This was a leaf-litter species found along streams in premontane and lower montane wet
forest. The species bred by direct development.
Major Threats While the species was clearly threatened habitat loss and degradation (largely resulting from the
conversion of forest to subsistence agricultural use), this does not explain the sudden disappearance of populations
from areas of pristine forest. It seems probable that factors related to declines in other montane frog species, such
as disease chytridiomycosis, also contributed to the species demise.
Conservation Measures This species has been recorded within the Parque Nacional Cusuco and Parque Nacional
Cerro Azul.
Notes on taxonomy: This species was previously included in the genus Eleutherodactylus (Crawford and Smith 2005).
Bibliography: Campbell, J.A. (1994b), Crawford, A.J. and Smith, E.N. (2005), McCranie, J.R. and Wilson, L.D. (1997a), McCranie, J.R.
and Wilson, L.D. (2002b), McCranie, J.R., Savage, J.M. and Wilson, L.D. (1989), Savage, J.M. (2000), Schmidt, K.P. (1933)
Data Providers: Gustavo Cruz, Larry David Wilson
MOUNT GLORIOUS TORRENT FROG
Habitat and Ecology Taudactylus diurnus was associated with permanent and temporary watercourses in montane
rainforests, tall open forest, notophyll vine forest and sclerophyll fern forest (Czechura and Ingram 1990). In addition,
animals were also found along watercourses in pure stands of the palm Archontophoenix cunninghamia, in exposed
areas, in gorges, in dense non-forest riparian vegetation (Lomandra longifolia, Carex neuroclamys, Elastostema
reticulatum and Blechnum nudum) and where the riparian vegetation had been slightly infested with Lantana camara
(Czechura and Ingram 1990). Permanent streams with rocky substrates were favoured, but this species also occurred
in permanent and ephemeral streams on gravel, clay, sand and soil substrates (Czechura and Ingram 1990). Active
frogs have been observed all year round, although less frequently during winter months (Czechura and Ingram 1990).
Breeding occurred in warm weather after or during heavy rain from late October to May, with a January to March
peak (Czechura and Ingram 1990; Meyer, Hines and Hero 2001d). Gravid females have been reported between No-
vember and May (Straughan and Lee 1966). Amplexus is inguinal and 24-36 eggs (2.2mm diameter) are deposited in
gelatinous clumps under rocks or branches in the water (Liem and Hosmer 1973; Watson and Martin 1973; Czechura
and Ingram 1990; Meyer, Hines and Hero 2001d). Tadpoles, illustrated by Liem and Hosmer (1973) and Watson and
Martin (1973), were found throughout the year.
Major Threats The reason(s) for the disappearance of this species remains unknown. Like Rheobatrachus silus,
logging has occurred in catchments occupied by the species (Hines, Mahony and McDonald 1999); however, the effect
of timber harvesting on the species has not been investigated. The species’ habitat is currently threatened by feral
pigs, invasion of weed species (especially mist flower) and altered stream flow and water quality due to upstream
disturbances (Hines, Mahony and McDonald 1999). Taudactylus diurnus was not found in areas along watercourses that
were heavily infested with Lantana camara or where the weeds Baccharis halimifilia and Agertina riparia (mist flower)
occurred (Czechura and Ingram 1990). The frogs were also absent from streams with very muddy water associated with
the activities of feral pigs (Czechura and Ingram 1990). From what is known from similar declines and disappearance
elsewhere in the world, the disease chytridiomycosis also must be suspected as a cause for the decline.
Conservation Measures The species’ habitat is fully protected within a National Park.
Bibliography: Czechura, G.V. and Ingram, G. (1990), Hines, H., Mahony, M. and McDonald, K. (1999), Ingram, G.J. (1980), Liem, D.S.
and Hosmer, W. (1973), McEvoy, J.S., McDonald, K.R. and Searle, A.K. (1979), Meyer, E., Hines, H. and Hero, J.-M. (2001d), Straughan,
I.R. and Lee, A.K. (1966), Tyler, M.J. (1997), Watson, G.F. and Martin, A.A. (1973)
Data Providers: Jean-Marc Hero, Sarah May, David Newell, Harry Hines, John Clarke, Ed Meyer
Habitat and Ecology There is no information on the habitat requirements of this species. It presumably bred on
wet rock surfaces near streams, like other members of the genus.
Major Threats The reasons for the extinction of this species are not known.
Conservation Measures It has not been recorded from any protected areas.
Notes on taxonomy: The five available museum specimens require further comparison with specimens of Nannophrys ceylonensis.
Bibliography: Clarke, B.T. (1983), Dutta, S.K. (1997), Dutta, S.K. and Manamendra-Arachchi, K. (1996)
Data Providers: Kelum Manamendra-Arachchi, Anslem de Silva
 Extinct and Extinct in the Wild Amphibian Species 139

EX Rana fisheri Stejneger, 1893 LAS VEGAS LEOPARD FROG

Order, Family: Anura, Ranidae
Country Distribution: United States of America (Extinct)
Geographic Range This species was known from a small number
of localities, elevation ca. 600m asl, in the northern portions of Las
Vegas Valley, Clark County, Nevada, USA (Jennings, Riddle and
Bradford 1995).
Population It was last seen in 1942 (Wright and Wright 1949) and is
now believed to be extinct (Jennings, Riddle and Bradford 1995).
Habitat and Ecology This frog was restricted to freshwater streams,
springs, seeps, and adjacent riparian habitat associated with the
Upper Las Vegas Valley (Wright and Wright 1949). Egg masses are
not known, but metamorphic individuals were collected in the same
habitats as those used by adults (Wright and Wright 1949).
Major Threats It is extinct evidently due to habitat loss resulting from spring capture and ground water pumping by the
growing city of Las Vegas (URS 1977), and exacerbated by the introduction of the Bullfrog Rana catesbeiana. Although
some suitable habitat persists within or near the former range of this species, only R. catesbeiana can be found.
Conservation Measures No conservation measures are needed; this species is extinct.
Notes on taxonomy: Since its description, Rana fisheri (Stejneger 1893) has been considered a distinct species (Linsdale 1940; Jennings,
Riddle and Bradford 1995), a subspecies of R. pipiens (Stebbins 1959) or a synonym of R. onca (Jennings 1988; Stebbins 2003). Morphological
analyses support the view that R. fisheri represents a separate species (Jennings, Riddle and Bradford 1995).
Bibliography: Behler, J.L. and King, F.W. (1979), Blackburn, L., Nanjappa, P. and Lannoo, M.J. (2001), Frost, D.R. (1985), Jennings, M.R.
(1988), Jennings, R.D., Riddle, B.R. and Bradford, D. (1995), Linsdale, J.M. (1940), Stebbins, R.C. (1954), Stebbins, R.C. (1959), Stebbins,
R.C. (1985b), Stebbins, R.C. (2003), Stejneger, L. (1893), Wright, A.H. and Wright, A.A. (1949)
Data Providers: Randy Jennings, Geoffrey Hammerson

EX Philautus adspersus (Günther, 1872)

Order, Family: Anura, Rhacophoridae Geographic Range This extinct Sri Lankan species is known only from the non-specific type locality of “Ceylon”
Country Distribution: Sri Lanka (Extinct) (not “Peradeniya” as sometimes stated) and from Nuwara Eliya, a resort town among mountains at 1,700-2,500m
asl (Manamendra-Arachchi and Pethiyagoda 2005). Only the site of Nuwara Eliya has been mapped here. Reports
of this frog occurring in India (Krishnamurthy and Sakunthala 1993) are believed to be erroneous (S.K. Dutta pers.
comm. 2002).
Population It is known only from two specimens, and was last collected around 1886. It is now thought to be extinct
© Madhava Meegaskumbura

since recent extensive field surveys of the amphibian fauna of Sri Lanka have failed to rediscover this frog.
Habitat and Ecology The habitat requirements of this species are not known. It presumably bred by direct develop-
ment.
Major Threats The threats that led to the extinction of this species are not known, though habitat loss through
conversion of land to agricultural use seems the most likely cause.
Conservation Measures It is not known to have occurred in any protected areas.
Bibliography: Bossuyt, F. and Dubois, A. (2001), Krishnamurthy, S.V. and Sakunthala, K. (1993), Manamendra-Arachchi, K. and
Pethiyagoda, R. (2005)
Data Providers: Kelum Manamendra-Arachchi, Sushil Dutta

EX Philautus dimbullae (Shreve, 1940)

Order, Family: Anura, Rhacophoridae Geographic Range This species is a Sri Lankan endemic, known only from the type locality of “Queenwood Estate,
Country Distribution: Sri Lanka (Extinct) Dimbulla, Ceylon”, at 1,500m, where it was collected in 1933 (Manamendra-Arachchi and Pethiyagoda 2005).
Population It is known only from the holotype. There have been no records since its original collection and the
species is now believed to be extinct because recent, extensive field surveys of the amphibian fauna of Sri Lanka,
including at the type locality, have not rediscovered this frog.
Habitat and Ecology The habitat requirements of this species are not known. It presumably bred by direct develop-
ment.
© Wildlife Heritage Trust

Major Threats The threats that resulted in the extinction of this species are not known, although presumably habitat
loss was a contributing factor.
Conservation Measures It has not been recorded from any protected areas.
Notes on taxonomy: This species is considered distinct from Philautus microtympanum, following Manamendra-Arachchi and
Pethiyagoda (2005).
Bibliography: Bossuyt, F. and Dubois, A. (2001), Dutta, S.K. and Manamendra-Arachchi, K. (1996), Manamendra-Arachchi, K. and
Pethiyagoda, R. (2005)
Data Providers: Kelum Manamendra-Arachchi, Anslem de Silva

EX Philautus eximius Shreve, 1940

Order, Family: Anura, Rhacophoridae Geographic Range This species is endemic to Sri Lanka, and is known only from the type locality of “Queenwood
Country Distribution: Sri Lanka (Extinct) Estate, Dimbulla, Ceylon”, at 1,500m asl, where it was collected in 1933 (Manamendra-Arachchi and Pethiyagoda
2005).
Population This species is known only from the holotype. There have been no sightings since 1933 and the species
is believed to now be extinct, because recent, extensive field surveys of the amphibian fauna of Sri Lanka, including
at the type locality, have not rediscovered this frog.
Habitat and Ecology The habitat requirements of this species are not known. It presumably bred by direct develop-
© Wildlife Heritage Trust

ment.
Major Threats The threats that resulted in the extinction of the species are not known, although presumably habitat
loss was a contributing factor.
Conservation Measures It has not been recorded from any protected areas.
Bibliography: Bossuyt, F. and Dubois, A. (2001), Manamendra-Arachchi, K. and Pethiyagoda, R. (2005)
Data Providers: Kelum Manamendra-Arachchi, Anslem de Silva

EX Philautus extirpo Manamendra-Arachchi and Pethiyagoda, 2005

Order, Family: Anura, Rhacophoridae Habitat and Ecology There is no information about the habitat requirements of this species. It presumably bred
Country Distribution: Sri Lanka (Extinct) by direct development.
Major Threats The threats that led to the extinction of this species are not known, though habitat loss seems the
Geographic Range This recently described species was originally collected in 1882 from an unspecified locality most likely cause.
on Sri Lanka (Manamendra-Arachchi and Pethiyagoda 2005). It is not possible to produce a meaningful distribution Conservation Measures It has not been recorded from any protected areas.
map for this species. Bibliography: Manamendra-Arachchi, K. and Pethiyagoda, R. (2005)
Population There have been no records since 1882, and it is presumed to be extinct, because extensive searches Data Providers: Kelum Manamendra-Arachchi, Rohan Pethiyagoda
have failed to rediscover this species.
140 Threatened Amphibians of the World

EX Philautus halyi Boulenger, 1904

Order, Family: Anura, Rhacophoridae Geographic Range This species is endemic to Sri Lanka, and is known only from the type locality of “Pattipola,
Country Distribution: Sri Lanka (Extinct) Ceylon [= Sri Lanka]”, where it was collected in 1899 (Manamendra-Arachchi and Pethiyagoda 2005).
Population It is known only from the holotype, and there have been no sightings since 1899. The species is believed
to now be extinct, because recent extensive field surveys of the amphibian fauna of Sri Lanka have not rediscovered
this frog.
Habitat and Ecology The habitat requirements of the species are not known. It presumably bred by direct develop-

© Madhava Meegaskumbura
ment.
Major Threats The causes of the extinction are not known, but presumably habitat loss was a contributing factor.
Conservation Measures It has not been recorded from any protected areas.
Notes on taxonomy: This species is considered distinct from Philautus leucorhinus following Manamendra-Arachchi and Pethiyagoda
(2005).
Bibliography: Manamendra-Arachchi, K. and Pethiyagoda, R. (2005)
Data Providers: Kelum Manamendra-Arachchi, Anslem de Silva

EX Philautus hypomelas (Günther, 1876)

Order, Family: Anura, Rhacophoridae
Country Distribution: Sri Lanka (Extinct)
Population It is known only from the type series. There have been no sightings of the species since it was described
Geographic Range This species is en- in 1876, and it is now believed to be extinct, because recent, extensive field surveys of the amphibian fauna of Sri
demic to Sri Lanka, and is known only Lanka have not rediscovered this frog.
from the imprecise type locality of “Ceylon Habitat and Ecology The habitat requirements of the species are not known. It presumably bred by direct develop-
© Madhava Meegaskumbura

[=Sri Lanka]” (Manamendra-Arachchi and ment.

Pethiyagoda 2005). It is not possible to map Major Threats The causes of the species’ extinction are not known, but presumably habitat loss was a contribut-
this species because the exact type locality ing factor.
is not known. Conservation Measures It has not been recorded from any protected areas.
Bibliography: Bossuyt, F. and Dubois, A. (2001), Manamendra-Arachchi, K. and Pethiyagoda, R. (2005)
Data Providers: Kelum Manamendra-Arachchi, Anslem de Silva

EX Philautus leucorhinus (Lichtenstein and Martens, 1856)

Order, Family: Anura, Rhacophoridae Population It is known only from the type specimen (collected before description in 1856), and it is now believed
Country Distribution: Sri Lanka (Extinct) to be extinct, because recent, extensive field surveys of the amphibian fauna of Sri Lanka have failed to rediscover
this frog.
Geographic Range This species is known Habitat and Ecology The habitat requirements of this species are not known. It presumably bred by direct develop-
only from the type locality “Ceylon” (= Sri ment.
Lanka) (Dutta and Manmendra-Arachchi Major Threats The causes of the species’ extinction are not known, but presumably habitat loss was a contribut-
1996). The type locality is too general to ing factor.
© Wildlife Heritage Trust

allow the production of a meaningful map Conservation Measures It has not been recorded from any protected areas.
for this species. Notes on taxonomy: Specimens recorded as Philautus temporalis and P. leucorhinus from the Western Ghats of India are now considered
to be P. wynaadensis. P. temporalis and P. leucorhinus are endemic species of Sri Lanka (S.D. Biju and K. Manamendra-Arachchi pers.
comm.; Bossuyt and Dubois 2001).
Bibliography: Bossuyt, F. and Dubois, A. (2001), Dutta, S.K. (1997), Dutta, S.K. and Manamendra-Arachchi, K. (1996), Manamendra-Arachchi,
K. and Pethiyagoda, R. (2005)
Data Providers: Kelum Manamendra-Arachchi, S.D. Biju, Anslem de Silva

EX Philautus malcolmsmithi (Ahl, 1927)

Order, Family: Anura, Rhacophoridae
Country Distribution: Sri Lanka (Extinct)
Geographic Range This species is an extinct Sri Lankan endemic, known only from the type locality of “Ceylon”
(Manamendra-Arachchi and Pethiyagoda 2005). It is not possible to produce a distribution map for this species
because the type locality is too imprecise.
Population The species is known only from the holotype specimen. There have been no sightings for over 70 years
and the species is now believed to be extinct. Recent, extensive field surveys of the amphibian fauna of Sri Lanka
have not rediscovered this frog.
Habitat and Ecology The habitat requirements of this species are not known. It presumably bred by direct develop-
ment.
Major Threats The cause of the species’ extinction is not known, but presumably habitat loss was a contributing
factor to its demise.
Conservation Measures It has not been recorded from any protected areas.
Notes on taxonomy: We consider this species to be distinct from Philautus leucorhinus following Manamendra-Arachchi and
Pethiyagoda (2005).
Bibliography: Ahl, E. (1927), Bossuyt, F. and Dubois, A. (2001), Manamendra-Arachchi, K. and Pethiyagoda, R. (2005)
Data Providers: Kelum Manamendra-Arachchi, Anslem de Silva

EX Philautus nanus (Günther, 1869)

Order, Family: Anura, Rhacophoridae is now believed to be extinct. Recent, extensive field surveys of the amphibian fauna of Sri Lanka have failed to
Country Distribution: Sri Lanka (Extinct) rediscover this frog.
Habitat and Ecology The habitat requirements of this species are not known. It presumably bred by direct develop-
Geographic Range This species is an ment.
extinct Sri Lankan endemic, known only Major Threats The reason for the species’ extinction is not known, but presumably habitat loss was a contributing
from the type locality of “southern Ceylon factor.
© Madhava Meegaskumbura

[=Sri Lanka]” (Manamendra-Arachchi and Conservation Measures It has not been recorded from any protected areas.
Pethiyagoda 2005). It is not possible to Notes on taxonomy: This species was resurrected from the synonymy of Philautus microtympanum by Dutta and Manamendra-Arachchi
produce a distribution map for this species (1996).
because the exact location of the type locality Bibliography: Bossuyt, F. and Dubois, A. (2001), Dutta, S.K. and Manamendra-Arachchi, K. (1996), Manamendra-Arachchi, K. and
is not known. Pethiyagoda, R. (2005)
Population It is known only from the Data Providers: Kelum Manamendra-Arachchi, Anslem de Silva
lectotype. There have been no records since
the species was described in 1869, and it

EX Philautus nasutus (Günther, 1868)
Order, Family: Anura, Rhacophoridae
Country Distribution: Sri Lanka (Extinct)
Geographic Range This species is an
extinct Sri Lankan endemic, known only from
the general type locality of “Ceylon [=Sri
© Madhava Meegaskumbura
Lanka]” (Manamendra-Arachchi and Pethi-
yagoda 2005). It is not possible to produce
a distribution map for this species because
the exact location of the type locality is not
known. Specimens previously attributed
to this species from India (Krishnamurthi
and Shakunthala 1993) are now believed
to be a separate taxon (Biju 2001), and
EX Philautus oxyrhynchus (Günther, 1872)
Order, Family: Anura, Rhacophoridae
Country Distribution: Sri Lanka (Extinct)
Geographic Range This species is an
extinct Sri Lankan endemic, known only
from the type locality of “Ceylon” (Mana-
© Madhava Meegaskumbura
mendra-Arachchi and Pethiyagoda 2005).
Peradeniya is not the correct type locality (K.
Manamendra-Arachchi pers. comm.). It is not
possible to produce a meaningful distribution
map for this species.
EX Philautus rugatus (Ahl, 1927)
Order, Family: Anura, Rhacophoridae
Country Distribution: Sri Lanka (Extinct)
Geographic Range This species is endemic to Sri Lanka, and is known only from the non-specific type locality of
“Ceylon (Farmlands)” or “Taralanda” (Manamendra-Arachchi and Pethiyagoda 2005). It is not possible to produce a
distribution map for this species because the exact location of the type locality is not known.
Population It is known only from the holotype specimen. There have been no sightings since the species was
described in 1927, and it is now believed to be extinct. Recent, extensive field surveys of the amphibian fauna of Sri
Lanka have not rediscovered this frog.
EX Philautus stellatus (Kelaart, 1853)
Order, Family: Anura, Rhacophoridae
Country Distribution: Sri Lanka (Extinct)
EX Philautus temporalis (Günther, 1864)
Order, Family: Anura, Rhacophoridae
Country Distribution: Sri Lanka (Extinct)
Geographic Range This species is endemic
to Sri Lanka, where it is known only from the
non-specific type locality of “Ceylon” (Mana-
© Madhava Meegaskumbura
mendra-Arachchi and Pethiyagoda 2005). It
is not possible to produce a distribution map
for this species because the exact location of
the type locality is not known.
Extinct and Extinct in the Wild Amphibian Species
Geographic Range This species is known only from the type local-
ity of Nuwara Eliya (06º 57’N, 80º 47’E) [Newera-Elllia], in Sri Lanka
(Manamendra-Arachchi and Pethiyagoda 2005).
Population It is known only from the lost holotype. There have
been no records since the species was described in 1853, and it is
now believed to be extinct. Recent, extensive field surveys of the
amphibian fauna of Sri Lanka, including at the type locality, have
failed to rediscover this frog.
Habitat and Ecology The habitat requirements of this species are
not known. It presumably bred by direct development.
Major Threats The threats that resulted in the extinction of this
species are not known, although presumably habitat loss was a
major factor.
141
specimens reported by Karunaratne (1998) are believed to belong to a different species (K. Manamendra-Arachchi
pers. comm.).
Population It is known only from the holotype. There have been no records since the species was described in
1869, and it is now believed to be extinct. Recent, extensive field surveys of the amphibian fauna of Sri Lanka have
not rediscovered this frog.
Habitat and Ecology The habitat requirements of this species are not known. It presumably bred by direct develop-
ment.
Major Threats The cause for the species’ extinction is not known, but presumably habitat loss was a contributing
factor.
Conservation Measures It has not been recorded from any protected areas.
Bibliography: Biju, S.D. (2001), Bossuyt, F. and Dubois, A. (2001), Dutta, S.K. (1997), Dutta, S.K. and Manamendra-Arachchi, K. (1996),
Karunaratne, S. (1998), Krishnamurthy, S.V. and Sakunthala, K. (1993), Manamendra-Arachchi, K. and Pethiyagoda, R. (2005)
Data Providers: Kelum Manamendra-Arachchi, Anslem de Silva
Population The species is known only from the lectotype. There have been no sightings since the species was
described in 1872, and it is now believed to be extinct. Recent, extensive field surveys of the amphibian fauna of Sri
Lanka have not rediscovered this frog.
Habitat and Ecology The habitat requirements of this species are not known. It presumably bred by direct develop-
ment.
Major Threats The reason for the species’ extinction is not known, but presumably habitat loss was a contributing
factor.
Conservation Measures It has not been recorded from any protected areas.
Notes on taxonomy: This species is considered distinct from Philautus leucorhinus following Manamendra-Arachchi and Pethiyagoda
(2005).
Bibliography: Bossuyt, F. and Dubois, A. (2001), Dutta, S.K. (1997), Dutta, S.K. and Manamendra-Arachchi, K. (1996), Manamendra-Arachchi,
K. and Pethiyagoda, R. (2005)
Data Providers: Kelum Manamendra-Arachchi, Anslem de Silva
Habitat and Ecology The habitat requirements of this species are not known. It presumably bred by direct develop-
ment.
Major Threats The causes of the species’ extinction are not known, but presumably habitat loss was a contribut-
ing factor.
Conservation Measures It has not been recorded from any protected areas.
Notes on taxonomy: This species is considered distinct from Philautus leucorhinus (Günther, 1858) following Manamendra-Arachchi
and Pethiyagoda (2005).
Bibliography: Ahl, E. (1927), Bossuyt, F. and Dubois, A. (2001), Manamendra-Arachchi, K. and Pethiyagoda, R. (2005)
Data Providers: Kelum Manamendra-Arachchi, Anslem de Silva
Conservation Measures It has not been recorded from any protected areas.
Notes on taxonomy: This species is known only from the now lost holotype.
Bibliography: Bossuyt, F. and Dubois, A. (2001), Kelaart, E.F. (1853), Manamendra-Arachchi, K. and Pethiyagoda, R. (2005)
Data Providers: Kelum Manamendra-Arachchi, Rohan Pethiyagoda, Sushil Dutta, Anslem de Silva
Population It is known only from the lectotype and type series, and has not been recorded since the original descrip-
tion in 1864. Recent, extensive field surveys of the amphibian fauna of Sri Lanka have failed to rediscover this frog
and it is now considered to be extinct.
Habitat and Ecology The habitat requirements of this species are not known. It presumably bred by direct develop-
ment.
Major Threats The causes of the species’ extinction are not known, but presumably habitat loss was a contribut-
ing factor.
Conservation Measures It has not been recorded from any protected areas.
Notes on taxonomy: Specimens formerly attributed to Philautus temporalis and P. leucorhinus from the Western Ghats are now
correctly assigned to P. wynaadensis. P. temporalis and P. leucorhinus are now considered to be endemic species of Sri Lanka (S.D. Biju
and K. Manamendra-Arachchi pers. comm.).
Bibliography: Biju, S.D. (2001), Bossuyt, F. and Dubois, A. (2001), Dutta, S.K. (1997), Inger, R.F. et al. (1984), Manamendra-Arachchi,
K. and Pethiyagoda, R. (2005)
Data Providers: S.D. Biju, Kelum Manamendra-Arachchi
142 Threatened Amphibians of the World

EX Philautus travancoricus (Boulenger, 1891)

Order, Family: Anura, Rhacophoridae Geographic Range This species is known only from the type locality of “Bodanaikanur, Travancore, at the foot of the
Country Distribution: India (Extinct) hills on the eastern side” in the Western Ghats of India. The altitudinal range is reported to be up to 400m asl.
Population It is known only from the holotype specimen, and was described in 1891. Extensive recent field surveys of
the area, and surrounding suitable habitat, have not relocated this species, and it is now considered to be extinct.
Habitat and Ecology It is believed to have been an arboreal species of tropical moist evergreen forest. It is presumed
to have been a direct developing species, like other species of the genus.
Major Threats The extinction of the species was caused by deforestation through conversion of land to agricultural
use and urban development.
Conservation Measures It has not been recorded from any protected area.
Bibliography: Biju, S.D. (2001), Boulenger, G.A. (1891), Dutta, S.K. (1997)
Data Providers: S.D. Biju
© S.D. Biju

At the time of going to press we received reports of the rediscovery of this species in the Western Ghats of India (S.D. Biju pers.
comm.).

EX Philautus variabilis (Günther, 1858)

Order, Family: Anura, Rhacophoridae
Country Distribution: Sri Lanka (Extinct)
Habitat and Ecology The habitat requirements of this frog are not known. It presumably bred by direct develop-
Geographic Range This species is known ment.
only from the general type locality of “Cey- Major Threats The cause of the species’ extinction is not known, but habitat loss seeems likely to have been a
lon” (= Sri Lanka). It is not possible to produce contributing factor.
a distribution map for this species because Conservation Measures It has not been recorded from any protected areas.
© Madhava Meegaskumbura

the exact location of the type collection Notes on taxonomy: Specimens formerly attributed to Philautus variabilis from the Western Ghats of India are misidentified P.
is not known (Manamendra-Arachchi and wynaadensis. P. variabilis is endemic to Sri Lanka, where it is now considered to be extinct (K. Manamendra-Arachchi and S.D. Biju
Pethiyagoda 2005). pers. comm.).
Population This species is known only from Bibliography: Biju, S.D. (2001), Bossuyt, F. and Dubois, A. (2001), Dutta, S.K. (1997), Manamendra-Arachchi, K. and Pethiyagoda, R.
the lectotype. Recent, extensive field surveys (2005), Ravichandran, M.S. (1996a), Ravishankar, D. et al. (2001), Vasudevan, K., Kumar, A. and Chellam, R. (2001)
of the amphibian fauna of Sri Lanka have not Data Providers: S.D. Biju, Kelum Manamendra-Arachchi, Anslem de Silva, Sushil Dutta
relocated this species and it is now believed
to be extinct.

EX Philautus zal Manamendra-Arachchi and Pethiyagoda, 2005

Order, Family: Anura, Rhacophoridae Habitat and Ecology The habitat requirements of this species are not known. It presumably bred by direct develop-
Country Distribution: Sri Lanka (Extinct) ment.
Major Threats The reason for the species’ extinction is not known, but presumably habitat loss was a contributing
Geographic Range This recently described species is a Sri Lankan endemic, known only from the general type factor.
locality of “Ceylon” (Manamendra-Arachchi and Pethiyagoda 2005). It is not possible to produce a distribution map Conservation Measures It has not been recorded from any protected areas.
for this species because the exact location of the type collection is not known. Bibliography: Manamendra-Arachchi, K. and Pethiyagoda, R. (2005)
Population It is known only from the holotype and two paratypes. There have been no records since the type collec- Data Providers: Kelum Manamendra-Arachchi, Anslem de Silva
tion (sometime before 1947) and the species is now believed to be extinct, because recent, extensive field surveys
of the amphibian fauna of Sri Lanka have failed to rediscover this frog.

EX Philautus zimmeri (Ahl, 1927)

Order, Family: Anura, Rhacophoridae Geographic Range This is an extinct Sri Lankan endemic, known Notes on taxonomy: This species is considered distinct from Philautus microtympanum following Manamendra-Arachchi and
Country Distribution: Sri Lanka (Extinct) only from the type locality of “Point de Galle, Ceylon”. Pethiyagoda (2005).
Population It is known only from the holotype specimen described Bibliography: Ahl, E. (1927), Bossuyt, F. and Dubois, A. (2001), Manamendra-Arachchi, K. and Pethiyagoda, R. (2005)
in 1927. There have been no records since this time and the species Data Providers: Kelum Manamendra-Arachchi, Anslem de Silva
is now believed to be extinct. Recent, extensive field surveys of the
amphibian fauna of Sri Lanka, including at the type locality, have
not rediscovered this frog.
Habitat and Ecology The habitat requirements of this species are
not known. It presumably bred by direct development.
Major Threats The reason for the species’ extinction is not known,
but presumably habitat loss contributed to its demise as the town of
Galle is heavily urbanized.
Conservation Measures It has not been recorded from any
protected areas.
 Extinct and Extinct in the Wild Amphibian Species
EX Rheobatrachus silus Liem, 1973
Order, Family: Anura, Rheobatrachidae
Country Distribution: Australia (Extinct)
CITES: Appendix II
© Michael J. Tyler
Geographic Range This species, an Australian endemic, was restricted to elevations between 350 and 800m asl in
the Blackall and Conondale Ranges in south-east Queensland (Hines, Mahony and McDonald 1999). The geographic
distribution of the species was less than 1,400km2 (map in Hines, Mahony and McDonald 1999). Rheobatrachus
silus inhabited streams in the catchments of the Mary, Stanley and Mooloolah River (Ingram 1983). It was thought
to have been first found in 1972 (Liem 1973), but Ingram (1991) reported a specimen collected in 1914 from the
Blackall Range.
Population The species declined rapidly and disappeared at about the same time as a sympatric species Taudactylus
diurnus (Czechura and Ingram 1990). Czechura and Ingram (1990) and Ingram (1990) state that the last frog was seen
in the wild in 1979 on the Conondale Range. However, Richards, McDonald and Alford (1993) reported the existence
of a specimen taken from the Blackall Range in 1981. Despite intensive searching, the species has not been located
since (Ingram and McDonald 1993; Hines, Mahony and McDonald 1999). In the laboratory, the last known individual
died in 1983 (Tyler and Davies 1985b). Ingram (1983) studied a population of the species in the headwaters of Bool-
oumba Creek, Conondale Range, and estimated that approximately 78 were present in 1976. No other estimates of
population size are available for the species. This species is now believed to be extinct.
Habitat and Ecology Rheobatrachus silus lived in rainforest, wet sclerophyll forest and riverine gallery open for-
est at 350m asl and was closely associated with watercourses and adjacent rock pools and soaks (Czechura 1991;
Meyer, Hines and Hero 2001e). These streams are mostly perennial, but in extremely dry years they may cease to
flow (Ingram 1983). The vegetation along the stream banks is usually closed forest or tall closed forest with emergent
eucalypts, although there are some sites in open forest with grassy ground cover (Ingram 1983). In spring and summer
individuals were usually found in or at the edge of rock pools, either amongst leaf-litter, under and between stones
EX Rheobatrachus vitellinus Mahony, Tyler and Davies, 1984
Order, Family: Anura, Rheobatrachidae
Country Distribution: Australia (Extinct)
CITES: Appendix II
© Michael J. Tyler
Geographic Range This species, an Australian endemic, was discovered in January 1984 (Mahony et al. 1984)
and was found exclusively in undisturbed rainforest in Eungella National Park, mid-eastern Queensland at altitudes
of 400-1,000m asl (Covacevich and McDonald 1993). The extent of occurrence of the species was less than 500km2
(map in McDonald 1990).
Population The species was considered common across its range until January 1985 when the first signs of decline
(reported by Winter and McDonald 1986) were observed at lower altitudes (i.e., about 400m asl) (McDonald 1990).
At higher altitudes the frogs remained common until March 1985 but were absent in June of that year (McDonald
1990). Despite continued efforts to locate the species, Rheobatrachus vitellinus has not been recorded again within
Eungella National Park or any other locations (Ingram and McDonald 1993; McDonald and Alford 1999). This species
is now considered to be extinct.
Habitat and Ecology It is an aquatic species largely restricted to the shallow section of fast-flowing creeks and
streams in rainforest. It is one of only two known species to brood its offspring within its stomach. Females deposited
143
SOUTHERN GASTRIC BROODING FROG
or in crevices around the edge (Ingram 1983). The species was also found under rock in shallow water in backwaters
and also the main flow of permanent watercourses (Ingram 1983; Czechura 1991). Searches of popular sites in winter
only recovered two frogs and it is assumed that the species hibernates in deep crevices in rocks or spaces between
rocks underwater during the colder months (Ingram 1983). Adult males tend to prefer deeper pools, whereas females
and juveniles may move to newly created pools after rain as long as these pools contained stones and/or leaf-litter
(Ingram 1983). The prerequisite for the use of pools by this species seems to be that the pool must be deep enough
for the frog to be able to sit with its head out of the water and be able to safely submerge (Ingram 1983). Individuals
will only sit fully exposed on the rocks during light rain (Ingram 1983). Rheobatrachus silus has never been recorded
from cleared riparian habitat. Breeding activity occurs between October and December (Ingram 1983). Males call from
rock crevices above pools (Ingram 1983). Females brood young within the stomach and give birth through the mouth
(Tyler and Carter 1982). Fertilized eggs or early stage larvae are presumably swallowed by the female and complete
their development in the stomach (Tyler and Carter 1982). The number of eggs in gravid females (approximately 40)
exceeds the number of juveniles found to occur in the stomach (21-26) (Tyler 1989). It is not known whether or not the
excess eggs are digested by the female or whether or not they are simply not swallowed (Tyler 1989). The production
of hydrochloric acid in the stomach of the female ceases during brooding (Tyler et al. 1983). Tadpoles develop in a
manner similar to the aquatic tadpoles of other species though, as they feed off egg yolk, the labial teeth are absent
and the intestines form at a later stage of development (Tyler 1989). After 6-7 weeks the females give birth to up to
25 young (Tyler and Davies 1983a). Young emerge from the female’s mouth as fully formed frogs and after four days
the digestive tract returns to normal and the female recommences feeding (Tyler and Davies 1983b). Ingram (1983)
reported minimum brooding periods from two individuals of 36 and 43 days and suggested that the duration was
such that females were unlikely to breed twice in one season.
Major Threats The reason(s) for the disappearance of this species remains unknown (Tyler and Davies 1985b).
Populations were present in logged catchments between 1972 and 1979. Although the species persisted in the
streams during these activities, the effects of timber harvesting on this aquatic species were never investigated.
Its habitat is currently threatened by feral pigs, invasion of weeds (especially mistflower Ageratina riparia), and
altered flow and water quality due to upstream disturbances. The habitat of the species is currently threatened by
feral pigs and the invasion of weeds (especially mistflower) (Hines, Mahony and McDonald 1999). However, from
what is known from similar declines and disappearances elsewhere in the world, the disease chytridiomycosis
must be suspected.
Conservation Measures The historical range of the species included several protected areas. Further research into
the cause of the decline of this species is needed.
Bibliography: Czechura, G.V. (1991), Czechura, G.V. and Ingram, G. (1990), Hines, H., Mahony, M. and McDonald, K. (1999), Ingram, G.J.
(1983), Ingram, G.J. (1990), Ingram, G.J. (1991), Ingram, G.J. and McDonald, K.R. (1993), Liem, D.S. (1973), Meyer, E., Hines, H. and Hero,
J.-M. (2001e), Richards, S.J., McDonald, K.R. and Alford, R.A. (1993), Tyler, M.J. (1989), Tyler, M.J. et al. (1983), Tyler, M.J. and Carter,
D.B. (1982), Tyler, M.J. and Davies, M. (1983a), Tyler, M.J. and Davies, M. (1983b), Tyler, M.J. and Davies, M. (1985b)
Data Providers: Ed Meyer, David Newell, Harry Hines, Sarah May, Jean-Marc Hero, John Clarke, Frank Lemckert
NORTHERN GASTRIC-BROODING FROG
their eggs, and then swallowed them. While in the stomach, tadpoles excreted some form of enzyme that inhibited
the female’s gastric digestion, and then proceded to develop into fully formed froglets. The froglets were then
regurgitated through the female’s mouth.
Major Threats The cause(s) of the population decline remain unknown. McDonald (1990) found no obvious evidence
that seasonal rarity, over-collecting, predation, drought, floods, habitat destruction, disease, heavy parasite loads
or stress due to handling for data collection were responsible for the population declines. Threats to the Eungella
National Park include fires that might sweep up the slopes of the mountain during harvesting of the sugar cane in the
fields below (Winter and McDonald 1986). Successive fires extend deeper into the rainforest leaving grassy ridges
that are devoid of trees (Winter and McDonald 1986). The rainforest is extremely narrow in places and continual fire
might eventually erode away entire sections of the forest or fragment the forest (Winter and McDonald 1986). Weeds
that encroach on the edges of the forest pose a similar threat (Winter and McDonald 1986). It was thought that it
might have been possible that the decline that was observed in 1984-1985 was a natural population fluctuation and
that residual individuals had retreated to hidden refuges (Winter and McDonald 1986; McDonald 1990). The extent
of such population fluctuations is unknown, but there is evidence of large swings in numbers of other Australian frogs
(McDonald 1990). However, despite continued efforts to locate the species, it has not been recorded within Eungella
National Park or any other locations since March 1985 (Ingram and McDonald 1993; Richards, McDonald and Alford
1993; Hero et al. 1998, 2002; McDonald and Alford 1999). Eungella National Park is subject to weed invasion on the
edges of the reserve (Winter and McDonald 1986). From what is known from similar declines and disappearance
elsewhere in the world, chytridiomycosis (present in at least some rainforest streams at Eungella) must be suspected
and a major causes of the extinction of this species.
Conservation Measures Its known range is within a protected area. A recovery plan for the species has been
prepared.
Bibliography: Cogger, H.G. (2000), Covacevich, J.A. and McDonald, K.R. (1993), Hero, J.-M. et al. (1998), Hero, J.-M. et al. (2002),
Ingram, G.J. and McDonald, K.R. (1993), Mahony, M., Tyler, M.J. and Davies, M. (1984), McDonald, K. and Alford, R. (1999), McDonald,
K.R. (1990), McDonald, K.R. and Tyler, M.J. (1984), Richards, S.J., McDonald, K.R. and Alford, R.A. (1993), Tyler, M.J. (1989), Tyler, M.J.
(1997), Winter, J. and McDonald, K. (1986)
Data Providers: Jean-Marc Hero, Keith McDonald, Ross Alford, Michael Cunningham, Richard Retallick
144 Threatened Amphibians of the World

EX Plethodon ainsworthi Lazell, 1998 AINSWORTH’S SALAMANDER

Order, Family: Caudata, Plethodontidae Geographic Range This species is known only from two specimens that were collected on 12 June 1964, two miles
Country Distribution: United States of America south of Bay Springs, Jasper County, Mississippi, USA (Lazell 1998).

Peabody Museum of Natural History, Yale University

(Extinct) Population No animals have been found in recent years despite survey work, and this species is considered to be
extinct..
Habitat and Ecology The specimens were collected in springhead litter (Lazell 1998). It was presumably a terrestrial
breeder with a direct development breeding strategy.
Major Threats Although the threats are not well known, it seems possible that habitat loss through deforestation
might have caused the species’ extinction.
Conservation Measures There are no conservation measures needed; this species is extinct.
Bibliography: Blackburn, L., Nanjappa, P. and Lannoo, M.J. (2001), Lazell, J. (1998)
Data Providers: Geoffrey Hammerson
© William Sacco,

EX Cynops wolterstorffi (Boulenger, 1905) YUNNAN LAKE NEWT

© Fei, L., Ye, C.-Y., Huang, Y.-Z. and Liu, M.-Y. 1999

Order, Family: Caudata, Salamandridae Geographic Range This species was restricted to Kunming Lake and the surrounding areas in Yunnan, China.
Country Distribution: China (Extinct) Population No animals can now be found and this species is considered to be extinct.
Habitat and Ecology The species formerly inhabited shallow lake waters and the adjacent irrigation channels,
ponds and marshes. Breeding and larval development took place in these aquatic habitats.
Major Threats The extinction of this species was related to habitat destruction and degradation resulting from general
pollution, land reclamation, domestic duck farming and the introduction of exotic fish and frog species.
Conservation Measures There are no conservation measures needed; this species is extinct.
Bibliography: He, X.R. (1998), Yang, D.-T. (1991b), Zhao, E.-M. (1998)
Data Providers: Yang Datong, Michael Wai Neng Lau
Globally Threatened
Amphibian Species
146 Threatened Amphibians of the World

ANURA

ARTHROLEPTIDAE

EN Arthroleptis crusculum Angel, 1950

Endangered B1ab(iii) Geographic Range This species is known only from Mount Nimba in Guinea; it presumably also occurs in the Liberian
Order, Family: Anura, Arthroleptidae and Ivorian parts of Mount Nimba, but this has not yet been confirmed. It has been recorded from 500-1,650m asl.
Country Distribution: Guinea Population There is no recent information on the population status of this species, but it was formerly recorded
Current Population Trend: Decreasing as being very common.
Habitat and Ecology It lives in high-altitude grassland, gallery forest, and on the edges of marshes. Like other
members of the genus, it presumably breeds by direct development and is not dependent upon water.
Major Threats The major threat to the species is habitat loss and degradation, since Mount Nimba is subject to
slash-and-burn agricultural activities and extensive iron-ore mining operations.
Conservation Measures The species occurs in the Mont Nimba Strict Nature Reserve, which was added to the
list of World Heritage Sites in 1981 (the adjacent part in Cote D’Ivoire was added in 1982). However, strengthened
and improved management of this site is needed, and further survey work is required to determine the current
population status of the species.
© Annika Hillers

Notes on taxonomy: We follow Poynton (2003c) in retaining the genus Schoutedenella only for Schoutedenella xenochirus, and assign
this species to its original genus, Arthroleptis. However, this species is very distinctive and should probably be transferred to the monotypic
genus Arthroleptulus (R.C. Drewes pers. comm.).
Bibliography: Angel, F. (1950), Perret, J.-L. (1991), Poynton, J.C. (2003c)
Data Providers: Mark-Oliver Rödel, Robert Drewes

EN Arthroleptis francei Loveridge, 1953

Endangered B1ab(iii) Geographic Range This species is known only from Mount Mulanje in southern Malawi and possibly from the
Order, Family: Anura, Arthroleptidae nearby Zomba Mountains. It is found as low as 700m asl in the Ruo Basin, but also occurs higher up on the mountain,
Country Distribution: Malawi including on the plateau, probably up to at least 2,500m asl.
Current Population Trend: Decreasing Population It is apparently common within its restricted range, but there have been no records over the last 30
years, probably due to a lack of fieldwork.
Habitat and Ecology This species inhabits both lowland and montane forest, including high-altitude cedar forest. It
is also found, but is probably much less common, in montane grassland and pine plantations. There is little informa-
tion on its adaptability to secondary habitats, but it is believed to favour undisturbed habitats. It lives in leaf-litter in
areas of very high rainfall. They sometimes climb trees up to at least 2m from the ground. It is presumed to breed by
direct development, though the site of egg deposition is not known.
Major Threats Although somewhat protected, the forest on Mount Mulanje continues to be lost in places as a result
of small-scale farming and extraction of wood.
© Colin Tilbury

Conservation Measures It occurs in the Mulanje Mountain Forest Reserve, and continued protection and man-
agement of this area is essential. Further survey work is required to determine the current population status of this
species.
Bibliography: Channing, A. (2001), Poynton, J.C. and Broadley, D.G. (1985a), Stevens, R.A. (1974), Stewart, M.M. (1967)
Data Providers: Lovemore Mazibuko, John Poynton

EN Arthroleptis nikeae Poynton, 2003

Endangered B1ab(iii) Geographic Range This species is known only from the Mafwemiro Catchment Forest Reserve in the Rubeho
Order, Family: Anura, Arthroleptidae Mountains in eastern Tanzania, at 1,900m asl. It presumably occurs more widely, at least within the northern part
Country Distribution: Tanzania of the Rubeho Mountains.
Current Population Trend: Decreasing Population It is a recently discovered species known only from two specimens collected in 2001.
Habitat and Ecology Both specimens were found on the floor of montane forest in a small valley, and one of them
was collected close to a stream. Like other members of the genus, it presumably breeds by direct development, and
is not dependent upon water.
Major Threats It is probably adversely affected by forest loss for agriculture, collection of wood and human
settlement.
Conservation Measures It occurs in the Mafwemiro Catchment Forest Reserve, which affords some level of pro-
© Michele Menegon

tection, but which is not managed for biodiversity conservation. Further survey work would be useful in determining
the species’ current population status.
Bibliography: Poynton, J.C. (2003a)
Data Providers: John Poynton, Kim Howell

VU Arthroleptis tanneri Grandison, 1983

Vulnerable B1ab(iii) Geographic Range This species is known only from the East and West Usambara Mountains, the Nguru Mountains,
Order, Family: Anura, Arthroleptidae and the Ukaguru Mountains in eastern and north-eastern Tanzania. It is a montane species, though its altitudinal
Country Distribution: Tanzania range is not yet fully known; the type specimen was collected at 1,530m asl.
Current Population Trend: Decreasing Population It appears to be an uncommon species.
Habitat and Ecology It lives in leaf-litter on the floor of montane forests, and is generally found in moister situations
(sometimes even aggregating alongside water). It probably cannot cope well with habitat loss. Like other members
of the genus, it presumably breeds by direct development.
Major Threats This species is probably adversely affected by forest loss due to agriculture, wood extraction,
and human settlement. Its habitat in the East Usambaras has recently come under serious threat as a result of the
activities of illegal gold miners.
Conservation Measures It occurs in the University of Dar es Salaam reserve at Mazumbai in the West Usambara
© Alan Channing

Mountains.
Bibliography: Grandison, A.G.C. (1983), Howell, K.M. (1993)
Data Providers: Kim Howell, John Poynton

CR Arthroleptis troglodytes Poynton, 1963
Critically Endangered B1ab(v)+2ab(v)
Order, Family: Anura, Arthroleptidae
Country Distribution: Zimbabwe
Current Population Trend: Decreasing
VU Arthroleptis xenodactylus Boulenger, 1909
Vulnerable B1ab(iii)
Order, Family: Anura, Arthroleptidae
Country Distribution: Tanzania
Current Population Trend: Decreasing
© Elizabeth Harper
CR Cardioglossa alsco Herrmann, Herrmann, Schmitz and Böhme, 2004
Critically Endangered B2ab(iii)
Order, Family: Anura, Arthroleptidae
Country Distribution: Cameroon
Current Population Trend: Decreasing
© Andreas Schmitz
EN Cardioglossa aureoli Schiøtz, 1964
Endangered B1ab(iii)+2ab(iii)
Order, Family: Anura, Arthroleptidae
Country Distribution: Sierra Leone
Current Population Trend: Decreasing
© Annika Hillers
Globally Threatened Amphibian Species
Geographic Range This species is known only from the western
Chimanimani Mountains in eastern Zimbabwe, where it occurs above
1,500m asl. It is likely to occur in the eastern Chimanimani Mountains
across the border in Mozambique.
Population It is probably a rare species. There have been no records
since this species was discovered in 1962 (when 16 specimens were
collected), despite survey efforts.
Habitat and Ecology Most of the specimens were collected in
sinkholes or caves, and a few were found in open montane grassland.
It presumably breeds by direct development.
Major Threats There is very little direct information available for
this poorly known species, and threats to the species are not well
understood. As an isolated, montane species, it could be at risk from
the effects of climate change.
Conservation Measures The area from which it has been recorded
is protected, primarily in the Chimanimani National Park. Surveys are
urgently needed to relocate this species.
147
CAVE SQUEAKER
Notes on taxonomy: We follow Poynton (2003c) in retaining the genus Schoutedenella only for Schoutedenella xenochirus, and
therefore assign this species to its original genus, Arthroleptis. There are serious taxonomic problems with the genera Arthroleptis and
Schoutedenella through much of Africa. In many cases, the available names can be referred only to museum specimens, not to animals
in the field. This is because the identification of these species frequently depends more on their vocalizations than their morphology.
Bibliography: Channing, A. (2001), Lambiris, A.J.L. (1989b), Poynton, J.C. (1963), Poynton, J.C. (1964b), Poynton, J.C. (2003c), Poynton,
J.C. and Broadley, D.G. (1985a)
Data Providers: John Poynton, Alan Channing
Geographic Range This species is endemic to Tanzania, and occurs on the island of Pemba, and on the mainland in
the East Usambara, Uluguru, and Udzungwa Mountains (Kihansi), and Mount Rungwe. It probably occurs more widely
in the Eastern Arc Mountains than current records suggest. It ranges from near sea level up to at least 2,000m asl.
Population It is a reasonably common species.
Habitat and Ecology It occurs in both lowland and montane forest, living in leaf-litter, under logs, and in the axils
of banana leaves. It is not known to what degree it can tolerate alteration of its natural habitat. It breeds by direct
development, and the eggs are laid in leaf-litter.
Major Threats It is probably adversely affected by forest loss for agriculture, wood extraction, and human settle-
ment. Its habitat in the East Usambara Mountains has recently come under serious threat as a result of the activities
of illegal gold miners.
Conservation Measures It occurs in the Amani Nature Reserve, but has not yet been recorded from Udzungwa
National Park (although it is likely to occur there).
Notes on taxonomy: Following Poynton (2003c), the genus Schoutedenella is retained only for Schoutedenella xenochirus, therefore we
assign this species to its original genus, Arthroleptis. There are major taxonomic problems with the genera Arthroleptis and Schoutedenella
through much of Africa. In many cases, the available names can be referred only to museum specimens and not to animals in the field.
This is because the identification of these species frequently depends more on their vocalizations than their morphology. It is likely that
more than one species is covered by this name.
Bibliography: Harper, E. and Vonesh, J.R. (2003), Howell, K.M. (1993), Poynton, J.C. (2003b), Poynton, J.C. (2003c)
Data Providers: Kim Howell, John Poynton
Geographic Range This recently described species is known only from the southern slopes of Mount Tchabal Mbabo
at 1,700-2,100m asl on the Adamawa Plateau in western Cameroon. It is unlikely to occur on the northern slopes, at
least not at the altitude of the type locality, because the forest habitat on the northern slopes at this altitude is very
different from the gallery forest on the southern slopes.
Population There is little information, although it is likely to be common within its small range, since 73 specimens
were collected close together, suggesting a high concentration of animals.
Habitat and Ecology It lives in montane gallery forest, and is associated with streams, in which it presumably
breeds by larval development. Individuals have been found under large stones around shallow pools adjacent to a
creek. They were calling at night during the dry season.
Major Threats The remaining forest habitat on Tchabal Mbabo is now confined to galleries and steep inaccessible
slopes, as a result of the clearing of forest for pasture. However, much of the forested habitat that can be easily
removed has now been cleared. The forest is also threatened by fire.
Conservation Measures This species is not known from any protected areas, though one has been recommended
for Tchabal Mbabo.
Notes on taxonomy: Records of Cardioglossa pulchra from Tchabal Mbabo (Amiet 1972a) now refer to this species (Herrmann et al.
2004).
Bibliography: Amiet, J.-L. (1972a), Herrmann, H.-W. et al. (2004)
Data Providers: Hans-Werner Herrmann
Geographic Range This species is known only from the hilly parts of the Freetown Peninsula in Sierra Leone,
although it possibly occurs a little more widely.
Population There has been no information on this species since it was last collected in 1963, but this is probably
due to lack of field surveys in this area. It was described as “not rare” on the Freetown Peninsula at the time of its
discovery.
Habitat and Ecology It has been found in forest, and in rural gardens near forest, living on rocks. It has been noted
as calling away from water, and so it is possible that, unlike other members of the genus, it does not require water
for breeding, although this has not been confirmed.
Major Threats There is likely to be extensive pressure on its forest habitat on the Freetown Peninsula primarily
for firewood collection.
Conservation Measures It is likely to be present in the Western Area Peninsula Forest Reserve, although much
more effective protection and management of this site is required. Additional survey work is urgently needed to
determine the current population status of this species.
Bibliography: Schiøtz, A. (1964a)
Data Providers: Arne Schiøtz, Mark-Oliver Rödel
148 Threatened Amphibians of the World

EN Cardioglossa melanogaster Amiet, 1972

Endangered B1ab(iii) Geographic Range This species is known from the mountains of eastern Nigeria and western Cameroon at
Order, Family: Anura, Arthroleptidae 1,200-2,000m asl. In Nigeria, it has been recorded from the Obudu Plateau, and in Cameroon it is known from
Country Distribution: Cameroon, Nigeria Mount Manenguba, the southern fringe of the Bamileke Plateau (at Foto and Mount Bana), Mount Nlonako, and the
Current Population Trend: Decreasing Bamenda Highlands (in the Bafut-Ngemba Forest Reserve). At Mwakoumel on Mount Manenguba, it co-exists with
Cardioglossa pulchra and C. venusta.
Population It is common on Mount Manenguba at about 1,600-1,700m asl.
Habitat and Ecology It lives in submontane and montane areas in forest and in dense secondary growth. Males
call from shady situations along streams. It is a stream-breeding species.
Major Threats It is at risk because of forest loss as a result of expanding human settlements, agricultural encroach-
© David C. Blackburn ment, and removal of wood.
Conservation Measures It does occur in the Bafut-Ngemba Forest Reserve, but additional habitat protection of the
remaining highland forests in Cameroon, particularly Mount Manenguba, and in Nigeria, is needed.
Bibliography: Amiet, J.-L. (1972a), Amiet, J.-L. (1972c), Amiet, J.-L. (1973b), Amiet, J.-L. (1973c), Amiet, J.-L. (1975), Amiet, J.-L.
(1981), Herrmann, H.-W. et al. (2005)
Data Providers: Jean-Louis Amiet, Arne Schiøtz

EN Cardioglossa oreas Amiet, 1972

Endangered B1ab(iii) Geographic Range This species is endemic to the mountains of western Cameroon at 1,900-2,650m asl, occurring
Order, Family: Anura, Arthroleptidae on Mount Manenguba, and on the Bamenda Highlands at the Bamboutos Mountains, the Bafut-Ngemba Forest
Country Distribution: Cameroon Reserve, and Mount Oku.
Current Population Trend: Decreasing Population It is common within its small range.
Habitat and Ecology It lives in montane forest, often in bamboo forest, and in degraded habitats with trees. They
are associated with areas around fast-flowing streams, where they sit on leaves, or hide in holes or under stones
and breed in streams.
Major Threats This species is seriously threatened by forest loss due to agricultural expansion and extraction of
wood, as well as by overgrazing and expansion of human settlements.
Conservation Measures The species occurs in the Bafut-Ngemba Forest Reserve, but is largely unprotected
elsewhere in its range. A conservation project has been conducted on Mount Oku for several years by BirdLife Inter-
© Václav Gvoždík

national, involving community management of the area by the local villages; this project needs to take into account
the conservation needs of this species.
Bibliography: Amiet, J.-L. (1972a), Amiet, J.-L. (1972c), Gartshore, M.E. (1986)
Data Providers: Jean-Louis Amiet, Arne Schiøtz

EN Cardioglossa pulchra Schiøtz, 1963

Endangered B1ab(iii) Geographic Range This species is known from the mountains of eastern Nigeria and western Cameroon at
Order, Family: Anura, Arthroleptidae 900-1,800m asl. In Nigeria, it has been recorded from the Obudu Plateau, and in Cameroon it is known from Mount
Country Distribution: Cameroon, Nigeria Manenguba, Mount Ngokham, the Bafut-Ngemba Forest Reserve, Batie, and Fongo-Tongo. At Mwakoumel on Mount
Current Population Trend: Decreasing Manenguba, it co-exists with C. melanogaster and C. venusta.
Population It is known to be common in the breeding season.
Habitat and Ecology It lives in submontane and montane forest along fast-flowing streams and also at high eleva-
tions along forest edges. It can survive with some deforestation and it breeds in streams.
Major Threats It is probably threatened by ongoing forest loss caused by agricultural encroachment, extraction of
wood, and expanding human settlements.
Conservation Measures It is found in the Bafut-Ngemba Forest Reserve in Cameroon and the Cross River National
© David C. Blackburn

Park in Nigeria. Continued protection of these areas, and further habitat protection of the remaining highland forests
in Cameroon, particularly Mount Manenguba, and in Nigeria, is needed.
Notes on taxonomy: Records of this species from Tchabal Mbabo refer to Cardioglossa alsco.
Bibliography: Amiet, J.-L. (1971b), Amiet, J.-L. (1972a), Amiet, J.-L. (1972c), Amiet, J.-L. (1973c), Amiet, J.-L. (1975), Schiøtz, A.
(1963)
Data Providers: Arne Schiøtz, Jean-Louis Amiet

EN Cardioglossa schioetzi Amiet, 1981

Endangered B1ab(iii) Geographic Range This species is endemic to the Oshie-Obudu Ridge in Cameroon and Nigeria. It is found from Acha
Order, Family: Anura, Arthroleptidae Tugi on Mount Oshie in Cameroon and from the Obudu Plateau in Nigeria. Its altitudinal range is 1,640-1,800m asl.
Country Distribution: Cameroon, Nigeria Population There is no information available on its population status, and the species is generally poorly known.
Current Population Trend: Decreasing Habitat and Ecology It lives in and around relict patches of montane forest, which are now fragmented, and also in
secondary vegetation in which no trees remain. It has been found on steep slopes of scree, boulders and clay, overgrown
with a dense cover of high grass and trees. It has also been found around streams, where it presumably breeds.
Major Threats It is threatened by habitat loss due to expanding agricultural activities and human settlements,
and overgrazing.
Conservation Measures It occurs in the Cross River National Park in Nigeria, but further measures are urgently
needed to protect the remaining patches of habitat in its very small range. Additional survey work would be useful
© Mary E. Gartshore

to determine the current population status of this species.

Bibliography: Amiet, J.-L. (1981), Gartshore, M.E. (1986)
Data Providers: Jean-Louis Amiet, Arne Schiøtz
 Globally Threatened Amphibian Species 149

CR Cardioglossa trifasciata Amiet, 1972

Critically Endangered B1ab(iii)+2ab(iii) Geographic Range This species is known only from the southern slopes of Mount Manenguba at 1,750-1,800m
Order, Family: Anura, Arthroleptidae asl in western Cameroon.
Country Distribution: Cameroon Population There is no information available on its current population status.
Current Population Trend: Decreasing Habitat and Ecology It has been found in and around a small stream running through dense secondary bush and
montane forest; it has not been recorded from primary forest, although it might well occur. The animals hide under
large rocks and small stones, and presumably breed in streams.
Major Threats Although the habitat of this species is not especially threatened, it is at severe risk given its small
range and vulnerability to agricultural encroachment, wood extraction, and expanding human settlements.
Conservation Measures The species is not known from any protected areas, and the remaining habitat in the
area needs to be protected as a matter of urgency. There is a need for further survey work to determine the current
population status of this species.
© Jean-Louis Amiet

Bibliography: Amiet, J.-L. (1972a), Amiet, J.-L. (1972c), Amiet, J.-L. (1973c), Amiet, J.-L. (1975)
Data Providers: Jean-Louis Amiet, Arne Schiøtz

EN Cardioglossa venusta Amiet, 1972

Endangered B1ab(iii) Geographic Range This species is known only from a small area of the mountains of western Cameroon at 950-1,350m
Order, Family: Anura, Arthroleptidae asl at Mount Manenguba, the Bamileke Plateau (at Fotabong, Fontem, and the Mbos Cliffs), Mount Nlonako, and the
Country Distribution: Cameroon Rumpi Hills. At Mwakoumel on Mount Manenguba, it co-exists with Cardioglossa pulchra and C. melanogaster.
Current Population Trend: Decreasing Population The species is largely unknown, and there is no information on its current population status.
Habitat and Ecology It lives in submontane forests near fast-flowing streams in hilly country with high rainfall. It
can survive in degraded, secondary habitat close to more mature forest and it breeds in streams.
Major Threats It is at risk due to forest loss caused by agricultural encroachment, extraction of wood, and expand-
ing human settlements.
Conservation Measures It may occur in the Rumpi Hills Forest Reserve, but this reserve is not well managed for
biodiversity conservation; improved and expanded habitat protection is urgently needed to ensure the survival of
this species.
© Andreas Schmitz

Bibliography: Amiet, J.-L. (1972a), Amiet, J.-L. (1972c), Amiet, J.-L. (1973c), Amiet, J.-L. (1975), Herrmann, H.-W. et al. (2005)
Data Providers: Arne Schiøtz, Jean-Louis Amiet

ASTYLOSTERNIDAE

VU Astylosternus diadematus Werner, 1898

Vulnerable B1ab(iii) Geographic Range This species is known only from western and south-western Cameroon, and possibly extreme
Order, Family: Anura, Astylosternidae eastern Nigeria, in hilly country and mountains, at 250-1,100m asl. There are records from Mount Cameroon, and
Country Distribution: Cameroon other mountains in Cameroon south of the higher parts of the Bamenda Highlands. Records of Astylosternus schioetzi
Current Population Trend: Decreasing from the northern part of Korup National Park, east to Nguti, and north to Mount Nta Ali (Lawson 1993) are based
on misidentifications and refer to this species (J.-L. Amiet pers. comm.).
Population It is abundant in suitable habitat.
Habitat and Ecology It lives in or near flowing water in lowland and submontane forest, and can survive in somewhat
degraded habitats. It breeds in streams and in small marshes criss-crossed with tiny watercourses. The tadpoles
live in clear, flowing water.
Major Threats The major threat to this species is habitat loss due to smallholder farming activities and logging.
© Mary E. Gartshore

Conservation Measures It occurs in the Korup National Park.

Notes on taxonomy: It is possible that animals resembling this species on Mount Manenguba and at Buea on Mount Cameroon belong
to as yet undescribed forms (Amiet 1977).
Bibliography: Amiet, J.-L. (1971b), Amiet, J.-L. (1973c), Amiet, J.-L. (1977), Amiet, J.-L. (1983a), Amiet, J.-L. (1987), Amiet, J.-L. (1989),
Herrmann, H.-W. et al. (2005), Lawson, D.P. (1993)
Data Providers: Jean-Louis Amiet

EN Astylosternus fallax Amiet, 1978 “1977”

Endangered B1ab(iii)+2ab(iii) Geographic Range This species is known only from three small areas of western and south-western Cameroon:
Order, Family: Anura, Astylosternidae between Yabassi and Nkongsamba, at low elevations (apparently, it is absent to the west of Nkongsamba); Mount
Country Distribution: Cameroon Yuhan in Korup National Park; and Mount Nta Ali in the Mamfe basin. It occurs mainly below 1,000m asl.
Current Population Trend: Decreasing Population It is a common species, certainly more so than Astylosternus laurenti.
Habitat and Ecology It lives and breeds in and near rivers and slow-flowing streams in lowland and hilly closed-
canopy forest. Males call from among dead leaves on the banks close to water.
Major Threats It is severely threatened by loss of habitat within its very small range, primarily due to human settle-
ment and agricultural encroachment.
Conservation Measures It occurs in Korup National Park, but further protection of the remaining forest habitat
is required.
Bibliography: Amiet, J.-L. (1977), Herrmann, H.-W. et al. (2005), Lawson, D.P. (1993)
© Jean-Louis Amiet

Data Providers: Jean-Louis Amiet

150 Threatened Amphibians of the World

EN Astylosternus laurenti Amiet, 1978 “1977”

Endangered B1ab(iii)+2ab(iii) Geographic Range This species is known only from a small part of south-western Cameroon at 400-850m asl,
Order, Family: Anura, Astylosternidae from the Ikenge Research Station in Korup National Park, to the south of the Rumpi Hills, Mount Manenguba and
Country Distribution: Cameroon the Bamileke Plateau. There are records from Ikenge, Fopouanga, Ekomtolo and Mahole. It is probably a little more
Current Population Trend: Decreasing widespread than current records indicate.
Population It is a common species.
Habitat and Ecology It lives and breeds in and near slow-flowing rivers and streams in lowland and hilly forest in
regions of high rainfall. It can live in dense secondary forest provided that there is a closed canopy. Males call from
rocks in and around the water.
Major Threats This species is severely threatened by habitat loss and degradation, primarily due to human settle-
© Mary E. Gartshore ment and agricultural encroachment.
Conservation Measures It occurs in Korup National Park, but further protection of other unprotected forest habitat,
such as Mount Manenguba, is required.
Bibliography: Amiet, J.-L. (1977), Lawson, D.P. (1993)
Data Providers: Jean-Louis Amiet

CR Astylosternus nganhanus Amiet, 1978 “1977”

Critically Endangered B1ab(iii)+2ab(iii) Geographic Range This species is known only from Mount Nganha Bibliography: Amiet, J.-L. (1977)
Order, Family: Anura, Astylosternidae on the Adamawa Plateau, at 1,400-1,700m asl, in Cameroon, to which Data Providers: Jean-Louis Amiet
Country Distribution: Cameroon it is probably endemic.
Current Population Trend: Decreasing Population It is known only from five specimens.
Habitat and Ecology This species is found along watercourses
in a few narrow gallery forests, and in seepage areas in nearby
grassland. Tadpoles, almost certainly of this species, have been
found in rock pools in streams.
Major Threats This poorly known species is probably at severe
risk from habitat loss due to smallholder farming activities and
subsistence wood extraction.
Conservation Measures It is not known from any protected areas
and protection of the remaining habitat in the range of the species
is urgently needed. Further survey work is required to establish the
current population status of the species.

EN Astylosternus perreti Amiet, 1978 “1977”

Endangered B1ab(iii) Geographic Range This species is known only from western Cameroon on the southern slopes of Mount Manenguba
Order, Family: Anura, Astylosternidae and from Mount Bana and Mount Nlonako on the Bamileke Plateau at an altitude of 1,200-1,400m asl. It presumably
© David C. Blackburn, courtesy of Project Exploration

Country Distribution: Cameroon occurs more widely in the Bamileke Plateau, and there is an unconfirmed record from the Rumpi Hills.
Current Population Trend: Decreasing Population It is common on Mount Manenguba.
Habitat and Ecology It lives in or near flowing water in lower montane and submontane forest, often in very steep,
precipitous areas close to torrents. It probably hides in holes during the day, and breeds in mountain streams.
Major Threats This species is experiencing severe habitat loss within its very small range. However, it is capable
of adapting to a measure of forest degradation.
Conservation Measures The species is not known from any protected areas, and habitat protection of the remaining
montane forests in western Cameroon is urgently needed.
Bibliography: Amiet, J.-L. (1977), Gartshore, M.E. (1986), Herrmann, H.-W. et al. (2005)
Data Providers: Jean-Louis Amiet

EN Astylosternus ranoides Amiet, 1978 “1977”

Endangered B1ab(iii)+2ab(iii) Geographic Range This species is known only from high altitudes (2,000-2,600m asl) in western Cameroon; there
Order, Family: Anura, Astylosternidae are records from the Bamboutos Mountains, Lake Oku, and Mount Neshele in the Bafut-Ngemba Forest Reserve.
Country Distribution: Cameroon Population There is no information on its population status.
Current Population Trend: Decreasing Habitat and Ecology It lives in grassy marshes, around lakes, and along streams and watercourses in montane
forest, montane shrubland, and montane grassland. Its breeding habitat is not fully known, but it appears to be less
tied to flowing water than other members of its genus.
Major Threats Overgrazing, forest clearance, and changes in water quality are probably the major threats to this
species.
Conservation Measures It occurs in the Bafut-Ngemba Forest Reserve, but is largely unprotected elsewhere in its
range. A conservation project has been conducted on Mount Oku for several years by BirdLife International, which
involves community management of the area involving the local villages, but which needs to take into account the con-
© Jean-Louis Amiet

servation needs of this species. Further survey work is needed to determine the species’ current population status.
Bibliography: Amiet, J.-L. (1977), Amiet, J.-L. (1989), Gartshore, M.E. (1986)
Data Providers: Jean-Louis Amiet
 Globally Threatened Amphibian Species 151

VU Astylosternus rheophilus Amiet, 1978 “1977”

Vulnerable B1ab(iii)+2ab(iii) Geographic Range This species is endemic to western Cameroon at altitudes of 1,300-2,450m asl. The nominate
Order, Family: Anura, Astylosternidae subspecies is recorded from Mount Manenguba, Mount Mbam, Santa, Mount Bana, Mount Ngokham, the Bamboutos
Country Distribution: Cameroon Mountains, and the Bafut-Ngemba Forest Reserve, and is likely also to occur on Mount Oku. The subspecies A. r.
Current Population Trend: Decreasing tchabelensis is known only from the southern slopes of Tchabal Mbaba, at 1,700-1,900m asl, in Cameroon, and might
occur on the Mambila Plateau in eastern Nigeria.
Population It is a very common species.
Habitat and Ecology It lives in both the submontane and montane zones, and is typically associated with small
streams in forest where it breeds. However, at higher altitudes above 2,000m asl it can also be found along streams
in montane grassland, sometimes bordered with trees. It seems able to tolerate minor disturbance of its habitat. The
subspecies A. r. tchabelensis lives in relict gallery forests in a generally deforested area.
Major Threats Although it is likely that it can tolerate a degree of habitat alteration, this species is probably declin-
© Václav Gvoždík

ing as a result of widespread habitat loss and degradation within its restricted range, mainly due to smallholder
agricultural activities, subsistence wood extraction, and human settlement.
Conservation Measures It occurs in the Bafut-Ngemba Forest Reserve.
Bibliography: Amiet, J.-L. (1977), Amiet, J.-L. (1989)
Data Providers: Jean-Louis Amiet

EN Astylosternus schioetzi Amiet, 1978 “1977”

Endangered B1ab(iii)+2ab(iii) Geographic Range This species is known only from two general areas in south-western Cameroon: Apouh and
Order, Family: Anura, Astylosternidae Koupongo, near Edéa, in southern Cameroon (these two localities are 50km apart, and are separated by the Sanaga
Country Distribution: Cameroon River). Records from the northern part of Korup National Park, east to Nguti, and north to Mount Nta Ali (Lawson 1993)
Current Population Trend: Decreasing are based on misidentifications and refer to Astylosternus diadematus (J.L. Amiet pers. comm.).
Population It appears to be a rare species occurring at low densities in the southern parts of its range.
Habitat and Ecology It lives in and near flowing water in lowland forest, and can survive in tall, secondary forest.
It breeds in small streams and in marshy depressions with very small, superficial streams.
Major Threats The major threat to the species is habitat loss and degradation.
Conservation Measures It might occur in the Douala-Edea Forest Reserve and Lac Ossa Wildlife Reserve. The
remaining forest habitat in south-western Cameroon is urgently in need of improved protection.
© David C. Blackburn

Bibliography: Amiet, J.-L. (1977), Lawson, D.P. (1993)

Data Providers: Jean-Louis Amiet

VU Leptodactylodon albiventris (Boulenger, 1905)

Vulnerable B1ab(iii) Geographic Range This species is known only from the western edge of the southern Cameroon plateau, though it
Order, Family: Anura, Astylosternidae might occur more widely than current records suggest. Its altitudinal range is 300-1,000m asl.
Country Distribution: Cameroon Population It is an uncommon species.
Current Population Trend: Decreasing Habitat and Ecology This is a species of lowland rainforest in hilly country. It is not known to what extent it can
adapt to forest disturbance. It breeds in streams, and the males call from rocky areas (under rocks or in cracks) close
to small streams.
Major Threats The major threat to this species is habitat loss from smallholder farming activities and logging.
Conservation Measures It is not known from any protected areas, though it might occur in Campo-Ma’an National
Park. There is a need for improved habitat protection at sites at where the species is known to occur.
Notes on taxonomy: Following Amiet and Dowsett-Lemaire (2000), Leptodactylodon bueanus is here considered a separate species
© Ignacio De La Riva

from L. albiventris (as opposed to a subspecies of L. albiventris following Amiet (1987)). Rödel and Pauwels (2003) provisionally assigned
records of L. albiventris from Equatorial Guinea (De la Riva 1994b) to L. stevarti.
Bibliography: Amiet, J.-L. (1980a), Amiet, J.-L. (1987), Amiet, J.-L. and Dowsett-Lemaire, F. (2000), Amiet, J.-L. and Schiøtz, A. (1972),
De la Riva, I. (1994b), Frétey, T. and Blanc, C.P. (2000), Rödel, M.-O. and Pauwels, O.S.G. (2003)
Data Providers: Jean-Louis Amiet

EN Leptodactylodon axillaris Amiet, 1971

Endangered B1ab(iii) Geographic Range This species is known from the Bamboutos Mountains in the Bamenda Highlands of western
Order, Family: Anura, Astylosternidae Cameroon, at altitudes of 2,300-2,700m asl. It probably also occurs on Mount Oku in the Bamenda Highlands.
Country Distribution: Cameroon Population It is most common above 2,400-2,450m asl.
Current Population Trend: Decreasing Habitat and Ecology It lives in bamboo forest and, in the rainy season, grazed montane grasslands where it hides
under rocks in seepage areas. It lives in drier areas than other members of its genus. It has never been found at its
breeding sites, but it is likely to breed in small streams in rocky areas.
Major Threats It is probably under severe threat from habitat loss and overgrazing. Like other high-altitude species,
it also could be at risk from climate change.
Conservation Measures This species is not known from any protected areas, and habitat protection is urgently
needed in the Bamboutos Mountains. In view of its tiny range and possible impacts of climate change, a captive-
© David C. Blackburn

breeding programme should probably be established.

Bibliography: Amiet, J.-L. (1971a), Amiet, J.-L. (1980a)
Data Providers: Jean-Louis Amiet
152 Threatened Amphibians of the World

VU Leptodactylodon bicolor Amiet, 1971

Vulnerable B1ab(iii) Geographic Range This species occurs on the southern and western edge of the Cameroon mountain range (ex-
Order, Family: Anura, Astylosternidae cluding Mount Cameroon), extending to the Obudu Plateau in eastern Nigeria. In Cameroon, it is known from Mount
Country Distribution: Cameroon, Nigeria Manenguba, Mount Nlonako, the Mbos Cliffs, Fotabong, Petit Diboum, Mount Bana, Acha Tugi near Oshie, Bafut
Current Population Trend: Decreasing near Bamenda, and the Rumpi Hills. Its altitudinal range is 950-1,750m asl.
Population It is generally common in suitable habitat.
Habitat and Ecology An inhabitant of forest in the submontane zone, usually near rocky streams and springs, or on
mossy talus slopes with fissures and caves. It avoids dry areas, living in places where there tends to be orographic
mists during the dry season. It can survive in somewhat degraded habitats. Breeding takes place in streams in rocky
areas.
© Mary E. Gartshore Major Threats The major threat to this species is forest loss due to smallholder farming activities and subsistence
wood extraction. The tadpoles are eaten locally by villagers in the Rumpi Hills.
Conservation Measures It occurs in the Cross River National Park in Nigeria. Further research is needed into the
harvest level of tadpoles of this species from the wild.
Bibliography: Amiet, J.-L. (1971a), Amiet, J.-L. (1980a), Amiet, J.-L. (1983a), Amiet, J.-L. and Schiøtz, A. (1972), Gartshore, M.E.
(1986)
Data Providers: Jean-Louis Amiet

VU Leptodactylodon boulengeri Nieden, 1910

Vulnerable B1ab(iii) Geographic Range This species is known from the mountains of western Cameroon. There are records from Banyo
Order, Family: Anura, Astylosternidae (near Batie), Petit Diboum, Mount Bana, Foto, Mbakang, Mount Nlonako and Bafut, with an isolated population to
Country Distribution: Cameroon the east on Mount Ngorro. It probably also occurs in the western section of the Adamawa Plateau, and perhaps also
Current Population Trend: Decreasing in Nigeria. It ranges from 800-1,450m asl.
Population It is a common species.
Habitat and Ecology It lives in degraded forest, dense bush and raffia palm verges along swamps bordering rocky
streams. Breeding takes place in streams.
Major Threats Although it can tolerate some habitat disturbance, it is probably threatened by habitat loss and
degradation due to smallholder farming activities and subsistence wood extraction throughout its range.
Conservation Measures It is not known to occur in any protected areas. There is a need for improved habitat
© Mary E. Gartshore

protection at sites at which this species is known to occur.

Bibliography: Amiet, J.-L. (1971a), Amiet, J.-L. (1971b), Amiet, J.-L. (1973b), Amiet, J.-L. (1975), Amiet, J.-L. (1980a), Amiet, J.-L. and
Schiøtz, A. (1972), Herrmann, H.-W. et al. (2005)
Data Providers: Jean-Louis Amiet

VU Leptodactylodon bueanus Amiet, 1981 “1980”

Vulnerable D2 Geographic Range This species is known only from the eastern slopes of Mount Cameroon, and from Bimbia Hill
Order, Family: Anura, Astylosternidae east of Limbe, in western Cameroon. Its altitudinal range is 200-1,000m asl.
Country Distribution: Cameroon Population It is abundant at many localities on the eastern slopes of Mount Cameroon.
Current Population Trend: Stable Habitat and Ecology It lives in and around streams and springs in forest, usually sheltering in rocky areas. Most of the
known localities are in degraded forest, but it requires dense vegetation in order to survive. Its absence above 1,000m
asl is probably related to the lack of streams, which it needs to breed, at high altitudes on Mount Cameroon.
Major Threats Despite its tolerance of secondary vegetation, populations are likely to be threatened by habitat loss
due to smallholder farming activities, subsistence wood extraction, and human settlement.
Conservation Measures It is not known to occur in any protected areas. There is a need for improved habitat
protection at sites where the species is known to occur.
© David C. Blackburn

Notes on taxonomy: Following Amiet and Dowsett-Lemaire (2000), Leptodactylodon bueanus is here treated as a separate species
from L. albiventris (as opposed to a subspecies of L. albiventris following Amiet (1987)).
Bibliography: Amiet, J.-L. (1980a), Amiet, J.-L. (1987), Amiet, J.-L. and Dowsett-Lemaire, F. (2000)
Data Providers: Jean-Louis Amiet

CR Leptodactylodon erythrogaster Amiet, 1971

Critically Endangered B1ab(iii) Geographic Range This species is known only from the south-eastern slopes of Mount Manenguba at 1,550-1,800m
Order, Family: Anura, Astylosternidae asl in western Cameroon.
Country Distribution: Cameroon Population It is reported to be abundant within its tiny range, and is most common at altitudes of 1,700-1,800m
Current Population Trend: Decreasing asl.
Habitat and Ecology It is found in submontane and lower montane forest, around springs and streams, living in
holes, humus, gravel, root masses and dense undergrowth. It can survive in open, disturbed forest. It presumably
breeds in streams, and co-exists with Leptodactylodon mertensi around 1,700m asl.
Major Threats Habitat loss is taking place on Mount Manenguba due to smallholder farming activities and sub-
sistence wood extraction, and while this species is tolerant of a degree of habitat disturbance, it is nonetheless at
severe risk in view of its tiny range.
Conservation Measures It is not currently recorded from any protected areas. There is an urgent need for improved
© Jean-Louis Amiet

protection of remaining forest habitats on Mount Manenguba.

Bibliography: Amiet, J.-L. (1970a), Amiet, J.-L. (1971b), Amiet, J.-L. (1973b), Amiet, J.-L. (1975), Amiet, J.-L. (1980a), Amiet, J.-L. and
Schiøtz, A. (1972)
Data Providers: Jean-Louis Amiet
 Globally Threatened Amphibian Species 153

EN Leptodactylodon mertensi Perret, 1959

Endangered B1ab(iii) Geographic Range This species is known only from the southern slopes of the Bamileke Plateau (at Mount Bana and
Order, Family: Anura, Astylosternidae Petit Diboum), Mount Nlonako and Mount Manenguba, western Cameroon. It occurs between 1,000 and 1,850m asl,
Country Distribution: Cameroon though only up to 1,700m asl on Mount Manenguba, above which it is replaced by Leptodactylodon erythrogaster.
Current Population Trend: Decreasing Population It is reported to be abundant on the southern and south-eastern slopes of Mount Manenguba.
Habitat and Ecology It lives in dense undergrowth in submontane and lower montane forest, and in the dense
herbage of raffia palm beds along streams. It can also survive in dense secondary forest habitats. The males call
near pools and riffles in small streams, or in waterlogged humus near springs. It avoids rocky areas and breeds in
small streams.
Major Threats Although this species can tolerate a degree of habitat modification, it is probably threatened by
forest clearance for smallholder farming.
Conservation Measures It is not known to occur in any protected areas, and protection of the remaining forest
© Andreas Schmitz

habitat at the sites where it occurs, particularly Mount Manenguba, is urgently needed.
Bibliography: Amiet, J.-L. (1970a), Amiet, J.-L. (1971b), Amiet, J.-L. (1980a), Amiet, J.-L. and Schiøtz, A. (1972), Herrmann, H.-W. et
al. (2005)
Data Providers: Jean-Louis Amiet

EN Leptodactylodon ornatus Amiet, 1971

Endangered B1ab(iii) Geographic Range This species comprises two subspecies, both endemic to western Cameroon: the nominate
Order, Family: Anura, Astylosternidae subspecies occurs on Mount Manenguba, Mount Nlonako, Mount Kupe, the Bonandam Hills and the Ebonji Hills at
Country Distribution: Cameroon 200-1,450m asl; Leptodactylodon ornatus permaculatus occurs on the southern and western slopes of the Bamileke
Current Population Trend: Decreasing Plateau at Fotabong, Foto, and the Mbos Cliffs, at 1,000-1,400m asl.
Population Generally, this species is somewhat uncommon, although L. o. permaculatus is not uncommon within
its small range.
Habitat and Ecology This is not a true montane species, but occurs at low and medium altitudes in hilly areas,
usually near mountains, in areas of high rainfall. It lives in wet lowland and submontane forest and also in partially
degraded forest. It frequents rocky areas that are found less frequently at lower altitudes. It breeds in fast-flowing
streams; the males call from fissures in rocks, or from under stones.
Major Threats Despite its tolerance of a degree of habitat modification, this species is probably threatened by
© Andreas Schmitz

forest clearance for smallholder farming.

Conservation Measures It is not known to occur in any protected areas, and protection of the remaining forest
habitat at the sites where it occurs, particularly Mount Manenguba, is urgently needed.
Bibliography: Amiet, J.-L. (1970a), Amiet, J.-L. (1980a), Amiet, J.-L. and Schiøtz, A. (1972), Herrmann, H.-W. et al. (2005)
Data Providers: Jean-Louis Amiet

EN Leptodactylodon perreti Amiet, 1971

Endangered B2ab(iii) Geographic Range This species is known only from the higher peaks of western Cameroon in the Bamenda
Order, Family: Anura, Astylosternidae Highlands, and further north-east at Tchabal Mbaba. In the Bamenda Highlands it is known from the Bamboutos
Country Distribution: Cameroon Mountains, Mount Mbam, Mount Ngokham, Mount Lefo, Mount Oku, and the Bafut-Ngemba Forest Reserve. Its
Current Population Trend: Decreasing altitudinal range is 1,200-2,650m asl.
Population It is reported to be common within its restricted range.
Habitat and Ecology This is mainly a species of the montane zone, but it also occurs in smaller numbers in the
submontane zone down to 1,200m asl. It is strictly confined to forest, and the males call from secluded sites and holes
in humus close to streams. At higher altitudes, it breeds in shallow marshes in forest traversed by small, superficial
streams. At lower altitudes it breeds in streams in rocky areas.
Major Threats It is seriously threatened by extensive forest loss, mainly from smallholder farming, throughout
its range.
© Wolfgang Böhme

Conservation Measures It occurs in the Bafut-Ngemba Forest Reserve, but remains largely unprotected elsewhere
in its range. BirdLife International has conducted a conservation project on Mount Oku for several years; this project
involves community management of the area involving the local villages, but it also needs to take into account the
conservation needs of this species.
Bibliography: Amiet, J.-L. (1971a), Amiet, J.-L. (1980a), Amiet, J.-L. (1983a), Amiet, J.-L. (1987), Amiet, J.-L. and Schiøtz, A. (1972)

VU Leptodactylodon polyacanthus Amiet, 1971

Vulnerable B1ab(iii) Geographic Range This species occurs in the western Cameroonian highlands, excluding Mount Cameroon, Mount
Order, Family: Anura, Astylosternidae Kupe, Mount Nlonako, Mount Manenguba and the Rumpi Hills. It also occurs on the Obdudu Plateau in eastern Nigeria.
Country Distribution: Cameroon, Nigeria There are two subspecies: L. p. polyacanthus occurs in the northern parts of the species’ range at 1,640-1,900m asl in
Current Population Trend: Decreasing the Bafut-Ngemba Forest Reserve, Acha Tugi on Mount Oshie, and the Obudu Plateau; and L. p. punctiventris occurs in
the southern part of the range at 1,000-1,400m asl on the wet southern and western slopes of the Bamileke Plateau
at Foto, Fotabong, Fongo-Tongo, the Mbos Cliffs, Petit Diboum and Mount Bana.
Population It is a common species.
Habitat and Ecology It lives in montane and submontane forest and degraded forest. During the breeding season
males can be found on wet clay, in rock crevices, and small streams; the females hide under stones. Breeding takes
place in streams and springs.
Major Threats Although it can tolerate degraded forest, it is likely to be threatened by ongoing forest loss due to
© Jean-Louis Amiet

smallholder farming activities and subsistence wood extraction.

Conservation Measures It occurs in the Bafut-Ngemba Forest Reserve.
Bibliography: Amiet, J.-L. (1970a), Amiet, J.-L. (1971a), Amiet, J.-L. (1971b), Amiet, J.-L. (1980a), Amiet, J.-L. and Schiøtz, A. (1972),
Gartshore, M.E. (1986)
Data Providers: Jean-Louis Amiet
154
EN Leptodactylodon stevarti Rödel and Pauwels, 2003
Endangered B1ab(iii)
Order, Family: Anura, Astylosternidae
Country Distribution: Equatorial Guinea, Gabon
Current Population Trend: Decreasing
VU Leptodactylodon ventrimarmoratus (Boulenger, 1904)
Vulnerable B1ab(iii)
Order, Family: Anura, Astylosternidae
Country Distribution: Cameroon
Current Population Trend: Decreasing
© Jean-Louis Amiet
EN Leptodactylodon wildi Amiet and Dowsett-Lemaire, 2000
Endangered B1ab(iii)
Order, Family: Anura, Astylosternidae
Country Distribution: Cameroon
Current Population Trend: Decreasing
© Chris Wild
VU Barbourula busuangensis Taylor and Noble, 1924
Vulnerable B1ab(iii)
Order, Family: Anura, Bombinatoridae
Country Distribution: Philippines
Current Population Trend: Decreasing
© Jaime Bosch
Geographic Range This species occurs on the islands of Busuanga, Culion, and Palawan, all in the western Philip-
pines. It probably occurs more widely than current records suggest, especially in areas between known sites on
Palawan. It is a lowland species occurring up to 300m asl, perhaps up to 500m asl.
Threatened Amphibians of the World
Geographic Range This species was discovered in the Monts de
Cristal in north-western Gabon at 460-550m asl. Specimens assigned
to this species have also been found at Monte Alen in Equatorial
Guinea, where it might occur up to 1,000m asl.
Population It has only recently been described, and only two
specimens are known from Gabon. Its population status in Equatorial
Guinea is difficult to assess, since it is hard to locate.
Habitat and Ecology It has been found only in primary forest under
rocks along streams in deep, narrow valleys. It breeds in fast-flow-
ing, rocky streams.
Major Threats It is at risk from habitat loss due to smallholder
farming activities and logging.
Conservation Measures It occurs in the Monte Alen National Park
in Equatorial Guinea and in the recently gazetted (2002) Monts de
Cristal National Park in Gabon.
BOMBINATORIDAE
Notes on taxonomy: We follow Rödel and Pauwels (2003) in provisionally assigning records of Leptodactylodon albiventris from
Equatorial Guinea (De la Riva 1994) to this newly described species.
Bibliography: De la Riva, I. (1994b), De la Riva, I., Bosch, J. and Marquez, R. (2003), Lasso, C.A. et al. (2002), Rödel, M.-O. and Pauwels,
O.S.G. (2003)
Data Providers: Mark-Oliver Rödel
Geographic Range This species occurs in the lowlands of south-western Cameroon, south of the Sanaga River,
at 50-1,150m asl.
Population It is not uncommon in its favoured habitat, though it is less common on the coastal plain.
Habitat and Ecology It is strictly a forest species, not occurring in secondary habitats. Outside the breeding season
it can be found among dead leaves on the forest floor. During the breeding season, males call from rocky areas by
streams and springs in forest, especially in valley heads where there are boulders under which tiny currents of water
flow. Breeding takes place in streams.
Major Threats The major threat to the species is habitat loss as a result of forest loss and fragmentation due to
logging.
Conservation Measures It might occur in a few protected areas, though this is not confirmed. There is a need for
improved habitat protection at sites where the species is known to occur.
Bibliography: Amiet, J.-L. (1970b), Amiet, J.-L. (1980a), Amiet, J.-L. (1987), Amiet, J.-L. and Perret, J.-L. (1969), Amiet, J.-L. and
Schiøtz, A. (1972)
Data Providers: Jean-Louis Amiet
Geographic Range This recently described species is known only from the vicinity of the villages of Kodmin and Edib
in the Bakossi Mountains, 1,000-1,350m asl, in western Cameroon, although it might be a bit more widespread.
Population It is generally an uncommon species.
Habitat and Ecology It lives in montane forest, often close to small streams; there is no information on its ability
to tolerate changes to its habitat. Breeding takes place in springs and rivulets.
Major Threats It is probably threatened by forest clearance for smallholder farming.
Conservation Measures It does not occur in any protected areas, and its remaining habitat in the Bakossi Mountains
requires urgent protection.
Bibliography: Amiet, J.-L. and Dowsett-Lemaire, F. (2000)
Data Providers: Jean-Louis Amiet
PHILIPPINE FLAT-HEADED FROG
Population It commonly occurs in large numbers where known, though the overall distribution is patchy and
fragmented.
Habitat and Ecology This aquatic species inhabits clear, unpolluted swift-flowing mountain streams and rivers in
lowland rainforests, where it usually floats near the surface of the water unless disturbed, when it will hide under
submerged rocks. The breeding strategy of this species remains unknown. Tadpoles have never been observed, and
eggs collected from gravid females were large, unpigmented, and few in number (Inger 1954), suggesting that it
could possibly reproduce by direct development (Brown and Alcala 1983). However to date breeding habits have not
yet been observed. It has not been found in open habitats outside forest.
Major Threats The lowland rainforest habitat of this species is generally protected on the island of Palawan.
However, some populations are threatened by habitat conversion, shifting agriculture, quarrying, large-scale mining,
and pollution of streams and rivers from agricultural effluents. The collection of frogs for the pet trade (including
internationally) is also a possible threat.
Conservation Measures There is a need for remaining intact lowland rainforest of Palawan to be designated as
protected areas. Further research is needed to establish the levels of offtake of this species from the wild for the
international pet trade; if proved to be a significant threat, then this species could also warrant listing by CITES.
Bibliography: Alcala, A.C. and Brown, W.C. (1985), Brown, R.M., Diesmos, A.C. and Alcala, A.C. (2001), Brown, W.C. and Alcala,
A.C. (1983), Frost, D.R. (1985), Infante, C.R. et al. (2002), Inger, R.F. (1954), Inger, R.F. (1999), Myers, G.S. (1943), Taylor, E.H. and Noble,
G.K. (1924)
Data Providers: Arvin Diesmos, Angel Alcala, Rafe Brown, Leticia Afuang, Genevieve Gee, Katie Hampson, Mae Leonida Diesmos,
Aldrin Mallari, Perry Ong, Dondi Ubaldo, Baldwin Gutierrez
 Globally Threatened Amphibian Species 155

EN Barbourula kalimantanensis Iskandar, 1978 BORNEAN FLAT-HEADED FROG

Endangered B2ab(iii) Geographic Range This species is known only from West Kalimantan in Indonesian Borneo, where it is presently
Order, Family: Anura, Bombinatoridae known only from two localities in the middle of the Kapuas River basin: Nanga Sayan (0° 44’S; 111° 40’E), 33km south
Country Distribution: Indonesia of Nanga Pinoh; and Sungai Kelawit (0° 37’S; 111° 47’E), in the Melawi River basin, about 1km upstream from Nanga
Current Population Trend: Decreasing Pintas. It presumably ranges at least a little more widely, but it probably has a restricted distribution.
Population It is known only from two specimens collected 20 years apart. Directed surveys, in suitable habitat,
close to the type locality have not recovered any further populations. This suggests that either the species has a very
limited distribution or that it exists in very small numbers (R.F. Inger pers. comm.).
Habitat and Ecology Both known specimens were collected in clear rocky rivers, about 20-50m wide, under large
rocks in midstream within tropical moist forest. The morphology of these frogs and the sites of collection indicate that
this species is fully aquatic, and is probably dependent upon clear water with a high oxygen content. Breeding ecology
is not known, but it possibly breeds by direct development. It has not been found in open areas outside forest.
© David Bickford

Major Threats The area where this species occurs is now very highly disturbed due to illegal gold mining, and the
rivers have become severely degraded as a result of siltation and also pollution with mercury waste (following its
use for gold extraction). It is probably also affected by siltation of its aquatic habitat following deforestation of the
surrounding land.
Conservation Measures It is not known from any protected areas, and effective conservation of large areas of
remaining forest habitat is urgently required to help prevent the siltation of rivers.
Bibliography: Iskandar, D.T. (1978), Iskandar, D.T. (1995)
Data Providers: Robert Inger, Indraneil Das, Robert Stuebing, Maklarin Lakim, Paul Yambun

VU Bombina fortinuptialis Tian and Wu, 1978 LARGE-SPINED BELL TOAD

© Michael Wai Neng Lau / Kadoorie Farm & Botanic Garden

Vulnerable B1ab(iii)+2ab(iii) Geographic Range This species is known only from Dayaoshan, Pingnan and Longsheng in Guangxi province,
Order, Family: Anura, Bombinatoridae China, from 1,200-1,640m asl.
Country Distribution: China Population It is very rare, with only a few specimens having been collected since it was described.
Current Population Trend: Decreasing Habitat and Ecology This species inhabits forests, and has not been found outside forested areas. It breeds in pools
and slow-flowing waters where the larvae also develop.
Major Threats The major threat is habitat destruction and degradation due to smallholder farming activities and
logging.
Conservation Measures This species occurs in several nature reserves, including the Dayaoshan National Nature
Reserve.
Bibliography: Fei, L. et al. (1999), MacKinnon, J. et al. (1996), Tian, W.S. and Wu, G.H. (1978)
Data Providers: Michael Wai Neng Lau, Yuan Zhigang

VU Bombina lichuanensis Ye and Fei, 1994 LICHUAN BELL TOAD

Vulnerable B1ab(iii) Geographic Range This species is known only from Beiyang, Lichuan in Hubei Province and Mabian County in
Order, Family: Anura, Bombinatoridae Sichuan Province, China, at around 1,830m asl. It probably occurs more widely than current records suggest, especially
© Fei, L., Ye, C.-Y., Huang, Y.-Z. and Liu, M.-Y. 1999

Country Distribution: China in areas between the two known sites.

Current Population Trend: Decreasing Population It is very rare.
Habitat and Ecology This species inhabits marshes in forested, mountainous areas. It breeds in pools where the
larvae also develop.
Major Threats The major threat to this species is forest loss due to small-scale agriculture and human settle-
ment.
Conservation Measures The population of this species in Hubei Province is probably within Lichuanxiaoheshuisha
Nature Reserve.
Bibliography: Fei, L. et al. (1999), MacKinnon, J. et al. (1996), Ye, C.-Y. and Fei, L. (1994)
Data Providers: Fei Liang, Zhao Wenge

VU Bombina microdeladigitora Liu, Hu and Yang, 1960 SMALL-WEBBED BELL TOAD

Vulnerable B2ab(iii) Hoang Lien Son mountain range near Sa Pa, Lao Cai Province, and from Ha Giang Province in the north. It has not
Order, Family: Anura, Bombinatoridae been found in other surveys further south in the Hoang Lien Son mountain range in Viet Nam. It probably occurs more
Country Distribution: China, Viet Nam widely than current records suggest, especially in areas between the known sites in China and those in Viet Nam. It
Current Population Trend: Decreasing has been recorded from 1,830-2,200m asl.
Population Across much of its range, this species is uncommon and difficult to find.
Habitat and Ecology It inhabits marshes in forests, grassland, and the surrounding areas. Males call from tree
holes, which is also where the eggs are laid (rather than in ponds); the water in these holes overflows during heavy
rains and the tadpoles are washed into streams.
Major Threats The major threat to this species is habitat loss due to the collection of wood for subsistence use,
and agricultural activities, including the planting of cardamom plantations (which are even found within Hoang
Lien Son National Park). In Viet Nam, an additional threat is posed by touristic activities on the Hoang Lien Son
© Nikolai L. Orlov

Geographic Range This species is known from China and Viet Nam. In China it is restricted to western Yunnan
(Jingdong, Yongde and Xishuangbanna) and in Viet Nam it is known from two localities, from high altitudes in the
Mountains.
Conservation Measures The range of this species overlaps with a few protected areas in China and Viet Nam,
including Mount Tay Con Linh II and Hoang Lien Son National Park in Viet Nam.
Notes on taxonomy: In the past, populations of this species in northern Viet Nam were considered to belong to Bombina maxima
(Ohler et al. 2000).
Bibliography: Bourret, R. (1942), Fei, L. et al. (1999), MacKinnon, J. et al. (1996), Ohler, A. et al. (2000), The Comprehensive Scientific
Expedition to the Qinghai-Xizang Plateau (1997), Tran, K. et al. (1992), Yang, D.-T. (1991b)
Data Providers: Peter Paul van Dijk, Steven Swan, Yang Datong, Annemarie Ohler
156 Threatened Amphibians of the World

BUFONIDAE

CR Adenomus dasi Manamendra-Arachchi and Pethiyagoda, 1998

Critically Endangered B1ab(iii)+2ab(iii) Geographic Range This species is restricted to a single locality “Moray Estate, Rajamally, near Mousakelle, alt.
Order, Family: Anura, Bufonidae 1,370m asl” in the Peak Wilderness Sanctuary of the Adam’s Peak area in Sri Lanka. It had previously been collected
Country Distribution: Sri Lanka in the late 1890s at the town Nuwara Eliya (1,710m asl), but is now considered to be extinct at this location (K.
Current Population Trend: Decreasing Manamendra-Arachchi pers. comm.).
Population This is a rare species, now known only from the Moray Estate. There is no information on its current
population status.
Habitat and Ecology It is associated with hill streams in tropical moist forest. Adults can be found on mossy wet
rocks within streams, and in riparian leaf-litter. Breeding and larval development take place in streams.
© V.A.M.P.K. Samarawickrama

Major Threats The extremely restricted distribution of this species makes it very susceptible to water pollution from
adjacent tea plantations and the possible negative impacts of tourism within the area.
Conservation Measures The species is present in the 224-km² Peak Wilderness Sanctuary. Additional survey work
is needed to determine the population status of the species.
Bibliography: Manamendra-Arachchi, K. and Pethiyagoda, R. (1998)
Data Providers: Kelum Manamendra-Arachchi, Anslem de Silva

EN Adenomus kelaartii (Günther, 1858)

Endangered B1ab(iii)+2ab(iii) Geographic Range This species is a Sri Lankan endemic, and is found over much of the south-west of the island.
Order, Family: Anura, Bufonidae It has been recorded at elevations between 30 and 1,230m asl.
Country Distribution: Sri Lanka Population It is fairly common where it occurs.
Current Population Trend: Decreasing Habitat and Ecology It is normally terrestrial, but also sometimes semi-arboreal, and is associated with hill streams
in tropical wet forest. Adults have been observed in rock crevices, leaf-litter and decaying logs, and within tree holes.
Larvae are found in permanent pools.
Major Threats The major threat to the species is habitat loss and degradation due to the clearance (clear cutting)
of forested areas, agrochemical pollution, and expansion of cardamom plantations.
Conservation Measures The species has been recorded from several protected areas, including the Peak Wilder-
ness Forest Reserve, Sinharaja World Heritage Site, Kanneliya Reserve Forest, Haycock Forest Reserve, Gilimale-
Eratne Forest Reserve and Kitulgala Forest Reserve. Continued and improved management of these reserves is
recommended.
© Peter Janzen

Bibliography: Dutta, S.K. and Manamendra-Arachchi, K. (1996), Haas, W., Lehr, E. and Köhler, G. (1997), Manamendra-Arachchi, K.
and Pethiyagoda, R. (1998)
Data Providers: Kelum Manamendra-Arachchi, Anslem de Silva, Deepthi Wickramasinghe

EN Altiphrynoides malcolmi (Grandison, 1978)

Endangered B1ab(iii)+2ab(iii) Geographic Range This species is endemic to the Bale Mountains at 3,200-4,000m asl, in Bale Province, Ethio-
Order, Family: Anura, Bufonidae pia.
Country Distribution: Ethiopia Population During the period 1971-1986, it was found to be common at several sites within its very limited geo-
Current Population Trend: Decreasing graphical range. The lack of recent records is due to the absence of survey work.
CITES: Appendix I Habitat and Ecology This species spans the transition from Schefflera-Hagenia-Hypericum forest to Afro-alpine
moorland. Egg strings are deposited amongst moist herbaceous vegetation where development proceeds to meta-
morphosis through a terrestrial larval stage.
Major Threats The main threat is probably timber extraction from high-altitude forest, and there is some evidence
that this may already have occurred at the type locality.
Conservation Measures At least one of the sites where this toad remains common lies within the Bale Mountains
National Park, although this protected area is not formally gazetted. Nonetheless, there is a long-running conservation
programme in the Bale Mountains National Park (led by the Ethiopian Wolf Conservation Programme). The priority
© Colin Tilbury

action for the species is the protection of remaining montane forest habitats from commercial and subsistence
exploitation. It is listed in Appendix I of CITES, though it is not present in international trade.
Bibliography: Grandison, A.G.C. (1978), Largen, M.J. (2001), Wake, M.H. (1980)
Data Providers: Malcolm Largen

CR Andinophryne colomai Hoogmoed, 1985

Critically Endangered B1ab(iii) Geographic Range This species is known only from the type locality (“Cabacera del Rio Baboso, cerca a Lita’, Carchi
Order, Family: Anura, Bufonidae Province, Ecuador”), and one other locality close by in the province of Carchi, in the north-western Andes of Ecuador.
Country Distribution: Ecuador It has been recorded from 1,180-1,400m asl.
Current Population Trend: Decreasing Population It is a very rare species, and there have been no records since September 1984. Subsequent visits to the
type locality have failed to find any individuals.
Habitat and Ecology The holotype was collected on a branch some 50cm above the ground in the forest of a small
creek at night (Hoogmoed 1985). A male was collected at night sitting on a tree trunk in a creek, while a female was
collected in the afternoon on the forest floor near a creek (Hoogmoed 1989).
Major Threats The type locality has been severely impacted by habitat destruction as a result of agriculture and
logging. In addition, spraying of herbicides in Colombia to control crops is polluting the species’ habitat.
Conservation Measures It is not known to occur in any protected areas. Survey work is needed to determine the
© Luis A. Coloma

population status of this species.

Bibliography: Hoogmoed, M.S. (1985), Hoogmoed, M.S. (1989)
Data Providers: Luis A. Coloma, Santiago Ron, Mario Yánez-Muñoz, Diego Cisneros-Heredia

EN Ansonia anotis Inger, Tan and Yambun, 2001
Endangered B1ab(iii)+2ab(iii)
Order, Family: Anura, Bufonidae
Country Distribution: Malaysia
Current Population Trend: Decreasing
© Paul Yambun
VU Ansonia fuliginea (Mocquard, 1890)
Vulnerable D2
Order, Family: Anura, Bufonidae
Country Distribution: Malaysia
Current Population Trend: Unknown
EN Ansonia guibei Inger, 1966
Endangered B1ab(iii)+2ab(iii)
Order, Family: Anura, Bufonidae
Country Distribution: Malaysia
Current Population Trend: Decreasing
© Indraneil Das
EN Ansonia latidisca Inger, 1966
Endangered B1ab(iii)+2ab(iii)
Order, Family: Anura, Bufonidae
Country Distribution: Indonesia, Malaysia
Current Population Trend: Decreasing
Globally Threatened Amphibian Species
Geographic Range This Bornean endemic is known only from
the montane forests of Kinabalu, Malaysia. Attempts to locate the
species on Gunung Mulu in Sarawak have not been successful. The
altitudinal range is 1,500-3,050m asl.
Population The current population status of this species is unknown.
It is rarely encountered.
Habitat and Ecology Most records are from moist montane and
sub-alpine forest above 2,500m asl. Adults are terrestrial, and breed-
ing is presumed to take place in forest streams.
Major Threats The major threat to this species in the future is likely
to be selective logging.
Conservation Measures It occurs in Kinabalu National Park, which
is well protected. Surveys of potentially suitable areas of habitat in
adjacent parts of Borneo are needed to determine whether or not this
species might occur elsewhere, and also to help better understand
its current population status.
Geographic Range This species is known only from two locations
in the western corner of Borneo: Mount Damus, near Sambas, in
Kalimantan (Indonesia), and Mount Penrissen, in western Sarawak
(Malaysia). It possibly occurs more widely than current records sug-
gest, especially in areas between the two known sites.
Population There are no estimates of population size. Only two
specimens are known and it was last collected over 50 years ago.
There are no recent records, probably due to lack of herpetological
work within its range.
Habitat and Ecology The known localities were primary, hilly
rainforest at the time the frogs were collected. The adults are
primarily terrestrial, and the larvae are found in forest streams. If it
is similar to other members of its genus, it is unlikely to be able to
adapt to modified habitats.
Major Threats The main threat to this species is habitat loss and
degradation primarily as a result of logging. The remaining suitable
157
Geographic Range This species is known only from two localities, Sayap Station and Sungai Purulon, Sabah (Borneo),
Malaysia. It presumably occurs elsewhere, in particular in areas between the confirmed localities. The altitudinal
range of this species is 300-1,000m asl.
Population The species is rarely encountered, and good estimates of population size are lacking.
Habitat and Ecology It is a terrestrial species of lowland and submontane tropical moist forest, and the larvae
inhabit clear forest streams. This species probably cannot adapt to modified habitats.
Major Threats The main threat to it is habitat loss and degradation as a result of the destruction of the forest through
logging and clearance for rice cultivation, and the consequent siltation of streams.
Conservation Measures The species has been recorded from Kinabalu National Park and the Crocker Range
National Park. Improved management of these areas and expanded protection of other remaining tracts of lowland
forest are essential.
Bibliography: Inger, R.F., Lian, T.-F. and Yambun, P. (2001), Inger, R.F., Tan, F.-L. and Yambun, P. (2001)
Data Providers: Robert Inger, Indraneil Das, Robert Stuebing, Maklarin Lakim, Paul Yambun
Notes on taxonomy: We follow Inger (1966) and Malkmus et al. (2002) in considering Ansonia altitudinis (Smith 1931) to be a synonym
of A. fuliginea.
Bibliography: Inger, R.F. (1960a), Inger, R.F. (1966), Inger, R.F. and Stuebing, R.B. (1997), Malkmus, R. et al. (2002), Matsui, M., Hikida,
T., and Namu, H. (1985)
Data Providers: Robert Inger, Indraneil Das, Robert Stuebing, Maklarin Lakim, Paul Yambun
Geographic Range This species is endemic to a portion of montane north-western Sabah (northern Borneo). A large
subpopulation exists at one corner of Kinabalu National Park between 1,600 and 2,000m asl. A small population has
also been found on Mount Trus Madi at 1,300m asl. Attempts to locate the species on Gunung Mulu in Sarawak have
not been successful. It has not been found at other montane sites south of Trus Madi.
Population It is sometimes locally very abundant, especially along streams.
Habitat and Ecology Adults disperse widely over the floor of montane and submontane forests. It breeds in small,
clear, rocky-bottomed streams and larvae live in torrents, clinging to rocks and feeding on lithophytes. It appears to
be unable to adapt to modified habitats.
Major Threats As Trus Madi is being actively logged, it is likely that the population there will be diminished, if not
driven to extinction. Destruction of the forest at Trus Madi will probably lead to the siltation of streams in which the
larvae develop. If the species occurs at other montane sites, these are likely to remain as insular isolates because
of the extensive deforestation taking place at lower elevations.
Conservation Measures The species occurs in Kinabalu National Park, which is well protected. However, the area
where this species is particularly abundant is increasingly subject to the impact of tourist activities. There is a particular
need to expand the existing protected area network (especially above 1,200m asl) south of Kinabalu National Park.
Bibliography: Inger, R.F. (1966), Inger, R.F. and Stuebing, R.B. (1997), Malkmus, R. (1994), Malkmus, R. et al. (2002), Malkmus, R., and
Kosuch, J. (2000)
Data Providers: Robert Inger, Indraneil Das, Robert Stuebing, Maklarin Lakim, Paul Yambun
habitat within its range has been almost entirely converted for recreational use (one of the two known sites, Mount
Penrissen, has recently been converted into a golf course), or converted to cultivated land. An additional threat is the
resultant sedimentation of streams (following logging) which results in the deterioration of breeding habitat.
Conservation Measures Effective preservation of hilly rainforest in the regions of the two known localities is
essential. In addition, further survey work in the hilly border area of Sarawak and Kalimantan is necessary to help
provide a better indication of the species’ current population status.
Bibliography: Inger, R.F. (1966)
Data Providers: Robert Inger, Indraneil Das, Robert Stuebing, Maklarin Lakim, Paul Yambun
158 Threatened Amphibians of the World

VU Ansonia mcgregori (Taylor, 1922)

Vulnerable B1ab(iii) Geographic Range This species is found in the mountains of central and western Mindanao Island in the Philippines.
Order, Family: Anura, Bufonidae It probably occurs more widely than current records suggest.
Country Distribution: Philippines Population It is fairly common.
Current Population Trend: Decreasing Habitat and Ecology It inhabits cool mountain streams and rivers in lower montane and lowland forests. It breeds
in streams, which is where the larvae also develop.
Major Threats The major threats include habitat loss and deforestation, due to agriculture and logging, and water
pollution.
Conservation Measures It is recorded from several protected areas on Mindanao, including Mount Malindang
National Park and Mount Apo and Mount Kitanlad Range Natural Parks. Conservation measures should include the
regulation and proper disposal of pesticides and herbicides, and the protection of the remaining rainforest, especially
riverine habitats and gallery forest, on Mindanao.
© Arvin C. Diesmos

Bibliography: Alcala, A.C. and Brown, W.C. (1985), Brown, R.M., Diesmos, A.C. and Alcala, A.C. (2001), Crombie, R.A. (n.d.), Frost, D.R.
(1985), Inger, R.F. (1954), Inger, R.F. (1999), Taylor, E.H. (1923)
Data Providers: Arvin Diesmos, Angel Alcala, Rafe Brown, Leticia Afuang, Genevieve Gee, Katie Hampson, Mae Leonida Diesmos,
Aldrin Mallari, Perry Ong, Dondi Ubaldo, Baldwin Gutierrez

VU Ansonia muelleri (Boulenger, 1887)

Vulnerable B1ab(iii) Geographic Range This species is found in the mountains of Mindanao and Dinagat Islands, in the Philippines.
© Rafe Brown, 2006. Courtesy of HerpWatch Philippines

Order, Family: Anura, Bufonidae Population It is fairly common where it occurs, but it has a patchy distribution.
Country Distribution: Philippines Habitat and Ecology It inhabits cool mountain streams and rivers in montane rainforest where it also breeds and
Current Population Trend: Decreasing the larvae develop.
Major Threats This species inhabits high-elevation forests, which are generally less threatened by habitat conver-
sion, agriculture, and human encroachment, and the pollution of streams and rivers. Regardless, some populations
of this species are subject to these threats.
Conservation Measures It is known from several protected areas, including Mount Malindang National Park.
Conservation measures must include the regulation and proper disposal of pesticides and herbicides, and the protec-
tion of the remaining rainforest, especially high-elevation riverine habitats and gallery forests, on Mindanao. The
taxonomic status of this species requires further study.
Bibliography: Alcala, A.C. and Brown, W.C. (1985), Brown, R.M., Diesmos, A.C. and Alcala, A.C. (2001), Crombie, R.A. (n.d.), Frost, D.R.
(1985), Inger, R.F. (1954), Inger, R.F. (1999), Taylor, E.H. (1923)
Data Providers: Arvin Diesmos, Angel Alcala, Rafe Brown, Leticia Afuang, Genevieve Gee, Katie Hampson, Mae Leonida Diesmos,
Aldrin Mallari, Perry Ong, Dondi Ubaldo, Baldwin Gutierrez

EN Ansonia ornata Günther, 1876

Endangered B1ab(iii) Geographic Range This species is known with certainty only from the type locality of “Brumagherries (Brahmagiri
Order, Family: Anura, Bufonidae Hills)” and adjoining areas of Coorg, Karnataka, in the southern Western Ghats. There are also unpublished reports of
Country Distribution: India this species from Kudremukh National Park (Karnataka), Silent Valley National Park and Wayanad Wildlife Sanctuary
Current Population Trend: Decreasing (Kerala) (Amphibian CAMP 2002). However, further survey work is needed to confirm that the species does occur at
these sites (S.D. Biju pers comm.) and these additional areas are currently not included in the map for this species.
It has been recorded at an altitude of around 1,000m asl.
Population It is an uncommon species.
Habitat and Ecology It is presumably restricted to tropical evergreen forest. Its breeding is not known, but it
presumably breeds by larval development in streams, like other members of its genus.
Major Threats The main threat to this species is continuing deforestation (mainly for the cultivation of coffee) in
the Coorg area and surroundings.
Conservation Measures This species is currently not known with certainty from any protected areas, and strength-
ened and expanded protection of the remaining forest habitat in this region of the Western Ghats is necessary. It is
© S.D. Biju

included as part of ongoing field studies begun in 1998 (S.D. Biju pers. comm.).
Notes on taxonomy: There are taxonomic difficulties with this species, and it is possible that Ansonia rubigina is a synonym of it
(S.D. Biju pers. comm.).
Bibliography: Biju, S.D. (2001), Chanda, S.K. and Deuti, K. (1997), Daniel, J.C. (1963), Daniels, R.J.R. (1991), Daniels, R.J.R. (1997),
Dutta, S.K. (1997), Günther, A. (1876), Ravi Kumar, M.V. (2000), Reddy, A.H.M. et al. (2003)
Data Providers: S.D. Biju, Sushil Dutta, M.S. Ravichandran

VU Ansonia penangensis Stoliczka, 1870

Vulnerable D2 Geographic Range This species is known only from Penang Island, Notes on taxonomy: There is much confusion surrounding the status of this species relative to Ansonia malayana. Populations that
Order, Family: Anura, Bufonidae Malaysia. Its altitudinal range is not known. have been assigned to this species on the Isthmus of Kra and from Fraser’s Hill in Malaysia are considered here to refer to A. malayana.
Country Distribution: Malaysia Population There is very little information. The collection of tadpoles However, it is not clear that these two species are distinct from each other, and further taxonomic work is needed. Records of A.
Current Population Trend: Stable on Penang Hill in 2004 represents the first records of the species penangensis from Sumatra, Indonesia, are now assigned to A. glandulosa.
in over 100 years; this absence of records might be a reflection of Bibliography: Dring, J.C.M. (1979), Leong, T.M. and Lim, K.K.P. (2003), Taylor, E.H. (1962)
inadequate surveying effort. Data Providers: Peter Paul van Dijk, Jeet Sukumaran, Indraneil Das, Norsham Yaakob, Leong Tzi Ming, Yodchaiy Chuaynkern
Habitat and Ecology This is a stream-breeding species that inhabits
rocky streams in rainforests.
Major Threats The forest habitat at the type locality is well pro-
tected; other potential threats are presently unknown.
Conservation Measures The hill forests on Penang Hill are cur-
rently protected as a catchment area for Georgetown, and the lower
reaches are within the Penang Botanical Gardens. Nonetheless, the
population status of this species requires careful monitoring, given
that it is known only from a single location.

EN Ansonia platysoma Inger, 1960
Endangered B1ab(iii)+2ab(iii)
Order, Family: Anura, Bufonidae
Country Distribution: Malaysia
Current Population Trend: Decreasing
© Indraneil Das
VU Ansonia rubigina Pillai and Pattabiraman, 1981
Vulnerable D2
Order, Family: Anura, Bufonidae
Country Distribution: India
Current Population Trend: Stable
© S.D. Biju
VU Ansonia siamensis Kiew, 1984
Vulnerable D2
Order, Family: Anura, Bufonidae
Country Distribution: Thailand
Current Population Trend: Unknown
VU Ansonia tiomanica Hendrickson, 1966
Vulnerable D2
Order, Family: Anura, Bufonidae
Country Distribution: Malaysia
Current Population Trend: Stable
© Jim McGuire
Globally Threatened Amphibian Species
Geographic Range This species is known from a single locality in
the Khao Chong Mountains of Peninsular Thailand. It presumably
occurs more widely within these mountains, as has been mapped.
The type locality is at an elevation of 300m asl.
Population It is apparently uncommon. A pair was collected in 1979,
and a further five individuals have been collected since (Kiew 1984b,
Matsui, Nabhitabhata and Panha 1998).
Habitat and Ecology It is restricted to lowland primary tropical
moist forest. It is a stream-breeding species.
Major Threats The habitat of this species appears to be reasonably
well protected at present, and there are currently no major threats.
However, given its restricted range, it is potentially at risk from
stochastic events and pollution from adjacent tourist sites.
Conservation Measures The known range is entirely within the
Khao Chong Mountains protected area. The population status of
this species requires careful monitoring, given that it is known only
from a single location.
159
Geographic Range This Bornean endemic occurs at a number of sites within Kinabalu National Park and at other
sites in the Crocker Range south of Kinabalu in Sabah (Malaysia); it is also known from Gunung Mulu National Park
in Sarawak. The altitudinal range in Kinabalu is 600-950m asl (Malkmus et al. 2002), and its overall altitudinal range
is 750-1,600m asl.
Population This species is abundant at most locations where it has been reported (such as the Crocker Range).
Habitat and Ecology Adults are found on the floor of montane forests and move to clear, rocky mountain creeks to
breed. The larvae cling to rocks in torrents in these streams and presumably feed on lithophytes. It appears not to
be able to survive in modified habitats.
Major Threats Logging in the submontane and montane forests could potentially lead to siltation of the streams
needed for larval development and result in loss of the lithophytes.
Conservation Measures The species is known to be present in the Kinabalu and Gunung Mulu National parks, both
of which are reasonably well managed; however, further protection of the remaining forest habitats, and continued
management of the existing protected areas, are needed. Additional surveys of potentially suitable sites in Kalimantan
are required to establish whether this species occurs there.
Bibliography: Inger, R.F. (1960a), Inger, R.F. (1966), Inger, R.F. and Stuebing, R.B. (1997), Malkmus, R. et al. (2002)
Data Providers: Robert Inger, Indraneil Das, Robert Stuebing, Maklarin Lakim, Paul Yambun
Geographic Range This species is present only in the Silent Valley and Wayanad areas of Kerala State in the southern
Western Ghats of India (Biju 2001) where it has been recorded at elevations between 1,000 and 1,200m asl.
Population It is a rare species, though populations are generally believed to be stable within the Silent Valley
National Park.
Habitat and Ecology It is restricted to tropical evergreen forest, and is associated with torrential streams and the
forest floor. Breeding presumably takes place by larval development in streams.
Major Threats There are currently no major threats to this species.
Conservation Measures It is currently considered to be adequately protected within the Silent Valley National
Park. Nonetheless, the population status of this species requires careful monitoring, given that it is known only
from two locations.
Notes on taxonomy: This species might be a synonym of Ansonia ornata (S.D. Biju pers. comm.).
Bibliography: Biju, S.D. (2001), Dutta, S.K. (1997), Pillai, R.S. and Pattabiraman, R. (1981)
Data Providers: S.D. Biju, Sushil Dutta, M.S. Ravichandran
Bibliography: Kiew, B.H. (1984b), Matsui, M., Nabhitabhata, J. and Panha, S. (1998)
Data Providers: Peter Paul van Dijk, Jeet Sukumaran, Norsham Yaakob, Leong Tzi Ming, Yodchaiy Chuaynkern
Geographic Range This species is known only from two localities on Tioman Island, Malaysia, at an altitude of
300-1,000m asl.
Population The population status of this species is unknown, but it is presumed to be stable at present.
Habitat and Ecology It is associated with boulders and waterfalls in caves. Breeding is presumed to take place
in streams.
Major Threats The species’ habitat is not currently under immediate threat, but the potential clearance of habitat
for tourism infrastructure is of concern for the future.
Conservation Measures Tioman Island has been designated as a protected area; continued enforcement of this
protected status is required. Safeguarding caves and other high-altitude habitats on Tioman Island is needed. The
population status of this species requires careful monitoring, given that it is known only from a single location.
Bibliography: Hendrickson, J.R. (1966)
Data Providers: Peter Paul van Dijk, Jeet Sukumaran, Norsham Yaakob, Leong Tzi Ming
160
VU Ansonia torrentis Dring, 1983
Vulnerable D2
Order, Family: Anura, Bufonidae
Country Distribution: Malaysia
Current Population Trend: Stable
EN Atelophryniscus chrysophorus McCranie, Wilson and Williams, 1989
Endangered A2ac; B1ab(iii,v)
Order, Family: Anura, Bufonidae
Country Distribution: Honduras
Current Population Trend: Decreasing
© James McCranie
CR Atelopus andinus Rivero, 1968
Critically Endangered A3ce
Order, Family: Anura, Bufonidae
Country Distribution: Peru
Current Population Trend: Decreasing
© Ted Kahn
CR Atelopus angelito Ardila-Robayo and Ruíz-Carranza, 1998
Critically Endangered A3ce
Order, Family: Anura, Bufonidae
Country Distribution: Colombia
Current Population Trend: Decreasing
Threatened Amphibians of the World
Geographic Range This species is known only from Gunung Mulu
in northern Sarawak, Borneo, Malaysia, at 1,800m asl, though it is
possible that it might occur in other montane forest blocks.
Population It is believed to be reasonably abundant.
Habitat and Ecology Specimens were observed calling beside a
small, clear, mountain stream with a steep gradient. Breeding is
presumed to take place in these forest streams.
Major Threats There are no current major threats to this species,
but if Gunung Mulu is indeed the only locality at which this species
occurs, then it might well be susceptible to stochastic threatening
processes.
Conservation Measures The species has been recorded from Gu-
nung Mulu National Park. Further survey work is needed to ascertain
whether or not this species is indeed confined to Gunung Mulu. In any
event, the population status of this species requires careful monitor-
ing given that it is currently known from only a single location.
Geographic Range This species is known only from near Valencia,
San Sebastián Municipality, in Cauca Department, Colombia, at
elevations of 2,900-3,000m asl. It might occur more widely than
current records suggest.
Population There is very little information available, but it was
collected as recently as 2000.
Habitat and Ecology This species is found on vegetation along
side streams in montane Andean forests and sub-páramo bush land.
There is no information known about breeding habitats, or its ability
to tolerate habitat disturbance.
Major Threats There have been serious declines of other high-eleva-
tion Atelopus species in the region suggesting that this species might
also be at risk. The cause of the declines is still not fully understood,
though chytridiomycosis is almost certainly involved, possibly in
combination with the effects of climate change.
Conservation Measures The type locality is within Parque Na-
Bibliography: Dring, J.C.M. (1984b)
Data Providers: Robert Inger, Indraneil Das, Robert Stuebing, Maklarin Lakim, Paul Yambun
Geographic Range This species occurs in the middle of the Cordillera Nombre de Dios along the Atlantic versant
of north-central Honduras at 750-1,760m asl. It has been collected at only two sites, 60km apart.
Population It is uncommon, and the population has been decreasing over the last 15 years.
Habitat and Ecology It lives in premontane and lower montane wet forest, and is found breeding in clear water
mountain streams.
Major Threats Major threats to this species include landslides in the upper reaches of streams (due to strong storms
and human activities) and slash-and-burn agriculture. As a montane, stream-breeding species, declines could also
be linked to chytridiomycosis.
Conservation Measures The upper elevations in the central Cordillera Nombre de Dios have been protected in
Pico Bonito National Park since 1988 and also in Refugio de Vida Silvestre Texiguat. Research is urgently needed
to determine whether or not this species has been affected by chytridiomycosis. If disease is shown to be a major
threat, then a captive-breeding programme may need to be established.
Bibliography: Lavilla, E.O. and de Sá, R. (2001), McCranie, J.R. and Wilson, L.D. (2002b), McCranie, J.R., Wilson, L.D. and Williams,
K.L. (1989), Wilson, L.D. and McCranie, J.R. (1998)
Data Providers: Gustavo Cruz, Larry David Wilson
Geographic Range This species is restricted to the upper Río Biabo Valley (northern versant of the Cordillera Azul)
(Departamento de San Martín), the Río Pisqui, (Departamento Loreto), and Río Cachiyacu (on the border of Departa-
mentos San Martín and Loreto), Peru. Its recorded altitudinal range is 1,000-2,000m asl.
Population There is no information on its current population status, but it has been seen as recently as 2004 near
Iquitos.
Habitat and Ecology It is a terrestrial species restricted to submontane tropical forest. Breeding is thought to take
place in streams. This species is presumed to be susceptible to habitat change and is therefore not expected to occur
in any modified or degraded habitats.
Major Threats There are no reports of chytridiomycosis impacting this Atelopus species, but it is presumed to be
susceptible to this pathogen, which is now causing amphibian declines in northern Peru. It is possible that populations
of this species at lower altitudes might be able to survive an outbreak of the disease.
Conservation Measures This species is present in the Parque Nacional Cordillera Azul. Given the susceptibility
of this species to chytridiomycosis, successful conservation measures are likely to require some form of disease
management programme and the maintenance of captive populations.
Bibliography: Alverson, W.S., Rodriguez, L.O. and Moskovits, D.K. (2001), Instituto Nacional de Recursos Naturales (INRENA) (2000),
La Marca, E. et al. (2005), Lötters, S. (1996), Lötters, S. et al. (2005), Lötters, S. and de la Riva, I. (1998), Pounds, J.A. et al. (2006), Rivero,
J.A. (1968b), The Field Museum of Natural History (2002)
Data Providers: Stefan Lötters, Antonio Salas, Ariadne Angulo, Javier Icochea, Robert Reynolds, Enrique La Marca
cional Natural Puracé, which might benefit in future from improved management. However, given the likely threat
of chytridiomycosis, disease management and captive-breeding programmes might be required. Research into this
species’ population status and ecological requirements is needed, in particular to determine if it occurs outside the
vicinity of the type locality.
Bibliography: Acosta-Galvis, A.R. (2000), Ardila-Robayo, M.C. and Ruiz-Carranza, P.M. (1998), La Marca, E. et al. (2005), Pounds, J.A.
et al. (2006)
Data Providers: María Cristina Ardila-Robayo, Wilmar Bolívar, Jose Vicente Rueda, Andrés Acosta-Galvis, John Lynch
 Globally Threatened Amphibian Species 161

CR Atelopus arsyecue Rueda-Almonacid, 1994

Critically Endangered A3ce Geographic Range This species is known only from the type locality: Parque Nacional Natural Sierra Nevada de
Order, Family: Anura, Bufonidae Santa Marta in the department of Cesar, Colombia. It has been recorded between 2.000 and 3,500m asl. It might
Country Distribution: Colombia occur a little more widely on the Sierra Nevada de Santa Marta.
Current Population Trend: Decreasing Population Only six specimens of this species are known, and there have been no records since 1991. It is not yet
clear whether or not this reflects lack of survey effort, or a genuine rarity or decline.
Habitat and Ecology It occurs in sub-Andean and Andean forests, sub-páramo and páramo. Egg chains are placed
in fast-flowing water, and the tadpoles develop in the water.
Major Threats The major threat is likely to be chytridiomycosis, leading to a catastrophic population decline, as
has occurred in many other montane species of Atelopus. Habitat loss caused by agricultural expansion and logging
© Juan Manuel Renjifo is also a major threat, as is pollution resulting from the fumigation of crops.
Conservation Measures Its range includes Parque Nacional Natural Sierra Nevada de Santa Marta. Further survey
work is required to determine the population status of this species, especially in light of the recent drastic declines
observed in other montane Atelopus species. Given the likely threat of chytridiomycosis, disease management and
captive-breeding programmes might be required.
Bibliography: Acosta-Galvis, A.R. (2000), La Marca, E. et al. (2005), Lötters, S. (1996), Pounds, J.A. et al. (2006), Rueda-Almonacid, J.V.
(1994b), Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and Lynch, J.D. (1996)
Data Providers: Jose Vicente Rueda, Andrés Acosta-Galvis, Martha Patricia Ramírez Pinilla, Mariela Osorno-Muñoz, Adolfo Amézquita,
María Cristina Ardila-Robayo

CR Atelopus arthuri Peters, 1973

Critically Endangered A2ace; Geographic Range This species is known from three localities on the Pacific versant of the Andes of Ecuador at an
B1ab(iii,v)+2ab(iii,v) altitude of 2,200-3,000m asl. The type locality is 15km north of Pallatanga, in Chimborazo Province. The second and
Order, Family: Anura, Bufonidae third localities are at Cashca Totoras and Las Guardias, respectively, in Bolivar Province.
Country Distribution: Ecuador Population This is an extremely rare species and it has not been recorded since 1988, despite searches, particularly
Current Population Trend: Decreasing at Cashca Totoras (Bustamante 2002). This suggests that a serious decline has taken place.
Habitat and Ecology It inhabits humid montane forest and sub-páramo. There is no specific information known about
breeding habits, though it is likely to be similar to other Atelopus species, with breeding taking place in streams.
Major Threats The major threat is likely to be chytridiomycosis, leading to a catastrophic population decline, as has
occurred in many other montane species of Atelopus. The disease has been confirmed in Gastrotheca pseustes in
the same area. Other major threats includes habitat loss due to agriculture, logging and infrastructure development
for human settlement, and agricultural pollution.
Conservation Measures The range of the species does not encompass any protected areas. However, it was
© Luis A. Coloma

protected in Bosque Protector Cashca Totoras (in Bolívar Province), where it disappeared. The species might be
extinct, and it might be too late for measures such as captive breeding; additional survey work is required to confirm
the continued persistence of this species in the wild.
Bibliography: Bustamante, M. R. (2002), La Marca, E. et al. (2005), Lötters, S. (1996), Merino-Viteri, A. (2001), Peters, J.A. (1973),
Pounds, J.A. et al. (2006), Ron, S.R. and Merino, A. (2000)
Data Providers: Santiago Ron, Luis A. Coloma, Martín R. Bustamante, Diego Cisneros-Heredia, Mario Yánez-Muñoz

CR Atelopus balios Peters, 1973

Critically Endangered A2ace Geographic Range This species is known from only four localities in Azuay, Cañar, and Guayas Provinces in the
Order, Family: Anura, Bufonidae Pacific lowlands of south-western Ecuador, from 200-460m asl.
Country Distribution: Ecuador Population This is now considered a very rare species. There have been no records since April 1995 despite
Current Population Trend: Decreasing repeated searches. It is no longer found at Río Patul (in Azuay province), where it used to be abundant (L. Coloma
pers. comm.).
Habitat and Ecology It is an inhabitant of lowland rainforest, and has been found on riverbanks. There is no specific
information known about breeding habits, though it is likely to be similar to other Atelopus species, with breeding
taking place in streams.
Major Threats This species seems to have disappeared, as with many other Atelopus species, likely due to chy-
tridiomycosis, although the altitude at which it occurs is low (the disease normally occurs at higher altitudes in the
tropics). Habitat degradation and loss, due to agriculture (crops and livestock) and logging, and pollution, continue
© Luis A. Coloma

to be very serious threats.

Conservation Measures The range of the species does not include any protected areas. As the species might be
extinct, it might be too late for conservation measures such as captive breeding; additional survey work is required
to confirm the continued persistence of this species in the wild.
Bibliography: Coloma, L.A. and Lötters, S. (1996), La Marca, E. et al. (2005), Lötters, S. (1996), Parker III, T.A. and Carr, J.L. (1992),
Peters, J.A. (1973), Pounds, J.A. et al. (2006)
Data Providers: Diego Cisneros-Heredia, Mario Yánez-Muñoz, Luis A. Coloma, Santiago Ron

CR Atelopus bomolochos Peters, 1973

Critically Endangered A2ace Geographic Range This species is found in the Cordillera Oriental in southern Ecuador, Azuay and Cañar Provinces,
Order, Family: Anura, Bufonidae between 2,500 and 2,800m asl, where it has been recorded from at least 15 localities. There was one record from
Country Distribution: Ecuador Parque Nacional Sangay. The specimens from the Departamento Piura, Peru, which referred to this species, are in
Current Population Trend: Decreasing fact specimens of an undescribed species (Coloma, Lötters and Salas 2000).
Population One individual was seen in 2002 in the Parque Nacional Sangay (D. Almeida pers. comm.), but otherwise
this formerly abundant species has disappeared from its range.
Habitat and Ecology It lives in humid montane forest, sub-páramo, and páramo (Lötters 1996). Breeding takes
place in streams.
Major Threats The major threat is likely to be chytridiomycosis, leading to a catastrophic population decline, as
has occurred in many other montane species of Atelopus. Dead and dying animals infected with the chytrid fungal
pathogen have been collected in Ecuador (Ron et al. 2003), and it was the first species (in 1980) in Central or South
© Luis A. Coloma

America confirmed to have chytridiomycosis. It tolerates some habitat destruction, and can be found near streams
in artificial grasslands. Introduced predators such as trout might threaten the species.
Conservation Measures The species has been recorded from Parque Nacional Sangay, which is a World Heritage Site.
The population status of this species urgently needs to be assessed; given the threat of chytridiomycosis, successful
conservation measures will probably need to include the maintenance of any surviving individuals in captivity.
Bibliography: Coloma, L.A., Lötters, S. and Salas, A.W. (2000), La Marca, E. et al. (2005), Lötters, S. (1996), Merino-Viteri, A. (2001),
Peters, J.A. (1973), Pounds, J.A. et al. (2006), Ron, S.R. et al. (2003), Ron, S.R. and Merino, A. (2000)
Data Providers: Luis A. Coloma, Santiago Ron, Stefan Lötters, Martín R. Bustamante, Andrés Merino-Viteri, Antonio Salas
162 Threatened Amphibians of the World

CR Atelopus boulengeri Peracca, 1904

Critically Endangered A3ce Geographic Range This species is known from six localities in the provinces of Morona-Santiago and Loja, in the
Order, Family: Anura, Bufonidae south-eastern versant of the Cordillera Oriental, the Cordillera de Cutucú, and the Cordillera del Cóndor, in eastern
Country Distribution: Ecuador Ecuador. It has been recorded from 800-2,000m asl.
Current Population Trend: Decreasing Population It is a rare species, and there have been no records since 1984, although some localities at which the
species is known to occur have not been well surveyed. It is not known precisely whether or not populations have
declined as observed in other Atelopus in the Ecuadorian Andes, though this seems likely.
Habitat and Ecology This species is an inhabitant of humid montane forest. There is no specific information
known about breeding habits, though it is likely to be similar to other Atelopus species, with breeding taking place
in streams.
Major Threats Agriculture, as well as mining and infrastructure development for human settlement, are major threats
to the species’ habitat, and much of the natural vegetation within its known distribution area has been cleared. Pol-
lution of streams is also a threat. It is almost certainly at severe risk from chytridiomycosis.
Conservation Measures The distribution range of this species overlaps with Parque Nacional Sangay, which is a
© Ted Kahn

World Heritage Site. Surveys are urgently needed to determine whether or not this species still persists within its
natural range. Given the threat of chytridiomycosis, successful conservation measures will probably need to include
the maintenance of any surviving individuals in captivity.
Bibliography: Duellman, W.E. and Lynch, J.D. (1988), La Marca, E. et al. (2005), Lötters, S. (1996), Peracca, M.G. (1904), Peters, J.A.
(1973), Pounds, J.A. et al. (2006)
Data Providers: Santiago Ron, Luis A. Coloma, Martín R. Bustamante, Diego Cisneros-Heredia, Ana Almandáriz, Manuel Morales

CR Atelopus carauta Ruíz-Carranza and Hernández-Camacho, 1978 RIO CARAUTA STUBFOOT TOAD
Critically Endangered A3ce Geographic Range This species is known from two localities: the Natural Las Orquideas. More research into the species’ distribution range, ecological requirements, and population
Order, Family: Anura, Bufonidae type locality of Parque Nacional Natural Las Orquideas, and from status is needed, in particular to determine if it occurs outside the vicinity of the two known localities; given the
Country Distribution: Colombia Murri in La Blanquita, both in Antioquia Department, in north-western threat of chytridiomycosis, successful conservation measures will probably need to include the maintenance of any
Current Population Trend: Decreasing Colombia, between 1,300 and 2,000m asl. surviving individuals in captivity.
Population It is uncommon. There do not appear to be any confirmed Bibliography: Acosta-Galvis, A.R. (2000), Cannatella, D.C. (1981), La Marca, E. et al. (2005), Lötters, S. (1996), Pounds, J.A. et al. (2006),
records since 1973, but it is not known if this is indicative of a decline, Ruiz-Carranza, P.M. and Hernández-Camacho, J.A. (1978), Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and Lynch, J.D. (1996)
or simply a lack of survey effort. Data Providers: Andrés Acosta-Galvis, Wilmar Bolívar, Fernando Castro, John Lynch
Habitat and Ecology It occurs on vegetation alongside streams
in sub-Andean forests. It has not been recorded outside forest. Its
breeding habits are not known, though breeding is likely to take
place in streams.
Major Threats The major threat is likely to be chytridiomycosis,
leading to a catastrophic population decline, as has occurred in many
other montane species of Atelopus. Forest destruction for the creation
of fruit plantations is occurring within the species’ range.
Conservation Measures The type locality is within Parque Nacional

CR Atelopus carbonerensis Rivero, 1972 VENEZUELAN YELLOW FROG

Critically Endangered A2ace; B2ab(v) Population Populations of this frog seem to be restricted to the type locality, where it was formerly abundant, but
Order, Family: Anura, Bufonidae is now extremely rare, and possibly even extinct. It was last recorded in 1998, despite intensive searches for the
Country Distribution: Venezuela species. Observations of population declines were made by La Marca (1995b).
Current Population Trend: Decreasing Habitat and Ecology This species is an inhabitant of cloud forest and it is found along streams. Surviving popula-
tions, if it is still extant, are restricted to an isolated patch of forest surrounded by pasturelands. It lays eggs chains
in streams, and the larvae develop in these streams.
Major Threats The first alert about the conservation status of the species was advanced by La Marca and Reinthaler
(1991). As with other Atelopus species, the most likely cause of the dramatic decline is chytridiomycosis, which was
confirmed in this species in 1988. Logging and agricultural expansion, both for crops and livestock, are major threats
to the species’ habitat. The recent introduction of Rana catesbeiana in places near the type locality poses the problem
of a new predator. With higher insolation as a result of climate change, this species could be at increased risk due to
© Pascual Soriani

Geographic Range This species is known from a single locality estimated to be less than 10km² in size (La Marca
1992) in the state of Mérida, in the Venezuelan Andes. In the past, it was most common at the Bosque de San Eusebio
(La Carbonera). It has been recorded from 2,000-2,800m asl.
its habit of basking in the sun. It has been recorded occasionally in the international pet trade although not at levels
thought to pose a major threat to the species.
Conservation Measures This species occurs in Parque Nacional Sierra de La Culata and a few unprotected areas
nearby (La Marca and Lötters 1997). Surveys to determine whether or not this species still survives are urgently needed.
Given the threat of chytridiomycosis, surviving individuals might need to be maintained in captivity.
Notes on taxonomy: This species was elevated to species status by La Marca (1983).
Bibliography: Barrio Amorós, C.L. (2004), La Marca, E. (1983), La Marca, E. (1992), La Marca, E. (1995b), La Marca, E. et al. (2005), La
Marca, E. and Lötters, S. (1997), La Marca, E. and Reinthaler, H.P. (1991), Lötters, S. (1996), Lötters, S., La Marca, E. and Vences, M. (2004),
Pounds, J.A. et al. (2006), Rivero, J.A. (1972), Rodríguez, J.P. and Rojas-Suárez, F. (1995), Torres, D.A. and Barrio, C.L. (2001)
Data Providers: Enrique La Marca, Argelia Rodríguez, Juan Elías García-Pérez

CR Atelopus carrikeri Ruthven, 1916

Critically Endangered A3ce Geographic Range This species is known from the páramos of the Sierra Nevada de Santa Marta, in the department
Order, Family: Anura, Bufonidae of Magdalena, Colombia. It has been recorded between 2,350 and 4,800m asl.
Country Distribution: Colombia Population The current population status of this species is unknown, but it was common in the past (1994). There
Current Population Trend: Decreasing has been no recent survey work due to security problems in the area.
Habitat and Ecology It occurs in sub-Andean and Andean forests, and páramo, and also the lower portion of
snowy areas. It can adapt to some modification of its habitat. It lays egg chains in streams, where the tadpoles
also develop.
Major Threats The most serious threat to this species is the risk of chytridiomycosis, which has had a devastat-
ing impact on other high-altitude Atelopus species. Climate change, habitat loss caused by agriculture, and crop
fumigation, are all also major threats.
© Juan Manuel Renjifo

Conservation Measures Its range includes Parque Nacional Natural Sierra Nevada de Santa Marta. Surveys to
determine the current population status of this species are urgently required. A captive-breeding programme might
need to be established.
Notes on taxonomy: We follow Coloma (2002) in considering Atelopus leoperezii as a synonym of this species.
Bibliography: Acosta-Galvis, A.R. (2000), Ardila-Robayo, M.C. and Acosta-Galvis, A. (2000a), Coloma, L.A. (2002), La Marca, E. et
al. (2005), Lötters, S. (1996), Pounds, J.A. et al. (2006), Rueda-Almonacid, J.V. (1994b), Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and
Hernández-Camacho, J.A. (1994), Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and Lynch, J.D. (1996)
Data Providers: Andrés Acosta-Galvis, Martha Patricia Ramírez Pinilla, Mariela Osorno-Muñoz, Jose Vicente Rueda, Adolfo Amézquita,
María Cristina Ardila-Robayo
 Globally Threatened Amphibian Species 163

EN Atelopus certus Barbour, 1923

Endangered A3ce Geographic Range This species is currently known only from the Cerro Sapo in south-western Darién Province,
Order, Family: Anura, Bufonidae Panama. It is suspected that the species is more widely distributed, and may range through much of Parque Nacional
Country Distribution: Panama Darién. It has been found at altitudes of 50-1,150m asl.
Current Population Trend: Decreasing Population This species is locally common within its small range, and has been recorded as recently as 2003.
Habitat and Ecology This is a terrestrial species of tropical montane and submontane forest. Breeding and larval
development takes place in forest streams.
Major Threats The main threats to the species are deforestation of habitat for agricultural use and general water
pollution. Congeners are known to have virtually disappeared throughout much of Mesoamerica, probably due to
chytridiomycosis, and this species might also be at risk from the disease, although it occurs at lower altitudes (the
disease normally occurs at higher elevations in the tropics).
Conservation Measures It has been recorded from Parque Nacional Darién. Further survey work is needed to
© Andrew Crawford

establish the limits of the species’ range, and additional research is needed to establish whether chytridiomycosis
might be a threat.
Bibliography: Ibáñez, R. et al. (2000), La Marca, E. et al. (2005), Lötters, S. (1996), Pounds, J.A. et al. (2006), Savage, J.M. (1972a),
Young, B. et al. (1999)
Data Providers: Frank Solís, Roberto Ibáñez, César Jaramillo, Querube Fuenmayor

CR Atelopus chiriquiensis Shreve, 1936

Critically Endangered A2ace Rica since 1996 and the species is now considered to be extinct in that country; it might also have disappeared in
Order, Family: Anura, Bufonidae Panama.
Country Distribution: Panama, Costa Rica Population This species was once considered locally abundant along streams. In Costa Rica this species has disap-
(Extinct) peared from its entire range, and there have been no sightings since 1996 despite many searches in appropriate mon-
Current Population Trend: Decreasing tane habitats. The population status in Panama shows evidence of a decline, with no records since the late 1990s.
Habitat and Ecology It is a diurnal, terrestrial species of stream margins in lower montane wet forest and rainforest.
Males are territorial and use an advertisement call to maintain a breeding site. Breeding and larval development for
this species takes place in forest streams (Savage 2002).
Major Threats Marked declines have been noticed in its extent of occurrence. In Costa Rica the decline and probable
extinction of this species has been linked to chytridiomycosis (Lips 1998), which was confirmed in this species in 1993
and 1994. Introduction of predatory trout, and general habitat loss both outside, and within protected areas, are also
threats to remaining populations. Climate change is considered to be a possible threat.
© David M. Dennis

Geographic Range This species is found in the lower montane zone of the Cordillera de Talamanca-Chiriqui axis of
Costa Rica (1,800-2,500m asl) and western Panama (1,400-2,100m asl)(Savage 2002). It has not been seen in Costa
Conservation Measures The range of the species is within the protected areas of Parque Nacional Chirripó and
Parque Internacional La Amistad. The threat of chytridiomycosis means that successful conservation measures will
probably need to include the maintenance of any surviving individuals in captivity.
Bibliography: Ibáñez, R. et al. (2000), La Marca, E. et al. (2005), Lindquist, E.D. and Swihart, D.W. (1997), Lips, K.R. (1998), Lips, K.R.
(1999), Lips, K.R., Green, D.E. and Papendick, R. (2003), Lips, K.R., Reeve, J.D. and Witters, L.R. (2003), Lötters, S. (1996), Pounds, J.A.
et al. (2006), Savage, J.M. (1972a), Savage, J.M. (2002), Young, B. et al. (1999)
Data Providers: Karen Lips, Roberto Ibáñez, Federico Bolaños, Gerardo Chaves, Frank Solís, Jay Savage, César Jaramillo, Querube
Fuenmayor, A. Castillo

CR Atelopus chocoensis Lötters, 1992

Critically Endangered A3ce Geographic Range This species is known from the type locality: Cerro del Ingles near San Jose del Palmor, close
Order, Family: Anura, Bufonidae to Serrania de los Paraguas, in Choco department, between 1,900 and 2,200m asl. It is also known from Boqueron in
Country Distribution: Colombia the Municipality of El Cairo, Valle del Cauca department, in Colombia. It might occur a little more widely.
Current Population Trend: Decreasing Population The current population status is unknown, and it is known from less than 10 specimens. The species
was last collected in 1998, and the area has most likely not been surveyed since then.
Habitat and Ecology A terrestrial species, it occurs in montane forest on the forest floor and near the forest edge.
It has not been recorded from anthropogenically disturbed habitats. Its breeding habits are not known, although
breeding is likely to take place in streams.
Major Threats Like other Atelopus species, it is presumably at serious risk from chytridiomycosis. Habitat loss due
to subsistence wood collection and clear-cutting, and forest clearance for the planting of illegal crops and cattle
ranching, is also a major threat.
Conservation Measures The range of the species includes the Parque Nacional Natural Tatamá. More research
© John D. Lynch

into the species’ range, ecological requirements, and population status is needed. The threat of chytridiomycosis
means that successful conservation measures will probably need to include the maintenance of any surviving
individuals in captivity.
Bibliography: Acosta-Galvis, A.R. (2000), La Marca, E. et al. (2005), Lötters, S. (1992a), Lötters, S. (1996), Pounds, J.A. et al. (2006)
Data Providers: Fernando Castro, Stefan Lötters, Jose Vicente Rueda

CR Atelopus chrysocorallus La Marca, 1994

Critically Endangered A2ace; Geographic Range This species is restricted to the type locality, near the village of Tostós in Trujillo State, in the
B1ab(iii,v)+2ab(iii,v) Venezuelan Andes. Its known range is less than 100km². It has been recorded from 2,000-2,700m asl.
Order, Family: Anura, Bufonidae Population It is a rare species, and there have been no records since it was discovered in 1988. A subsequent
Country Distribution: Venezuela survey failed to record this species again, suggesting a possible decline, as is the case with many other Venezuelan
Current Population Trend: Decreasing Atelopus species.
Habitat and Ecology It is a diurnal, photophilic species found on the floor of cloud forest. It lays egg chains in
streams, where the tadpoles also develop.
Major Threats The most critical threat to this species is probably chytridiomycosis. However, the type locality lies
within an unprotected forested area that has been altered at a fast rate, mainly due to subsistence wood collection,
and conversion to crop lands and pasture lands. With higher insolation as a result of climate change, this species
could be at increased risk due to its habit of basking in the sun. La Marca and Lötters (1997) indicated that severe
floods might have affected populations of this species.
Conservation Measures Its range does not include any protected areas, and surveys are urgently needed to confirm
the continued existence of this species. Given the threat of chytridiomycosis, successful conservation measures will
© Ted Kahn

probably need to include the maintenance of any surviving individuals in captivity.

Bibliography: Barrio Amorós, C.L. (2004), La Marca, E. (1994b), La Marca, E. et al. (2005), La Marca, E. and Lötters, S. (1997), Lötters,
S. (1996), Pounds, J.A. et al. (2006), Rodríguez, J.P. and Rojas-Suárez, F. (1995)
Data Providers: Enrique La Marca, Juan Elías García-Pérez
164 Threatened Amphibians of the World

CR Atelopus coynei Miyata, 1980

Critically Endangered A2ace Geographic Range This species is known from the provinces of Pichincha, Imbabura, and Carchi on the Pacific
Order, Family: Anura, Bufonidae versant of the Andes in north-western Ecuador. It has been recorded from 600-1,380m asl.
Country Distribution: Ecuador

© Ted Kahn, based on a photograph by Ken Miyati

Population It is a rare species and has not been recorded since September 1984. It has probably undergone a
Current Population Trend: Decreasing serious population decrease.
Habitat and Ecology This species is an inhabitant of humid north-western Andean montane forest. It appears
to be able to adapt to secondary forest. It lays its eggs in swift-flowing streams and rivers. It has typical Atelopus
tadpoles, attached to rocks.
Major Threats The most critical threat to this species is probably chytridiomycosis. Agriculture, both crops and
livestock, as well as logging and infrastructure development for human settlement, are major threats to the species’
habitat.
Conservation Measures The range of this species includes the Reserva Ecológica Cotacachi-Cayapas. Surveys are
needed to establish whether or not this species still survives in the wild. Given the threat of chytridiomycosis, successful
conservation measures will probably need to include the maintenance of any surviving individuals in captivity.
Bibliography: La Marca, E. et al. (2005), Lötters, S. (1996), Miyata, K. (1980), Pounds, J.A. et al. (2006)
Data Providers: Santiago Ron, Luis A. Coloma, Martín R. Bustamante, Diego Cisneros-Heredia, Ana Almandáriz, Mario Yánez-Muñoz

CR Atelopus cruciger (Lichtenstein and Martens, 1856) RANCHO GRANDE HARLEQUIN FROG
Critically Endangered A2ace 1995; La Marca and Lötters 1997; Manzanilla and La Marca 1999; Lötters, La Marca and Vences 2004). Recently
Order, Family: Anura, Bufonidae (2004), a single small population of A. cruciger has been found just south of the town of Cata within the limits of the
Country Distribution: Venezuela 107,000-ha Parque Nacional Henri Pittier, in cloud forest at 600m asl.
Current Population Trend: Decreasing Habitat and Ecology It is a diurnal species usually found on rocks of rivulets or the surroundings, where they can
climb on to plants up to 1.5m above ground. The general habitat is humid forest in montane and lowland environ-
ments. It breeds along swift-flowing streams. The recently rediscovered population was found by a cascading
mountain stream in cloud forest.
Major Threats The major cause of the observed population decline of this species is chytridiomycosis, which was
© Francisco José López-López

Geographic Range This species is restricted to several localities in the northern and southern versants of the
Cordillera de la Costa of Venezuela (Estadoes Aragua, Carabobo, Miranda, Vargas, Yaracuy and the Distrito Federal)
and recently from the Cerro Azul (Estado Cojedes) (Rivas Fuenmayor 1998) which suggests that the species might be
present throughout the entire mountainous area of the central coastal range (Lötters, La Marca and Vences 2004).
It has been recorded from 30-2,200m asl.
Population Although this species was once abundant, it has undergone an extreme decline, to the point that despite
extensive surveys no specimens have been seen since 1986 and there are no museum records after 1988 (La Marca
confirmed in 1986 (Bonaccoroso et al. 2003). Pollution by acid rain could be another possible threat, given the vicinity
of the species to the large concentration of industries generating polluting gases in the nearby area of Valencia-
Maracay. Droughts and flash floods could be a further potential threat, as well as over collecting for scientific or
pet trade purposes.
Conservation Measures Many of the known localities are within the Parque Nacional Henri Pittier, Parque Nacional
Rancho Grande, and Parque Nacional San Esteban. Monitoring of the populations, establishment of a captive-breeding
population, and disease management are all urgently required.
Notes on taxonomy: This species was recently redescribed by Lötters, La Marca and Vences (2004) following the discovery that the
original type material represents Atelopus varius of Central America. The former junior synonym A. vogli is considered to be a distinct
species following Lötters, La Marca and Vences (2004).
Bibliography: Barrio Amorós, C.L. (2004), Bonaccorso, E. et al. (2003), Guayasamin, J.M. et al. (2002), La Marca, E. (1995b), La Marca,
E. et al. (2005), La Marca, E. and Lötters, S. (1997), Lichtenstein, M.H.C. and Von Martens, E.C. (1856), Lötters, S. (1996), Lötters, S., La
Marca, E. and Vences, M. (2004), Manzanilla, J. (2001), Manzanilla, J. et al. (1995), Manzanilla, J. and La Marca, E. (1999), Manzanilla,
J. and La Marca, E. (2004), Pounds, J.A. et al. (2006), Rivas Fuenmayor, G. (1998), Rodríguez, J.P. and Rojas-Suárez, F. (1995), Solano
de Chacín, H. (1968)
Data Providers: Jesús Manzanilla, Enrique La Marca, Ronald Heyer, Ernesto Fernández-Badillo

EN Atelopus dimorphus Lötters, 2003

Endangered A3e Geographic Range This species is known only from the western versant of the Cordillera Azul, Huánuco Department,
Order, Family: Anura, Bufonidae Peru, where it has been observed between 1,650 and 1,800m asl.
Country Distribution: Peru Population It is currently known only from museum specimens, and was last recorded in 1980.
Current Population Trend: Decreasing Habitat and Ecology The general habitat in the area of collection is humid mountain forest. It is presumed to be
a stream-breeding species.
Major Threats The threats to the species within its range on the Cordillera Azul have not been recorded. However,
like other members of its genus, it is assumed to be at severe risk from chytridiomycosis.
Conservation Measures It is not known to occur in any protected areas. In view of the likely threat of chytridiomy-
cosis, it is a very high priority to conduct surveys to locate this species and determine its current population status;
a captive-breeding programme may also need to be established.
Bibliography: La Marca, E. et al. (2005), Lötters, S. (2003), Lötters, S. et al. (2005), Pounds, J.A. et al. (2006)
© Rainer Schulte

Data Providers: Stefan Lötters

CR Atelopus ebenoides Rivero, 1963

Critically Endangered A2ace+3ce Population The northern population had not been recorded since 1995, until its remarkable rediscovery in early
Order, Family: Anura, Bufonidae May of 2006 in the highlands of Boyacá . The southern populations were last recorded in 1992, though there have
Country Distribution: Colombia not been any recent surveys.
Current Population Trend: Decreasing Habitat and Ecology It occurs on vegetation, mosses and in leaf-litter along streams in Andean forests and páramos.
It has not been recorded from disturbed habitat. The tadpoles develop in streams.
Major Threats The major threat is likely to be chytridiomycosis, leading to a catastrophic population decline, as
has occurred in many other montane species of Atelopus. Habitat loss caused by agricultural expansion (cattle
ranching and the planting of illegal crops), as well as pollution from the fumigation of crops, and water source loss
are all threats.
© Rocha, C. GESA - UPTC

Geographic Range This species occurs in the southern part of the Colombian Andes in Cauca and Huila Departments,
with a separate northern population (Atelopus ebenoides marinkellei) at the Páramo de las Papas, in the Department
of Boyacá, in the Cordillera Oriental of Colombia. Its altitudinal range is 2,500-4,700m asl.
Conservation Measures It occurs in the Parque Nacional Natural Puracé. Survey work has recently confirmed the
persistence of the northern population, but additional surveys are needed to ascertain the population status of the
southern population. Given the threat of chytridiomycosis, successful conservation measures will probably need to
include the maintenance of any surviving individuals in captivity. Research is also needed to confirm the taxonomic
status of the northern populations of the species.
Notes on taxonomy: The northernmost population of this species is sometimes regarded as a separate species (Atelopus
marinkellei).
Bibliography: Ardila-Robayo, M.C. and Acosta-Galvis, A. (2000b), Cochran, D.M. and Goin, C.J. (1970), La Marca, E. et al. (2005), Lötters,
S. (1996), Pounds, J.A. et al. (2006), Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and Lynch, J.D. (1996)
Data Providers: Alberto Cadena, Andrés Acosta-Galvis, Wilmar Bolívar, John Lynch
 Globally Threatened Amphibian Species 165

CR Atelopus elegans (Boulenger, 1882)

Critically Endangered A2ace Population In Ecuador, it has declined very seriously and has disappeared in the last ten years, with no records since
Order, Family: Anura, Bufonidae November 1994, apart from one record in November 2002 from Bogotá River in Esmeralda Province. For example, in
Country Distribution: Colombia, Ecuador a recent survey, it was not found 5km north-west of Alluriquín (in Pichincha Province), where it used to be abundant
Current Population Trend: Decreasing (L. Coloma pers. comm.). On Gorgona Island, the species is still very common.
Habitat and Ecology It lives in lowland and submontane humid rainforest, and has not been found in degraded
habitats. It breeds in streams.
Major Threats The decline in Ecuador is unexplained, and is possibly due to chytridiomycosis, although its elevational
range is rather low for this disease. It is also impacted by habitat destruction and degradation, due to agriculture,
logging, and human settlement.
Conservation Measures In Ecuador, its range overlaps with the Reserva Ecológica Cotacachi-Cayapas. In Colombia
it occurs in the Parque Nacional Natural Isla Gorgona. It is an urgent priority to locate the Ecuadorian population and
© Luis A. Coloma

Geographic Range This species occurs in north-western Ecuador in Esmeraldas, Imbabura, and Pichincha Provinces,
at 300-1,140m asl, and on Gorgona Island, a small island 30km off the Colombian coast on the Pacific side, where
it occurs close to sea level.
to implement emergency ex-situ conservation measures. Close monitoring of the population on Gorgona Island is
needed, since, as far as is known, chytrid has not yet been recorded from this population.
Notes on taxonomy: The population on Gorgona Island (i.e., Atelopus gracilis) should be removed from the synonymy of Atelopus
elegans (S. Lötters pers. comm.), but is here provisionally retained in A. elegans pending revision.
Bibliography: Ardila-Robayo, M.C. and Acosta-Galvis, A. (2000b), Boulenger, G.A. (1882b), Cochran, D.M. and Goin, C.J. (1970), Coloma,
L.A. and Ron, S.R. (2001), La Marca, E. et al. (2005), Lötters, S. (1996), Peters, J.A. (1973), Pounds, J.A. et al. (2006), Rivero, J.A. (1963b),
Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and Lynch, J.D. (1996), Urbina, J.N. and Londono, M.C. (2003)
Data Providers: Luis A. Coloma, Santiago Ron, Wilmar Bolívar, Diego Cisneros-Heredia, Stefan Lötters

CR Atelopus erythropus Boulenger, 1903

Critically Endangered A3ce Geographic Range This species is known only from the vicinity of the type locality of Cordillera Carabaya on the
Order, Family: Anura, Bufonidae Amazonian versant of Departamento Puno, Peru. Records of this species from Departamento Huanuco and Depar-
Country Distribution: Peru tamento Ucayali (Rodriguez, Cordova and Icochea 1993) require further investigation. It has an altitudinal range of
Current Population Trend: Decreasing 1,800-2,500m asl.
Population Its population is unknown, but it might be declining. It has been recorded as recently as 2003.
Habitat and Ecology This species’ habitat is cloud forest on the Amazonian versant of the Peruvian Andes. The
species is not expected to be tolerant of habitat degradation. Breeding is presumed to take place in streams.
Major Threats The major threat is likely to be chytridiomycosis, leading to a catastrophic population decline, as
has occurred in many other montane species of Atelopus. This disease has not yet been recorded in this species, but
incidence of the disease is known to be spreading in northern Peru.
© Alessandro Catenazzi

Conservation Measures This species is not present in any protected areas. If chytridiomycosis is indeed shown to be
a threat to this species, then the maintenance of populations in captivity will be a needed conservation measure.
Bibliography: De la Riva, I. et al. (2000), La Marca, E. et al. (2005), Lötters, S. (1996), Lötters, S. (2003), Lötters, S. and de la Riva, I.
(1998), Pounds, J.A. et al. (2006), Rodríguez, L.O., Cordova, J.H. and Icochea, J. (1993)
Data Providers: Ariadne Angulo, Karl-Heinz Jungfer, Robert Reynolds, Javier Icochea, Stefan Lötters, Roberto Ibáñez, Juan Carlos
Chaparro-Auza, César Aguilar Puntriano

CR Atelopus eusebianus Rivero and Granados, 1993

Critically Endangered A3ce Geographic Range This species is known from a few locations all within close proximity in the Cauca Department,
Order, Family: Anura, Bufonidae on the western slope of the central Andes, Totoro-Malvasa, in Colombia, between 2,820 and 3,250m asl.
Country Distribution: Colombia Population One recent survey found no individuals, although another survey in 2004 did record several individuals.
Current Population Trend: Decreasing It is not known to what extent the species is in overall decline.
Habitat and Ecology It occurs along streams on vegetation in páramo, and has not been recorded from disturbed
habitat. Breeding and larval development takes place in streams.
Major Threats The most serious risk to this species is chytridiomycosis, which has had a devastating impact on
other high-altitude species of Atelopus. Other, localized threats include habitat loss caused by agricultural expansion
© Francisco José López-López

(cattle ranching and the planting of illegal crops), water extraction, and fumigation of illegal crops. Climate change
might also be impacting the species.
Conservation Measures It occurs in the Parque Nacional Natural Puracé. More research into the species’ range,
ecological requirements, and population status is needed, and in particular surveys are needed to determine whether
or not the species is currently in decline as has been evidenced in other high-elevation stream-dwelling Atelopus
species in the region. A captive-breeding programme might need to be established for this species.
Bibliography: Acosta-Galvis, A.R. (2000), Ardila-Robayo, M.C. and Acosta-Galvis, A. (2000b), La Marca, E. et al. (2005), Lötters, S. (1996),
Pounds, J.A. et al. (2006), Rivero, J.A. and Granados-Diaz, H.G. (1993), Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and Lynch, J.D. (1996)
Data Providers: Jose Vicente Rueda, Maria Isabel Herrera, Fernando Castro, Andrés Acosta-Galvis, Enrique La Marca, Wilmar Bolívar,
John Lynch

CR Atelopus exiguus Boettger, 1892

Critically Endangered A3ce Geographic Range This species is known from the type locality, Laguna Llaviuco, in Azuay province, and from
Order, Family: Anura, Bufonidae the nearby vicinity in the Cordillera Occidental of southern Ecuador. It is known from eight localities, and has been
Country Distribution: Ecuador recorded from 3,150-3,850m asl.
Current Population Trend: Decreasing Population This historically abundant species was recorded in May 1995, at a time when no decline was recorded
(Coloma, Lötters and Salas 2000). Since then, there has been little survey work, but it is expected that the species
has undergone a population decline across its range, although it is still known to survive in the wild (La Marca et
al. 2005).
Habitat and Ecology It is an inhabitant of sub-páramo and páramo, but is also known to occur in pastureland around
the Laguna Llaviuco. Breeding and larval development take place in streams.
Major Threats The most serious risk to this species is chytridiomycosis, which has had a devastating impact on other
high-altitude species of Atelopus. Agriculture, both crops and livestock, as well as climate change and the construction
of dams are major threats to the species’ habitat. Invasive trout species might prey on tadpoles of this species.
© Eduardo Toral

Conservation Measures The range of this species includes Parque Nacional Cajas and Parque Nacional Mazan.
The population status of this species urgently needs to be assessed and, given the threat of chytridiomycosis, the
maintenance of individuals in captivity is a recommended conservation measure. It is also necessary to control trout
populations within the protected areas.
Notes on taxonomy: This species was removed from the synonymy of Atelopus ignescens by McDiarmid (1971).
Bibliography: Boettger, O. (1892), Coloma, L.A., Lötters, S. and Salas, A.W. (2000), La Marca, E. et al. (2005), Lötters, S. (1996), McDiarmid,
R. (1971), Merino-Viteri, A. (2001), Pounds, J.A. et al. (2006), Ron, S.R. and Merino, A. (2000)
Data Providers: Eduardo Toral, Manuel Morales, Diego Cisneros-Heredia, Luis A. Coloma, Santiago Ron, Martín R. Bustamante
166 Threatened Amphibians of the World

CR Atelopus famelicus Rivero and Morales, 1995 “1992”

Critically Endangered A2ace Geographic Range This species is known from two localities: the Parque Nacional Natural Munchique, where it has been recorded from the private protected area of the El Tambito
Order, Family: Anura, Bufonidae type locality (La Costa), and El Tambito, both localities on the western Proselva Foundation. Further survey work is required to determine the current population status of this species in the
Country Distribution: Colombia slope of the Cordillera Occidental of the Andes, in Cauca Department, wild. Given the threat of chytridiomycosis, any surviving individuals might need to be maintained in captivity.
Current Population Trend: Decreasing Colombia, between 1,300 and 1,580m asl. Notes on taxonomy: This species was previously confused with Atelopus longirostris and Atelopus lynchi, according to Rivero and
Population It is known from only a few specimens, and, despite Morales (1995).
survey work, has not been recorded since 1993. Bibliography: Acosta-Galvis, A.R. (2000), La Marca, E. et al. (2005), Lötters, S. (1996), Pounds, J.A. et al. (2006), Rivero, J.A. and Morales,
Habitat and Ecology It is a terrestrial species found in montane V.R. (1995), Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and Lynch, J.D. (1996), Ruiz-Carranza, P.M., Vélez and Ardila-Robayo, M.C. (1995)
forest, and which has not been recorded outside forest. Its breeding Data Providers: Wilmar Bolívar, Andrés Acosta-Galvis, John Lynch, Stefan Lötters
habits are not known, but presumably it breeds in streams like other
species of the genus.
Major Threats The most serious risk to this species is chytridio-
mycosis, which has had a devastating impact on other high-altitude
species of Atelopus. There is also a threat of pollution resulting
from fumigation of the illegal crops grown in Parque Nacional
Natural Munchique.
Conservation Measures The two localities are on the border of

CR Atelopus farci Lynch, 1993

Critically Endangered A2ace; B2ab(iii,v) Geographic Range This species is known from only one site: Granjas del Padre Luna, Municipio de Alban, western
Order, Family: Anura, Bufonidae slope of the Cordillera Oriental, in Cundinamarca Department, Colombia, at 2,090m asl.
Country Distribution: Colombia Population This species was known to be abundant up until around 1995. Six subsequent visits and some 80 hours
Current Population Trend: Decreasing of searching during the course of 2002-2003 turned up only one tadpole, perhaps indicative of a serious decline.
Habitat and Ecology It occurs in streams in cloud forest. It also reproduces in rapid streams.
Major Threats The most serious risk to this species is chytridiomycosis, which has had a devastating impact on other
high-altitude species of Atelopus. Habitat loss is also a major threat to this species, mainly caused by agricultural
expansion and human settlement.
Conservation Measures This species is not known to occur in any protected area, and there is an urgent need
for improved habitat protection at its only known locality. Further survey work is required to determine the current
© Juan Manuel Renjifo

population status of this species. Given the probable threat of chytridiomycosis, disease management and captive-
breeding programmes might be required.
Bibliography: La Marca, E. et al. (2005), Lötters, S. (1996), Lynch, J.D. (1993b), Pounds, J.A. et al. (2006), Ruiz-Carranza, P.M., Ardila-
Robayo, M.C. and Lynch, J.D. (1996)
Data Providers: Jose Vicente Rueda, Wilmar Bolívar, Andrés Acosta-Galvis

VU Atelopus flavescens Duméril and Bibron, 1841

Vulnerable A3ce Geographic Range This species is known from north-eastern coastal and central French Guiana. It occurs from
Order, Family: Anura, Bufonidae sea level up to 300m asl.
Country Distribution: French Guiana Population This is a common species, although it is not common in Mont Grand Matoury and Matoury region. It
Current Population Trend: Stable has been recorded as recently as 2000.
Habitat and Ecology This species is known from the proximity of fast-flowing small streams in lowland primary
forest. It has not been recorded outside primary forest. Embryonic and larval development occurs in water.
Major Threats There are no present threats to this species, but future threats will most likely include shifting
agriculture and clear-cutting of the forests. Although illegal, there is still some collection for the pet trade, but at
present this is not a major threat to the species. This species probably occurs at elevations below the altitude at
which chytridiomycosis seems prevalent, though this nonetheless remains a potential future threat.
Conservation Measures The range of this species includes Matoury Nature Reserve and Kaw Nature Reserve.
© Renaud Boistel

Continued population monitoring is required, especially in light of the potential threat of chytridiomycosis.
Notes on taxonomy: Atelopus spumarius barbotini has been suggested to represent a junior synonym of this species (Kok 2000).
Bibliography: Boistel, R., Grosjean. S. and Lötters, S. (2005), Duméril, A.M.C. and Bibron, G. (1841), Kok, P.J.R. (2000), La Marca, E. et
al. (2005), Lescure, J. (1974), Lescure, J. (1981a), Lescure, J. and Marty, C. (2000), Lötters, S. (1996), Lynch, J.D. (1993b), McDiarmid, R.W.
(1973), Mudde, P. (1996), Pounds, J.A. et al. (2006), Schmidt, M. (1999), van den Nieuwenhuizen, A. (2003a), van den Nieuwenhuizen,
A. (2003b)
Data Providers: Jean Lescure, Christian Marty, Renaud Boistel, Stefan Lötters, Enrique La Marca, Robert Reynolds, Marinus Hoogmoed,
Ross MacCulloch, Philippe Gaucher, Stefan Lötters

VU Atelopus franciscus Lescure, 1974

Vulnerable A3ce Geographic Range This species is known from central coastal French Guiana. It has been recorded from 5-200m
Order, Family: Anura, Bufonidae asl.
Country Distribution: French Guiana Population It is a locally common species (Lescure and Marty 2001), and was recorded as recently as 2000.
Current Population Trend: Stable Habitat and Ecology This species is found near fast-flowing small streams and creeks in lowland rainforest; it is
not known from any disturbed habitats. Eggs are laid in the water and the tadpoles adhere to rocks.
Major Threats There are no current major threats. However, it is potentially at risk from chytridiomycosis, although
it occurs at low altitudes (which might afford it a degree of protection from this disease). An illegal international pet
trade exists for this species, but it is not a threat to the species as a whole.
Conservation Measures The range of the species includes several nature reserves. Continued population monitoring
is required, especially in light of the potential threat of chytridiomycosis.
Bibliography: Boistel, R., Grosjean. S. and Lötters, S. (2005), La Marca, E. et al. (2005), Lescure, J. (1974), Lescure, J. and Marty, C.
© Christian Marty

(2000), Lötters, S. (1996), Pounds, J.A. et al. (2006)

Data Providers: Jean Lescure, Christian Marty, Marga Born, Renaud Boistel, Robert Reynolds, Marinus Hoogmoed, Ross MacCulloch,
Philippe Gaucher, Stefan Lötters
 Globally Threatened Amphibian Species 167

CR Atelopus galactogaster Rivero and Serna, 1993 “1991”

Critically Endangered A3ce Geographic Range This species is known from two localities within Parque Nacional Natural Paramillo, in Antioquia
Order, Family: Anura, Bufonidae Department, on the northern slope of the western Andes, in Colombia, at 1,500m asl.
Country Distribution: Colombia Population This species is known only from ten specimens; however, there has been no recent survey work within
Current Population Trend: Unknown its range.
Habitat and Ecology Both of the known localities are within primary forest, but no other information is available
on the species’ habitat requirements or breeding habits.
Major Threats The most serious risk to this species is chytridiomycosis, which has had a devastating impact on
other high-altitude species of Atelopus.
Conservation Measures Both known localities of the species are within Parque Nacional Natural Paramillo. Further
research into the species’ range, ecological requirements, and population status is needed, in particular to determine
if it occurs outside its current known range. Given the likely threat of chytridiomycosis, disease management and
© César Landazábal

captive-breeding programmes might be required.

Bibliography: Acosta-Galvis, A.R. (2000), La Marca, E. et al. (2005), Lötters, S. (1996), Pounds, J.A. et al. (2006), Rivero, J.A. and Serna,
M.A. (1993), Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and Lynch, J.D. (1996)
Data Providers: Jose Vicente Rueda, Stefan Lötters, John Lynch

CR Atelopus glyphus Dunn, 1931

Critically Endangered A3ce Geographic Range This species occurs in eastern Panama, in the Serranía de Pirre, and also the Chocó of Colombia,
Order, Family: Anura, Bufonidae at 884-1,500m asl.
Country Distribution: Colombia, Panama Population It is considered to be generally common within its known range. It was still common in September 2002
Current Population Trend: Decreasing in the Serranía de Pirre, above Cana, in eastern Panama (R. Ibáñez pers. obs.).
Habitat and Ecology It is a terrestrial species of tropical montane forest, with breeding and larval development taking
place in forest streams. There is no information on whether or not this species can survive in degraded habitats.
Major Threats The major threat is likely to be a future catastrophic decline, due to chytridiomycosis, as has occurred
in many other species of Atelopus. Additional threats include habitat loss due to agricultural development (including the
planting of illegal crops), logging, and human settlement, and pollution resulting from the spraying of illegal crops.
Conservation Measures The species has been recorded from two protected areas: Parque Nacional Darién (a World
Heritage Site) in Panama and Parque Nacional Natural los Katíos in Colombia. In view of the severe risk of chytridio-
© Andrew Crawford

mycosis, the status of this species should be closely monitored, and ex-situ populations should be established.
Bibliography: Ibáñez, R. et al. (2000), La Marca, E. et al. (2005), Lötters, S. (1996), Pounds, J.A. et al. (2006), Ruiz-Carranza, P.M.,
Ardila-Robayo, M.C. and Lynch, J.D. (1996), Savage, J.M. (1972a), Young, B. et al. (1999)
Data Providers: Roberto Ibáñez, Frank Solís, César Jaramillo, Querube Fuenmayor, Stefan Lötters, Jose Vicente Rueda, Andrés
Acosta-Galvis

CR Atelopus guanujo Coloma, 2002

Critically Endangered A2ace Geographic Range This species is known only from the type locality and its immediate vicinity (Guaranda, Gallo
Order, Family: Anura, Bufonidae Rumi) in the Chimbo Basin of the Cordillera Occidental of Ecuador, in the province of Bolívar. These localities are
Country Distribution: Ecuador between 2,600 and 2,923m asl in the upper Río Chimbo Valley (Coloma 2002).
Current Population Trend: Decreasing Population It is a rare species, and the most recent record dates from April 1988 (Coloma 2002). Since then, survey
efforts have been unsuccessful in finding any individuals, suggesting a serious population decline and possible
extinction.
Habitat and Ecology This species is an inhabitant of humid cloud forest. Frogs have also been collected in disturbed
montane cloud forest areas (Coloma 2002). There is no specific information on breeding habits, but it is likely to be
similar to other Atelopus species, with breeding and larval development taking place in streams.
Major Threats The most likely cause of the species’ population decline is chytridiomycosis. Habitat loss is also a
major threat, due to agriculture (crops, livestock, and wood plantations), logging, and infrastructure development for
© Luis A. Coloma

human settlement. Invasive species such as dogs, cats, and chickens also prey on this species.
Conservation Measures The known range of the species does not include any protected areas. Surveys are needed
to establish whether or not this species still persists in the wild. In view of the severe threat of chytridiomycosis, any
surviving populations should be maintained in captivity.
Bibliography: Coloma, L.A. (2002), La Marca, E. et al. (2005), Merino-Viteri, A. (2001), Pounds, J.A. et al. (2006), Ron, S.R. and Merino,
A. (2000)
Data Providers: Luis A. Coloma, Santiago Ron, Martín R. Bustamante, Diego Cisneros-Heredia, Ana Almandáriz

CR Atelopus guitarraensis Osorno Muñoz, Ardila-Robayo and Ruíz-Carranza, 2001

Critically Endangered A3ce Geographic Range This species is only known to occur in the Bibliography: La Marca, E. et al. (2005), Osorno Muñoz, M., Ardila-Robayo, M.C. and Ruiz-Carranza, P.M. (2001a), Osorno Muñoz, M.,
Order, Family: Anura, Bufonidae Laguna de Guitara, Parque Nacional Natural Sumapaz, Meta Depart- Ardila-Robayo, M.C. and Ruiz-Carranza, P.M. (2001b), Pounds, J.A. et al. (2006)
Country Distribution: Colombia ment, Colombia, at 3,400m asl. Data Providers: Mariela Osorno-Muñoz, María Cristina Ardila-Robayo, Andrés Acosta-Galvis
Current Population Trend: Decreasing Population It is rare, being recorded only from two expeditions
conducted in 1975 and 1990. There has been no subsequent survey
work for this species.
Habitat and Ecology It lives and reproduces in streams in páramo
habitats.
Major Threats The most serious risk to this species is chytridio-
mycosis, which has had a devastating impact on other high-altitude
species of Atelopus. Its habitat is threatened by fires caused by
cattle ranchers.
Conservation Measures This species occurs in the Parque Nacional
Natural Sumapaz. Further work is required to determine the popula-
tion status of this species; given the threat of chytridiomycosis,
a captive-breeding programme might need to be established for
this species.
168 Threatened Amphibians of the World

CR Atelopus halihelos Peters, 1973

Critically Endangered A2ace; B1ab(iii)+2ab(iii) Geographic Range This species is known only from the Cordillera Cutucú, in Morona-Santiago Province, in southern
Order, Family: Anura, Bufonidae Ecuador, around 1,975m asl.
Country Distribution: Ecuador Population Although sampling effort through the species’ range is insufficient, it is likely that this species has
Current Population Trend: Decreasing undergone a population decline (as seen in other montane species of Atelopus). It was last recorded in 1984, and is
perhaps no longer present at the type locality.
Habitat and Ecology This species is an inhabitant of humid montane forest. There is no specific information on their
breeding habits, but these are likely to be similar to other Atelopus species, with breeding and larval development
taking place in streams.
Major Threats The most serious risk to this species is chytridiomycosis, which has had a devastating impact on
other high-altitude Atelopus species. Agriculture, both crops and livestock, as well as logging and infrastructure
development for human settlement are major threats to the species’ habitat.
Conservation Measures It is not known to occur in any protected areas. Surveys to confirm the continued exis-
tence of this species are needed; given the threat of chytridiomycosis, surviving individuals should be maintained
© Ted Kahn

in captivity.
Bibliography: Duellman, W.E. and Lynch, J.D. (1988), La Marca, E. et al. (2005), Lötters, S. (1996), Peters, J.A. (1973), Pounds, J.A.
et al. (2006)
Data Providers: Santiago Ron, Luis A. Coloma, Martín R. Bustamante

CR Atelopus laetissimus Ruíz-Carranza, Ardila-Robayo and Hernández-Camacho, 1994

Critically Endangered A3ce Geographic Range This species is known from the slopes of south-east Cuchilla San Lorenzo, in the north-west
Order, Family: Anura, Bufonidae sector of Parque Nacional Natural Sierra Nevada de Santa Marta (in the department of Magdalena), and the adjacent
Country Distribution: Colombia El Dorado Nature Reserve, in Colombia. It has been recorded from 1,900-2,880m asl.
Current Population Trend: Decreasing Population This was a common species when last recorded in 1992. A population was rediscovered on 22 May 2006
when staff of the El Dorado Nature Reserve located two individuals in a small forested stream.
© Fundación ProAves / www.proaves.org

Habitat and Ecology This species is an inhabitant of sub-Andean forests. It can also adapt to some modification
of its habitat. It lays egg chains in streams, where the tadpoles also develop.
Major Threats The most serious risk to this species is chytridiomycosis, which has had a devastating impact on other
high-altitude Atelopus species. Climate change, habitat loss due to agriculture, and pollution from the fumigation
of illegal crops are also major threats.
Conservation Measures Its range includes Parque Nacional Natural Sierra Nevada de Santa Marta, and the
adjacent El Dorado Nature Reserve established in March 2006 to secure one of the last forested valleys for this and
other threatened amphibian and bird species. Further surveys are needed to determine the current population status
of this species. Disease management and captive-breeding programmes might be required, particularly if it is found
that the species has suffered extensive declines due to chytridiomycosis.
Bibliography: Acosta-Galvis, A.R. (2000), La Marca, E. et al. (2005), Lötters, S. (1996), Pounds, J.A. et al. (2006), Ruiz-Carranza, P.M.
and Ardila-Robayo, M.C. (1991), Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and Hernández-Camacho, J.A. (1994), Ruiz-Carranza, P.M.,
Ardila-Robayo, M.C. and Lynch, J.D. (1996)
Data Providers: Martha Patricia Ramírez Pinilla, Mariela Osorno-Muñoz, Jose Vicente Rueda, Adolfo Amézquita, María Cristina
Ardila-Robayo

EN Atelopus limosus Ibáñez, Jaramillo and Solís, 1995

Endangered B1ab(iii) Geographic Range This species is endemic to the eastern Atlantic versant of central Panama, although it is believed
Order, Family: Anura, Bufonidae to occur much more widely. It is a low-altitude species, occurring at 10-730m asl.
Country Distribution: Panama Population This species is reasonably common at a number of localities, and there is little evidence of a decline.
Current Population Trend: Decreasing However, at Filo de Santa Rita, Provincia de Colón, while relatively abundant in October 2000, only one individual
was seen in December 2002 (R. Ibáñez pers. obs.).
Habitat and Ecology This is a terrestrial species of tropical lowland forest. Breeding and larval development takes
place in forest streams.
Major Threats The deforestation of habitat for agricultural use and general infrastructure development, as well as
water pollution and stream sedimentation, are the main threats to this species. A number of congeners have recently
disappeared over much of their range due to the effects of chytrid, although this species probably occurs below the
altitude at which chytridiomycosis is prevalent.
Conservation Measures The species has been recorded from Parque Nacional Chagres, but expanded habitat
© Brian Kubicki

protection is recommended. Further survey work is needed to establish the limits of the species’ range, and close
monitoring of existing populations is necessary.
Bibliography: Ibáñez, R. et al. (2000), Ibanez-D.R., Jaramillo, C.A. and Solis, F.A. (1995), La Marca, E. et al. (2005), Lötters, S. (1996),
Pounds, J.A. et al. (2006), Savage, J.M. (1972a), Young, B. et al. (1999)
Data Providers: Frank Solís, Roberto Ibáñez, César Jaramillo, Querube Fuenmayor

EN Atelopus longibrachius Rivero, 1963

Endangered A3ce; B1ab(iii) Geographic Range This species is known only from the region Conservation Measures It occurs in Parque Nacional Natural Munchique. In view of the severe risk of chytridio-
Order, Family: Anura, Bufonidae of the type locality: El Tambo, near Guisito, at 800m asl, Cauca mycosis, it is a high priority to conduct surveys to determine the current population status of this species, as well as
Country Distribution: Colombia Department, Colombia. Lötters (1996) suggested that the elevation to initiate a captive-breeding programme.
Current Population Trend: Decreasing given for the type locality (300m asl) was a typographic error in the Bibliography: Acosta-Galvis, A.R. (2000), Cannatella, D.C. (1981), La Marca, E. et al. (2005), Lötters, S. (1996), Pounds, J.A. et al. (2006),
original publication. Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and Lynch, J.D. (1996)
Population There is no recent information on the population status Data Providers: Andrés Acosta-Galvis, Wilmar Bolívar, Stefan Lötters
of this species.
Habitat and Ecology It lives on the ground in tropical humid forests,
and presumably breeds in streams.
Major Threats The major threat is likely to be chytridiomycosis, as
has occurred in many other species of Atelopus (although its occur-
rence at lower altitudes might afford it some level of protection from
the disease). Additional threats include deforestation for agricultural
development, illegal crops, logging and human settlement, and pol-
lution resulting from the spraying of illegal crops.
 Globally Threatened Amphibian Species 169

CR Atelopus lozanoi Osorno-Muñoz, Ardila-Robayo and Ruíz-Carranza, 2001

Critically Endangered A2ace; B2ab(v) Geographic Range This species is known only from the type locality in the Páramo de Palacio, Cundinamarca,
Order, Family: Anura, Bufonidae Colombia, at an altitude of 3,000-3,300m asl.
Country Distribution: Colombia Population This species was once abundant, but it has not been seen since 1993, despite repeated visits to the
Current Population Trend: Decreasing site.
Habitat and Ecology It lives in páramo and does not tolerate habitat destruction. Breeding and larval development
take place in fast-flowing streams.
Major Threats Although not proven, the most likely cause of the decline of this species is chytridiomycosis, which
has had a devastating impact on other high-altitude species of Atelopus. It is also likely to be threatened by fires set
by cattle ranchers. In addition, introduced trout prey upon tadpoles in streams.
© Maria Cristina Ardila Conservation Measures Occurs in Parque Nacional Natural Chingaza. Survey work is needed to determine the
population status of this species; given the threat of chytridiomycosis, successful conservation measures will probably
need to include the maintenance of any surviving individuals in captivity. There is also a need to control populations
of trout within the National Park.
Bibliography: La Marca, E. et al. (2005), Osorno Muñoz, M., Ardila-Robayo, M.C. and Ruiz-Carranza, P.M. (2001b), Pounds, J.A. et
al. (2006)
Data Providers: Mariela Osorno-Muñoz, Adolfo Amézquita, Andrés Acosta-Galvis

CR Atelopus lynchi Cannatella, 1981

Critically Endangered A3ce; B1ab(iii,iv,v) Geographic Range This species is endemic to the Pacific slopes of northern Ecuador (Maldonado in the Province
Order, Family: Anura, Bufonidae of Carchi), between 800 and 1,410m asl. Records of this species from the western slope of the Cordillera Occidental,
Country Distribution: Ecuador Valle del Cauca Department, in Colombia refer to an unidentified, probably un-named species.
Current Population Trend: Decreasing Population This species has disappeared from Ecuador, and there have been no records from this country since
1984.
Habitat and Ecology It lives on the border between lowland tropical rainforests and montane forests. It is terrestrial,
presumably breeding in streams.
Major Threats The major threat is likely to be chytridiomycosis, which has caused major declines in many other
species of Atelopus. Additional threats include deforestation for agricultural development (including the planting of
illegal crops), logging, and human settlement, and pollution resulting from the fumigation of illegal crops.
Conservation Measures This species is not known from any protected areas. Survey work is urgently required to
determine whether or not this species still persists, and, in view of the severe risk of chytridiomycosis, any surviving
© John D. Lynch

populations should immediately form part of an ex-situ management programme.

Notes on taxonomy: Rivero and Serna (1993 “1991”) doubted that specimens from Valle del Cauca, Colombia, were correctly assigned
to this species.
Bibliography: Acosta-Galvis, A.R. (2000), Cannatella, D.C. (1981), La Marca, E. et al. (2005), Lötters, S. (1996), Merino-Viteri, A. (2001),
Pounds, J.A. et al. (2006), Rivero, J.A. and Morales, V.R. (1995), Rivero, J.A. and Serna, M.A. (1993), Ruiz-Carranza, P.M., Ardila-Robayo,
M.C. and Lynch, J.D. (1996)
Data Providers: Santiago Ron, Luis A. Coloma, Martín R. Bustamante, Wilmar Bolívar, Stefan Lötters, Juan Manuel Renjifo, Jose
Vicente Rueda

CR Atelopus mandingues Osorno-Muñoz, Ardila-Robayo and Ruíz-Carranza, 2001

Critically Endangered A3ce Geographic Range This species is known only from the type Bibliography: La Marca, E. et al. (2005), Osorno Muñoz, M., Ardila-Robayo, M.C. and Ruiz-Carranza, P.M. (2001a), Pounds, J.A. et
Order, Family: Anura, Bufonidae locality: Reserva Biológica Carpanta, Cundinamarca, Colombia, at al. (2006)
Country Distribution: Colombia an altitude of 2,900-3,350m asl. Data Providers: Mariela Osorno-Muñoz, Adolfo Amézquita
Current Population Trend: Unknown Population This species was uncommon until 1990. A single pair
was recorded in 1992; however, due to guerrilla activity, no one has
subsequently returned to the original locality to look for it.
Habitat and Ecology It lives in páramo and cloud forest, and does not
tolerate habitat destruction. It reproduces in fast-flowing streams.
Major Threats The most serious risk to this species is chytridio-
mycosis, which has had a devastating impact on other high-altitude
species of Atelopus.
Conservation Measures It occurs in the Reserva Biológica Car-
panta, which borders Parque Nacional Natural Chingaza. Survey work
is urgently needed to determine the population status of this species.
In view of the severe risk of chytridiomycosis, ex-situ populations
might need to be established.

CR Atelopus mindoensis Peters, 1973

Critically Endangered A2e Geographic Range This species is known from Pichinca and Esmeraldas provinces, in the north-western versant of
© Natural History Museum and Biodiversity Research Center,

Order, Family: Anura, Bufonidae the Andes of Ecuador, between 700 and 2,200m asl. It is known from more than ten localities.
Country Distribution: Ecuador Population There is no current information on the population status of this species; it was last recorded in 1989,
Current Population Trend: Decreasing despite survey efforts, suggesting a serious population decrease.
Habitat and Ecology This species is an inhabitant of lowland rainforest and humid montane forest. Eggs are laid
in strings in streams, and tadpoles attach themselves to rocks.
Major Threats The major threat is likely to be a catastrophic decline, due to chytridiomycosis, as has occurred in
many other species of Atelopus. Agriculture, both crops and livestock, as well as logging, are also major threats to
the species’ habitat, and very little suitable habitat remains.
Conservation Measures The range of this species overlaps Reserva Ecológica Cotacachi-Cayapas and just reaches
the northern limit of Reserva Ecológica Los Illinizas. Additional survey work is urgently needed to determine whether
University of Kansas

or not this species still persists. In view of the severe risk of chytridiomycosis, surviving individuals might need to
form the basis for the establishment of an ex-situ population.
Bibliography: La Marca, E. et al. (2005), Lötters, S. (1996), Lötters, S. (2001), Miyata, K. (1980), Peters, J.A. (1973), Pounds, J.A. et
al. (2006)
Data Providers: Luis A. Coloma, Santiago Ron, Martín R. Bustamante, Mario Yánez-Muñoz, Diego Cisneros-Heredia, Ana Almandáriz
170 Threatened Amphibians of the World

CR Atelopus minutulus Ruíz-Carranza, Hernandez-Camacho and Ardila-Robayo, 1988

Critically Endangered A3ce Geographic Range This species lives on the eastern slope of the eastern Andes, Meta Department, Villavicencio,
Order, Family: Anura, Bufonidae Acacias, Vareda Portachuelo, via Manzanares, Colombia, at 1,370-1,560m asl.
Country Distribution: Colombia Population This species was abundant in the early 1980s, but was last seen in 1985 (despite subsequent surveys
Current Population Trend: Decreasing in 1987). No one has returned to look for this species since.
Habitat and Ecology It is an inhabitant of cloud forest, with breeding and larval development taking place in streams.
It tolerates natural disturbances such as landslides, but not significant opening up of its habitat.
Major Threats It occurs in a very restricted area that is severely threatened by habitat destruction, particularly due to
cattle ranching. However, the most serious risk to this species is chytridiomycosis, which has had a devastating impact
on other high-altitude Atelopus species.
© Maria Cristina Ardila Conservation Measures This species is not known to occur in any protected areas. Surveys are urgently needed
to determine the population status of the species. In view of the severe risk of chytridiomycosis, ex-situ populations
might need to be established.
Bibliography: Acosta-Galvis, A.R. (2000), Cocroft, R.B. et al. (1990), La Marca, E. et al. (2005), Lötters, S. (1996), Pounds, J.A. et al.
(2006), Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and Lynch, J.D. (1996), Ruiz-Carranza, P.M., Hernández-Camacho, J.A. and Ardila-
Robayo, M.C. (1988)
Data Providers: Jose Vicente Rueda, María Cristina Ardila-Robayo, Andrés Acosta-Galvis, Mariela Osorno-Muñoz, Ruth Adriana
Maldonado-Silva

CR Atelopus monohernandezi Ardila-Robayo, Osorno-Muñoz and Ruíz-Carranza, 2001

Critically Endangered A2e Geographic Range This species is found in Santuario de Fauna y mine the biology and population status of this species. Given the threat of chytridiomycosis, successful conservation
Order, Family: Anura, Bufonidae Flora Guanentá Alto Río Fonce, Río Cañaverales, Santander Depart- measures will probably need to include the maintenance of any surviving individuals in captivity.
Country Distribution: Colombia ment, Colombia, at 1,700-2,000m asl. Bibliography: Ardila-Robayo, M.C., Osorno-Muñoz, M. and Ruiz-Carranza, P.M. (2002), La Marca, E. et al. (2005), Pounds, J.A. et al.
Current Population Trend: Decreasing Population During the course of multiple visits undertaken in (2006)
1979-1982, this species was found to be common. There have been Data Providers: María Cristina Ardila-Robayo, Mariela Osorno-Muñoz
no subsequent surveys for the species, with the exception of one
survey in 1992 when this species was not recorded, suggesting a
serious decrease.
Habitat and Ecology It is an inhabitant of cloud forest, with
breeding and larval development taking place in streams. It toler-
ates disturbed forest.
Major Threats The most serious risk to this species is chytridio-
mycosis, which has had a devastating impact on other high-altitude
species of Atelopus.
Conservation Measures It occurs in the Santuario de Fauna y Flora
Guanentá Alto Río Fonce. Further survey work is required to deter-

CR Atelopus mucubajiensis Rivero, 1972

Critically Endangered A2ace; B2ab(v) Population It is a very rare species. This is the only Venezuelan Andean amphibian for which an ongoing population
Order, Family: Anura, Bufonidae monitoring programme exists. The population seems to have experienced a drastic decline in the last 15 years or so,
Country Distribution: Venezuela to the point that no individuals of this species were recorded since 1993 (adults) or 1994 (larvae), until recent surveys
Current Population Trend: Decreasing confirmed that a few individuals (23 in total) continue to survive in the wild (Barrio-Amorós 2004).
Habitat and Ecology This species is an inhabitant of páramo and cloud forest, the highest environments in the
Venezuelan Andes, resembling alpine tundra, but with daily temperature extremes. It is usually found within grasses
and frailejones (Espeletia spp.), and along streams. It probably lives in the shrubs of the sub-páramo environment. It
is photophilic and lays egg chains in streams, where the tadpoles also develop.
Major Threats The major threat is likely to be chytridiomycosis, which has resulted in catastrophic declines of
many other species of Atelopus. Chytridiomycosis was confirmed in this species in 1988. Introduced trout, introduced
conifers, fires caused by humans, over collection, and agriculture and infrastructure development for human settle-
© Enrique La Marca

Geographic Range This species has a very restricted distribution of only a few square kilometres at the type
locality, in the Páramo de Mucubají, in the Sierra de Santo Domingo, in the Venezuelan Andes. It has an altitudinal
range of 2,300-3,500m asl.
ment are all major threats. With higher insolation as a result of climate change, this species could be at increased
risk due to its habit of basking in the sun.
Conservation Measures Most of the range of this species is within the Parque Nacional Sierra Nevada. A new
monitoring project for this species has recently begun (www.andigena.org). In view of the threat of chytridiomycosis,
an ex-situ population might need to be established.
Bibliography: Barrio Amorós, C.L. (2004), Barrio-Amorós, C.L. (2004), La Marca, E. (1995b), La Marca, E. et al. (2005), La Marca, E. and
Lötters, S. (1997), La Marca, E. and Reinthaler, H.P. (1991), Lötters, S. (1996), Pounds, J.A. et al. (2006), Rivero, J.A. (1972), Rodríguez,
J.P. and Rojas-Suárez, F. (1995)
Data Providers: Enrique La Marca, Suleima Santiago, Stefan Lötters, Juan Elías García-Pérez, César Luis Barrio Amorós

CR Atelopus muisca Rueda-Almonacid and Hoyos, 1992 “1991”

Critically Endangered A2ace; B2ab(v) Geographic Range This species occurs in Parque Nacional Natural Chingaza, Colombia, between 2,900 and
Order, Family: Anura, Bufonidae 3,350m asl.
Country Distribution: Colombia Population This species was common up until 1996, but four expeditions in 1998 and 2003 failed to turn up a single
Current Population Trend: Decreasing individual at any of the previous known localities.
Habitat and Ecology It occurs in Andean cloud forests and páramos. Breeding and larval development take place
in streams.
Major Threats The major threat is likely to be chytridiomycosis, which has resulted in catastrophic declines of many
other species of Atelopus. Cattle grazing is also a threat to the habitat of this species, and introduced trout have a
© Juan Manuel Renjifo

negative impact on reproductive success.

Conservation Measures This species occurs in the Parque Nacional Natural Chingaza. Further survey work is
required to determine whether or not this species still persists in the wild. In view of the threat of chytridiomycosis,
any surviving individuals might need to form the basis for the establishment of an ex-situ population.
Bibliography: Acosta-Galvis, A.R. (2000), La Marca, E. et al. (2005), Lötters, S. (1996), Pounds, J.A. et al. (2006), Rueda-Almonacid, J.V.
and Hoyos, J.M. (1991), Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and Lynch, J.D. (1996)
Data Providers: Jose Vicente Rueda, Wilmar Bolívar, Adolfo Amézquita, Andrés Acosta-Galvis
 Globally Threatened Amphibian Species 171

CR Atelopus nahumae Ruíz-Carranza, Ardila-Robayo and Hernández-Camacho, 1994

Critically Endangered A3ce Geographic Range This species is known from the slopes of south-east Cuchilla San Lorenzo, in the north-west
Order, Family: Anura, Bufonidae sector of Parque Nacional Natural Sierra Nevada de Santa Marta, and the adjacent El Dorado Nature Reserve, in the
Country Distribution: Colombia department of Magdalena, Colombia. It has been recorded between 1,900 and 2,800m asl.
Current Population Trend: Unknown Population This was a common species when it was last recorded in 1992. It was rediscovered in 2006, following
survey work in the recently established El Dorado Nature Reserve.

© Conservacion Internacional-Colombia
Habitat and Ecology This species is an inhabitant of sub-Andean forests, and can also tolerate some degree of
habitat disturbance. It lays egg chains in streams, where the tadpoles also develop.
Major Threats The most serious risk to this species is chytridiomycosis, which has had a devastating impact on
other high-altitude species of Atelopus. Other major threats include climate change, habitat loss caused by agriculture
(illegal crops) and logging, and pollution caused by the fumigation of illegal crops.
Conservation Measures The range of this species encompasses the Parque Nacional Natural Sierra Nevada de Santa
Marta; the adjacent El Dorado Nature Reserve was established in March 2006 to secure one of the last forested valleys
for this and other threatened amphibian and bird species. Surveys are needed to determine the current population
status of this species and, given the threat of chytridiomycosis, a captive-breeding programme might be required.
Bibliography: Acosta-Galvis, A.R. (2000), La Marca, E. et al. (2005), Lötters, S. (1996), Pounds, J.A. et al. (2006), Ruiz-Carranza, P.M.,
Ardila-Robayo, M.C. and Hernández-Camacho, J.A. (1994), Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and Lynch, J.D. (1996)
Data Providers: Martha Patricia Ramírez Pinilla, Mariela Osorno-Muñoz, Jose Vicente Rueda, Adolfo Amézquita, María Cristina
Ardila-Robayo

CR Atelopus nanay Coloma, 2002

Critically Endangered A2ace; B2ab(v) Geographic Range This species is known only from páramo habitats near Laguna Toreadora, in the Cordillera
Order, Family: Anura, Bufonidae Occidental, in Azuay Province, Ecuador (Coloma 2002) at around 4,000m asl.
Country Distribution: Ecuador Population The current population status of this species is not known. It has not been recorded since July 1989,
Current Population Trend: Decreasing despite surveys within the range, suggesting a serious population decrease.
Habitat and Ecology This species is an inhabitant of herbaceous páramo. Some specimens have been found active
close to streams and springs in an area of many interconnected pools, while others have been found on land under
rocks (Coloma 2002). There is no specific information available on their breeding habits, but they are likely to be
similar to other Atelopus species, with breeding and larval development taking place in streams.
Major Threats The major threat is likely to be chytridiomycosis, leading to a catastrophic population decline as has
occurred in many other montane species of Atelopus. Agriculture (both crops and livestock), as well as logging and
infrastructure development for human settlement, are major threats to the species’ habitat. Invasive alien species
© Luis A. Coloma

are also a problem.

Conservation Measures The type locality of the species is within Parque Nacional Cajas. Further survey work
is required to determine whether or not this species survives in the wild. Given the threat of chytridiomycosis, any
surviving individuals might need to form the basis for the establishment of an ex-situ population.
Bibliography: Coloma, L.A. (2002), La Marca, E. et al. (2005), Merino-Viteri, A. (2001), Pounds, J.A. et al. (2006), Ron, S.R. et al.
(2003)
Data Providers: Luis A. Coloma, Santiago Ron, Martín R. Bustamante, Stefan Lötters

CR Atelopus nepiozomus Peters, 1973

Critically Endangered A3ce Geographic Range This species is known from six localities, on the eastern flanks of the Andes in Morona-Santiago
Order, Family: Anura, Bufonidae Province, Ecuador, at 2,000-3,450m asl.
Country Distribution: Ecuador Population The population status of this species is not known, but sampling effort through its range is insufficient.
Current Population Trend: Unknown It was last recorded in September 1985.
Habitat and Ecology This species is an inhabitant of sub-páramo and páramo. There is no specific information
available on breeding, but it is likely to be similar to other Atelopus species, with breeding and larval development
taking place in streams.
Major Threats The most serious risk to this species is chytridiomycosis, which has had a devastating impact on other
high-altitude Atelopus species. Habitat loss and degradation as a result of mining and logging is a major threat.
Conservation Measures The range of this species slightly overlaps with Parque Nacional Podocarpus, although it
has, to date, not been recorded from inside the park. Further survey work is required to determine the abundance of
this species. In view of the severe risk of chytridiomycosis, ex-situ populations might need to be established.
Bibliography: La Marca, E. et al. (2005), Lötters, S. (1996), Merino-Viteri, A. (2001), Peters, J.A. (1973), Pounds, J.A. et al. (2006)
© Ted Kahn

Data Providers: Luis A. Coloma, Santiago Ron, Martín R. Bustamante, Stefan Lötters, Diego Cisneros-Heredia

CR Atelopus nicefori Rivero, 1963

Critically Endangered A3ce Geographic Range This species is known only from the type local- to determine the population status of this species, and to establish whether or not it occurs outside the vicinity of the
Order, Family: Anura, Bufonidae ity: Alto de Caicedo, in Antioquia Department, Colombia, between type locality. In view of the threat of chytridiomycosis, ex-situ populations might need to be established.
Country Distribution: Colombia 1,800 and 2,670m asl. Notes on taxonomy: Specimens discussed by Cochran and Goin (1970) were subsequently named as Atelopus echeverrii.
Current Population Trend: Unknown Population The current population status of this species is unknown, Bibliography: Acosta-Galvis, A.R. (2000), Cochran, D.M. and Goin, C.J. (1970), Cocroft, R.B. et al. (1990), La Marca, E. et al. (2005),
but it was not uncommon when last collected. There has not been Lötters, S. (1996), Pounds, J.A. et al. (2006), Rivero, J.A. (1963b), Rivero, J.A. and Serna, M.A. (1985), Ruiz-Carranza, P.M., Ardila-Robayo,
any recent survey work for this species. M.C. and Lynch, J.D. (1996)
Habitat and Ecology It occurs along forested streams in sub-An- Data Providers: Stefan Lötters, Jose Vicente Rueda, John Lynch
dean and Andean forests. It is restricted to forest habitats requiring
canopy cover over the stream to maintain suitable conditions. Breed-
ing and larval development take place in streams.
Major Threats The most serious risk to this species is chytridio-
mycosis, which has had a devastating impact on other high-altitude
species of Atelopus. Forests within the range of the species are being
cleared for the planting of fruit trees, as well as illegal crops.
Conservation Measures The known range of the species is close
to Parque Nacional Natural Orquideas. Further survey work is needed
172 Threatened Amphibians of the World

CR Atelopus oxyrhynchus Boulenger, 1903

Critically Endangered A2ace Geographic Range This species has a very restricted distribution, in Conservation Measures Some subpopulations occur within the Parque Nacional Sierra Nevada and Parque Nacional
Order, Family: Anura, Bufonidae cloud forests in the vicinities of the city of Mérida, in Mérida State, Sierra de la Culata. Surveys are needed to establish whether or not this species still persists in its natural range.
Country Distribution: Venezuela in the Cordillera de Mérida in the Venezuelan Andes. Its elevation In view of the threat of chytridiomycosis, surviving individuals might need to form the basis for the establishment
Current Population Trend: Decreasing ranges from 2,100-3,500m asl. of an ex-situ population.
Population It is now an extremely rare species, and although found Bibliography: Barrio Amorós, C.L. (2004), Boulenger, G.A. (1903), La Marca, E. (1992), La Marca, E. (1995b), La Marca, E. et al. (2005),
in good numbers in 1978 and 1985, it was last recorded in 1994. La Marca, E. and Lötters, S. (1997), La Marca, E. and Reinthaler, H.P. (1991), Lötters, S. (1996), Pounds, J.A. et al. (2006), Rodríguez, J.P.
Subsequent survey attempts to locate this species have failed to find and Rojas-Suárez, F. (1995)
any individuals, suggesting a serious population decrease. Data Providers: Enrique La Marca, Irwin García, Rubén Albornoz, Juan Elías García-Pérez
Habitat and Ecology This species is an inhabitant of montane cloud
forests. It is photophillic and lays egg chains in streams, where the
tadpoles also develop.
Major Threats The major threat is likely to be chytridiomycosis,
leading to a catastrophic population decline, as has occurred in many
other montane species of Atelopus. Habitat loss and degradation is
also a major threat, due to agriculture (crops, livestock, and planta-
tions), logging (in the past), and mining.

CR Atelopus pachydermus (Schmidt, 1857)

Critically Endangered A2ace Geographic Range This species occurs on the eastern versant of the Andes of Ecuador in Napo Province. It was
Order, Family: Anura, Bufonidae reported from humid montane forests in the Departamentos of Amazonas, Cajamarca and La Libertad, Peru, by
Country Distribution: Ecuador Rodríguez, Cordova and Icochea (1993), but these specimens refer to an unnamed species (S. Lötters pers. comm.).
Current Population Trend: Decreasing It might be present in Colombia, because the type specimens were shipped from Buenaventura, Colombia. It occurs
at around 2,755-3,300m asl.
© Natural History Museum and Biodiversity

Population This formerly abundant species has completely disappeared from its known range, and the last record
is from 1996. Surveys at Papallacta undertaken since 1993 have failed to record this species.
Research Center, University of Kansas

Habitat and Ecology It occurs near streams in páramo and sub-páramo, and breeding takes place in streams.
Major Threats The major threat is likely to be chytridiomycosis, leading to a catastrophic population decline, as has
occurred in many other montane species of Atelopus.
Conservation Measures The geographic range of this species in Ecuador overlaps Reserva Ecológica Cayambe-Coca,
Reserva Ecológica Antisana, Parque Nacional Llanganates, and Parque Nacional Sangay. Survey work is needed to
determine whether or not this species still persists. In view of the threat of chytridiomycosis, surviving individuals
might need to form the basis for the establishment of an ex-situ population.
Notes on taxonomy: The species treated here (and by previous authors) as A. pachydermus is in fact unnamed. True Atelopus pachydermus
are from an unknown locality, probably in the Peruvian Andes (S. Lötters pers. comm.).
Bibliography: Acosta-Galvis, A.R. (2000), La Marca, E. et al. (2005), Lötters, S. (1996), Merino-Viteri, A. (2001), Peters, J.A. (1973), Pounds,
J.A. et al. (2006), Rodríguez, L.O., Cordova, J.H. and Icochea, J. (1993), Ron, S.R. and Merino, A. (2000), Schmidt, O. (1857)
Data Providers: Wilmar Bolívar, Luis A. Coloma, Santiago Ron, Stefan Lötters, Enrique La Marca

CR Atelopus pedimarmoratus Rivero, 1963

Critically Endangered A3ce Geographic Range This species is known only from the type locality: San Isidro, Municipio de Ubala, Cundinamarca
Order, Family: Anura, Bufonidae Department, Colombia, at 2,600-3,100m asl.
Country Distribution: Colombia Population There have been no reports of this species since its original description in 1963, but there appear to
Current Population Trend: Unknown have been no attempts to relocate it.
Habitat and Ecology It is an inhabitant of cloud forest, with breeding and larval development taking place in
streams.
Major Threats The major threat is likely to be chytridiomycosis, leading to a catastrophic population decline, as
has occurred in many other montane species of Atelopus. Habitat destruction due to potato farming is also a threat
to this species.
Conservation Measures The species has not been recorded from any protected area, although it might occur in the
Reserva Forestal Protectora El Predio Río Sucio, in the Municipio de Gachalá. Survey work is necessary to determine
© César Landazábal

the current population status of the species. In view of the threat of chytridiomycosis, surviving individuals might
need to be taken into captivity.
Notes on taxonomy: Lötters (1996), suggested that the Cauca localities (by Cochran and Goin 1970) were doubtful on geographic
grounds.
Bibliography: Acosta-Galvis, A.R. (2000), Cochran, D.M. and Goin, C.J. (1970), La Marca, E. et al. (2005), Lötters, S. (1996), Pounds,
J.A. et al. (2006), Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and Lynch, J.D. (1996)
Data Providers: Jose Vicente Rueda, María Cristina Ardila-Robayo, Mariela Osorno-Muñoz, Ruth Adriana Maldonado-Silva, Enrique
La Marca

CR Atelopus peruensis Gray and Cannatella, 1985

Critically Endangered A2ace was) present at Charco (3,700m asl), Huari, Departamento Ancash, and also in the Departamento Piura. Its altitudinal
Order, Family: Anura, Bufonidae range is from 2,800-4,200m asl.
Country Distribution: Peru Population This species was very abundant until 1992, but there appears to have been a decline since then, and several
Current Population Trend: Unknown populations, one of them at Cajamarca-Celendín, are believed to have disappeared (R. Schulte pers. comm.).
Habitat and Ecology It inhabits puna and “sub-puna” (high altitude) habitats, with scattered tussock grass and
Baccharis sp. They are sensitive to habitat degradation. Breeding takes place in streams, with larvae adhering to the
undersides of large (30-50cm) rocks in swiftly flowing streams at a depth of approximately 30cm.
Major Threats The major threat is likely to be chytridiomycosis, leading to a catastrophic population decline, as
has occurred in many other montane species of Atelopus. The chytrid pathogen has yet to be found in this species,
but the disease is known to be spreading in northern Peru, and the declines already reported are likely to be related
to this disease. It has previously been reported in the pet trade, although this appears to have stopped. A population
in Cajamarca was lost due to water contamination from activities at a nearby gold mine.
© Jörn Köhler

Geographic Range This species is known only from the northern Peruvian Andes near the surroundings of Celendin,
Abra Comulica, San Miguel de Pallaques and Province of Hualgayoc, all in the Departamento Cajamarca; it is (or
Conservation Measures This species occurs in Parque Nacional Huascarán, and might be present in Reserva
Nacional Callipuy, and Santuario Nacional Callipuy. Disease management and captive-breeding programmes appear
necessary given the threat of chytridiomycosis.
Bibliography: Gray, P. and Cannatella, D.C. (1985), INRENA (Instituto Nacional de Recursos Naturales, Peru) (2001), Instituto Nacional de
Recursos Naturales (INRENA) (2000), La Marca, E. et al. (2005), Lötters, S. (1996), Lötters, S. et al. (1990), Lötters, S. et al. (2005), Lötters,
S. and Köhler, J. (1998), Mebs, D. et al. (1995), Pounds, J.A. et al. (2006), Rodríguez, L.O., Cordova, J.H. and Icochea, J. (1993)
Data Providers: Rainer Schulte, Antonio Salas, Ariadne Angulo, Stefan Lötters, Ulrich Sinsch, Alfonso Miranda Leiva

CR Atelopus petriruizi Ardila-Robayo, 1999
Critically Endangered A3ce
Order, Family: Anura, Bufonidae
Country Distribution: Colombia
Current Population Trend: Unknown
CR Atelopus pictiventris Kattan, 1986
Critically Endangered A3ce
Order, Family: Anura, Bufonidae
Country Distribution: Colombia
Current Population Trend: Unknown
© Gustavo Kattan
CR Atelopus pinangoi Rivero “1980” (1984)
Critically Endangered A2ac; B1ab(iii,v)
Order, Family: Anura, Bufonidae
Country Distribution: Venezuela
Current Population Trend: Decreasing
CR Atelopus planispina Jiménez de la Espada, 1875
Critically Endangered A2ace
Order, Family: Anura, Bufonidae
Country Distribution: Ecuador
Current Population Trend: Decreasing
© Luis A. Coloma
Geographic Range This species is known from five localities in the eastern slopes of the Ecuadorian Andes, from
Volcán Reventador to the south, to Cordillera de Cutucú. It has been recorded from 1,000-2,000m asl.
Globally Threatened Amphibian Species
Geographic Range This species occurs in Parque Nacional Natural
Los Picachos, Varedas la Esperanza y Andalucia, Caquetá Depart-
ment, Colombia, between 1,750 and 2,500m asl. It might occur a
little more widely.
Population It is only known from four specimens collected between
1994 and 1998; there has been no subsequent survey work for the
species.
Habitat and Ecology It occurs and breeds in streams in premontane
moist forest.
Major Threats The most serious risk to this species is chytridio-
mycosis, which has had a devastating impact on other high-altitude
species of Atelopus. Habitat loss due to the planting of illegal crops
is a threat to this species.
Geographic Range This species is only known from the type
locality, “Piñango”, at 2,920m asl, and from a second population
a few kilometres away, in cloud forest near the village of Pinango,
Mérida State, in the Andes of Venezuela. It has been recorded from
2,300-2,920m asl.
Population This is a rare species. The last recorded observation
was in 1997, despite repeated searches, thus suggesting a serious
population decrease.
Habitat and Ecology This species is an inhabitant of montane
cloud forest. It is photophilic and lays egg chains in streams, where
the tadpoles also develop.
Major Threats The major threat is likely to be chytridiomycosis,
which has also caused major declines in many other montane spe-
cies of Atelopus. Habitat loss due to logging and agriculture (cattle
ranching, crops) -- the type locality was completely destroyed by
the mid-1980s -- and introduced trout, are also major threats to the
173
Conservation Measures It occurs in Parque Nacional Natural Los Picachos. Survey work is required to determine
the current population status of the species. Given the threat of chytridiomycosis, surviving individuals might need
to form the basis for an ex-situ population.
Bibliography: Acosta-Galvis, A.R. (2000), Ardila-Robayo, M.C. (1999), La Marca, E. et al. (2005), Pounds, J.A. et al. (2006)
Data Providers: María Cristina Ardila-Robayo, John Lynch
Geographic Range This species is known from Alto Pance (Corea), Municipality of Cali, in the Valle del Cauca
Department on the eastern slope of the Cordillera Occidental, in Colombia, at 2,600m asl.
Population This species is known from only a few specimens, and was last recorded in 1996.
Habitat and Ecology It is a terrestrial and diurnal species, with all specimens having been found in artificially cleared
forest. Its breeding habits are not known, but it presumably breeds in streams like other species in the genus.
Major Threats The most serious threat to this species is chytridiomycosis, which has had a devastating impact on
other high-altitude species of Atelopus.
Conservation Measures The type locality is within the Parque Nacional Natural Farallones de Cali. Further survey
work is required to determine the biology and current population status of this species. Considering that chytridio-
mycosis is the likely overwhelming threat, surviving individuals might need to form the basis for the establishment
of an ex-situ population.
Bibliography: Acosta-Galvis, A.R. (2000), Kattan, G. (1986), La Marca, E. et al. (2005), Lötters, S. (1996), Pounds, J.A. et al. (2006),
Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and Lynch, J.D. (1996)
Data Providers: Stefan Lötters, John Lynch
GREEN AND RED VENTER HARLEQUIN TOAD
species. With increased insolation as a result of climate change this species could be at enhanced risk due to its
sub-basking habits.
Conservation Measures The range of the species does not include any protected areas. Surveys are required to
establish whether or not this species still survives. In view of the threat of chytridiomycosis, surviving individuals
might need to form the basis for the establishment of an ex-situ population.
Bibliography: Barrio Amorós, C.L. (2004), La Marca, E. (1994d), La Marca, E. (1995b), La Marca, E. et al. (2005), La Marca, E. and Lötters,
S. (1997), La Marca, E. and Reinthaler, H.P. (1991), Lötters, S. (1996), Péfaur, J.E. and Rivero, J.A. (2000), Pounds, J.A. et al. (2006), Rivero,
J.A. (1984), Rodríguez, J.P. and Rojas-Suárez, F. (1995), Vial, J.L. and Saylor, L. (1993)
Data Providers: Enrique La Marca, Juan Elías García-Pérez
Population The population of this species has declined dramatically. The last record was in July 1985 (an amplec-
tant pair) despite repeated visits to known (El Reventador) or inferred localities (within its extent of occurrence)
(Bustamante 2002).
Habitat and Ecology This species is an inhabitant of humid montane forest. There is no specific information known
about breeding, but it is likely to be similar to other Atelopus species, with breeding and larval development taking
place in streams.
Major Threats The major threat is likely to be chytridiomycosis, and which has also led to catastrophic population
declines in many other montane species of Atelopus. Habitat loss due to agriculture (both crops and livestock),
logging, mining, and infrastructure development for human settlement, as well as agricultural pollution, are also
major threats.
Conservation Measures Its distribution range overlaps with Parque Nacional Sumaco Napo-Galeras, Parque Nacional
Sangay, Reserva Ecológica Antisana, Parque Nacional Llanganates, and possibly Reserva Ecológica Cayambe-Coca.
Surveys are needed to establish whether or not this species still survives. In view of the threat of chytridiomycosis,
surviving individuals might need to form the basis for the establishment of an ex-situ population.
Notes on taxonomy: The taxonomic status of A. planispina and A. palmatus was discussed by Coloma (1997). Recent unpublished
infromation suggests that A. palmatus might be a valid species known from the type locality (S. Lötters pers. comm.).
Bibliography: Bustamante, M. R. (2002), Coloma, L.A. (1997), Jiménez de la Espada, M. (1875), La Marca, E. et al. (2005), Lötters, S.
(1996), Peters, J.A. (1973), Pounds, J.A. et al. (2006)
Data Providers: Santiago Ron, Luis A. Coloma, Martín R. Bustamante, Diego Cisneros-Heredia, Stefan Lötters, Enrique La Marca
174 Threatened Amphibians of the World

CR Atelopus pulcher (Boulenger, 1882)

Critically Endangered A2ace Geographic Range This species is distributed along the lower Andean versant in the upper Río Huallaga drainage,
Order, Family: Anura, Bufonidae Departamentos San Martín and Loreto, Peru. It is present at approximately 600-900m asl. There are populations
Country Distribution: Peru resembling this species from the north-eastern versant of Peru and adjacent Ecuador, but these are not included in
Current Population Trend: Decreasing this account until their taxonomic status can be more conclusively verified.
Population The current population status of this species is not known, although it was recorded as recently as July
2004. It is now considered to be relatively rare, and has disappeared from many known localities.
Habitat and Ecology This is a diurnal and terrestrial species of lowland and premontane tropical forest. Breeding
takes place in streams. It is not known how adaptable the species is to habitat modification.
Major Threats The threats to this species include the potentially devastating impacts of chytridiomycosis (the disease
is already present in northern Peru) which was confirmed in this species in 2003. Other threats include localized habitat
loss through conversion to agricultural land.
© Ariadne Angulo

Conservation Measures The species occurs in Parque Nacional Cordillera Azul. A captive-breeding program for
this species has been established, and there is ongoing research investigating the biology of a population of this
species that apparently has chytridiomycosis.
Notes on taxonomy: This species was removed from the synonymy of A. spumarius by Lötters et al. (2002b).
Bibliography: La Marca, E. et al. (2005), Lötters, S. et al. (2002b), Lötters, S. et al. (2005), Pounds, J.A. et al. (2006)
Data Providers: Stefan Lötters

CR Atelopus quimbaya Ruíz-Carranza and Osorno-Muñoz, 1994

Critically Endangered A3ce Geographic Range This species is known only from the vicinity of the type locality on the border of Risaralda and
Order, Family: Anura, Bufonidae Quindio Departments, Colombia, between 2,200 and 2,900m asl. It might occur more widely than current records
Country Distribution: Colombia suggest.
Current Population Trend: Unknown Population This is a rare species, and was last recorded in 1992.
Habitat and Ecology It is found on vegetation along streams and away from streams in sub-Andean and Andean
forests; it has not been recorded outside forest habitat. Breeding and larval development take place in streams.
Major Threats The most serious risk to this species is chytridiomycosis, which has had a devastating impact on
other high-altitude species of Atelopus.
Conservation Measures It might occur within a protected area (Parque Regional Natural Ucumarí, in Risaralda
Department). Further survey work is required to determine the population status of this species and whether or not
© Pedro Ruiz Carranza

if it occurs outside the vicinity of the type locality. Given the threat of chytridiomycosis, recommended conservation
measures will probably need to include the maintenance of any surviving individuals in captivity.
Bibliography: Acosta-Galvis, A.R. (2000), La Marca, E. et al. (2005), Lötters, S. (1996), Pounds, J.A. et al. (2006), Ruiz-Carranza, P.M.
and Osorno-Muñoz, M. (1994), Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and Lynch, J.D. (1996)
Data Providers: Wilmar Bolívar, Andrés Acosta-Galvis, John Lynch

CR Atelopus reticulatus Lötters, Haas, Schick and Böhme, 2002

Critically Endangered A3ce Geographic Range This species is known only from the type locality on the eastern versant of the Cordillera Azul
Order, Family: Anura, Bufonidae (Departamento Ucayali), ca. 3km by road after Divisoria on the Tingo Mariá-Pucallpa road, Peru. It was collected at
Country Distribution: Peru an elevation of 1,600m asl.
Current Population Trend: Decreasing Population There is no information on the population of this species. It appears that the last record was in 1992,
but the lack of subsequent observations might be due to lack of survey work.
Habitat and Ecology It is recorded from primary montane tropical forest; it is not known whether or not it occurs in
modified habitats. Breeding is believed to take place in streams, like other species in the genus.
Major Threats The major threat is likely to be chytridiomycosis, leading to a catastrophic population decline, as has
occurred in many other montane species of Atelopus. The chytrid pathogen has yet to be found in this species, but
the disease is known to be spreading in northern Peru. Its habitat might also be threatened by small-scale agriculture
(e.g., cocoa and tea), but this requires further confirmation.
© Rainer Schulte

Conservation Measures The presence of this species in the Parque Nacional Cordillera Azul is possible, but is
not confirmed. Further survey work is required to determine the current population status of this species. Given the
threat of chytridiomycosis, recommended conservation measures will probably need to include the establishment
of a captive-breeding programme.
Bibliography: La Marca, E. et al. (2005), Lötters, S. et al. (2002a), Lötters, S. et al. (2005), Pounds, J.A. et al. (2006)
Data Providers: Stefan Lötters, Jorge Luis Martinez, Rainer Schulte

CR Atelopus seminiferus Cope, 1874

Critically Endangered A3ce Geographic Range This species is known only from the type locality and the immediate vicinity, from between
Order, Family: Anura, Bufonidae Balsa Puerto and Moyobamba (San Martin), Peru. It occurs at 1,000-2,000m asl.
Country Distribution: Peru Population There is no information on population status or abundance, although it was recorded as recently as
Current Population Trend: Decreasing 2004.
Habitat and Ecology This is a terrestrial species of montane primary forests on the eastern slopes of the Andes. It
is a stream-breeding species and not believed to be present in modified habitats.
Major Threats The major threat is likely to be chytridiomycosis, leading to a catastrophic population decline, as
has occurred in many other montane species of Atelopus; however, the chytrid pathogen has not yet been positively
confirmed in this species.
Conservation Measures This species is present in the Bosque de Protección Alto Mayo. Given the possible threat
of chytridiomycosis, disease management and captive-breeding programmes might be required, as is further survey
© Rainer Schulte

work to establish the population status of this species.

Bibliography: La Marca, E. et al. (2005), Lötters, S. (1996), Lötters, S. et al. (2005), Pounds, J.A. et al. (2006), Rodríguez, L.O., Cordova,
J.H. and Icochea, J. (1993)
Data Providers: Rainer Schulte, Antonio Salas, Ariadne Angulo, Stefan Lötters
 Globally Threatened Amphibian Species 175

CR Atelopus senex Taylor, 1952

Critically Endangered A2ace Geographic Range This species occurs in humid montane forest in central Costa Rica in the Cordilleras Central and
Order, Family: Anura, Bufonidae Talamanca in Costa Rica from 1,100-2,200m asl.
Country Distribution: Costa Rica Population This species was formerly abundant but has not been seen since 1986 despite repeated searches. It was
Current Population Trend: Decreasing formerly abundant on the slopes of Volcán Barva, but is now believed extinct there (Savage 2002).
Habitat and Ecology It occurs and reproduces in stream margins in premontane rainforest and lower montane rain-
forest. It is a diurnal, stream-breeding species, and used to be found in great concentrations during the reproductive
period from July to August (Savage 2002).
Major Threats The major threat is likely to be chytridiomycosis, leading to a catastrophic population decline, as has
occurred in many other montane species of Atelopus. Other threats to this species might include climate change,
© Wayne Van Devender collecting for the pet trade, and possibly pollution.
Conservation Measures The range of this species is protected by both Parque Nacional Tapantí and Parque Nacional
Braulio Carrillo (although it is now believed extinct in the latter area). Further survey work is required to determine
whether or not this species still persists. Given the threat of chytridiomycosis, surviving individuals might need to
form the basis for the establishment of an ex-situ population.
Bibliography: Barrantes, U. (1986), Cocroft, R.B. et al. (1990), La Marca, E. et al. (2005), Lötters, S. (1996), Pounds, J.A. et al. (2006),
Savage, J.M. (2002)
Data Providers: Federico Bolaños, Gerardo Chaves, Uriel Barrantes

CR Atelopus sernai Ruíz-Carranza and Osorno-Muñoz, 1994

Critically Endangered A2ace Geographic Range This species is known from the type locality, Serrania Las Baldias, and from one other locality,
Order, Family: Anura, Bufonidae Belnira, Paramo Morron, both of which are in Antioquia Department, in Colombia, between 2,800 and 3,100m asl.
© John Jairo Estrada, courtesy of Museo de Herpetología

Country Distribution: Colombia Population It is not a common species, and recent surveys have not found any specimens at the type locality; a decline
Current Population Trend: Decreasing has also been observed at the other known locality. The most recent record of the species was apparently in 2000.
Habitat and Ecology It occurs on vegetation along streams in natural forest edge in the high Andes, and has not
been recorded from disturbed habitats. Breeding and larval development take place in streams.
Major Threats The major threat is likely to be chytridiomycosis, leading to a catastrophic population decline, as has
occurred in many other montane species of Atelopus. Other major threats include habitat loss, due to agricultural
expansion (illegal crops), water pollution from agriculture, and possibly climate change.
Conservation Measures The range of the species is not within any protected areas. Further survey work is required
Universidad de Antioquia

to determine the population status of this species and the limits of its range. In view of the threat of chytridiomycosis,
the status of this species should be closely monitored, and ex-situ populations should be established.
Bibliography: Ardila-Robayo, M.C. and Acosta-Galvis, A. (2000b), La Marca, E. et al. (2005), Pounds, J.A. et al. (2006), Ruiz-Carranza,
P.M. and Osorno-Muñoz, M. (1994), Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and Lynch, J.D. (1996)
Data Providers: Jose Vicente Rueda, Mariela Osorno-Muñoz, María Cristina Ardila-Robayo, Ruth Adriana Maldonado-Silva, Wilmar
Bolívar, Fernando Castro, John Lynch

CR Atelopus simulatus Ruíz-Carranza and Osorno-Muñoz, 1994

Critically Endangered A2ace Geographic Range This species is known from the central Andes of Colombia, from Cauca and Tolima Departments,
Order, Family: Anura, Bufonidae between 2,500 and 3,000m asl.
Country Distribution: Colombia Population Before 1999 this was a common species, but it has only been recorded once since then (2001) at a single
Current Population Trend: Decreasing locality despite numerous searches. A serious decline appears to have taken place.
Habitat and Ecology It occurs on the ground near streams and also in open areas; it has not been recorded from
© Fundación ProAves / www.proaves.org

anthropogenically disturbed habitats. The tadpoles develop in streams.

Major Threats The major threat is likely to be chytridiomycosis, leading to a catastrophic population decline, as
has occurred in many other montane species of Atelopus. Habitat loss due to agricultural expansion (including the
planting of illegal crops), as well as water pollution from agriculture, are major threats to the species. Climate change
might also be a threat, but this needs to be investigated further.
Conservation Measures It is not known whether or not this species occurs in any protected areas. Further survey
work is needed to determine whether or not this species still survives. Given the threat of chytridiomycosis, recom-
mended conservation measures should include the establishment of a captive-breeding programme.
Bibliography: Ardila-Robayo, M.C. and Acosta-Galvis, A. (2000b), La Marca, E. et al. (2005), Pounds, J.A. et al. (2006), Ruiz-Carranza,
P.M. and Osorno-Muñoz, M. (1994), Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and Lynch, J.D. (1996)
Data Providers: Alonso Quevedo Gil, Jose Vicente Rueda, Wilmar Bolívar

CR Atelopus sonsonensis Vélez-Rodríguez and Ruíz-Carranza, 1997

Critically Endangered A3ce Geographic Range This species is known only from the type locality: Mesopotamia, near Páramo de Sonsón, in the
Order, Family: Anura, Bufonidae municipality of Sonsón, Vereda Caunzal, on the eastern side of the Cordillera Central, about 15km to the east of the
Country Distribution: Colombia municipality of Argelia, in Colombia. It was recorded at 1,500m asl, and might occur more widely.
Current Population Trend: Unknown Population The population status is unknown. This is a relatively newly described species, but it apparently has not
been recorded since it was first discovered in 1996.
Habitat and Ecology It occurs on the ground in the under-storey of forest along streams; it has not been recorded
from disturbed habitats. Breeding and larval development take place in streams.
Major Threats The major threat is likely to be chytridiomycosis, leading to a catastrophic population decline, as
has occurred in many other montane species of Atelopus. Habitat loss due to smallholder farming activities and the
planting of illegal crops, and water pollution from agriculture, are major threats to the species.
Conservation Measures This species is not known from any protected areas. More research into the species’ range
and population status is needed, in particular to determine if it occurs outside the vicinity of the type locality. In view
© John D. Lynch

of the threat of chytridiomycosis, ex-situ populations might need to be established.

Bibliography: Acosta-Galvis, A.R. (2000), La Marca, E. et al. (2005), Pounds, J.A. et al. (2006), Vélez-Rodríguez and Ruiz-Carranza,
P.M. (1997)
Data Providers: Wilmar Bolívar, Fernando Castro
176 Threatened Amphibians of the World

CR Atelopus sorianoi La Marca, 1983 SCARLET HARLEQUIN TOAD

Critically Endangered A2ace; B2ab(iii,v) geographic range of any Venezuelan Atelopus species, and lives at an altitude of 2,400-2,718m asl.
Order, Family: Anura, Bufonidae Population At the time of its discovery, this species was particularly abundant at the type locality. However, it is
Country Distribution: Venezuela now extremely rare, or perhaps even extinct; the last record of the species was in 1990.
Current Population Trend: Decreasing Habitat and Ecology It is an inhabitant of montane cloud forests. It is photophilic and lays egg chains in streams,
where the tadpoles also develop.
Major Threats The major threat is likely to be chytridiomycosis (which was confirmed in this species in 1988), and
which has led to a catastrophic population decline, as has been observed in many other montane Atelopus species.
Clear cutting has greatly reduced the amount of available habitat for this species. Climate change might also be a
threat. With higher insolation as a result of climate change, this species could be at increased risk due to its habit
of basking in the sun.
Conservation Measures The small range of this species does not include any protected areas. Surveys are needed
© Pascual Soriani

Geographic Range This species is known from a single stream in an isolated cloud forest, the Paramito de San
Francisco, near the town of Guaraque, in Mérida State, Cordillera de Mérida, in Venezuela. It has the most restricted
to establish whether or not this species still survives, and a project aimed at evaluating their population status and
at assessing environmental and climate conditions of the cloud forests of Mérida State in the Venezuelan Andes
(where populations of this toad might occur in remote ridges and valleys) has been initiated. Ex-situ conservation
action might also be required.
Bibliography: Barrio Amorós, C.L. (2004), La Marca, E. (1983), La Marca, E. (1995b), La Marca, E. et al. (2005), La Marca, E. and Lötters,
S. (1997), La Marca, E. and Reinthaler, H.P. (1991), Pounds, J.A. et al. (2006), Rodríguez, J.P. and Rojas-Suárez, F. (1995), Vial, J.L. and
Saylor, L. (1993)
Data Providers: Enrique La Marca, Argelia Rodríguez, Juan Elías García-Pérez

VU Atelopus spumarius Cope, 1871

Vulnerable A3ce Dios in Peru (A. Salas, in litt. to E. La Marca), but there is no recent information.
Order, Family: Anura, Bufonidae Habitat and Ecology It lives on the floor of terra firme tropical rainforest and in the leaf-litter near running streams.
© Ron Gagliardo, courtesy of Atlanta Botanical Gardenf

Country Distribution: Brazil, Colombia, Ecuador,
French Guiana, Guyana, Peru, Suriname
Current Population Trend: Decreasing
Geographic Range This species occurs in the Amazonian lowlands of Colombia, Ecuador and eastern Peru, to
Amazonas, Para, Amapa (Brazil), and the Guianas (Frost 1985; but see taxonomic notes). It ranges from sea level
to 600m asl.
Population It is locally abundant. It is unlikely to be declining in most of its range, but in Ecuador it is thought to
have declined significantly and there are no records since November 1994. It was formerly abundant at Madre de
It appears to be restricted to undisturbed habitats. Breeding takes place in fast-flowing streams.
Major Threats Threats to this species include forest loss due to agriculture, and logging and clear cutting, especially
in eastern Amazônia. It occurs below the altitude at which chytridiomycosis is normally a problem, but declines have
nevertheless taken place in Ecuador and probably also Peru, and it is possible that animals from Iquitos in Peru suc-
cumbed to this disease (R. Schulte pers. comm. to E. La Marca).
Conservation Measures It occurs in many protected areas across its range. Further research is necessary to determine
the degree of threat posed by chytridiomycosis. A captive-breeding program for this species has been initiated.
Notes on taxonomy: Cocroft et al. (1990) and Lötters et al. (2002b) suggested that Atelopus spumarius might be a complex of more
than one species. Lescure and Marty (2001) recognize two subspecies: A. s. barbotini Lescure 1981; and A. s. hoogmoedi Lescure 1973
without arguments and in contrast to their earlier opinion. Lötters et al. (2002b) suggested that A. spumarius occurs in the upper Amazon
Basin only (Peru, Colombia, Brazil); populations from southern Peru, Ecuador and central Brazil might refer to undescribed taxa within A.
spumarius sensu lato. Populations from the Guianas and eastern Amazonia might be treated as A. hoogmoedi complex (maybe within
A. spumarius sensu lato). Atelopus spumarius hoogmoedi is a synonym of A. spumarius under Lescure and Gasc (1986). This form is
understood by Lescure and Gasc (1986) to be more or less A. spumarius sensu lato; however, A. spumarius hoogmoedi is not conspecific
with A. spumarius sensu stricto (S. Lötters pers. comm.)
Bibliography: Boulenger, G.A. (1882a), Cocroft, R.B. et al. (1990), Coloma, L.A. and Ron, S.R. (2001), Cope, E.D. (1871), Frost, D.R. (1985),
Gascon, C. (1989), La Marca, E. et al. (2005), Lescure, J. (1973), Lescure, J. (1981b), Lescure, J. and Gasc, J.P. (1986), Lescure, J. and Marty,
C. (2000), Lötters, S. (1996), Lötters, S. et al. (2002b), Lötters, S. et al. (2005), Pounds, J.A. et al. (2006), Rivero, J.A. (1968b), Rodríguez,
L.O. and Duellman, W.E. (1994), Ron, S.R. (2001), Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and Lynch, J.D. (1996)
Data Providers: Claudia Azevedo-Ramos, Santiago Ron, Luis A. Coloma, Martín R. Bustamante, Antonio Salas, Rainer Schulte, Stefan
Lötters, Ariadne Angulo, Fernando Castro, Jean Lescure, Christian Marty, Enrique La Marca, Marinus Hoogmoed

VU Atelopus spurrelli Boulenger, 1914

Vulnerable A3ce Geographic Range This species occurs in the Colombian Pacific lowlands, in Valle de Cauca, Risaralda, and Choco
Order, Family: Anura, Bufonidae Departments, between 50 and 500m asl.
Country Distribution: Colombia Population It is a locally abundant species, and has apparently not shown any dramatic population declines. It has
Current Population Trend: Decreasing been recorded as recently as 2001.
© Mauricio Rivera-Correa, courtesy of Museo de

Habitat and Ecology It lives in leaf-litter in humid forests close to water sources, and is presumed to breed in
streams. It has been found in primary and secondary forest, but not in heavily degraded areas.
Major Threats The major threats are deforestation for agricultural development, the planting of illegal crops, logging,
Herpetología Universidad de Antioquia

and human settlement, and pollution resulting from the spraying of illegal crops. Since it occurs at very low altitudes,
it might not be as prone to chytridiomycosis as other members of its genus.
Conservation Measures It occurs in several protected areas. Continued population monitoring is required, especially
in light of the potential threat of chytridiomycosis.
Bibliography: Acosta-Galvis, A.R. (2000), Daly, J.W. et al. (1994), Goldberg, S.R. and Bursey, C.R. (2003), La Marca, E. et al. (2005),
Lötters, S. (1996), Pounds, J.A. et al. (2006), Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and Lynch, J.D. (1996)
Data Providers: Wilmar Bolívar, Stefan Lötters

CR Atelopus subornatus Werner, 1899

Critically Endangered A3ce Geographic Range This species is known from two localities, some 20km apart, in the northern Colombian Andes:
Order, Family: Anura, Bufonidae Alto San Miguel and Aguadita, Municipios de Sibaté, and Fusagasuga, Cundinamarca Department, between 2,000
Country Distribution: Colombia and 2,800m asl.
Current Population Trend: Decreasing Population This species was common until the time of the last record in 1999, although there have been no sub-
sequent surveys for the species.
Habitat and Ecology It occurs in pristine and disturbed cloud forests. Breeding and larval development take place
in streams.
Major Threats The major threat is likely to be chytridiomycosis, leading to a catastrophic population decline, as has
occurred in many other montane species of Atelopus. Habitat loss due to agricultural expansion, and water pollution
caused by run-off from pig farms, are also serious threats faced by this species.
Conservation Measures It is not known to occur in any protected area. Survey work is needed to determine the
population status of this species; in view of the threat of chytridiomycosis, ex-situ populations might need to be
© John D. Lynch

established.
Notes on taxonomy: This species was removed from the synonymy of Atelopus ignescens by Lynch (1986).
Bibliography: Acosta-Galvis, A.R. (2000), Ardila-Robayo, M.C. and Acosta-Galvis, A. (2000a), La Marca, E. et al. (2005), Lötters, S.
(1989), Lötters, S. (1996), Lynch, J.D. (1968), Lynch, J.D. (1986c), Pounds, J.A. et al. (2006), Ruiz-Carranza, P.M., Ardila-Robayo, M.C.
and Lynch, J.D. (1996)
Data Providers: María Cristina Ardila-Robayo, Jose Vicente Rueda, Mariela Osorno-Muñoz, Ruth Adriana Maldonado-Silva
 Globally Threatened Amphibian Species 177

CR Atelopus tamaense La Marca, García-Pèrez and Renjifo, 1989
Critically Endangered A3ce
Order, Family: Anura, Bufonidae
Country Distribution: Colombia, Venezuela
Current Population Trend: Decreasing
TAMÁ HARLEQUIN FROG
Geographic Range This species is known from the Páramo de Tamá, is a very high priority to conduct surveys to relocate this species and determine its current population status, as well
on the Venezuelan-Colombian border, in Apure State, Venezuela. It as to initiate a captive-breeding programme.
has been heard in Colombia from Venezuela, along the stream that Notes on taxonomy: The correct name of this species is Atelopus tamaense (a noun in apposition, rather than an adjective) (E. La
marks the border between the two countries. Its known altitudinal Marca pers. comm.)
range is 2,950-3,200m asl. Bibliography: Barrio Amorós, C.L. (2004), La Marca, E. (1992), La Marca, E. et al. (2005), La Marca, E. and Lötters, S. (1997), La Marca,
Population This species has not been seen since 1987, but there E., Garcia-Perez, J.E. and Renjifo, J.M. (1990), Pounds, J.A. et al. (2006), Rodríguez, J.P. and Rojas-Suárez, F. (1995)
have not been any surveys within its range since this time. Data Providers: Enrique La Marca, Juan Elías García-Pérez
Habitat and Ecology It lives in high-elevation sphagnum bogs in
páramo habitat. The tadpoles have been found in streams.
Major Threats The major threat is likely to be chytridiomycosis,
leading to a catastrophic population decline, as has occurred in
many other montane species of Atelopus. The only likely threat to
its habitat is fire.
Conservation Measures It occurs in the Parque Nacional El Tamá
in Venezuela, and Parque Nacional Natural Tamá in neighbouring
Colombia. In view of the risk of infections with the chytrid fungus, it

VU Atelopus tricolor Boulenger, 1902 THREE-COLOURED HARLEQUIN TOAD

Vulnerable A3ce Peru; and five specimens from 40km north Caranavi, Buena Vista Hills, Nor Yungas Province, La Paz Department,
Order, Family: Anura, Bufonidae Bolivia (Lavilla et al. 1997).
Country Distribution: Bolivia, Peru Population It is still common within its range, and the declines associated with other species of Atelopus have not
Current Population Trend: Decreasing yet been noted in this species. It has been recorded as recently as 2003.
Habitat and Ecology This species inhabits wet primary montane forest, corresponding to the Yungas Forest (De la
Riva et al. 2000, Köhler 2000a). Breeding takes place in streams. Köhler (2000a) observed this species perching, at
night, at 0.3-1.2m elevation, in vegetation in disturbed primary forest, and secondary growth along roadsides. Males
call in small groups of 4-10 individuals.
Major Threats The major threats are habitat loss, due to smallholder farming activities (coffee, coca, chili peppers),
and pollution, as a result of increased stream sedimentation. Chytridiomycosis has yet to be found in this species, but
is a potential major threat in the future. Introduced predatory trout might be a threat in some areas.
Conservation Measures In Bolivia, it is present in the Parque Nacional Carrasco, Parque Nacional Madidi and Pilon
© Jörn Köhler

Geographic Range This species is found in the eastern Andes in Peru and Bolivia, at elevations of 600-2,500m
asl. The type locality is the Marcapata Valley in Peru, in the Amazonian slopes of the eastern Andes (south-eastern
Cuzco Department). Other Peruvian records are from areas close to the Puno Department, at 1,700-2,100m asl
(Duellman 1979, Köhler 2000a). In Bolivia there are records from La Paz Department, in the Yungas region, and in
the south of the Chapare’s region, at Cochabamba Department, at the eastern slope of the Andes (Fugler 1984, De la
Riva 1990, Reynolds and Foster 1992, Köhler 2000a). The larval description of A. tricolor comes from 12 samples of
Pucartambo-Atalaya, 68km (by the road) north-east Union Bridge, at Tachila River at 1,700m asl, in Cuzco Department,
Lajas. In Peru, it is present in Manu Biosphere Reserve. Continued population monitoring is required, especially in
light of the potential threat of chytridiomycosis.
Notes on taxonomy: Reynolds and Foster (1992) indicated that Atelopus rugulosus is a synonym of A. tricolor. Subsequently, Lötters
and De la Riva (1998) confirmed this and pointed out that in evaluating the types of A. rugulosus from south-east Peru, and A. wilimani
from Bolivia, both are synonyms of A. tricolor.
Bibliography: Boulenger, G.A. (1902), de la Riva, I. (1990), De la Riva, I. et al. (2000), Donoso-Barros, R. (1969), Duellman, W.E. (1979),
Fugler, C.M. (1984), Instituto Nacional de Recursos Naturales (INRENA) (2000), Köhler, J. (2000a), La Marca, E. et al. (2005), Lavilla, E.O.
and de Sá, R. (2001), Lavilla, E.O., de Sa, R.O. and de la Riva, I. (1997), Lötters, S. and de la Riva, I. (1998), Noble, G.K. (1921), Pounds,
J.A. et al. (2006), Reynolds, R. and Foster, M. (1992)
Data Providers: Antonio Salas, Roberto Ibáñez, Alessandro Catenazzi, Juan Carlos Chaparro-Auza, Ariadne Angulo, Steffen Reichle,
Jörn Köhler, Ignacio De la Riva, Stefan Lötters, Claudia Cortez, Wilfredo Arizabal

CR Atelopus varius (Lichtenstein and von Martens, 1856) HARLEQUIN FROG

Critically Endangered A2ace individuals were found. In Panama, some populations have declined, but others persist. For example, Lips (1999)
Order, Family: Anura, Bufonidae reported a mass mortality in Fortuna, Provincia de Chiriquí, which affected this species in December 1996-January
Country Distribution: Costa Rica, Panama 1997. This site was visited again in February 1998, but the species was not found (Ibáñez 1999). It has been recorded
Current Population Trend: Decreasing in Panama as recently as November 2002, but it is believed to still be in serious decline.
Habitat and Ecology It is a terrestrial species of humid lowland and montane forest; specimens recorded at lowland
rainforest localities were all found along high-gradient, rocky streams, in hilly areas (Savage 2002). It is associated
with small fast-flowing streams and is often found along the banks and sitting out on rocks in streams; at night
they sleep in crevices or low vegetation. They forrmely occurred in large concentrations during the dry season, from
December to May (Savage 2002). Eggs are laid in water and are probably attached to rocks.
Major Threats The major threat is likely to be chytridiomycosis, which has led to catastrophic population declines in
many other montane species of Atelopus. Museum specimens of this species have been found to have chytrid fungi.
One specimen collected in 2003 from the only known site at which the species survives in Costa Rica tested positive
© Brian Kubicki

Geographic Range This species is known from both Atlantic and Pacific versants of the cordilleras of Costa Rica and
western Panama, up to 2,000m asl. It is also present on outlying ridges and hills down to 16m asl at a few lowland
sites. The species is absent from the lowlands of the Pacific north-west (Savage 2002). Records from Colombia are
in error. Recent declines have reduced the Costa Rican range to only one known locality, Fila Chonta, 10km north-
west of the city of Quepos.
Population Over 100 populations of this species were known from Costa Rica where it was often quite common.
Drastic declines began in Monteverde in 1988 and the species was thought to have been extirpated in Costa Rica by
1996. However, after nearly eight years during which the species was thought to be extinct, a remnant population
was discovered near Quepos, on the Pacific coastal range, in 2003, and was surveyed again in 2005 when more
for chytrid infection, and the disease was also confirmed in individuals in 1986, 1990, 1992 and 1997. Other threats
to the species include habitat loss due to the destruction of natural forests, and predation by introduced trout. The
only known site in Costa Rica is under serious threat of a landslide that could potentially destroy the entire stream
section where they are presently found. It was collected by the thousands in the 1970s and shipped to Germany as
part of the international pet trade.
Conservation Measures The species is present in three protected areas in Panama, and was previously found in a
number of Costa Rican protected areas. Ex-situ conservation measures are now needed to ensure the future survival
of this species, and a captive-breeding program has been initiated.
Bibliography: Crump, M.L. (1986), Crump, M.L. and Pounds, J.A. (1989), Ibáñez, R. (1999), Ibáñez, R. et al. (2000), La Marca, E. et al.
(2005), Lips, K.R. (1999), Lips, K.R., Green, D.E. and Papendick, R. (2003), Lips, K.R., Reeve, J.D. and Witters, L.R. (2003), Lötters, S. (1996),
Pounds, J.A. et al. (2006), Pounds, J.A. and Crump, M.L. (1987), Pounds, J.A. and Crump, M.L. (1994), Pounds, J.A., Fogden, M.P.L. and
Campbell, J.H. (1999), Savage, J.M. (1972a), Savage, J.M. (2002), Young, B. et al. (1999), Zippel, K. (2005)
Data Providers: Alan Pounds, Robert Puschendorf, Federico Bolaños, Gerardo Chaves, Martha Crump, Frank Solís, Roberto Ibáñez, Jay
Savage, César Jaramillo, Querube Fuenmayor, Karen Lips
178 Threatened Amphibians of the World

CR Atelopus walkeri Rivero, 1963

Critically Endangered A3ce Geographic Range This species is known from the northern slope Conservation Measures The range of the species includes Parque Nacional Natural Sierra Nevada de Santa Marta.
Order, Family: Anura, Bufonidae of the Parque Nacional Natural Sierra Nevada de Santa Marta in Surveys are needed to determine the current population status of this species. Disease management and captive-
Country Distribution: Colombia the Department of Magdalena, and from the Guatapuri river basin breeding programmes are recommended.
Current Population Trend: Decreasing in the department of Cesar, in northern Colombia, between 1,500 Bibliography: Acosta-Galvis, A.R. (2000), La Marca, E. et al. (2005), Lötters, S. (1996), Pounds, J.A. et al. (2006), Ruiz-Carranza, P.M.,
and 2,900m asl. Ardila-Robayo, M.C. and Lynch, J.D. (1996)
Population This species was considered to be common when it was Data Providers: Martha Patricia Ramírez Pinilla, Mariela Osorno-Muñoz, Jose Vicente Rueda, Adolfo Amézquita, María Cristina
last recorded in 1992. There have been no further surveys within its Ardila-Robayo
range since then.
Habitat and Ecology It occurs in sub-Andean and Andean forests. It
can also adapt to some modification of its habitat. It lays egg chains
in streams, where the tadpoles also develop.
Major Threats The most serious risk to this species is chytridio-
mycosis, which has had a devastating impact on other high-altitude
Atelopus species. Habitat fragmentation, primarily due to agriculture,
and pollution, resulting from agricultural practices and the fumigation
of crops, are also major threats.

CR Atelopus zeteki Dunn, 1933 PANAMANIAN GOLDEN FROG

Critically Endangered A2ace populations have been declining catastrophically due to chytridiomycosis, and the well-known El Copé population
Order, Family: Anura, Bufonidae collapsed and disappeared over the course of a few months in late 2004 (K. Lips pers. comm.). The chytridiomycosis
Country Distribution: Panama epidemic is spreading from west to east through Panama, and populations in the eastern part of its range are now
Current Population Trend: Decreasing at severe risk of disappearing.
CITES: Appendix I Habitat and Ecology It is a terrestrial species of tropical montane forest, with breeding and larval development
taking place in forest streams.
Major Threats The major threat is likely to be chytridiomycosis, which has led to catastrophic population declines in
many other species of montane Atelopus. The deforestation of habitat for both agriculture and general infrastructure
development, water pollution, and over collection for the pet trade are also threats to this species. In 2003, a road
from Sorá to El Valle was opened along the ridge of the Cordillera Central, resulting in heavy sedimentation of most
streams on the Pacific and Caribbean slopes, which has negatively affected a significant portion of the habitat used
by this species (R. Ibañez, in litt. to E. La Marca).
Conservation Measures This species is protected in Panama by national legislation (as Atelopus varius zetek) decree
© Paul Crump

Geographic Range This species is endemic to Panama, occurring east of the main Tabasará ridge in Provincias
Coclé and Panamá at 335-1,315m asl.
Population This species was reasonably common at a number of localities, and has been recorded as recently as
2005, but it is apparently less abundant north of El Copé, in comparison with observations in 1980. It is very rare or
extinct on Cerro Campana. It has been extinct in the El Valle de Antón for approximately 40 years. In recent years,
No. 23 of January 30, 1967. It has been recorded from the protected areas of Parque Nacional Altos de Campana
and Parque Nacional Omar Torrijos Herrera. A successful captive-breeding programme involving many zoos in North
America is in place, although no re-introductions will be made until existing threats can be addressed. Steps are also
under way to establishing a captive colony in Panama.
Bibliography: Cocroft, R.B. et al. (1990), Daly, J.W. et al. (1994), Ibáñez, R. et al. (2000), La Marca, E. et al. (2005), Lindquist, E.D. (1995),
Lindquist, E.D. and Hetherington, T.E. (1996), Lindquist, E.D. and Hetherington, T.E. (1998a), Lindquist, E.D. and Hetherington, T.E. (1998b),
Lindquist, E.D., Hetherington, T.E. and Badgley, D.D. (1999), Lötters, S. (1996), Pounds, J.A. et al. (2006), Ramos, C.W., Pimentel, N. and
Martinez-Cortes, V. (2002), Young, B. et al. (1999), Zippel, K. (2002)
Data Providers: Karen Lips, Frank Solís, Roberto Ibáñez, César Jaramillo, Querube Fuenmayor

CR Bufo amabilis Pramuk and Kadivar, 2003

Critically Endangered B1ab(iii) Geographic Range This species is known only from elevations of 2,050-2,200m asl in the Loja Basin, an inter-
Order, Family: Anura, Bufonidae Andean valley in Loja Province, Ecuador (Pramuk and Kadivar 2003). It is likely to have a restricted distribution, and
Country Distribution: Ecuador recent surveys confirm this.
Current Population Trend: Decreasing Population Past collections indicate that the species was fairly common at areas nearby creeks, even near planta-
tions; however, surveys undertaken between 1989 and 2001 failed to find the species. It appears that it has not been
collected since 1968, and a serious decrease might have taken place.
Habitat and Ecology The species has been collected in small pools and irrigation canals. Little is known of its habitat
requirements or ecology, but breeding is presumed to take place in freshwater by larval development.
Major Threats The apparent declines of this species might in part be due to the modification of much of the Loja
basin area for agriculture, urbanization, and other regional development. It appears that populations of this toad
in the area surrounding Provincia Loja have been severely affected by human activities (Pramuk and Kadivar 2003).
Disease might also be a factor but there is no evidence to confirm this.
© John D. Lynch

Conservation Measures The species is not known to occur within a protected area. More exhaustive searches are
needed to determine the population status of this species.
Bibliography: Pramuk, J.B. and Kadivar, F. (2003)
Data Providers: Luis A. Coloma

EN Bufo amatolicus Hewitt, 1925 AMATOLA TOAD

Endangered B1ab(ii,iii,iv,v)+2ab(ii,iii,iv,v) Geographic Range This species is known only from the Winterberg and Amatola Mountains, between Katberg and
Order, Family: Anura, Bufonidae Keiskammahoek, in Eastern Cape Province, South Africa, at 1,400-1,800m asl.
Country Distribution: South Africa Population It is locally moderately common, and congregates in large numbers to breed.
Current Population Trend: Decreasing Habitat and Ecology It breeds in shallow temporary pools and seepages in high-altitude moist grasslands, and is
absent from forests and plantations.
Major Threats The main threats are deforestation, overgrazing, and fires; over the last 20 years, about 20% of its
habitat has been lost to plantations.
Conservation Measures It occurs in the Hogsback Indigenous Forest and the Kubisi Indigenous Forest. Continued and
strengthened protection of these areas, and expanded protection of other remaining forest habitat, are necessary.
Bibliography: Branch, W.R. (1988), Channing, A. (2001), Minter, L.R. et al. (2004), Passmore, N.I. and Carruthers, V.C. (1995), Tandy, M.
and Keith, R. (1972), Wager, V.A. (1986)
© Richard Boycott

Data Providers: Leslie Minter, James Harrison

 Globally Threatened Amphibian Species 179

VU Bufo aucoinae O’Neill and Mendelson, 2004

Vulnerable B1ab(iii) Geographic Range This species is known from 25-500m asl in far north-western Panama and western Costa Rica,
Order, Family: Anura, Bufonidae from the Golfo Dulce area, north to the Rio Baru near Dominical.
Country Distribution: Costa Rica, Panama Population It is not abundant.
Current Population Trend: Decreasing Habitat and Ecology A nocturnal and terrestrial species that belongs to a clade of frogs considered intolerant of
habitat disturbance (J. Mendelson pers. comm.). Adults are restricted to primary forest, but can migrate across de-
graded habitat to large streams and rivers where they breed at the end of the wet season (Mid-December to January)
(Savage 2002; O’Neill and Mendelson 2004). It has not been found in forest patches < 10ha in size, but they also do
not occur in all large forest patches in the region. It is hypothesized that the secondary limiting factor is proximity
of appropriate breeding streams to occupied forest patch. The breeding stream requirements appear to be a wide,
low-gradient, rocky stream with small isolated pools with sun exposure.
Major Threats Habitat loss and fragmentation is a significant threat in western Costa Rica, due in particular to
© Twan Leenders

cattle ranching and plantation forestry. Loss of access to breeding streams and poisoning of rivers for the harvest of
freshwater shrimp are also problems (M. Ryan pers. comm.).
Conservation Measures The species is known from Corcovado, Braulio Carrillo, and Carara National Parks in
Costa Rica.
Notes on taxonomy: This species has recently been separated from Bufo melanochlorus (O’Neill and Mendelson 2004).
Bibliography: O’Neill, E.M. and Mendelson, J.R. (2004), Savage, J.M. (2002)
Data Providers: Joseph Mendelson

EN Bufo beddomii Günther, 1875

Endangered B1ab(iii) Geographic Range This species is believed to be endemic to the southern Western Ghats of India, and has been
Order, Family: Anura, Bufonidae recorded from Kudremukh, Ponmudi and Kalakad. Specimens collected in Maharashtra are misidentifications of Bufo
Country Distribution: India c.f. scaber (S.D. Biju pers. comm.). It has an altitudinal range of 1,000-1,500m asl.
Current Population Trend: Decreasing Population This is a rare species, and there are no estimates of population size or trends. Populations are fragmented,
and the species is believed to be declining throughout much of its range.
Habitat and Ecology It is a terrestrial species primarily associated with montane tropical moist forest, and is
presumed to breed in streams. Individuals have been recorded in abandoned eucalyptus plantations. It has not been
recorded from agricultural areas.
Major Threats The main threat is the conversion of areas of suitable habitat to intensive plantations (tea, eucalyptus
and Acacia), and industrial development and urbanization around Ponmudi.
© S.U. Saravanakumar

Conservation Measures It has been recorded from the Kalakad-Mundanthurai Tiger Reserve (Tamil Nadu) and
Kudremukh National Park (Karnataka). The species has been the focus of recent studies (S.D. Biju 1998-2001).
Notes on taxonomy: Many museum specimens assigned to Bufo beddomi are misidentifications (Biju 2001).
Bibliography: Biju, S.D. (2001), Dutta, S.K. (1997), Krishnamurthy, S.V. and Hussain (2000), Ravichandran, M.S. (1996a), Ravichandran,
M.S., and Pillai, R.S. (1990), Ravichandran, M.S., and Pillai, R.S. (1990), Vasudevan, K., Kumar, A. and Chellam, R. (2001)
Data Providers: S.D. Biju, Sushil Dutta, M.S. Ravichandran Karthikeyan Vasudevan, S.P. Vijayakumar, Chelmala Srinivasulu, Gajanan
Dasaramji Bhuddhe

EN Bufo brauni Nieden, 1911

Endangered B1ab(iii,v) Geographic Range This species occurs in the West and East Usambaras, the Ulugurus, and the Udzungwa Mountains
Order, Family: Anura, Bufonidae in eastern Tanzania, from 750m asl (perhaps lower) to 1,800m asl.
Country Distribution: Tanzania Population It is not an uncommon species. However, there is evidence that it is becoming more scarce.
Current Population Trend: Decreasing Habitat and Ecology It lives in submontane and montane forest zones, occurring in leaf-litter on the forest floor,
and breeding in streams. It tolerates limited habitat disturbance, but is not found in open habitats, except when
these are close to forest.
Major Threats It is probably adversely affected by forest loss for agriculture, collection of wood, and human settle-
ment, and its habitat in the East Usambaras has recently come under serious threat as a result of the activities of
illegal gold miners.
Conservation Measures It occurs in Amani Nature Reserve and Udzungwa National Park, but expanded and
© Michele Menegon

strengthened protection of these and other forest reserves in the Eastern Arc mountains is necessary. The species
requires close population monitoring.
Bibliography: Harper, E. and Vonesh, J.R. (2003), Howell, K.M. (1993), Poynton, J.C. (1998), Poynton, J.C. (2003b), Tandy, M. and
Keith, R. (1972)
Data Providers: Kim Howell, John Poynton, Mills Tandy

EN Bufo caeruleostictus Günther, 1859

Endangered A2c Geographic Range This species is known from several localities in western Ecuador in the north-western Andean
Order, Family: Anura, Bufonidae montane forest and western Ecuador moist forests. It has been recorded from 40-2,000m asl.
Country Distribution: Ecuador Population It is a rare species.
Current Population Trend: Decreasing Habitat and Ecology It is found from the lower Pacific Andean slopes up to cloud forests. In Las Pampas, the habitat
of this species was described as being close to rivers/creeks in low montane cloud forest (Hoogmoed 1989). There
is no information about its breeding habitats although it is presumed to be a larval developer.
Major Threats The major threat to the species is habitat loss due to agricultural expansion, both for crops and
livestock ranching, as well as logging and wood plantations. The invasive bullfrog (Rana catesbeiana) is probably
also competing with this species.
Conservation Measures The species is likely to be present in several protected areas. Further research is needed into
the breeding biology of this species, and the possible adverse effects of competition with the invasive bullfrog.
© Luis A. Coloma

Bibliography: Günther, A.C.L.G. (1859), Hoogmoed, M.S. (1989)

Data Providers: Diego Cisneros-Heredia, Ana Almandáriz, Mario Yánez-Muñoz, Luis A. Coloma, Santiago Ron
180 Threatened Amphibians of the World

EN Bufo californicus Camp, 1915 ARROYO TOAD

Endangered A2ac located very small populations in four creeks in south-western Riverside County (Temecula, Arroyo Seco, San Mateo,
Order, Family: Anura, Bufonidae and Tenaja creeks) (USFWS 1994). The single recent record in San Bernardino County is from Deep Creek in the San
Country Distribution: Mexico, United States Bernardino National Forest. It is also still extant in north-western Baja California.
of America Population The total estimated breeding population is less than 3,000 individuals (USFWS, Federal Register, 6 May
Current Population Trend: Decreasing 1998). Only six of the 22 extant populations south of Ventura are known to contain more than a dozen adults (USFWS
1994). It has been extirpated from an estimated 75% of its former range in the USA (Sweet, in USFWS 1993). Reproduc-
tive success has been poor in recent years (USFWS 1993). There is very little recent information on the trends in this
species in Mexico. It occurs in at least a couple dozen sites, but viable populations may remain in only five drainages
(USFWS 1993). It is known in California from 22 river basins in nine counties (USFWS 1999).
Habitat and Ecology This species inhabits washes, streams, and arroyos, and adjacent uplands (in desert and
shrubland). It is also found on sandy banks in riparian woodlands (willow, cottonwood, sycamore, and/or coast live
oak) in California. It is also found along rivers that have shallow gravelly pools adjacent to sandy terraces (USFWS
© Andrew Borcher

Geographic Range This species formerly ranged from San Luis Obispo County, California, USA, south to north-west-
ern Baja California in Mexico; see Gergus, Sullivan and Malmos 1997 for southernmost record. It is now apparently
extirpated in San Luis Obispo County; populations persist in headwater areas of streams in Santa Barbara, Ventura,
Los Angeles, Riverside, and San Diego counties; recent sightings of scattered individuals have been reported from
Orange, San Bernardino, and southern Imperial counties (USFWS 1994). The majority of the remaining populations in
Santa Barbara and Ventura counties are in Los Padres National Forest (five viable populations); Sespe Creek in Ventura
County has the largest known population. Other populations occur in the Sisquoc, Santa Ynez, and upper and lower
Piru drainages (USFWS 1994). It also occurs in San Diego County along the Santa Margarita, Guejito, Sweetwater,
Vallecito, San Luis Rey, Santa Ysabel, Witch, Cottonwood, Temescal, Agua Caliente, Santa Maria, Lusardi, Pine Valley,
Noble, Kitchen, Long Potrero, Upper San Diego, San Vicente, and Morena drainages; populations in the Temescal,
Agua Caliente, Pine Valley, and Cottonwood drainages may be considered viable (USFWS 1994). It was recently
recorded in Whitewater Canyon in Riverside County (Patten and Myers, 1992, Herpetol. Rev. 23:122). Recent surveys
1993). Adults obtain shelter by burrowing into sandy soil. It lays eggs among gravel, leaves, or sticks, or on mud or
clean sand, at the bottom of shallow quiet waters of streams or shallow ponds, in areas with little or no emergent
vegetation. Newly metamorphosed individuals remain near pools for up to several weeks (until the pools dry up).
Major Threats Threats include habitat degradation (mainly through urbanization, dam construction and ill-timed water
releases, agriculture, road construction, off-road vehicle use, overgrazing, and mining activities, and also via drought and
wildfires), recreational use of habitat (which causes habitat degradation and direct mortality), predation by introduced
fishes and bullfrogs (Rana catesbeiana), and small population sizes (see USFWS 1993 and 1994 for further details).
Conservation Measures USFWS (2001) designated as critical habitat approximately 182,360 acres in 22 units
in Monterey, Santa Barbara, Ventura, Los Angeles, San Bernardino, Riverside, Orange, and San Diego counties.
Critical habitat includes stream and river courses, riparian areas, and adjacent terrace and upland habitats. It does
not include existing structures or other developed areas. See recovery plan (USFWS 1999). In Mexico this species is
found within San Pedro Martir National Park.
Bibliography: Behler, J.L. and King, F.W. (1979), Blackburn, L., Nanjappa, P. and Lannoo, M.J. (2001), Collins, J.T. (1991), Gergus, E.W.A.
(1998), Gergus, E.W.A., Grismer, L.L., and Beaman, K. (1997), Gergus, E.W.A., Sullivan, B.K. and Malmos, K.B. (1997), Price, A.H. and
Sullivan, B.K. (1988), Stebbins, R.C. (1954), Stebbins, R.C. (1972), Stebbins, R.C. (1985b), Sullivan, B.K. (1992), U.S. Fish and Wildlife
Service (1993), U.S. Fish and Wildlife Service (1994a), U.S. Fish and Wildlife Service (1999b), U.S. Fish and Wildlife Service (2000g),
U.S. Fish and Wildlife Service (2001a)
Data Providers: Geoffrey Hammerson, Georgina Santos-Barrera

EN Bufo canorus Camp, 1916 YOSEMITE TOAD

Endangered A2ae breed every other year or once every three years. It persists in meadow habitats degraded by cattle as well as in
Order, Family: Anura, Bufonidae lakes stocked with non-native trout.
Country Distribution: United States of America Major Threats Leading hypotheses for the declines are disease (chytridiomycosis), airborne contaminants, and
Current Population Trend: Decreasing livestock grazing. An examination of preserved specimens from a 1970 die-off found multiple pathogens, but no single
pathogen was present in more than 25% of the specimens, suggesting that the animals suffered from suppressed
immune systems (Green and Kagarise Sherman 2001). Davidson, Shaffer and Jennings (2002) found a weak pattern
between declines at sites and amount of agricultural land upwind (suggesting that windborne agrochemicals may
have contributed to declines). Livestock grazing may have detrimental impacts on Yosemite Toads through trampling,
© Andreas & Christel Nöllert

Geographic Range This species is endemic to the state of California, USA. It is found only in the high sierra from
the Blue Lakes region north of Ebbets Pass (Alpine County) south to Spanish Mountain area (Fresno County), and is
found at elevations of 1,460-3,630m asl.
Population The total adult population size is unknown but is likely to be at least a few thousand. Declines, some
in seemingly pristine environments, occurred in the eastern Sierra Nevada between the early 1970s and early 1990s
(Kagarise Sherman and Morton 1993). Although still distributed over most of its original range, and many populations
have active breeding and recruitment (Shaffer et al. 2000), the species has declined or disappeared from more than
50% of the sites from which it has been recorded (Jennings and Hayes 1994; Drost and Fellers 1996). USFWS (2000)
reviewed additional evidence of declines in distribution and abundance.
Habitat and Ecology It inhabits wet mountain meadows and borders of forests, and obtains shelter in rodent bur-
rows as well as in dense vegetation. This species breeds in shallow edges of snowmelt pools and ponds or along
edges of lakes and slow-moving streams. Some breeding sites dry up before larvae metamorphose. Females may
alteration of meadow habitat, and possible lowered water quality (D. Martin pers. comm.). Other factors that may
have contributed to declines are the 1980s California drought, and predation by introduced trout. Ultraviolet radiation
is not suspected to be a major contributor to declines based on fieldwork by Sadinski (pers. comm.).
Conservation Measures Most of the habitat of this species is confined within protected areas including: Yosemite
National Park, Kings Canyon National Park, wilderness areas of Eldorado, Inyo, Stanislaus, and Sierra National Forest,
with the highest number of populations recorded in Sierra National Forest followed by Stanislaus National Forest.
Off-highway vehicle use, pack stock and cattle grazing still occur in National Forests, as well as protected wilderness
areas; only pack stock use continues in Yosemite National Park and Kings Canyon National Park. Because declines
have occurred in pristine areas in parks, no occurrences can be regarded as adequately protected. This species has
been federally petitioned for listing under the Endangered Species Act in March 2000 by the Pacific Rivers Council
and Centre for Biological Diversity. In December 2002 the US Fish and Wildlife Service published a decision in the
Federal Register that placed the toad on the “warranted-but-precluded” list due to higher priority listings. Further
taxonomic work is required to determine the status of this species relative to B. exsul.
Notes on taxonomy: Molecular data suggest that Bufo exsul is possibly conspecific with this species.
Bibliography: Behler, J.L. and King, F.W. (1979), Blackburn, L., Nanjappa, P. and Lannoo, M.J. (2001), Bradford, D.F., Swanson, C. and
Gordon, M.S. (1992), Camp, C.L. (1916), Cunningham, J.D. (1963), Davidson, C., Shaffer, H.B. and Jennings, M.R. (2002), Drost, C.A. and
Fellers, G.M. (1996), Frost, D.R. (1985), Green, D.E. and Kagarise Sherman, C. (2001), Grinnel, J. and Storer, T.I. (1924), Jennings, M.R.
and Hayes, M.P. (1994), Kagarise Sherman, C. and Morton, M.L. (1993), Karlstrom, E.L. (1962), Karlstrom, E.L. (1973), Karlstrom, E.L.
and Livezy, R.L. (1955), Martin, D.L (1991), Mullally, D.P (1953), Shaffer, H.B. et al. (2000), Sherman, C.K. (1980), Stebbins, R.C. (1972),
Stebbins, R.C. (1985a), Stebbins, R.C. (2003), U.S. Fish and Wildlife Service (2000b)
Data Providers: Geoffrey Hammerson, Rob Grasso, Carlos Davidson

EN Bufo cataulaciceps Schwartz, 1959 CUBAN PINE TOAD

Endangered B1ab(iii) Geographic Range This species has a restricted range in the Conservation Measures The range of the species includes the Reserva Ecológica los Indios.
Order, Family: Anura, Bufonidae lowlands of the Isla de Juventud and extreme western Cuba. It has Bibliography: Díaz, L.M. (1998a), Hedges, S.B. (1993), Hedges, S.B. (1999), Hedges, S.B. (2001), Henderson, R.W. and Powell, R (1999),
Country Distribution: Cuba been recorded from sea level up to 50m asl. Henderson, R.W. and Powell, R. (2001), Schwartz, A. (1959b), Schwartz, A. and Henderson, R.W. (1991)
Current Population Trend: Decreasing Population It can be very common at breeding aggregations, but Data Providers: Blair Hedges, Luis Díaz
is otherwise hard to find.
Habitat and Ecology It inhabits savannah habitat, with pinewood
and palms, with sandy soils. Males call from temporary pools (rain
puddles), flooded pastures, and other shallow still water (which is
also where the eggs are laid). It is an explosive breeder that breeds
after heavy rains.
Major Threats The main threats to this species are habitat loss and
degradation due to intensive agriculture and the extraction of sand
for the glass industry. The invasive Dichrostachys cinerea (Marabu),
a thorny, fast-growing woody species native to Africa that was
introduced to the Caribbean in the 19th century, also poses a threat
to native habitat, and is difficult and expensive to control.
 Globally Threatened Amphibian Species 181

EN Bufo cavifrons Firschein, 1950 MOUNTAIN TOAD

Endangered B1ab(iii)+2ab(iii) Geographic Range This species occurs only in the Sierra de Los Bibliography: Mendelson III, J.R. (1997a), Mendelson III, J.R. (1998a), Porter, K. (1963)
Order, Family: Anura, Bufonidae Tuxtlas and a few other localities near the Santa Marta volcano, in Data Providers: Georgina Santos-Barrera, Oscar Flores-Villela
Country Distribution: Mexico southern Veracruz, Mexico. It is known from 200-1,600m asl.
Current Population Trend: Decreasing Population This is a naturally rare species.
Habitat and Ecology It inhabits tropical pine-oak forests with
abundant leaf-litter. It is a stream-breeding amphibian.
Major Threats The major threat to this species is loss and degrada-
tion of its pine-oak forest habitat due to agricultural activities (crops
and livestock), wood extraction, and infrastructure development.
Conservation Measures Although this species can be found in pro-
tected areas, the protection of these areas is not always guaranteed,
and some are subject to human disturbance; improved management
of these is urgently needed. This species is protected by Mexican
law under the “Special Protection” category (Pr).

CR Bufo chavin Lehr, Kohler, Aguilar and Ponce, 2001

Critically Endangered B2ab(iii,v) Geographic Range This species is only recorded from Palma Pampa and Cordillera de Carpish, Departamento Huá-
Order, Family: Anura, Bufonidae nuco, Peru. It is believed to have a genuinely restricted distribution, with an altitudinal range of 2,600-3,072m asl.
Country Distribution: Peru Population This species is uncommon and probably declining.
Current Population Trend: Decreasing Habitat and Ecology It is a species of primary cloud forest on the eastern Andean slopes of central Peru. It has also
been found in disturbed areas at the forest edge. Its breeding habitat is not known, but it may breed in bromeliads,
presumably by larval development.
Major Threats Deforestation for agriculture (potatoes) is a major threat. Agrochemical use has also contributed to
the decline of this species at known sites.
Conservation Measures The species is not known to be present in protected areas, and its remaining habitat is in
urgent need of protection. The current population status of this species also needs to be determined.
Bibliography: Lehr, E. (2002), Lehr, E. et al. (2001), Lehr, E., Rodriguez, D. and Cordova, J.H. (2002)
Data Providers: Ariadne Angulo, Edgar Lehr, César Aguilar Puntriano
© Edgar Lehr

EN Bufo claviger Peters, 1863

Endangered B1ab(iii)+2ab(iii) Geographic Range This species is only known with certainty from forest habitats is clearly necessary.
Order, Family: Anura, Bufonidae northern Bengkulu Province, Sumatra, Indonesia, where it occurs Bibliography: Iskandar, D.T. and Colijn, E. (2000)
Country Distribution: Indonesia at low altitudes. It has not been found in West Sumatra, despite Data Providers: Djoko Iskandar, Mumpuni
Current Population Trend: Decreasing attempts to locate it there. There is an old record from Nias Island
but there are doubts that the locality of this specimen has been
correctly recorded.
Population It is probably rare, since a number of expeditions
have collected only a few specimens, and it probably occurs at
low densities.
Habitat and Ecology It lives in lowland forest, but it is not known
whether or not it can adapt to secondary habitats. It probably breeds
by larval development in slow-flowing streams.
Major Threats The primary threat is likely to be habitat conversion
due to smallholder farming and logging activities.
Conservation Measures Although it is known to occur in forest
in a recreational area for tourists, expanded protection of lowland

VU Bufo corynetes Duellman and Ochoa, 1991

Vulnerable D2 Geographic Range This species’ appears to be restricted to a west-facing slope descending into the Amazonian
Order, Family: Anura, Bufonidae lowlands of Abra Malaga and San Luis, Provincia Urubamba (Departamento Cuzco), Peru. Its altitudinal range is
Country Distribution: Peru 2,000-3,200m asl.
© Natural History Museum and Biodiversity Research

Current Population Trend: Stable Population It is a very rare species.

Habitat and Ecology This is a forest edge species specifically restricted to the zone of puna grassland directly
adjacent to montane forest; it is not known whether or not it can occur in modified habitats. Its breeding habitat is
not known, though breeding probably takes place in water.
Major Threats There are not believed to be any major threats to this species at present; however, it might be
susceptible to habitat loss in the future.
Center, University of Kansas

Conservation Measures It is not known from any protected areas. There is a need for close monitoring of the
population status of this species.
Bibliography: Cordova, J.H. (1999), Duellman, W.E. and Ochoa, O. (1991), Duellman, W.E. and Schülte, R. (1992), Instituto Nacional de
Recursos Naturales (INRENA) (2000), Rodríguez, L.O., Cordova, J.H. and Icochea, J. (1993)
Data Providers: Ariadne Angulo, Wilfredo Arizabal, Jesús Córdova-Santa Gadea
182 Threatened Amphibians of the World

CR Bufo cristatus Wiegmann, 1833 LARGE-CRESTED TOAD

Critically Endangered B1ab(i,ii,iii,iv,v)+2ab Geographic Range This species is found in the central Sierra Madre Oriental, Mexico, in the states of Puebla
(i,ii,iii,iv,v) and Veracruz. However, it only survives in northern Puebla at Apulco, at 1,400m asl, and Barranca de Xocoyolo, at
Order, Family: Anura, Bufonidae about 1,300m asl.
Country Distribution: Mexico Population This species was believed to be extinct and has only recently been rediscovered at Apulco and Barranca
Current Population Trend: Decreasing de Xocoyolo, where it is uncommon. There have been no recent records of the species at any of the previously known
localities in Veracruz, and it is believed to be extinct in these sites.
Habitat and Ecology It inhabits pristine cloud forests (dominated by oak) and breeding takes place in streams by
larval development.
Major Threats The disappearance and disturbance of the pine-oak forests, as well as the pollution and desiccation
of streams, are the major threats to this species. The habitat has been completely destroyed in the localities where
this species appears to have been extirpated.
Conservation Measures Protection of the original pine-oak forest and measures to avoid the pollution of streams
from anthropogenic activities are required to protect this species. This species is protected by Mexican law under
© Ronald Altig

the “Special Protection” category (Pr).

Bibliography: Brandt, T. and Buschmann, H. (2004), Gleed-Owen, C. (1996), Mendelson III, J.R. (1997a), Mendelson III, J.R. (1998b),
Mendelson III, J.R. and Canseco-Márquez, L. (1998)
Data Providers: Georgina Santos-Barrera, Luis Canseco-Márquez

VU Bufo cycladen Lynch and Smith, 1966

Vulnerable A2a; B1ab(v) Geographic Range This species occurs in a relatively narrow altitudinal band around 750-1,000m asl on the Pacific
Order, Family: Anura, Bufonidae slope of the Sierra Madre del Sur in Guerrero and Oaxaca states, Mexico. Most specimens have been collected around
Country Distribution: Mexico the private hacienda of Agua del Obispo, along the old highway between Tierra Colorada and Chilpancingo, Guerrero.
Current Population Trend: Decreasing One other series is known from Putla de Guerrero, about 150km east on the same slope of the Sierra Madre del Sur,
Oaxaca. The distribution is presumed to be continuous between these two localities at mid elevations, and might
well continue further east or west as this region is undersampled for amphibians.
Population Field notes and collections by E.H. Taylor, H.M. Smith, and W.E. Duellman in the 1940s, 1950s and 1960s
indicate that this species was common in the Agua del Obispo area. Approximately five weeks of fieldwork there during
2000 and 2003 resulted in the discovery of just a single individual, indicating a probable serious decline.
Habitat and Ecology This species is known from the ecotone of scattered small pines and brush between lower
altitude tropical deciduous forest and higher altitude pine-oak forest. The species is presumed to be a larval de-
© J.R. Mendelson

veloper.
Major Threats Although this species is believed to be tolerant of some degree of habitat alteration, such as selec-
tive logging, large-scale agricultural clearance and agrochemicals are likely to cause significant impacts. There is
no direct evidence that chytridiomycosis is a threat to this species, but chytrid is known from this locality (Lips et al.
2004) and the drastic population decline is consistent with such an impact.
Conservation Measures This species is not known from any protected areas, but might occur in the lower reaches
of the Omilterne Reserve. There is a need for improved habitat protection at sites where this species is known to
occur. Further research is required to investigate the potential threat posed by chytridiomycosis.
Notes on taxonomy: This species was removed from the synonymy of Bufo coccifer by Mendelson et al. (2005).
Bibliography: Lips, K.R. et al. (2004), Lynch, J.D. and Smith, H.M. (1966), Mendelson III, J.R. et al. (2005)
Data Providers: Joseph Mendelson

EN Bufo djohongensis Hulselmans, 1977

Endangered B2ab(iii) Geographic Range This species is known only from the Adamawa Notes on taxonomy: The taxonomic status of this species is uncertain. It was described as a subspecies of Bufo funereus (Hulselmans
Order, Family: Anura, Bufonidae Plateau in north-central Cameroon. 1977), and was considered a distinct species by Joger (1982) and Böhme and Schneider (1987). J.-L. Amiet (pers. comm.) considers that
Country Distribution: Cameroon Population There is very little information on its population status, it is a junior synonym of Bufo villiersi, an opinion not shared by U. Joger (pers. comm.).
Current Population Trend: Decreasing and it has only been recorded on a few occasions. Bibliography: Böhme, W. and Schneider, B. (1987), Hulselmans, J.L.J. (1977), Joger, U. (1982)
Habitat and Ecology It lives in gallery forests in a landscape of Data Providers: Ulrich Joger, Mills Tandy, Jean-Louis Amiet
montane grassland and wooded savannah, and has been found
breeding in streams.
Major Threats It is probably suffering from continuing degradation
of its montane habitats as a result of expanding agriculture, overgraz-
ing by livestock, wood extraction, fire, and human settlement.
Conservation Measures It has not been recorded from any
protected areas, and some form of protection of the remaining
habitat on the Adamawa Plateau is urgently needed. Further survey
work is necessary to determine the current population status of
this species.

VU Bufo empusus (Cope, 1862) CUBAN SMALL-EARED TOAD

Vulnerable B2ab(iii) Geographic Range This species has a wide but patchy distribu- Conservation Measures It occurs in several protected areas. There is a need to control the spread of invasive
Order, Family: Anura, Bufonidae tion in lowland Cuba and the Isla de Juventud. It ranges from sea species within these existing protected areas.
Country Distribution: Cuba level up to 70m asl. Bibliography: Hedges, S.B. (1993), Hedges, S.B. (1999), Hedges, S.B. (2001), Henderson, R.W. and Powell, R (1999), Henderson, R.W.
Current Population Trend: Decreasing Population It is abundant during breeding, but otherwise burrows and Powell, R. (2001), Schwartz, A. and Henderson, R.W. (1991)
underground and is rarely seen. Data Providers: Blair Hedges, Luis Díaz
Habitat and Ecology It inhabits xeric and mesic lowland forests
and savannahs. The males call from flooded ditches and large tem-
porary pools of rainwater, and the eggs are laid in still water. It is
an explosive breeder, though outside the breeding season it remains
in burrows below ground.
Major Threats The major threats to this species are habitat loss
and degradation, due to agriculture (it occurs in an area of intense
agricultural production), and pollution from agricultural pesticides.
The spread of the invasive plant species Dichrostachys cinerea is
also threatening this species’ native habitat.
 Globally Threatened Amphibian Species
VU Bufo exsul Myers, 1942
Vulnerable D2
Order, Family: Anura, Bufonidae
Country Distribution: United States of America
Current Population Trend: Stable
© Kenny Wray
Geographic Range This species’ range includes several springs feeding Deep Springs Lake, in Deep Springs Valley,
Inyo County, California, USA: Buckhorn Spring, Corral Spring and adjacent ponds, and Bog Mound Springs at about
1,520m; also Antelope Springs at about 1,710m (Murphy, Simandle and Becker 2003, Stebbins 2003). Apparently,
this species was introduced to a flowing well in Saline Valley, Death Valley National Park, Inyo County, California
(Murphy, Simandle and Becker 2003), and also introduced at Batchelder Spring, Westgard Pass, Inyo County, but is
possibly extirpated there.
Population The population of this species was regarded as more or less stable in the early 1970s (Bury, Dodd and
CR Bufo fastidiosus (Cope, 1875)
Critically Endangered A2ace
Order, Family: Anura, Bufonidae
Country Distribution: Costa Rica, Panama
Current Population Trend: Decreasing
© Karen Lips
CR Bufo fluviaticus Schwartz, 1972
Critically Endangered B2ab(iii) Geographic Range This species has a very restricted range (only
Order, Family: Anura, Bufonidae two known localities) in north-western Dominican Republic. Its
Country Distribution: Dominican Republic altitudinal range is 150-175m asl.
Current Population Trend: Decreasing Population It is presumably very rare. It has not been encountered
by any herpetologist in the three decades since it was discovered,
including by herpetologists who have collected extensively on the
island, as recently as 2003 (M. Hernandez pers. comm.).
Habitat and Ecology It is found in xeric habitats with broadleaf
gallery forest, typically in close proximity to streams. Males call
from shallow running water, and eggs are laid in still water. It is
not known whether or not animals stay close to streams or spread
out into xeric habitats.
Major Threats The species is threatened by habitat destruction from
agriculture (crops and livestock) and subsistence wood collection.
EN Bufo fractus Schwartz, 1972
Endangered B1ab(iii) Geographic Range This species has a fragmented distribution in
Order, Family: Anura, Bufonidae eastern Dominican Republic, and is known from six to eight locations
Country Distribution: Dominican Republic very close together. The exact elevational range is not known but it
Current Population Trend: Decreasing is definitely known from only below 500m asl.
Population Althought this species was formerly known to be
abundant at a breeding aggregation near the town of Higuey in 1972,
extensive surveys in the region in 2003 did not find any individuals
(M. Hernandez pers. comm.).
Habitat and Ecology So far it has only been recorded from degraded
forest; however, it is difficult to find outside the breeding season and
so may just not have been recorded from primary forest yet. Eggs are
laid in temporary pools, and the larvae develop in the pools.
Major Threats The main threat to the species is habitat loss and
degradation largely due to intensive agriculture, but also due to human
settlement and livestock grazing. The development of new tourism
facilities nearby poses an additional threat to the existing habitat.
183
BLACK TOAD
Fellers 1980), and no significant change was reported in 1990 (California Department of Fish and Game 1990, Schui-
erer and Anderson 1990). The population at Antelope Springs (apparently introduced) was reported to have died out
(Stebbins 1985b), but is still referred to by both Murphy, Simandle and Becker (2003) and Stebbins (2003). Historical
population data is as follows: over 4,000 individuals (1971 survey), more than 80,000 individuals (1977 estimate). The
population at Corral Spring(s) was reported in 1980 as more than 22,000 (Sherman 1980).
Habitat and Ecology Its primary habitat is watercourses/marshes (grass, sedge, dwarf bulrush, and watercress),
formed by water flow from springs, and surrounded by desert with low bushes. Adults are more aquatic than other
toad species in California. Adults prefer habitats with short plant cover and unobstructed access to still or slowly
flowing water (Schuierer and Anderson 1990). It retreats to rodent burrows or other refuges in winter. It breeds in
shallow marsh and pond waters (Schuierer and Anderson 1990).
Major Threats At present, there do not appear to be any major threats to this species. However, potential future
threats include habitat destruction from irrigation schemes or other factors resulting in water table alteration,
recreational vehicle use, cattle overgrazing, and predation by carp.
Conservation Measures The Department of Fish and Game has purchased 719 acres to protect the habitat at Deep
Springs; Deep Spring College (owner of property at Buckhorn and Antelope springs) has fenced an area to exclude
livestock and is manipulating irrigation water to minimize impacts on breeding toads, eggs, and larvae (California
DF&G 1990). Other sites are on BLM land, but the level of protection is uncertain.
Notes on taxonomy: Molecular data suggest that this species might be conspecific with Bufo canorus, but this requires further
investigation.
Bibliography: Behler, J.L. and King, F.W. (1979), Blackburn, L., Nanjappa, P. and Lannoo, M.J. (2001), Bury, R.B., Dodd, Jr., C.K. and
Fellers, G.M. (1980), California Department of Fish and Game (CDF&G). (1990), Frost, D.R. (1985), Murphy, J.F., Simandle, E.T. and Becker,
D.E. (2003), Myers, G.S. (1942c), Schuierer, F.W. (1962), Schuierer, F.W. (1963), Schuierer, F.W. (1972), Shaffer, H.B. et al. (2000), Sherman,
C.K. (1980), Stebbins, R.C. (1985b), Stebbins, R.C. (2003)
Data Providers: Geoffrey Hammerson
Geographic Range This species occurs on both slopes of the southern Cordillera de Talamanca of Costa Rica and
the Atlantic slope of immediately adjacent Panama, from 760-2,100m asl (Savage 2002).
Population The species was once abundant in Costa Rica, but it has undergone marked declines since the 1980s.
There is no information available on the population size or abundance of this species in Panama, but it is presumed
to have declined.
Habitat and Ecology Its habitat is premontane and lower montane rainforest. It is a diurnal terrestrial species that
is largely fossorial, and has been excavated from leaf-litter along stream banks in August. Juveniles have been found
on rocky stream margins throughout the year. This species is an explosive breeder, with reproduction taking place in
ephemeral pools after heavy rain in late April to May; eggs are laid in the shallow pools (Savage 2002).
Major Threats The main cause of the population decline is likely to be chytridiomycosis. In addition, there is general
habitat loss across the range due to agriculture and timber extraction.
Conservation Measures The species is known to occur in Parque Internacional La Amistad, a transboundary pro-
tected area between Panama and Costa Rica. Given the threat of chytridiomycosis, the implementation of a disease
management and captive-breeding programme is recommended.
Bibliography: Graybeal, A. and de Queiroz, K. (1992), Ibáñez, R. et al. (2000), Lips, K.R. (1998), Lips, K.R. and Krempels, D.M. (1995),
Savage, J.M. (2002), Young, B. et al. (1999)
Data Providers: Federico Bolaños, Frank Solís, Roberto Ibáñez, Gerardo Chaves, Jay Savage, César Jaramillo, Querube Fuenmayor
HISPANIOLAN CRESTLESS TOAD
Conservation Measures It is not known to occur in any protected areas. Additional survey efforts are required to
determine whether or not this species survives in the wild.
Bibliography: Hedges, S.B. (1993), Hedges, S.B. (1999), Hedges, S.B. (2001), Henderson, R.W. and Powell, R (1999), Henderson, R.W.
and Powell, R. (2001), Powell, R. et al. (2000), Powell, R. and Pregill, G.K. (1991), Schwartz, A. (1972), Schwartz, A. and Henderson,
R.W. (1991)
Data Providers: Blair Hedges, Sixto Inchaustegui, Marcelino Hernandez, Robert Powell
EASTERN CRESTED TOAD
Conservation Measures It is not known to occur in any protected areas, and protection of the existing habitat is
urgently needed.
Notes on taxonomy: This species was elevated from subspecific status under Bufo guentheri by Powell (1993).
Bibliography: Hedges, S.B. (1993), Hedges, S.B. (1999), Hedges, S.B. (2001), Henderson, R.W. and Powell, R (1999), Henderson, R.W.
and Powell, R. (2001), Powell, R. (1992), Powell, R. (1993), Powell, R. et al. (2000), Schwartz, A. (1972), Schwartz, A. and Henderson,
R.W. (1991)
Data Providers: Blair Hedges, Sixto Inchaustegui, Marcelino Hernandez, Robert Powell
184 Threatened Amphibians of the World

EN Bufo gallardoi Carrizo, 1992

Endangered B1ab(iii) Geographic Range This species occurs in Jujuy, Ledesma and Calilegua in Argentina from 1,000-1,700m asl.
Order, Family: Anura, Bufonidae Population It is a rare species, but it is nevertheless recorded on a regular basis.
Country Distribution: Argentina Habitat and Ecology It is restricted to montane forests (Yungas). It occurs on the ground and reproduces in slow-
Current Population Trend: Decreasing flowing streams where the larvae also develop. Its tolerance to habitat disturbance is not known.
Major Threats The main threats include selective logging and clear cutting of primary forests, the introduction of
predatory fish (trout), and the alteration of watersheds.
Conservation Measures Some populations are protected in Parque Nacional Calilegua, which was created to
preserve a representative part of the Yungas forest. Strengthened protection of this reserve, and expanded protection
of other remaining habitat, is recommended.
Bibliography: Cochran, D.M. (1955), Lavilla, E.O. et al. (2000), Lavilla, E.O. and Cei, J.M. (2001), Vaira, M. (2002)
Data Providers: Esteban Lavilla
© Marcos Vaira

EN Bufo gemmifer Taylor, 1940 JEWELED TOAD

Endangered B1ab(iii)+2ab(iii) Geographic Range This species is known only from central coastal
Order, Family: Anura, Bufonidae Guerrero, Mexico, in north-western Acapulco city.
Country Distribution: Mexico Population It is thought to be rare.
Current Population Trend: Decreasing Habitat and Ecology The primary habitat is xeric and deciduous
forest, and it has never been recorded out of the lowlands near
Acapulco. It is a stream-breeding amphibian.
Major Threats The main threat is habitat loss due to agriculatural
expansion, wood extraction, and the expansion of plantations.
Conservation Measures Protection of the suburban and tropical
dry areas around Acapulco represents the only chance for the survival
of this species. It is protected by Mexican law under the “Special
Protection” category (Pr).
Notes on taxonomy: This species is currently under taxonomic review.
Bibliography: Porter, K. (1963)
Data Providers: Georgina Santos-Barrera, Oscar Flores-Villela

VU Bufo guentheri Cochran, 1941

Vulnerable B2ab(iii) Geographic Range This species is widely but patchily distributed in southern, central, and northern and north-western
Order, Family: Anura, Bufonidae Hispaniola (both Haiti and Dominican Republic): Plaine du Nord-Valle de Cibao, Valle de San Juan, and Plaine du Cul
Country Distribution: Dominican Republic, Haiti de Sac-Valle de Neiba. The altitudinal range is from 40m below sea level up to 107m asl.
Current Population Trend: Decreasing Population In suitable habitats, this species can be found all year round. However, according to B. Hedges (pers.
comm.) the species is infrequently encountered.
Habitat and Ecology It occurs in dry lowland valleys, in both mesic and xeric areas, retreating under rocks and logs.
Males call from rain-flooded areas, and eggs are laid in still water. The species breeds year round in some areas. It
is occasionally found near water with heavy concentrations of sulphur.
Major Threats The major threats to this lowland species are habitat loss, due to both livestock grazing and selective
logging, and agricultural pollution.
© Alejandro Sanchez

Conservation Measures It occurs in several protected areas.

Bibliography: Hedges, S.B. (1993), Hedges, S.B. (1999), Hedges, S.B. (2001), Henderson, R.W. and Powell, R (1999), Henderson, R.W.
and Powell, R. (2001), Powell, R. (1992), Schwartz, A. (1972), Schwartz, A. and Henderson, R.W. (1991)
Data Providers: Blair Hedges, Sixto Inchaustegui, Marcelino Hernandez, Robert Powell

VU Bufo gundlachi Ruibal, 1959 CUBAN HIGH-CRESTED TOAD

Vulnerable B2ab(iii) Geographic Range This species is widespread but patchily dis- Bibliography: Centro Nacional de Areas Protegidas (CNAP) (2002), Hedges, S.B. (1993), Hedges, S.B. (1999), Hedges, S.B. (2001),
Order, Family: Anura, Bufonidae tributed in Cuba, including the Isla de Juventud, at low elevations Henderson, R.W. and Powell, R (1999), Henderson, R.W. and Powell, R. (2001), Schwartz, A. and Henderson, R.W. (1991)
Country Distribution: Cuba (sea level up to 70m asl). Data Providers: Blair Hedges, Luis Díaz
Current Population Trend: Decreasing Population It is abundant during the breeding season, but otherwise
is rarely seen since it burrows underground.
Habitat and Ecology Mainly a forest species, but it also occurs in
xeric and mesic grasslands, and diurnally retreats in burrows near
rocks. Males call while floating in rain-flooded pools and the eggs
are laid in still water. This species is an explosive breeder.
Major Threats The major threat to this species is habitat loss
and degradation, due to agriculture (it occurs in an area of intense
agricultural production), and pollution from agricultural pesticides.
The spread of the invasive plant species Dichrostachys cinerea is
also a threat to the species’ native habitat.
Conservation Measures It occurs in several protected areas.
There is a need to control the spread of invasive species within
existing protected areas.
 Globally Threatened Amphibian Species
CR Bufo holdridgei Taylor, 1952
Critically Endangered A2ace; B1ab(v)
Order, Family: Anura, Bufonidae
Country Distribution: Costa Rica
Current Population Trend: Decreasing
© Wayne Van Devender
EN Bufo houstonensis Sanders, 1953
Endangered C2a(ii)
Order, Family: Anura, Bufonidae
Country Distribution: United States of America
Current Population Trend: Decreasing
© Danté Fenolio/www.anotheca.com
Geographic Range Historically, this species ranged across the central coastal region of Texas. Houston toads
disappeared from the Houston area (Harris, Fort Bend and Liberty counties) during the 1960s following an extended
drought and the rapid urban expansion of the city of Houston. Although this species has been found in nine additional
counties (Austin, Bastrop, Burleson, Colorado, Lavaca, Lee, Leon, Milam, Robertson) as recently as the 1990s, several
of these populations have not been seen since they were first discovered. Of the few remaining populations, the
largest is in Bastrop County.
Population At least 2000 adults occur in Bastrop County; unknown numbers probably persist in seven other counties
(http://www.tpwd.state.tx.us/nature/endang/htoad.htm; Seal 1994). Recent trend analyses suggest that Houston toads
are declining in Bastrop State Park, which lies near the centre of its critical habitat in Bastrop County.
Habitat and Ecology Houston toads are restricted to areas with sandy, friable soil such as loblolly pine forest,
mixed deciduous forest, post oak savannah, and coastal prairie. Breeding may occur from late January to late June,
but usually earlier than May, in rain pools, flooded fields, roadside ditches, and natural or man-made ponds. Optimal
habitats are non-flowing, fishless pools that persist for at least 60 days (long enough for larvae to metamorphose).
Houston toads are nocturnal, spending daylight hours in burrows, buried in sand, or under leaf-litter, pine duff, or
surface objects.
Major Threats Habitat conversion poses the most serious threat to the Houston toad. Several populations were
eliminated with the expansion of Houston, and the largest remaining population in Bastrop County is also under
intense and immediate threat from urbanization and recreational over-development. Many Houston toads are killed
each year by automobiles. Roadway mortality will increase as human populations continue to increase within the
species’ habitat and as the habitat continues to be dissected by more roads. Road construction further isolates
populations and disrupts or prevents the movement of individual toads between populations. This movement of
toads is necessary to maintain gene flow, and thus genetic diversity, and to supplement small or declining local
populations. It is possible to build roads with underpasses or other structures that allow toads and other wildlife to
pass safely beneath the roads. While converting woodlands to pastures or ploughed fields destroys Houston toad
EN Bufo ibarrai Stuart, 1954
Endangered B1ab(iii)
Order, Family: Anura, Bufonidae
Country Distribution: Guatemala, Honduras
Current Population Trend: Decreasing
© Twan Leenders
185
Geographic Range This species occurs on the Volcán Barva, Cordillera Central, Costa Rica, at 200-2,200m asl
(Savage 2002).
Population It was formerly common in appropriate habitat and during the breeding season at the onset of the rainy
season (2,765 males were seen visiting two pools in an eight-day period; Nowak and Robinson 1975). It has not been
seen since 1986 despite repeated searching throughout its range, and it is possible that it is now extinct.
Habitat and Ecology It lives in lower montane rainforest. It is a fossorial species, and may be found under surface
debris within the forest during periods of heavy rain, but otherwise concentrates on mossy stream banks during dry
periods. It is an explosive breeder that lays its eggs in forest floor pools, as well as in man-made drainage ditches
(Savage 2002).
Major Threats Although not proven, the main cause of the population decline is likely to be chytridiomycosis perhaps
in synergy with the effects of climate change.
Conservation Measures The entire known range of this species is protected in Parque Nacional Braulio Carrillo,
20km north-east of the capital of San José. Surveys are urgently required to determine whether or not the species
survives and, in view of the threat of chytridiomycosis, surviving individuals might need to form the basis for the
establishment of an ex-situ population.
Bibliography: Novak, R.M. and Robinson, D.C. (1975), Savage, J.M. (2002)
Data Providers: Federico Bolaños, Gerardo Chaves
HOUSTON TOAD
habitat and favours the proliferation of other toad species, certain agricultural practices can be beneficial to Houston
toads. These include maintaining low to moderate numbers of livestock to avoid overgrazing, protecting pond habitat
from livestock and predatory fish, planting native bunchgrasses instead of sod-forming grasses such as Bermuda
grass (which are difficult for the toads to move through), and conserving large blocks of woodlands. Certain forestry
practices may benefit the Houston toad, while others, such as clear cutting, are harmful. Thinning and burning have
been shown to benefit some species of amphibians and reptiles by opening up the forest canopy and allowing more
sunlight to reach the forest floor. This practice encourages the growth of vegetation and, in turn, increases insect
numbers. This may be beneficial to the Houston toad. Other threats that often appear in conjunction with the factors
outlined above include prolonged drought and the presence of fire ants, an unwelcome species from Brazil. Fire ants
have been observed preying on toadlets as they leave their breeding pond. Fire ants thrive in open, sunny areas where
the soil has been disturbed and woody vegetation uprooted, as in agricultural fields and urban areas. Protecting large
forested areas is one of the most effective deterrents to fire ants. Where fire ant control with pesticides is necessary,
mounds should be treated individually, rather than broadcasting the chemicals, to avoid impacting other invertebrates
that the Houston toad eats (see http://ifw2es.fws.gov/HoustonToad).
Conservation Measures The Houston toad was the first amphibian granted protection under the U.S. Endangered
Species Act. A critical habitat was designated in 1978 in Bastrop and Burleson counties, in areas supporting the
largest populations known at that time. However, the population within critical habitat in Burleson County has not
been seen since 1983. In the 1970s, the state of Texas acquired land within designated critical habitat in Bastrop
County adjacent to Buescher and Bastrop state parks to aid in conservation. Additionally, an effort was started in
1978 by the Houston Zoo to identify remaining Houston toad populations and supplement them or establish new
populations in protected areas using wild-caught adults, naturally deposited eggs, or captive-reared juveniles and
adults. However, new populations were not established in spite of introducing over 500,000 individuals (adults,
juveniles, larvae) into sites at the Attwater Prairie Chicken National Wildlife Refuge. Research is urgently needed
to determine the status of Houston toad populations outside Bastrop County and promote conservation efforts in
these areas. Research is also critical to determine which management practices are most conducive to the Houston
toad and the ecosystem on which it depends. The Houston Toad Recovery Plan was published by the U.S. Fish and
Wildlife Service (USFWS) in 1984. The Texas Parks and Wildlife Department and the USFWS have jointly prepared
a brochure for private landowners who wish to implement their agricultural practices in ways that are compatible
with the needs of the Houston toad and the Texas Forest Service has formed a committee to develop management
practices that protect the Houston toad and its habitat. Additionally, the USFWS is working with community leaders,
private landowners, and conservation organizations to develop and implement a regional Habitat Conservation Plan
for Bastrop County, which would provide for the issuance of endangered species permits that allow development to
proceed while ensuring permanent habitat protection. The USFWS also has established a fund with the National Fish
and Wildlife Foundation to assist in local habitat protection efforts for the Houston toad.
Bibliography: Bartlett, R.D. and Bartlett, P.P. (1999), Blackburn, L., Nanjappa, P. and Lannoo, M.J. (2001), Brown, L.E. (1971), Brown,
L.E. (1973), Brown, L.E. et al. (1984), Bury, R.B., Dodd, Jr., C.K. and Fellers, G.M. (1980), Campbell, L. (1995), Conant, R. and Collins, J.T.
(1991), Dixon, J.R. (2000), Dodd, Jr, C.K. and Seigel, R.A. (1991), Frost, D.R. (1985), Gaston, M.A., Forstner, M.R.J. and Dixon, J.R. (2001),
Hillis, D.M., Hillis, A.M. and Martin, R.F. (1984), Jacobson, N.L. (1989), Kennedy, J.P. (1961), Matthews, J.R. and Moseley, C.J. (eds)
(1990), Quinn, H.R. and Mengden, G. (1984), Seal, U.S. (1994), U.S. Fish and Wildlife Service, U.S. Fish and Wildlife Service (1980b),
U.S. Fish and Wildlife Service (1990a)
Data Providers: Geoffrey Hammerson, Donald Shepard
Geographic Range This species occurs at moderate and intermediate elevations of central and southern Guatemala
and the Sierra de las Minas, eastern Guatemala, from 1,360-1,980m asl. Its range has also recently been extended
into contiguous regions of Honduras in the western ranges of the Southern Cordillera Region in the departments of
Intibuca, Lemipira and Ocotepeque, up to 2,020m asl (Mendelson et al. 2005). However, McCranie and Wilson (2002)
refer all Honduran material to Bufo coccifer.
Population This species persists in small numbers in appropriate habitat. Nearly continuous field work throughout
the range of this species in Guatemala between 1989 and 1998 produced less than 10 observations.
Habitat and Ecology It occurs in pine-oak or premontane or lower montane moist forest, and breeds in ponds,
marshes and wet meadows. Males have been observed calling from the banks of ponds.
Major Threats Its range is severely fragmented due to habitat loss, primarily due to agricultural activities. Formerly
robust populations in Guatemala are now extirpated following severe alteration of montane wetlands and cloud
forests, and may well have succumbed to the effects of chytrid, which has been documented in other anuran species
in the same area (Mendelson et al. 2004).
Conservation Measures A portion of the range previously occurred in the Reserva de la Biósfera Sierra de las Minas
in Guatemala, but it has now been extirpated from this region. Protection of the remaining habitat of this species is
urgently required. Given the threat of chytrid, this species requires close population monitoring.
Bibliography: Campbell, J.A. (2001), McCranie, J.R. and Wilson, L.D. (2002b), Mendelson III, J.R. (2001), Mendelson III, J.R. et al.
(2004), Mendelson III, J.R. et al. (2005)
Data Providers: Manuel Acevedo, Bruce Young
186 Threatened Amphibians of the World

EN Bufo inyangae Poynton, 1963 INYANGA TOAD

Endangered B1ab(iii) Geographic Range This species in known only from the Nyanga Mountains in eastern Zimbabwe. It presumably
Order, Family: Anura, Bufonidae occurs in adjacent Mozambique, but there have been no records here, probably because of a lack of herpetological
Country Distribution: Zimbabwe work in this area. It has been found at 2,400-2,560m asl, but probably also occurs at slightly lower elevations.
Current Population Trend: Decreasing Population It is rather cryptic and hard to find, but it seems to be fairly common within its very limited range.
Habitat and Ecology It inhabits montane grassland with exposed bare granite. The animals hide under stones, in
cracks in the granite, and in rodent burrows. It breeds in temporary pools, and larvae have been seen moving across
wet granite faces.
Major Threats The high-altitude habitat of this species has remained relatively intact up until now, although it might
be at increasing risk from wood plantations, overgrazing by livestock, and human settlement.
Conservation Measures It occurs in the Nyanga National Park; continued and strengthened mangement of this
protected area is needed.
© Richard Boycott

Bibliography: Channing, A. (2001), Lambiris, A.J.L. (1989b), Poynton, J.C. (1963), Poynton, J.C. (1964b), Poynton, J.C. and Broadley,
D.G. (1988), Tandy, M. and Keith, R. (1972)
Data Providers: John Poynton, Mills Tandy

VU Bufo justinianoi Harvey and Smith, 1994

Vulnerable B1ab(iii) Geographic Range This species is found in the Bolivian Andes, in Cochabamba, Santa Cruz and La Paz Departments.
Order, Family: Anura, Bufonidae It is know from the type locality, El Chape, in Florida Province, Santa Cruz Department, at an altitude of 2,050m asl,
Country Distribution: Bolivia and from Campamento Fortaleza, in Carrasco Province, Cochabamba Department, from 1,875-2,220m asl, and from
Current Population Trend: Decreasing four localities in Nor Yungas Province, in La Paz Department, from 1,440-2,250m asl.
Population The population status of this species is not known.
Habitat and Ecology It inhabits wet montane forest including cloud and Yungas forest. Köhler (2000a) suggests
that it is probably restricted to primary forest. The larvae develop in streams.
Major Threats The major threat to this species is habitat destruction and degradation caused primarily by agri-
culture.
Conservation Measures Its range includes Parque Nacional Carrasco and Parque Nacional Amboro.
Bibliography: De la Riva, I. et al. (2000), Harvey, M. and Smith, E. (1994), Köhler, J. (2000a)
Data Providers: Claudia Cortez, Steffen Reichle, Ignacio De la Riva, Jörn Köhler
© Jörn Köhler

EN Bufo kotagamai Feonando and Dayawansa, 1994

Endangered B1ab(iii)+2ab(iii) Geographic Range This species is restricted to three sites in south-western Sri Lanka (Kitulgala Forest Reserve,
Order, Family: Anura, Bufonidae Messana Forest Reserve and Sinharaja World Heritage Site). It ranges from 150-1,070m asl.
Country Distribution: Sri Lanka Population Little is known about the population status of this species, but it is not common.
Current Population Trend: Decreasing Habitat and Ecology A terrestrial species associated with wet leaf-litter, rocks and other ground cover, close to
streams in tropical wet montane forest, its breeding biology has not been recorded, but it presumably takes place in
water, probably in streams, by larval development.
Major Threats Habitat loss (small-scale agriculture) and human disturbance (local tourism) are the primary threats
to this species.
© Madhava Meegaskumbura

Conservation Measures It has been recorded from several protected areas, including the Kitulgala Forest Reserve,
Messana Forest Reserve and the Sinharaja World Heritage Site.
Bibliography: Dubois, A. and Ohler, A. (1999), Dutta, S.K. and Manamendra-Arachchi, K. (1996), Manamendra-Arachchi, K. and
Pethiyagoda, R. (1998)
Data Providers: Kelum Manamendra-Arachchi, Anslem de Silva, Deepthi Wickramasinghe

EN Bufo koynayensis Soman, 1963

Endangered B1ab(iii) Geographic Range This species is known only from two localities (Koyna and Aboli) in the Western Ghats of
Order, Family: Anura, Bufonidae Maharashtra, India. Specimens have been recorded at elevations between 900 and 1,200m asl.
Country Distribution: India Population It is a rare species, and is uncommon at the type locality.
Current Population Trend: Decreasing Habitat and Ecology This is a terrestrial toad of moist to wet evergreen forest, and dry riparian grassland. Its breeding
has not been recorded, but it presumably takes place in water, probably in streams, by larval development.
Major Threats The main threat to this species is habitat loss due to agriculture and clear cutting of forests.
Conservation Measures It has been recorded from the Konya Wild Life Sanctuary, but expanded protection of
forest habitats in the species’ range is needed. Further research is needed into the range and breeding biology of
this species.
Notes on taxonomy: The taxonomy and relationships of this species need re-evaluation (Biju 2001).
Bibliography: Biju, S.D. (2001), Dutta, S.K. (1997), Giri, V. and Chaturvedi, N. (2001), Grandison, A.G.C. and Daniel, J.C. (1964), Soman,
P.W. (1963)
© K. Jayaram

Data Providers: S.D. Biju, Sushil Dutta, Robert Inger

 Globally Threatened Amphibian Species 187

EN Bufo kumquat Das and Lim, 2001

Endangered B1ab(iii) Geographic Range This newly described species was found on the west coast of Peninsular Malaysia. It is known
Order, Family: Anura, Bufonidae only from the type locality, Sabak Bernam, and from the North Selangor Peat Swamp, though it possibly occurs
Country Distribution: Malaysia more widely.
Current Population Trend: Decreasing Population It is locally abundant, but it is possibly hard to find when not breeding (which may be one explanation
for its recent discovery).
Habitat and Ecology It is a peat swamp specialist. It is thought be be an explosive breeder, and presumably breeds
by larval development.
Major Threats The major threat is drainage and reclamation of peat swamps for agricultural purposes, and part of
its type locality has already been destroyed.
Conservation Measures This species has not been recorded from any protected areas, making habitat protection
an urgent priority.
© Jeet Sukumaran

Bibliography: Das, I. and Lim, K.P.P. (2001)

Data Providers: Indraneil Das, Jeet Sukumaran, Norsham Yaakob

CR Bufo lemur (Cope, 1868) PUERTO RICAN CRESTED TOAD

Critically Endangered A2a; B1ab(v)+2ab(v); Geographic Range This species has a very restricted range, and is known from only a handful of localities along
C2a(ii) the north and south coasts of Puerto Rico and Virgin Gorda Island. In recent years it has been recorded from only one
Order, Family: Anura, Bufonidae location on the south coast of Puerto Rico. It has been recorded from sea level up to 50m asl.
Country Distribution: Virgin Islands (British), Population The north coast population has not been recorded since 1992, and it is most likely extirpated in this area. It
Puerto Rico was last recorded on Virgin Gorda Island in 1964, and other surveys since then have not located any individuals. Since
Current Population Trend: Decreasing 1992, there has only been one known population remaining. In 1984, there were 900 mature individuals recorded in
this population; subsequently, in 1998, there were only 215 mature individuals recorded (of which 34 were females),
in 2002 only 100 mature individuals, and in 2003 only 80 mature individuals were recorded (R. Joglar pers. comm.).
Habitat and Ecology It is a terrestrial species found in semi-arid, rocky areas of seasonal evergreen forest. Eggs
are laid in permanent or temporary pools of water, streams, or small dams for livestock.
Major Threats Infrastructure development for human settlement is a major threat, particularly on the north coast. In the
south of its range some breeding pools were deliberately drained to clear parking space for visitors to the beach.
© Alejandro Sanchez

Conservation Measures The last known population occurs entirely within the Guanica National Forest, which is a
well-managed area. Captive breeding has been successful, and after many years a re-introduction program in Puerto
Rico seems to be showing some success, with re-introduced captive-bred animals now returning to the constructed
ponds where they were first released (Zippel 2005).
Bibliography: Hedges, S.B. (1993), Henderson, R.W. and Powell, R (1999), Henderson, R.W. and Powell, R. (2001), Joglar, R.L. (1999),
Rivero, J.A. (1998), Rivero, J.A. and Seguí Crespo, D. (1992), Zippel, K. (2005)
Data Providers: Blair Hedges, Rafael Joglar, Richard Thomas, Luis J. Villanueva-Rivera, Neftalí Ríos-Lopez

EN Bufo leucomyos McCranie and Wilson, 2000

Endangered B1ab(iii) Geographic Range Disjunct populations of this species occur on the Atlantic slope of north-central Honduras at
Order, Family: Anura, Bufonidae 0-1,600m asl.
Country Distribution: Honduras Population The population is apparently stable in appropriate habitat and the species is recorded on a regular
Current Population Trend: Decreasing basis.
Habitat and Ecology It can be found in lowland moist, and premontane and lower montane wet, forest; all specimens
have been collected from broadleaf forests. The larvae apparently develop in streams.
Major Threats A major threat to this species is the impact of landslides on upper clear water streams, as a result of
strong storms and human activities taking place in the lower portions of the streams. Habitat loss due to slash-and-burn
agriculture, smallholder farming, logging, and expanding human settlements is also a significant threat.
Conservation Measures Part of the range of this species is protected in La Muralla and Pico Bonito National Parks,
the Lancetilla Botanical Garden, and the Reserva de Visda Silvestre Texiguat.
© Twan Leenders

Bibliography: McCranie, J.R. and Wilson, L.D. (2000a), McCranie, J.R. and Wilson, L.D. (2002b)
Data Providers: Larry David Wilson, Gustavo Cruz

EN Bufo longinasus Stejneger, 1905 CUBAN LONG-NOSED TOAD

Endangered B2ab(iii) Geographic Range This species is only known from three very small Conservation Measures Its range includes several protected areas, although these areas do not provide sufficient
Order, Family: Anura, Bufonidae distributional pockets (Pinar del Río Province, Sierra de Trinidad, protection for the species. Improved management of these reserves, and strengthened protection of the remaining
Country Distribution: Cuba and Sierra del Guaso) in Cuba, and each of these may turn out to be forest habitat, is required.
Current Population Trend: Decreasing separate species. It has been recorded from 100-820m asl. Bibliography: Centro Nacional de Areas Protegidas (CNAP) (2002), Hedges, S.B. (1993), Hedges, S.B. (1999), Hedges, S.B. (2001),
Population It is generally an uncommon species, although one Henderson, R.W. and Powell, R (1999), Henderson, R.W. and Powell, R. (2001), Moreno, L.V. et al. (1999), Schwartz, A. and Henderson,
population in the west is known to be very abundant. However, R.W. (1991)
the eastern population has not been found again since it was first Data Providers: Blair Hedges, Luis Díaz
described in the early 1900s.
Habitat and Ecology It is found in upland pinewoods and mesic
broadleaf forest, and is terrestrial by day but may be arboreal at night.
It has not been recorded outside forest habitat, and is always found
by streams. Males call while floating on the surface of water. Eggs
are laid in streams, where the larvae also develop.
Major Threats There is ongoing habitat loss in the upland wood-
lands in which it occurs from clear-cutting and subsistence logging,
charcoaling, fires, and agricultural expansion.
188 Threatened Amphibians of the World

VU Bufo macrocristatus Firschein and Smith, 1957

Vulnerable B1ab(iii)
Order, Family: Anura, Bufonidae
Country Distribution: Guatemala, Mexico
Current Population Trend: Decreasing
Geographic Range This species’ geographic range extends from Los Bibliography: Campbell, J.A. (2001), Mendelson III, J.R. (1997b)
Chimalapas region in south-central Oaxaca, Mexico, south and east Data Providers: Georgina Santos-Barrera, Manuel Acevedo
along the Atlantic versant as far as the Sierra de los Cuchumatanes
in Guatemala. It occurs at 1,000-1,600m asl.
Population It is rare across its range.
Habitat and Ecology This species is mainly associated with high
humid areas in cloud forest and, in Mexico, pine-oak-Liquidambar
forests. It breeds in streams.
Major Threats The major threats to this species are habitat loss, due
to agriculture and human settlement, and water pollution.
Conservation Measures The conservation, expansion and man-
agement of cloud forest are key measures to preserve this species
since it has not been recorded from any of the Biosphere reserves
in Chiapas, Mexico. It also does not occur in any protected area
in Guatemala.

VU Bufo microtympanum Boulenger, 1882

Vulnerable B1ab(iii)+2ab(iii) Geographic Range This is a poorly known species that is presumed Notes on taxonomy: Several specimens bearing this name, and which are deposited in different museums in India, do not belong to
Order, Family: Anura, Bufonidae to be endemic to the southern Western Ghats of India. A record from this species (Biju 2001).
Country Distribution: India Maharashtra (Yazdani and Mahabal 1976) requires confirmation. Ma- Bibliography: Biju, S.D. (2001), Chanda, S.K. (2002), Dubois, A. and Ohler, A. (1999), Dutta, S.K. (1997), Manamendra-Arachchi, K. and
Current Population Trend: Decreasing namendra-Arachchi and Pethiyagoda (1998) clarified that it does not Pethiyagoda, R. (1998), Vasudevan, K., Kumar, A. and Chellam, R. (2001), Yazdani, G.M. and Mahabal, A. (1976)
occur in Sri Lanka. The altitudinal range is from 1,400-2,100m asl. Data Providers: S.D. Biju, Sushil Dutta, Karthikeyan Vasudevan, S.P. Vijayakumar, Chelmala Srinivasulu, Gajanan Dasaramji Bhuddhe
Population It is a rare species.
Habitat and Ecology A terrestrial species associated with montane
‘shola’ (grassland-forest mosaic) and tropical moist forest habitat; it
has not been recorded from modified habitats. It may be found in the
vicinity of water bodies, but the breeding habitat is not known.
Major Threats The major threat to this species is habitat loss due to
the conversion of forested areas to cultivated land (including timber
and non-timber plantations), future dam development projects, road
construction, and urbanization.
Conservation Measures It has been recorded from the Kalakkad
Mundauthurai Tiger Reserve (Tamil Nadu) and Silent Valley National
Park (Kerala).

EN Bufo nelsoni Stejneger, 1893 AMARGOSA TOAD

Endangered C2a(i) 1 March 1996). Surveys at 20 sites since 1990 yielded the following results: apparently extirpated from one spring,
Order, Family: Anura, Bufonidae decreased abundance at four springs, fluctuating but relatively constant populations at 15 sites (USFWS, Federal
Country Distribution: United States of America Register, 1 March 1996; see also unpublished 1993 and 1994 reports by Hoff, prepared for the USFWS, Reno, Nevada).
Current Population Trend: Decreasing Over the past few decades, the species reportedly has declined greatly from its former range and abundance (Altig
and Dodd 1987), but more recent surveys found that distribution and abundance were greater than previously known
(USFWS, Federal Register, 1 March 1996).
Habitat and Ecology Usually found near water at desert springs and outflow. Vegetation bordering water consists
of cottonwood trees, cattails, and sedges. May congregate at streetlights to feed on attracted insects (Burroughs
1999). Eggs and larvae develop in spring waters (open areas with little vegetation at LaFleur).
Major Threats Factors that may be adversely affecting the toad and its habitat include the effects of variable rainfall
on small populations, livestock and feral burro grazing and trampling, off-road vehicle use, grading for flood control,
activities related to commercial development, non-native predators (catfish, crayfish, bullfrog (Rana catesbeiana)),
© Kenny Wray

Geographic Range This species occurs in riparian habitats associated with the Amargosa River, tributary springs
of the Amargosa River in Oasis Valley, and isolated spring systems near Beatty, Nye County, Nevada, USA (USFWS,
Federal Register, 1 March 1996).
Population The total adult population size is uncertain but is likely to be at least several hundred. Thousands were
reported in 1958; estimates of the size of the metamorphosed population at 10 sites in 1993 and 1994 ranged from
30-130, but some sites that were probably occupied have not been surveyed in recent years (USFWS, Federal Register,
water pollution, and water diversion (Froglog, December 1994). Trampling of larvae by cattle may be negatively af-
fecting the LaFleur population, which also may be threatened by road widening (of Route 95). Expansion of non-native
salt cedar may degrade habitat (Burroughs 1999).
Conservation Measures The majority of available habitat is on private property. BLM has initiated protection (Area
of Critical Environmental Concern) for all occupied sites under their jurisdiction. TNC purchased a 60-ha ranch near
Beatty for experimental habitat management (Burroughs 1999). Nye County is a co-operator with state and federal
agencies in a conservation agreement (Burroughs 1999).
Bibliography: Altig, R. and Dodd, C.K. Jr. (1987), Blackburn, L., Nanjappa, P. and Lannoo, M.J. (2001), Burroughs, M. (1999), Bury,
R.B., Dodd, Jr., C.K. and Fellers, G.M. (1980), Feder, J.H. (1977), Stebbins, R.C. (1951), Stebbins, R.C. (1985b), U.S. Fish and Wildlife
Service (1989a)
Data Providers: Geoffrey Hammerson

EN Bufo nesiotes Duellman and Toft, 1979

Endangered B1ab(iii) Geographic Range This species is known only from the vicinity of
Order, Family: Anura, Bufonidae the type locality of Laguna, west slope of Serrania de Sira, Depar-
Country Distribution: Peru tamento Huánuco, Peru. The type specimen was recorded at 1,280m
Current Population Trend: Decreasing asl, but the species may well occur from 600 to around 2,000m asl
(as another specimen is believed to have been found at 600m asl). It
presumably ranges much more widely than has been mapped, but it
is nonetheless likely to be endemic to the Serrania de Sira.
Population There is no information on the population status of
this species.
Habitat and Ecology Their habitat is premontane and montane
forest on an isolated mountain ridge of the upper Amazon Basin.
Nothing is known about their breeding strategy, although it has
been suggested that this may be a species that breeds by direct
development. It is not known to what degree they can withstand
any anthropogenic disturbance.
Major Threats Although the restricted area that the species inhabits is relatively inaccessible, there is some
disturbance due to farming activity and human settlements. In addition, the lower parts of the range are being
selectively logged.
Conservation Measures This species is quite likely to occur in Reserva Comunal El Sira. Continued maintenance
of this, and other remaining habitat in the Serrania de Sira, is necessary.
Notes on taxonomy: Duellman and Toft (1979) described this species from only one specimen (a female), and suggested this species
could have direct development.
Bibliography: Duellman, W.E. and Toft, C.A. (1979), Rodríguez, L.O., Cordova, J.H. and Icochea, J. (1993)
Data Providers: Ariadne Angulo, Karl-Heinz Jungfer, Javier Icochea
 Globally Threatened Amphibian Species 189

EN Bufo noellerti Manamendra-Arachchi and Pethiyagoda, 1998

Endangered B1ab(iii) Geographic Range This species has a restricted distribution in south-west Sri Lanka, and has been recorded at
Order, Family: Anura, Bufonidae elevations of between 150 and 460m asl.
Country Distribution: Sri Lanka Population It is not a common species, and populations may be declining proportionately with a loss in forest
Current Population Trend: Decreasing habitat.
Habitat and Ecology It is a terrestrial species found in, and around, tropical lowland moist forest. Adults have been
recorded from rubber plantations, tea estates, and domestic gardens close to the forest edge only. The breeding
habitat is not known.
Major Threats The main threats to the species are habitat loss, due to selective logging, and agrochemical pol-

© Madhava Meegaskumbura
lution.
Conservation Measures It has been recorded from the Kanneliya Forest Reserve, Gilimale-Eratne Forest Reserve and
the Sinharaja World Heritage Site; the continued management of these for biodiversity conservation is important.
Bibliography: Dubois, A. and Ohler, A. (1999), Dutta, S.K. and Manamendra-Arachchi, K. (1996), Manamendra-Arachchi, K. and
Pethiyagoda, R. (1998)
Data Providers: Kelum Manamendra-Arachchi, Anslem de Silva

VU Bufo nyikae Loveridge, 1953

Vulnerable D2 Geographic Range This species is known only from the Nyika Plateau in northern Malawi and north-eastern Zambia.
Order, Family: Anura, Bufonidae It is a high-altitude species, although its precise altitudinal range is not known.
Country Distribution: Malawi, Zambia Population It is probably reasonably common within its small range.
Current Population Trend: Stable Habitat and Ecology It is apparently associated mainly with montane forest, and with wet, boggy places in grassland,
and is never far from trees. It breeds in small, shallow pools.
Major Threats Most, if not all, of the range of this species is in a protected area, and so it is probably not seriously
threatened at present.
Conservation Measures It occurs in the Nyika National Park. This species requires close population monintoring
given that it is known from only a single location.
Notes on taxonomy: This species was removed from the synonymy of Bufo lonnbergi by Poynton (1997), where it had been placed
© David C. Blackburn

by Grandison (1972).
Bibliography: Broadley, D.G. (1971), Channing, A. (2001), Grandison, A.G.C. (1972b), Poynton, J.C. (1964a), Poynton, J.C. (1997), Poynton,
J.C. and Broadley, D.G. (1988), Stewart, M.M. (1967), Stewart, M.M. and Wilson, V.J. (1966), Tandy, M. and Keith, R. (1972)
Data Providers: Mills Tandy, Lovemore Mazibuko, Alan Channing, John Poynton

EN Bufo pantherinus Smith, 1828 WESTERN LEOPARD TOAD

Endangered B1ab(ii,iii,iv,v)+2ab(ii,iii,iv,v) Geographic Range This species is known only from a very small area of the Western Cape Province in South
Order, Family: Anura, Bufonidae Africa, ranging from the Cape Peninsula eastward to beyond Gansbaai in the Pearly Beach area. It occurs only at low
Country Distribution: South Africa elevations, within 10km of the sea.
Current Population Trend: Decreasing Population It is a locally common species.
Habitat and Ecology It occurs in large wetlands, vleis, dams, and sluggish water in lowland fynbos heathland, as
well as in altered habitats, such as farmland, suburban gardens, and urban open areas, although always in close
proximity to freshwater habitats. It breeds in permanent waterbodies, and occasionally temporary waterbodies that
retain water well into summer, and has a preference for deep water, with floating plants.
Major Threats Although it is tolerant of a degree of habitat alteration, it is probably being negatively impacted by
increased urbanization and agricultural expansion. Road kills may also be an important factor.
Conservation Measures It occurs in the Agulhas National Park and in the northern limits of Cape of Good Hope
© Andrew Turner

Nature Reserve, as well as in various local reserves, However, much of its habitat remains unprotected, such that
improved protection of this habitat is necessary.
Bibliography: Channing, A. (2001), Cherry, M.I. (1992), Cunningham, M. and Cherry, M.I. (2000), Eick, B.N., Harley, E.H. and Cherry,
M.I. (2001), Minter, L.R. et al. (2004), Passmore, N.I. and Carruthers, V.C. (1995), Poynton, J.C. and Lambiris, A.J.L. (1988), Tandy, M.
and Keith, R. (1972)
Data Providers: Leslie Minter, James Harrison

CR Bufo peripatetes Savage, 1972

Critically Endangered A3ce Geographic Range This species is known only from two sites: iomycosis, and are now possibly extinct. This disease must be considered a very serious threat to B. peripatetes as
Order, Family: Anura, Bufonidae Parque Internacional La Amistad, a transboundary protected area be- well, and the disease has already passed through the range of this species. Destruction of available forest habitat
Country Distribution: Panama tween Costa Rica and Panama, and Cerro Bollo, in the western central (generally caused by the driving of cattle through La Amistad) is a threat to the species.
Current Population Trend: Decreasing cordillera of Panama. It has been recorded at 1,500-1,856m asl. Conservation Measures There are no specific conservation measures currently in place, but further research is
Population There is no information available on the population size needed into the range, ecology, and population status of this species. In view of the threat of chytridiomycosis, the
or abundance of this species. It can still be found above Boquete on status of this species should be closely monitored, and an ex-situ population should be established.
the trail to Almirante and Cerro Bollo on the border between Chiriquí Bibliography: Ibáñez, R. et al. (2000), Savage, J.M. (1972b), Savage, J.M. and Donnelly, M.A. (1992), Young, B. et al. (1999)
and Bocas del Toro Provinces. Data Providers: Frank Solís, Roberto Ibáñez, César Jaramillo, Querube Fuenmayor, Karen Lips
Habitat and Ecology This is a largely unknown terrestrial spe-
cies of tropical montane forest. There is no information available
on reproduction or breeding habitats, though it presumably takes
place in water. Assuming that it is similar to Bufo fastidiosus and
B. holdridgei, it is likely to emerge unpredictably to breed, and is
probably very hard to find.
Major Threats The closely related species Bufo fastidiosus and B.
holdridgei have both declined dramatically, probably due to chytrid-
190
EN Bufo perplexus Taylor, 1943
Endangered B1ab(iii)
Order, Family: Anura, Bufonidae
Country Distribution: Mexico
Current Population Trend: Decreasing
VU Bufo perreti Schiøtz, 1963
Vulnerable D2
Order, Family: Anura, Bufonidae
Country Distribution: Nigeria
Current Population Trend: Unknown
© Arne Schiøtz
VU Bufo quechua Gallardo, 1961
Vulnerable B1ab(iii)
Order, Family: Anura, Bufonidae
Country Distribution: Bolivia
Current Population Trend: Decreasing
© Arturo Muñoz
VU Bufo rubropunctatus Guichenot, 1848
Vulnerable A2c
Order, Family: Anura, Bufonidae
Country Distribution: Argentina, Chile
Current Population Trend: Decreasing
© Andrés Morya
Threatened Amphibians of the World
Geographic Range This species is found in the Tepalcatepec Basin
and surrounding areas in south-western Michoacán, south of the
Balsas River in Guerrero, Mexico. It probably occurs more widely
than currently known.
Population This is a rare species.
Habitat and Ecology It inhabits the Pacific lowlands in areas of
seasonal tropical forest, and lives under rocks and logs close to
streams, breeding in pools.
Major Threats The main threats to this species are infrastructure
development and agricultural expansion, coupled with changes in
the management of agricultural habitat.
Conservation Measures It is not found within any protected
area, and protection of the species’ original forest habitat is
urgently needed.
Bibliography: Blair, W.F. (1972), Duellman, W.E. (1961)
Data Providers: Georgina Santos-Barrera, Luis Canseco-Márquez
Geographic Range This species has been recorded only from the Idanre Hills, in Ondo Province, south-western
Nigeria. It is very likely to occur in other similar habitats in south-western Nigeria. There have been no recent records
of the species, presumably because of a lack of herpetological work within its range.
Population It is reported to be very common within its small known range, and tadpoles are abundant during the
wet season.
Habitat and Ecology It is closely associated with gneiss domes or inselbergs in the forest zone, though is absent
from similar inselbergs in the savannah belt. It lives terrestrially in patches of shrubby vegetation and forest on the
gneiss domes. The eggs are probably laid on soil, and the tadpoles then disperse by crawling out on the wet, sloping
(often almost vertical) rock.
Major Threats Its rocky habitat is largely inaccessible and useless to humans, so this species is probably not facing
any serious threats at present. However, it is intrinsically at risk because of its restricted range.
Conservation Measures It is not known from any protected areas, and there is a need for improved protection
of sites where this species is known to occur. There is also a need for population monitoring given that it is known
from only a single location.
Bibliography: Schiøtz, A. (1963), Tandy, M. and Keith, R. (1972)
Data Providers: Arne Schiøtz, Mills Tandy
Geographic Range This species is known from the eastern slopes of the Bolivian Andes, in Cochabamba and Santa
Cruz Departments. It has also been recorded in Chapare Province, Cochabamba Department and Caballero Province,
Santa Cruz Department in Bolivia (Köhler 2000a; Cortez 2001). It has been recorded from 1,900-2,300m asl.
Population The population status of this species is not known.
Habitat and Ecology A terrestrial species inhabiting wet montane forest including cloud and Yungas forest. The
eggs are laid in lotic waters (Köhler 2000a).
Major Threats Major threats to this species’ habitat include agriculture (mainly from smallholder farmers) and
agricultural pollution. Köhler (2000a) points out that it is very common to find this species infected by parasites, with
visible red pustules that, according to De la Riva (1997), are caused by the larvae of a trombidioid mite. However, it
is not known whether or not the infestations have a negative effect on the species.
Conservation Measures Its range includes Parque Nacional Carrasco and Parque Nacional Amboro.
Notes on taxonomy: Harvey and Smith (1993) pointed out that Bufo echinodes, described by Reynolds and Foster (1992), is a synonym
of B. quechua. This species is possibly a complex of more than one species.
Bibliography: Cortez, C. (2000), De la Riva, I. (1997), De la Riva, I. et al. (2000), Gallardo, J.M. (1961b), Harvey, M. and Smith, E. (1993),
Hoogmoed, M.S. (1990), Köhler, J. (2000a), Mercadal-de Barrio, I.T. and Barrio, A. (1978), Reynolds, R. and Foster, M. (1992)
Data Providers: Claudia Cortez, Steffen Reichle, Ignacio De la Riva, Jörn Köhler
Geographic Range This species is endemic to the temperate forests of Chile and Argentina. In Chile, the distribution
extends from Lanalhue Lake (37° 55’S; 73° 19’W), Arauco Province, to Palena 43°S (approximately). In Argentina, the
species is restricted to southern Río Negro Province and northern Chubut Province (including a single record from
Parque Nacional Los Alerces). It has an altitudinal range of 200-800m asl.
Population The northernmost population in Chile is declining, while the Argentinean subpopulations seem to be
stable, with successful reproduction with recruitment recently verified (Vidoz and Ubeda 2000). It is abundant during
the breeding season when courtship aggregations are formed.
Habitat and Ecology It inhabits humid to xeric forests, as well as open environments, some of them with a certain
degree of disturbance (Vidoz and Ubeda 2000). Breeding occurs in shallow temporary ponds adjacent to rivers,
reservoirs and lakes. Adults live in small holes or under bushes during the day.
Major Threats The major threat is degradation and fragmentation of habitat, due to agriculture, afforestation with
exotic species (pine), overgrazing by cattle, and human settlements. Tourism development also appears to be a threat
to the habitat of this species.
Conservation Measures The species is protected by Parque Nacional Lago Puelo and Parque Nacional Los Alerces
(a single record) in Argentina, and Parque Nacional Puyehue and Parque Nacional Vicente Pérez Rosales in Chile.
Bibliography: Cei, J.M. (1962), Cei, J.M. (1980), Formas, J.R. (1979), Formas, J.R. (1995), Formas, J.R. and Pugín, E. (1978), Gallardo,
J.M. (1962a), Gallardo, J.M. (1965), Glade, A. (1993), Lavilla, E.O. et al. (2000), Servicio Agrícola Ganadero (1998), Veloso, A. and Navarro,
J. (1988), Vidoz, F. and Ubeda, C.A. (2000)
Data Providers: Carmen Úbeda, Alberto Veloso, Herman Núñez, Néstor Basso

VU Bufo rumbolli Carrizo, 1992
Vulnerable B1ab(iii)
Order, Family: Anura, Bufonidae
Country Distribution: Argentina
Current Population Trend: Decreasing
© Arturo Muñoz
EN Bufo sclerocephalus Mijares-Urrutia and Arends, 2001
Endangered B1ab(iii,v)
Order, Family: Anura, Bufonidae
Country Distribution: Venezuela
Current Population Trend: Decreasing
VU Bufo scorteccii Balletto and Cherchi, 1970
Vulnerable D2
Order, Family: Anura, Bufonidae
Country Distribution: Yemen
Current Population Trend: Stable
EN Bufo spiculatus Mendelson, 1997
Endangered B1ab(iii)
Order, Family: Anura, Bufonidae
Country Distribution: Mexico
Current Population Trend: Decreasing
Globally Threatened Amphibian Species 191
Geographic Range This species occurs in Calilegua, Ledesma Department, Jujuy Province and Arasayal, Orán
Department, Salta Province, Argentina, from 700-1,700m asl. It is expected to occur more widely, and might be
present in Bolivia.
Population Although it is a rare species, it is nonetheless collected regularly.
Habitat and Ecology It is a terrestrial species occurring in montane forests (Yungas). Its tolerance to habitat distur-
bance is not known. It reproduces in slow-flowing streams where the tadpoles also develop.
Major Threats Selective logging and clear-cutting of primary forests are the main threat to this species, although it
is also being affected by the introduction of predatory fish (trout) and the alteration of watersheds.
Conservation Measures It is present in Parque Nacional Calilegua and Parque Nacional Baritú.
Bibliography: Carrizo, G.R. (1991), De la Riva, I. et al. (2000), Lavilla, E.O. et al. (2000), Lavilla, E.O. and Cei, J.M. (2001), Vaira, M.
(2002)
Data Providers: Esteban Lavilla
Geographic Range This species is known from the Sierra de San awareness of the local people regarding the use of this species is recommended. Further survey work is also needed
Luis, in Falcón State, Venezuela, where it has been recorded from to determine the current population status of the species.
1,150-1,500m asl. Bibliography: Barrio Amorós, C.L. (2004), Mijares-Urrutia, A. and Arends, A. (2001)
Population The most recent specimens were collected in 1992, and Data Providers: Abraham Mijares, Enrique La Marca
no additional specimens have been found since then (although this
may be due to a lack of fieldwork within its range).
Habitat and Ecology It occurs on vegetation in cloud forest. Some
individual males have been observed calling in slow-moving water,
and the larvae probably develop in water.
Major Threats The habitat of the species is under intense pressure
from agriculture and livestock farming, even within Parque Nacional
Juan Crisóstomo Falcón. The species is also used in local medicinal
use for the preparation of treatments against dermal herpes.
Conservation Measures Although it occurs in Parque Nacional
Juan Crisóstomo Falcón, this area is poorly managed for conservation
and is in urgent need of more effective management. Education and
Geographic Range This species is possibly restricted to Wadi
al Khalili near Mafhaq (1,550m asl) on a high plateau in northern
Yemen.
Population It is probably common within its restricted range.
Habitat and Ecology It is found within the wadi, and shelters in the
surrounding vegetation. Breeding takes place in pools.
Major Threats There are no current threats to this species, although
its restricted range makes it vulnerable to stochastic events.
Conservation Measures It is not known whether or not this spe-
cies occurs in protected areas, or whether or not it is protected by
national legislation. It requires close population monitoring given
that is known from only one location.
Bibliography: Balletto, E., Cherchi, M.A. and Gasperetti, J. (1985)
Data Providers: Andrew Gardner, Theodore Papenfuss, Steven Anderson,
Matthias Stöck, Sergius Kuzmin
Geographic Range This species is known from two localities in cloud forest patches in the Sierra de Juárez and the Sierra Mixe can guarantee the preservation of suitable condi-
northern Oaxaca in the Sierra de Juárez in Mexico, and another allo- tions for the survival of this species. Further survey work is necessary to determine the current population status
patric population has been recorded in the Sierra Mixe, south-central of this species.
Oaxaca, Mexico; additional survey work is likely to result in its discov- Bibliography: Mendelson III, J.R. (1997a)
ery at additional locations. The type locality is at 1,570m asl. Data Providers: Georgina Santos-Barrera, Luis Canseco-Márquez
Population There is no information available on the population
status of this species.
Habitat and Ecology It appears that this species is confined to high
elevations covered with primary cloud forests and lowland rainforest
beneath the cloud forest. It breeds in permanent streams.
Major Threats The main threats are the fragmentation and
disturbance of the cloud and rainforest habitat, which results in
the disappearance of streams and a decrease in the humidity of
the leaf-litter.
Conservation Measures The range of the species is not within
any protected area, and only the preservation and restoration of
192 Threatened Amphibians of the World

CR Bufo sumatranus Peters, 1871

Critically Endangered B1ab(iii) Geographic Range This species is known only from a tiny area on Major Threats Since its rediscovery, there has been extensive and very rapid habitat destruction for rice paddies in
Order, Family: Anura, Bufonidae the island of Sumatra, Indonesia. The type locality is unspecified. It its only known locality, leading to the siltation of streams, which might affect its ability to breed.
Country Distribution: Indonesia is apparently confined to Lubuk Selasi, at the head of the Terusan Conservation Measures The species is not known from any protected area (it has not been recorded from a con-
Current Population Trend: Decreasing River at about 1,000m asl, on the borders of three districts, Padang servation forest about 10km from the only known locality). A community-based initiative involving local NGOs and
Pariaman, Solok and Pesisir Selatan. Despite searches, it has not communities is urgently needed to save the only known population of this species.
been found above 1,260m, or below 800m asl. Surveys in several Bibliography: Dubois, A. and Ohler, A. (1999), Iskandar, D.T. and Colijn, E. (2000)
other parts of Sumatra have failed to find this species, and it is likely Data Providers: Djoko Iskandar, Mumpuni
to have a very restricted distribution.
Population This species was rediscovered in 2001 after a gap of
141 years. At present, it is only known from one area, but is relatively
abundant within its tiny range.
Habitat and Ecology It has been found along a small, clear stream
with a width of 15m in secondary forest. No evidence of breeding
has been found, but it presumably breeds by larval development in
streams. It is not known whether or not the species is dependent
on forest, though this is likely.

EN Bufo tacanensis Smith, 1952

Endangered B1ab(iii) Geographic Range This species occurs at intermediate elevations along the Pacific versant of eastern Chiapas
Order, Family: Anura, Bufonidae (Unión de Juarez), Mexico to western Guatemala, at elevations of 1,500-1,700m asl.
Country Distribution: Guatemala, Mexico Population This is a rare species throughout its range, and there are no recent records of it in either Mexico or
Current Population Trend: Decreasing Guatemala, despite searches.
Habitat and Ecology This species has been recorded in foothills in premontane tropical forest. It is probably a
stream breeder.
Major Threats The alteration of the original forested areas by local people has severely impacted the only known
habitat for this species. Furthermore, as a montane stream-breeding bufonid, it is possible that the species may be
at risk of chytridiomycosis.
Conservation Measures Conservation of the forested areas along the foothills of the Tacana volcano is necessary
to ensure the preservation of this species. Fortunately, some programs of protection and restoration of the forests in
© Sean M. Rovito

the area have been implemented. In view of the severe risk of chytridiomycosis, the status of this species should be
closely monitored, and ex-situ populations should be established if chytrid is shown to be a threat.
Bibliography: Smith, W.P. (1952)
Data Providers: Georgina Santos-Barrera, Oscar Flores-Villela, Manuel Acevedo, Antonio Muñoz Alonso

CR Bufo taiensis Rödel and Ernst, 2000

Critically Endangered B2ab(iii) Geographic Range This species in known only from south-western Côte d’Ivoire. It possibly occurs in adjacent
Order, Family: Anura, Bufonidae Liberia.
Country Distribution: Côte d’Ivoire Population It is clearly a very rare species, and is known from only four specimens in an area that has been
Current Population Trend: Decreasing thoroughly surveyed.
Habitat and Ecology It is known only from primary rainforest. There is no information on its breeding, but if like B.
tuberosus, then breeding is likely to take place in small forest streams.
Major Threats There is ongoing forest loss in south-western Côte d’Ivoire, due to agriculture, timber extraction,
and human settlement.
Conservation Measures This species has been recorded from Taï National Park. Further survey work is required to de-
termine the biology and population status and trends of this species, as well as the limits of its distribution range.
© Mark-Oliver Rödel

Bibliography: Rödel, M.-O. (2000b), Rödel, M.-O. and Ernst, R. (2000), Tandy, M. and Perret, J.-L. (2000)
Data Providers: Mark-Oliver Rödel, Jean-Luc Perret, Mills Tandy

VU Bufo taladai Schwartz, 1960 CUBAN SPOTTED TOAD

Vulnerable B1ab(iii) Geographic Range This species is found in central and eastern Cuba from sea level up to 560m asl.
Order, Family: Anura, Bufonidae Population It is very common in suitable habitat.
Country Distribution: Cuba Habitat and Ecology It is found in lowland mesic broadleaf forests and cultivated fields (providing farming is not
Current Population Trend: Decreasing too intensive). Males call in streams and shallow creeks, and eggs are laid in permanent water.
Major Threats The major threat to this species is habitat loss due to intensive agriculture (livestock, sugarcane),
charcoaling, and nickel mining. Pollution from agricultural pesticides is also a threat.
Conservation Measures This species occurs in many protected areas.
Bibliography: Centro Nacional de Areas Protegidas (CNAP) (2002), Hedges, S.B. (1993), Hedges, S.B. (1999), Hedges, S.B. (2001),
Henderson, R.W. and Powell, R (1999), Henderson, R.W. and Powell, R. (2001), Schwartz, A. (1960b), Schwartz, A. and Henderson,
R.W. (1991)
© David Ortiz Martinez

Data Providers: Blair Hedges, Luis Díaz

 Globally Threatened Amphibian Species
EN Bufo tutelarius Mendelson, 1997
Endangered B1ab(iii,v) Geographic Range This species occurs on the Pacific versant of
Order, Family: Anura, Bufonidae Mexico from the Chimalapas region in south-eastern Oaxaca south
Country Distribution: Guatemala, Mexico through Chiapas to extreme western Guatemala, at elevations of
Current Population Trend: Decreasing 1,000-2,000m asl.
Population It is uncommon but still present in one Oaxaca locality,
with no evidence of a decline. In Chiapas, it has declined and is now
rare, though it still persists. Apparently, it is rare in Guatemala, as
recent expeditions within its range have not recorded this species,
indicating a decline there also.
Habitat and Ecology This species is closely associated with
streams, which provide breeding habitat, in cloud forest; it also
occurs in pine-oak (broadleaf) forest, and does tolerate moderate
forest disturbance.
Major Threats The major threat to this species has been the loss
and fragmentation of suitable forest habitat as a result of livestock
ranching and agricultural activities, and wood extraction. Since this
VU Bufo uzunguensis Loveridge, 1932
Vulnerable B1ab(iii)
Order, Family: Anura, Bufonidae
Country Distribution: Tanzania
Current Population Trend: Decreasing
© Michele Menegon
EN Bufo villiersi Angel, 1940
Endangered B1ab(iii)
Order, Family: Anura, Bufonidae
Country Distribution: Cameroon
Current Population Trend: Decreasing
© Jean-Louis Amiet
EN Bufoides meghalayanus (Yazdani and Chanda, 1971)
Endangered B1ab(iii) Geographic Range This species is known only from two sites,
Order, Family: Anura, Bufonidae the type locality “Mawblang, ca. 5km. southeast of Cherrapunji
Country Distribution: India town (Khasi-Jaintia Hills District, Meghalaya), latitude 25° 15’N
Current Population Trend: Decreasing and longitude 91° 44’E, altitude above mean sea-level 4,369 ft.
or ca. 1,330 meters”, India, and a second, recently discovered site
in Mizoram, Bangladesh. It probably occurs at least a little more
widely than current records suggest, especially in areas between
the two known sites.
Population Little is known about this species other than the original
description, but it is considered to be rare.
Habitat and Ecology The habitat is primarily wet forest floor,
although adjacent scrubland and grassland are also used. The adults
are to some degree arboreal. Breeding occurs on both pandanas and
the ground. Larvae are found in water holes, presumably in both
trees and on the ground.
193
is a montane, stream-breeding bufonid, it is possible that it may be affected by chytridiomycosis.
Conservation Measures It occurs in Reserva de la Biósfera La Sepultura and Reserva de la Biósfera El Triunfo,
but further protection of the remaining habitat in the range of this species is needed. In view of the severe risk of
chytridiomycosis, the status of this species should be closely monitored and ex-situ populations should be established
if chytrid is shown to be a threat.
Bibliography: Campbell, J.A. (2001), Mendelson III, J.R. (1997b), Mendelson III, J.R., Ustach, P.C. and Montes de-Oca, A.N. (1999)
Data Providers: Manuel Acevedo, Luis Canseco-Márquez, Antonio Muñoz Alonso
Geographic Range This species occurs in the Udzungwa Mountains and Southern Highlands of Tanzania, from
Dabaga south-west to Nyamwanga. It is found above 1,800m asl.
Population It appears to be locally abundant in suitable habitat.
Habitat and Ecology It is a species of swampy montane grassland. Its ability to adapt to habitat modification is
unknown, though most records are from pristine grassland. It breeds in wet areas in the bottom of gentle valleys
on high plateaus.
Major Threats The montane grasslands of southern Tanzania are being destroyed and fragmented as a result of
afforestation, overgrazing, agricultural expansion, and human settlement, which is a major threat to this species.
Conservation Measures It has not been recorded in any protected areas, and there is a need for improved protec-
tion of sites where the species is known to occur. It might be present in the Udzungwa National Park, but it is not
clear whether it ranges that far north.
Bibliography: Grandison, A.G.C. (1972b), Howell, K.M. (1993), Loveridge, A. (1932a), Poynton, J.C. (1998), Tandy, M. and Keith, R.
(1972)
Data Providers: Kim Howell, John Poynton, Mills Tandy, Michele Menegon
Geographic Range This species is known from the mountains of western Cameroon at 1,200-2,500m asl, where
it occurs on Mount Manenguba, the Bamileke Plateau (at Djuttitsa, Batie and Bangwa), the Bamboutos Mountains,
and Riboa (between the Adamawa and Mambilla Plateaus). It might occur more widely, and if Bufo djohongensis
proves to be a synonym, then its range will extend to the east of the Adamawa Plateau.
Population It is not a well-known species, though it is probably moderately common in suitable habitats.
Habitat and Ecology It lives along fast-flowing streams in montane grassland, sometimes with forest strips, and
hides in holes during the day. It breeds in slow-flowing streams bordered with trees.
Major Threats Although it can probably cope with some disruption to its habitat, it is probably suffering from
continuing degradation of its montane habitats as a result of smallholder farming activities, livestock ranching, wood
extraction, and human settlement.
Conservation Measures It is not known to occur in any protected areas. Protection of the remaining highland forests
in Cameroon, particularly Mount Manenguba, is urgently needed.
Notes on taxonomy: It is possible that Bufo djohongensis is a synonym of this species (J.-L. Amiet pers. comm.).
Bibliography: Amiet, J.-L. (1973b), Amiet, J.-L. (1976c), Perret, J.-L. (1971b), Tandy, M. and Keith, R. (1972)
Data Providers: Jean-Louis Amiet, Mills Tandy
Major Threats The main threat is clear-cutting of the species’ forest habitat.
Conservation Measures It has been reported from the Mizoram Wildlife Sanctuary, but additional habitat protec-
tion is necessary.
Bibliography: Chanda, S.K. (1994), Chanda, S.K. (2002), Dutta, S.K. (1997), Pillai, R.S. and Yazdani, G.M. (1973)
Data Providers: Sushil Dutta, Mohammed Firoz Ahmed
194 Threatened Amphibians of the World

VU Capensibufo rosei Hewitt, 1926 ROSE’S MOUNTAIN TOAD

Vulnerable B1ab(ii,iii,iv)+2ab(ii,iii,iv) Geographic Range This species occurs only in south-western South Africa, where it is restricted to the Cape
Order, Family: Anura, Bufonidae Peninsula and to the mountains south-west of the Breede River. Its altitudinal range is 60-1,600m asl, with more
Country Distribution: South Africa than 80% of localities being above 400m asl.
Current Population Trend: Decreasing Population It is locally common at breeding sites, and large dense breeding aggregations can sometimes be found.
However, it is absent from some apparently suitable sites.
Habitat and Ecology It is a species of mountain fynbos heathland, and does not survive in altered habitats. Breeding
takes place in temporary rock pools in weathered sandstone.
Major Threats The main threat to this species is the loss of its fynbos habitat, mainly because of the spread of
alien vegetation and frequent burning.
Conservation Measures Much of the range of this species is within protected areas, including the Cape Peninsula
National Park and several State Forests.
© Andrew Turner

Bibliography: Branch, W.R. (1988), Channing, A. (2001), De Villiers, A.L. (1997), Grandison, A.G.C. (1980), Minter, L.R. et al. (2004),
Passmore, N.I. and Carruthers, V.C. (1995), Tandy, M. and Keith, R. (1972)
Data Providers: Leslie Minter, Alan Channing, James Harrison

CR Churamiti maridadi Channing and Stanley, 2002

Critically Endangered B1ab(iii) Geographic Range This species is known only from the Ukaguru Mountains in eastern Tanzania. The only
Order, Family: Anura, Bufonidae specimens were collected at 1,840m asl. It has not been found in other, better-surveyed parts of the Eastern Arc
Country Distribution: Tanzania mountain chain.
Current Population Trend: Decreasing Population Only two female specimens are known.
Habitat and Ecology The only specimens were collected in dry montane forest. The large toe pads suggest that it is
arboreal. Its breeding is unknown, but the presence of large numbers of pigmented eggs in the only known specimens
suggests that it is neither a live-bearer, nor a direct developer.
Major Threats The forests in the Ukaguru Mountains are poorly protected, and threatened by agricultural encroach-
ment and human settlement.
Conservation Measures This species occurs in the Mamiwa-Kisara Forest Reserve. Further survey work is required
© William T. Stanley

to determine the biology and population status and trends of this species and the limits of its range.
Bibliography: Channing, A. and Stanley, W.T. (2002)
Data Providers: Kim Howell, Alan Channing

EN Dendrophryniscus carvalhoi Izecksohn, 1994 “1993”

Endangered B1ab(iii) Geographic Range This species is known from only from two localities at around 800m asl in the state of Espírito
Order, Family: Anura, Bufonidae Santo, southern Brazil: the type locality, Santa Tereza; and from nearby at Fundao. It almost certainly occurs a little
Country Distribution: Brazil more widely.
Current Population Trend: Decreasing Population It appears to be a rare species.
Habitat and Ecology It lives in the leaf-litter of primary and secondary forests, but not in more degraded habitats,
and is dependent upon bromeliads in which it lays its eggs and in which the the larvae develop.
Major Threats The major threats are habitat loss due to agricultural expansion, livestock grazing, clear-cutting, the
collection of bromeliads, and human settlement. The population at Fundao is threatened by coffee cultivation.
Conservation Measures It might occur in the Estacion Biólogica Santa Lucia, and Reserva Biólogica Augusto
© Miguel Trefaut Rodrigues

Ruschi, but this is not confirmed. It has been found in forest on private land, and bromeliads from this area have
been introduced into nearby protected areas. Since it is quite possible that these bromeliads contained larvae of the
species, it may have become inadvertently established in these protected areas.
Bibliography: Izecksohn, E. (1993)
Data Providers: Débora Silvano, Oswaldo Luiz Peixoto

EN Didynamipus sjostedti Andersson, 1903

Endangered B1ab(iii) of 200-1,250m asl, and also from the Kendonge Forest Reserve to the north of Mount Cameroon, the Mokoko Forest
Order, Family: Anura, Bufonidae Reserve north-west of Mount Cameroon close to the border with Nigeria, and from Baro just outside the north-western
Country Distribution: Cameroon, Equatorial border of Korup National Park. Although it can be expected between the known localities in Cameroon, there has been
Guinea, Nigeria extensive herpetological fieldwork in this area, which is perhaps indicative of a patchy distribution.
Population There have been very few records of this species, but it is common on the southern slopes of Mount Camer-
oon, and is especially numerous at around 1,000m asl. There is also a healthy population in the Makoko Forest Reserve,
and it is locally extremely abundant in the Oban Hills. There is no recent information on its status on Bioko.
Habitat and Ecology It lives on forest edges and in clearings in moist forest from the lowlands to the submontane
zone. They form aggregations of 5-40 individuals including males, females and juveniles. They are most often seen
sitting on wet leaves of low herbaceous vegetation. It has also been found in selectively logged forest on the edge of
© Mary E. Gartshore

Current Population Trend: Decreasing
Geographic Range This species is known from extreme south-western Cameroon in the general area of Mount
Cameroon and surrounding forests, and from old specimens at 400-600m asl near Basile on Bioko Island, Equatorial
Guinea. It has recently been discovered in the Oban Hills in Nigeria (M. Gartshore pers. comm.). In Cameroon there
are records from Mount Cameroon (especially from the southern slopes) where it has been recorded at an altitude
small farms. Its breeding habits are not known, but it is suspected to have a viviparous mode of reproduction, since
it is most closely related to the West African genus Nimbaphrynoides.
Major Threats The main threat to the species is habitat loss primarily due to agricultural expansion, wood extrac-
tion, and human settlement.
Conservation Measures It is recorded from the Makoko and Kendonge Forest Reserves, and probably occurs in
Korup National Park. It also has recently been found in the Cross River National Park in Nigeria.
Bibliography: Amiet, J.-L. (1976b), Gartshore, M.E. (1986), Gartshore, M.E. (1999), Grandison, A.G.C. (1981), Lawson, D.P. (1993),
Mertens, R. (1965)
Data Providers: Mary Gartshore, Jean-Louis Amiet, Robert Drewes
 Globally Threatened Amphibian Species 195

CR Leptophryne cruentata (Tschudi, 1838) BLEEDING TOAD

Critically Endangered A2ac Geographic Range This species occurs only on Mount Pangrango, Mount Gedeh and Curug Luhur, Jawa Barat,
Order, Family: Anura, Bufonidae Java, Indonesia, at altitudes between 1,000m and 2,000m asl.
Country Distribution: Indonesia Population In 1976, this species was abundant within its small range. In 1987, it was very rare following the eruption
Current Population Trend: Decreasing of Mount Galunggung. There were no records from the early 1990s until 2003, when one individual was sighted from
the Cibeureum Waterfall. It appears to have undergone a major decline.
Habitat and Ecology It lives in the boundary zone between moist lowland and montane forest. It breeds in very
slow-moving, intermittent streams in forest where the larvae also develop.
Major Threats It appears to have declined drastically due to a volcanic eruption. However, its decline is also remi-
niscent of similar disappearances of montane stream-breeding amphibians in other parts of the moist tropics, and so
chytridiomycosis cannot be ruled out (although this disease has not so far been recorded in this region).
Conservation Measures This species occurs in the Gunung Gede Pangrango National Park. Surveys are needed to
© Djoko Iskandar

locate this species and a captive-breeding programme might need to be established.

Bibliography: Iskandar, D.T. and Colijn, E. (2000)
Data Providers: Djoko Iskandar, Mumpuni

EN Melanophryniscus devincenzii Klappenbach, 1968

Endangered B2ab(iii) Geographic Range This species occurs in the southern part of Misiones Province, Argentina, and Rivera, Tacuarembó,
Order, Family: Anura, Bufonidae and Cerro Largo Departments in Uruguay at elevations of 150-350m asl.
Country Distribution: Argentina, Uruguay Population It is very rare in Misiones (Argentina) where males have been found (in 1998 and 2003) in amplexus
Current Population Trend: Decreasing with females of other species, possibly because of a lack of conspecifics. The population status of the species in
Uruguay is unknown.
Habitat and Ecology It occurs in open areas with rocky outcrops, but also occurs in the ecotones between grass-
lands and forests of Astronium balansae. In Uruguay, it has been found in eucalyptus plantations. Nothing is known
about its breeding habits, other than that males have been found in amplexus in temporary streams and the larvae
presumably develop in streams.
Major Threats The pollution of soil and water due to agricultural practices is a known threat to this species. In
Misiones, it is also threatened by the conversion of native habitat to pine plantations.
Conservation Measures It occurs in small provincial parks in Argentina, including Fachinal and the Campos San
Juan private reserves. Broad-scale conservation actions are necessary to combat the effects of pollution on its
© Axel Kwet

breeding habitat.
Bibliography: Langone, J.A. (1994), Lavilla, E.O. et al. (2000), Lavilla, E.O. and Cei, J.M. (2001), Lavilla, E.O., Barrionuevo, S. and Baldo,
D. (2002), Maneyro, R. and Langone, J.A. (2001)
Data Providers: Esteban Lavilla, Diego Baldo, Jose Langone

VU Melanophryniscus dorsalis (Mertens, 1933)

Vulnerable B1ab(iii)+2ab(iii) Geographic Range This species occurs along the coast of Brazil in Santa Catarina and Rio Grande de Sul states,
Order, Family: Anura, Bufonidae from 0-20m asl.
Country Distribution: Brazil Population It was previously collected rather extensively, but there have only been a few recent collections and the
Current Population Trend: Decreasing species is believed to have undergone declines in recent years.
Habitat and Ecology It occurs on sand dunes and nearby areas, and reproduces in temporal pools.
Major Threats The major threat to this species is habitat loss and degradation due to the conversion of habitat for
beaches, urbanization, and off-road recreational vehicle use.
Conservation Measures It occurs in Guarita Municipal Park, but there is a need for improved protection of habitat
at other sites at which this species is known to occur.
Bibliography: Garcia, P.C.A. and Vinciprova, G. (2003)
Data Providers: Paulo Garcia
© Axel Kwet

VU Melanophryniscus macrogranulosus Braun, 1973

Vulnerable B1ab(iii) Geographic Range This species is known only from the type locality: Torres, Rio Grande do Sul State, Brazil. The
Order, Family: Anura, Bufonidae type locality is listed as a municipality, which has since been divided into several municipalities.
Country Distribution: Brazil Population It has not been recorded since 1960.
Current Population Trend: Decreasing Habitat and Ecology There is no information about its ecology or biology in the literature. It presumably breeds
in water.
Major Threats There is no information on threats to this species, although there has been extensive habitat loss in
its range due to urbanization in the last 30 years and there is probably very little habitat remaining.
Conservation Measures There are no protected areas near where this species was collected. It is categorized as
Vulnerable on the Brazil national list.
Bibliography: Braun, P.C. (1973), Braun, P.C. and Braun, C.A.S. (1980)
© Marcos Di Bernardo

Data Providers: Débora Silvano, Paulo Garcia

196 Threatened Amphibians of the World

VU Melanophryniscus montevidensis (Phillippi, 1902)

Vulnerable A2ac; B1ab(iii,iv,v) Geographic Range This species is restricted to coastal Uruguay (Montevideo, Canelones, Maldonado, and Rocha
Order, Family: Anura, Bufonidae Departments) and adjacent Brazil (one locality in Rio Grande do Sul state). It occurs at sea level.
Country Distribution: Brazil, Uruguay Population This species is in decline in some areas and extirpated in others. Nevertheless, it is still common at
Current Population Trend: Decreasing several localities.
Habitat and Ecology It is a diurnal species inhabiting coastal sand dunes. It is an explosive breeder, and may be
found in large numbers in temporary pools after heavy rains; the larvae develop in these pools. It is not tolerant of
habitat disturbance.
Major Threats The major threat to this species is habitat loss due to human settlements, exotic tree plantations,
and the drainage of wetlands.
Conservation Measures It does not occur in any protected areas, and there is a need for improved protection of
sites at which this species is known to occur.
Notes on taxonomy: Elevated to full species status by Klappenbach and Langone (1992).
© Axel Kwet

Bibliography: Klappenbach, M.A. and Langone, J.A. (1992), Langone, J.A. (1994), Maneyro, R. and Langone, J.A. (2001), Tedros, M.,
Kolenc, F. and Borteiro, C. (2001)
Data Providers: Jose Langone

VU Melanophryniscus orejasmirandai Prigioni and Langone, 1986

Vulnerable D2 Geographic Range The type locality of this species is Sierra de Bibliography: Langone, J.A. (1994), Maneyro, R. and Langone, J.A. (2001), Prigioni, C.M. and Langone, J.A. (1986)
Order, Family: Anura, Bufonidae Ánimas, Maldonado Province, Uruguay, at an elevation of 350-500m Data Providers: Esteban Lavilla, Jose Langone
Country Distribution: Uruguay asl. It occurs in an area of less than 10km².
Current Population Trend: Stable Population It is common at its only known locality, and has not
undergone any change in population size in 10 years of monitoring
from 1990 to 2000.
Habitat and Ecology It occurs in grasslands and rocky outcrops,
and reproduces in permanent small streams. Its tolerance to habitat
disturbance is unknown.
Major Threats Although there are currently no major threats to the
species, habitat loss and degradation due to wood plantations and
touristic activities might result in declines in the future.
Conservation Measures It does not occur in any protected areas.
This species requires continued close population monitoring particu-
larly since it is known only from a single location.

VU Metaphryniscus sosae Señaris, Ayarzaguena and Gorzula, 1994

Vulnerable D2 Geographic Range This species is known only from the type locality,
Order, Family: Anura, Bufonidae on top of Cerro Marahuaca, in Amazonas state (03° 40’N; 65° 27’W),
Country Distribution: Venezuela at 2,600m asl, in Venezuela.
Current Population Trend: Stable Population The population status of this species is not known.
Habitat and Ecology It inhabits high montane tepui habitat.
It breeds by direct development, perhaps with internal fertilization.
Major Threats There are no current major threats, but its re-
stricted range makes it susceptible to threatening processes such
as wildfire.
Conservation Measures There is a population within Parque
Nacional Duida-Marahuaca. There is a need for close population
monitoring of this species, particularly since it is restricted to a
single location.
Bibliography: Barrio Amorós, C.L. (2004), La Marca, E. (1997), Señaris, J.C.,
Ayarsagüena, J. and Gorzula, S. (1994)
Data Providers: Enrique La Marca, Celsa Señaris

CR Nectophrynoides asperginis Poynton, Howell, Clarke and Lovett, 1999 “1998” KIHANSI SPRAY TOAD
Critically Endangered B1ab(ii,iii,v)+2ab( by mid-January 2004, only three individuals could be seen and just two males were heard calling. There were a few
ii,iii,v) records of calling animals during the rest of 2004, and an unconfirmed report from January 2005, but there are no
Order, Family: Anura, Bufonidae records of any individuals since, despite surveys, and the species might now be extinct in the wild.
Country Distribution: Tanzania Habitat and Ecology It lives only in soaked herbaceous vegetation in the spray zone of the Kihansi Falls. It breeds
Current Population Trend: Decreasing by internal fertilization, the females retaining the larvae internally in the oviduct until little toadlets are born.
© Timothy A. Herman / Toledo Zoological Society

CITES: Appendix I
Geographic Range This species is only known from the Kihansi Falls, in the Kihansi Gorge, in the Udzungwa
Mountains, eastern Tanzania, at 600-940m asl. Its global range covers a tiny area of just two hectares around the
Kihansi Falls, and searches for it around other waterfalls on the escarpment of the Udzungwa Mountains have not
located any additional populations.
Population It was formerly abundant in a tiny area, with a population of around 17,000 animals. Reports indicate
that the species fluctuated naturally in its population size. The population appeared to be at a high in May 1999, at a
low in 2001 and 2002 (when the population was estimated at 1,000 animals), and at a high again in June 2003, when
perhaps as many as 20,000 individuals survived. However, subsequently the population went into steep decline, and
Major Threats The serious decline of this species appears related to the Lower Kihansi Hydropower Project, involving
the construction of a dam in 2000 upstream on the Kihansi River, which cut off 90% of the original water flow to the
gorge, thereby hugely reducing the volume of spray, particularly in the dry season, as well as altering the vegetational
composition. An artificial gravity-fed sprinkler system was set up to mimic the natural spray of the Kihansi ecosystem
with the remaining water flow. Unfortunately, the sprinkler system was not ready by the time the water was cut off in
2000, and by the time the sprinklers came on nine months later the ecosystem had already dried up (see Krajick 2006).
Later, during the dry season in 2003, the artificial sprinkler system failed for a while. Around this time, the fungal
disease chytridiomycosis was confirmed in dead animals of this species, and this disease is probably responsible for
the final population crash. It is possible that the drought caused by the failure of the sprinkler system resulted in stress
to the animals that rendered them susceptible to the disease. There are also reports that the 2003 crash coincided
with a brief opening of the dam’s floodgates to flush sediments; tests showed that these contained pesticides used
in maize farming operations upstream, in concentrations sufficient to kill the toads (Krajick 2006).
Conservation Measures It is not known from any protected areas. Sufficient minimum bypass flow from the dam
is required to maintain the spray habitat. An artificial sprinkler system is in place, and this is especially important in
the dry season. Captive breeding is ongoing in Toledo and New York Bronx Zoos. The captive population has fluctu-
ated as husbandry problems have been encountered and addressed (animals were initially plagued with various
infections and nutritional deficiencies), but currently stands at about 300 individuals. Investigations are urgently
needed to determine whether or not any individuals survive in the wild, and whether or not chytridiomycosis is still
present in any remaining individuals.
Bibliography: Krajick, K. (2006), Poynton, J.C. (1998), Poynton, J.C. (2003b), Poynton, J.C. et al. (1998), Quinn, C.H. et al. (2005),
Zippel, K. (2005)
Data Providers: John Poynton, Kim Howell, Alan Channing, Simon Loader, Michele Menegon
 Globally Threatened Amphibian Species 197

EN Nectophrynoides cryptus Perret, 1971

Endangered B1ab(iii) Geographic Range This species is known only from the northern Conservation Measures It occurs in the Uluguru North Forest Reserve, but this area is not generally managed
Order, Family: Anura, Bufonidae part of the Uluguru Mountains in Tanzania where it has been recorded for biodiversity conservation. Improved management of this reserve, and the protection of other remaining forest
Country Distribution: Tanzania at 600-2,200m asl. habitat in the Ulugurus, is necessary.
Current Population Trend: Decreasing Population There have been no records of this species since the Bibliography: Howell, K.M. (1993), Perret, J.-L. (1971a), Perret, J.-L. (1972), Poynton, J.C. (1998)
CITES: Appendix I original collections of the species in 1926 and 1927. In view of the Data Providers: Simon Loader, John Poynton, Kim Howell
amount of herpetological work that has taken place in this area, it
would appear to be a rare species.
Habitat and Ecology The type series was collected in forest,
secondary forest, banana cultivation and bamboo. From this informa-
tion, it seems that it can tolerate a degree of habitat disturbance,
but probably not the complete opening of its habitat that has taken
place in some parts of the Ulugurus. It is ovoviviparous, with internal
fertilization, the females retaining the larvae internally in the oviduct
until the birth of the toadlets.
Major Threats The habitat of this species is probably being lost to
agricultural encroachment, wood extraction, and expanding human
settlements, especially at lower altitudes.

EN Nectophrynoides minutus Perret, 1972

Endangered B1ab(iii) Geographic Range This species is known only from the Uluguru Bibliography: Howell, K.M. (1993), Perret, J.-L. (1971a), Perret, J.-L. (1972), Poynton, J.C. (1998)
Order, Family: Anura, Bufonidae and Rubeho Mountains in eastern Tanzania, at 1,200-1,500m asl, Data Providers: Simon Loader, John Poynton, Kim Howell
Country Distribution: Tanzania and probably higher.
Current Population Trend: Decreasing Population It appears to be a reasonably common species.
CITES: Appendix I Habitat and Ecology It has been found only in undisturbed forest,
and appears not to tolerate habitat alteration. It is ovoviviparous, with
internal fertilization and the females retaining the larvae internally
in the oviduct until the toadlets are born.
Major Threats Its habitat is probably being lost, especially at lower
altitudes, due to agricultural encroachment, wood extraction, and
expanding human settlements.
Conservation Measures It occurs in the Uluguru North and Uluguru
South Forest Reserves, and also in the Ukwiva Forest Reserve in the
Rubeho Mountains. These forest reserves are in need of continued
and strengthened management to ensure the maintenance of
remaining forest habitat.

CR Nectophrynoides poyntoni Menegon, Salvidio and Loader, 2004

Critically Endangered B1ab(iii)+2ab(iii) Geographic Range This species is known only from the Mkalazi Valley at 1,200m in the Udzungwa Scarp Forest
Order, Family: Anura, Bufonidae Reserve, Udzungwa Mountains, in eastern Tanzania. Surveys of other parts of the Udzungwa Forest Reserve have
Country Distribution: Tanzania not located this species.
Current Population Trend: Decreasing Population It is reasonably common within its tiny range, though much less numerous than the sympatric Nec-
CITES: Appendix I tophrynoides tornieri.
Habitat and Ecology All specimens were found in moist submontane rainforest, close to a stream. At night they
can be found on leaves 60-160cm above the ground, and during the day they hide on the ground under fallen trees
and coarse wood debris. Its breeding strategy is unknown, but it is assumed to be a live-bearer, like other member
of its genus, with internal fertilisation, giving birth to tiny toadlets.
Major Threats Its habitat is probably being lost due to agricultural encroachment, wood extraction, and expanding
human settlements.
© Michele Menegon

Conservation Measures This species occurs in the Udzungwa Scarp Forest Reserve, but not in any well-protected
areas. The population status and trends of this species require monitoring.
Bibliography: Menegon, M., Salvidio, S. and Loader, S.P. (2004)
Data Providers: Michele Menegon, Simon Loader

EN Nectophrynoides pseudotornieri Menegon, Salvidio and Loader, 2004

Endangered B1ab(iii) Geographic Range This species is known only from the Uluguru conservation and is in need of improved management. Further survey work is needed to determine the current
Order, Family: Anura, Bufonidae North Forest Reserve on the eastern slopes of the northern part of population status of this species.
Country Distribution: Tanzania the Uluguru Mountains, eastern Tanzania, at 1,080-1,345m asl. It Bibliography: Menegon, M., Salvidio, S. and Loader, S.P. (2004)
Current Population Trend: Decreasing appears to have a very restricted distribution. Data Providers: Michele Menegon, Simon Loader
CITES: Appendix I Population There is no information on the population status of
this species; it is known only from two specimens, collected in
1996 and 2000.
Habitat and Ecology This species is known only from tall, submon-
tane rainforest. Its breeding biology is unknown, but it is assumed
to be ovoviviparous, like other member of its genus, with internal
fertilization, giving birth to tiny toadlets.
Major Threats The submontane forest on the eastern slopes of
the Uluguru Mountains has been extensively cleared, mainly due
to agricultural encroachment, wood extraction, and expanding
human settlements.
Conservation Measures It occurs in the Uluguru North Forest
Reserve, but this area is not generally managed for biodiversity
198 Threatened Amphibians of the World

EN Nectophrynoides vestergaardi Menegon, Salvidio and Loader, 2004

Endangered B1ab(iii) Geographic Range This species is known only from the West Usambara Mountains in north-eastern Tanzania.
Order, Family: Anura, Bufonidae There are records from the Shume-Magamba Forest Reserve, the Mazumbai Forest Reserve, and the Ambangulu
Country Distribution: Tanzania Estate between 1,230 and 2,000m asl.
Current Population Trend: Decreasing Population There is little information available on its population status. However, the fact that 23 specimens have
CITES: Appendix I been found widely over the West Usambara Mountains, despite limited survey effort, suggests that it is not uncom-
mon in suitable habitat within its small range.
Habitat and Ecology All records have been from montane and submontane forest, including in the ecotone
between forest and ericaceous vegetation. It is probably terrestrial, and some animals were found inside a rotten
log. Its breeding is unknown, but it is assumed to be ovoviviparous, like other members of its genus, with internal
fertilization, giving birth to tiny toadlets.
Major Threats In some parts of the West Usambaras its habitat is probably being lost, especially due to agricultural
© Martin Vestergaard

encroachment, commercial logging, wood extraction, and expanding human settlements.

Conservation Measures It occurs in the University of Dar es Salaam’s forest reserve at Mazumbai, but additional
protection of the habitat in the West Usambara Mountains is needed.
Bibliography: Menegon, M., Salvidio, S. and Loader, S.P. (2004)
Data Providers: Michele Menegon, Simon Loader

VU Nectophrynoides viviparus (Tornier, 1905)

Vulnerable B1ab(iii) Geographic Range This species occurs in the Uluguru and Udzungwa Mountains and in the Southern Highlands of
Order, Family: Anura, Bufonidae eastern and southern Tanzania. It is a montane species, ranging from 1,350-2,800m asl.
Country Distribution: Tanzania Population It is relatively common.
Current Population Trend: Decreasing Habitat and Ecology It is a terrestrial species living in forest, bamboo, and grasslands at the forest edge. It has
© David Moyer / Wildlife Conservation Society

CITES: Appendix I been found in maize cultivation, but needs to be close to the forest edge, and probably cannot tolerate complete
opening up of its habitat. It breeds by internal fertilization, the females retaining the larvae internally in the oviduct
until little toadlets are born.
Major Threats The major threat to this species is habitat loss, especially at lower altitudes, due to agricultural
encroachment, wood extraction and expanding human settlements.
Conservation Measures It occurs in several forest reserves, but has not been found in any well-protected area. It
might occur in the Udzungwa National Park, but has thus far not been recorded.
Bibliography: Howell, K.M. (1993), Menegon, M. (2000), Perret, J.-L. (1971a), Perret, J.-L. (1972), Poynton, J.C. (1998), Poynton, J.C.
(2003b)
Data Providers: Simon Loader, John Poynton, Kim Howell

CR Nectophrynoides wendyae Clarke, 1988

Critically Endangered B1ab(iii)+2ab(iii) Geographic Range This species is known from one tiny area, in the Udzungwa Scarp Forest Reserve, above Chita,
Order, Family: Anura, Bufonidae on the escarpment of the Udzungwa Mountains, in eastern Tanzania. It is found from 1,500-1,650m asl. It has not
Country Distribution: Tanzania been found at other localities in the Udzungwa Mountains.
Current Population Trend: Decreasing Population The species is quite common in one tiny area, measuring roughly 300m x 300m.
CITES: Appendix I Habitat and Ecology It is a species of montane moist forest, living on the ground. It is not known whether or not it
can survive in anthropogenically disturbed habitats, since the only known site is in undisturbed forest. It breeds by
internal fertilization, the females retaining the larvae internally in the oviduct until little toadlets are born.
Major Threats Its habitat is probably being lost due to agricultural encroachment, wood extraction, and expanding
human settlements.
Conservation Measures This species occurs in the Udzungwa Scarp Forest Reserve, but not in any well-protected
areas. The population status and trends of this species require monitoring.
© Michele Menegon

Bibliography: Clarke, B.T. (1988), Howell, K.M. (1993), Poynton, J.C. (1998), Poynton, J.C. (2003b)
Data Providers: Michele Menegon, Simon Loader, John Poynton, Kim Howell

CR Nimbaphrynoides liberiensis Xavier, 1979 “1978”

Critically Endangered B1ab(iii) Geographic Range This species is known only from the Mount Nimba region in Liberia. It presumably also occurs in
Order, Family: Anura, Bufonidae adjacent parts of Guinea and perhaps also in Côte d’Ivoire, but there have not yet been any records. It is a montane
Country Distribution: Liberia species, and the type locality is at 1,290m asl.
Current Population Trend: Decreasing Population It is apparently extremely common in its small range, but there do not appear to have been any recent
CITES: Appendix I records, presumably due to a lack of fieldwork in the area.
Habitat and Ecology It is apparently a species of forest and forest edge close to savannah. It is a viviparous species,
with the female nourishing the young internally prior to the birth of small toadlets.
Major Threats Habitat loss and degradation due to the mining of bauxite is the biggest threat to this species.
There is probably also ongoing loss of forest due to wood extraction. The species is inherently at risk because of
its small range.
Conservation Measures This species presumably occurs in the Mount Nimba Strict Nature Reserve, which is a World
Heritage Site. Survey work is necessary to determine the current population status and trends of this species.
© Laura Sandberger

Bibliography: Grandison, A.G.C. (1981), Perret, J.-L. (1971a), Perret, J.-L. (1972), Xavier, F. (1978)
Data Providers: Mark-Oliver Rödel, Arne Schiøtz
 Globally Threatened Amphibian Species 199

CR Nimbaphrynoides occidentalis (Angel, 1943)

Critically Endangered B1ab(iii) Geographic Range This species is known only from the Mount Nimba region in Guinea and Côte d’Ivoire. It
Order, Family: Anura, Bufonidae presumably also occurs in Liberia, but there have not yet been any records. It is a montane species, occurring above
Country Distribution: Côte d’Ivoire, Guinea 1,000m asl.
Current Population Trend: Decreasing Population It is apparently an abundant species within its small range, and a survey in 2003 succeeded in locating
CITES: Appendix I several individuals, even during the cold season when they are supposed to be inactive.
Habitat and Ecology It is a species of montane grassland. It is a viviparous species, with the female nourishing the
young internally prior to the birth of small toadlets.
Major Threats Habitat loss and degradation due to the mining of iron ore/bauxite is the biggest threat to this spe-
cies, and new mining sites have been selected recently for mining in the Guinean part of Mount Nimba. There is a
risk that mining could destroy the entire range of the species. Fires in the montane grassland might be a threat. The
species is inherently at risk because of its small range.
Conservation Measures This species presumably occurs in the Mount Nimba Strict Nature Reserve, which is a
© Piotr Naskrecki

World Heritage Site. Survey work is necessary to monitor the population status of this species.
Bibliography: Lamotte, M. (1959), Lamotte, M. and Sanchez-Lamotte, C. (1999), Perret, J.-L. (1971a), Perret, J.-L. (1972), Xavier, F.
(1978), Xavier, F., Zuber-Vogeli, M. and Le Quang, Y. (1970)
Data Providers: Mark-Oliver Rödel, Arne Schiøtz

VU Oreophrynella cryptica Señaris, 1995 “1993”

Vulnerable D2 Geographic Range This species is restricted to the type locality:
Order, Family: Anura, Bufonidae “Sector este, cima del Auyan-tepui, Estado Bolívar”, in Venezuela
Country Distribution: Venezuela (05° 53’ 36”N; 62° 29’ 12”W), at 1,750m asl.
Current Population Trend: Stable Population It is a rare species.
Habitat and Ecology It is an inhabitant of high montane tepui
habitat. Its breeding habitat is unknown, but it is presumed to breed
by direct development.
Major Threats Although there are no current major threats, its
restricted range makes it more susceptible to threatening processes,
such as wildfire and disturbance of the habitat by tourists.
Conservation Measures The species’ range is within Parque
Nacional Canaima. There is a need for close population monitoring
of this species given that it is restricted to a single location.
Bibliography: Barrio Amorós, C.L. (2004), Barrio, C. (1998), La Marca, E. (1997),
Señaris, J.C. (1995)
Data Providers: Enrique La Marca, Celsa Señaris

VU Oreophrynella huberi Diego-Aransay and Gorzula, 1987

Vulnerable D2 Geographic Range This species is restricted to the type locality, Cerro
Order, Family: Anura, Bufonidae El Sol, to the north-east of the Auyán-tepui (0.6km² at 06° 06’N; 62°
Country Distribution: Venezuela 32’W), in Bolívar state, Venezuela, at an elevation of 1,700m asl.
Current Population Trend: Stable Population It is a very rare species.
Habitat and Ecology It is a diurnal toad that is found in thick herba-
ceous vegetation on peat bogs of high montane tepui environments.
It breeds by direct development.
Major Threats Although there are no current major threats, its re-
stricted range makes it more susceptible to threatening processes.
Conservation Measures The type locality of this species is within
Parque Nacional Canaima. There is a need for close population moni-
toring of this species given that it is restricted to a single location.
Bibliography: Barrio Amorós, C.L. (2004), Diego-Aransay, A. and Gorzula, S.
(1987), La Marca, E. (1992), Péfaur, J.E. and Rivero, J.A. (2000), Señaris, J.C.,
Ayarsagüena, J. and Gorzula, S. (1994), Vial, J.L. and Saylor, L. (1993)
Data Providers: Enrique La Marca, Celsa Señaris

VU Oreophrynella macconnelli Boulenger, 1900

Vulnerable D2 Geographic Range This species is known from the base of Mount Bibliography: Barrio Amorós, C.L. (2004), Barrio, C. (1998), Frost, D.R. (1985), Gines, H. (1959), La Marca, E. (1992), La Marca, E. (1995b),
Order, Family: Anura, Bufonidae Roraima on the border of Guyana and Venezuela, at an elevation La Marca, E. (1997), McDiarmid, R. (1971), McDiarmid, R. and Gorzula, S. (1989), Péfaur, J.E. and Rivero, J.A. (2000), Rivero, J.A. (1961),
Country Distribution: Guyana, Venezuela of 1,060m asl. It is not absolutely certain which country the type Señaris, J.C., Ayarsagüena, J. and Gorzula, S. (1994), Vial, J.L. and Saylor, L. (1993)
Current Population Trend: Stable specimen comes from, but it is presumed that the species is in fact Data Providers: Marinus Hoogmoed, Celsa Señaris
present on both sides of the border.
Population The population status of this species is not known.
Habitat and Ecology It is an inhabitant of tropical moist submon-
tane forest. Its breeding habitat is unknown, but it is presumed to
breed by direct development.
Major Threats Threats to this species are unknown, although its
range is in relatively pristine habitat at present.
Conservation Measures It is not known from any protected
areas. A well-managed protected area would help to ensure the
conservation of this range-restricted species. There is also a need
for close population monitoring given that it is apparently restricted
to a single location.
200 Threatened Amphibians of the World

VU Oreophrynella nigra Señaris, Ayarzaguena and Gorzula, 1994

Vulnerable D2 Geographic Range This species is known from Tepuis Kukenan and Yuruaní, in Bolívar State, Venezuela, at 2,300-
Order, Family: Anura, Bufonidae 2,700m asl.
Country Distribution: Venezuela Population It is a common species.
Current Population Trend: Stable Habitat and Ecology This is a diurnal frog that can be found on rocks in montane tepui environments. It is a com-
munally breeding species, with breeding taking place by direct development in galleries within peat bogs.
Major Threats Although there are no current major threats, its restricted range makes it more susceptible to
threatening processes such as wildfire.
Conservation Measures It is recorded from Yuruaní National Monument. There is a need for close population
monitoring of this species given that it is restricted to a single location.
© Karl-Heinz Jungfer Bibliography: Barrio Amorós, C.L. (2004), La Marca, E. (1997), McDiarmid, R. and Gorzula, S. (1989), Señaris, J.C., Ayarsagüena, J.
and Gorzula, S. (1994), Solano, H. (1989)
Data Providers: Enrique La Marca, Celsa Señaris

VU Oreophrynella quelchii (Boulenger, 1895)

Vulnerable D2 Geographic Range This species is restricted to the summit of Mount Roraima in Venezuela, Guyana and Brazil, and
Order, Family: Anura, Bufonidae from Wei-Assipo-Tepui in Guyana. It has been recorded from 2,300-2,800m asl.
© César Luis Barrio-Amorós / Fundación AndígenA

Country Distribution: Brazil, Guyana, Venezuela Population It is common on the summit of Mount Roraima, and its population is probably stable.
Current Population Trend: Stable Habitat and Ecology A diurnal toad usually found on open rock surfaces in high montane tepui environments. It
breeds by direct development.
Major Threats Although there are no current major threats, the restricted range of this species makes it more vulner-
able to threatening processes, such as disturbance by tourists (who frequently handle the animals).
Conservation Measures It is protected in Monumento Natural Los Tepuyes in Venezuela, and Parque Nacional Monte
Roraima in Brazil. There is a need for increased education among tourists to make them aware of the importance of
not handling these animals in the wild. Close population monitoring is also required, particularly since this species
is known only from a single location.
Bibliography: Barrio Amorós, C.L. (2004), Frost, D.R. (1985), Gines, H. (1959), Gorzula, S. and Señaris, J.C. (1998), La Marca, E. (1992), La
Marca, E. (1995b), La Marca, E. (1997), McDiarmid, R. (1971), McDiarmid, R. and Gorzula, S. (1989), Péfaur, J.E. and Rivero, J.A. (2000),
Rivero, J.A. (1961), Señaris, J.C., Ayarsagüena, J. and Gorzula, S. (1994), Solano, H. (1989), Vial, J.L. and Saylor, L. (1993)
Data Providers: Marinus Hoogmoed, Celsa Señaris

VU Oreophrynella vasquezi Señaris, Ayarzaguena and Gorzula, 1994

Vulnerable D2 Geographic Range This species is restricted to the summit of
Order, Family: Anura, Bufonidae Ilú-tepui (05° 25’N; 60° 58’W), in Bolívar State, Venezuela, where it
Country Distribution: Venezuela has been recorded from 2,450-2,650m asl.
Current Population Trend: Stable Population The population status of this species is not known.
Habitat and Ecology A diurnal, high-montane toad found in
tepui habitats. It breeds by direct development, and lays its eggs
under rocks.
Major Threats Although there are no current major threats, the
restricted range of this species makes it more vulnerable to threaten-
ing processes, such as wildfire.
Conservation Measures It most likely occurs within Parque Nacio-
nal Canaima. Close population monitoring of this species is required,
particularly since it is known only from a single location.
Bibliography: Barrio Amorós, C.L. (2004), Barrio, C. (1998), La Marca, E. (1997),
Señaris, J.C., Ayarsagüena, J. and Gorzula, S. (1994)
Data Providers: Enrique La Marca, Celsa Señaris

EN Osornophryne antisana Hoogmoed, 1987

Endangered B1ab(iii,v) Geographic Range This species is known only from the type locality, Antisana, Napo Province, Ecuador, and from
Order, Family: Anura, Bufonidae the volcano of Quilindana. In total, it is known from only four localities. It has been recorded between 3,400 and
Country Distribution: Ecuador 4,000m asl.
Current Population Trend: Decreasing Population It appears to be a rare, and locally uncommon species; however, this may be an artefact of inappropriate
survey techniques.For example, when digging into the soil under the páramo plants, dozens of specimens have been
found (M. Yanez and D.F. Cisneros-Heredia pers. comm.). The population of this species is thought to be declining.
Habitat and Ecology It is terrestrial, and both diurnal and nocturnal, inhabiting wet sub-páramo habitats. The
© Mario Humberto Yánez-Muñoz

holotype was collected in an area of meadows with isolated patches of forest. Creeks and swamps are the typical
landscape on the upper limit of the wet altitudinal forest (Hoogmoed 1987). It breeds by direct development, and
lays its eggs on soil under vegetation.
Major Threats The major threat to this species is habitat loss due to agricultural activities, both crops and livestock;
climate change may also be impacting the species.
Conservation Measures This species occurs in the Reserva Ecológica Antisana, the Reserva Ecológica Cayambe-
Coca, and Parque Nacional Llanganates.
Notes on taxonomy: The identity of the specimens from the northern part of Parque Nacional Llanganates requires confirmation.
Bibliography: Hoogmoed, M.S. (1987)
Data Providers: Luis A. Coloma, Santiago Ron, Diego Cisneros-Heredia, Mario Yánez-Muñoz
 Globally Threatened Amphibian Species 201

EN Osornophryne guacamayo Hoogmoed, 1987 GUACAMAYO PLUMP TOAD

Endangered B1ab(iii) Geographic Range This species is known from the Cordillera de Guacamayos in north-eastern Ecuador (in Napo
Order, Family: Anura, Bufonidae Provice) and from the Valle del Sibundoy in southern Colombia. It probably occurs more widely than current records
Country Distribution: Colombia, Ecuador suggest, especially in areas between known sites. Its altitudinal range is 2,100-3,500m asl.
Current Population Trend: Decreasing Population It is locally uncommon.
Habitat and Ecology It lives in very wet tropical cloud forest, and is usually found on low vegetation and in leaf-litter;
specimens have been collected along a roadside, on moss in rock crevices. It is found only in undisturbed habitats.
Breeding is by direct development.
Major Threats The major threat is habitat loss as a result of deforestation for agricultural development, the planting
of illegal crops, fire, logging activities, and human settlement. An additional threat is pollution resulting from the
spraying of illegal crops.
Conservation Measures The range of this species in Ecuador overlaps several protected areas, including the
© Luis A. Coloma

Reserva Ecológica Cayambe-Coca, the Reserva Ecológica Antisana and Parque Nacional Sumaco in Ecuador. It has
not been found in any protected areas in Colombia. Improved and expanded habitat protection is urgently needed to
ensure the persistence of this species in its montane forest habitat.
Notes on taxonomy: The Colombian populations have different skin texture and some morphological differences compared to Ecuadorian
populations. More research is needed to determine whether or not this is in fact two separate species (D.F. Cisneros-Heredia pers.
comm.).
Bibliography: Gluesenkamp, A.G. and Acosta, N. (2001), Hoogmoed, M.S. (1987), Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and Lynch,
J.D. (1996)
Data Providers: Luis A. Coloma, Santiago Ron, Diego Cisneros-Heredia, Ana Almandáriz, Wilmar Bolívar

EN Osornophryne percrassa Ruíz-Carranza and Hernández-Camacho, 1976 HERVEO PLUMP TOAD

Endangered B1ab(iii) Geographic Range This species is known from the central páramos in Tolima, Caldas and Quindio Departments,
Order, Family: Anura, Bufonidae on the eastern flank of the central Andes, in Colombia, between 2,700 and 3,700m asl.
Country Distribution: Colombia Population It is not a common species.
Current Population Trend: Decreasing Habitat and Ecology It occurs on leaf-litter and rocks on the ground in Andean forests and páramos, and has not
been recorded from anthropogenic habitats. It is a direct developing species.
Major Threats The species is threatened by habitat loss and fragmentation, mainly due to the expansion of
agriculture, and pollution from the fumigation of illegal crops. Climate change is also a potential threat to this
high-elevation species.
Conservation Measures Its range includes several protected areas, but improved and expanded protection of the
montane habitat of this species is necessary.
Bibliography: Acosta-Galvis, A.R. (2000), Ardila-Robayo, M.C. and Acosta-Galvis, A. (2000b), Gluesenkamp, A.G. (1995), Ruiz-Carranza,
© John D. Lynch

P.M. and Hernández-Camacho, J.A. (1976b)

Data Providers: Wilmar Bolívar, John Lynch

VU Osornophryne sumacoensis Gluesenkamp, 1995

Vulnerable D2 Geographic Range This species is known only from around a small Bibliography: Gluesenkamp, A.G. (1995)
Order, Family: Anura, Bufonidae crater lake on the eastern slopes of Volcán Sumaco, in Napo Province, Data Providers: Luis A. Coloma, Santiago Ron, Diego Cisneros-Heredia, Ana Almandáriz
Country Distribution: Ecuador Ecuador, at 2,500m asl.
Current Population Trend: Stable Population It is locally common.
Habitat and Ecology It is an inhabitant of the cloud forest sur-
rounding the lake. This forest is made up of bamboo (Chusquea
sp.), Ficus and other trees up to 20m tall. There is an abundance of
epiphytes in the area. It is terrestrial, both diurnal and nocturnal, and
specimens have been found under leaf-litter by day (Gluesenkamp
1995). It breeds by direct development, and lays its eggs on soil
under vegetation.
Major Threats There are no current major threats to the species,
although eruption of Volcán Sumaco poses a future potential threat
to its only known site.
Conservation Measures It is present in Parque Nacional Sumaco.
There is a need for close population monitoring of this species,
particularly given its restriction to a single location.

EN Osornophryne talipes Cannatella, 1986 CANNATELLA’S PLUMP TOAD

Endangered B1ab(iii) Geographic Range This species ranges from the type locality in Imbabura Province, Ecuador, north to Narino and Cauca
Order, Family: Anura, Bufonidae Departments in southern Colombia. It probably occurs a little more widely than current records suggest, especially in
Country Distribution: Colombia, Ecuador areas between known sites. The type locality is at 3,400m asl, and it occurs at similar altitudes in Colombia.
Current Population Trend: Decreasing Population It appears to be a rare species; however, this may be an artefact of inappropriate survey techniques.
For example, when digging into the soil under páramo plants, dozens of specimens have been found(M. Yanez and
D.F. Cisneros-Heredia pers. comm.).
Habitat and Ecology It has been found only in undisturbed habitats in dense upper montane forest, with trees
© Mario Humberto Yánez-Muñoz

reaching heights of as much as 10m, and in sub-páramo bushland. It breeds by direct development.
Major Threats The major threat is habitat loss as a result of deforestation for agricultural development, the plant-
ing of illegal crops, fire, logging, and human settlement. Pollution resulting from the spraying of illegal crops is an
additional threat.
Conservation Measures It is not known from any protected areas in Ecuador, but it occurs in Parque Nacional
Natural Purace in Colombia. It is recommended that the existing protected areas network be expanded to incorporate
remaining patches of high-elevation montane forest.
Bibliography: Cannatella, D.C. (1986), Lynch, J.D. (1981a), Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and Lynch, J.D. (1996)
Data Providers: Luis A. Coloma, Santiago Ron, Taran Grant, Diego Cisneros-Heredia, Ana Almandáriz
202 Threatened Amphibians of the World
EN Pedostibes tuberculosus Günther, 1875
Endangered B1ab(iii)
Order, Family: Anura, Bufonidae
Country Distribution: India
Current Population Trend: Decreasing
© S.U. Saravanakumar
EN Pelophryne albotaeniata Barbour, 1938
Endangered B1ab(iii) Geographic Range This species is found only on Palawan Island,
Order, Family: Anura, Bufonidae in the Philippines. The type locality was at an elevation of 1,500m
Country Distribution: Philippines asl.
Current Population Trend: Decreasing Population It is apparently rare, and there have been no field
observations of this species for more than 40 years, though this may
be an artefact of limited survey work.
Habitat and Ecology It inhabits arboreal microhabitats in riverine
habitat in montane and lowland forests. Its breeding biology
is unrecorded, but it presumably takes place in water by larval
development.
Major Threats The loss of lowland rainforest, mainly due to
agricultural activities and selective logging, is a major threat to
this species.
Conservation Measures Although the forests on Palawan are,
in general, reasonably well protected in much of the island, more
effective protection of the remaining rainforest on the island, espe-
EN Pelophryne api Dring, 1984
Endangered B1ab(iii)
Order, Family: Anura, Bufonidae
Country Distribution: Malaysia
Current Population Trend: Decreasing
© Indraneil Das
VU Pelophryne guentheri (Boulenger, 1882)
Vulnerable B1ab(iii) Geographic Range This species is known from south-western
Order, Family: Anura, Bufonidae and north-eastern Sarawak (Malaysia) and Serasan (north-western
Country Distribution: Indonesia, Malaysia Kalimantan, Indonesia), and might also occur in north-western Kali-
Current Population Trend: Decreasing mantan (Indonesia) and south-western Sabah (Malaysia), in Borneo.
It probably occurs more widely than current records suggest. It is a
lowland species present at elevations below 200m asl.
Population There is no information on the population status of
this species.
Habitat and Ecology It occurs in the leaf-litter of hilly lowland
primary moist forest. The large eggs are probably deposited in very
small rain pools.
Major Threats The major threat to this species is habitat loss due
to logging. The northern part of the range now consists of disturbed
habitat (largely deforested) and it is unclear whether or not the
species still survives there.
Geographic Range This species is endemic to the Western Ghats of India, where it has a very fragmented range.
It occurs at elevations of 300-1,800m asl.
Population There is no reliable information available on the population status of this species, although it is locally
uncommon and believed to be declining.
Habitat and Ecology It is a semi-arboreal species generally associated with montane moist evergreen forest.
Breeding takes place on the ground at stream banks, and the larvae develop in water.
Major Threats It is threatened by the conversion of its forest habitat to non-timber plantations (including coffee
and tea), the collection of timber and wood for subsistence use by local people, and the construction of roads and
dams.
Conservation Measures It has been recorded from several protected areas, including the Konya Wild Life Sanctu-
ary (Maharashtra), Cotigao Wild Life Sanctuary (Goa), Indira Ghandi National Park and Kalakad-Mundanthurai Tiger
Reserve (Tamil Nadu), Ponmudi Hills and Silent Valley National Park (Kerala). The species is the focus of ongoing
studies from 1998 to the present (S. D. Biju pers. comm.).
Bibliography: Biju, S.D. (2001), Chanda, S.K. (2002), Das, I. and Whittakar, R. (1988), Das, I. and Whittakar, R. (1990), Dutta, S.K. (1997),
Günther, A. (1876), Inger, R.F. et al. (1984), Pillai, R.S. (1986)
Data Providers: S.D. Biju, Sushil Dutta, Robert Inger, Vivek Ashok Gour-Broome
cially riverine habitats and gallery forests, is necessary. Further survey work is also needed to establish the current
population status of this species.
Bibliography: Alcala, A.C. and Brown, W.C. (1985), Brown, R.M., Diesmos, A.C. and Alcala, A.C. (2001), Frost, D.R. (1985), Inger, R.F.
(1954), Inger, R.F. (1960b), Inger, R.F. (1999), Taylor, E.H. (1923)
Data Providers: Arvin Diesmos, Angel Alcala, Rafe Brown, Leticia Afuang, Genevieve Gee, Katie Hampson, Mae Leonida Diesmos,
Aldrin Mallari, Perry Ong, Dondi Ubaldo, Baldwin Gutierrez
Geographic Range This species is endemic to Borneo where it is known only from the type locality (Gunung Mulu
National Park) in northern Sarawak, Malaysia; however, it might occur a little more widely. It has an altitudinal
range of 65-1,200m asl.
Population The population status and abundance of the species is not known.
Habitat and Ecology All known specimens have been found in primary rainforest in limestone karst areas. The
larvae are endotrophic and occur in small pools on the forest floor.
Major Threats It is not certain whether this species is entirely restricted to karst areas, but, if so, then its distribution
is likely to be extremely patchy and localized, thereby exposing this species to the hazards of stochastic events.
Conservation Measures The only known locality for the species is protected in Gunung Mulu National Park, a
52,000-ha park that is the most studied tropical karst area in the world. Further survey work is requried to determine
the current population status of the species at this locality, and whether it might be present in other karst areas
of Borneo.
Bibliography: Dring, J.C.M. (1984b), Dring, J.C.M. (1984b), Inger, R.F. and Stuebing, R.B. (1997)
Data Providers: Robert Inger, Indraneil Das, Robert Stuebing, Maklarin Lakim, Paul Yambun
Conservation Measures The species is present in Kubah National Park, although the southern section is facing
increasing pressure from encroaching logging activities. There is a need for improved habitat protection of sites at
which this species is known to occur.
Bibliography: Inger, R.F. (1966), Leong, T.M., Grismer, L. and Mumpuni (2002)
Data Providers: Robert Inger, Indraneil Das, Robert Stuebing, Maklarin Lakim, Paul Yambun
 Globally Threatened Amphibian Species 203

VU Pelophryne lighti (Taylor, 1920)

Vulnerable B1ab(iii) Geographic Range This species is patchily distributed in Bohol, Samar, Leyte and Mindanao in the Philippines. It
Order, Family: Anura, Bufonidae probably occurs a little more widely than current records suggest.
Country Distribution: Philippines Population It is locally common.
Current Population Trend: Decreasing Habitat and Ecology It inhabits arboreal microhabitats in riverine areas in montane and lowland forests, and is able
to survive in slightly disturbed habitats. It presumably breeds in water by larval development.
Major Threats The major threats to this species include the loss of lowland rainforest (as a result of agriculture and
logging), and the pollution of mountain streams and rivers due to agricultural effluents and mine tailings.
Conservation Measures This species is present in several protected areas. Conservation measures must include
the regulation and proper disposal of pesticides, herbicides, and mine tailings and the protection of the remaining
rainforest, especially riverine habitats and gallery forests.
Bibliography: Alcala, A.C. and Brown, W.C. (1985), Frost, D.R. (1985), Inger, R.F. (1960b), Inger, R.F. (1999), Taylor, E.H. (1921)
© Arvin C. Diesmos

Data Providers: Arvin Diesmos, Angel Alcala, Rafe Brown, Leticia Afuang, Genevieve Gee, Katie Hampson, Mae Leonida Diesmos,
Aldrin Mallari, Perry Ong, Dondi Ubaldo, Baldwin Gutierrez

VU Pelophryne misera (Mocquard, 1890)

Vulnerable B1ab(iii) Geographic Range This species is known only from mountains in Sabah and north-eastern Sarawak, in northern
Order, Family: Anura, Bufonidae Borneo, Malaysia. It might also occur in the mountains of northern Kalimantan, Indonesia, but has not yet been recorded
Country Distribution: Malaysia from there. It probably occurs more widely than current records suggest. It is present in forests above 1,500m asl.
Current Population Trend: Decreasing Population It is difficult to observe, but is considered locally abundant in montane forests (Malkmus et al. 2002).
Habitat and Ecology All known observations were made in montane ‘elfin’ type forests. Adults are largely terrestrial
and are found in leaf-litter, rock crevices, and holes in the ground. Breeding takes place in small, temporary pools on
the forest floor, and tadpoles are endotrophic.
Major Threats Habitat loss is probably the most serious threat, due mainly to logging. Logging is occurring at
© Andreas & Christel Nöllert

Gunung Murud, but is below the altitudinal range of this species. The development of recreation and tourism facili-
ties is a potential threat.
Conservation Measures The range of the species includes Kinabalu National Park. Other suitable montane forests,
particularly those of Trus Madi, require more effective protection.
Bibliography: Inger, R.F. (1966), Inger, R.F. and Stuebing, R.B. (1997), Malkmus, R. et al. (2002)
Data Providers: Robert Inger, Indraneil Das, Robert Stuebing, Maklarin Lakim, Paul Yambun

VU Pelophryne rhopophilius Inger and Stuebing, 1996

Vulnerable B1ab(iii) Geographic Range This species is known only from south-western and north-eastern Sarawak (Malaysia), but prob-
Order, Family: Anura, Bufonidae ably also occurs in western Kalimantan (Indonesia), in Borneo. It probably occurs more widely than current records
Country Distribution: Malaysia suggest. It is present at elevations around 800m asl.
Current Population Trend: Decreasing Population It is moderately abundant within its restricted range.
Habitat and Ecology Males have been seen calling from low shrubs in submontane and montane mossy forest.
Eggs are probably deposited in very small rain pools.
Major Threats The major threat to the species is ongoing habitat loss due to agriculture and logging.
Conservation Measures The species’ range includes Gunung Mulu National Park and Lanjak Entimau Wildlife
Sanctuary.
Bibliography: Inger, R.F. and Stuebing, R.B. (1996), Inger, R.F. and Stuebing, R.B. (1997)
Data Providers: Robert Inger, Indraneil Das, Robert Stuebing, Maklarin Lakim, Paul Yambun
© Djoko Iskandar

EN Pelophryne scalptus (Liu and Hu, 1973)

© Michael Wai Neng Lau / Kadoorie Farm & Botanic Garden

Endangered B1ab(iii) Geographic Range This species is restricted to the hilly areas in southern Hainan Province, China, from 350-
Order, Family: Anura, Bufonidae 1,400m asl.
Country Distribution: China Population There is no information available on its population status.
Current Population Trend: Decreasing Habitat and Ecology It inhabits evergreen broadleaf forests. Males call near small streams, which are probably
also the breeding sites.
Major Threats The species is particularly susceptible to habitat destruction and degradation that is taking place pri-
marily due to smallholder farming activities and small-scale wood extraction from expanding human settlements.
Conservation Measures The range of this species includes several protected areas, but expanded and improved
protection of the remaining forest habitat is necessary.
Bibliography: Fei, L. et al. (1999), MacKinnon, J. et al. (1996), Ye, C.-Y, Fei, L. and Hu, S.Q. (1993)
Data Providers: Michael Wai Neng Lau, Shi Haitao
204 Threatened Amphibians of the World

EN Rhamphophryne macrorhina Trueb, 1971

Endangered B1ab(iii)+2ab(iii) Geographic Range This species is known from the type locality: Santa Rita, in Antioquia Department, Colombia,
Order, Family: Anura, Bufonidae between 1,890 and 1910m asl. It is also known from two other locations in Colombia: Guatape (Mesopotamia) and
Country Distribution: Colombia Amalfi (Anori) all in Antioquia Department, in the central Andes, between 1,800 and 1,900m asl. There is no suitable

© Brian C. Bock, courtesy of Museo de Herpetología

Current Population Trend: Decreasing habitat available to the species between these known localities.
Population It is a common species.
Habitat and Ecology It occurs in the leaf-litter of sub-Andean and Andean forests, and is restricted to primary and
good secondary forest. It most likely breeds by direct development like other species in the genus.
Major Threats Habitat fragmentation and loss, due to the expansion of agriculture, timber extraction, desiccation,
and the fumigation of illegal crops, are all major threats to the species’ habitat. Climate change may also be a threat,
Universidad de Antioquia given that this species occurs at relatively high elevations.
Conservation Measures The range of the species includes the Reserva La Forzosa, but other tracts of montane
forest remain in urgent need of formal protection.
Bibliography: Acosta-Galvis, A.R. (2000), Grant, T. (1998), Grant, T. (1999), Trueb, L. (1971)
Data Providers: Wilmar Bolívar, John Lynch

EN Rhamphophryne nicefori (Cochran and Goin, 1970)

Endangered B1ab(iii)+2ab(iii) Geographic Range This species is known from the type locality, “El Chaquiro, [department of] Antioquia, Colombia”
Order, Family: Anura, Bufonidae (amended to “Hacienda Palmas, El Chaquiro, Antioquia Department, Colombia, 2,670m elevation”), and from nearby
Country Distribution: Colombia locations.
© Juan Camilo Arredondo-S, courtesy of Museo de

Current Population Trend: Decreasing Population It is a common species within its relatively small range.
Habitat and Ecology It occurs in high-altitude grassland, and has not been recorded from anthropogenic habitats.
It breeds by direct development.
Hespetología Universidad de Antioquia

Major Threats The main threat is habitat loss due to expanding pastures for cattle grazing. Climate change may
be a threat in the future.
Conservation Measures The range of this species does not include any protected areas, such that habitat protec-
tion remains the most urgent conservation action required. Further survey work is needed to determine whether it
occurs outside the vicinity of the type locality.
Bibliography: Cochran, D.M. and Goin, C.J. (1970), Trueb, L. (1971)
Data Providers: Wilmar Bolívar, John Lynch

CR Rhamphophryne rostrata (Noble, 1920)

Critically Endangered B1ab(iii)+2ab(iii) Geographic Range This species is known only from the type Conservation Measures The type locality is not within a protected area. Further survey work is needed to establish
Order, Family: Anura, Bufonidae locality: “Santa Rita Creek, fourteen miles north of the village of whether or not the species still occurs at the type locality or in any location outside the type locality, as the species
Country Distribution: Colombia Mesopotamia in the southern part of the Department of Antioquia, might already be extinct in view of the scale of habitat destruction in the general vicinity.
Current Population Trend: Decreasing Colombia”, at 2,472m asl. It is unlikely to range more widely. Bibliography: Acosta-Galvis, A.R. (2000), Cochran, D.M. and Goin, C.J. (1970), Grant, T. (1999), Graybeal, A. and Cannatella, D.C. (1995),
Population This species is known from only two specimens collected Lynch, J.D. and Renjifo, J.M. (1990), Noble, G.K. (1920), Rivero, J.A. and Castaño, C. (1990), Ruiz-Carranza, P.M., Ardila-Robayo, M.C.
in 1914. Searches of the type locality since then have not found it, and Lynch, J.D. (1996), Trueb, L. (1971)
and it is possible that it is now extinct. Data Providers: Wilmar Bolívar, John Lynch
Habitat and Ecology The type locality was forest when the
specimens were collected, but the exact habitat requirements of the
species are still unknown. It presumably breeds by direct develop-
ment like other species in the genus.
Major Threats At the type locality there has been significant logging
in the past, and there is now high human population density in this
area with accompanying increased infrastructure development for
human settlement. Agriculture, including the planting of illegal crops,
is also a threat in the area as well as fumigation of illegal crops.

VU Spinophrynoides osgoodi (Loveridge, 1932)

Vulnerable B1ab(iii)+2ab(iii) Geographic Range This species is endemic to the mountains of south-central Ethiopia (Arsi, Balé, Sidamo and
Order, Family: Anura, Bufonidae Gamo Gofa Provinces) at 1,950-3,520m asl. With the exception of the population in the Gughe Mountains, all records
Country Distribution: Ethiopia are from east of the Rift Valley.
Current Population Trend: Decreasing Population It is locally common within its limited range.
CITES: Appendix I Habitat and Ecology It is essentially a species of montane forest, though perhaps extending marginally into
open moorland. It has been observed breeding in a small, probably temporary, pool in a grassy glade surrounded by
Hypericum woodland.
Major Threats It is threatened by environmental degradation resulting from human settlement, specifically the
destruction of forests through both subsistence and commercial exploitation.
Conservation Measures The species is relatively common in the Bale Mountains National Park, but there is a need
for improved protection of other sites at which this species has been recorded.
Bibliography: Grandison, A.G.C. (1978), Largen, M.J. (2001)
Data Providers: Malcolm Largen
© M.J. Largen
 Globally Threatened Amphibian Species 205

EN Stephopaedes anotis (Boulenger, 1907) CHIRINDA TOAD

Endangered B1ab(iii) Geographic Range This species is known only from the Chirinda Forest in eastern Zimbabwe and the Dombe Forest
Order, Family: Anura, Bufonidae in adjacent Mozambique. In the Chirinda Forest it occurs at approximately 900-1,300m asl.
Country Distribution: Mozambique, Zimbabwe Population It is apparently common in its small range.
Current Population Trend: Decreasing Habitat and Ecology It lives in the leaf-litter of evergreen forest, often hiding inside or beneath rotten logs. The
eggs are laid in pockets of water between the buttress roots of Chrysophyllum gorungosanum trees, or in water-filled
grooves on the trunks of fallen trees. It is not found outside forest.
Major Threats Its forest habitat is threatened by agriculture, wood extraction, and human settlement.
Conservation Measures It occurs in the Gungunyana Forest Reserve in Zimbabwe. The Chirinda Forest is admin-
istered by the Forestry Commission and is one of the best researched forests in Zimbabwe, but the area remains in
need of strengthened protection.
Bibliography: Channing, A. (1978), Channing, A. (1993), Channing, A. (2001), Poynton, J.C. (1964b), Poynton, J.C. and Broadley, D.G.
© Richard Boycott

(1988), Poynton, J.C. and Clarke, B.T. (1999), Tandy, M. and Keith, R. (1972), Taylor, P. (1973)
Data Providers: John Poynton, Alan Channing

EN Stephopaedes howelli Poynton and Clarke, 1999

Endangered B1ab(iii) Geographic Range This species, which is endemic to Tanzania, is known only from the Mrora Forest on Mafia
Order, Family: Anura, Bufonidae Island, and from the Jozani Forest on Zanzibar (and from an unmapped, unspecified locality on the east of Zanzibar).
Country Distribution: Tanzania All records are from close to sea level.
Current Population Trend: Decreasing Population It is probably an uncommon species.
Habitat and Ecology It is terrestrial, occurring in lowland coastal forest. Some of its remaining habitat is degraded,
and it seems to display some adaptability to living in such habitats. Its breeding behaviour is unknown, but it might
breed in puddles in tree roots like Stephopaedes anotis.
Major Threats Its habitat is being degraded rapidly on Mafia for agriculture, wood extraction, and human settle-
ments.
Conservation Measures It possibly occurs in the Mafia Marine Park, but this remains to be confirmed, and improved
protection of remaining suitable habitat on Mafia is a priority. On Zanzibar, the Jozani Forest, the largest tract of forest
© Alan Channing

remaining on the main island, has recently had its protection status upgraded to that of a national park.
Bibliography: Howell, K.M. (1993), Poynton, J.C. (1998), Poynton, J.C. (2003b), Poynton, J.C. and Clarke, B.T. (1999)
Data Providers: John Poynton, Kim Howell

EN Stephopaedes usambarae Poynton and Clarke, 1999

Endangered B1ab(iii) Geographic Range This species is known only from the foothills of
Order, Family: Anura, Bufonidae the East Usambara Mountains in north-eastern Tanzania. All records
Country Distribution: Tanzania have been from below 410m asl.
Current Population Trend: Decreasing Population It is not uncommon within its very small known range.
Habitat and Ecology It is a terrestrial species of lowland forest,
but also survives in mildly disturbed, selectively logged forest. Its
breeding behaviour is unknown, but it might breed in puddles in tree
roots (like Stephopaedes anotis).
Major Threats There is ongoing loss of its habitat due to agricultural
encroachment, wood extraction, and human settlement.
Conservation Measures It occurs in the Kwamgumi, Segoma and
Mtai Forest Reserves, but these are not well protected and are in
need of improved management.
Bibliography: Harper, E. and Vonesh, J.R. (2003), Poynton, J.C. (1998), Poynton,
J.C. and Clarke, B.T. (1999)
Data Providers: John Poynton, Kim Howell

EN Werneria bambutensis (Amiet, 1972)

Endangered B1ab(iii)+2ab(iii) Geographic Range This species is known only from the mountains of western Cameroon, where it occurs on Mount
Order, Family: Anura, Bufonidae Manenguba and in the Bamenda Highlands in the Bamboutos Mountains and Mount Oku. It also occurs at Santa on
Country Distribution: Cameroon Mount Neshele. Its altitudinal range is from 1,750-2,600m asl, but it rarely occurs below 2,100-2,200m asl.
Current Population Trend: Decreasing Population It is not a common species. In 1985, it was reported as possibly nearing extinction in the Bamboutos
Mountains.
Habitat and Ecology It is found on flat rocks in fast-flowing streams at high elevations, typically in montane forest
patches. It breeds in streams, and it seems that after reproduction the adults disperse into forest patches, open bamboo
glades, and montane grassland. The larvae live in fast, even torrential, water. During the dry season the adults may
live in water, hiding under stones. It seems to occur at higher altitudes than other members of its genus, living at a
higher altitude than both Werneria tandyi and W. mertensiana on Mount Manenguba.
© David C. Blackburn

Major Threats It is probably threatened by forest clearance and degradation resulting from overgrazing, fires, and
cultivation.
Conservation Measures The species might occur in the Bafut-Ngemba Forest Reserve, but is largely unprotected
elsewhere in its range; additional protection of the remaining highland forests in Cameroon, particularly Mount
Manenguba, is needed. A conservation project has been conducted on Mount Oku for several years by BirdLife
International, involving community management of the area involving the local villages. This project needs to take
into account the conservation needs of this species.
Bibliography: Amiet, J.-L. (1972b), Amiet, J.-L. (1976b), Amiet, J.-L. (1989), Gartshore, M.E. (1986), Rödel, M.-O. et al. (2004)
Data Providers: Jean-Louis Amiet
206 Threatened Amphibians of the World

CR Werneria iboundji Rödel, Schmitz, Pauwels and Böhme, 2004

Critically Endangered B1ab(iii)+2ab(iii)
Order, Family: Anura, Bufonidae
Country Distribution: Gabon
Current Population Trend: Decreasing
Geographic Range This species is known only from two specimens temperature, humidity, and available food, and so have serious consequences for this species which, like its congeners,
from the type locality at 560m asl on the east flank of Mount Iboundji, is dependent on high humidity.
Massif du Chaillu, Offoué-Onoy Department, Ogooué-Lolo Province, Conservation Measures Mount Iboundji has been proposed as a Biodiversity Sanctuary, due to its botanical and
Gabon. Searches at a second waterfall on Mount Iboundji and at herpetological values, but no protection status has been granted to date.
many other waterfalls in Gabon, revealed no further records of this Bibliography: Rödel, M.-O. et al. (2004)
species. However, it is possible that it occurs at other waterfalls, Data Providers: Olivier Pauwels
particularly in the Chaillu Massif, although it must be localized.
Population Unlike most congeners, which are often locally very
abundant, this species appears to be very rare; only two individuals
could be located in 20 person-hours of searching.
Habitat and Ecology This species is known only from among rocks
at the edge of a plunge pool at the base of a large waterfall in lowland
forest. It is presumed to be a larval developer.
Major Threats The main threat to the species is logging, which
is ongoing, but has not yet reached the waterfall basin where the
species occurs. Even if logging occurs nearby, it is likely to modify

EN Werneria mertensiana (Amiet, 1976)

Endangered B2ab(iii) Geographic Range This species is known from Cameroon where it has been recorded from Mounts Manenguba,
Order, Family: Anura, Bufonidae Nlonako, Kupe and Nta Ali, and at Fotabong in western Cameroon, and on the mountains around Yaoundé (Kala and
Country Distribution: Cameroon Mbam-Minkoum) in southern Cameroon. There is also one specimen recorded from Equatorial Guinea on Monte
Current Population Trend: Decreasing Alén, although this may be another species and requires confirmation. Its altitudinal range is from 800-1,050m asl.
It probably also occurs on the southern and western slopes of the Bamileke Plateau, and perhaps also on the Obudu
Plateau in eastern Nigeria.
Population It has been found to be reasonably common in suitable habitat within its small range.
Habitat and Ecology It is associated with rocks in streams and waterfalls in forest and degraded secondary habi-
tats at the lower limit of the submontane zone, although it has also been found in leaf-litter away from water. On
Mount Manenguba it occurs at a lower altitude than Werneria tandyi, although their ranges might overlap slightly.
© Mary E. Gartshore

It breeds in streams.
Major Threats This species is presumably affected by habitat loss due to agricultural encroachment and human
settlement.
Conservation Measures In Cameroon, the species occurs in a few forest reserves, such as Nta Ali Forest Reserve;
however, the higher elevation forest habitats in western Cameroon, particularly Mount Manenguba, are in urgent
need of improved protection. The specimen recorded from Equatorial Guinea was found in Monte Alén National Park
in Equatorial Guinea, although confirmation of the taxonomic identity of this population is needed.
Bibliography: Amiet, J.-L. (1972b), Amiet, J.-L. (1976b), De la Riva, I. (1994b), Gartshore, M.E. (1986), Herrmann, H.-W. et al. (2005),
Lasso, C.A. et al. (2002), Lawson, D.P. (1993), Rödel, M.-O. et al. (2004)
Data Providers: Jean-Louis Amiet

EN Werneria preussi (Matschie, 1893)

Endangered B1ab(iii) Geographic Range This species is known from the lower slopes of Mount Cameroon at 700-1,200m asl, from Mount
Order, Family: Anura, Bufonidae Kupe around 900m asl, as well as a highly disjunct population from the highlands of west-central Togo (“Bismarkberg”),
Country Distribution: Cameroon, Togo although this record may be in error. There have been no records from Togo for more than 100 years.It possibly has
Current Population Trend: Decreasing a wider range across the Cameroonian highlands than is currently known.
Population It is common, perhaps even abundant, in suitable habitat on Mount Cameroon. A survey in eastern Ghana
in 2001 very close to the Togo highlands failed to recover this species.
Habitat and Ecology It is associated with rocky steams and waterfalls in submontane forest and degraded secondary
habitats. The lack of permanent streams at high elevations on Mount Cameroon probably imposes an altitudinal limit
on the species in this part of its range, since it breeds in streams and lives exclusively in and around water.
Major Threats It is presumably at risk from forest loss due to agricultural encroachment and human settlement.
Conservation Measures It does not occur in any protected areas, and protection of remaining forest habitat on
© Wolfgang Böhme

Mount Cameroon and Mount Kupe is urgently needed.An ecotourism project has been established at Mount Kupe
for some time, which aims to reduce hunting and agricultural expansion, and the area has been proposed as a Strict
Nature Reserve (Réserve Écologique Intégrale). Further survey work is needed to confirm the existence and status
of the population in Togo.
Notes on taxonomy: Studies are needed to determine whether or not specimens from Cameroon and Togo belong to the same species
(M.-O. Rödel pers. comm.).
Bibliography: Amiet, J.-L. (1971b), Amiet, J.-L. (1972b), Amiet, J.-L. (1976b), Amiet, J.-L. (1987), Grandison, A.G.C. (1981), Rödel, M.-O.
et al. (2004), Rödel, M.-O. and Agyei, A.C. (2003)
Data Providers: Jean-Louis Amiet, Mark-Oliver Rödel

EN Werneria submontana Rödel, Schmitz, Pauwels and Böhme, 2004

Endangered B1ab(iii) Geographic Range This species is known from Nyasoso on the south-west side of Mount Kupe at 910m asl, and from
Order, Family: Anura, Bufonidae 800-1,200m asl in the Bakossi Mountains (including the Mwendelengo Mountains), Cameroon. A specimen collected
Country Distribution: Cameroon in the vicinity of Nkongsamba (Maholé, 10km north-west of Tombel, Bakossi Forest Reserve, at 300-350m asl), may
Current Population Trend: Decreasing also belong to this species, as may specimens from Mofako in the Rumpi Hills. However, it is not believed likely to
occur much more widely, given its specific habitat preferences and lack of habitat in the general area.
Population This species is often locally abundant.
Habitat and Ecology On Mount Kupe, this species was found in a transition zone between good quality secondary
forest and undisturbed primary forest. The species has been found during the day along a stream, under rocks in
a partly dried-up river basin, and on stony ground between wet, very low vegetation in the spray zone of a small,
artificial waterfall. It is presumed to be a larval developer.
Major Threats The habitat of this species is being steadily deforested for cultivation (particularly since human popu-
© Andreas Schmitz

lations in the area are growing quickly) and several logging companies hope to start large-scale logging operations
soon in the Bakossi/Mwendelengo Mountains. Even if deforestation does not eliminate the habitat of this species,
it is likely to significantly alter temperature, humidity, and available food, and so have serious consequences for this
species which, like its congeners, is dependent on high humidity. Near Nyassosso, the household use of detergents
in rivers is also a potential threat.
Conservation Measures An ecotourism project has been established at Mount Kupe for some time, and aims to
reduce hunting and agricultural expansion. In addition, a Strict Nature Reserve (Réserve Écologique Intégrale) has
been proposed for the area. A management plan has been in preparation with the aim of designating most of the
Bakossi Mountains as Protection Forest (55,000ha).
Bibliography: Rödel, M.-O. et al. (2004)
Data Providers: Mark-Oliver Rödel, Andreas Schmitz
 Globally Threatened Amphibian Species 207

EN Werneria tandyi (Amiet, 1972)

Endangered B2ab(iii) Geographic Range This species is known only from Mount Manenguba and the Rumpi Hills in western Cameroon,
Order, Family: Anura, Bufonidae though its range is not well understood. Its altitudinal range on Mount Manenguba is from 1,300-1,750m asl, and
Country Distribution: Cameroon the single specimen from the Rumpi Hills was collected at 1,000m asl.
Current Population Trend: Decreasing Population It is common within its limited range.
Habitat and Ecology It lives on the verges of fast-flowing streams in submontane forest and degraded secondary
habitats, where several individuals may cluster together on rocks in the splash zone. On Mount Manenguba it lives at
a higher altitude than Werneria merteniana, but at lower elevations than W. bambutensis. It breeds in streams.
Major Threats It is presumably at risk from forest loss due to agricultural encroachment and human settlement.
Conservation Measures It may occur in the Rumpi Hills Forest Reserve, although this is not managed for biodiversity
© David C. Blackburn conservation; increased protection of the montane forest habitats in western Cameroon is rather urgently needed,
particularly on Mount Manenguba.
Bibliography: Amiet, J.-L. (1972b), Amiet, J.-L. (1976b), Gartshore, M.E. (1986), Rödel, M.-O. et al. (2004)
Data Providers: Jean-Louis Amiet

CR Wolterstorffina chirioi Boistel and Amiet, 2001

Critically Endangered B1ab(iii)+2ab(iii) Geographic Range This recently described species is known only from the summit of Mount Oku at 3,000m asl in
Order, Family: Anura, Bufonidae the Bamenda Highlands in western Cameroon. In view of the absence of suitable vegetation at other localities in
Country Distribution: Cameroon Cameroon, this species is probably endemic to this area.
Current Population Trend: Decreasing Population The population status of this species is unknown.
Habitat and Ecology It lives in Afro-Alpine vegetation and grassland at the summit of Mount Oku. Its breeding
habits are unknown, but since it lives above the water limit on Mount Oku, it is presumably either a live-bearer or a
direct-developer laying eggs on the ground.
Major Threats The habitat at the summit of Mount Oku is threatened by overgrazing and fire. Because it is a high-
altitude species with a very limited range, it is potentially affected by climate change.
Conservation Measures A conservation project has been conducted on Mount Oku for several years by BirdLife
© David C. Blackburn

International, involving community management of the area by the local villages. This project needs to take into
account the conservation needs of this species. A captive-breeding programme should be considered in view of pos-
sible effects of climate change. Survey work is necessary to determine the current population status and breeding
habits of this species.
Bibliography: Boistel, R. and Amiet, J.-L. (2001)
Data Providers: Jean-Louis Amiet

EN Wolterstorffina mirei (Perret, 1971)

Endangered B1ab(iii)+2ab(iii) Geographic Range This species is known from western Cameroon on Mount Oku (at 2,500m asl) and Mount Meletan
Order, Family: Anura, Bufonidae at 2,200-2,700m asl in the Bamboutos Mountains. Both localities are within the Bamenda Highlands. There is some
Country Distribution: Cameroon doubt as to the validity of the record from Mount Oku (J.L. Amiet pers. comm.).
Current Population Trend: Decreasing Population It was reported to be abundant on Mount Meletan.
Habitat and Ecology This is a species of montane grassland that retreats to areas near streams and small watercours-
es in bamboo forest in the dry season. Its breeding habits are unknown, though it is likely to breed in streams.
Major Threats This species is almost certainly at risk from fire, overgrazing, and forest loss due to agriculture.
Habitat damage is very severe in the Bamboutos Mountains.
Conservation Measures A conservation project has been conducted on Mount Oku for several years by BirdLife
International, involving community management of the area by the local villages. This project needs to take into
© David C. Blackburn

account the conservation needs of this species (assuming that it occurs there).
Bibliography: Boistel, R. and Amiet, J.-L. (2001), Perret, J.-L. (1971a), Perret, J.-L. (1972)
Data Providers: Jean-Louis Amiet

VU Wolterstorffina parvipalmata (Werner, 1898)

Vulnerable B1ab(iii) Geographic Range This species is known from eastern Nigeria on the Obudu Plateau and the higher parts of the
Order, Family: Anura, Bufonidae Oban Hills, and from a number of montane localities in western Cameroon, including Mount Cameroon, the Rumpi
Country Distribution: Cameroon, Nigeria Hills, Mount Kupe, Mount Manenguba, the Mbos Cliffs, and Foto. It is also known from Mount Kala and Mount
Current Population Trend: Decreasing Mbam-Minkoum near Yaoundé in southern Cameroon. Its altitudinal range is 800-2,000m asl.
Population It is generally uncommon.
Habitat and Ecology It is confined to montane forest near streams and small waterfalls, and requires forest with a
closed canopy. It moves further from watercourses during the rainy season. It presumably normally breeds in streams,
though tadpoles have been found in a water tank.
Major Threats The main threat is habitat loss, primarily due to smallholder farming activities, which is continuing
over much of its range.
© Mary E. Gartshore

Conservation Measures It occurs in the Cross River National Park in Nigeria, but there is a need for improved
protection of other sites at which the species is known to occur.
Bibliography: Amiet, J.-L. (1971b), Amiet, J.-L. (1987), Amiet, J.-L. and Perret, J.-L. (1969), Boistel, R. and Amiet, J.-L. (2001), Herrmann,
H.-W. et al. (2005), Perret, J.-L. (1971a), Perret, J.-L. (1972)
Data Providers: Jean-Louis Amiet, Mary Gartshore
208 Threatened Amphibians of the World

CENTROLENIDAE

EN Centrolene audax (Lynch and Duellman, 1973)

Endangered B1ab(iii)+2ab(iii) Geographic Range This species is known from four localities on the Amazonian slopes of the Andes in Río Salado
Order, Family: Anura, Centrolenidae and the Cascada de San Rafael, Napo Province, Ecuador, at altitudes of 1,300-1,700m asl.
Country Distribution: Ecuador Population It is known to be rare; it was last recorded in 1979 and searches in 1999-2001 failed to turn up any
Current Population Trend: Decreasing individuals (Bustamante 2002). However, the sampling effort throughout its range has probably been insufficient.
Habitat and Ecology It lives in cloud forest. The holotype was found in a bromeliad in the spray zone of a waterfall
and on a cliff (Lynch and Duellman 1973). It lays its eggs on vegetation, and the larvae develop in water.
Major Threats Habitat destruction and degradation, as a result of agriculture and logging, are the main threats to
© Mario Humberto Yánez-Muñoz

this species. It is possible that chytridiomycosis may also have contributed to the decline.
Conservation Measures Its distribution may overlap marginally with the Reserva Ecológica Cayambe-Coca. Im-
proved protection of the montane forest habitat in the range of this species is necessary. Given the possible threat
of chytridiomycosis, surveys are needed to determine this species’ current population status.
Bibliography: Bustamante, M. R. (2002), Lynch, J.D. and Duellman, W.E. (1973), Ruiz-Carranza, P.M. and Lynch, J.D. (1991a)
Data Providers: Luis A. Coloma, Santiago Ron, Diego Cisneros-Heredia

EN Centrolene azulae (Flores and McDiarmid, 1989)

Endangered B1ab(iii)+2ab(iii) Geographic Range This species is known only from Fundo Azul. Further research into the distribution range and population status of this little-known species is needed, while
Order, Family: Anura, Centrolenidae Nuevo Mundo (at 1,500m asl), a cloud forest of the Cordillera Azul the potential impacts of climate change and possible infection with the chytrid fungus require investigation.
Country Distribution: Peru in Huanuco Department, central Peru. It probably occurs a little Bibliography: Alverson, W.S., Rodriguez, L.O. and Moskovits, D.K. (2001), Duellman, W.E. and Schülte, R. (1993), Flores, G. and McDiarmid,
Current Population Trend: Decreasing more widely than is currently known, but it is believed to have a R.W. (1989), Rodríguez, L.O., Cordova, J.H. and Icochea, J. (1993)
restricted range. Data Providers: Lily Rodríguez, Jorge Luis Martinez, Daniel Neira, Wilfredo Arizabal, Edgar Lehr, César Aguilar Puntriano
Population It is rare and poorly known.
Habitat and Ecology It is associated with stream habitats in
montane tropical forest, where the trees are generally widely
spaced and separated by 2-3m tall shrubland. Within this there are
also wet depressions and small ravines, with slightly taller (4-9m)
elfin forest. Eggs are deposited on leaves above streams, and larvae
develop in the streams.
Major Threats The principal threat to this species is habitat loss
due to agricultural expansion from smallholder farming and human
settlement.
Conservation Measures It may occur in Parque Nacional Cordillera

CR Centrolene ballux (Duellman and Burrowes, 1989)

Critically Endangered A2ac; B2ab(iii,iv,v) Geographic Range This species is known from three nearby localities in Saloya River Valley in Ecuador (Pichincha
Order, Family: Anura, Centrolenidae Province), at 1,700-2,010m asl, and from the Pacific versant of Colombia, in the Reserva Natural La Planada, 1,780m
Country Distribution: Colombia, Ecuador asl, Nariño Department.
Current Population Trend: Decreasing Population In Ecuador, the most recent record is from 1989, and the species is no longer present in the Saloya Valley,
and is apparently not in other nearby valleys. It has apparently declined seriously. In Colombia, many specimens were
collected when it was first recorded, but there have been no recent surveys.
Habitat and Ecology It lives on vegetation next to streams in humid upper montane forest. It breeds in streams and
is probably not tolerant of degraded habitats.
Major Threats The most likely cause of the severe decline of this species is the movement of the cloud layer up the
mountain sides as a result of climate change, resulting in reduced humidity within the altitudinal range of the spe-
© Ignacio De La Riva

cies (probably exacerbated by habitat fragmentation). Additional likely threats include: deforestation for agricultural
development (including illegal crops), fire, logging, and human settlement; introduction of alien predatory fish species
in streams; and pollution resulting from the spraying of illegal crops. Chytridiomycosis also cannot be ruled out.
Conservation Measures It occurs in the privately owned Reserva La Planada in Colombia, and could be in the
binational reserve near Planada. Surveys are urgently needed to determine whether or not this species still survives,
and, if necessary, an ex-situ captive population should be established.
Bibliography: Acosta-Galvis, A.R. (2000), Duellman, W.E. and Burrowes, P.A. (1989), Marquez, R., De la Riva, I. and Bosch, J. (1996),
Ruiz-Carranza, P.M. and Lynch, J.D. (1991a), Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and Lynch, J.D. (1996)
Data Providers: Wilmar Bolívar, Luis A. Coloma, Santiago Ron, Diego Cisneros-Heredia, Erik Wild

EN Centrolene fernandoi Duellman and Schulte, 1993

Endangered B1ab(iii)+2ab(iii) Geographic Range This species is currently known only from the type locality on the west slopes of Abra Tangarana
Order, Family: Anura, Centrolenidae (1,080m asl) north-east of the San Juan de Pacaysapa, Departemento San Martin, in northern Peru. It is likely to occur
Country Distribution: Peru more widely, but is believed to have a restricted distribution.
© Natural History Museum and Biodiversity Research

Current Population Trend: Decreasing Population It is an uncommon species.

Habitat and Ecology The area in which this species occurs supports lower montane rainforest and cloud forest,
whereas the intervening valleys are extensively cultivated. All individuals of this arboreal species were found on
the upper surface of trees 1.5-2.0m above a small stream in a narrow ravine at night. Eggs are laid on leaves, and
larvae develop in streams This species needs the specific narrow microhabitat of flat damp areas with a constant
source of water.
Center, University of Kansas

Major Threats The main threats to the species are unclear, but habitat loss, as a result of the expansion of agricultural
activities, wood extraction, and human settlement, is likely to be the main threat.
Conservation Measures The species is not present in any protected areas, and some form of formal habitat protec-
tion is urgently needed along the west slopes of Abra Tangarana.
Bibliography: Duellman, W.E. and Schülte, R. (1993), Rodríguez, L.O., Cordova, J.H. and Icochea, J. (1993)
Data Providers: Lily Rodríguez, Jorge Luis Martinez, Javier Icochea, Ariadne Angulo, Edgar Lehr, Daniel Neira, César Aguilar
Puntriano
 Globally Threatened Amphibian Species 209

VU Centrolene geckoideum Jiménez de la Espada, 1872

Vulnerable A3c; B1ab(iii) Population It is localized, but can be conspicuous in its microhabitat. In Ecuador, the population at Quebrada Zapadores
Order, Family: Anura, Centrolenidae (Pichincha Province) appears to have disappeared.
Country Distribution: Colombia, Ecuador Habitat and Ecology This species inhabits cloud forest, where it can be found on vegetation next to running water or
Current Population Trend: Decreasing on rocks in streams and behind waterfalls. The eggs are placed on boulders in the splash zone of fast-flowing streams
and waterfalls. It is a very territorial species, and the males guard the eggs. It is very susceptible to deforestation,
and does not survive in degraded habitats.
Major Threats The major threat is habitat loss and deforestation, as a result of agricultural development (particularly
the planting of illegal crops), logging, and human settlement. Other threats include the introduction of alien predatory

© Francisco José López-López

Geographic Range This species occurs in all three Cordilleras of Colombia from Antioquia, Caldas and Boyaca
Departments, south to the north-western Andean slopes of Ecuador in Carchi and Pichincha Provinces. It has been
recorded between 1,750 and 2,500m asl.
fish in streams, and pollution resulting from the spraying of illegal crops. Like some other species in its family, it might
also be affected by the movement of the cloud layer up the mountain sides as a result of climate change, resulting
in reduced humidity within the altitudinal range of the species (probably exacerbated by habitat fragmentation).
Chytridiomycosis also cannot be ruled out as a potential threat.
Conservation Measures It occurs in several protected areas in Colombia, but none in Ecuador. Further introduc-
tions of predatory alien fish should be prevented. The species is in need of close population monitoring given the
potential threat of chytridiomycosis.
Notes on taxonomy: In spite of its Amazonian type locality, this species is in fact known only from the Andes.
Bibliography: Acosta-Galvis, A.R. (2000), Goin, C.J. (1964), Grant, T., Bolivar-G., W. and Castro, F. (1998), Jiménez de la Espada, M.
(1872), Lynch, J.D., Ruiz-Carranza, P.M. and Rueda-Almonacid, J.V. (1983), Rueda-Almonacid, J.V. (1994a), Ruiz-Carranza, P.M. and Lynch,
J.D. (1991a), Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and Lynch, J.D. (1996)
Data Providers: Wilmar Bolívar, Luis A. Coloma, Santiago Ron, Diego Cisneros-Heredia, Erik Wild, Mario Yánez-Muñoz

CR Centrolene gemmatum (Flores, 1985)

Critically Endangered B2ab(iii) Geographic Range This species is known only from the type locality: San Francisco de Las Pampas, Cotopaxi
Order, Family: Anura, Centrolenidae Province, Ecuador, at 1,800m asl.
Country Distribution: Ecuador Population It has not been found since it was first described in the early 1980s. Intensive searching at Bosque
Current Population Trend: Decreasing Integral Otonga (near the type locality) has not turned up any individuals. However, the type region is large, and the
species might have been overlooked.
Habitat and Ecology It is found near streams in cloud forest. Reproduction is presumed to be via eggs laid on
vegetation, with the tadpoles developing in water.
Major Threats Habitat destruction and degradation, due to small-scale agriculture and subsistence wood collection,
is the primary threat to this species, but it might also have been impacted by chytridiomycosis.
Conservation Measures The species is not known to occur in any protected areas. Further survey work is required
to determine whether or not this species still survives.
© Luis A. Coloma

Bibliography: Flores, G. (1985a), Ruiz-Carranza, P.M. and Lynch, J.D. (1991a)

Data Providers: Luis A. Coloma, Santiago Ron

CR Centrolene heloderma (Duellman, 1981)

Critically Endangered A2ac Geographic Range This species occurs on the western versant of the Cordillera Occidental in Colombia in Cauca,
Order, Family: Anura, Centrolenidae Valle del Cauca and Risaralda Departments, south to the Tandayapa and Saloya Valleys, in Pichincha Province, Ecuador
Country Distribution: Colombia, Ecuador (where it has been recorded from four localities). Its altitudinal range is 1,900-2,400m asl.
© Natural History Museum and Biodiversity Research

Current Population Trend: Decreasing Population It has disappeared from all known localities in Ecuador, the most recent record dating back to March
1979, although they might occur elsewhere in this country. In Colombia, it is very rare, and was last recorded in 1996.
Overall, it appears to have undergone a serious decline.
Habitat and Ecology It inhabits the upper elevations of cloud forest, surviving only in mature forest. It breeds in
streams, with the eggs laid on leaves overhanging the water.
Major Threats The most likely cause of the severe decline of this species is the movement of the cloud layer up the
Center, University of Kansas

mountain sides as a result of climate change, resulting in reduced humidity within the altitudinal range of the spe-
cies (probably exacerbated by habitat fragmentation). Additional likely threats include: deforestation for agricultural
development (including illegal crops), fire, logging, and human settlement; introduction of alien predatory fish species
in streams; and pollution resulting from the spraying of illegal crops. Chytridiomycosis also cannot be ruled out.
Conservation Measures This species probably has been recorded in several protected areas in Colombia, but is
not recorded from any in Ecuador. Surveys are urgently needed to determine whether or not this species still survives,
and, if necessary, an ex-situ captive population should be established.
Bibliography: Duellman, W.E. (1981), Ruiz-Carranza, P.M. and Lynch, J.D. (1991a), Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and Lynch,
J.D. (1996)
Data Providers: Luis A. Coloma, Santiago Ron, Diego Cisneros-Heredia, Taran Grant, Wilmar Bolívar

VU Centrolene hesperium (Cadle and McDiarmid, 1990)

Vulnerable B1ab(iii) Geographic Range This species is known only from forested probably susceptible to localized habitat loss as a result of livestock farming and selective wood extraction. It might
Order, Family: Anura, Centrolenidae slopes at elevations of 1,500-1,800m asl on the Pacific versant of also be vulnerable to the effects of localized climate change and infection with the chytrid fungus.
Country Distribution: Peru the Cordillera Central in the department of Cajamarca in northern Conservation Measures It is not known to occur in any protected areas, and there is clearly a need for improved
Current Population Trend: Decreasing Peru. Its range is incompletely known, and it presumably occurs habitat protection of sites at which this species is known to occur. Further research is needed to help better under-
much more widely, but nevertheless probably has a somewhat stand the threats to this species, and close monitoring of the population is recommended given the potential risk
restricted distribution. of chytridiomycosis.
Population It is believed to be a common species; the type series Bibliography: Cadle, J.E. and McDiarmid, R.W. (1990), Rodríguez, L.O., Cordova, J.H. and Icochea, J. (1993)
is represented by 67 specimens. Data Providers: Lily Rodríguez, Jorge Luis Martinez, César Aguilar Puntriano, Daniel Neira, Edgar Lehr
Habitat and Ecology Associated with streams in montane forest.
At night, individuals were perched on the upper surfaces of leaves
over streams; others were observed during the day resting vertically
on upper leaf surfaces over streams where breeding occurred. Eggs
are deposited on leaves above permanent and temporary streams,
and larvae develop in the streams. It is not known if the species can
adapt to modified habitats.
Major Threats The threats to the species are not well known. It is
210 Threatened Amphibians of the World

EN Centrolene lynchi (Duellman, 1980)

Endangered B2ab(iii,iv,v) range is 1,100-1,600m asl.
Order, Family: Anura, Centrolenidae Population It is known to have undergone declines in Ecuador, but has been seen as recently as August 2001. It
Country Distribution: Colombia, Ecuador appears to have disappeared entirely from some regions, such as the Río Faisanes area.
Current Population Trend: Decreasing Habitat and Ecology It lives in cloud forest, including secondary forest, along streams, where it needs forest
vegetation overhanging water, but it is not found in degraded habitats. It breeds in streams, with the eggs laid on
leaves overhanging the water.
Major Threats The most likely cause of the severe decline of this species is the movement of the cloud layer up

© Mario Humberto Yánez-Muñoz

Geographic Range This species is known only from two general areas: on the western versant of the Cordillera
Occidental in Risaralda Department, Colombia; and seven localities on the western versant of the Andes in Ecuador in
Pichincha Province. It probably occurs between these areas, but it is a rare species and hard to detect. Its altitudinal
the mountain sides as a result of climate change, resulting in reduced humidity within the altitudinal range of the
species (probably exacerbated by habitat fragmentation). Additional likely threats include: deforestation due to
agricultural development, the planting of illegal crops, fire, logging, and human settlement; introduction of alien
predatory fish species in streams; and pollution resulting from the spraying of illegal crops. Chytridiomycosis also
cannot be ruled out.
Conservation Measures Its distribution overlaps with the Reserva Ecológica Los Illinizas in Ecuador, but it is not
known from any protected areas in Colombia, and further protection of the cloud forest habitat of this species is neces-
sary. Further research is needed to ascertain whether or not chytrid is a threat to this species; given the multitiude of
current threats, including the impact of predatory fish, a captive-breeding programme may need to be established.
Notes on taxonomy: This species and Centrolene scirtetes may be conspecific (D. Cisneros-Heredia pers. comm.).
Bibliography: Duellman, W.E. (1980), Ruiz-Carranza, P.M. and Lynch, J.D. (1991a), Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and Lynch,
J.D. (1996)
Data Providers: Luis A. Coloma, Santiago Ron, Erik Wild, Diego Cisneros-Heredia

EN Centrolene mariae (Duellman and Toft, 1979)

Endangered B1ab(iii) Geographic Range This species is known only from the Serrania and human settlement. In addition, the lower parts of the Serrania de Sira are being selectively logged.
Order, Family: Anura, Centrolenidae de Sira at 2,000m asl, extending north-north-west to south-east Conservation Measures It may occur in the Reserva Comunal El Sira. Further research is needed to determine the
Country Distribution: Peru between two large northward flowing rivers, the Río Pachitea on current population status of this little-known species.
Current Population Trend: Decreasing the west and the Río Ucayali on the east, in Huanuco Department, Bibliography: Cannatella, D.C. and Duellman, W.E. (1982), Duellman, W.E. and Schülte, R. (1993), Duellman, W.E. and Toft, C.A. (1979),
Peru, at 1,550-2,000m asl. It presumably occurs more widely, but it Rodríguez, L.O., Cordova, J.H. and Icochea, J. (1993)
nevertheless probably has a restricted distribution. Data Providers: Lily Rodríguez, Jorge Luis Martinez, Erik Wild, Wilfredo Arizabal, Edgar Lehr, César Aguilar Puntriano
Population The population status of this species is not known (its
description was based on a single specimen).
Habitat and Ecology The fauna of the Serrania de Sira area
apparently represent isolated relicts from Andean relatives that
spread across the lowlands during periods of climatic depression
in the Pleistocene. The area is cloud forest, and characterized by
the presence of many bromeliads and moss. Eggs are deposited on
leaves, and larvae develop in streams.
Major Threats Although the area that the species inhabits is rela-
tively inaccessible, there is some habitat disturbance due to farming

VU Centrolene peristictum (Lynch and Duellman, 1973)

Vulnerable B1ab(iii,iv,v)+2ab(iii,iv,v) Geographic Range This species occurs on the Pacific slopes of the Andes in Ecuador (in Pichincha Province, where
Order, Family: Anura, Centrolenidae it is known from three localities), north to the northern extent of the western slope of the Cordillera Occidental in
Country Distribution: Colombia, Ecuador Colombia. Its recorded altitudinal range is 1,780-1,820m in Colombia, but has been found down to 1,350m in Ecuador.
Current Population Trend: Decreasing The Ecuadorian population appears to be isolated from that in Colombia.
Population In Ecuador, the species seems to have undergone a severe decline, and it has not been recorded in recent
years. It remains uncommon in Colombia.
Habitat and Ecology It is a species of undisturbed cloud forest, and has not been found in secondary habitats. It
breeds in streams, with the eggs laid on leaves overhanging the water.
Major Threats The most likely cause of the severe decline of this species is the movement of the cloud layer up the
mountain sides as a result of climate change, resulting in reduced humidity within the altitudinal range of the species
(probably exacerbated by habitat fragmentation). Additional likely threats include deforestation, due to agricultural
development (including the planting of illegal crops), fire, logging, and human settlement; the introduction of alien
© John D. Lynch

predatory fish species in streams; and pollution resulting from the spraying of illegal crops. Chytridiomycosis also
cannot be ruled out.
Conservation Measures It occurs in three Natural National Parks in Colombia: Farallones de Cali, Munchique, and
Tatama. It is not known from any protected areas in Ecuador. There is a need for further survey work in Ecuador to
establish the current population status of the species in the wild.
Bibliography: Lynch, J.D. and Duellman, W.E. (1973), Ruiz-Carranza, P.M. and Lynch, J.D. (1991a), Ruiz-Carranza, P.M., Ardila-Robayo,
M.C. and Lynch, J.D. (1996)
Data Providers: Luis A. Coloma, Santiago Ron, John Lynch, Diego Cisneros-Heredia, Erik Wild

EN Centrolene petrophilum Ruíz-Carranza and Lynch, 1991

Endangered B1ab(iii) Geographic Range This species occurs on the eastern flank of
Order, Family: Anura, Centrolenidae the northern portion of the Cordillera Oriental, Boyacá Department,
Country Distribution: Colombia Colombia, between 1,600 and 2,200m asl.
Current Population Trend: Decreasing Population It is a common species.
Habitat and Ecology It occurs on vegetation next to or above
streams in cloud forest, and can also occur in secondary forest. It lays
its eggs on rocks in streams, and the larvae develop in the water.
Major Threats Deforestation due to agricultural activities (live-
stock ranching, and the cultivation of crops) is the main threat to
this species.
Conservation Measures It does not occur in any protected areas,
and there is a need for the protection of existing cloud forest frag-
ments, such as the site known as “Pajarito”.
Bibliography: Acosta-Galvis, A.R. (2000), Ruiz-Carranza, P.M. and Lynch,
J.D. (1991b)
Data Providers: María Cristina Ardila-Robayo, Jose Vicente Rueda
 Globally Threatened Amphibian Species 211

EN Centrolene pipilatum (Lynch and Duellman, 1973)

Endangered B1ab(iii,iv,v)+2ab(iii,iv,v) Geographic Range This species occurs on the Amazonian slopes of the Andes in Río Salado, Río Azuelo and 16km
Order, Family: Anura, Centrolenidae from Santa Rosa in Napo and Sucumbíos Provinces, Ecuador, at 1,300-1,740m asl.
Country Distribution: Ecuador Population First collected in 1971, this species was last recorded in 1977. Several trips to known localities where

© Natural History Museum and Biodiversity Research

Current Population Trend: Decreasing the species occurs have not turned up any individuals (e.g. Bustamante 2002); however, the area of available habitat
in the vicinity of the type locality is large, and the sampling effort has probably been insufficient.
Habitat and Ecology It is found near streams in cloud forest, and has been recorded over 3m above the ground.
Eggs are presumably laid on vegetation, with the larvae developing in water.
Major Threats The main threat to the species is habitat loss and degradation mainly due to smallholder farming
and livestock ranching. It is possible that it may also have been affected by chytridiomycosis.
Center, University of Kansas Conservation Measures It probably occurs in the Reserva Ecológica Cayambe-Coca. Further survey work is needed
to determine the current population status of this species.
Bibliography: Bustamante, M. R. (2002), Lynch, J.D. and Duellman, W.E. (1973), Ruiz-Carranza, P.M. and Lynch, J.D. (1991a)
Data Providers: Luis A. Coloma, Santiago Ron, Diego Cisneros-Heredia

CR Centrolene puyoense (Flores and McDiarmid, 1989)

Critically Endangered B1ab(iii) Geographic Range This species is known only from the type local- Conservation Measures The species is not recorded from any protected area. Survey work is urgently required to
Order, Family: Anura, Centrolenidae ity: Puyo, Pastaza Province, Ecuador, at 1,000-1,050m asl. It is likely determine the current population status of this species and the precise limits of its range.
Country Distribution: Ecuador to occur in other nearby localities, but its distribution is probably Notes on taxonomy: Duellman and Schulte (1993) transferred this species to the genusCentrolene from Cochranella.
Current Population Trend: Decreasing very limited in extent. Bibliography: Duellman, W.E. and Schülte, R. (1993), Flores, G. and McDiarmid, R.W. (1989), Ruiz-Carranza, P.M. and Lynch, J.D.
Population It has not been seen since the early 1980s, but there (1991a)
has not been any survey work for the species. However, there is no Data Providers: Luis A. Coloma, Santiago Ron
suitable habitat left at the type locality, so its continued survival
depends upon undiscovered populations occurring nearby in suit-
able habitat.
Habitat and Ecology It lives in lower montane forest. The holotype
was collected in the evening, after it had rained the previous day and
night (Flores and McDiarmid 1989). Reproduction is presumed to be
via eggs laid on vegetation, with the tadpoles developing in water.
Major Threats Habitat destruction and degradation due to agricul-
ture and logging is the primary threat to this species. By 1996, the
forest had been cleared at the type locality.

VU Centrolene quindianum Ruíz-Carranza and Lynch, 1995

Vulnerable D2 Geographic Range This species is known from Filandia and Salinto in Quindio Department, and from Ucumari and
Order, Family: Anura, Centrolenidae Pereira in Risaralda Department, on the western flank of the Cordillera Central, between 1,900 and 2,050m asl, in
Country Distribution: Colombia Colombia.
Current Population Trend: Stable Population It is a very common species.
Habitat and Ecology It occurs on vegetation next to streams in sub-Andean forests. Eggs are laid on leaves over-
hanging the water and when they hatch the tadpoles drop into the water below where they develop further. They
require gallery forest to lay their eggs, hence they do not tolerate much habitat disturbance.
Major Threats Although there are no major threats to the species overall at present, habitat loss and degradation,due
to the expansion of cattle raising, and water pollution, are potential future threats. The invasive Rana catesbeiana
is also a threat, since it eats the tadpoles of this species.
Conservation Measures Its range includes a few protected areas. There is a need for close monitoring of the
population status of this species.
© John D. Lynch

Bibliography: Acosta-Galvis, A.R. (2000), Ruiz-Carranza, P.M. and Lynch, J.D. (1995b), Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and
Lynch, J.D. (1996)
Data Providers: Erik Wild, John Lynch

VU Centrolene robledoi Ruíz-Carranza and Lynch, 1995

Vulnerable B1ab(iii) Geographic Range This species occurs north of the Cordillera Central in the department of Antioquia, on the eastern
Order, Family: Anura, Centrolenidae flanks of the Cordillera Central in the department of Risaralda, and on the western flank of the Cordillera Central in
Country Distribution: Colombia the department of Caldas, in Colombia. It has been recorded from 800-2,800m asl.
Current Population Trend: Decreasing Population It is a common species.
Habitat and Ecology It occurs on vegetation next to streams in cloud forest. It lays its eggs on leaves, and when
hatched the tadpoles drop in to the water below. It has also been recorded from secondary forests near water.
Major Threats The major threat is habitat loss and degradation due to the expansion of agriculture, coupled with
water source loss, and pollution from the fumigation of crops.
Conservation Measures Its range includes Reserva Regional Bosques de Florencia in the department of Caldas.
Bibliography: Ardila-Robayo, M.C. and Acosta-Galvis, A. (2000b), Ruiz-Carranza, P.M. and Lynch, J.D. (1995b), Ruiz-Carranza, P.M.,
© Andrew Crawford

Ardila-Robayo, M.C. and Lynch, J.D. (1996)

Data Providers: Martha Patricia Ramírez Pinilla, Mariela Osorno-Muñoz, Jose Vicente Rueda, Adolfo Amézquita, María Cristina
Ardila-Robayo
212 Threatened Amphibians of the World

VU Centrolene tayrona Ruíz-Carranza and Lynch, 1991

Vulnerable B1ab(iii) Geographic Range This species is known from the Sierra Nevada de Santa Marta, in Magdalena Department,
Order, Family: Anura, Centrolenidae Colombia. It has been recorded between 980 and 1,790m asl.
Country Distribution: Colombia Population It is a common species.
Current Population Trend: Decreasing Habitat and Ecology It occurs on vegetation next to water sources in sub-Andean forests and cloud forests. It lays
its eggs on leaves, and when hatched the tadpoles fall in to the water below where they develop further.
Major Threats The major threat is habitat loss due to agriculture (both crops and livestock), as well as water pol-
lution due to the fumigation of crops.
Conservation Measures Its range includes Parque Nacional Natural Sierra Nevada de Santa Marta.
Notes on taxonomy: This species was considered to be a synonym of Hylopsis platycephalus Werner, 1894, by Lynch (1981b), but this
was not accepted by McDiarmid and Savage (1984).
Bibliography: Acosta-Galvis, A.R. (2000), Lynch, J.D. (1981b), McDiarmid, R.W. and Savage, J.M. (1984), Ruiz-Carranza, P.M. and Lynch,
J.D. (1991b), Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and Lynch, J.D. (1996)
© Taran Grant

Data Providers: John Lynch, Martha Patricia Ramírez Pinilla, Mariela Osorno-Muñoz, Jose Vicente Rueda, Adolfo Amézquita, María
Cristina Ardila-Robayo

VU Cochranella adiazeta Ruíz-Carranza and Lynch, 1991

Vulnerable B1ab(iii) Geographic Range This species occurs on the eastern slope of the Cordillera Oriental in Cundinamarca, Santander
Order, Family: Anura, Centrolenidae and Tolima Departments, Colombia, between 1,130 and 2,060m asl.
Country Distribution: Colombia Population It is common, with several recent records.
Current Population Trend: Decreasing Habitat and Ecology It occurs on vegetation next to streams in premontane humid forest; it also occurs in secondary
forest. It lays its eggs on vegetation, and the tadpoles develop in water.
© Conservacion Internacional-Colombia

Major Threats The major threat is habitat loss due to agricultural expansion (cattle ranching and crops).
Conservation Measures It occurs in the Santuario de Fauna y Flora Guanentá Alto Río Fonce.
Bibliography: Acosta-Galvis, A.R. (2000), Ruiz-Carranza, P.M. and Lynch, J.D. (1991d), Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and
Lynch, J.D. (1996)
Data Providers: Jose Vicente Rueda, Martha Patricia Ramírez Pinilla

CR Cochranella anomala (Lynch and Duellman, 1973)

Critically Endangered B2ab(iii) Geographic Range This species is known only from the type locality: Río Azuela, Napo Province, in northern
Order, Family: Anura, Centrolenidae Ecuador, at 1,740m asl. It is likely to occur a little more widely, but surveys elsewhere have not recorded it, and so
Country Distribution: Ecuador its distribution is unlikely to be extensive.
© Natural History Museum and Biodiversity Research

Current Population Trend: Decreasing Population It has not been seen since its original discovery (one individual) in 1971. A few return visits to the type
locality have not recorded it.
Habitat and Ecology It lives in cloud forest. The holotype was found on a mossy limb of a bush about 1.5m above a cas-
cading rivulet at night (Lynch and Duellman 1973). Its breeding is unknown, but is likely to take place in streams.
Major Threats At the type locality, patches of forest have been cleared for agriculture and livestock farming.
Conservation Measures The distribution range of this species appears to overlap with Reserva Ecológica Cay-
Center, University of Kansas

ambe-Coca. Survey work is required to determine the population status of this species and to ascertain the limits
of its range.
Bibliography: Lynch, J.D. and Duellman, W.E. (1973), Ruiz-Carranza, P.M. and Lynch, J.D. (1991a)
Data Providers: Luis A. Coloma, Santiago Ron

VU Cochranella armata Lynch and Ruíz-Carranza, 1996

Vulnerable D2 Geographic Range This species is known only from near Boquerón at the base of Cerro Ingles, near Vereda La
Order, Family: Anura, Centrolenidae Amarillas, Município El Cairo, in Valle del Cauca Department, Colombia, at 2,160m asl. Even though every stream
Country Distribution: Colombia within 5km of the type locality has been sampled, the species has not been recorded, suggesting that it has a very
Current Population Trend: Stable restricted range.
Population It was reportedly a common species when collected.
Habitat and Ecology Specimens have been collected on vegetation next to an open stream in primary forest. Its
breeding habits are not known, but it is likely to breed in streams.
Major Threats There are no major threats, but given its small range it is susceptible to stochastic threatening
processes.
Conservation Measures The type locality is within a private reserve, which is part of a larger network of reserves.
Maintenance of the habitat of this restricted-range species is essential to ensure its persistence, and it requires
close population monitoring given its restriction to a single location.
© John D. Lynch

Bibliography: Acosta-Galvis, A.R. (2000), Lynch, J.D. and Ruiz-Carranza, P.M. (1996a)
Data Providers: Erik Wild, John Lynch
 Globally Threatened Amphibian Species 213

VU Cochranella balionota (Duellman, 1981)

Vulnerable B2ab(iii,iv,v) Geographic Range This species occurs between 400 and 800m might also be affected by the movement of the cloud layer up the mountain sides as a result of climate change, resulting
Order, Family: Anura, Centrolenidae along the western slope of the Cordillera Occidental of Colombia, in reduced humidity within the altitudinal range of the species (probably exacerbated by habitat fragmentation).
Country Distribution: Colombia, Ecuador from El Tambito in Cauca Department, south to Ecuador. In Ecuador, Conservation Measures Its presence has not been confirmed from any protected areas, though it might occur in
Current Population Trend: Decreasing it is known only from the type locality: Mindo, in Pichincha Province, Parque Nacional Natural Munchique in Colombia. There is clearly a need for improved habitat protection of sites at
at 1,540m asl. which this species is known to occur.
Population It is reasonably common in Colombia, but there are no Bibliography: Acosta-Galvis, A.R. (2000), Duellman, W.E. (1981), Ruiz-Carranza, P.M. and Lynch, J.D. (1991a), Ruiz-Carranza, P.M.,
recent records from Ecuador. Ardila-Robayo, M.C. and Lynch, J.D. (1996)
Habitat and Ecology It is an arboreal species, living on vegetation Data Providers: Wilmar Bolívar, Luis A. Coloma, Santiago Ron, Diego Cisneros-Heredia, John Lynch, Erik Wild
next to streams in humid lowland tropical forest and cloud forest. It
seems to require closed forest. It breeds in streams, with the eggs
laid on leaves overhanging water.
Major Threats The major threat is habitat loss as a result of defor-
estation for agricultural development (including the planting of illegal
crops), logging, and human settlement. Other threats include the
introduction of alien predatory fish, and pollution resulting from the
spraying of illegal crops. Like some other centrolenids, this species

VU Cochranella cochranae (Goin, 1961)

Vulnerable B1ab(iii) Geographic Range This species is found on the lower Amazonian slopes of the Andes in northern and southern
Order, Family: Anura, Centrolenidae Ecuador (Sucumbios, Pastaza and Zamora Chinchipe Departments), at elevations of 1,100-1,600m asl. It might occur
Country Distribution: Ecuador a little more widely.
© Natural History Museum and Biodiversity Research

Current Population Trend: Decreasing Population This species is noted as being uncommon across most of its range.
Habitat and Ecology It lives in montane rainforest along streams with steep gradients (Lynch and Duellman 1973).
Reproduction is presumed to occur via eggs laid on vegetation, with the tadpoles developing in the water.
Major Threats Habitat destruction and degradation is a threat to this species, mainly as a result of smallholder
farming activities and clear-cutting of forests.
Conservation Measures Its distribution range overlaps with Parque Nacional Sumaco Napo-Galeras, Reserva
Center, University of Kansas

Ecológica Antisana, Parque Nacional Podocarpus and Parque Nacional Llanganates.

Bibliography: Goin, C.J. (1961), Lynch, J.D. and Duellman, W.E. (1973), Ruiz-Carranza, P.M. and Lynch, J.D. (1991a)
Data Providers: Luis A. Coloma, Santiago Ron, Diego Almeida

VU Cochranella daidalea Ruíz-Carranza and Lynch, 1991

Vulnerable B1ab(iii) Geographic Range This species occurs on the western flank of the Cordillera Oriental in Cundinamarca and Santander
Order, Family: Anura, Centrolenidae Departments, Colombia, between 1,630 and 2,060m asl.
Country Distribution: Colombia Population It is common, and has recently been collected.
Current Population Trend: Decreasing Habitat and Ecology It occurs on vegetation near streams in premontane and cloud forest, and has been recorded
from secondary forest. It lays its eggs on vegetation, and the tadpoles develop in the water.
© Conservacion Internacional-Colombia

Major Threats The main threat to this species is habitat fragmentation and loss due to agriculture (cattle raising
and crops).
Conservation Measures It occurs in the Santuario de Fauna y Flora Guanentá Alto Río Fonce.
Bibliography: Acosta-Galvis, A.R. (2000), Ruiz-Carranza, P.M. and Lynch, J.D. (1991c), Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and
Lynch, J.D. (1996)
Data Providers: María Cristina Ardila-Robayo, Jose Vicente Rueda

VU Cochranella garciae Ruíz-Carranza and Lynch, 1995

Vulnerable B1ab(iii) Geographic Range This species is known from Cauca, Huila and Bibliography: Ruiz-Carranza, P.M. and Lynch, J.D. (1995b), Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and Lynch, J.D. (1996)
Order, Family: Anura, Centrolenidae Tolima (Combeima) Departments, on the eastern slope of the Cordil- Data Providers: Alonso Quevedo Gil, John Lynch
Country Distribution: Colombia lera Central, in Colombia, between 1,900 and 3,030m asl.
Current Population Trend: Decreasing Population It is a relatively common species.
Habitat and Ecology It occurs on vegetation alongside streams
in sub-Andean and Andean forests. The eggs are laid on leaves
overhanging water and when they hatch the tadpoles drop in to
the water below where they develop further. They require gallery
forest to lay their eggs and hence are very sensitive to disturbance
of their forest habitat.
Major Threats The main threat to this species is habitat loss as a
result of clear-cutting of forests and agricultural encroachment, due
to the planting of illegal crops and cattle raising.
Conservation Measures The range of the species includes a
few protected areas, such as Parque Nacional Natural Nevado
del Huila.
214 Threatened Amphibians of the World

VU Cochranella griffithsi (Goin, 1961)

Vulnerable B1ab(iii) Geographic Range This species occurs on the Pacific slopes of the Cordillera Occidental in Colombia, in Antioquia,
Order, Family: Anura, Centrolenidae Choco, Cauca, Valle del Cauca, Nariño and Risaralda Departments, south to Cotopaxi Province in adjacent Ecuador. It
Country Distribution: Colombia, Ecuador is also known from the eastern slopes of the Cordlillera Central in the Department of Caldas, Colombia. In Ecuador it

© Natural History Museum and Biodiversity Research

Current Population Trend: Decreasing has been recorded from only five sites covering a relatively small area. It has been recorded at 1,780-2,650m asl.
Population It is very common in Colombia, but uncommon in Ecuador where it appears to have disappeared from
Quebrada Zapadores in Pichincha Province, where it once occurred.
Habitat and Ecology It occurs on vegetation next to streams in montane forests, and is perhaps a little more arboreal
than some other related species. It is not found in degraded habitats, but does occur in selectively logged forest. It
breeds in streams, with the eggs laid on leaves overhanging the water.
Center, University of Kansas Major Threats The major threat is habitat loss as a result of deforestation for agricultural development (including
the planting of illegal crops), logging, and human settlement. Other threats include the introduction of alien preda-
tory fish, and pollution resulting from the spraying of illegal crops. Like some other centrolenids, this species might
also be affected by the movement of the cloud layer up the mountain sides as a result of climate change, resulting
in reduced humidity within the altitudinal range of the species (probably exacerbated by habitat fragmentation).
Chytridiomycosis is a possible future threat.
Conservation Measures It occurs in several protected areas in Colombia, while in Ecuador its range overlaps with
Reserva Ecológica Los Illinizas.
Bibliography: Acosta-Galvis, A.R. (2000), Goin, C.J. (1961), Morales, M. et al. (2002), Ruiz-Carranza, P.M. and Lynch, J.D. (1991a),
Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and Lynch, J.D. (1996)
Data Providers: Wilmar Bolívar, Luis A. Coloma, Santiago Ron, Mario Yánez-Muñoz, Taran Grant

EN Cochranella luminosa Ruíz-Carranza and Lynch, 1995

Endangered B1ab(iii)+2ab(iii) Geographic Range This species is known from three localities in Bibliography: Ruiz-Carranza, P.M. and Lynch, J.D. (1995a)
Order, Family: Anura, Centrolenidae Antioquia Department, on the western flank of the western Andes Data Providers: Erik Wild, John Lynch
Country Distribution: Colombia in Colombia, between 1,140 and 1,430m asl.
Current Population Trend: Decreasing Population It is not a rare species but it is difficult to collect as it
occurs high in the canopy.
Habitat and Ecology It is restricted to primary forest, occurring
next to streams. It lays its eggs on the top of leaves in the canopy
and when hatched the larvae drop into the stream below where
they develop further.
Major Threats The main threats are habitat loss and fragmenta-
tion caused by the expansion of livestock ranching activities, timber
extraction for subsistence use, and the planting of illegal crops.
Conservation Measures The range of the species includes
Parque Nacional Natural Orquideas. However, additional habitat
protection is required for this species, given its reliance on intact
primary forest.

EN Cochranella mache Guayasamin and Bonaccorso, 2004

Endangered B1ab(iii) Geographic Range This species is known only from around 510m asl near Río Aguacatal in the Reserva Biológica
Order, Family: Anura, Centrolenidae Bilsa in the Montañas de Mache of Esmeraldas Province in the Chocó region of the north-western Ecuadorian lowlands.
Country Distribution: Ecuador The species is likely to occur in other localities within the surrounding Mache Chindul Ecological Reserve, but may
Current Population Trend: Decreasing be restricted to the Mache-Chindul Mountains.
Population It appears to be rare, although this may be due to the fact that the species generally inhabits the forest
canopy and may be hard to locate.
Habitat and Ecology It is known from evergreen foothill forest, and individuals have been found at night on vegeta-
tion 1-1.5m above the ground near streams. The species is presumed to be a larval developer.
© Juan Manuel Guayasamin

Major Threats The Mache Chindul Ecological Reserve represents the last sizeable block of forest in the northern
coastal hills of north-western Ecuador. Unfortunately, the area is under significant pressure from logging.
Conservation Measures The known range of this species is encompassed by the 2,500-ha Reserva Biológica Bilsa,
managed by Fundación Jatun Sacha, which, in turn, is located within the Mache Chindul Ecological Reserve.
Bibliography: Guayasamin, J.M. and Bonaccorso, E. (2004)
Data Providers: Juan Guayasamin

EN Cochranella megacheira (Lynch and Duellman, 1973)

Endangered B1ab(iii,iv,v)+2ab(iii,iv,v) Geographic Range This species occurs on the Amazonian slopes of the Andes in Ecuador (at four locations in Napo
Order, Family: Anura, Centrolenidae Province: near Santa Rosa, Río Azuela, Río Salado, and Guacamayos), and also from the Amazonian slopes of the
Country Distribution: Colombia, Ecuador Cordillera Oriental, in Putumayo Department, Colombia. It has been recorded at 1,300-1,750m asl.
© Natural History Museum and Biodiversity Research

Current Population Trend: Decreasing Population It was most recently seen in Ecuador in 2000, and it appears to be very rare, having apparently disap-
peared from Río Azuela. There is no recent information from Colombia.
Habitat and Ecology It inhabits mature, closed cloud forest, where it has been found on the leaves and stems of
bushes and trees overhanging streams. It does not occur in degraded areas. It breeds in streams, with the eggs laid
on leaves overhanging the water.
Major Threats The major threats include: deforestation for agricultural development, the planting of illegal crops,
Center, University of Kansas

logging, and human settlement; introduction of alien predatory fish in streams; and pollution resulting from the spraying
of illegal crops. Like some other centrolenids, this species might also be affected by the movement of the cloud layer
up the mountain slopes as a result of climate change, resulting in reduced humidity within the altitudinal range of the
species (probably exacerbated by habitat fragmentation). Chytridiomycosis may present a possible future threat.
Conservation Measures It has not been found in any protected areas, and protection of the cloud forest habitat of
this species is urgently needed. Survey work is necessary to ascertain the current population status of this species;
given the nature of some of the threats, an ex situ population may need to be established.
Bibliography: Lynch, J.D. and Duellman, W.E. (1973), Ruiz-Carranza, P.M. and Lynch, J.D. (1991a), Ruiz-Carranza, P.M., Ardila-Robayo,
M.C. and Lynch, J.D. (1996)
Data Providers: Wilmar Bolívar, Luis A. Coloma, Santiago Ron, Ana Almandáriz, Taran Grant
 Globally Threatened Amphibian Species 215

VU Cochranella posadae Ruíz-Carranza and Lynch, 1995

Vulnerable B1ab(iii) Geographic Range This species is known from Caldas (Samaná), Cauca (Inza) and Huila (San Jose Isnos) Departments
Order, Family: Anura, Centrolenidae on the eastern flank of the Cordillera Central in Colombia, between 1,100 and 2,800m asl. It presumably occurs more
Country Distribution: Colombia widely, particularly at localities between the two currently known areas.
Current Population Trend: Decreasing Population It is a common species.
Habitat and Ecology It occurs in sub-Andean and Andean forests, on vegetation alongside streams. Eggs are laid on

© Fundación ProAves / www.proaves.org

leaves over-hanging water and when they hatch the tadpoles drop into the water below where they develop further.
They require gallery forest to lay their eggs, and hence are sensitive to any disturbance of their forest habitat.
Major Threats Threats to this species include habitat fragmentation and loss, due to the expansion of agriculture
(including the planting of illegal crops) and timber extraction, and water pollution.
Conservation Measures The range of the species does not include any protected areas, and there is a need for
improved habitat protection of sites where this species is known to occur.
Bibliography: Acosta-Galvis, A.R. (2000), Ruiz-Carranza, P.M. and Lynch, J.D. (1995b), Ruiz-Carranza, P.M. and Lynch, J.D. (1997), Ruiz-
Carranza, P.M., Ardila-Robayo, M.C. and Lynch, J.D. (1996)
Data Providers: Erik Wild, John Lynch

VU Cochranella prasina (Duellman, 1981)

Vulnerable B1ab(iii) Geographic Range This species is known from Valle del Cauca Department, northwards through Choco and Risaralda
Order, Family: Anura, Centrolenidae departments to Antioquia Department, on the western flank of the western Andes, in Colombia, between 900 and
Country Distribution: Colombia 1,450m asl. It probably occurs more widely.
© Natural History Museum and Biodiversity Research

Current Population Trend: Decreasing Population It can be locally common in suitable habitat.
Habitat and Ecology It occurs in rainforests and sub-Andean forest, and is often found on vegetation along streams
(being restricted to streams with canopy cover). It lays its eggs on the upper surface of leaves and when hatched the
tadpoles drop into the stream below.
Major Threats The major threats are habitat fragmentation and loss, due to the expansion of agriculture (cattle
raising and planting of illegal crops), and water pollution.
Center, University of Kansas

Conservation Measures The range of the species includes Parque Nacional Natural Orquideas.
Bibliography: Acosta-Galvis, A.R. (2000), Ruiz-Carranza, P.M. and Lynch, J.D. (1991a), Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and
Lynch, J.D. (1996)
Data Providers: Fernando Castro, Taran Grant, Erik Wild

VU Cochranella punctulata Ruíz-Carranza and Lynch, 1995

Vulnerable B1ab(iii) Geographic Range This species is known from Tolima, Antioquia and Caldas Departments on the eastern flank of
Order, Family: Anura, Centrolenidae the Central Cordillera, between 500 and 930m asl, in Colombia. It probably occurs more widely.
Country Distribution: Colombia Population It is a very common species.
Current Population Trend: Decreasing Habitat and Ecology It occurs in sub-Andean forests, on vegetation alongside streams. Eggs are laid on leaves
© Mauricio Rivera-Correa, courtesy of Museo de

over-hanging water and when they hatch the tadpoles drop in to the water below where they develop further. They
require gallery forest to lay their eggs, and hence are very sensitive to habitat disturbance.
Major Threats The major threats are habitat fragmentation and loss, due to the expansion of agriculture (cattle
Herpetología Universidad de Antioquia

raising and planting of illegal crops), and water pollution. A small white fly lays its eggs within the eggs of this
species, resulting in egg mortality.
Conservation Measures The range of this species does not include any protected areas, hence there is a need for
improved habitat protection at sites where it is known to occur.
Bibliography: Acosta-Galvis, A.R. (2000), Ruiz-Carranza, P.M. and Lynch, J.D. (1995b), Ruiz-Carranza, P.M. and Lynch, J.D. (1997), Ruiz-
Carranza, P.M., Ardila-Robayo, M.C. and Lynch, J.D. (1996)
Data Providers: Alonso Quevedo Gil, John Lynch

VU Cochranella resplendens (Lynch and Duellman, 1973)

Vulnerable B1ab(iii) Geographic Range This species is only known from the type locality in Ecuador (Santa Cecilia) and from one
Order, Family: Anura, Centrolenidae location in Colombia (south-western Putumayo), but might occur more widely than these records suggest. It occurs
Country Distribution: Colombia, Ecuador between 300 and 400m asl.
Current Population Trend: Decreasing Population This is thought to be a rare species in Colombia, but there is no information on population status in Ecuador.
Habitat and Ecology It occurs in lowland tropical primary and secondary rainforests, and sub-Andean forests, on
vegetation next to running water sources like streams. The holotype was found in primary rainforest (Lynch and Du-
ellman 1973). It has not been recorded from anthropogenic habitats. The eggs are laid on leaves above the streams,
and when hatched the tadpoles drop into the stream below.
Major Threats The species is threatened by habitat fragmentation and habitat loss, as a result of agricultural
expansion (including the planting of illegal crops) and timber extraction, and water pollution.
Conservation Measures It is not known from any protected areas in Colombia or Ecuador, and there is a need for
improved habitat protection at sites where this species is known to occur.
© John D. Lynch

Bibliography: Acosta-Galvis, A.R. (2000), Guayasamin, J.M. and Bonaccorso, E. (2004), Lynch, J.D. and Duellman, W.E. (1973), Ruiz-
Carranza, P.M. and Lynch, J.D. (1991a), Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and Lynch, J.D. (1996)
Data Providers: Fernando Castro, Jose Vicente Rueda, Wilmar Bolívar, Ruth Amanda Estupinan, Luis A. Coloma, Santiago Ron, Ana
Almandáriz
216 Threatened Amphibians of the World

VU Cochranella riveroi (Ayarzaguena, 1992)

Vulnerable D2 Geographic Range This species is endemic to Cerro Aracamuni, in
Order, Family: Anura, Centrolenidae Amazonas State, Venezuela, at an elevation of 1,600m asl.
Country Distribution: Venezuela Population It is a rare species.
Current Population Trend: Stable Habitat and Ecology It lives and reproduces (by larval development)
within terrestrial bromeliads in montane forest.
Major Threats There are no current major threats, but the re-
stricted range of this species makes it vulnerable to threatening
processes.
Conservation Measures The population occurs in a Natural Monu-
ment within the Parque Nacional Serrania de la Neblina. There is a
need for close population monitoring of this species given that it is
known only from a single location.
Bibliography: Ayarzaguena, J. (1992), Barrio Amorós, C.L. (2004), Barrio, C.
(1998), Duellman, W.E. (1993), Duellman, W.E. (1999), Gorzula, S. and Señaris,
J.C. (1998), La Marca, E. (1997), Lynch, J.D. and Ruiz-Carranza, P.M. (1996a),
Myers, C.W. and Donnelly, M.A. (1997)
Data Providers: Enrique La Marca, Celsa Señaris

VU Cochranella rosada Ruíz-Carranza and Lynch, 1997

Vulnerable B1ab(iii) Geographic Range This species is known from Tolima (Falan near Mariqutia), Antioquia (Anori), and Caldas Depart-
Order, Family: Anura, Centrolenidae ments, between 1,100 and 2,000m asl, in Colombia. It is believed to occur more widely.
Country Distribution: Colombia Population It is a common species in Tolima, but is thought to be uncommon in Antioquia.
Current Population Trend: Decreasing Habitat and Ecology It occurs in sub-Andean forests on vegetation alongside streams. Eggs are laid on leaves
over-hanging water and when they hatch the tadpoles drop into the water below where they develop further. They
© Fundación ProAves / www.proaves.org

require gallery forest to lay their eggs and hence are sensitive to any habitat disturbance.
Major Threats The major threats are habitat fragmentation and loss, due to agricultural expansion (including planting
of illegal crops) and timber extraction, and water pollution.
Conservation Measures The range of the species does not include any protected areas, and there is a need for
improved habitat protection at the sites where it is known to occur.
Bibliography: Acosta-Galvis, A.R. (2000), Ruiz-Carranza, P.M. and Lynch, J.D. (1997)
Data Providers: Alonso Quevedo Gil, John Lynch

VU Cochranella ruizi Lynch, 1993

Vulnerable B1ab(iii) Geographic Range This species is known from both slopes of the western Andes, between Cauca and Risaralda
Order, Family: Anura, Centrolenidae Departments, and on the western slope of the central Andes, in Colombia, between 2,100 and 2,470m asl.
Country Distribution: Colombia Population It is not a particularly common species.
Current Population Trend: Decreasing Habitat and Ecology It occurs on vegetation next to streams in sub-Andean forests. It lays its eggs on leaves above
the water and when hatched the tadpoles drop into the stream below where they develop further. It is restricted to
gallery forest due to its requirement for overhanging branches for breeding habitat.
Major Threats The major threats are habitat loss, due to agricultural development (including the planting of illegal
crops), logging, and human settlement, and pollution resulting from the spraying of illegal crops.
Conservation Measures The range of the species includes a few protected areas, such as the Parque Nacional
Natural Tatamá.
© Martha Silva-Velasco

Bibliography: Lynch, J.D. (1993a), Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and Lynch, J.D. (1996)
Data Providers: Fernando Castro, John Lynch, Erik Wild

VU Cochranella savagei Ruíz-Carranza and Lynch, 1991

Vulnerable B1ab(iii) Geographic Range This species occurs on the western slope of the Cordillera Central in the departments of Quin-
Order, Family: Anura, Centrolenidae dio and Risaralda, on the eastern slope of the Cordillera Central in the department of Caldas, and on the western
Country Distribution: Colombia slope of the Cordillera Occidental, in the department of Valle del Cauca, in Colombia. It has been recorded from
Current Population Trend: Decreasing 1,400-2,410m asl.
© Paul D. A. Gutiérrez-C, courtesy of Museo de

Population It is a common species.

Habitat and Ecology It occurs in sub-Andean forests on vegetation next to running water such as streams. The
eggs are laid on leaves, and when hatched the tadpoles fall into the water below where they develop further. It is
Herpetología Universidad de Antioquia

also recorded from secondary forests.

Major Threats The major threat is forest loss due to agriculture, both crops and livestock, and pollution (as a result
of the fumigation of crops).
Conservation Measures Its range includes the Reserva Regional Bosques de Florencia and the Parque Nacional
Natural Farallones de Cali.
Bibliography: Acosta-Galvis, A.R. (2000), Ruiz-Carranza, P.M. and Lynch, J.D. (1991b), Ruiz-Carranza, P.M. and Lynch, J.D. (1991c),
Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and Lynch, J.D. (1996)
Data Providers: Wilmar Bolívar, Martha Patricia Ramírez Pinilla, Mariela Osorno-Muñoz, Jose Vicente Rueda, Adolfo Amézquita,
María Cristina Ardila-Robayo
 Globally Threatened Amphibian Species 217

EN Cochranella saxiscandens Duellman and Schulte, 1993

Endangered B1ab(iii) Geographic Range This species is known only from one locality (two sites in close proximity to each other) at 800m
Order, Family: Anura, Centrolenidae asl on the south slope of the north-west-south-east range of the mountains immediately to the north of Tarapoto
Country Distribution: Peru Province, San Martin Department, in northern Peru. It probably occurs more widely, but it is still likely to have a

© Natural History Museum and Biodiversity Research

Current Population Trend: Decreasing restricted distribution.
Population It is a common species at the type locality, and was described from a series of 31 adult males.
Habitat and Ecology The type locality is a rocky gorge into which two streams plunge. The sides of the gorge are
nearly vertical rocky walls, and the floor of the gorge is littered with huge boulders. The species was found perched
on ferns on boulders, but most were on the boulders within 10cm of the waterline at night. They breed in streams,
with eggs laid on leaves above water.
Center, University of Kansas Major Threats The main threat to this species is habitat loss due to agriculture, wood extraction, and human settle-
ment, as well as the impacts of tourism. The potential impacts of localized climate change and possible infection
with the chytrid fungus require further investigation.
Conservation Measures It is not known to occur in any protected areas, but the type locality at least is in need of
some formal protection or management.
Bibliography: Duellman, W.E. and Schülte, R. (1993), Rodríguez, L.O., Cordova, J.H. and Icochea, J. (1993)
Data Providers: Lily Rodríguez, Jorge Luis Martinez, Ariadne Angulo, Karl-Heinz Jungfer, Wilfredo Arizabal

VU Cochranella siren (Lynch and Duellman, 1973)

Vulnerable B1ab(ii,iii,iv,v) Geographic Range This species occurs on the Amazonian slopes of the eastern Andes in Napo Province, Ecuador
Order, Family: Anura, Centrolenidae (at elevations of 1,310-1,700m asl); in Putumayo Department, Colombia; and in Ayacucho Province, southern Peru.
Country Distribution: Colombia, Ecuador, Peru The Peruvian specimens might refer to a distinct species (Duellman and Schülte 1993).
© Natural History Museum and Biodiversity Research

Current Population Trend: Decreasing Population In Ecuador, it has disappeared from the type locality (Río Salado) and several other localities, but one
specimen was seen in Azuela in 1998.
Habitat and Ecology It lives near streams in pre-montane forest (Lynch and Duellman 1973; QCAZ database). It lays
its eggs on vegetation, and the tadpoles develop in the water.
Major Threats Habitat destruction and degradation is a threat to this species, due mainly to the activities of
smallholder farmers and as a result of logging. Given its narrow altitudinal range, its decline in Ecuador might be
Center, University of Kansas

related to the impacts of climate change.

Conservation Measures It occurs in the Reserva Ecológica Cayambe-Coca in Ecuador.
Bibliography: Duellman, W.E. and Schülte, R. (1993), Lynch, J.D. and Duellman, W.E. (1973), Rodríguez, L.O., Cordova, J.H. and Icochea,
J. (1993), Ruiz-Carranza, P.M. and Lynch, J.D. (1991a)
Data Providers: Luis A. Coloma, Santiago Ron, Diego Cisneros-Heredia

VU Cochranella susatamai Ruíz-Carranza and Lynch, 1995

Vulnerable B1ab(iii) Geographic Range This species is known from the departments of Tolima, Caldas and Antioquia, on the eastern
Order, Family: Anura, Centrolenidae flank of the Cordillera Central, in Colombia, between 400 and 1,650m asl. It is likely to occur a little more widely
Country Distribution: Colombia than these records suggest.
Current Population Trend: Decreasing Population It is a common species.
Habitat and Ecology It occurs in sub-Andean forests on vegetation along streams. Eggs are laid on leaves overhanging
© Fundación ProAves / www.proaves.org

water and when they hatch the tadpoles drop into the water below where they develop further. They require gallery
forest to lay their eggs, and hence are sensitive to any habitat disturbance.
Major Threats The major threats are habitat fragmentation and loss, due to agricultural expansion (including planting
of illegal crops) and timber extraction, and water pollution.
Conservation Measures The range of the species does not include any protected areas, and there is a need for
improved habitat protection at sites where it is known to occur.
Bibliography: Acosta-Galvis, A.R. (2000), Ruiz-Carranza, P.M. and Lynch, J.D. (1995b), Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and
Lynch, J.D. (1996)
Data Providers: Erik Wild, John Lynch

VU Cochranella xanthocheridia Ruíz-Carranza and Lynch, 1995

Vulnerable B1ab(iii) Geographic Range This species is known from the departments
Order, Family: Anura, Centrolenidae of Risaralda and Antioquia on the western flank of the Cordillera
Country Distribution: Colombia Occidental, in Colombia, between 800 and 2,060m asl.
Current Population Trend: Decreasing Population It is a common species.
Habitat and Ecology It occurs on vegetation next to streams in
sub-Andean forests. It lays its eggs on the top of leaves and when
hatched the tadpoles drop into the water below where they develop
further. It has not been recorded outside forest habitat.
Major Threats The major threats are habitat fragmentation and loss,
due to agricultural expansion (including planting of illegal crops) and
timber extraction, and water pollution.
Conservation Measures The range of the species includes Parque
Nacional Natural Orquideas.
Bibliography: Acosta-Galvis, A.R. (2000), Ruiz-Carranza, P.M. and Lynch,
J.D. (1995a)
Data Providers: Fernando Castro, John Lynch
218 Threatened Amphibians of the World

VU Hyalinobatrachium antisthenesi (Goin, 1963)

Vulnerable B1ab(iii) Geographic Range This species is known from the northern versants of the Venezuelan Coastal Range, in the
Order, Family: Anura, Centrolenidae central region, and from the Cerro Azul, in Cojedes State, in the Serranía del interior. It has an altitudinal range

© César Luis Barrio-Amorós / Fundación AndígenA

Country Distribution: Venezuela between 220 and 1,200m asl.
Current Population Trend: Decreasing Population The current population status of this species is not known.
Habitat and Ecology It occurs in cloud forests and gallery forests. It lays its eggs on either side of leaves overhanging
streams, and when hatched the larvae fall into the stream below where they develop further.
Major Threats Threats to this species are unknown, but the main threat is likely to be habitat loss due to smallholder
farming activities, logging, burning, and human settlements. The Venezuelan Coastal Range is a highly modified
habitat.
Conservation Measures Some populations are within the Parque Nacional Henri Pittier and Rancho Grande in
Aragua State. Further survey work is needed to determine the population status of this species.
Bibliography: Barrio Amorós, C.L. (2004), Goin, C.J. (1963), La Marca, E. (1992), La Marca, E. (1997), Rivero, J.A. (1968a), Ruiz-Carranza,
P.M. and Lynch, J.D. (1991a)
Data Providers: Enrique La Marca, Jesús Manzanilla, Celsa Señaris

EN Hyalinobatrachium cardiacalyptum McCranie and Wilson, 1997

Endangered B1ab(iii,v)+2ab(iii,v) Geographic Range This species is found in west-central Honduras Notes on taxonomy: This species was reported from the Reserva de la Biosfera Río Plátano as Centrolenella fleischmanni by Cruz
Order, Family: Anura, Centrolenidae along the middle portion of the Río Patuca, western Olancho and (1978).
Country Distribution: Honduras Quebrada Las Marias near La Colonia, north-western Olancho, at Bibliography: Cruz Diaz, G.A. (1978), McCranie, J.R. and Wilson, L.D. (1997b), McCranie, J.R. and Wilson, L.D. (2002b)
Current Population Trend: Decreasing elevations of 100-700m asl. Data Providers: Gustavo Cruz, Larry David Wilson
Population It is moderately common but the population is de-
creasing.
Habitat and Ecology It occurs and breeds along streams in lowland
moist forest and premontane wet forest.
Major Threats The main threats to the species are deforestation,
forest fires, and water pollution. In addition, much of its range
is threatened by the development of the Patuca 2 hydroelectric
project.
Conservation Measures It occurs in Parque Nacional Río Patuca,
and it has been collected at the edge of the Río Platano and the
Asang Launa Tawahka Biosphere Reserves.

CR Hyalinobatrachium crybetes McCranie and Wilson, 1997

Critically Endangered B1ab(iii)+2ab(iii) Geographic Range This species is known only from the vicinity Bibliography: McCranie, J.R. and Wilson, L.D. (1997b), McCranie, J.R. and Wilson, L.D. (2002b)
Order, Family: Anura, Centrolenidae of Catacamas, Olancho department, east-central Honduras, at an Data Providers: Gustavo Cruz, Larry David Wilson
Country Distribution: Honduras elevation of 500-680m asl.
Current Population Trend: Decreasing Population It is rare and is known only from a small section of a
disturbed stream margin. It has not been seen since the first specimen
was collected in 1967. McCranie and Wilson (1997b), collecting at
the same locality, did not find this species.
Habitat and Ecology It was collected in vegetation along a small
stream, in lowland moist forest. It presumably lays eggs in vegetation
overhanging streams.
Major Threats Habitat loss due to agriculture, logging, and human
settlement is the major threat to this species.
Conservation Measures It is not known to occur in any protected
areas. Further survey work is required to determine whether or not
this species is still extant in the wild.

EN Hyalinobatrachium esmeralda Ruíz-Carranza and Lynch, 1998

Endangered B1ab(iii) Geographic Range This species is known from two sites 50km Bibliography: Barrera-Rodriguez, M. (2000), Ruiz-Carranza, P.M. and Lynch, J.D. (1998)
Order, Family: Anura, Centrolenidae apart, in Municipio Pajarito and Municipio Garagoa, in Boyacá De- Data Providers: Jose Vicente Rueda, Andrés Acosta-Galvis
Country Distribution: Colombia partment, on the Cordillera Oriental of the Colombian Andes, between
Current Population Trend: Decreasing 1,600 and 1,750m asl, though it might occur more widely.
Population It is a common species.
Habitat and Ecology It occurs on vegetation near streams in cloud
forest; the eggs are laid on the vegetation, and the larvae develop
in the water. It also occurs in secondary forest.
Major Threats The main threat to this species is habitat loss
caused by the expansion of cattle ranching and the planting of il-
legal crops. Pollution, resulting from the fumigation of illegal crops,
is also a threat.
Conservation Measures It does not occur in any protected areas,
making habitat protection a much-needed conservation action for
this species.
 Globally Threatened Amphibian Species 219

VU Hyalinobatrachium fragile (Rivero, 1985)

Vulnerable B1ab(iii) Geographic Range This species is known from several localities in the Cordillera de la Costa, including La Sierra in
Order, Family: Anura, Centrolenidae Cojedes State, and Sierra de Aroa in Yaracuy State, in Venezuela. It has been recorded from 100-700m asl.
Country Distribution: Venezuela Population It is probably an uncommon species.
Current Population Trend: Decreasing Habitat and Ecology It inhabits montane forest environments, where it occurs along streams. It lays its eggs on
the under side of leaves overhanging streams, and when hatched the larvae fall into the stream below where they
develop further.
Major Threats The major threat to this species is habitat loss due to agriculture (both crops and livestock), as well
as logging. Avalanches are a potential threat along the coast, especially during heavy rainfall periods.

© Francisco José López-López

Conservation Measures It is present in several protected areas, including Parque Nacional Henri Pittier and
Rancho Grande.
Notes on taxonomy: In the past, some specimens of this species were considered to be Hyalinobatrachium fleishmanni.
Bibliography: Barrio Amorós, C.L. (2004), La Marca, E. (1992), La Marca, E. (1997), Myers, C.W. and Donnelly, M.A. (1997), Rivero, J.A.
(1985), Ruiz-Carranza, P.M. and Lynch, J.D. (1991a)
Data Providers: Enrique La Marca, Celsa Señaris

EN Hyalinobatrachium guairarepanensis Señaris, 2001

Endangered B1ab(iii) Geographic Range This species is known from the central part of Conservation Measures Its range includes Parque Nacional El Avila and Parque Nacional Macarao. Further research
Order, Family: Anura, Centrolenidae the Venezuelan coastal mountain range on the Cordillera de la Costa, is required to determine the reasons for its decline in pristine habitat. If disease is shown to be a major threat, then
Country Distribution: Venezuela at elevations between 720 and 1,000m asl. surviving individuals may need to form the basis for the establishment of an ex-situ population.
Current Population Trend: Decreasing Population The population of this species is in decline, even in Bibliography: Barrio Amorós, C.L. (2004), Rivero, J.A. (1961), Rivero, J.A. (1968a), Señaris, J.C. (2001)
pristine habitats. It is associated with Atelopus cruciger, which Data Providers: Enrique La Marca, Celsa Señaris
has disappeared from almost all of its range, probably because of
chytridiomycosis.
Habitat and Ecology It occurs along streams in seasonal (semi-
caducifolious) forests. It lays its eggs on leaves overhanging streams,
and when hatched the larvae fall into the stream below where they
develop further.
Major Threats Agriculture, logging, water pollution, and infrastruc-
ture development for human settlement are all major threats to the
species’ habitat. However, it is declining even in pristine habitats,
suggesting that chytridiomycosis, or some other disease, may be
playing a role in the observed declines.

VU Hyalinobatrachium ibama Ruíz-Carranza and Lynch, 1998

Vulnerable B1ab(iii) Geographic Range This species occurs on the western slopes of the Cordillera Oriental in Norte de Santander
Order, Family: Anura, Centrolenidae and Santander Departments, Colombia, between 1,600 and 2,050m asl. It probably occurs a little more widely than
Country Distribution: Colombia current records suggest.
Current Population Trend: Decreasing Population It is common, and there are many recent records.
Habitat and Ecology It occurs in riparian habitats in old-growth forests. It lays its eggs on vegetation, and the
© Conservacion Internacional-Colombia

tadpoles develop in streams.

Major Threats Habitat loss caused by agricultural expansion (cattle ranching, planting of illegal crops, and espe-
cially onion farming) is the main threat to this species. Pollution, resulting from the fumigation of illegal crops, is
also a threat.
Conservation Measures It occurs in Santuario de Fauna y Flora Guanentá, Alto Río Fonce and Area Natural Unica
Los Estoraques.
Bibliography: Ruiz-Carranza, P.M. and Lynch, J.D. (1991b), Ruiz-Carranza, P.M. and Lynch, J.D. (1998)
Data Providers: Jose Vicente Rueda, Mariela Osorno-Muñoz

VU Hyalinobatrachium orientale (Rivero, 1968)

Vulnerable B1ab(iii) Geographic Range The nominate subspecies is known only from the eastern sector of the Venezuelan Coastal Range,
Order, Family: Anura, Centrolenidae with records from Serranía del Interior, Macizo de Turimiquire, and Serranía de Paria in the states of Monagas and
Country Distribution: Trinidad and Tobago, Sucre, at elevations of 190-1,200m asl. The subspecies H. o. tobagoensis is recorded from north-eastern Tobago in
Venezuela Trinidad and Tobago. Reports from Colombia are incorrect.
Current Population Trend: Decreasing Population It is a common species across its range.
Habitat and Ecology A nocturnal species that lives on Heliconia vegetation over-hanging cascading streams about
2-6m above the water in tropical rainforest. The eggs are laid on the underside of leaves, and the larvae develop in
the water. It has been found only in mature, closed-canopy forest.
Major Threats In Venezuela, although some of its habitat is within protected areas, other areas are being lost due
to human settlement and agricultural expansion. There are no known threats to the species in Tobago.
© Juan Elías García-Pérez

Conservation Measures It occurs in the Parque Nacional Peninsula de Paria in Venezuela. The rainforest in which
this species occurs in Tobago is protected in the Little Tobago Wildlife Sanctuary. Taxonomic work is required to
investigate whether or not populations of this species from Tobago represent a distinct species.
Notes on taxonomy: It is possible that two species might be included in Hyalinobatrachium orientale in Venezuela, one being the
species from the type locality, and the other an undescribed form from Macizo del Turimiquire. Animals on Tobago might also belong
to a different species.
Bibliography: Barrio Amorós, C.L. (2004), Cannatella, D.C. and Lamar, W.W. (1986), Duellman, W.E. (1977), Hardy, Jr, J.D. (1982),
Hardy, Jr, J.D. (1984), La Marca, E. (1992), La Marca, E. (1995b), La Marca, E. (1997), Murphy, J.C. (1997), Myers, C.W. and Donnelly,
M.A. (1997), Rivero, J.A. (1968a), Rivero, J.A. (1985), Ruiz-Carranza, P.M. and Lynch, J.D. (1991a), Ruiz-Carranza, P.M., Ardila-Robayo,
M.C. and Lynch, J.D. (1996)
Data Providers: Celsa Señaris, Enrique La Marca, Jerry Hardy
220 Threatened Amphibians of the World

EN Hyalinobatrachium pallidum (Rivero, 1985)

Endangered B1ab(iii) Geographic Range This species is known from Guacharaquita, in E. (1992), La Marca, E. (1997), Mijares-Urrutia, A. (1990), Myers, C.W. and Donnelly, M.A. (1997), Rivero, J.A. (1985), Ruiz-Carranza,
Order, Family: Anura, Centrolenidae the state of Táchira, in the Venezuelan Andes at 1,768m asl, and from P.M. and Lynch, J.D. (1991a)
Country Distribution: Venezuela Mundo Nuevo, in the state of Cojedes, at 396m asl. Data Providers: Enrique La Marca, Jesús Manzanilla
Current Population Trend: Decreasing Population It is not a common species.
Habitat and Ecology It occurs in evergreen dry forest in the Andes,
and humid forests in the southern central part of the Venezuelan
coastal mountain tange. Eggs are laid on the underside of leaves
in humid forest near streams, into which the larvae then fall to
complete their development.
Major Threats The habitat of the Andean populations has suffered
from intensive exploitation and degradation in the past, and these
populations are now almost extirpated.
Conservation Measures The range of this species does not include
any protected areas. Surveys are needed to determine the current
population status of the Andean population.
Bibliography: Barrio Amorós, C.L. (2004), Duellman, W.E. (1999), La Marca,

EN Hyalinobatrachium pellucidum (Lynch and Duellman, 1973)

Endangered B1ab(iii)+2ab(iii) Geographic Range This species is known from three localities: Río Azuela, Río Reventador and Río Salado, all in
Order, Family: Anura, Centrolenidae Napo Province on the Amazonian slopes of the Ecuadorian Andes, at 1,740m asl.
Country Distribution: Ecuador Population It was last recorded in 1979, when five specimens were collected from one site. The Río Azuela site has
© Natural History Museum and Biodiversity Research

Current Population Trend: Decreasing been revisited several times without locating any individuals.
Habitat and Ecology It lives in cloud forest. The holotype was found on the leaf of a herb over a small stream at
night (Lynch and Duellman 1973). Reproduction is presumed to occur via eggs laid on vegetation, with the larvae
developing in streams.
Major Threats Habitat destruction and degradation is the main threat to this species, primarily due to smallholder
farming and logging. At this stage it is not clear whether or not this species might have declined within pristine
Center, University of Kansas

habitats.
Conservation Measures It is not known to occur in any protected areas, and protection of remaining cloud forest habi-
tats is sorely needed. Further survey work is necessary to determine the current population status of this species.
Bibliography: Lynch, J.D. and Duellman, W.E. (1973), Ruiz-Carranza, P.M. and Lynch, J.D. (1991a)
Data Providers: Luis A. Coloma, Santiago Ron, Diego Cisneros-Heredia

VU Hyalinobatrachium revocatum (Rivero, 1985)

Vulnerable B1ab(iii) Geographic Range This species is known from the southern Bibliography: Barrio Amorós, C.L. (2004), Duellman, W.E. (1993), La Marca, E. (1997), Rivero, J.A. (1961), Rivero, J.A. (1968a), Rivero,
Order, Family: Anura, Centrolenidae slopes of the central part of Cordillera de la Costa, and Serranía J.A. (1985), Ruiz-Carranza, P.M. and Lynch, J.D. (1991a)
Country Distribution: Venezuela del Interior, in the Venezuelan coastal range, at elevations between Data Providers: Enrique La Marca, Celsa Señaris
Current Population Trend: Decreasing 1,200 and 1,800m asl.
Population It is a common frog in undisturbed habitats.
Habitat and Ecology It occurs along streams, and is usually found
on overhanging vegetation up to 2m above the stream. The eggs are
laid on the upper and under sides of leaves, and when hatched the
larvae fall into the stream below where they develop further.
Major Threats The major threat is habitat loss, due to agriculture
(both crops and livestock), logging, and infrastructure development
for human settlement. Agricultural pollution is also having an impact
on populations.
Conservation Measures Some populations lie within the El Avila,
Macarao and Henri Pittier and Rancho Grande National Parks.

DENDROBATIDAE

CR Aromobates nocturnus Myers, Paolillo and Daly, 1991 SKUNK FROG

Critically Endangered A2a; B2ab(v) Geographic Range This species is known only from the type Nacional Dinira. Surveys are urgently needed to determine the population status of this species.
Order, Family: Anura, Dendrobatidae locality: about 2km ESE by air from Agua de Obispo, in the state of Bibliography: Barrio Amorós, C.L. (2001), Barrio Amorós, C.L. (2004), La Marca, E. (1992), La Marca, E. (1996b), Myers, C.W., Paolillo,
Country Distribution: Venezuela Trujillo, Venezuela, at 2,250m asl (9.42’N, 70.05’W). A. and Daly, J.W. (1991)
Current Population Trend: Decreasing Population It is a very rare species that appears to have declined. Data Providers: Enrique La Marca, Juan Elías García-Pérez
Several attempts to find it in recent years have failed, and it is still
known only from the time it was first described.
Habitat and Ecology It occurs in small cold-water streams in cloud
forest. The eggs are laid on land and the male protects the eggs.
When hatched the male carries the larvae on his back to water
where they develop further.
Major Threats Habitat loss and degradation due to agriculture (both
crops and livestock) and road construction, is the major threat to the
species. The locality is near the headwaters of a stream and thus
could have been affected by a drought in the past. The species could
also potentially be affected by disease, such as chytridiomycosis.
Conservation Measures There is a population within the Parque
 Globally Threatened Amphibian Species 221

CR Colostethus anthracinus Edwards, 1971

Critically Endangered A2ac Geographic Range This species occurs in a narrow altitudinal Conservation Measures The range of this species overlaps Parque Nacional Cajas and its southern limit is adjacent
Order, Family: Anura, Dendrobatidae zone (2,710-3,500m asl) on the Cordillera Oriental (Cordillera Real to the northern limit of Parque Nacional Podocarpus. Surveys are urgently needed to establish the population status
Country Distribution: Ecuador or Central), and the Mazán River in southern Ecuador. There are of this species.
Current Population Trend: Decreasing records from six localities. Bibliography: Coloma, L.A. (1995), Edwards, S.R. (1971)
Population The most recent record of this species is from December Data Providers: Luis A. Coloma, Santiago Ron, Taran Grant, Manuel Morales
1991. An extensive search in Mazán in 1995 (where the species
was abundant in 1986) turned up no individuals. It appears to have
undergone a serious decline.
Habitat and Ecology It lives in páramo, very humid montane for-
est, and lower humid montane forest (Coloma 1995). Reproduction
probably occurs by the females laying eggs on the ground, and the
males bringing the tadpoles to streams for development.
Major Threats The primary threat to this species is habitat loss and
degradation due to agriculture, and logging. The possible disappear-
ance of the species could be related to chytridiomycosis.

VU Colostethus awa Coloma, 1995

Vulnerable B1ab(iii) Geographic Range This species is known from the western Andean slopes and in the western Pacific lowlands of
Order, Family: Anura, Dendrobatidae north-western Ecuador. It occurs from 265-1,220m asl.
Country Distribution: Ecuador Population It is not uncommon.
Current Population Trend: Decreasing Habitat and Ecology It inhabits humid sub-montane tropical forest, and lays its eggs outside water on leaf-litter or
low vegetation. The adults then carry the tadpoles on their back to nearby streams where they develop further.
Major Threats The major threats are habitat loss due to agriculture (both crops and livestock) and logging, and
agricultural pollution.
Conservation Measures Its range overlaps Reserva Ecológica Cotacachi-Cayapas, and slightly overlaps Reserva
© H. Mauricio Ortega-Andrade

Ecológica Mache-Chindul and Reserva Ecológica Los Illinizas.

Bibliography: Coloma, L.A. (1995), Morales, M. et al. (2002), Parker III, T.A. and Carr, J.L. (1992)
Data Providers: Luis A. Coloma, Santiago Ron, Mario Yánez-Muñoz, Diego Cisneros-Heredia, Ana Almandáriz

EN Colostethus cevallosi Rivero, 1991

Endangered B1ab(iii) Geographic Range This species is known from the eastern side of the Andes, in Ecuador, in two general areas: four
Order, Family: Anura, Dendrobatidae localities between 480 and 970m asl in Pastaza Province in central Ecuador, and Centro Shuar Yawi, at 920-1,040m
Country Distribution: Ecuador asl, in Zamora Chinchipe Province in south-eastern Ecuador. Records from Peru require confirmation.
Current Population Trend: Decreasing Population It was discovered in Zamora Chinchipe Province in 2003. However, surveys in Pastaza Province have not
turned up any additional records of this species since it was first discovered.
Habitat and Ecology It occurs in eastern Cordillera Real montane forests and napo moist forests. The distribution
lies mainly within very humid premontane forest and pluvial premontane forest. As with other members of the genus,
it probably breeds on the ground, under rocks or in leaf-litter, and the larvae are carried by the parent to streams.
Most members of this genus do not adapt well to anthropogenic disturbance.
Major Threats The main threat to the species is habitat loss and degradation due to agriculture, involving both crops
and livestock, as well as logging and infrastructure development for human settlement.
Conservation Measures It is not known to occur in any protected areas, and some form of formal habitat protection
© Morley Read

is urgently needed. Further survey work is necessary to determine the status of the the subpopulation in Pastaza
Province.
Bibliography: Almendariz, A. (1991), Coloma, L.A. (1995), Rivero, J.A. (1991)
Data Providers: Luis A. Coloma, Diego Almeida, Fernando Nogales, Ana Almandáriz, Diego Cisneros-Heredia, Karl-Heinz Jungfer,
Santiago Ron

VU Colostethus chalcopis Kaiser, Coloma and Gray, 1994 MARTINIQUE VOLCANO FROG
Vulnerable D2 Geographic Range This species is found on the south-eastern slope Bibliography: Breuil, M. (2004), Hedges, S.B. (1999), Hedges, S.B. (2001), Henderson, R.W. and Powell, R (1999), Henderson, R.W. and
Order, Family: Anura, Dendrobatidae of Montagne Pelee, Martinique, at 500m asl. Powell, R. (2001), Kaiser, H. and Altig, R. (1994), Kaiser, H. and Henderson, R.W. (1994), Kaiser, H., Coloma, L.A. and Gray, H.M. (1994),
Country Distribution: Martinique Population It is relatively common in its tiny range. Schwartz, A. and Henderson, R.W. (1991)
Current Population Trend: Stable Habitat and Ecology It lives in grassy areas on lava flows. Eggs Data Providers: Blair Hedges, Beatrice Ibéné, Michel Breuil, Robert Powell
are laid on the ground in a nest under leaves and guarded by the
adults, and presumably the adults transport the larvae to water, as
with other members of the genus.
Major Threats Although there are no current major threats to the
species, its small range render it at severe risk of stochastic events,
especially from volcanic eruptions.
Conservation Measures It does not occur in any protected areas,
and there is a need for improved protection of the Montagne Pelee.
The species requires close population monitoring, given that it is
restricted to a single location.
Notes on taxonomy: It has been suggested that this species is not native to
Martinique, but is synonymous with another species from South America (M.
Breuil pers. comm.).
222 Threatened Amphibians of the World

CR Colostethus delatorreae Coloma, 1995

Critically Endangered B2ab(iii,iv,v) C2a(ii) Geographic Range This species is known only from three nearby localities at elevations of 2,340-2,700m on the
Order, Family: Anura, Dendrobatidae western slopes of the Andes in Carchi Province, in extreme north-western Ecuador.
Country Distribution: Ecuador Population It is locally uncommon. Fifty calling males were recorded in 2003 in Moran, Carchi Province. However,
Current Population Trend: Decreasing it appears to have disappeared at all other localities.
Habitat and Ecology It occurs in wetlands and bogs, and in areas of cattle ranching surrounded by forest remnants.
It lays its eggs on the ground, and the males carry the tadpoles to water where they develop further.
Major Threats Habitat destruction and degradation is a significant threat to this species, due mainly to agriculture

© Mario Humberto Yánez-Muñoz

(including livestock ranching) and logging. It is possible that this species has been impacted by chytridiomycosis,
which has been confirmed in other frog species within its range.
Conservation Measures The species is not known to occur in any protected areas, and there is a need for urgent
habitat protection at sites at which it is known to survive.
Bibliography: Coloma, L.A. (1995)
Data Providers: Luis A. Coloma, Santiago Ron, Mario Yánez-Muñoz

CR Colostethus dunni (Rivero, 1961)

Critically Endangered A2ace Geographic Range This species is known from the central part
Order, Family: Anura, Dendrobatidae of the Venezuelan Coastal Range, in the vicinity of Caracas, from
Country Distribution: Venezuela around 800 to at least 1,520m asl. Its presence has been suggested,
Current Population Trend: Decreasing but not confirmed, in Quebrada de Caurimare, Miranda State, also
in the vicinity of Caracas.
Population This species was once considered one of the more
common species of the Caracas Valley; however, comprehensive
surveys in 1992-1993 and, more recently, in 2004 have failed to find
any populations in these areas. It is not known when exactly the
decline began, but the decline of Atelopus cruciger, another formerly
common frog from the same region, has been documented. It is
possible that Colostethus dunni has experienced a similar decline.
There are no known extant populations in the wild at present, even
in suitable habitat.
Habitat and Ecology It inhabits seasonal montane forests. Its
breeding habits are unknown, although the larvae are likely to
develop in water, like other species of the genus.
Major Threats The major threats to the species include habitat loss due to urban development and fires, and pollu-
tion. The cause(s) of the recent dramatic decline have not been established, but it is charecteristic of declines seen
in other species impacted by chytridiomycosis.
Conservation Measures Some populations might be protected in Parque Nacional El Avila, near Caracas. Research
into the cause(s) of the population decline is urgently needed, as are surveys to determine the current status of
populations.
Bibliography: Barrio Amorós, C.L. (2004), Barrio, C. (1998), La Marca, E. (1992), La Marca, E. (1997), La Marca, E. (2004), Rivero, J.A.
(1961)
Data Providers: Enrique La Marca, Jesús Manzanilla, Celsa Señaris

CR Colostethus edwardsi Lynch, 1982

Critically Endangered A2ac; B1ab(iii,iv,v)+ Geographic Range This species is known from two localities in the Cordillera Oriental, La Cueva de las Moyas and
2ab(iii,iv,v) the Páramo de Cruz Verde, Cundinamarca Department, Colombia, between 3,030 and 3,300m asl.
Order, Family: Anura, Dendrobatidae Population It was last seen in 1996. Three visits to the type locality have failed to turn up this species, and it is
Country Distribution: Colombia likely that it has declined severely due to complete habitat alteration.
Current Population Trend: Decreasing Habitat and Ecology It occurs in streams within caves and crevices. Breeding and larval development takes place
in streams.
Major Threats It is threatened by habitat loss due to conversion to timber forestry and water pollution caused by
urban runoff.
Conservation Measures The species is not known to occur in any protected areas. Surveys are needed to determine
the current population status of the species.
Bibliography: Lynch, J.D. (1982b), Rivero, J.A. (1988), Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and Lynch, J.D. (1996)
Data Providers: Jose Vicente Rueda
© John D. Lynch

EN Colostethus elachyhistus Edwards, 1971

Endangered A2ac; B2ab(iii,iv,v) Geographic Range This species occurs on the western slopes of the Andes in southern Ecuador, at elevations of
Order, Family: Anura, Dendrobatidae 850-2,760m asl. It also occurs on the Pacific and Amazonian slopes of the Andes in the Huancabamba depression,
Country Distribution: Ecuador, Peru Piura Department, northern Peru, at 600-1,800m asl.
Current Population Trend: Decreasing Population It is an uncommon species. In Ecuador, populations from Loja and nearby localities at the higher altitudes
of its range seem to be extinct, but lowland populations are still extant with records as recently as 2003.
Habitat and Ecology It lives in dry and humid lowland and premontane forest, and occurs near streams, especially
in dry forests. The larvae presumably develop in streams.
Major Threats The cause of the declines at high elevations, which have taken place even in suitable habitat, could
be due to chytridiomycosis. Habitat destruction and degradation, in particular due to agricultural activities, is also a
threat to this species, and the species may also be impacted by invasive alien species and pollution.
Conservation Measures It does not occur in any protected areas, and some form of formal habitat protection is
urgently needed. Further survey work is needed to determine whether any animals survive around Loja and other
© John D. Lynch

nearby localities.
Bibliography: Coloma, L.A. (1995), Duellman, W.E. and Wild, E.R. (1993), Edwards, S.R. (1971), Rodríguez, L.O., Cordova, J.H. and
Icochea, J. (1993)
Data Providers: Javier Icochea, Luis A. Coloma, Santiago Ron
 Globally Threatened Amphibian Species 223

VU Colostethus humilis Rivero, 1978

Vulnerable D2 Geographic Range This species is known from the type locality, Conservation Measures One population is within Parque Nacional Guaramacal (at the limit of the range).
Order, Family: Anura, Dendrobatidae in the vicinity of the town of Boconó, in Trujillo State, Venezuela, at Bibliography: Barrio Amorós, C.L. (2004), Barrio-Amorós, C.L. and Garcia-Porta, J. (2003), García-Pérez, J.E. (1999), La Marca, E. (1992),
Country Distribution: Venezuela an altitude of 1,470m asl, and from a recently discovered population La Marca, E., Vences, M. and Lötters, S. (2002), Péfaur, J.E. and Rivero, J.A. (2000), Rivero, J.A. (1978), Vial, J.L. and Saylor, L. (1993)
Current Population Trend: Stable found inside the Parque Nacional Guaramacal, also close to Boconó. Data Providers: Enrique La Marca, Juan Elías García-Pérez, César Luis Barrio Amorós
It is also now known from the states of Barinas and Portugues. It has
been recorded from 600-1,800m asl.
Population It is locally common.
Habitat and Ecology It occurs in seasonal montane forest, and
cloud forests. It is associated with temporary ponds, where the larvae
presumably develop, but exact reproductive habits are unknown. It is
probably able to tolerate minor habitat disturbance.
Major Threats Human settlement, and associated infrastructure
development, is a threat to some populations. The lagoon at the
type locality has already disappeared. It might also be affected
by drought.

CR Colostethus jacobuspetersi Rivero, 1991

Critically Endangered B2ab(i,ii,iii,iv,v) Geographic Range This species was known from the inter-Andean valleys of Guayllabamba, Toachi, Chimbo, and
Order, Family: Anura, Dendrobatidae Cañar, and on the western slopes of the Andes in northern and central Ecuador, where it occupied areas between
Country Distribution: Ecuador 1,500 and 3,800m and was known from more than ten localities.
© Natural History Museum and Biodiversity Research

Current Population Trend: Decreasing Population There are no records of this species since the 1960s. Bustamante (2002) provides data of its absence at
Bosque Protector Cashca Totoras (Provincia Bolívar), where it used to be a common frog. It is possible that undetected
relict populations might still survive.
Habitat and Ecology It lives in mid- to high-elevation humid montane forest (Coloma 1995). It lays its eggs on the
ground, and males carry the eggs to small streams for development.
Major Threats The primary threat to this species is habitat destruction and degradation due to agriculture, clear
Center, University of Kansas

cutting, and human settlement; virtually no habitat remains where this species used to occur. Invasive alien species
are also a threat.
Conservation Measures The distribution range of the species overlaps Reserva Ecológica Los Illinizas, Reserva
Geobotánica Pululahua, and Reserva de Producción Faunística Chimborazo. Surveys are urgently required to determine
the population status and continued persistence of this species.
Bibliography: Bustamante, M. R. (2002), Coloma, L.A. (1995), Rivero, J.A. (1991)
Data Providers: Luis A. Coloma, Santiago Ron

CR Colostethus juanii Morales, 1994

Critically Endangered B2ab(i,iii) Geographic Range This species is known only from the area of the type locality, Villavicencio, in the foothills of the
Order, Family: Anura, Dendrobatidae Orinoco region, Meta Department, Colombia, at 580m asl, where it occurs in a single botanical garden surrounded
Country Distribution: Colombia by a substantial urban area.
Current Population Trend: Decreasing Population It is scarcely distributed in its tiny area of occupancy. There are few specimens known, and it was last
seen in 2002. Searches in outlying areas have not turned up any individuals.
Habitat and Ecology It is known from small patches of forest and from a botanical garden; it has not been found in
larger tracts of forest in nearby areas. It is a terrestrial and diurnal species, and it lays its eggs in the leaf-litter. The
males probably take care of the eggs and then move the larvae to small streams for development.
Major Threats Habitat fragmentation and destruction caused by the expansion of human settlement, and water
pollution, are the pimary threats to this species.
Conservation Measures This species is not recorded from any protected areas, and there is an urgent need for
protection of its remaining habitat. Further survey work is needed to determine the current population status of this
© John D. Lynch

species.
Bibliography: Morales, V.R. (1994)
Data Providers: Andrés Acosta-Galvis, Jose Vicente Rueda

EN Colostethus kingsburyi (Boulenger, 1918)

Endangered B1ab(iii) Geographic Range This species is known from a narrow altitudinal zone (1,140-1,300m asl) on the eastern slopes
Order, Family: Anura, Dendrobatidae of the Andes in the vicinity of Volcán Reventador and in the Río Pastaza trench, in Napo, Orellana and Pastaza
Country Distribution: Ecuador Provinces, Ecuador.
© Natural History Museum and Biodiversity Research

Current Population Trend: Decreasing Population It was formally abundant, at least judging by the large numbers of specimens in museums, but there
is little recent information.
Habitat and Ecology It occurs in premontane forest, where the annual mean precipitation is 2,000-4,000mm and the
annual mean temperature is 18-24°C. Reproduction probably occurs by the females laying the eggs on the ground,
with the males taking the larvae to streams for their development.
Major Threats Habitat destruction and degradation, mainly due to agricultural expansion (involving crops, plantations
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and livestock) and logging, is the main threat to this species. Its narrow altitudinal range might leave it vulnerable
to the effects of climate change.
Conservation Measures Its distribution range overlaps with several protected areas, including Parque Nacional
Sumaco Napo-Galeras, Parque Nacional Llanganatis, the Reserva Ecológica Antisana, and the Reserva Ecológica
Cayambe-Coca. This species is in need of close population monitoring.
Notes on taxonomy: Molecular data suggest that Colostethus kingsburyi is a species complex (L.A. Coloma pers. comm.).
Bibliography: Andersson, L.G. (1945), Boulenger, G.A. (1918), Coloma, L.A. (1995), Edwards, S.R. (1974b), Edwards, S.R. (1985)
Data Providers: Luis A. Coloma, Santiago Ron, Manuel Morales
224
CR Colostethus leopardalis Rivero, 1978
Critically Endangered A2ace; B2ab(v)
Order, Family: Anura, Dendrobatidae
Country Distribution: Venezuela
Current Population Trend: Decreasing
EN Colostethus mandelorum (Schmidt, 1932)
Endangered B1ab(iii)
Order, Family: Anura, Dendrobatidae
Country Distribution: Venezuela
Current Population Trend: Decreasing
EN Colostethus mertensi (Cochran and Goin, 1964)
Endangered B1ab(iii)
Order, Family: Anura, Dendrobatidae
Country Distribution: Colombia
Current Population Trend: Decreasing
VU Colostethus murisipanensis La Marca, 1997
Vulnerable D2
Order, Family: Anura, Dendrobatidae
Country Distribution: Venezuela
Current Population Trend: Stable
Threatened Amphibians of the World
Geographic Range This species is known only from a very restricted Conservation Measures There is a population within Parque Nacional Sierra Nevada (Mérida State), in the Ven-
area near Páramo de Mucubají, Sierra de Santo Domingo, in estado ezuelan Andes. Surveys are required to determine the population status of this species. Given the potential risk of
Mérida, in the Venezuelan Andes, from 2,435-3,300m asl. disease, there might be a need for the establishment of a captive-breeding programme.
Population It has not been seen in at least a decade, and its range Notes on taxonomy: We follow La Marca (1994a) in assigning this species to the genus Colosthethus, not Nephelobates.
is very much reduced. Bibliography: Barrio Amorós, C.L. (2001), Barrio Amorós, C.L. (2004), La Marca, E. (1994a), La Marca, E. and Lötters, S. (1997), La
Habitat and Ecology It inhabits páramo grassland and sub-páramo Marca, E. and Reinthaler, H.P. (1991), Lötters, S. (1996), Mijares-Urrutia, A. (1991), Péfaur, J.E. and Rivero, J.A. (2000), Rivero, J.A.
shrubland environments and cloud forest, the highest environments (1976), Vial, J.L. and Saylor, L. (1993)
in the Venezuelan Andes, resembling alpine tundra, but with daily Data Providers: Enrique La Marca, Juan Elías García-Pérez
temperature extremes. It is usually found within grasses and frail-
ejones (Espeletia spp.), usually along and within streams.
Major Threats Introduced trout and possibly disease might be
threatening this species. This species shares its habitat with Atelopus
mucubajiensis, a páramo frog well known for having experienced a
dramatic population decline (La Marca and Reinthaler 1994; Lotters
1996; La Marca and Lotters 1997) probably due to chytridiomycosis,
and so this disease must also be suspected in C. leopardalis.
Geographic Range This species is restricted to the Cerro Bibliography: Barrio Amorós, C.L. (2004), Hardy, J.D. (1984), La Marca, E. (1993), La Marca, E. (1997), Péfaur, J.E. and Rivero, J.A. (2000),
Turimiquire, a mountainous region shared between Monagas, Sucre, Rivero, J.A. (1961), Rivero, J.A. (1982b), Vial, J.L. and Saylor, L. (1993)
and Anzoátegui States, in eastern Venezuela. It has been recorded Data Providers: Enrique La Marca, Jesús Manzanilla
from 1,900-2,630m asl.
Population It is a common species.
Habitat and Ecology It inhabits cloud forest and sub-páramo shru-
bland. The eggs are laid on land and when hatched the male carries
the larvae on his back to water, where they develop further.
Major Threats The main threat is habitat loss due to agriculture,
involving both crops and livestock, as well as road construction for
telecommunications installations.
Conservation Measures Its range includes a protected area for
a dam project (Zona Protectora Macizo Turimiquire), but the habitat
of this species is in need of much better protection.
Geographic Range This species is known from Cerro Munchique, Bibliography: Cochran, D.M. and Goin, C.J. (1964), Cochran, D.M. and Goin, C.J. (1970), Rivero, J.A. (1988), Ruiz-Carranza, P.M.,
and Quintana (near Popayan in Cauca Department), between 2,100 Ardila-Robayo, M.C. and Lynch, J.D. (1996)
and 2,350m asl, in Colombia. Data Providers: John Lynch, Fernando Castro
Population It is a common species.
Habitat and Ecology It occurs on the ground next to streams in cloud
forests. It lays its eggs on the ground, which (when hatched), are
transported on the backs of the male to temporary ponds where they
develop further. It has not been recorded outside forest habitat.
Major Threats The major threats are deforestation for agricul-
tural development, the planting of illegal crops, logging, and human
settlement.
Conservation Measures The range of the species includes Parque
Nacional Natural Munchique.
Geographic Range This species is known only from the type local-
ity, stated to be “Murisipan-Tepui (05 53N, 62 04W)”, in the state of
Bolívar, Venezuela, at 2,350m asl.
Population It is known from only one specimen.
Habitat and Ecology The single specimen was collected under
a rock in tropical montane forest. Breeding habits are unknown,
although the larvae probably develop in streams.
Major Threats There are no current major threats, but fires, caused
by lightning, are a potential threat.
Conservation Measures Its range does not include any protected
areas. There is a need for further survey work to determine the
current population status of this species, and to monitor its status,
given that it is known only from one site.
Bibliography: Barrio Amorós, C.L. (2004), La Marca, E. (1996b), La Marca,
E. (1997)
Data Providers: Enrique La Marca, Celsa Señaris
 Globally Threatened Amphibian Species 225

VU Colostethus olfersioides (A. Lutz, 1925)

Vulnerable A2ac Geographic Range This species is known from Rio de Janeiro State, northwards along the coast to Alagoas State,
Order, Family: Anura, Dendrobatidae in south-eastern Brazil. It occurs up to about 400m asl.
Country Distribution: Brazil Population It used to be a very common species, but it has recently declined and is now absent from several locali-
Current Population Trend: Decreasing ties at which it was previously common.
Habitat and Ecology This diurnal species occurs in the leaf-litter of primary and secondary forest. It can also tolerate
significant habitat disturbance and is found in gardens and plantations. They lay their eggs in temporary pools, and
after they hatch they are carried on the backs of the adults to streams where they complete their development.
Major Threats Habitat loss, due to deforestation and agricultural development, is ongoing, but probably has not
been severe enough to cause the declines that have been observed. Chytridiomycosis could be implicated, although
the species occurs well below the elevation at which conditions are normally suitable for chytrid.
Conservation Measures It is known to occur in several protected areas, such as Parque Nacional da Serra da
Bocaina. Research to determine the causes of the current decline is urgently needed.
© Ivan Sazima

Notes on taxonomy: Colostethus carioca, C. capixaba and C alagoanus are probably synonyms of this species (V. Verdade pers.
comm.).
Bibliography: Cochran, D.M. and Goin, C.J. (1970), Eterovick, P.C. et al. (2005), Izecksohn, E. and Carvalho-e-Silva, S.P. (2001), Lutz, B.
(1954), Rivero, J.A. (1988), Verdade, V.K. (2001)
Data Providers: Sergio Potsch de Carvalho-e-Silva, Vanessa Verdade

VU Colostethus pulchellus (Jiménez de la Espada, 1871)

Vulnerable A2ace; B1ab(i,ii,iii, iv,v) Geographic Range This species occurs in the inter-Andean páramos of southern Colombia (in Nariño and Cauca
Order, Family: Anura, Dendrobatidae Departments) and northern Ecuador (south to Cotapaxi Province), and on the eastern slopes of the Andes from southern
Country Distribution: Colombia, Ecuador Colombia south to the headwaters of the Río Pastaza in Ecuador. Its altitudinal range is 1,590-2,970m asl.
Current Population Trend: Decreasing Population It has disappeared from sites where it used to be abundant in Ecuador, but is still common at Cosanga
in Napo Province. It remains common in Colombia.
Habitat and Ecology An inhabitant of both dry and humid montane forests. It is a terrestrial species, and breeding
takes place in streams. It can be found in secondary growth near streams, but not in open areas.
Major Threats The major threat to this species is habitat loss as a result of agricultural development (including
© Francisco José López-López

the planting of illegal crops), logging, and human settlement; introduction of alien predatory fish in streams, and
pollution resulting from the spraying of illegal crops, are additional threats. There is heavy deforestation as a result
of wood extraction at Cosanga, where the species survives in Ecuador. The cause of declines observed in Ecuador is
not clear, but could be due to habitat loss, climate change, and/or chytridiomycosis.
Conservation Measures In Ecuador, its geographic range overlaps with the Parque Nacional Sumaco Napo-Galeras,
Reserva Ecológica Antisana, Parque Nacional Llanganates, and the Reserva Ecológica Cayambe-Coca. It occurs in
several protected areas in Colombia. The species requires close population monitoring, especially in view of the
potential risk of chytridiomycosis.
Notes on taxonomy: The populations from the Carchi basin might represent a distinct species (Coloma 1995).
Bibliography: Coloma, L.A. (1995), Jiménez de la Espada, M. (1875), Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and Lynch, J.D. (1996)
Data Providers: Wilmar Bolívar, Luis A. Coloma, Santiago Ron

EN Colostethus ranoides (Boulenger, 1918)

Endangered B1ab(i,ii,iii,iv,v)+2ab(i,ii,iii,iv,v) Geographic Range This species occurs north of Villavicencio, in introduced bullfrogs could eliminate the species from remaining forest fragments. Further research and survey
Order, Family: Anura, Dendrobatidae the eastern foothills of the Andes, Meta Department, Colombia, at work is required to better understand its current population status and in order to advance appropriate conservation
Country Distribution: Colombia 450-800m asl. measures for this species.
Current Population Trend: Decreasing Population It is rare, and was last seen in 2001, despite recent Notes on taxonomy: This species was removed from the synonymy of Colostethus brunneus by Silverstone (1971).
survey work in its range. Bibliography: Rivero, J.A. (1988), Rivero, J.A. and Serna, M.A. (1988), Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and Lynch, J.D. (1996),
Habitat and Ecology It is a terrestrial, diurnal species that occurs Silverstone, P.A. (1971)
in foothill forest and lays its eggs in leaf-litter. The males probably Data Providers: Andrés Acosta-Galvis, Jose Vicente Rueda, Fernando Castro
take care of the eggs, and then move the larvae to small streams
for their continued development.
Major Threats Habitat loss caused by agricultural expansion
is the main threat to this species, and oil extraction may pose a
future threat. It is also threatened by the introduced bullfrog, Rana
catesbeiana.
Conservation Measures Although it occurs in several small
private reserves, ex situ populations should be established because
the entire range of this species is nearly entirely deforested and

CR Colostethus ruizi Lynch, 1982

Critically Endangered B1ab(iii,v)+2ab(iii,v) Geographic Range This species is known from Fusagasuga, Bibliography: Lynch, J.D. (1982b), Rivero, J.A. (1988), Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and Lynch, J.D. (1996)
Order, Family: Anura, Dendrobatidae between Alto de San Miguel and Aguadita, on the western flank of Data Providers: Wilmar Bolívar, Jose Vicente Rueda
Country Distribution: Colombia the eastern Andes, Cundinamarca Department, Colombia, between
Current Population Trend: Decreasing 2,410 and 2,469m asl.
Population It is rare, and is known from only six individuals. Its
range has been completely deforested, except for one private tract
of land 11ha in size. Four expeditions since 1995 have failed to turn
up any individuals of this species.
Habitat and Ecology It lives on the forest floor and along streams
in cloud forest; it is not found outside old-growth forest. Its breeding
habits are not documented, but it is assumed that breeding and larval
development take place in streams.
Major Threats Deforestation for the planting of crops and cattle
grazing is the primary threat to this species.
Conservation Measures The species is not known to occur in any
protected areas. Surveys are urgently needed to determine whether
or not this species still persists in the wild.
226 Threatened Amphibians of the World

EN Colostethus ruthveni Kaplan, 1997

Endangered B1ab(iii) Geographic Range This species is known from the lower slopes of the north-western portion of the Sierra Nevada
Order, Family: Anura, Dendrobatidae de Santa Marta, in Magdalena Department, Colombia. It occurs between 680 and 1,540m asl.
Country Distribution: Colombia Population It is a common species.
Current Population Trend: Decreasing Habitat and Ecology It occurs along streams in cloud forest and dry tropical forests near the Caribbean coast. The
eggs are laid in bromeliads, and the larvae are then carried on the back of the adults to water where they develop
further.
Major Threats The major threat is habitat loss and degradation due to agriculture, involving both crops and livestock,
as well as logging, agricultural pollution, and infrastructure development.
Conservation Measures The species occurs in Parque Nacional Natural Sierra Nevada de Santa Marta and in the
adjacent, recently established El Dorado Nature Reserve.
Bibliography: Kaplan, M. (1997)
Data Providers: John Lynch, Martha Patricia Ramírez Pinilla, Mariela Osorno-Muñoz, Jose Vicente Rueda, Adolfo Amézquita, María
© Taran Grant

Cristina Ardila-Robayo

EN Colostethus saltuensis Rivero, 1978

Endangered B1ab(iii) Geographic Range Until recently, this species was only known from the type locality at 830m asl, on the road from
Order, Family: Anura, Dendrobatidae La Fria tio Michelena, in Táchira State, Venezuela. However, another population was found close by at a slightly
Country Distribution: Venezuela higher elevation (about 1,500m asl).
Current Population Trend: Decreasing Population It is not a common species; some populations are still found along the Andean piedmont facing Lake
Maracaibo.
© Conservacion Internacional-Colombia

Habitat and Ecology It occurs in humid lower montane Andean forests. The eggs are laid on land and the male
protects the eggs until they hatch when he carries the larvae on his back to water, where they develop further.
Major Threats The main threat is habitat loss and degradation due to agriculture, involving both crops and livestock,
as well as logging, water pollution, and infrastructure development for human settlement.
Conservation Measures It does not occur in any protected areas, and some form of formal protection is urgently
required to conserve the habitat of this species.
Bibliography: Barrio Amorós, C.L. (2004), La Marca, E. (1992), La Marca, E. (1997), Rivero, J.A. (1978)
Data Providers: Enrique La Marca, Juan Elías García-Pérez

EN Colostethus toachi Coloma, 1995

Endangered B1ab(iii) Geographic Range This species is known from nine localities in north-western Ecuador, from elevations of 200-
Order, Family: Anura, Dendrobatidae 1,410m asl.
Country Distribution: Ecuador Population It is a locally common species.
Current Population Trend: Decreasing Habitat and Ecology It occurs in humid tropical forest and very humid premontane forest (Coloma 1995), as well
as lowland grasslands. It lays its eggs on land in leaf-litter or under rocks, and the larvae are then carried to water
by the adults, where they develop further.
Major Threats The major threat is habitat loss and degradation due to agriculture, for both crops and livestock, as
© Mario Humberto Yánez-Muñoz

well as logging and agricultural pollution.

Conservation Measures Its range overlaps with the Reserva Ecológica Cotacachi-Cayapas.
Bibliography: Coloma, L.A. (1995), Morales, M. et al. (2002)
Data Providers: Diego Cisneros-Heredia, Ana Almandáriz, Mario Yánez-Muñoz, Luis A. Coloma, Santiago Ron

VU Colostethus vergeli (Hellmich, 1940)

Vulnerable D2 Geographic Range This species occurs on the western flank of the Cordillera Oriental, in Fusagasuga, Cundinamarca
Order, Family: Anura, Dendrobatidae Department, and Icononso, Tolima Department, Colombia, at 1,500-1,800m asl.
Country Distribution: Colombia Population It is known to be an abundant species, although no one has looked for it since 1994.
Current Population Trend: Stable Habitat and Ecology It occurs near streams in cloud forest, and seems able to tolerate some minor habitat disturbance.
The eggs are laid on land, and the males carry the tadpoles to streams where they develop further.
© Fundación ProAves / www.proaves.org

Major Threats There are no current known threats to this species, and it is somewhat resilient to habitat disturbance.
Nonetheless, its small range renders it susceptible to stochastic threatening processes.
Conservation Measures It does not occur in any protected areas. The species requires close population monitoring
given its very restricted range.
Notes on taxonomy: This species was considered a synonym of Colostethus fuliginosus by Edwards (1974b).
Bibliography: Edwards, S.R. (1971), Edwards, S.R. (1974a), Rivero, J.A. and Serna, M.A. (1988), Ruiz-Carranza, P.M., Ardila-Robayo,
M.C. and Lynch, J.D. (1996)
Data Providers: Adolfo Amézquita, Jose Vicente Rueda
 Globally Threatened Amphibian Species 227

CR Colostethus vertebralis (Boulenger, 1899)

Critically Endangered A2ace Geographic Range This species occurs in the inter-Andean valleys and on the western slopes of the Andes in
Order, Family: Anura, Dendrobatidae southern Ecuador, at elevations of 1,770-3,500m asl.
Country Distribution: Ecuador Population The most recent record of this species is from 1997 (QCAZ database). It was abundant in the Bosque
Current Population Trend: Decreasing Protector Mazán in 1986, but one year of searching in 1995 yielded only a single specimen. It was also abundant in
Laguna Busa in 1987. Recent expeditions to the range have turned up no specimens, and no surviving populations
are known.
Habitat and Ecology It occurs in cloud forest, in ponds in open areas and in streams. Eggs are placed on the ground,
and the males carry the tadpoles to streams where they develop further.
Major Threats The primary threat threat to this species is habitat destruction and degradation, due to agriculture
and clear cutting, but this does not account for the observed decline, and chytridiomycosis is implicated as the likely
cause of its apparent disappearance.
Conservation Measures The range of this species overlaps Parque Nacional Cajas and slightly overlaps Parque
© Eduardo Toral

Nacional Sangay. Survey work is urgently needed to determine whether or not this species survives in its natural
range.
Bibliography: Boulenger, G.A. (1899), Coloma, L.A. (1995)
Data Providers: Luis A. Coloma, Santiago Ron, Manuel Morales, Ana Almandáriz

VU Colostethus wayuu Acosta, Cuentas and Coloma, 1999

Vulnerable D2 Geographic Range This species is known only from the Parque Bibliography: Acosta, A.R., Cuentas, D.A. and Coloma, L.A. (1999)
Order, Family: Anura, Dendrobatidae Nacional Natural de Macuira, on the Sierranía of Macuira, in the Data Providers: Andrés Acosta-Galvis, Taran Grant
Country Distribution: Colombia Department of Guajira, Colombia. It has an altitudinal range of
Current Population Trend: Unknown 210-780m asl. It occurs in an isolated area of forest habitat in an
otherwise arid region, so it is very unlikely to occur anywhere else.
Population It is abundant in the rainy season.
Habitat and Ecology It lives in tropical forest with dry vegetation,
and can be found in secondary forest. The eggs are deposited on the
forest floor and the male carries the hatched tadpoles to streams
and ponds to complete their development.
Major Threats There are no current major threats to the species,
and its only known population is within a national park. However, its
restricted range renders it susceptible to threatening processes.
Conservation Measures It has been recorded from the Parque
Nacional Natural de Macuira, Colombia. The population of this
species requires close monitoring, particularly given that is known
only from a single location.

EN Cryptophyllobates azureiventris (Kneller and Henle, 1985)

Endangered B1ab(iii) Geographic Range This species is known only from the vicinity of the type locality of Tarapoto-Yurimaguas Road,
Order, Family: Anura, Dendrobatidae 700m asl (San Martín Department), Peru. It is presumed to have a restricted distribution.
Country Distribution: Peru Population It is not very common.
Current Population Trend: Decreasing Habitat and Ecology Its habitat is lowland primary tropical rainforest, and eggs are laid under leaf-litter and the
larvae are transported to small streams. Nothing is known about its adaptability to modified habitats.
Major Threats The major threat to the species is general habitat loss resulting from various human activities;
local people use the area for recreation and there are lots of settlements, with agricultural development, in lower
areas. The species has been recorded in trade. Specimens presumably originated from illegal collection by tourists,
although they may have been captive bred. However, the species has not been seen in the commercial trade over
the last fifteen years.
© Karl-Heinz Jungfer

Conservation Measures It is not present in any protected areas, and there is an urgent need to both encourage the
protection of remaining lowland habitat as well as establish a conservation management plan that includes some
form of legislation concerning trade.
Bibliography: Kneller, M. and Henle, K. (1985), Lötters, S., Jungfer, K.-H. and Widmer, A. (2000)
Data Providers: Javier Icochea, Ariadne Angulo, Karl-Heinz Jungfer, Stefan Lötters, Wilfredo Arizabal, Jorge Luis Martinez

CR Dendrobates abditus (Myers and Daly, 1976)

Critically Endangered A2ac; B1ab(iii) Geographic Range This species is only known from the eastern base of Volcán Reventador, south-west of the
Order, Family: Anura, Dendrobatidae Río Azuela bridge on the Quito-Lago Agrio road, in Napo Province, at 1,700m asl, on the Amazonian versant of the
Country Distribution: Ecuador Andes in Ecuador.
Current Population Trend: Decreasing Population It no longer survives at its only known locality, but there is a small chance that it survives elsewhere in
© Natural History Museum and Biodiversity Research

CITES: Appendix II localities that have not yet been surveyed.

Habitat and Ecology The type locality is a forested ridge, which has a relatively low canopy and is exceedingly
dense in most places. This forest is cool and very damp, with a conspicuous moss layer and many epiphytes on the
tree trunks. Specimens have been found by day, on or close to the ground, in the forest and in adjacent new clearings.
The eggs are laid on the ground, and the tadpoles are carried on the back of the adults one by one to bromeliads,
where they develop.
Center, University of Kansas

Major Threats Patches of forest at the type locality have been cleared for agriculture and livestock farming. The
species also could have been affected by the synergistic effects of chytridiomycosis and climate change.
Conservation Measures The distribution range of the species might overlap with Reserva Ecológica Cayambe-Coca,
but its presence there has not been confirmed. There is an urgent need for further survey work to determine whether
or not this species might survive at other sites near the type locality.
Bibliography: Myers, C.W. (1987), Myers, C.W. and Daly, J.W. (1976a)
Data Providers: Luis A. Coloma, Santiago Ron
228 Threatened Amphibians of the World

VU Dendrobates altobueyensis (Silverstone, 1975)

Vulnerable D2 Geographic Range This species has a very restricted range around INDERENA No. 39 of 9 July, 1985, forbids the collection of Dendrobates spp. from the wild for breeding (or other)
Order, Family: Anura, Dendrobatidae Alto del Buey, on the Serrania Baudo, in Choco Department, Colombia, purposes.
Country Distribution: Colombia between 980 and 1,070m asl. Bibliography: Myers, C.W. (1987), Rueda-Almonacid, J.V. (1999), Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and Lynch, J.D. (1996),
Current Population Trend: Stable Population There is little information on its population status. Silverstone, P.A. (1975)
CITES: Appendix II Habitat and Ecology It lives on the ground and in bromeliads in Data Providers: Wilmar Bolívar, Taran Grant
very humid pre-montane forests in the Pacific region of Colombia.
There is no information on breeding, but it is probable that the
eggs are laid in leaf-litter, and that the tadpoles are carried by the
adults to bromeliads. There is no information on its adaptability to
secondary habitats.
Major Threats There is very little information on threats to this
species, though it is likely that some populations are threatened by
habitat loss due mainly to smallholder agricultural activities, logging,
and human settlement. It might also be collected for the pet trade.
Conservation Measures It occurs in the Parque Nacional Natural
Utria. Management practices that could allow a commercial, sus-
tainable harvest of this species should be investigated. Decree

EN Dendrobates arboreus Myers, Daly and Martínez, 1984 POLKADOT POISON FROG
Endangered B1ab(iii) Geographic Range This species is endemic to the western cordilleras and Atlantic lowlands of Panama. It occurs
Order, Family: Anura, Dendrobatidae up to at least 1,120m asl.
Country Distribution: Panama Population There is no information available on the population status of this species.
Current Population Trend: Decreasing Habitat and Ecology This is a phytotelmic canopy species of humid lowland and montane forest, with breeding
CITES: Appendix II taking place in bromeliads.
Major Threats The main threat is habitat loss due to logging, livestock ranching and human settlement. The level
of trade in this species is not known. It is potentially at risk from chytridiomycosis.
Conservation Measures It has been recorded from the protected areas of Parque Internacional La Amistad and
Bosque Protector Palo Seco. Further research and survey work is needed to ascertain the possible negative impacts
of trade and chytrid.
Bibliography: Ibáñez, R. et al. (2000), Myers, C.W., Daly, J.W. and Martinez, V. (1984), Young, B. et al. (1999)
Data Providers: Frank Solís, Roberto Ibáñez, César Jaramillo, Querube Fuenmayor
© Marcos Guerra

VU Dendrobates azureus Hoogmoed, 1969 BLUE POISON FROG

Vulnerable D2 Geographic Range This species is known only from the vicinity of the type locality: “Sipaliwini, forest-island on
Order, Family: Anura, Dendrobatidae western slope Vier Gebroeders Mountain, 2°N; 55° 58’W, Surinam.” The type locality is at around 350m asl.
Country Distribution: Suriname Population It is a common species.
Current Population Trend: Stable Habitat and Ecology It is usually seen on rocks along streams in naturally occurring primary forest fragments (for-
CITES: Appendix II est/savannah mosaic). Males call from leaves on trees or from the ground. A clutch of 2-6 eggs are laid in leaf-litter
and attended to by the male, but also sometimes by the female. Males carry the tadpoles on their back to streams
where they develop further.
Major Threats It is not significantly threatened in its tiny range, but fire probably has an impact on its forest habitat
(though rumours from European dart frog collectors that a fire had rendered the wild populations extinct in 1992
were proved to be unfounded). Illegal collection of the species for the international pet trade probably is no longer
a significant threat, since the species is extensively bred in captivity.
Conservation Measures The range of the species is wholly within Sipaliwini Protected Area. In 1997 the National
Aquarium in Baltimore, in the USA, initiated a captive-breeding program, which has been very successful and now
© Paddy Ryan

involves more than 20 other institutions.

Bibliography: Hoogmoed, M.S. (1969), Zippel, K. (2005)
Data Providers: Robert Reynolds, Marinus Hoogmoed, Ross MacCulloch, Philippe Gaucher

EN Dendrobates bombetes (Myers and Daly, 1980) CAUCA POISON FROG

Endangered B1ab(iii) Geographic Range This species is known from both flanks of the western Andes, in the department of Valle del
Order, Family: Anura, Dendrobatidae Cauca, and from the western flank of the central Andes, in the departments of Quindio and Risaralda, in Colombia,
Country Distribution: Colombia between 1,580 and 2,100m asl.
Current Population Trend: Decreasing Population It is a common species, but it is believed to be declining.
CITES: Appendix II Habitat and Ecology It occurs on the lowest stratum and on bromeliads in cloud forests and very dry forests. Eggs
are laid in the leaf-litter, and the larvae develop in bromeliads.
Major Threats Habitat loss and degradation caused by agricultural expansion, timber extraction, crop fumigation,
and the removal of bromeliads is a major threat. This species is also collected illegally for the pet trade.
Conservation Measures It is not known to occur in any protected areas, and habitat protection is urgently needed.
Decree INDERENA No. 39 of 9 July, 1985, forbids the collection of Dendrobates spp. from the wild for breeding (or
other) purposes. It is also necessary to review international legislation regarding the illegal trade of this species.
© Juan Manuel Renjifo

Bibliography: Myers, C.W. (1987), Myers, C.W. and Daly, J.W. (1980), Rueda-Almonacid, J.V. (1999), Ruiz-Carranza, P.M., Ardila-Robayo,
M.C. and Lynch, J.D. (1996)
Data Providers: Martha Patricia Ramírez Pinilla, Mariela Osorno-Muñoz, Jose Vicente Rueda, Adolfo Amézquita, María Cristina
Ardila-Robayo
 Globally Threatened Amphibian Species 229

VU Dendrobates granuliferus Taylor, 1958 GRANULAR POISON FROG

Vulnerable B1ab(iii) Geographic Range This species occurs in the lowlands of south-western Costa Rica, adjacent south-western
Order, Family: Anura, Dendrobatidae Panama, and possibly in south-eastern Costa Rica, from 20-100m asl.
Country Distribution: Costa Rica, Panama Population It is common in Costa Rica.
Current Population Trend: Decreasing Habitat and Ecology A diurnal, terrestrial species most often found in relatively undisturbed humid lowland for-
CITES: Appendix II est, but also recorded from secondary forest and plantations. Eggs are deposited on the forest floor and the female
transports the tadpoles to bromeliads to complete metamorphosis.
Major Threats The major threat is habitat loss due to agriculture, selective logging, and human settlement. The
current level of offtake of specimens for the international trade is unknown.
Conservation Measures It is present in several protected areas, including Parque Nacional Corcovado.
Bibliography: Ibáñez, R. et al. (2000), Meyer, E. (1992), Meyer, E. (1993), Meyer, E. (1996), Savage, J.M. (2002), Silverstone, P.A. (1975),
Young, B. et al. (1999)
Data Providers: Frank Solís, Roberto Ibáñez, Gerardo Chaves, Jay Savage, César Jaramillo, Querube Fuenmayor
© Paddy Ryan

CR Dendrobates lehmanni Myers and Daly, 1976 LEHMANN’S POISON FROG

Critically Endangered B2ab(iii) drainage, Department of Valle del Cauca; and from Alto del Oso near San José del Palmar in the Department of Choco.
Order, Family: Anura, Dendrobatidae Its overall altitudinal rage is between 600 and 1,200m asl. Records from the Serrania de Baudo in the Department
Country Distribution: Colombia of Choco (not mapped) require confirmation.
Current Population Trend: Decreasing Population It is a locally common species in its tiny range.
CITES: Appendix II Habitat and Ecology It lives mostly on the ground in submontane tropical rainforests, but it can also be found perch-
ing on leaves up to 60cm above ground. It is not found in heavily degraded areas, but does occur in mature secondary
forest. The eggs are laid on the ground, and the tadpoles are carried on the backs of the females to bromeliads, where
they develop. The females feed the tadpoles on unfertilized eggs.
Major Threats The major threats are habitat loss and degradation, as a result of agricultural development (illegal
crops), logging, and human settlement, and pollution, resulting from the spraying of illegal crops. It occasionally
appears in the illegal pet trade.
© Karl-Heinz Jungfer

Geographic Range This species is known with certainty only from two localities in Colombia on the western slopes
of the Cordillera Occidental: west of Dagua at 850-1,200m on the south-facing versant of the upper Río Anchicayá
Conservation Measures The species occurs in the Parque Nacional Natural Farallones de Cali. Management practices
that could allow a commercial, sustainable harvest of this species should be investigated. Decree INDERENA No. 39
of 9 July, 1985, forbids the collection of Dendrobates spp. from the wild for breeding (or other) purposes.
Notes on taxonomy: Lötters (1992b) doubted the distinction of this species from Dendrobates histrionicus.
Bibliography: Bauer, L. (1979), Bechter, R. (1978), Fenolio, D. (1994), Garraffo, H.M. et al. (2001), Lötters, S. (1992b), Lötters, S. (1996),
Lötters, S. et al. (1999), Lötters, S. and Widmer, A. (1997), Myers, C.W. and Daly, J.W. (1976b), Myers, C.W., Daly, J.W. and Martinez,
V. (1984), Rueda-Almonacid, J.V. (1999), Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and Lynch, J.D. (1996), Schoop, E. (2002), van Dijk, W.
(2001), Zimmermann, H. and Zimmermann, E. (1980), Zimmermann, H. and Zimmermann, E. (1981)
Data Providers: Wilmar Bolívar, Fernando Castro, Stefan Lötters

EN Dendrobates mysteriosus Myers, 1982 MARAÑÓN POISON FROG

Endangered B1ab(iii) Geographic Range This species is currently known with certainty only from a single location in the vicinity of the
Order, Family: Anura, Dendrobatidae town of Santa Rosa at the foothills of the Cordillera del Cóndor (Cajamarca Department), north-eastern Peru, at
Country Distribution: Peru around 1,000m asl.
Current Population Trend: Decreasing Population The population status of this species is not known. Much of the area is deforested and populations are
CITES: Appendix II only known from forest remnants that remain on some steep cliffs and slopes.
Habitat and Ecology It has only been recorded from primary premontane forest, with breeding taking place in
bromeliads.
Major Threats The main threat is habitat clearance for agricultural use (including pastureland and coffee cultivation).
Recently, a few specimens have also been reported in international trade.
Conservation Measures Reportedly present in the Cordillera del Cóndor Reserve, this species is also protected
by national legislation. Further research into the distribution and population status of the species is necessary, and
© Karl-Heinz Jungfer

there is a need to monitor its presence in international trade.

Bibliography: Myers, C.W. (1982), Rodríguez, L.O., Cordova, J.H. and Icochea, J. (1993), Schmidt, O. (1857), Schmidt, O. (1858), Schulte,
R. (1990), Schulte, R. (1999)
Data Providers: Javier Icochea, Karl-Heinz Jungfer, Stefan Lötters, Wilfredo Arizabal, Jorge Luis Martinez

VU Dendrobates opisthomelas Boulenger, 1899 ANDEAN POISON FROG

Vulnerable B1ab(iii) Geographic Range This species occurs from the central and western Andes, in Antioquia Department, to the eastern
Order, Family: Anura, Dendrobatidae slope of the central Andes, in Caldas Department, Colombia. It has been recorded between 1,160 and 2,200m asl. It
Country Distribution: Colombia might occur a little more widely than current records suggest.
Current Population Trend: Decreasing Population It is a common species.
CITES: Appendix II Habitat and Ecology It occurs on the ground in leaf-litter in montane forest, although it may also climb on vegetation
© Mauricio Rivera-Correa, courtesy of Museo de

in the interior of primary forest. It lays its eggs in leaf-litter and the tadpoles develop in bromeliads.
Major Threats The major threats to this species are habitat loss/degradation, due to agriculture, logging and fire,
and pollution as a result of the fumigation of crops. It is also harvested illegally for the international pet trade.
Herpetología Universidad de Antioquia

Conservation Measures It is not known to occur in any protected areas, and there is clearly a need for improved
habitat protection at sites where this species is known to occur. Measures need to be put in place to ensure levels of
offtake of this species from the wild are sustainable. Decree INDERENA No. 39 of 9 July, 1985, forbids the collection
of Dendrobates spp. from the wild for breeding (or other) purposes.
Bibliography: Myers, C.W. (1987), Rueda-Almonacid, J.V. (1999), Ruiz-Carranza, P.M. and Ramírez Pinilla, M. (1992), Ruiz-Carranza,
P.M., Ardila-Robayo, M.C. and Lynch, J.D. (1996), Silverstone, P.A. (1975)
Data Providers: Martha Patricia Ramírez Pinilla, Mariela Osorno-Muñoz, Jose Vicente Rueda, Adolfo Amézquita, María Cristina
Ardila-Robayo
230 Threatened Amphibians of the World

EN Dendrobates sirensis Aichinger, 1991

Endangered B1ab(iii) Geographic Range This species is known only from the Cordillera Bibliography: Aichinger, M. (1991)
Order, Family: Anura, Dendrobatidae El Sira (in the departments of Huanuco, Pasco, Ucayali), Peru, at an Data Providers: Javier Icochea, Karl-Heinz Jungfer, Ariadne Angulo
Country Distribution: Peru altitude of 750-1,000m asl. This is a single isolated mountain, so the
Current Population Trend: Decreasing species is probably restricted to this area.
CITES: Appendix II Population Nothing is known of its population status, although it
is likely to be rare.
Habitat and Ecology It is a diurnal species of primary montane
cloud forest, and is presumed to use phytotelms for breeding. It is
not known if it can occur in secondary or modified habitats.
Major Threats Although the restricted area that the species inhabits
is relatively inaccessible, there is some disturbance due to farming
and human settlement. In addition, the lower parts of the range are
subject to selective logging.
Conservation Measures It is present in the Reserva Comunal El
Sira, an indigenous people’s reserve. Strengthened management may
be necessary to ensure the maintenance of this species’ habitat.

EN Dendrobates speciosus O. Schmidt, 1857 SPLENDID POISON FROG

Endangered B1ab(iii) Geographic Range This species is a Panamanian endemic of the western Cordillera Central adjacent to Costa Rica.
Order, Family: Anura, Dendrobatidae It is present at around 1,370m asl.
Country Distribution: Panama Population The species was formerly considered to be common, but its current population status is not known.
Current Population Trend: Decreasing Habitat and Ecology It is a terrestrial species of humid lowland and montane forest, with breeding taking place
CITES: Appendix II in plants.
Major Threats The major threat is habitat loss due to logging and human settlement. It is also subject to collection
for the pet trade.
Conservation Measures The species has been recorded from the protected areas of Bosque Protector Palo Seco
and Parque Internacional La Amistad. Further research is needed into the current population status of the species,
and some form of management or legislation needs to be put in place in order to control the level of offtake for the
pet trade.
© Karl-Heinz Jungfer

Bibliography: Edwards, M.W., Daly, J.W. and Myers, C.W. (1988), Ibáñez, R. et al. (2000), Jungfer, K.H. (1985), Myers, C.W., Daly, J.W.
and Martinez, V. (1984), Oostveen, H. (1981), Savage, J.M. (1968b), Silverstone, P.A. (1975), Young, B. et al. (1999)
Data Providers: Frank Solís, Roberto Ibáñez, César Jaramillo, Querube Fuenmayor

CR Dendrobates steyermarki Rivero, 1971

Critically Endangered B2ab(iii) Geographic Range This species is restricted to Cerro Yapacana, at elevations from 600-1,300m asl, in Amazonas
Order, Family: Anura, Dendrobatidae state, Venezuela.
Country Distribution: Venezuela Population According to Gorzula and Señaris (1999) this frog is exceedingly common in bromeliads on the summit,
Current Population Trend: Decreasing and it is possible to collect hundreds in just a few hours. It is still found in great numbers.
CITES: Appendix II Habitat and Ecology It occurs in terrestrial bromeliads in montane tepui forest. Breeding probably takes place in
bromeliads.
Major Threats Intensive open gold mining in the area is a major threat to the species’ habitat. Barrio and Fuentes
(1999) reported the illegal collection and exportation of some 150 specimens to Germany, and other collections for
scientific purposes. Wildfires might also be a threat to the species’ habitat.
Conservation Measures Cerro Yapacana is a Venezuelan natural monument, like most other tepuis in the region.
Surveys are necessary to monitor the population trends of this species. Measures are also required to combat illegal
© Karl-Heinz Jungfer

trade of this species, and to regulate the mining activities so that the survival of the species is not jeopardised.
Bibliography: Barrio Amorós, C.L. (2001), Barrio Amorós, C.L. (2004), Barrio, C.L. and Fuentes, O. (1999), Frost, D.R. (1985), Gorzula, S.
(1988), Gorzula, S. and Cerda, J. (1979), Gorzula, S. and Señaris, J.C. (1998), La Marca, E. (1992), La Marca, E. (1995b), La Marca, E. (1997),
Myers, C.W. (1987), Rivero, J.A. (1971), Rodríguez, J.P. and Rojas-Suárez, F. (1995), Silverstone, P.A. (1975), Walls, J.G. (1994)
Data Providers: Enrique La Marca, Celsa Señaris

VU Dendrobates viridis (Myers and Daly, 1976) GREEN POISON FROG

Vulnerable B1ab(iii) Geographic Range This species occurs on the western slopes of the Cordillera Occidental of Colombia, from Río
Order, Family: Anura, Dendrobatidae Anchicaya in Valle del Cauca Department, to Río Saija in Cauca Department, between 100 and 1,200m asl.
Country Distribution: Colombia Population It is very common.
Current Population Trend: Decreasing Habitat and Ecology It lives on the ground and in bromeliads in primary and good secondary, lowland and sub-
CITES: Appendix II montane forests. The tadpoles presumably are deposited in bromeliads.
Major Threats The major threats are deforestation due to agricultural development (including planting of illegal
crops), logging, and human settlement, and pollution resulting from the spraying of crops. Unlike other species of
the genus, it has not been recorded in international trade.
Conservation Measures It occurs in the Parque Nacional Natural Farallones. Decree INDERENA No. 39 of 9 July,
1985, forbids the collection of Dendrobates spp. from the wild for breeding (or other) purposes.
Bibliography: Myers, C.W. (1987), Myers, C.W. and Daly, J.W. (1976b), Rueda-Almonacid, J.V. (1999), Ruiz-Carranza, P.M., Ardila-
© Pedro Ruiz-Carranza

Robayo, M.C. and Lynch, J.D. (1996)

Data Providers: Wilmar Bolívar, Fernando Castro
 Globally Threatened Amphibian Species 231

EN Dendrobates virolinensis (Ruíz-Carranza and Ramírez-Pinilla, 1992)

Endangered B1ab(iii) Geographic Range This species occurs in the Santuaro de Fauna y Flora Guanentá, Alto Río Fonce, Río Cañaverales
Order, Family: Anura, Dendrobatidae and El Reloj; and also in Municipio de Socorro, Santander Department, Colombia, at 1,700-2,00m asl.
Country Distribution: Colombia Population It is common, with many recent records.
Current Population Trend: Decreasing Habitat and Ecology It lives on the floor of cloud forest, and only occurs in secondary forest if there is a lot of leaf-
CITES: Appendix II litter and bromeliads (since it lays its eggs in leaf-litter and the males bring the larvae to the bromeliads).
Major Threats Habitat fragmentation and loss is the main threat, and is taking place primarily due to agricultural
expansion. This species is also found in the pet trade.
Conservation Measures Although it occurs in the Santuaro de Fauna y Flora Guanentá, improved habitat protection
is required, and there is a need to monitor and regulate the offtake of this species for international trade. Decree
© Luis Fernando Cáceres INDERENA No. 39 of 9 July, 1985, forbids the collection of Dendrobates spp. from the wild for breeding (or other)
purposes.
Bibliography: Amezquita, A. (1995), Perez, G., Ruiz-Carranza, P.M. and Ramirez-Pinilla, M.P. (1992), Rueda-Almonacid, J.V. (1999),
Ruiz-Carranza, P.M. and Ramírez Pinilla, M. (1992), Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and Lynch, J.D. (1996), Vences, M. et al.
(2000b)
Data Providers: Adolfo Amézquita, Jose Vicente Rueda

VU Epipedobates cainarachi Schulte, 1989 CAINARACHI POISON FROG

Vulnerable B1ab(iii) Geographic Range This species is known only from the type locality and adjacent areas including the Tarapoto-
Order, Family: Anura, Dendrobatidae Yurimaguas road (San Martín Department) in Peru. It probably occurs more widely than is currently known. It has
Country Distribution: Peru been recorded up to 600m asl.
Current Population Trend: Decreasing Population There is nothing known about the population status of this species.
CITES: Appendix II Habitat and Ecology It inhabits lowland tropical moist forest and “rolling hills”, and can be found in primary
and possibly slightly degraded habitats. The breeding habitat is not known, although the larvae are presumably
deposited in water.
Major Threats The major threat is the loss of forest habitat through agriculture (coffee), livestock production, and
subsistence wood collection. The species has not been recorded from international trade.
Conservation Measures It is not recorded from any protected areas. Improved habitat protection is required at
sites where this species is known to occur.
© Karl-Heinz Jungfer

Bibliography: Jungfer, K.-H. (1989), Rodriguez, L. and Myers, C.W. (1993), Schulte, R. (1989)
Data Providers: Javier Icochea, Karl-Heinz Jungfer

CR Epipedobates ingeri (Cochran and Goin, 1970)

Critically Endangered B1ab(iii) Geographic Range This species is known only from the type Notes on taxonomy: Jungfer, Lötters and Jörgens (2000) suggest this species should revert to the genus Dendrobates.
Order, Family: Anura, Dendrobatidae locality: Asarrio, on the Pescado River in the Amazonian lowlands, Bibliography: Jungfer, K.-H., Lötters, S. and Jörgens, D. (2000), Rueda-Almonacid, J.V. (1999), Silverstone, P.A. (1976)
Country Distribution: Colombia Caqueta Department, Colombia, at 200m asl. It has not been found Data Providers: Andrés Acosta-Galvis, Jose Vicente Rueda
Current Population Trend: Decreasing elsewhere, and probably has a very small range.
CITES: Appendix II Population It is known from a single specimen, which was described
in 1970. Recent work in the region has not turned up any new
specimens. Given the level of habitat destruction that has taken
place within its range, it could be extinct.
Habitat and Ecology The area where it was collected was lowland
forest. It is probably a terrestrial and diurnal species, laying its eggs
in leaf-litter. The males probably take care of the eggs and then move
the larvae to pools of water in epiphytes for development.
Major Threats The entire region where the type specimen was
collected has been deforested for cattle farming since 1978.
Conservation Measures There are no protected areas near the type
locality. A search for any remaining populations is urgently needed
before the species is declared extinct.

CR Epipedobates planipaleae Morales and Velazco, 1998 OXAPAMPA POISON FROG

Critically Endangered B1ab(iii) Geographic Range This recently described species is known only Bibliography: Morales, V.R. and Velazco, P.M. (1998)
Order, Family: Anura, Dendrobatidae from the type locality of Llamaquizú stream, close to Oxapampa town, Data Providers: Javier Icochea, Edgar Lehr, Karl-Heinz Jungfer, Stefan Lötters
Country Distribution: Peru at 2,010m asl (Departamento: Pasco), Peru. It is thought to have a
Current Population Trend: Decreasing genuinely restricted distribution.
CITES: Appendix II Population It is a very rare species, and is known only from the
type series.
Habitat and Ecology It is recorded from montane tropical rainforest.
Its breeding habitat is not known.
Major Threats The major threat to this species is habitat loss due
to expanding agricultural activities.
Conservation Measures Although Parque Nacional Yanachaga-
Chemillen is very near to the type locality, populations of the species
have not been reported from this protected area; there might be a
need to extend the boundaries of this park to include the known
type locality. The species is protected by national legislation in Peru.
Further surveys are needed to determine the current population status
of the species, and captive-breeding programmes might be required
to ensure its persistence.
232 Threatened Amphibians of the World

EN Epipedobates tricolor (Boulenger, 1899)

Endangered B1ab(iii,v) Geographic Range This species is known only from seven localities on the Andean slopes of the Bolívar Province
Order, Family: Anura, Dendrobatidae in central Ecuador, where it occurs at elevations of approximately 1,000-1,769m asl.
Country Distribution: Ecuador Population It appears to have been declining in the northern portion of its range in Ecuador, where some popula-
Current Population Trend: Decreasing tions have disappeared.
CITES: Appendix II Habitat and Ecology It lives in montane forest near streams. Eggs are laid in the leaf-litter, and the males carry
the larvae to running and standing water for development. It is not known if this species can adapt to habitat
modification.
Major Threats The major threat is agrochemical pollution of waterways, and habitat loss due to smallholder farming
activities. It is also collected for medicinal use in Ecuador. Some of the declines appear to have taken place in suitable
habitat, so chytridiomycosis cannot be ruled out as a possible threat.
Conservation Measures It is not known with certainty if this species occurs in any protected areas, and expanded
protection of the remaining montane forest within the range of the species’ is recommended. The possible impact of
© Wouter Beukema

overharvesting for medicinal use, and commercial trade in wild specimens, requires further investigation.
Notes on taxonomy: This species and Epipedobates anthonyi were recently separated by Schulte (1999). We follow Graham et al.
(2004) in restricting the distribution of E. tricolor to central Ecuador.
Bibliography: Boulenger, G.A. (1899), Graham, C.H. et al. (2004), Schulte, R. (1999), Silverstone, P.A. (1976)
Data Providers: Luis A. Coloma

CR Mannophryne caquetio Mijares-Urrutia and Arends, 1999

Critically Endangered B1ab(iii,v)+2ab(iii,v) Geographic Range This species is known from the type locality, the main threat. Water extraction is a threat at the type locality. Pollution and/or parasites might be causing what
Order, Family: Anura, Dendrobatidae stated to be “toma de agua de Mapararí” in the municipality of Feder- appear to be problems in the development or incomplete development of the limbs. It is potentially at risk from
Country Distribution: Venezuela ación, in Sierra de Churuguara, in the state of Falcón, Venezuela (ap- chytridiomycosis.
Current Population Trend: Decreasing prox. 10.47N, 69.25W), where it has been recorded around 800m asl. Conservation Measures It occurs in the Parque Nacional Cueva de la Quebrada del Toro. Survey work is neces-
An additional population in Parque Nacional Cueva de la Quebrada sary to determine the population status and trends of this species. Captive breeding might be required if the risk of
del Toro was found in May 2002 (J. Manzanilla pers. comm.). chytridiomycosis proves real.
Population There has been an observed decline in the population Bibliography: Barrio Amorós, C.L. (2004), Manzanilla, J., García-París, M. and La Marca, E. (2002), Manzanilla, J., La Marca, E. and
of this species. However, in May 2002, specimens were collected García-París, M. (2003), Mijares-Urrutia, A. and Arends, A. (1999b)
at the type locality and a new population was also found in Parque Data Providers: Enrique La Marca, Jesús Manzanilla, Abraham Mijares
Nacional Cueva de la Quebrada del Toro.
Habitat and Ecology It is a terrestrial, diurnal frog found in decidu-
ous lower montane tropophilous forests. The larvae are presumably
carried on the backs of the males to streams, as with other members
of the genus.
Major Threats Habitat loss and degradation, due primarily to agri-
culture (crops and livestock), and general disturbance by humans, is

EN Mannophryne collaris (Boulenger, 1912)

Endangered B1ab(iii,v)+2ab(iii,v) Geographic Range This species is known from the Andes in the state of Mérida, in Venezuela. It occurs from
Order, Family: Anura, Dendrobatidae 200-1,800m asl.
© César Luis Barrio-Amorós / Fundación AndígenA

Country Distribution: Venezuela Population It is a locally abundant species, but appears to be in decline in some parts of its range. La Marca (1995a)
Current Population Trend: Decreasing noted population declines in this species in several localities in the Andes of Venezuela.
Habitat and Ecology It inhabits seasonal (semi-deciduous) montane forests. The eggs are laid on land and the male
protects the eggs. When hatched the male carries the larvae on his back to water where they develop further.
Major Threats The main threat to the species is habitat destruction and alteration, due primarily to agriculture (for
both crops and livestock), infrastructure development for human settlement in the vicinity of the city of Merida, and
tourism development. In addition, pollution may also be contributing to the observed declines.
Conservation Measures Its range does not include any protected areas, and expanded protection of the remaining
montane forest within the range of the species is recommended.
Bibliography: Barrio Amorós, C.L. (2001), Barrio Amorós, C.L. (2004), Barrio, C.L. and Fuentes, O. (1999), Dole, J. (1974), Durant, P. and
Dole, J. (1975), Frost, D.R. (1985), Gines, H. (1959), Gottberg, C., Ramoni, P. and Praderio, M.J. (2002), La Marca, E. (1992), La Marca, E.
(1994e), La Marca, E. (1995a), La Marca, E. (1995b), La Marca, E. (1997), Lutz, A. (1927), Manzanilla, J., García-París, M. and La Marca,
E. (2002), Manzanilla, J., La Marca, E. and García-París, M. (2003), Péfaur, J.E. and Rivero, J.A. (2000), Rivero, J.A. (1961), Vial, J.L.
and Saylor, L. (1993)
Data Providers: Enrique La Marca, Juan Elías García-Pérez, Jesús Manzanilla

VU Mannophryne cordilleriana La Marca, 1994

Vulnerable D2 Geographic Range This species is known from a few localities in (polymelia, amelia) were found at the type locality (J. Manzanilla and E. La Marca pers. comm.) where significant
Order, Family: Anura, Dendrobatidae the vicinity of Santo Domingo, in the state of Mérida, in the Venezu- pollution of the stream was also observed.
Country Distribution: Venezuela elan Andes. It has been recorded from 1,300-1,950m asl. Conservation Measures Its range does not include any protected areas. Improved habitat protection is required at
Current Population Trend: Stable Population It is a common species. sites where this species is known to occur. There is also a need for close population monitoring, particularly given
Habitat and Ecology It inhabits montane humid forest. The eggs its small range and the potential risk of chytridiomycosis.
are laid on land and the male protects the eggs. When the eggs Bibliography: Barrio Amorós, C.L. (2004), La Marca, E. (1994e), La Marca, E. (1995a), La Marca, E. (1995b), La Marca, E. (1997), Manzanilla,
are hatched, the male carries the larvae on his back to the streams J., García-París, M. and La Marca, E. (2002), Manzanilla, J., La Marca, E. and García-París, M. (2003)
where they develop further. It seems to be quite tolerant of habitat Data Providers: Enrique La Marca, Juan Elías García-Pérez, Jesús Manzanilla
disturbance, and is sometimes found in very polluted streams (but
see Major Threats).
Major Threats Its habitat is impacted by agriculture, wood col-
lection, and human settlement, although this does not seem to be
having an immediate impact on the species. It is potentially at risk
from chytridiomycosis (since the species has been proven to carry
this fungus, though it does not appear to show any symptoms of
the disease). In May 2004, some specimens with malformations
 Globally Threatened Amphibian Species 233

CR Mannophryne lamarcai Mijares-Urrutia, 1999

Critically Endangered B1ab(iii) Geographic Range This species is restricted to the Cerro Socopó in any protected areas, and its remaining habitat is in urgent need of protection. This species requires close monitoring,
Order, Family: Anura, Dendrobatidae the municipality of Mauroa, in the state of Falcón, Venezuela, at 600- particularly given the potential threat of chytridiomycosis.
Country Distribution: Venezuela 1,250m asl. It might occur more widely than is currently known. Notes on taxonomy: This frog was tentativelly assigned to Mannophryne herminae, but its taxonomic status needs further research
Current Population Trend: Decreasing Population It is a rare species that is probably declining rapidly (La Marca 1999).
because of habitat loss. It was last collected in 2004. Bibliography: Barrio Amorós, C.L. (2004), La Marca, E. (1994e), Manzanilla, J., García-París, M. and La Marca, E. (2002), Manzanilla, J.,
Habitat and Ecology It is a diurnal species that has been found La Marca, E. and García-París, M. (2003), Mijares-Urrutia, A. and Arends, A. (1999a), Mijares-Urrutia, A. and Arends, A. (1999b)
living among grasses in a small marsh along an unpaved road, and Data Providers: Abraham Mijares, Enrique La Marca, Jesús Manzanilla
in calm parts of a small stream. Males have been observed calling
from river margins. The larvae are presumably carried on the backs
of the males to streams where they develop further, as with other
members of the genus.
Major Threats The area around the site of its description was
formerly cloud forest, but the forest was cleared to establish pasture-
lands for cattle farming. Climate change, acid rain, and pollution are
also threats. It is potentially at risk from chytridiomycosis.
Conservation Measures The range of this species does not include

CR Mannophryne neblina (Test, 1956)

Critically Endangered B1ab(v)+2ab(v) Geographic Range This species is known only from the type locality mycosis cannot be ruled out as a threat.
Order, Family: Anura, Dendrobatidae and the nearby vicinity, at elevations between 900 and 1,100m asl, in Conservation Measures The range of this species includes Parque Nacional Henri Pittier. Additional surveys are
Country Distribution: Venezuela Paso Portachuelo, near Estación Biológica de Rancho Grande, Parque required to establish whether or not this species is still extant, particularly since there is still suitable habitat in its
Current Population Trend: Decreasing Nacional Henri Pittier, Aragua State, Venezuela. natural range.
Population It has not been recorded since its discovery 50 years Bibliography: Barrio Amorós, C.L. (2004), Edwards, S.R. (1974b), Gines, H. (1959), La Marca, E. (1992), La Marca, E. (1994e), La Marca,
ago, and so it might have declined and possibly even disappeared. E. (1995a), La Marca, E. and Lötters, S. (1997), Manzanilla, J. et al. (1995), Myers, C.W., Paolillo, A. and Daly, J.W. (1991), Rivero, J.A.
In the last 10 years, searches for the species have been unsuc- (1961), Test, F.H. (1956), Test, F.H., Sexton, O.J. and Heatwole, H. (1966)
cessful. This is one of the largest species of the genus, occurring Data Providers: Enrique La Marca, Jesús Manzanilla
in one of the best-studied places in Venezuela, and so it is of great
concern that no further populations or specimens of this frog have
been discovered.
Habitat and Ecology It inhabits cloud forest. Breeding habits are
unknown, although it probably lays eggs on the forest floor and
adults carry tadpoles to the stream where they develop further, like
other species of the genus.
Major Threats Threats to this species are unknown, but chytridio-

CR Mannophryne olmonae (Hardy, 1983) BLOODY BAY POISON FROG

Critically Endangered A2ae; B1ab(v) Geographic Range This species is restricted to the upland areas of north-eastern Tobago Island and Little Tobago
Order, Family: Anura, Dendrobatidae Island, Trinidad and Tobago. It is found at elevations of between 120 and 360m asl.
Country Distribution: Trinidad and Tobago Population Populations of this species have declined significantly over the last 10 years, prior to which it was
Current Population Trend: Decreasing thought to be relatively abundant.
Habitat and Ecology This is a tropical forest species. Males of this species call from rocks in streams during the
daytime. The eggs are laid on land; hatching tadpoles (11-19 in number) are carried to streams by the male where
they complete their development. It is not thought to occur in degraded habitat.
Major Threats The causes of the current decline in this species are unknown. Areas of suitable habitat remain
and there are no known human impacts on the species. It is possible that the species might have been impacted by
chytridiomycosis, though this has not been proven.
Conservation Measures The rainforest in which this species occurs in Tobago is protected in Little Tobago Wildlife
© Ryan P. Mannette

Sanctuary. Survey work is required to monitor the population status and trends of this species, and research into the
causes of the decline of this species is needed; a captive breeding programme might need to be established if the
threat of chytridiomycosis proves real.
Bibliography: Hardy, Jr, J.D. (1982), Hardy, Jr, J.D. (1983), Manzanilla, J., García-París, M. and La Marca, E. (2002), Manzanilla, J., La
Marca, E. and García-París, M. (2003), Mertens, R. (1970a), Mertens, R. (1972), Murphy, J.C. (1997)
Data Providers: Jerry Hardy

EN Mannophryne riveroi (Donoso-Barros, 1964)

Endangered B1ab(iii) Geographic Range This species is known from Cerro Azul, Macuro, and Cerro El Humo, 10km north of Macuro, in
Order, Family: Anura, Dendrobatidae Península de Paria, Sucre State, Venezuela. It has been recorded from 400-1,000m asl.
© César Luis Barrio-Amorós / Fundación AndígenA

Country Distribution: Venezuela Population The population status of this species is not known, although specimens were collected in Parque Nacional
Current Population Trend: Decreasing Peninsula de Paria as recently as May 2004.
Habitat and Ecology It lives along streams in lowland humid forest up to cloud forest habitat. A clutch of eggs
is deposited in the leaf-litter of the forest floor, and then after hatching the larvae are carried to water where they
develop further.
Major Threats The major threat is habitat loss due to selective logging and small-scale agriculture. Chytridiomycosis
represents a potential future threat.
Conservation Measures Although most of the range of this species is within a protected area (Parque Nacional
Peninsula de Paria), this reserve is under pressure of deforestation (by the local people). Strengthened management
and protection of this site is necessary to safeguard this species.
Bibliography: Ayarzaguena, J. and Señaris, J.C. (1993), Barrio Amorós, C.L. (2004), Donoso-Barros, R. (1964), Edwards, S.R. (1974b),
Frost, D.R. (1985), La Marca, E. (1992), La Marca, E. (1994e), La Marca, E. (1995a), La Marca, E. (1995b), Manzanilla, J., García-París,
M. and La Marca, E. (2002), Manzanilla, J., La Marca, E. and García-París, M. (2003), Myers, C.W., Paolillo, A. and Daly, J.W. (1991),
Rivero, J.A. (1988)
Data Providers: Enrique La Marca, Jesús Manzanilla
234 Threatened Amphibians of the World

VU Mannophryne trinitatis (Garman, 1887) YELLOW-THROATED FROG

Vulnerable B1ab(iii) Population In Venezuela, it is considered to be common, while on Trinidad it is considered to be very common with
Order, Family: Anura, Dendrobatidae generally stable populations.
Country Distribution: Trinidad and Tobago, Habitat and Ecology Adults are found along densely shaded, narrow, shallow, slow-flowing, clear water streams
Venezuela in undisturbed montane and moist forests, where they engage in elaborate courtship and defensive behaviours. A
Current Population Trend: Decreasing terrestrial species, it lays its eggs in leaf-litter and on rocks near streams and the adults carry the tadpoles on their
backs to the deeper stream pools. The populations in Tamana caves in central Trinidad are known to use the cave
ponds for their tadpoles. Tadpoles in other populations have also been found in still temporary pools far from streams.
The exact deposition site appears to be dependent on the presence of predators, with adults migrating significant
distances in search of predator free pools. These animals depend on the insects that eat bat guano for food.
Major Threats In Venezuela, the species is threatened by habitat degradation and destruction resulting from
small-scale agriculture. On Trinidad, it is locally threatened by domestic, agricultural, and some industrial water
© Karl-Heinz Jungfer

Geographic Range This species is present in the Northern and Central Ranges of the island of Trinidad, in Trinidad
and Tobago. In Venezuela, populations are present in Península de Paria (Sucre state), Serranía del Turimiquire (Sucre-
Monagas states), and Serranía del interior, Miranda, Guarico, and Anzoatigui states. In Venezuela the species occurs
between 40 and 1,525m asl, and on Trinidad it can be found from montane areas down to sea level.
pollution. On the southern slopes of the Northern Range in Trinidad, deforestation has resulted in fragmentation of
the species’ habitat.
Conservation Measures A number of protected areas are present within the range of this species. In Venezuela, it
is known to occur in the Parque Nacional Peninsula de Paria and Parque Nacional el Guacharo.
Notes on taxonomy: This form is probably a complex of more than one species. The populations in Venezuela almost certainly belong
to a different, undescribed, species (J. Manzanilla pers. comm.).
Bibliography: Alemán, C. (1952), Barrio Amorós, C.L. (2004), Edwards, S.R. (1974b), Frost, D.R. (1985), Jowers, M.J. and Downie, J.R.
(2004), Kenny, J.S. (1969), La Marca, E. (1992), La Marca, E. (1994e), La Marca, E. (1995a), La Marca, E. (1997), Manzanilla, J., García-
París, M. and La Marca, E. (2002), Manzanilla, J., La Marca, E. and García-París, M. (2003), Mole, R.R. and Urich, F.W. (1894), Murphy,
J.C. (1997), Parker, H.W. (1933a)
Data Providers: Enrique La Marca, Jesús Manzanilla, Jerry Hardy

EN Mannophryne yustizi (La Marca, 1989)

Endangered B1ab(iii) Geographic Range This species is known mainly from Yacambú and Terepaima National Parks in Lara State,
Order, Family: Anura, Dendrobatidae Venezuela. It has been recorded from 1,200-1,800m asl.
© César Luis Barrio-Amorós / Fundación AndígenA

Country Distribution: Venezuela Population It is a common species and there is currently no evidence of any decline. Healthy populations were found
Current Population Trend: Decreasing in April 2003 in Parque Nacional Yacambú.
Habitat and Ecology It occurs along streams and on the forest floor of montane moist and cloud forests. The eggs
are laid on land and the male protects the eggs. When they hatch, the male carries the larvae on his back to water
where they develop further.
Major Threats In some parts of the range, populations are affected by the extensive clearing of land for coffee
plantations (and specifically by the agro-chemicals associated with the plantations).
Conservation Measures Although it is protected in two Venezuelan Andean national parks (Yacambú and Terepaima
National Parks), some populations (in Cubiro and Guarico) live outside protected areas. Expanded protection of the
montane forest within the species’ range is recommended.
Bibliography: Barrio Amorós, C.L. (2004), La Marca, E. (1989), La Marca, E. (1992), La Marca, E. (1994e), La Marca, E. (1995a), La Marca,
E. (1995b), La Marca, E. (1997), Manzanilla, J., García-París, M. and La Marca, E. (2002), Manzanilla, J., La Marca, E. and García-París,
M. (2003), Myers, C.W., Paolillo, A. and Daly, J.W. (1991), Yustiz, E. (1996)
Data Providers: Enrique La Marca, Juan Elías García-Pérez, Jesús Manzanilla

EN Nephelobates alboguttatus (Boulenger, 1903)

Endangered B1ab(iii,v)+2ab(iii,v) Habitat and Ecology It occurs in clear, fast-flowing streams in Andean cloud forests. The eggs are laid on land
Order, Family: Anura, Dendrobatidae and the male protects the eggs. When they are hatched, the male carries the larvae on his back to water where
© César Luis Barrio-Amorós / Fundación AndígenA

Country Distribution: Venezuela they develop further.

Current Population Trend: Decreasing Major Threats Some of the original environments occupied by this frog in the vicinities of the city of Mérida, in the
Cordillera de Mérida, have been drastically changed by human activities, particularly due to agricultural expansion,
for both crops and livestock. Much of the remaining habitat is severely fragmented. Introduced trout prey on the
larvae of this species. However, this species is also declining in undisturbed habitats, which suggests an additional
threat, possibly disease (such as chytridiomycosis).
Conservation Measures Some populations are protected within national parks (Sierra Nevada and Sierra de
la Culata). However, the existing protected areas network requires expansion to ensure the protection of suitable
cloud forest habitats for this species. Further research is required to establish the reasons for the species’ decline in
apparently suitable habitat, and to determine whether chytrid poses a threat.
Notes on taxonomy: Colostethus inflexus was synonymized with this species by Frost (1985), but was revived in Myers, Paolillo and
Daly (1991) and La Marca (1997). Although officially still in the synonymy of Nephelobates alboguttatus, C. inflexus might prove to be
a distinct species.
Bibliography: Barrio Amorós, C.L. (2004), Frost, D.R. (1985), La Marca, E. (1984), La Marca, E. (1985b), La Marca, E. (1992), La Marca, E.
(1994a), La Marca, E. (1995b), La Marca, E. (1997), Lutz, A. (1927), Myers, C.W., Paolillo, A. and Daly, J.W. (1991), Péfaur, J.E. and Rivero,
Geographic Range This species is known from the Andes of Mérida State, Venezuela, from 1,600-3,090m asl. J.A. (2000), Piñero, J. and Durant, P. (1993), Piñero, J. and La Marca, E. (1996), Rivero, J.A. (1961), Vial, J.L. and Saylor, L. (1993)
Population It is a rare species and has been in decline in recent years. Data Providers: Enrique La Marca, Juan Elías García-Pérez

EN Nephelobates duranti (Pefaur, 1985)

Endangered B1ab(iii)+2ab(iii) Geographic Range This species is only known from the type locality Bibliography: Barrio Amorós, C.L. (2004), Diaz, A., Péfaur, J. and Durant, P. (1997), Frost, D.R. (1985), La Marca, E. (1992), La Marca, E.
Order, Family: Anura, Dendrobatidae and the vicinity: Páramo de la Culata, in the district of Libertador, in (1994a), La Marca, E. (1995b), La Marca, E. (1997), Mijares-Urrutia, A. and La Marca, E. (1997b), Péfaur, J.E. (1985), Rivero, J.A. (1988),
Country Distribution: Venezuela the state of Mérida, Venezuela, at 2,600-3,000m asl. Vial, J.L. and Saylor, L. (1993)
Current Population Trend: Decreasing Population It is an uncommon species. Data Providers: Enrique La Marca, Juan Elías García-Pérez
Habitat and Ecology It inhabits clear, fast-flowing streams in An-
dean cloud forests and sub-páramo shrubland. The eggs are laid on
land and the male protects the eggs. When they are hatched, the male
carries the larvae on his back to water where they develop further.
Major Threats Livestock farming is a major threat to the species’
habitat, which is now severely fragmented. Introduced trout also
prey on the larvae of this species.
Conservation Measures A population probably occurs within
Parque Nacional Sierra de La Culata. Continued and strengthened
management of this area, and expansion of the existing protected
areas network to include other tracts of montane forest habitat,
are necessary.
 Globally Threatened Amphibian Species 235

EN Nephelobates haydeeae (Rivero, 1976)

Endangered B1ab(iii,v)+2ab(iii,v) Geographic Range This species is known from the Venezuelan areas network to include other tracts of montane forest habitat in the Venezuelan Andes, are necessary.
Order, Family: Anura, Dendrobatidae Andean states of western Mérida and eastern Táchira. It has been Bibliography: Barrio Amorós, C.L. (2004), Frost, D.R. (1985), La Marca, E. (1992), La Marca, E. (1994a), La Marca, E. (1995b), La Marca, E.
Country Distribution: Venezuela recorded between 1,825 and 2,670m asl. (1997), Mijares-Urrutia, A. and La Marca, E. (1997b), Myers, C.W., Paolillo, A. and Daly, J.W. (1991), Péfaur, J.E. and Rivero, J.A. (2000),
Current Population Trend: Decreasing Population It is an uncommon species and is in decline. Rivero, J.A. (1976), Rivero, J.A. (1988), Vial, J.L. and Saylor, L. (1993)
Habitat and Ecology It occurs in clear, fast-flowing streams in Data Providers: Enrique La Marca, Juan Elías García-Pérez
Andean cloud forests, and in the puddles alongside the streams,
and is sometimes hidden inside introduced “kikuyo” pasture grasses
(Pennisetum clandestinum). The eggs are laid on land and the male
protects the eggs. When they are hatched, the male carries the larvae
on his back to water where they develop further.
Major Threats Most of the species’ original habitat has now been
lost to agricultural activities (involving livestock and crops), and the
remaining habitat is severely fragmented.
Conservation Measures The species’ range may be partially
within Parque Nacional Los Paramos. Continued and strengthened
management of this area, and expansion of the existing protected

EN Nephelobates mayorgai (Rivero, 1978)

Endangered B1ab(iii)+2ab(iii) Geographic Range This species is known from several localities within Mérida State, on the road from Mérida to
Order, Family: Anura, Dendrobatidae La Azulita, in Venezuela, at elevations of 1,700-2,400m asl.
© César Luis Barrio-Amorós / Fundación AndígenA

Country Distribution: Venezuela Population It is a locally common species.

Current Population Trend: Decreasing Habitat and Ecology It is found along mountain streams in cloud forests. The eggs are laid on land and the male
protects the eggs. When they are hatched, the male carries the larvae on his back to water where they develop
further.
Major Threats The main threats are agriculture, involving both crops and livestock, as well as agricultural water
pollution. The introduced bullfrog (Rana catesbeiana) also poses a threat.
Conservation Measures The range of this species does not include any protected areas; protection of remaining
tracts of forest habitat in the Venezuelan Andes is necessary.
Bibliography: Barrio Amorós, C.L. (2004), La Marca, E. (1992), La Marca, E. (1994a), La Marca, E. (1995b), La Marca, E. (1997), Mijares-
Urrutia, A. and La Marca, E. (1997a), Myers, C.W., Paolillo, A. and Daly, J.W. (1991), Péfaur, J.E. and Rivero, J.A. (2000), Rivero, J.A.
(1978), Rivero, J.A. (1988), Vial, J.L. and Saylor, L. (1993)
Data Providers: Enrique La Marca, Juan Elías García-Pérez

CR Nephelobates meridensis (Dole and Durant, 1972)

Critically Endangered B2ab(iii) Geographic Range This species is known from the type locality, Chorotal, 15km south-east of la Azulita, and the
Order, Family: Anura, Dendrobatidae nearby vicinity, in Mérida State, Venezuela, between 1,880 and 2,400m asl.
© César Luis Barrio-Amorós / Fundación AndígenA

Country Distribution: Venezuela Population It is an uncommon species.

Current Population Trend: Decreasing Habitat and Ecology It inhabits streams in cloud forest. The eggs are laid on land and the male protects the eggs;
when they hatch, the male carries the larvae on his back to the water where they develop further.
Major Threats The primary threats to this species are habitat loss due to agriculture (crops and livestock) and
agricultural pollution. The invasive bullfrog Rana catesbeiana is also a threat.
Conservation Measures The range of this species does not include any protected areas, and there is an urgent
need to ensure that the remaining habitat at the type locality is maintained.
Bibliography: Barrio Amorós, C.L. (2004), Dole, J. and Durant, P. (1972), Frost, D.R. (1985), La Marca, E. (1992), La Marca, E. (1994a),
La Marca, E. (1995b), La Marca, E. (1997), Mijares-Urrutia, A. and La Marca, E. (1997a), Myers, C.W., Paolillo, A. and Daly, J.W. (1991),
Péfaur, J.E. and Rivero, J.A. (2000), Rivero, J.A. (1988), Vial, J.L. and Saylor, L. (1993)
Data Providers: Enrique La Marca, Juan Elías García-Pérez

EN Nephelobates molinarii (La Marca, 1985)

Endangered B1ab(ii,iii)+2ab(ii,iii) Geographic Range This species is only known from a single zone near Bailadores, in Mérida State, Venezuela, at
Order, Family: Anura, Dendrobatidae an altitude of 1,800-2,600m asl.
© César Luis Barrio-Amorós / Fundación AndígenA

Country Distribution: Venezuela Population It is an uncommon species.

Current Population Trend: Decreasing Habitat and Ecology It inhabits dry evergreen montane forest, where it lives along slow-flowing streams. The eggs
are laid on land and the male protects the eggs. When they are hatched, the male carries the larvae on his back to
water where they develop further.
Major Threats Habitat destruction and degradation, caused primarily by intensive agriculture, has reduced the area
of occupancy of this species to less than 10km².
Conservation Measures Its range does not include any protected areas, although it is possible that it might
occur in Parque Nacional Paramos El Batallon y La Negra. Continued and strengthened management of this area,
and expansion of the existing protected areas network to include other montane forest habitat in the Venezuelan
Andes, are necessary.
Bibliography: Barrio Amorós, C.L. (2004), La Marca, E. (1985a), La Marca, E. (1985b), La Marca, E. (1992), La Marca, E. (1994a), La Marca,
E. (1997), Myers, C.W., Paolillo, A. and Daly, J.W. (1991), Péfaur, J.E. and Rivero, J.A. (2000), Rivero, J.A. (1988)
Data Providers: Enrique La Marca, Juan Elías García-Pérez
236
EN Nephelobates orostoma (Rivero, 1976)
Endangered B1ab(ii,iii)+2ab(ii,iii)
Order, Family: Anura, Dendrobatidae
Country Distribution: Venezuela
Current Population Trend: Decreasing
EN Nephelobates serranus (Pefaur, 1985)
Endangered B1ab(ii,iii)+2ab(ii,iii)
Order, Family: Anura, Dendrobatidae
Country Distribution: Venezuela
Current Population Trend: Decreasing
EN Phyllobates terribilis Myers, Daly and Malkin, 1978
Endangered B1ab(iii)
Order, Family: Anura, Dendrobatidae
Country Distribution: Colombia
Current Population Trend: Decreasing
CITES: Appendix II
© Jörn Köhler
EN Phyllobates vittatus (Cope, 1893)
Endangered B1ab(iii)
Order, Family: Anura, Dendrobatidae
Country Distribution: Costa Rica
Current Population Trend: Stable
CITES: Appendix II
© Tobias Eisenberg
Threatened Amphibians of the World
Geographic Range This species is known with certainty only from
the type locality in Táchira State, Venezuela (Boca del Monte, Camino
del Pregonero), from 2,300-2,800m asl. Other Táchira localities listed
in Barrio (1999) are unconfirmed.
Population It is an uncommon species.
Habitat and Ecology It inhabits mountain streams in cloud forest.
The eggs are laid on land and the male protects the eggs. When
they are hatched, the male carries the larvae on his back to water
where they develop further.
Major Threats The major threats are agriculture, involving both
crops and livestock, and agricultural pollution, and the area of oc-
cupancy of this species is now less than 20km².
Conservation Measures It is not known to occur in any protected
areas, and protection of remaining tracts of forest habitat in the
Venezuelan Andes is necessary.
Bibliography: Barrio Amorós, C.L. (2004), Frost, D.R. (1985), La Marca, E.
Geographic Range This species is only known from the type local-
ity, Vía El Morro, in the district of Libertador, in the state of Mérida,
Venezuela, from 1,800-2,300m asl.
Population It is an uncommon species.
Habitat and Ecology It inhabits mountain streams in cloud forest.
The eggs are laid on land and the male protects the eggs. When
they are hatched, the male carries the larvae on his back to water
where they develop further.
Major Threats The major threats are agriculture, involving both
crops and livestock, and predation on larvae by invasive trout; the
area of occupancy of this species is now less than 20km².
Conservation Measures A population may be protected within
Parque Nacional Sierra Nevada. Continued and strengthened man-
agement of this area, and expansion of the existing protected areas
network to include other montane forest habitat in the Venezuelan
Andes, are necessary.
(1992), La Marca, E. (1994a), La Marca, E. (1995b), La Marca, E. (1997), Mijares-Urrutia, A. and La Marca, E. (1997a), Péfaur, J.E. (1985),
Rivero, J.A. (1976), Rivero, J.A. (1988)
Data Providers: Enrique La Marca, Juan Elías García-Pérez
Bibliography: Barrio Amorós, C.L. (2004), La Marca, E. (1992), La Marca, E. (1994a), La Marca, E. (1995b), La Marca, E. (1997), Mijares-
Urrutia, A. and La Marca, E. (1997a), Myers, C.W., Paolillo, A. and Daly, J.W. (1991), Péfaur, J.E. (1985), Rivero, J.A. (1988)
Data Providers: Enrique La Marca, Juan Elías García-Pérez
GOLDEN POISON FROG
Geographic Range This species is known only from tiny areas on the Pacific coast of Colombia on the Río Saija
drainage, in Cauca Department, occurring up to 200m asl.
Population It is extremely common in its tiny range.
Habitat and Ecology It lives on the ground in humid forests, and is only known from primary forest. It is not known
whether or not it can adapt to secondary habitats. The eggs are laid on the ground and the males transport the
larvae to permanent pools.
Major Threats The major threats are deforestation for agricultural development, the planting of illegal crops, logging,
and human settlement, and pollution resulting from the spraying of illegal crops. It is very occasionally reported in
international trade in small numbers.
Conservation Measures It does not occur in any protected areas, and the protection of part of this species’ lowland
forest habitat is recommended. Management practices that could allow a commercial, sustainable harvest of this
species should be investigated. Decree INDERENA No. 39 of 9 July, 1985, forbids the collection of Phyllobates spp.
from the wild in Colombia for breeding (or other) purposes.
Bibliography: Lötters, S. et al. (1997), Myers, C.W., Daly, J.W. and Malkin, B. (1978), Rueda-Almonacid, J.V. (1999), Ruiz-Carranza,
P.M., Ardila-Robayo, M.C. and Lynch, J.D. (1996)
Data Providers: Wilmar Bolívar, Stefan Lötters
GOLFODULCEAN POISON FROG
Geographic Range This species is known from the lowlands of the Golfo Dulce region of south-western Costa Rica,
from 20-550m asl (Savage 2002), and has also recently been recorded from Dominical in the Provincia de Puntarenas
of Costa Rica (Ryan 2002). It is expected to occur in parts of immediately adjacent south-western Panama.
Population This species is moderately common and regularly recorded (although extremely rare in Dominical).
Habitat and Ecology This is a diurnal, terrestrial species associated with streams in primary lowland moist and
wet forests. Eggs are usually deposited on leaves above the ground; the male carries hatching larvae to small pools
to complete their development (Savage 2002).
Major Threats This species is threatened by forest clearance for agricultural land and tree plantations. Water
pollution caused by contamination from gold mining activities is also a threat, as is potential over-collection of
adults for the pet trade.
Conservation Measures Most of the species’ range is in three protected areas in Costa Rica, including Parque
Nacional Corcavado. There is a need for strengthened management of these sites, and expanded protection to include
other remnant forest patches in Costa Rica. Management practices that could allow a commercial, sustainable harvest
of this species for the pet trade should be investigated.
Bibliography: Ryan, M. (2002), Savage, J.M. (1968b), Savage, J.M. (1976), Savage, J.M. (2002), Silverstone, P.A. (1976), Weygoldt,
P. (1987), Zimmermann, H. (1982)
Data Providers: Frank Solís, Roberto Ibáñez, Gerardo Chaves, Jay Savage, César Jaramillo, Querube Fuenmayor, Federico Bolaños
 Globally Threatened Amphibian Species 237

DISCOGLOSSIDAE

VU Alytes dickhilleni Arntzen and García-París, 1995 BETIC MIDWIFE TOAD

Vulnerable B2ab(iii,iv) Population Populations of this species are very fragmented, many of them confined to isolated mountains and valleys.
Order, Family: Anura, Discoglossidae It is relatively common in the Alcaraz, Segura, and Cazorla mountains, but it is rare in drier mountains (Filabres, Baza,
Country Distribution: Spain Gádor), where it is associated with springs. Populations in drier areas can consist of only a few adults.
Current Population Trend: Decreasing Habitat and Ecology The species is present in pine and oak forests, most often on calciferous substrate, in open,
very rocky landscapes. Adults occur in rock fissures and on stones next to water sources. Reproduction and larval
development takes place in permanent mountain streams, man-made reservoirs and cattle troughs, and the larvae
may take a long time to mature. Almost all known breeding habitats are human-modified water bodies.
Major Threats The species is threatened by loss of suitable breeding habitat as a result of excessive water
withdrawal, droughts, and modernization of agricultural practices leading to the abandonment of cattle troughs and
other man-made water sources. A potential future threat is the fungal disease chytridiomycosis, which has already
© Jeroen Speybroeck

impacted the related Alytes obstetricans in Spain.

Geographic Range This species is restricted to the mountains of south-eastern Spain. It occurs at altitudes of
700-2,140m asl (Sierra Nevada, Almería).
Conservation Measures This species is listed on Appendix III of the Berne Convention. It is listed in regional Red
Data Books and is present in the protected areas of Parque Nacional Sierra Morena, Parque Nacional de Sierra
Nevada, and the Natural Park of Cazorla, Segura y las Villas. Protection measures in Castilla-La Mancha, Andalusia,
such as restoration and construction of new breeding habitats, are under way.
Bibliography: Arnold, E.N. (2003), Arntzen, J.W. and García-París, M. (1995), Benavides, J. et al. (2000), Fromhage, L., Vences, M. and
Veith, M. (2004), García-Cardenete, L. et al. (2003), Gasc, J.-P. et al. (eds.) (1997), Márquez, R. and Bosch, J. (1996), Martínez-Solano, I.
et al. (2003), Martínez-Solano, I. et al. (2004), Pleguezuelos, J.M. (1997), Pleguezuelos, J.M., Márquez, R. and Lizana, M. (2002)
Data Providers: Jaime Bosch, Miguel Tejedo, Miguel Lizana,Iñigo Martínez-Solano, Alfredo Salvador, Mario García-París, Ernesto
Recuero Gil, Jan Willem Arntzen, Rafael Marquez, Carmen Diaz Paniagua

VU Alytes muletensis (Sanchiz and Adrover, 1979 “1977”) MALLORCAN MIDWIFE TOAD
Vulnerable D2 Habitat and Ecology It is currently found only in small streams deeply carved into limestone mountains. The pres-
Order, Family: Anura, Discoglossidae ence of the species is positively associated with steep slopes. Breeding takes place in the small streams that persist
Country Distribution: Spain (Native and as pools in summer. A few populations occur by man-made water sources (cattle troughs, containers, rain tanks
Reintroduced) etc.) in open mountainous country; these are within the river basins of nearby canyon-living populations. Animals
Current Population Trend: Increasing are generally found in rock crevices and under stones. This species does not tolerate serious habitat degradation.
The distribution of predators on the species is negatively associated with elevation, and reproductive success is
positively associated with elevation.
Major Threats The major threats are predation by the introduced Viperine Snake (Natrix maura), and competition
for space with Perez’s Frog (Rana perezi). Development of tourism and human settlements, specifically the increased
need for water resources (including damming and canalization of streams), is an additional threat. The threats are
not likely to decrease, and so the current recovery programme needs to be continued more or less indefinitely. One
© Jeroen Speybroeck

Geographic Range This species is restricted to the Sierra Tramuntana of northern Mallorca, Balearic Islands, Spain.
The present altitudinal range is from 10-850m asl. Its area of occupancy is less than 10km², but slowly increasing as
a result of intensive conservation action.
Population The population is approximately 500-1,500 adult pairs (Gasc et al. 1997; Arnold 2003). There are ap-
proximately 25, mostly isolated, populations. The total population is slowly increasing following coordinated recovery
efforts, following a long period of decline and near extinction. The current increase, which probably started around
the time that the first re-introductions were made in 1989, has been maintained even during years of drought,
notably in 1999-2000. The population trend in this species is monitored through annual tadpole counts, the counts
for 2004 (over 30,000 tadpoles) being the highest on record. The increase in numbers in established populations is
not dependent upon continued re-introductions. However, it is unlikely that new populations would be established
without re-introductions.
isolated re-introduced population was impacted by an unidentified non-fungal disease in 2002 which killed some
tadpoles. This disease did not recur in 2003 and 2004.
Conservation Measures The species is protected by sub-national and national legislation. It is listed on Appendix II of
the Berne Convention, on Annexes II and IV of the EU Natural Habitats Directive, and on the national and sub-national
Red Data Books. It is present in the protected areas of the Tramuntana mountains. The Balearic Government and
Jersey Wildlife Preservation Trust have undertaken captive-breeding, re-introduction and other conservation initiatives.
At least 10 populations have been successfully reintroduced. Re-introductions of animals from the Jersey Wildlife
Preservation Trust stopped in 2002, but a new captive-breeding facility now exists on Mallorca, and re-introductions
are expected to resume. However, as a result of the recent discovery of disease, a recommendation was made in 2004
to the Balearic Government to halt the re-introduction programme. A new recovery programme for the species is now
being developed. A systematic programme is in place to remove Natrix maura from the range of the species.
Bibliography: Alcover, J.A. et al. (1984), Arnold, E.N. (2003), Arntzen, J.W. and García-París, M. (1995), Bush, S. (1996), Fromhage, L.,
Vences, M. and Veith, M. (2004), Gasc, J.-P. et al. (eds.) (1997), Halliday, T. (1992), Lea, J., Dyson, M. and Halliday, T. (2002), Martínez-
Solano, I. et al. (2004), Mejias, R. and Amengual, J. (2000), Moore, R.D., Griffiths, R.A. and Román, A. (2004), Pleguezuelos, J.M. (1997),
Pleguezuelos, J.M., Márquez, R. and Lizana, M. (2002), Roca, V. et al. (1998), Román, A. and Mayol, J. (1997), Schley, L., Griffiths, R.A.
and Román, A. (1998)
Data Providers: Joan Mayol Serra, Richard Griffiths, Jaime Bosch, Trevor Beebee, Benedikt Schmidt, Miguel Tejedo, Miguel Lizana,
Iñigo Martínez-Solano, Alfredo Salvador, Mario García-París, Ernesto Recuero Gil, Jan Willem Arntzen

HELEOPHRYNIDAE

CR Heleophryne hewitti Boycott, 1988 HEWITT’S GHOST FROG

Critically Endangered B2ab(ii,iii,iv,v) Geographic Range This species appears to be restricted to four perennial rivers (Geelhoutboom, Martin’s, Klein
Order, Family: Anura, Heleophrynidae and Diepkloof) with their headwaters in the Elandsberg mountains, in the Eastern Cape province, South Africa. Its
Country Distribution: South Africa altitudinal range is 400-550m asl.
Current Population Trend: Decreasing Population It is an uncommon and declining species.
Habitat and Ecology It is a species of fynbos heathland and grassy fynbos. Only very small remnants of fynbos
survive within its range, so very little non-breeding habitat survives. It breeds in fast-flowing perennial rivers and
streams with rocky beds in the upper reaches of the Elandsberg. Adults and tadpoles are found beneath submerged
and partly submerged rocks in these streams, and occasionally at the edge of small waterfalls and cascades. The
tadpoles take two years to develop.
Major Threats The main threats are loss of suitable non-breeding and breeding habitat as a result of afforestation
with exotic pine plantations, fires, erosion, siltation of streams, dams, and road building. Introduced predatory fish
© Richard Boycott

are probably also a threat.

Conservation Measures The species is not known to occur in any protected areas, and the maintenance of its
remaining breeding and non-breeding habitat is essential. There is also a need for continued monitoring of known
populations and survey work for other populations.
Bibliography: Boycott, R.C. (1988), Branch, W.R. (1988), Channing, A. (2001), Minter, L.R. et al. (2004), Passmore, N.I. and Carruthers,
V.C. (1995)
Data Providers: Leslie Minter, Alan Channing, James Harrison
238 Threatened Amphibians of the World

CR Heleophryne rosei Hewitt, 1925 TABLE MOUNTAIN GHOST FROG

Critically Endangered B1ab(ii,iii,v)+2ab(ii,iii,v) Geographic Range This species is endemic to the southern, eastern, and marginally western (and historically
Order, Family: Anura, Heleophrynidae northern), slopes of Table Mountain, in the Western Cape province, extreme south-western South Africa. It occurs
Country Distribution: South Africa at 240-1,060m asl. Detailed information on the localities from which it has been recorded on Table Mountain are
Current Population Trend: Decreasing provided by Minter et al. (2004).
Population It is a rare and elusive species that appears to be declining and survives in low population densities;
for example, it is estimated that the number of tadpoles in the Skeleton Gorge has decreased by 50% since 1980
(Minter et al. 2004).
Habitat and Ecology It lives in forest and fynbos heathland, breeding in clear perennial streams in forested gorges,
valleys and ravines on Table Mountain. Non-breeding adults have been found in damp, sheltered habitat well away
© Atherton de Villiers from streams, including in caves. The tadpoles require longer than 12 months to complete metamorphosis, and so it
is important that there is perennial water to allow the larvae to develop.
Major Threats The main threats are the spread of alien vegetation, frequent fires, and construction of water stor-
age reservoirs on the mountain affecting the consistency of stream-flow. Intensive eco-tourism is also a potential
threat.
Conservation Measures The whole of this species’ range is incorporated in the Table Mountain National Park,
part of the Cape Floristic Region World Heritage Site. A monitoring programme by Western Cape Nature Conserva-
tion is in place.
Bibliography: Baard, E.H.W. (1989), Boycott, R.C. and de Villiers, A.L. (1986), Branch, W.R. (1988), Channing, A. (2001), De Villiers, A.L.
(1997), McLachlan, A. (1978), Minter, L.R. et al. (2004), Passmore, N.I. and Carruthers, V.C. (1995)
Data Providers: Leslie Minter, Alan Channing, James Harrison

HEMISOTIDAE

VU Hemisus guttatus (Rapp, 1842) SPOTTED SNOUT-BURROWER

Vulnerable B2ab(ii,iii,iv) Geographic Range This species, which is known only from South Africa, occurs in southern Mpumalanga, and
Order, Family: Anura, Hemisotidae central and eastern KwaZulu-Natal, south to Durban on the coast. The northernmost coastal record is from Hluhluwe.
Country Distribution: South Africa It ranges from sea level up to over 1,000m on the summit of the Lebombo Mountains. It has not been recorded from
Current Population Trend: Decreasing Swaziland, but it presumably occurs in this country.
Population It is a rare species.
Habitat and Ecology It inhabits grassland and savannah. It breeds in seasonal pans, swampy areas, and in pools
near rivers. It nests in burrows in wet soil by temporary water, and tadpoles move to water to develop.
Major Threats The main threats include: habitat loss due to afforestation, sugar cane cultivation, and urbanization;
alien fish introductions; and invasive alien plants lowering the water table.
Conservation Measures It occurs in the Great St Lucia Wetlands Park, the Hluhluwe-Umfolozi Game Park, and
© Atherton de Villiers

other protected areas.

Bibliography: Alexander, G.J. (1990), Channing, A. (2001), Lambiris, A.J.L. (1989a), Laurent, R.F. (1972), Minter, L.R. et al. (2004),
Passmore, N.I. and Carruthers, V.C. (1995), Wager, V.A. (1986)
Data Providers: Leslie Minter, Alan Channing, James Harrison

HYLIDAE

EN Agalychnis annae Duellman, 1963 BLUE-SIDED TREEFROG

Endangered A2abe Population Since the late 1980s, it has disappeared from pristine areas, including in protected areas such as Parque
Order, Family: Anura, Hylidae Nacional Tapantí and the Reserva Biológica Monteverde, where it was once common. It remains common only in
Country Distribution: Costa Rica highly altered habitats in metropolitan San José.
Current Population Trend: Decreasing Habitat and Ecology This is a nocturnal species that lives in premontane moist and wet forests and rainforest,
and tolerates disturbance to its habitat. It remains the most abundant species in San José and suburbs near heavily
polluted streams, especially in shade-grown coffee plantations and gardens. It breeds in streams.
Major Threats Despite the apparent adaptability of this species, it is nonetheless subject to unconfirmed factors
that have caused amphibian faunas to decline in certain locations in Central America, in particular the fungal disease,
chytridiomycosis. It is possible that this species survives only in polluted areas because the chytrid fungus is more
susceptible to pollution than the frog. The few, known remaining populations of this species are threatened by an intro-
duced fish (Xiphophorus hellerii) that preys on the larvae. This species is also found in the international pet trade.
© Twan Leenders

Geographic Range This species occurs in the northern Cordillera de Talamanca, Cordillera de Tilarán and Cordil-
lera Central, Costa Rica, at 780-1,650m. However, it has disappeared from most parts of its range, surviving mainly
around San José only.
Conservation Measures Research is needed to determine whether or not this species can survive only in polluted
areas, because of the ineffectiveness of the chytrid fungus in such environments. If this proves to be the case,
then well-meaning conservation measures to abate water pollution could unintentionally lead to the extinction of
this species. Given the nature of the threats, it may be worth considering the establishment of a captive-breeding
programme for this species.
Bibliography: Duellman, W.E. (2001), Pounds, J.A. et al. (1997), Proy, C. (1993), Proy, C. (2000), Savage, J.M. (2002), Villa, J. and
Townsend, D.S. (1983)
Data Providers: Alan Pounds, Federico Bolaños, Gerardo Chaves
 Globally Threatened Amphibian Species 239

VU Agalychnis litodryas (Duellman and Trueb, 1967) PINK-SIDED TREEFROG

Vulnerable B1ab(iii) Geographic Range This species is known from the single locality of Río Tuira at Río Mono in Darién Province,
Order, Family: Anura, Hylidae Panama. It has been recorded from five localities in Ecuador and is presumed to occur in Colombia, although it has
Country Distribution: Ecuador, Panama not yet been recorded. It ranges from 100-1,000m asl.
Current Population Trend: Decreasing Population The species is known only from a single record in Panama. It is considered to be rare in Ecuador.
Habitat and Ecology An arboreal species of humid lowland forest. The single specimen in Panama was collected
on a bush in a swamp. The holotype was found at night perched about one and a half meters above the ground in
a bush at the edge of a swamp (Duellman and Trueb 1967). A gliding species, the Ecuadorian specimens have been
heard calling from bushes overhanging a shallow pond in a banana grove at night (Duellman 2001). It deposits its
eggs on the leaves floating on ponds, and larvae jump into the pond and develop in the water. Since it depends on
© Karl-Heinz Jungfer the forest canopy, it is not very adaptable to disturbance.
Major Threats Habitat loss and degradation is a major threat in Ecuador, due to agriculture, timber, and human
settlement.
Conservation Measures The species occurs in Parque Nacional Darién in Panama, and Reserva Ecológica
Cotacachi-Cayapas in Ecuador. Further survey work is needed to determine whether or not this species occurs in
Colombia, as is predicted.
Bibliography: Duellman, W.E. (2001), Duellman, W.E. and Trueb, L. (1967), Ibáñez, R. et al. (2000), Young, B. et al. (1999)
Data Providers: Luis A. Coloma, Santiago Ron, Karl-Heinz Jungfer, Diego Cisneros-Heredia, Ana Almandáriz, Frank Solís, Roberto
Ibáñez, César Jaramillo, Querube Fuenmayor

CR Agalychnis moreletii (Duméril, 1853) MORELET’S TREEFROG

Critically Endangered A3e Geographic Range This species occurs from north-eastern Puebla state and south-central Veracruz state, Mexico,
Order, Family: Anura, Hylidae to north-western Honduras on the Atlantic versant; and from south-central Guerrero state, Mexico, to central El
Country Distribution: Belize, El Salvador, Salvador on the Pacific versant, at elevations of 300-1,500m asl.
Guatemala, Honduras, Mexico Population It was formerly locally abundant in some locations in Chiapas state, Mexico, El Salvador and Guatemala.
Current Population Trend: Decreasing However, recent surveys in Guerrero, Oaxaca and Chiapas, Mexico, indicate that it has disappeared from all the sites
surveyed. It is uncommon, but occasionally found in breeding aggregations in Belize and Honduras. In Guatemala and
Honduras, the population is declining due to habitat destruction.
Habitat and Ecology It lives in lowland to montane moist forests on mountain slopes. It occurs in both pristine and
disturbed habitats. Breeding takes place in small intermittent or permanent waterbodies.
Major Threats Chytridiomycosis is probably the main cause of the disappearance of populations in Mexico, and the
species is now probably seriously at risk from this disease. Habitat destruction due to subsistence and smallholder
agriculture is also a threat to this species, which also was formerly common in the pet trade.
© Twan Leenders

Conservation Measures This species occurs in a number of protected areas throughout its range. Continued survey
work is needed to monitor the population status of this species, and particularly to determine whether or not the reason
for the apparent decline is due to chytridiomycosis. A captive-breeding programme might need to be established.
Bibliography: Duellman, W.E. (2001), Lee, J.C. (1996), Lips, K.R. et al. (2004), McCranie, J.R. and Wilson, L.D. (2002b)
Data Providers: Georgina Santos-Barrera, Julian Lee, Manuel Acevedo, Larry David Wilson

EN Argenteohyla siemersi (Mertens, 1937)

Endangered B2ab(iii) Geographic Range This species occurs in Argentina in north-west Corrientes Province (A. s. pederseni) and Entre
Order, Family: Anura, Hylidae Rios and Buenos Aires Provinces (A. s. siemersi). In Uruguay, it occurs in two localities in San Jose and Rocha states
Country Distribution: Argentina, Uruguay (A. s. siemersi). It occurs at elevations of 0-70m asl.
Current Population Trend: Decreasing Population The nominate subspecies was known to be uncommon, and the last record is from 1982. All populations
have disappeared from Argentina and Uruguay, the reason for which is not well understood. However, in 1983, the
Paraná had the largest flood of the century and the population here subsequently disappeared and has not been
found since, despite many attempts to relocate it. The subspecies A. s. pederseni is rare but was recorded in 2002
and 2003, and this population at least appears to be stable.
Habitat and Ecology A. s. pederseni occurs in leaf axils of terrestrial Aechmae bromeliads in gallery forests along
main river systems. It reproduces in temporary pools close to the bromeliads. The nominate subspecies occurs and
reproduces in wetlands and river deltas (Río Paraná), and tolerates moderate habitat disturbance.
Major Threats The main threat to this species is habitat destruction caused by fires used to expand or maintain
pastures and pine forestry plantations. Some populations are also likely to be threatened by the “Hidrovía Paraná-
Paraguay” project (the transformation of the Paraguay-Paraná-Uruguay-La Plata river system into a 3,400-km long
© Jorge Abel Cespedez

shipping canal). In Uruguay, this species is threatened by draining of wetlands for agricultural expansion.
Conservation Measures Some populations of A. s. pederseni occur in the 17,600-ha Parque Nacional Mburucuya.
The existing protected areas network requires expansion to ensure the protection of lowland riverine and wetland
habitats. Further survey work is necessary to determine the status of populations of the nominate subspecies.
Notes on taxonomy: There are two subspecies: Argenteohyla siemersi siemresi and A. s. pederseni.
Bibliography: Alvarez, B.B. et al. (2002), Alvarez, B.B. et al. (2003), Céspedez, J.A. (2000), Langone, J.A. (1994), Lavilla, E.O. et al. (2000),
Lavilla, E.O. and Cei, J.M. (2001), Maneyro, R. and Langone, J.A. (2001)
Data Providers: Esteban Lavilla, Jorge Céspedez, Diego Baldo, Boris Blotto, Jose Langone

CR Bokermannohyla izecksohni (Jim and Caramaschi, 1979)

Critically Endangered B1ab(iii,v)+2ab(iii,v) Geographic Range This species is known only from Rubião Júnior, Botucatu, in the state of São Paulo, Brazil.
Order, Family: Anura, Hylidae Population Surveys to locate this species in recent years have not found any individuals.
Country Distribution: Brazil Habitat and Ecology It was originally found on vegetation or on forest leaf-litter near streams. The type specimen
Current Population Trend: Decreasing was found in a temporary pond, which has subsequently disappeared. It breeds in temporary ponds.
Major Threats The type locality of the species has been destroyed by agriculture and human settlement.
Conservation Measures The current known range of the species is not within any protected area. Further survey
work is needed to determine if it occurs outside the vicinity of the type locality, and if it even still persists at the
type locality.
Notes on taxonomy: This species was previously included in the genus Hyla but has recently been moved to the genus Bokermannohyla
(Faivovich et al. 2005).
Bibliography: Faivovich, J. et al. (2005), Jim, J. and Caramaschi, U. (1979)
Data Providers: Miguel Trefaut Rodrigues, Carlos Alberto Gonçalves da Cruz
© Ivan Sazima
240 Threatened Amphibians of the World

EN Bromeliohyla bromeliacia (K. Schmidt, 1933)

Endangered A2ace Geographic Range This species occurs on the Atlantic versant of north-western Chiapas State, Mexico; central and
Order, Family: Anura, Hylidae eastern Guatemala; the Maya Mountains of Belize; and north-western Honduras, at 350-1,790m asl.
Country Distribution: Belize, Guatemala, Population It is known from single specimens in Belize and Mexico. It is uncommon in Guatemala, but it continues
Honduras, Mexico to occur in suitable habitat, although some populations are in serious decline. In fact, some higher elevation popula-
Current Population Trend: Decreasing tions in Guatemala, where most of the range occurs, appear to have declined sharply although the species has not
disappeared; populations at lower elevations appear stable. It is relatively common in Honduras.
Habitat and Ecology It is found in bromeliads in premontane and lower montane wet forest. It breeds by larval
development. It is also found in leaf sheaths on banana trees, and tolerates some level of habitat disturbance.
Major Threats The main threat is deforestation, due to agriculture (both crops and livestock), and water pollution
which is taking place at higher elevations. The declines observed at high elevations, which have taken place even in
suitable habitat, could be due to chytridiomycosis.
Conservation Measures It is found in Parque Nacional Montaña de Cusuco and Parque Nacional Cerro Azul in
© Silviu Petrovan

Honduras, and in the Reserva de la Biósfera Sierra de la Minas and Biotopo del Quetzal in Guatemala. Further research
is necessary to ascertain whether chytrid is a threat to this species.
Notes on taxonomy: This species was previously included in the genus Hyla but has recently been moved to the new genus Bromeliohyla
(Faivovich et al. 2005).
Bibliography: Campbell, J.A. (1998), Duellman, W.E. (2001), Faivovich, J. et al. (2005), Martinez-Coronel, M., Ramirez-Bautista, A. and
Vidal-Lopez, R. (1995), McCranie, J.R. and Wilson, L.D. (2002b), Schmidt, K.P. (1933)
Data Providers: Gustavo Cruz, Manuel Acevedo, Julian Lee

CR Bromeliohyla dendroscarta (Taylor, 1940)

Critically Endangered A2ace Geographic Range This species is found on the Atlantic slopes of the Sierra Madre Oriental and associated ranges
Order, Family: Anura, Hylidae in Mexico. It is also found in central Veracruz and northern Oaxaca in Mexico. It occurs from 450-1,900m asl.
Country Distribution: Mexico Population This has always been an uncommon species, but it appears to have gone into serious decline, and has
Current Population Trend: Decreasing not been recorded since 1974. Recent surveys to locate it have been unsuccessful, and it might now be extinct.
Society for the Study of Amphibians and Reptiles

Habitat and Ecology This species inhabits cloud forest habitats. It breeds and takes refuge in bromeliads.
Major Threats This species has disappeared in suitable habitat, probably due to chytridiomycosis. Deforestation and
© Hylid Frogs of Middle America, 2001,

alteration of the cloud forest are probably also major threats, as it requires bromeliads in mature trees to survive.
Conservation Measures The range of this species does not include any protected area. Additional survey work is
urgently needed to determine whether or not this species is still extant in its natural range. It is protected by Mexican
law under the “Special Protection” category (Pr).
Notes on taxonomy: This species was previously included in the genus Hyla but has recently been moved to the new genus Bromeliohyla
(Faivovich et al. 2005).
Bibliography: Duellman, W.E. (2001), Faivovich, J. et al. (2005), Lips, K.R. et al. (2004)
Data Providers: Georgina Santos-Barrera, Luis Canseco-Márquez

CR Charadrahyla altipotens (Duellman, 1968)

Critically Endangered A2ace Geographic Range This species is known from the Pacific slopes of the Sierra Madre del Sur de Oaxaca, and north
Order, Family: Anura, Hylidae to the towns of San Gabriel and San Sebastián, in south-western Oaxaca, Mexico. It might occur a little more widely
Country Distribution: Mexico than current records suggest.
Current Population Trend: Decreasing Population This has always been a rare species, but it appears to have gone into serious decline, and has not been
Society for the Study of Amphibians and Reptiles

recorded since the 1960s. Recent surveys to locate it have been unsuccessful, and it might now be extinct.
Habitat and Ecology This species occurs in pine-oak and cloud forests, and prefers rocky streams with abundant
© Hylid Frogs of Middle America, 2001,

vegetation as microhabitat. It presumably breeds in streams.

Major Threats This species has disappeared in suitable habitat, probably due to chytridiomycosis. The disappear-
ance of the fragments of cloud forest in Oaxaca, the most fragile habitat in Mexico, is also threatening this species.
At present this area is under extreme pressure from human population growth.
Conservation Measures The range of this species does not include any protected areas. While additional survey
work is urgently needed to determine whether or not this species is still extant in its natural range, protection of
the remaining cloud forest fragments is important to preserve the humid habitats for this species as well as other
amphibians. It is listed by the Mexican government in the category “Special Protection” (Pr).
Notes on taxonomy: This species was previously included in the genus Hyla but has recently been moved to the new genus Charadrahyla
(Faivovich et al. 2005).
Bibliography: Duellman, W.E. (2001), Faivovich, J. et al. (2005), Lips, K.R. et al. (2004)
Data Providers: Georgina Santos-Barrera, Luis Canseco-Márquez

EN Charadrahyla chaneque (Duellman, 1961)

Endangered B1ab(iii) Geographic Range This species is known from only three localities east of the Isthmus of Tehuantepec in northern
Order, Family: Anura, Hylidae Chiapas and eastern Oaxaca, Mexico. The type locality is at 1,690m asl.
Country Distribution: Mexico Population This is a common species within its restricted range.
Current Population Trend: Decreasing Habitat and Ecology It only inhabits high-elevation cloud forests with cascading mountain streams. It presumably
Society for the Study of Amphibians and Reptiles

breeds in streams.
Major Threats Habitat loss due to logging represents the main threat to this species.
© Hylid Frogs of Middle America, 2001,

Conservation Measures Its range includes the Reserva de la Biósfera Sierra del los Tuxtlas, but expanded protec-
tion of cloud forest habitat in this region is needed. This species is protected by Mexican law under the “Special
Protection” category (Pr).
Notes on taxonomy: This species was previously included in the genus Hyla but has recently been moved to the new genus Charadrahyla
(Faivovich et al. 2005).
Bibliography: Duellman, W.E. (2001), Faivovich, J. et al. (2005), Mendelson III, J.R. and Campbell, J.A. (1999), Perez-Higareda,
G. (1981b)
Data Providers: Antonio Muñoz Alonso, Luis Canseco-Márquez
 Globally Threatened Amphibian Species 241

VU Charadrahyla nephila (Mendelson and Campbell, 1999)

Vulnerable B1ab(iii) Geographic Range This species is known from Sierra de Juárez and Sierra Mixe, north-central Oaxaca, Mexico. A
Order, Family: Anura, Hylidae specimen has also been collected from Los Tuxtlas, Veracruz, Mexico, but this record is in doubt. It probably occurs
Country Distribution: Mexico more widely than records suggest. It occurs at elevations of 680-2,256m asl.
Current Population Trend: Decreasing Population This is a common species.
Habitat and Ecology It inhabits mesic cloud forest, and is commonly found in or near streams and low vegetation,
and presumably breeds in streams.
Major Threats The high rate of disturbance of the cloud forest is the main threat to this species. Tadpoles have been
found in southern Mexico with loss of keratinised mouthparts, suggesting that chytridiomycosis might be involved.
Conservation Measures The range of this species does not include any protected areas, and urgent protection
of the forests along the Sierra de Juárez and Sierra Mixe is required. The species is in need of close population
monitoring, particularly if chytrid is shown to be a genuine threat.
© J.R. Mendelson

Notes on taxonomy: This species was previously included in the genus Hyla, but has recently been moved to the new genus Charadrahyla
(Faivovich et al. 2005).
Bibliography: Duellman, W.E. (2001), Faivovich, J. et al. (2005), Lips, K.R. et al. (2004), Mendelson III, J.R. and Campbell, J.A. (1999)
Data Providers: Georgina Santos-Barrera, Luis Canseco-Márquez

VU Charadrahyla taeniopus (Günther, 1901)

Vulnerable B1ab(iii) Geographic Range This species occurs in north-eastern Hidalgo, southward through northern Puebla to central
Order, Family: Anura, Hylidae Veracruz, Mexico. It probably occurs more widely than current records suggest, especially in areas between known
Country Distribution: Mexico sites. It occurs at intermediate elevations.
Current Population Trend: Decreasing Population This is a common species. Several surveys conducted in the last few years in the Hidalgo-Veracruz region
Society for the Study of Amphibians and Reptiles

report healthy populations along its range.

Habitat and Ecology This species inhabits cloud forests characterized by moderate to low temperatures and high
© Hylid Frogs of Middle America, 2001,

humidity. It is always associated with streams and epiphytic plants. It presumably breeds in streams.
Major Threats Transformation of the forest into cultivated areas affects the presence of trees and epiphytic plants,
the preferred habitat of this species. In some towns in Veracruz, this species is consumed as a traditional cooked
dish; however, there is no information on the level of offtake.
Conservation Measures The range of this species does not include any protected areas, and there is clearly a need
for protection of the remaining cloud forest remnants in this area. It is listed as “Threatened” (Amenazada) by the
Mexican government. Further research is needed to determine the levels of offtake of this species from the wild.
Notes on taxonomy: This species was previously included in the genus Hyla but has recently been moved to the new genus Charadrahyla
(Faivovich et al. 2005).
Bibliography: Duellman, W.E. (2001), Faivovich, J. et al. (2005)
Data Providers: Georgina Santos-Barrera, Luis Canseco-Márquez

CR Charadrahyla trux (Adler and Dennis, 1972)

Critically Endangered B1ab(iii,v)+2ab(iii,v) Geographic Range This species is found on only on the slopes of Cerro Teótepec, in the Sierra Madre del Sur, in
Order, Family: Anura, Hylidae Guerrero, Mexico, at 1,760-2,415m asl.
Country Distribution: Mexico Population This is a rare species. Recent attempts to locate it have been unsuccessful, suggesting a population
Current Population Trend: Decreasing decline, and it might even be extinct.
Society for the Study of Amphibians and Reptiles

Habitat and Ecology This species inhabits pine-oak and bamboo-tree fern forests. It is commonly found associated
with cascading mountain streams where it breeds.
© Hylid Frogs of Middle America, 2001,

Major Threats It has possibly been impacted by chytridiomycosis. In addition, the transformation of the forests along
the transect between Filo de Caballo and Atoyac to open agricultural lands and farmlands is having a detrimental
impact on the remaining habitat of this species.
Conservation Measures The range of this species does not include any protected areas. While additional survey
work is urgently needed to determine whether or not this species is still extant in its natural range, a program to
protect the forest areas surrounding the Cerro Teótepec is also recommended. This species is listed as “Threatened”
(Amenazada) by the Mexican government.
Notes on taxonomy: This species was previously included in the genus Hyla but has recently been moved to the new genus Charadrahyla
(Faivovich et al. 2005).
Bibliography: Duellman, W.E. (2001), Faivovich, J. et al. (2005), Lips, K.R. et al. (2004)
Data Providers: Georgina Santos-Barrera, Luis Canseco-Márquez

CR Dendropsophus amicorum (Mijares-Urrutia, 1998)

Critically Endangered B1ab(iii) Geographic Range This species is known from a single specimen Notes on taxonomy: This species was previously included in the genus Hyla but has recently been moved to the resurrected genus
Order, Family: Anura, Hylidae taken at Cerro Socopó, 84km north-west of Carora, in Falcón state, Dendropsophus (Faivovich et al. 2005).
Country Distribution: Venezuela Venezuela (10.28N, 70.48W), at 1,250m asl. It is assumed to be Bibliography: Barrio Amorós, C.L. (2004), Faivovich, J. et al. (2005), Mijares-Urrutia, A. (1998)
Current Population Trend: Decreasing restricted to the Cerro Socopó. Data Providers: Enrique La Marca, Abraham Mijares
Population The population status of this species is not known.
Habitat and Ecology The Cerro Socopó is a relict of cloud forest
surrounded by semi-arid vegetation. It presumably breeds by larval
development, but the site of egg deposition is not known.
Major Threats All of the area of Cerro Socopó has been severely
affected by forest destruction for agriculture (crops and livestock),
as well as by agricultural pollution.
Conservation Measures The range of this species does not include
any protected areas, and the species’ habitat is in urgent need of
protection. Additional survey work is needed to determine the biology
and population status and trends of this species.
242 Threatened Amphibians of the World

EN Dendropsophus gryllatus (Duellman, 1973)

Endangered B1ab(iii) Geographic Range This species is known from the Pacific lowlands of north-western Ecuador, from 200-500m asl.
Order, Family: Anura, Hylidae It is known from three localities, but with further sampling is likely to occur more widely.
Country Distribution: Ecuador Population It is a rare species.

© Natural History Museum and Biodiversity Research

Current Population Trend: Decreasing Habitat and Ecology This is originally a lowland forest species. However, specimens have been collected at a large
duckweed-covered pond in a banana plantation outside forest; a few individuals were calling from bushes at the
edge of the water, while others were found on grasses overhanging shallow muddy pools (Duellman 1973). It breeds
in ponds, and probably deposits its eggs in the water.
Major Threats The main threat to this species is habitat loss, and much of the natural vegetation within its range has
been cleared. While its occurrence in banana plantations may suggest that the species can survive in altered habitats,
Center, University of Kansas farming practices have changed radically over the last 20 years, especially with the use of pesticides, and cultivation
that is more intensive. The soil is very fertile, and banana plantations do very well; however, these plantations are
increasingly intensively managed and the use of pesticides poses an increasing threat.
Conservation Measures It is not known to occur in any protected areas. There is an urgent need for protection of
remaining lowland forest habitat in this part of Ecuador.
Notes on taxonomy: This species was previously included in the genus Hyla but has recently been moved to the resurrected genus
Dendropsophus (Faivovich et al. 2005).
Bibliography: Duellman, W.E. (1973), Faivovich, J. et al. (2005)
Data Providers: Luis A. Coloma, Santiago Ron, Diego Cisneros-Heredia, Ana Almandáriz

EN Dendropsophus meridensis (Rivero, 1961)

Endangered B1ab(iii,v) Geographic Range This species is known from the Andes of Mérida State, in Venezuela. It has been recorded from
Order, Family: Anura, Hylidae 1,200-2,400m asl. Records from the Sierra del Turimiquire, in Sucre and Monagas States, are misidentifications.
© César Luis Barrio-Amorós / Fundación AndígenA

Country Distribution: Venezuela Population This is an uncommon species that is estimated to have undergone a significant decline over the past
Current Population Trend: Decreasing ten years.
Habitat and Ecology It is an inhabitant of small ponds in cloud forests. It lays its eggs directly in water, where
the larvae also develop.
Major Threats The main threats are agriculture, involving both crops and livestock, as well as agricultural pollu-
tion. The recent introduction of Rana catesbeiana has probably been at least parly responsible for the decline in
populations of this frog.
Conservation Measures Although most of the species’ range is outside protected areas, it may occur in Parque
Nacional La Culata and/or Parque Nacional Sierra Nevada. Remaining cloud forest habitats in the Venezuelan Andes
are in urgent need of protection.
Notes on taxonomy: This species was previously included in the genus Hyla but has recently been moved to the resurrected genus
Dendropsophus (Faivovich et al. 2005).
Bibliography: Barrio Amorós, C.L. (2004), Duellman, W.E. (1977), Duellman, W.E. (1989b), Faivovich, J. et al. (2005), La Marca, E. (1992),
La Marca, E. (1995b), La Marca, E. (1997), Péfaur, J.E. and Rivero, J.A. (2000), Rivero, J.A. (1961), Vial, J.L. and Saylor, L. (1993)
Data Providers: Enrique La Marca

VU Dendropsophus stingi (Kaplan, 1994)

Vulnerable D2 Geographic Range This species is found on the eastern slope of Bibliography: Faivovich, J. et al. (2005), Kaplan, M. (1994)
Order, Family: Anura, Hylidae the Cordillera Oriental in Boyacá Department, Colombia, at elevations Data Providers: María Cristina Ardila-Robayo, Jose Vicente Rueda
Country Distribution: Colombia of 2,000-2,020m asl.
Current Population Trend: Increasing Population This species is abundant and there are many recent
records. Some populations even appear to be increasing.
Habitat and Ecology It occurs and reproduces in shrubs in flooded
pastures, marshes, and temporary pools.
Major Threats There are no current major threats; however, its small
range renders it susceptible to stochastic threatening processes.
Conservation Measures It does not occur in any protected areas.
There is a need for close population monitoring of this species
considering its small range.
Notes on taxonomy: This species was previously included in the genus
Hyla but has recently been moved to the resurrected genus Dendropsophus
(Faivovich et al. 2005).

EN Duellmanohyla chamulae (Duellman, 1961)

Endangered B1ab(iii) Geographic Range This species occurs above 1,600m asl on the northern slopes of the central highlands of Chiapas,
Order, Family: Anura, Hylidae Mexico, from Jitotol to Soluschiapa.
Country Distribution: Mexico Population This is not a rare species.
Current Population Trend: Decreasing Habitat and Ecology It inhabits and breeds in cascading mountain streams in cloud forest areas.
Society for the Study of Amphibians and Reptiles

Major Threats The main threat is deforestation and the disturbance of cloud forest remnants (at present, moun-
tainous cloud forests are the most impacted habitats in Mexico). Chytridiomycosis may also also pose apotential
© Hylid Frogs of Middle America, 2001,

threat to this species.

Conservation Measures The range of this species does not include any protected areas, and the protection
and restoration of forested highland areas of northern Chiapas, Mexico, is needed. Further research is required to
determine whether chytrid might pose a threat to this species. It is protected by Mexican law under the “Special
Protection” category (Pr).
Bibliography: Campbell, J.A. and Smith, E.N. (1992), Duellman, W.E. (2001)
Data Providers: Georgina Santos-Barrera, Antonio Muñoz Alonso
 Globally Threatened Amphibian Species 243

EN Duellmanohyla ignicolor (Duellman, 1961)

Endangered B1ab(iii)+2ab(iii) Geographic Range This species is known from the Sierra de Juárez, northern Oaxaca, Mexico, at elevations of
Order, Family: Anura, Hylidae 680-1,850m asl.
Country Distribution: Mexico Population This species is relatively uncommon and is known only from a few localities.
Current Population Trend: Decreasing Habitat and Ecology It is a stream-breeding amphibian that requires the presence of streams and humid micro-

Society for the Study of Amphibians and Reptiles

habitats.
Major Threats The main threat to this species is the disturbance and desiccation of streams in cloud forest. In

© Hylid Frogs of Middle America, 2001,

addition, larvae with keratinized mouthparts have been found in southern Mexico, which suggests infection with
chytridiomycosis.
Conservation Measures Urgent protection of the Sierra de Juárez area is suggested since there are no existing
protected areas in the region. In view of what appears to be a genuine risk of chytridiomycosis, the status of this
species should be closely monitored, and ex-situ populations may need to be established. This species is protected
by Mexican law under the “Special Protection” category (Pr).
Bibliography: Campbell, J.A. and Smith, E.N. (1992), Duellman, W.E. (2001), Lips, K.R. et al. (2004)
Data Providers: Georgina Santos-Barrera, Antonio Muñoz Alonso, Luis Canseco-Márquez

EN Duellmanohyla lythrodes (Savage, 1968)

Endangered B1ab(iii) Geographic Range This species is known from the humid lowlands of the Atlantic versant in southern Costa Rica
Order, Family: Anura, Hylidae and adjacent north-western Panama, from 170-440m asl (Savage 2002).
Country Distribution: Costa Rica, Panama Population It is a rare species known only from three specimens.
Current Population Trend: Decreasing Habitat and Ecology The few known specimens have been collected on low vegetation (approximately 3m above
Society for the Study of Amphibians and Reptiles

ground) within humid lowland forest. The species may be associated with headwater streams (Savage 2002). Larval
development is presumed to take place in these streams.
© Hylid Frogs of Middle America, 2001,

Major Threats The major threat to the species is general habitat loss due to deforestation for smallholder farming
activities, logging, and human settlement. Chytridiomycosis is also a potential threat to this species.
Conservation Measures It has not been recorded from any protected areas, and the lowland forest habitat of this
species is in urgent need of protection. Further research is needed to determine the current population status of this
species, and whether it is at risk of infection with chytrid.
Bibliography: Campbell, J.A. and Smith, E.N. (1992), Duellman, W.E. (1970), Duellman, W.E. (2001), Ibáñez, R. et al. (2000), Savage,
J.M. (1968a), Savage, J.M. (2002), Young, B. et al. (1999)
Data Providers: Frank Solís, Roberto Ibáñez, Gerardo Chaves, Jay Savage, César Jaramillo, Querube Fuenmayor

VU Duellmanohyla rufioculis (Taylor, 1952)

Vulnerable B1ab(iii) Geographic Range This species occurs on the Caribbean and Pacific slopes of the mountains of Costa Rica, at
Order, Family: Anura, Hylidae elevations of 775-1,580m asl.
Country Distribution: Costa Rica Population It is regularly encountered in suitable habitat throughout its range.
Current Population Trend: Decreasing Habitat and Ecology It lives in premontane wet forest and rainforest. A nocturnal stream breeder, males call
throughout the year but are most active from August to December. The species sometimes aggregates around small,
shallow streams or seeps draining into streams. Tadpoles are found in quiet pools.
Major Threats Major threats include deforestation, stream alteration, and pollution. Chytridiomycosis is also a
potential threat.
Conservation Measures Its range includes several national parks, such as Braulio Carrillo, and other protected areas.
The species is in need of close population monitoring, particularly if chytrid is shown to represent a genuine threat.
Bibliography: Campbell, J.A. and Smith, E.N. (1992), Pounds, J.A. et al. (1997), Savage, J.M. (2002)
Data Providers: Federico Bolaños, Gerardo Chaves
© Brian Kubicki

CR Duellmanohyla salvavida (McCranie and Wilson,1986)

Critically Endangered B2ab(iii,v) Geographic Range This species occurs in Sierra de Nombre de Dios and Montana Macuzal, Atlantica and Yoro
Order, Family: Anura, Hylidae departments, north-central Honduras, at elevations of 90-1,400m asl.
Country Distribution: Honduras Population It is uncommon and all populations are believed to be declining.
Current Population Trend: Decreasing Habitat and Ecology It lives in lowland moist forest and premontane wet forest. It has been collected along shallow,
slow-moving streams. It lays eggs in vegetation over streams.
Major Threats Habitat loss and degradation due to subsistence and smallholder agriculture, and logging, is the
main threat to this species. Landslides on the upper clear water streams (caused by strong storms and human activi-
ties in the lower portions of the streams), water pollution, and fires are also threats. Chytridiomycosis presents a
potential threat to this species.
Conservation Measures It occurs within the Parque Nacional Pico Bonito and the Refugio de Vida Silvestre
Texiguat. There is a need for close monitoring of the population status of this species, particularly given the threat
© James McCranie

of chytridiomycosis.
Bibliography: Campbell, J.A. and Smith, E.N. (1992), McCranie, J.R. and Wilson, L.D. (1986), McCranie, J.R. and Wilson, L.D. (1990),
McCranie, J.R. and Wilson, L.D. (2002b), Wilson, L.D. and McCranie, J.R. (1998)
Data Providers: Gustavo Cruz, Larry David Wilson
244 Threatened Amphibians of the World

VU Duellmanohyla schmidtorum (Stuart, 1954)

Vulnerable B1ab(iii)+2ab(iii) Geographic Range This species is known from extreme eastern Oaxaca (Chimalapas) and extreme south-western
Order, Family: Anura, Hylidae Chiapas, Mexico, and adjacent Guatemala.
Country Distribution: Guatemala, Mexico Population It is not very common.
Current Population Trend: Decreasing Habitat and Ecology This species only inhabits montane cloud forest from moderate to high elevations in almost

Society for the Study of Amphibians and Reptiles

pristine environments. It is presumed to breed in streams.
Major Threats The major threat is habitat loss due to deforestation and infrastructure development. However,

© Hylid Frogs of Middle America, 2001,

chytridiomycosis is also a potential threat.
Conservation Measures The range of this species includes Reserva de la Biósfera El Triunfo. There is a need for
close population monitoring, particularly if chytrid is shown to represent a genuine threat. This species is protected
by Mexican law under the “Special Protection” category (Pr).
Bibliography: Campbell, J.A. (2001), Campbell, J.A. and Smith, E.N. (1992), Duellman, W.E. (2001)
Data Providers: Georgina Santos-Barrera, Antonio Muñoz Alonso

CR Duellmanohyla soralia (Wilson and McCranie, 1985)

Critically Endangered B2ab(iii,v) Geographic Range This species occurs in Sierra de Omoa and Espiritu Santo in north-western Honduras and north-
Order, Family: Anura, Hylidae eastern Guatemala, between 40 and 1,570m asl.
Country Distribution: Guatemala, Honduras Population Formerly, it was moderately common in appropriate habitat. Currently, though, the population is declin-
Current Population Trend: Decreasing ing throughout its range.
Habitat and Ecology It is found on low vegetation along streams in lowland moist forest, and premontane and
lower montane wet forest. Breeding and larval development take place in streams.
Major Threats The main threats to the species are habitat loss due to agriculture (for uses such as pasture) and
wood extraction, and water pollution. Chytridiomycosis is also a potential threat.
Conservation Measures In Honduras, it is known to occur in Parque Nacional Cusuco and in the Parque Nacional
Cerro Azul, both formally protected since 1987. Surveys are required to monitor the population status and trends of
© Franklin E. Castañeda

this species, particularly given the potential threat of chytridiomycosis.

Bibliography: Campbell, J.A. and Smith, E.N. (1992), McCranie, J.R. and Wilson, L.D. (2002b), Wilson, L.D. and McCranie, J.R. (1985)
Data Providers: Gustavo Cruz, Manuel Acevedo, Larry David Wilson

CR Duellmanohyla uranochroa (Cope, 1875)

Critically Endangered A2ace Geographic Range This species is known from the cordilleras of Costa Rica and western Panama (300-1,450m asl).
Order, Family: Anura, Hylidae In Costa Rica, the species occurs on the Atlantic versant at 656-1,740m asl and on the Pacific slope at 880-1,600m
Country Distribution: Costa Rica, Panama asl (Savage 2002).
Current Population Trend: Decreasing Population It was formerly common but has disappeared from its entire Costa Rican range except for a remnant
population at Monteverde where it has declined substantially since the late 1980s. It is now infrequently seen in
Costa Rica, and it is presumed to have declined in Panama.
Habitat and Ecology It inhabits humid lowland and montane forest. Adults are usually found in the proximity of
streams. During the day, individuals hide in vegetation, especially in leaf axils of epiphytes and terrestrial aroids. At
night, moderately dense congregations have been found along small, fast-flowing streams. Males call from dense
vegetation several metres away from the streams, from 0.5-3.0m above ground. Larvae develop in quiet pools, but
sometimes adhere to large rocks in the stream bottom when the streams rise following heavy rains (Savage 2002).
Major Threats Museum specimens have been found to be infected with chytrid fungi, and although it requires
© Brian Kubicki

confirmation, it is likely that infection with this pathogen, perhaps in combination with climate change, is responsible
for the current population decline. Habitat loss due to smallholder farming activities is also a threat to this species.
Conservation Measures Although there are no specific conservation measures in place, this species has been
recorded from at least three protected areas in Panama, and more than three protected areas in Costa Rica. Further
survey work is required to monitor the population status and trends of this species, and to ascertain clearly whether
or not chytrid poses a threat or not. A captive-breeding programme might need to be established.
Bibliography: Duellman, W.E. (2001), Ibáñez, R. et al. (2000), Savage, J.M. (1968a), Savage, J.M. (2002), Young, B. et al. (1999)
Data Providers: Alan Pounds, Gerardo Chaves, Frank Solís, Roberto Ibáñez, Jay Savage, César Jaramillo, Querube Fuenmayor

CR Ecnomiohyla echinata (Duellman, 1962)

Critically Endangered A2ace Geographic Range This species is known only from the type locality Conservation Measures This species is not known to occur in any protected areas, and urgent protection of the cloud
Order, Family: Anura, Hylidae at around 2,000m asl on the northern slopes of the Sierra de Juárez, in forest areas in Sierra de Juárez is recommended. A survey to determine whether or not this species is still extant in
Country Distribution: Mexico the vicinity of Vista Hermosa town, north-central Oaxaca, Mexico. the wild is also required. It is protected by Mexican law under the “Special Protection” category (Pr).
Current Population Trend: Decreasing Population This has always been a rare species, but it appears Notes on taxonomy: This species was previously included in the genus Hyla but has recently been moved to the new genus Ecnomiohyla
to have gone into serious decline, and has not seen recorded since (Faivovich et al. 2005).
1962. Recent surveys to locate it have been unsuccessful, and it Bibliography: Duellman, W.E. (2001), Faivovich, J. et al. (2005), Lips, K.R. et al. (2004)
might now be extinct. Data Providers: Georgina Santos-Barrera, Luis Canseco-Márquez
Habitat and Ecology This species is restricted to cloud forest, and is
highly dependent on humid habitats and in particular, the vegetation
along low streams as well as epiphytic plants that serve as a refuge.
It is a stream-breeding species.
Major Threats This species has disappeared in suitable habitat,
probably due to chytridiomycosis. The disappearance or disturbance
of the unique cloud forest environment, primarily through selective
logging, at Sierra de Juárez is also a serious threat to the survival
of this species as well as many other amphibians, especially those
associated with humid habitats.
 Globally Threatened Amphibian Species 245

EN Ecnomiohyla fimbrimembra (Taylor, 1948)

Endangered B1ab(iii) Geographic Range This species occurs on the slopes of the Cordillera de Tilarán, Cordillera Central, and Cordillera
Order, Family: Anura, Hylidae de Talamanca of Costa Rica and western Panama, from 750-1,900m asl (Savage 2002). It almost certainly occurs
Country Distribution: Costa Rica, Panama more widely than has been mapped, and is also probably found between known sites.
Current Population Trend: Decreasing Population Because this is a high-canopy species, it is overlooked, and it is therefore hard to assess its population
status.
Habitat and Ecology This is a nocturnal canopy species associated with primary humid premontane and lower
montane forest. Breeding, egg deposition and larval development take place within tree-holes.
Major Threats The main threat is general habitat loss as a result of deforestation due primarily to the clearing of
land for livestock ranching.
Conservation Measures The species has been recorded from Parque Internacional La Amistad, Panama, and in
more than three protected areas within Costa Rica.
© Jessica Alison

Notes on taxonomy: This species was previously included in the genus Hyla but has recently been moved to the new genus Ecnomiohyla
(Faivovich et al. 2005).
Bibliography: Duellman, W.E. (2001), Faivovich, J. et al. (2005), Hayes, M.P., Pounds, J.A. and Robinson, D.C. (1986), Ibanez, D.R. et al.
(1991), Ibáñez, R. et al. (2000), Savage, J.M. (1981), Savage, J.M. (2002), Young, B. et al. (1999)
Data Providers: Frank Solís, Roberto Ibáñez, Gerardo Chaves, Jay Savage, César Jaramillo, Querube Fuenmayor

VU Ecnomiohyla miliaria (Cope, 1886)

Vulnerable B1ab(iii) Geographic Range This species is known from humid lowlands and premontane slopes from south-eastern Nicaragua
Order, Family: Anura, Hylidae to south-eastern Costa Rica on the Atlantic versant (20-900m asl), and on the Pacific versant in humid premontane
Country Distribution: Costa Rica, Nicaragua, areas of south-western Costa Rica and western and central Panama, at 1,000-1,330m asl (Savage 2002). It is expected
Panama to have a wider distribution than is currently known.
Current Population Trend: Decreasing Population There is no information on the population status of this rarely seen, high-canopy species.
Habitat and Ecology This is a nocturnal canopy species of primary humid lowland and montane forest. Eggs are
deposited in tree holes, which is also where the larvae develop.
Major Threats The major threat is loss of primary forest habitat due to agricultural expansion, logging, and human
settlement.
Conservation Measures The species has been recorded from several protected areas in both Panama and Costa
Rica. Further surveys are required to fully determine the range of this species.
Notes on taxonomy: This species was previously included in the genus Hyla but has recently been moved to the new genus Ecnomiohyla
(Faivovich et al. 2005). We follow Duellman (2001) and Savage (2002) in assigning records from Colombia and Ecuador to Ecnomiohyla
© Brian Kubicki

phantasmagoria.
Bibliography: Duellman, W.E. (1970), Duellman, W.E. (2001), Faivovich, J. et al. (2005), Ibáñez, R. et al. (2000), McCranie, J.R.,
Townsend, J.H. and Wilson, L.D. (2003), Pounds, J.A. et al. (1997), Savage, J.M. (2002), Wilson, L.D., McCranie, J.R. and Williams, K.L.
(1985), Young, B. et al. (1999)
Data Providers: Frank Solís, Roberto Ibáñez, Gerardo Chaves, Jay Savage, César Jaramillo, Querube Fuenmayor

EN Ecnomiohyla minera (Wilson, McCranie and Williams, 1985)

Endangered B1ab(iii) Geographic Range This species is known from two disjunct populations, in the Municipio de Purulhá, in the
Order, Family: Anura, Hylidae department of Baja Verapaz, and the Sierra de Santa Cruz, Izabal Department, both in Guatemala, at 700-1,830m
Country Distribution: Guatemala asl. It has been reported, but not collected, from the Reserva de la Biósfera Sierra de las Minas (this locality is not
Current Population Trend: Decreasing mapped here).
Society for the Study of Amphibians and Reptiles

Population It has always been rare, and has not been seen since 1994, although there have been no recent searches
specifically targeting this species.
© Hylid Frogs of Middle America, 2001,

Habitat and Ecology It occurs in the vegetation of pristine cloud forest, and is assumed to breed in tree cavities
or bromeliads.
Major Threats The main threat to this species is habitat loss, primarily due to agriculture, logging, and expanding
ornamental plant farms.
Conservation Measures It occurs close to Biotopo del Quetzal, and possibly occurs in the Reserva de la Biósfera
Sierra de las Minas. The cloud forest habitat of this species is in need of further protection.
Notes on taxonomy: This species was previously included in the genus Hyla but has recently been moved to the new genus Ecnomiohyla
(Faivovich et al. 2005).
Bibliography: Campbell, J.A. (1998), Faivovich, J. et al. (2005), Lee, J.C. (1996), Wilson, L.D., McCranie, J.R. and Williams, K.L. (1985)
Data Providers: Manuel Acevedo, Bruce Young

EN Ecnomiohyla phantasmagoria (Dunn, 1943)

Endangered B1ab(iii) Geographic Range This species is known only from two localities Conservation Measures It is not known from any protected areas, and habitat protection is urgently required. Further
Order, Family: Anura, Hylidae nearly 1,000km apart from each other: Río Cauca, near the border of survey work is necessary to determine the current population status of this species and the limits of its range.
Country Distribution: Colombia, Ecuador Antioquia and Bolivar Departments in north-western Colombia; and Notes on taxonomy: This species was previously included in the genus Hyla but has recently been moved to the new genus
Current Population Trend: Decreasing extreme north-western Esmeraldas Province, in Ecuador, at 500m. Ecnomiohyla (Faivovich et al. 2005). We follow Duellman (2001) and Savage (2002) in considering this as a species distinct from
It is likely to occur between these two localities. Ecnomiohyla miliaria.
Population It appears to be an extremely rare species, though Bibliography: Duellman, W.E. (2001), Faivovich, J. et al. (2005), Rueda-Almonacid, J.V. (1999), Savage, J.M. (2002)
this might be an artefact of it being a canopy species that is very Data Providers: Karl-Heinz Jungfer, Juan Manuel Renjifo
hard to detect.
Habitat and Ecology It is an inhabitant of the canopy of rainforest,
and probably breeds in tree holes.
Major Threats The major threats are likely to be deforestation for
agricultural development, planting of illegal crops, logging, and hu-
man settlement, and pollution resulting from the spraying of illegal
crops. Both localities from which is has been recorded are seriously
threatened, with the only known site in Colombia particularly threat-
ened by the expansion of livestock farming.
246 Threatened Amphibians of the World

CR Ecnomiohyla salvaje (Wilson, McCranie and Williams, 1985)

Critically Endangered B1ab(iii) Geographic Range This species is known from Cerro del Mono near Le Union in eastern Guatemala close to the
Order, Family: Anura, Hylidae border with Honduras; and nearby at Quebrada Grande, Department de Copan, in extreme western Honduras. Its
Country Distribution: Guatemala, Honduras altitudinal range is 1,370-1,450m asl.
Current Population Trend: Decreasing Population This is a poorly known species. Only two adult specimens have been collected from Honduras and one

Society for the Study of Amphibians and Reptiles

juvenile specimen from Guatemala. It is extremely rare and is probably in decline. Recent visits to the Honduran site
have failed to find the species.

© Hylid Frogs of Middle America, 2001,

Habitat and Ecology It inhabits lower montane wet forest, where it breeds in tree cavities.
Major Threats The primary threat to the species is severe deforestation, due to coffee cultivation and livestock grazing.
The only known site in Honduras is highly disturbed, but the habitat is in better condition at the Guatemalan site.
Conservation Measures It is not known from any protected areas, although it is on edge of the Parque Nacional
Cerro Azul in Honduras. Maintenance and/or protection of the species’ remaining intact habitat is recommended.
Notes on taxonomy: This species was previously included in the genus Hyla but has recently been moved to the new genus Ecnomiohyla
(Faivovich et al. 2005).
Bibliography: Campbell, J.A. (2001), Duellman, W.E. (2001), Faivovich, J. et al. (2005), McCranie, J.R. and Wilson, L.D. (2002b), Wilson,
L.D., McCranie, J.R. and Williams, K.L. (1985)
Data Providers: Gustavo Cruz, Larry David Wilson, Randy McCranie, Manuel Acevedo

CR Ecnomiohyla valancifer (Firschein and Smith, 1956)

Critically Endangered B2ab(iii) Geographic Range This species is found in the Sierra de Los Tuxtlas at Volcán San Martin, southern Veracruz,
Order, Family: Anura, Hylidae Mexico. It is known from 500-1,180m asl. There is also one specimen from Toledo District in Belize that has been
Country Distribution: Mexico assigned to this species, but further taxonomic work is required to confirm the identity of this population, which
Current Population Trend: Decreasing probably belongs to another species.
Society for the Study of Amphibians and Reptiles

Population This is a rare species, and it is known from only two specimens collected at the type locality.
Habitat and Ecology This species inhabits tropical rainforest on the slopes of Volcán San Martin. It is probably a
© Hylid Frogs of Middle America, 2001,

stream-breeding species.
Major Threats Disturbance and transformation of the original forest habitat, primarily due to agricultural develop-
ment, is a major threat to this species in its range (outside Reserva de la Biósfera Los Tuxtlas).
Conservation Measures The range of the species includes Reserva de la Biósfera Los Tuxtlas, which is quite
well managed. However, protection and restoration of the Volcán Sán Martin and surrounding areas is urgently
required to ensure the species’ persistence. This species is protected by Mexican law under the “Special Protec-
tion” category (Pr).
Notes on taxonomy: This species was previously included in the genus Hyla but has recently been moved to the new genus Ecnomiohyla
(Faivovich et al. 2005).
Bibliography: Duellman, W.E. (2001), Faivovich, J. et al. (2005), Lee, J.C. (2000)
Data Providers: Julian Lee, Oscar Flores-Villela

EN Exerodonta catracha (Porras and Wilson, 1987)

Endangered B1ab(iii)+2ab(iii) Geographic Range This species occurs in the Sierra de Opalaca and the Sierra de Montecillos, Montana de La
Order, Family: Anura, Hylidae Sierra, in south-west Honduras, at 1,800-2,160m asl.
Country Distribution: Honduras Population It is considered moderately common.
Current Population Trend: Decreasing Habitat and Ecology It lives in arboreal bromeliads in lower montane moist forest, and breeds in ponds.
Society for the Study of Amphibians and Reptiles

Major Threats The main threat to the species is deforestation primarily due to agricultural expansion (involving both
crops and livestock), logging, and human settlement.
© Hylid Frogs of Middle America, 2001,

Conservation Measures The range of this species includes the Reserva de la Biósfera Sierra de Montecillos.
Expanded protection of the montane forests of Honduras is necessary.
Notes on taxonomy: This species was previously included in the genus Hyla but has recently been moved to the resurrected genus
Exerodonta (Faivovich et al. 2005).
Bibliography: Faivovich, J. et al. (2005), McCranie, J.R. and Wilson, L.D. (2002b), McCranie, J.R., Wilson, L.D. and Williams,
K.L. (1993b), Porras, L. and Wilson, L.D. (1987)
Data Providers: Gustavo Cruz, Larry David Wilson, Franklin Casteñeda

EN Exerodonta chimalapa (Mendelson and Campbell, 1994)

Endangered B1ab(iii) Geographic Range This species is known only from the Chimalapas region that includes the Sierra Atravesada in
Order, Family: Anura, Hylidae south-eastern Oaxaca and adjacent Chiapas, Mexico, at 850-1500m asl.
Country Distribution: Mexico Population It is known to be common.
Current Population Trend: Decreasing Habitat and Ecology This stream-breeding amphibian inhabits low or moderate altitude streams in cloud forest or
dry pine-oak forest, and takes refuge in bromeliads.
Major Threats The main threat is deforestation resulting from logging, and the subsequent dessication of forest
streams.
Conservation Measures It does not occur in any protected areas; as with many other montane cloud and pine-oak for-
est patches in Mexico, there is an urgent need for the protection of the forest remnants in the Chimalapas region.
Notes on taxonomy: This species was previously within the genus Hyla but has recently been moved to the resurrected genus
Exerodonta (Faivovich et al. 2005).
Bibliography: Duellman, W.E. (2001), Faivovich, J. et al. (2005), Mendelson III, J.R. and Campbell, J.A. (1994)
© Thomas Bille

Data Providers: Antonio Muñoz Alonso, Luis Canseco-Márquez

 Globally Threatened Amphibian Species 247

VU Exerodonta juanitae (Snyder, 1972)

Vulnerable B1ab(iii) Geographic Range This species is known from two disjunct populations: one in San Vicente de Benitez, in the
Order, Family: Anura, Hylidae Sierra Madre del Sur, Guerrero, and the other in San Gabriel Mixtepec, Oaxaca, also in the Sierra Madre del Sur,
Country Distribution: Mexico south-western Mexico. It is found at elevations of 750-1,080m asl in Guerrero, and at elevations of 580-1,530m asl
Current Population Trend: Decreasing in Oaxaca. It might occur more widely.

Society for the Study of Amphibians and Reptiles

Population This species is not common. Recent surveys suggest that it has undergone a decline in numbers.
Habitat and Ecology This species only inhabits montane forest at moderate elevations. It breeds in temporary

© Hylid Frogs of Middle America, 2001,

streams.
Major Threats The destruction and degradation of the original forest cover (mainly due to logging) in Guerrero, is
the major threat to this species, and most of the localities where the species occurs have been severely impacted by
human activities. Recently documented declines might be due to chytridiomycosis.
Conservation Measures The range of this species is not within any protected areas, and there is a need for improved
habitat protection at sites where this species is known to occur. The Government of Mexico considers it “Threatened”
(Amenazada). Close population monitoring is required, especially given the potential threat of chytridiomycosis.
Notes on taxonomy: This species was previously included in the genus Hyla but has recently been moved to the resurrected genus
Exerodonta (Faivovich et al. 2005).
Bibliography: Duellman, W.E. (2001), Faivovich, J. et al. (2005), Lips, K.R. et al. (2004)
Data Providers: Georgina Santos-Barrera, Luis Canseco-Márquez

VU Exerodonta melanomma (Taylor, 1940)

Vulnerable B1ab(iii) Geographic Range This species is found only on the Pacific slopes of the Sierra Madre del Sur in Guerrero and
Order, Family: Anura, Hylidae Oaxaca, Mexico. It probably occurs more widely, especially in intervening areas between currently known sites. It
Country Distribution: Mexico occurs at elevations of 900 and 2,000m asl.
Current Population Trend: Decreasing Population This is a common species.
Society for the Study of Amphibians and Reptiles

Habitat and Ecology It inhabits cloud forest and pine-oak forest, and is almost exclusively found in forest streams
and the surrounding vegetation. Animals seek refuge in bromeliads during the dry season. Breeding takes place in
© Hylid Frogs of Middle America, 2001,

streams.
Major Threats The major threat is habitat loss and disturbance due to expanding agriculture (crops and livestock),
logging, and human settlement. Tadpoles have been found in southern Mexico with loss of keratinized mouthparts,
which suggests that chytridiomycosis might be a threat.
Conservation Measures The range of this species does not include any protected areas, and there is a need for
improved protection, and/or restoration, of pine-oak forests at sites where this species is known to occur. Close
population monitoring is required, especially given the potential threat of chytridiomycosis. It is protected by Mexican
law under the “Special Protection” category (Pr).
Notes on taxonomy: This species was previously included in the genus Hyla but has recently been moved to the resurrected genus
Exerodonta (Faivovich et al. 2005).
Bibliography: Campbell, J.A. and Duellman, W.E. (2000), Duellman, W.E. (2001), Faivovich, J. et al. (2005), Lips, K.R. et al. (2004)
Data Providers: Georgina Santos-Barrera, Luis Canseco-Márquez

CR Exerodonta perkinsi (Campbell and Brodie, 1992)

Critically Endangered B1ab(iii,v) Geographic Range This species is known only from the type locality (approximately 15° 53’N; 91° 16’W) on the
Order, Family: Anura, Hylidae northern slope of the Sierra de Cuchumatanes, Guatemala, at 1,050-1,080m asl.
Country Distribution: Guatemala Population It is rare and declining. Only 20 specimens have been collected and it is becoming harder to find.
Current Population Trend: Decreasing Habitat and Ecology It inhabits subtropical rainforest, breeding in temporary pools and slow-flowing streams, and
Society for the Study of Amphibians and Reptiles

also in the axils of elephant-ear plants. It can survive in heavily degraded habitats, providing that the habitat is not
opened up too much.
© Hylid Frogs of Middle America, 2001,

Major Threats The major threat is habitat loss due to human settlement (returning refugees), which is associated
with expanding small-scale agriculture and wood collection. Chytridiomycosis also cannot be ruled out as a cause
of the observed decline.
Conservation Measures The species is not known to occur in any protected areas, and protection of its remaining
intact habitat is recommended. Survey work is required to determine the current population status of this species.
Notes on taxonomy: This species was previously included in the genus Hyla but has recently been moved to the resurrected genus
Exerodonta (Faivovich et al. 2005).
Bibliography: Campbell, J.A. (2001), Campbell, J.A. and Brodie, E.D., Jr. (1992), Duellman, W.E. (2001), Faivovich, J. et al. (2005)
Data Providers: Manuel Acevedo, Eric Smith

VU Exerodonta pinorum (Taylor, 1937)

Vulnerable B1ab(iii,v) Geographic Range This species is known from the Pacific slopes of the Sierra Madre del Sur, in central Guerrero,
Order, Family: Anura, Hylidae and central and south-western Oaxaca, Mexico. It probably occurs at more localities than have been mapped within
Country Distribution: Mexico its general range.
Current Population Trend: Decreasing Population This is a rare species, and it appears to have undergone declines.
Society for the Study of Amphibians and Reptiles

Habitat and Ecology It occurs in cloud and pine-oak forests at 700-1,070m asl and is commonly found in shallow
water or on low vegetation along mountain streams. It breeds in streams.
© Hylid Frogs of Middle America, 2001,

Major Threats The high rate of habitat disturbance, due to agricultural activities within forested areas in Mexico,
and specifically in Oaxaca and Guerrero, is the main threat to this species. Recent declines might also be due to
chytridiomycosis.
Conservation Measures The range of this species does not include any protected areas, and improved protection
of the forests of south-western Oaxaca and the central highlands of Guerrero is urgently needed. Close population
monitoring is required, especially given the potential threat of chytridiomycosis. This species is protected by Mexican
law under the “Special Protection” category (Pr).
Notes on taxonomy: This species was previously included in the genus Hyla but has recently been moved to the resurrected genus
Exerodonta (Faivovich et al. 2005).
Bibliography: Campbell, J.A. and Brodie, E.D., Jr. (1992), Duellman, W.E. (2001), Faivovich, J. et al. (2005), Lips, K.R. et al. (2004)
Data Providers: Georgina Santos-Barrera, Luis Canseco-Márquez
248 Threatened Amphibians of the World

VU Exerodonta xera (Mendelson and Campbell, 1994)

Vulnerable B1ab(iii) Geographic Range This species is known from south-western Zapotitlán de las Salinas, central Puebla, and northern
Order, Family: Anura, Hylidae Oaxaca, Mexico. It might occur more widely than is currently known. It occurs at about 1,500m asl.
Country Distribution: Mexico Population It is a common species.
Current Population Trend: Decreasing Habitat and Ecology This stream-breeding amphibian inhabits temporal streams with rocks and arid scrub vegetation
that provide suitable microhabitats. It seeks refuge in bromeliads during the dry season.
Major Threats The major threat is the ongoing disturbance and loss of the species’ arid tropical scrub habitat due
to infrastructure development, partly due to the construction of tourist facilities.
Conservation Measures The range of this species includes Reserva de la Biósfera del Valle de Tehuacán-Cui-
catlán.
Notes on taxonomy: This species was previously included in the genus Hyla but has recently been moved to the resurrected genus
Exerodonta (Faivovich et al. 2005).
Bibliography: Canseco-Márquez, L., Gutiérrez-Mayén, G. and Mendelson, J.R. III (2003), Duellman, W.E. (2001), Faivovich, J. et al.
© Tim Burkhardt

(2005)
Data Providers: Georgina Santos-Barrera, Luis Canseco-Márquez

CR Hyla bocourti Mocquard, 1899

Critically Endangered A2ace Geographic Range This species is found in the highlands of Alta and Baja Verapaz, Guatemala, at 1,300-1,536m
Order, Family: Anura, Hylidae asl.
Country Distribution: Guatemala Population Campbell and Lawson (1992) report that the species can be abundant during the early part of the rainy
Current Population Trend: Decreasing season. Numerous recent visits to the range of this species have failed to turn up any individuals, suggesting a
Society for the Study of Amphibians and Reptiles

serious decline.
Habitat and Ecology It has been found in open, grassy meadows when they become flooded during the early part of
© Hylid Frogs of Middle America, 2001,

the rainy season. It has also been found beneath sheaths of banana plants and in a bromeliad. This species appears
to tolerate some level of habitat disturbance. Breeding takes place in temporary pools.
Major Threats A major threat to this species is the ornamental plants industry, which requires heavy use of
pesticides and fungicides because the cut flowers and ferns exported to first world countries have to be completely
free of insects or blemishes. In areas with many of these ornamental plant farms, the streams are typically heavily
contaminated. Another cause of the observed declines, which have taken place even in suitable habitat, could be
chytridiomycosis.
Conservation Measures This species probably occurs in the Biotopo del Quetzal. Surveys are needed to determine
the population status of this species, and the reasons for the observed decline. It is also necessary to ensure that
ornamental plant farms adhere to protocols to ensure the safe and proper disposal of pesticides.
Bibliography: Campbell, J.A. and Lawson, D.P. (1992), Duellman, W.E. (2001), Mendelson III, J.R. et al. (2004)
Data Providers: Manuel Acevedo, Bruce Young

CR Hyla chlorostea Reynolds and Foster, 1992

Critically Endangered B1ab(iii)+2ab(iii) Geographic Range This species occurs on the oriental slope of the Andes, and is known only from the type local-
Order, Family: Anura, Hylidae ity: “Parjacti, 83.2km by road north-eastern Cochabamba, on road to Villa Tunari, Chapare Province, Department of
Country Distribution: Bolivia Cochabamba, Bolivia, elevation approximately 2,044m” (Reynolds and Foster 1992).
Current Population Trend: Decreasing Population The population status of this species is not known, and it might be extinct. The areas surrounding the
type locality have been well surveyed in recent years, but the species has not been recorded again since its original
collection in the 1970s.
Habitat and Ecology It is an arboreal forest species, reported in the Yungas forest (Reynolds and Foster 1992; De
la Riva et al. 2000). There is no information on breeding, though it is likely to be a larval developer in water, like
other species of the genus.
Major Threats The quality of the species’ habitat continues to decline due to agriculture, logging, and infrastructure
© Ignacio De La Riva

development for human settlement.

Conservation Measures The species is not known from any protected areas. This species is a priority for urgent
survey work in order to determine whether or not it is still extant.
Notes on taxonomy: This species is considered incerta sedis, and is presently left in the genus Hyla pending resolution (Faivovich et
al. 2005). De la Riva et al. (2000) registered a slight doubt as to the taxonomic validity of the species.
Bibliography: De la Riva, I. et al. (2000), Duellman, W.E., De la Riva, I. and Wild, E. (1997), Faivovich, J. et al. (2005), Reynolds, R.
and Foster, M. (1992)
Data Providers: Claudia Cortez, Steffen Reichle, Ignacio De la Riva, Jörn Köhler

VU Hyla walkeri Stuart, 1954

Vulnerable B1ab(iii) Geographic Range This species occurs in the highlands of central Chiapas, Mexico, south-west to the Sierra de
Order, Family: Anura, Hylidae los Cuchumatanes, Guatemala. In Guatemala, it occurs on the central Plateaus and in the south-east (1,450-2,340m
Country Distribution: Guatemala, Mexico asl). It probably occurs more widely than current records suggest.
Current Population Trend: Decreasing Population It is uncommon in Guatemala and has not been seen since 1995. Extensive recent surveys in the last few
years indicate that this species is nearing extirpation in this country. It is still abundant in Chiapas.
Habitat and Ecology It occurs in pine-oak and pine-fir forests and high grasslands. Breeding takes place in
temporary ponds.
Major Threats Loss of habitat due to agriculture, logging, and human settlement is the major threat.
Conservation Measures It occurs in the Parque Nacional Lagunas de Montebello in Mexico, but is not recorded
from any protected areas in Guatemala.
Bibliography: Stuart, L.C. (1954)
© Sean M. Rovito

Data Providers: Georgina Santos-Barrera, Luis Canseco-Márquez, Manuel Acevedo, Antonio Muñoz Alonso
 Globally Threatened Amphibian Species 249

EN Hylomantis lemur (Boulenger, 1882)

Endangered A2ae Costa Rica the species is now only known with certainty from three sites: Fila Asuncion, 15km south-west of Limón
Order, Family: Anura, Hylidae (an abandoned farm); in a forested area near Parque Nacional Barbilla near Siquirres (where one female has been
Country Distribution: Colombia, Costa Rica, found); and from Guayacán (in Limón Province). The first of these three sites is the only site known to have a large
Panama breeding population. All other previously known Costa Rican populations of this species have disappeared including
Current Population Trend: Decreasing those in Monteverde, San Ramon, Braulio Carrillo, and Tapantí. Its altitudinal range is 440-1,600m asl.
Population It was once considered to be a reasonably common species in Costa Rica, but most populations have
recently disappeared. The species is still considered to be reasonably common in lower elevation parts of Panama
(where, for example, there are recent records from Palmarazo), but a decline has been recorded in the Reserva
Forestal Fortuna, Chiriquí, and there are no records from this site since 1999. There is no recent population informa-
tion from Colombia.
Habitat and Ecology It is a nocturnal treefrog associated with sloping areas in humid lowland and montane primary
forest, and is not found in degraded habitats. The eggs are usually deposited on leaf surfaces and the larvae are
washed off or fall into water below the site of oviposition.
Major Threats The massive declines noted in this species are probably due to chytridiomycosis. However, general
habitat loss remains a threat, and this is especially the case in Costa Rica where deforestation by squatters threatens
© Ron Holt, courtesy of Atlanta Botanical Garden

Fila Asunción, one of the three known remaining populations.

Conservation Measures Within Costa Rica, the former range included several national parks and other protected
areas; there may be a remaining population within Parque Nacional Barbilla but surveys are needed to confirm this.
Geographic Range This species occurs in The species is known to be present within at least six Panamanian protected areas, but it is not known from any
Costa Rica and Panama, and marginally in protected areas in Colombia. A successful captive breeding program began in 2001 at the Altanta Botanical Garden,
Colombia. It occurs predominantly on the which has since transferred individuals to other zoos to continue these captive breeding efforts.
Atlantic versant from the vicinity of Tiláran, Notes on taxonomy: This species was previously within the genus Phyllomedusa but has recently been moved to the genus Hylomantis
Guanacaste Province, Costa Rica, to western (Faivovich et al. 2005).
Panama; the disjunct Pacific slope records Bibliography: Cannatella, D.C. (1980), Duellman, W.E. (2001), Faivovich, J. et al. (2005), Ibáñez, R. et al. (2000), Jungfer, K.-H. and
are from north-western Costa Rica and Weygoldt, P. (1994), Myers, C.W. and Duellman, W.E. (1982), Pounds, J.A. et al. (1997), Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and
south-western, central, and extreme eastern Lynch, J.D. (1996), Savage, J.M. (2002), Young, B. et al. (1999), Zippel, K. (2005)
Panama in the Darien area, where it extends Data Providers: Frank Solís, Roberto Ibáñez, Jay Savage, César Jaramillo, Querube Fuenmayor, Brian Kubicki, Alan Pounds, Gerardo
marginally across the border into Colombia. In Chaves, Karl-Heinz Jungfer

EN Hyloscirtus charazani (Vellard, 1970)

Endangered B1ab(iii) Geographic Range This species is found on the eastern slopes of the Andes, and is known from the type locality in
Order, Family: Anura, Hylidae Charazani and the nearby Pomasani River, in Huasawaico, Saavradra Province, in the department of La Paz (Aparicio
Country Distribution: Bolivia 1999), Bolivia. There is an additional record from near Waleke, in Bautista Savedra Province, Charazani Canton. The
Current Population Trend: Decreasing elevation range is 2,700-3,200m asl.
Population It is abundant at the type locality.
Habitat and Ecology It is an aquatic and terrestrial species, inhabiting streams in the inter-Andean valleys (De la
Riva et al. 2000). It is a stream breeder, and the larva is morphologically indistinguishable from larvae of Hyloscirtus
armatus sensu lato, exhibiting stream adaptation through a robust body with strong tail musculature, enlarged oral
disc, increased number of labial tooth rows, and complete marginal papillae (Lötters et al. 2005).
Major Threats The major threat to this species is water pollution: the only known population occurs in a stream
© Ignacio De La Riva

used by the local communities living nearby (S. Reichle pers. comm.).
Conservation Measures The Área Natural de Manejo Integrado Nacional Apolobamba probably protects the popula-
tion, although the type locality is in a community land-use zone of the reserve. Broad-scale actions are necessary to
deal with the threat of water pollution.
Notes on taxonomy: This species was previously included in the genus Hyla but has recently been moved to the resurrected genus
Hyloscirtus (Faivovich et al. 2005). It was rediscovered by J. Aparicio and S. Reichle in 1997 at the type locality.
Bibliography: Aparicio, J. (1999), De la Riva, I. et al. (2000), Duellman, W.E., De la Riva, I. and Wild, E. (1997), Faivovich, J. et al. (2005),
Köhler, J. (2000a), Lötters, S. et al. (2005), Vellard, J. (1970)
Data Providers: Claudia Cortez, Steffen Reichle, Ignacio De la Riva, Jörn Köhler

EN Hyloscirtus colymba (Dunn, 1931)

Endangered A2ace Geographic Range This species is known from the Atlantic slopes of south-eastern Costa Rica to central Panama,
Order, Family: Anura, Hylidae from sea level to 1,116m asl, and also from the Pacific slopes of eastern Panama (Savage 2002).
Country Distribution: Costa Rica, Panama Population Little is known about the population size or abundance of this species. It has disappeared from the
Current Population Trend: Decreasing Reserva Forestal Fortuna, Chiriquí, Panama, but is still present at El Cope in Panama.
Habitat and Ecology It inhabits cloud forest or humid lowland and montane forest, where they have been found
along streams. The species breeds in swift streams; eggs are deposited under rocks, and larvae cling to rocks by
means of an oral disk (Savage 2002).
Major Threats The decline in the Reserva Forestal Fortuna, Panama, is probably due to chytridiomycosis, and this
is likely to be the most serious threat to the species. It is probably also impacted by the destruction of natural forests
for the planting of crops, smallholder livestock ranching, and logging.
© Ignacio De La Riva

Conservation Measures It is known from several protected areas in Panama, and a single park in Costa Rica
(Parque Internacional La Amistad). In view of the risk of chytridiomycosis, the status of this species should be closely
monitored, and ex-situ populations should be established.
Notes on taxonomy: This species was previously within the genus Hyla but has recently been moved to the resurrected genus
Hyloscirtus (Faivovich et al. 2005).
Bibliography: Duellman, W.E. (2001), Faivovich, J. et al. (2005), Ibáñez, R. et al. (2000), Lips, K.R. (1999), Savage, J.M. (2002), Young,
B. et al. (1999)
Data Providers: Frank Solís, Roberto Ibáñez, Gerardo Chaves, Jay Savage, César Jaramillo, Querube Fuenmayor
250 Threatened Amphibians of the World

EN Hyloscirtus denticulentus (Duellman, 1972)

Endangered B1ab(iii) Geographic Range This species occurs on the western slope of the Cordillera Oriental, in Boyacá and Santander
Order, Family: Anura, Hylidae Departments, Colombia, at 1,630-2,400m asl.
Country Distribution: Colombia Population It is known to be common, although there has been no survey work for this species since 1987.
Current Population Trend: Decreasing Habitat and Ecology It lives in streams in cloud forests, and also in secondary forest. The eggs are laid on vegeta-
tion, and the larvae develop in water.
Major Threats Habitat loss due to agriculture and cattle ranching, and predation by introduced trout, represent the
major threats to this species.
Conservation Measures It occurs in one protected area, the Santuario de Fauna y Flora Guanentá Alto Río Fonce,
where work has been under way by The Nature Conservancy in conjunction with Fundación Natura and other partners
to support the expansion of the existing reserve, as well as the the creation of a new public protected area.
Notes on taxonomy: This species was previously included in the genus Hyla but has recently been moved to the resurrected genus
© John D. Lynch

Hyloscirtus (Faivovich et al. 2005). Ruíz-Carranza, Ardila-Robayo and Lynch (1996) noted that one of the paratypes of this species is
referrable to Hyloscirtus palmeri.
Bibliography: Duellman, W.E. (1972), Faivovich, J. et al. (2005), Ruiz-Carranza, P.M. and Lynch, J.D. (1982), Ruiz-Carranza, P.M., Ardila-
Robayo, M.C. and Lynch, J.D. (1996)
Data Providers: María Cristina Ardila-Robayo, Jose Vicente Rueda

VU Hyloscirtus lindae (Duellman and Altig, 1978)

Vulnerable B1ab(iii) Geographic Range This species occurs on the Amazonian slopes of the Andes in southern Colombia (in Caqueta
Order, Family: Anura, Hylidae and Putumayo Departments) and Ecuador (south to Morona Province). It ranges from 2,000-2,500m asl.
© Natural History Museum and Biodiversity Research

Country Distribution: Colombia, Ecuador Population It is a common species.

Current Population Trend: Decreasing Habitat and Ecology It lives in upper humid montane forest, and it also survives in pastureland and other altered
habitats. However, although it is adaptable, it probably cannot tolerate extremely severe habitat clearance, leading
to a very open landscape. It is associated with creeks and breeds in streams.
Major Threats The major threats are habitat loss from agricultural development, planting of illegal crops, logging,
and human settlement, and pollution resulting from the spraying of illegal crops. The species has a narrow altitudinal
range, and lives in habitats where catastrophic extinctions have occurred in other frog species with stream-dwelling
Center, University of Kansas

tadpoles, probably due to chytridiomycosis.

Conservation Measures It occurs in several protected areas in Ecuador, including Parque Nacional Llanganates and
Parque Nacional Sangay, but is apparently not recorded from any in Colombia. There is a need for close population
monitoring of this species, given the potential threat of chytridiomycosis.
Notes on taxonomy: This species was previously included in the genus Hyla but has recently been moved to the resurrected genus
Hyloscirtus (Faivovich et al. 2005).
Bibliography: Duellman, W.E. and Altig, R. (1978), Duellman, W.E. and Hillis, D.M. (1990), Faivovich, J. et al. (2005), Ruiz-Carranza,
P.M. and Lynch, J.D. (1982), Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and Lynch, J.D. (1996)
Data Providers: Diego Almeida, Wilmar Bolívar, Luis A. Coloma, Santiago Ron

EN Hyloscirtus lynchi (Ruíz-Carranza and Ardila-Robayo, 1991)

Endangered B1ab(iii) Geographic Range This species occurs on the western slope of the Cordillera Oriental in the department of Santander,
Order, Family: Anura, Hylidae Colombia, at 2,540-2,700m asl. It may be more widely distributed than existing records indicate.
Country Distribution: Colombia Population It is known to be common, although there has been no survey work for this species since 1998.
Current Population Trend: Decreasing Habitat and Ecology It lives in streams in cloud forest, but only in old growth forest. The eggs are laid on vegeta-
tion, and the tadpoles larvae in streams.
Major Threats Habitat loss due to agriculture and cattle ranching, and predation by introduced trout, are the major
threats to this species.
Conservation Measures It does not occur in any protected areas, and there is an urgent need for expanded protec-
tion of remaining forest habitats in the Colombian Andes.
Notes on taxonomy: This species was previously within the genus Hyla but has recently been moved to the resurrected genus
Hyloscirtus (Faivovich et al. 2005).
Bibliography: Faivovich, J. et al. (2005), Ruiz-Carranza, P.M. and Ardila-Robayo, M.C. (1991), Ruiz-Carranza, P.M., Ardila-Robayo, M.C.
and Lynch, J.D. (1996)
Data Providers: María Cristina Ardila-Robayo, Jose Vicente Rueda
© John D. Lynch

EN Hyloscirtus pantostictus (Duellman and Berger, 1982)

Endangered B2ab(iii) Geographic Range This species is known only from the Nudo de Pasto area on the eastern slopes of the Andes
Order, Family: Anura, Hylidae in southern Colombia and northern Ecuador. In Ecuador, it has been recorded only in the extreme north-eastern
Country Distribution: Colombia, Ecuador páramos around Sánta Barbara on the Cordillera Oriental on the border with Colombia. It has been recorded at
Current Population Trend: Decreasing about 1,950-2,700m asl.
Population It is a rare species.
Habitat and Ecology A cloud forest species, it has also been found in partially cleared forest, where frogs were
found at night on branches of trees above a stream. However, its adaptability to disturbed habitats is not well known.
It breeds in streams.
Major Threats The major threats are likely to be deforestation due to agricultural development, planting of illegal
crops, logging, and human settlement, and pollution resulting from the spraying of illegal crops. All known localities in
Ecuador are subject to severe human disturbance. The species lives in montane habitats where catastrophic declines
© Luis A. Coloma

have affected other frogs with stream-dwelling larvae, probably as a result of chytridiomycosis.
Conservation Measures It is not known from any protected areas, and there is an urgent need to expand the
protected area coverage in the Colombian Andes. Research is needed to determine the current population status of
this species and to ascertain whether chytrid poses a threat.
Notes on taxonomy: This species was previously within the genus Hyla but has recently been moved to the resurrected genus
Hyloscirtus (Faivovich et al. 2005).
Bibliography: Duellman, W.E. and Berger, T.J. (1982), Duellman, W.E. and Hillis, D.M. (1990), Faivovich, J. et al. (2005), Ruiz-Carranza,
P.M. and Lynch, J.D. (1982), Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and Lynch, J.D. (1996)
Data Providers: Wilmar Bolívar, Luis A. Coloma, Santiago Ron, Diego Cisneros-Heredia
 Globally Threatened Amphibian Species 251

EN Hyloscirtus piceigularis (Ruíz-Carranza and Lynch, 1982)

Endangered B1ab(iii) Geographic Range This species occurs on the western flank of Notes on taxonomy: This species was previously included in the genus Hyla, but has recently been moved to the resurrected genus
Order, Family: Anura, Hylidae the Cordillera Oriental in the departments of Cundinamarca and Hyloscirtus (Faivovich et al. 2005).
Country Distribution: Colombia Santander, Colombia, at 1,750-2,000m asl. Bibliography: Faivovich, J. et al. (2005), Ruiz-Carranza, P.M. and Lynch, J.D. (1982), Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and
Current Population Trend: Decreasing Population It was common in Virolín in 1987; however, there has Lynch, J.D. (1996)
been no survey work for the species since. Data Providers: María Cristina Ardila-Robayo, Jose Vicente Rueda
Habitat and Ecology It lives in streams in old growth cloud forest. It
lays its eggs on vegetation, and the larvae develop in the water.
Major Threats Habitat loss due to agriculture and cattle ranch-
ing, and predation by introduced trout, are the major threats to
this species.
Conservation Measures It occurs in the Santuario de Fauna y Flora
Guanentá Alto Río Fonce. Continued and strengthened management
of this reserve, and expanded protection of cloud forest elsewhere
in the range of this species is needed.

VU Hyloscirtus platydactylus (Boulenger, 1905)

Vulnerable A2ace; B1ab(iii,v) Geographic Range This Andean species occurs in the Cordillera de Merida in the states of Barinas, Merida and
Order, Family: Anura, Hylidae Tachira, and from Sierra de Perijá in Zulia State, Venezuela, at 1,600-3,000m asl. It is also known from the eastern
© César Luis Barrio-Amorós / Fundación AndígenA

Country Distribution: Colombia, Venezuela slopes of Cordillera Oriental in Colombia in the department of Norte de Santander, at 1,050m asl. Records from Sierra
Current Population Trend: Decreasing del Turimiquire in Sucre and Monagas states are in error.
Population Formerly an abundant species, there are few recent records, in part due to lack of survey efforts in
its range. However, some populations, such as those north of the city of Mérida on Monte Zerpa, Venezuela, have
experienced declines in suitable habitat (E. La Marca pers. obs.). However, in 2001 it was reportedly found without
difficulty in Venezuela in brooks of cloud forest (Barrio Amorós 2001).
Habitat and Ecology It lives in cloud forests, and is able to tolerate minor habitat disturbance. It is usually found
along streams, where it breeds.
Major Threats The main threat is deforestation, especially the conversion of cloud forests to pasture lands. However,
declines that have taken place in suitable habitat suggest another threatening process might be operating, possibly
chytridiomycosis.
Conservation Measures It occurs in the Sierra Nevada and Sierra de la Culata National Parks in Venezuela, and
probably also occurs in other protected areas. Close population monitoring of this species is required, particularly if
chytrid is shown to be a genuine threat.
Notes on taxonomy: This species was previously included in the genus Hyla but has recently been moved to the resurrected genus
Hyloscirtus (Faivovich et al. 2005). It is possibly confused with Hyloscirtus bogotensis.
Bibliography: Barrio Amorós, C.L. (2001), Barrio Amorós, C.L. (2004), Duellman, W.E. (1972), Faivovich, J. et al. (2005), La Marca, E. (1985b),
La Marca, E. (1992), La Marca, E. (1994c), La Marca, E. (1995b), La Marca, E. (1997), Péfaur, J.E. and Rivero, J.A. (2000), Ruiz-Carranza,
P.M. and Lynch, J.D. (1982), Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and Lynch, J.D. (1996), Vial, J.L. and Saylor, L. (1993)
Data Providers: Enrique La Marca, Juan Elías García-Pérez

EN Hyloscirtus psarolaimus (Duellman and Hillis, 1990)

Endangered B1ab(iii) Geographic Range This Andean species from Colombia and Ecuador occurs on the Amazonian slopes of the Cordil-
Order, Family: Anura, Hylidae lera Oriental from Putumayo Department in southern Colombia south to Morona Province in southern Ecuador. It lives
Country Distribution: Colombia, Ecuador at elevations of 1,950-2,660m asl.
Current Population Trend: Decreasing Population There is very little information, since there are only a few records of this species.
Habitat and Ecology It is a species of cloud forest. Specimens have been found on stems and branches of bushes
near, or over, streams at night. There is no information on its adaptability to secondary habitats. It presumably
breeds in streams.
Major Threats The major threats are likely to be deforestation due to agricultural development, the planting of
illegal crops, logging, and human settlement, and pollution resulting from the spraying of illegal crops. The species
also lives in montane habitats where catastrophic declines have affected other frogs with stream-dwelling larvae,
probably as a result of chytridiomycosis.
© Luis A. Coloma

Conservation Measures In Ecuador, its geographic range overlaps with the Reserva Ecológica Cayambe-Coca, the
Reserva Ecológica Antisana, Parque Nacional Llanganates, and Parque Nacional Sangay. It is not known from any
protected areas in Colombia. Further research is necessary to determine the current population status of this species
in the wild, and whether chytrid poses a threat.
Notes on taxonomy: This species was previously included in the genus Hyla but has recently been moved to the resurrected genus
Hyloscirtus (Faivovich et al. 2005).
Bibliography: Duellman, W.E. and Altig, R. (1978), Duellman, W.E. and Hillis, D.M. (1990), Faivovich, J. et al. (2005), Ruiz-Carranza,
P.M. and Ardila-Robayo, M.C. (1991), Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and Lynch, J.D. (1996)
Data Providers: Wilmar Bolívar, Luis A. Coloma, Santiago Ron

CR Hyloscirtus ptychodactylus (Duellman and Hillis, 1990)

Critically Endangered B2ab(iii) Geographic Range This species is only known from Pilaló at 2,320m protected areas, and its remaining intact habitat is in urgent need of protection. Survey work is urgently required to
Order, Family: Anura, Hylidae in Cotopaxi Provice, and nearby areas in Ecuador. determine the current population status of the species.
Country Distribution: Ecuador Population Recent efforts to locate the species at the type local- Notes on taxonomy: This species was previously included in the genus Hyla but has recently been moved to the resurrected genus
Current Population Trend: Decreasing ity were unsuccessful, but more surveys are needed before it can Hyloscirtus (Faivovich et al. 2005).
be concluded whether or not this species has declined in suitable Bibliography: Duellman, W.E. and Hillis, D.M. (1990), Faivovich, J. et al. (2005)
habitats. Data Providers: Luis A. Coloma, Santiago Ron
Habitat and Ecology It has been found only in disturbed cloud
forest, where adults have been found by day. There is no informa-
tion on its ability to survive in heavily degraded habitats. Breeding
takes place in streams.
Major Threats At the type locality, patches of forest have been
cleared for agriculture and livestock farming, possibly posing a seri-
ous threat to the survival of the species. The species lives in habitats
where catastrophic extinctions have affected other frogs with stream-
dwelling tadpoles, probably as a result of chytridiomycosis.
Conservation Measures This species is not known from any
252 Threatened Amphibians of the World

EN Hyloscirtus simmonsi (Duellman, 1989)

Endangered B1ab(iii) Geographic Range This species is known from Antioquia to Valle del Cauca Departments, on the western flank of
Order, Family: Anura, Hylidae the Cordillera Occidental, in Colombia, from 1,100-2,000m asl.

© Natural History Museum and Biodiversity Research

Country Distribution: Colombia Population It is a common species.
Current Population Trend: Decreasing Habitat and Ecology It is found on low vegetation alongside streams in primary forest and secondary forest. It can
tolerate some habitat destruction as long as vegetation remains alongside the stream. Eggs are laid on the top of
leaves above the stream and when hatched the larvae drop into the stream below where they then develop further.
Major Threats The major threat to this species is habitat loss and degradation due to smallholder farming and
subsistence logging, and water pollution from the spraying of illegal crops.
Conservation Measures The range of the species includes several protected areas. Taxonomic research is needed
Center, University of Kansas to determine whether or not this represents a complex of species.
Notes on taxonomy: This species was previously included in the genus Hyla but has recently been moved to the resurrected genus
Hyloscirtus (Faivovich et al. 2005). This form might represent a species complex.
Bibliography: Duellman, W.E. (1989b), Faivovich, J. et al. (2005), Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and Lynch, J.D. (1996)
Data Providers: Fernando Castro, John Lynch

EN Hyloscirtus staufferorum (Duellman and Coloma, 1993)

Endangered B1ab(iii) Geographic Range This species is known only from two localities in Napo Province, Ecuador: Volcán Sumaco, and
Order, Family: Anura, Hylidae the Cordillera de Guacamayos. It probably occurs a little more widely. Its altitudinal range is 2,040-2,500m asl.
Country Distribution: Ecuador Population There is no information on the population status of this species.
Current Population Trend: Decreasing Habitat and Ecology It lives in very humid, low cloud forest, and breeds in streams. There is no information on its
adaptability to secondary habitats.
Major Threats The main threat is forest clearance due to agriculture, livestock farming, and wood extraction. It lives
in montane habitats where catastrophic declines have affected other frogs with stream-dwelling larvae, probably
as a result of chytridiomycosis.
Conservation Measures The geographic range of this species overlaps with Parque Nacional Sumaco. Continued
and strengthened management of this reserve, and expanded protection of cloud forest elsewhere in the range of
this species, are needed. Further survey work is necessary to determine its current population status, and whether
© Luis A. Coloma

chytrid poses a threat.

Notes on taxonomy: This species was previously included in the genus Hyla but has recently been moved to the resurrected genus
Hyloscirtus (Faivovich et al. 2005).
Bibliography: Duellman, W.E. and Coloma, L.A. (1993), Faivovich, J. et al. (2005)
Data Providers: Luis A. Coloma, Santiago Ron

VU Hyloscirtus torrenticola (Duellman and Altig, 1978)

Vulnerable B1ab(iii) Geographic Range This species occurs on the eastern slopes of the Andes in Ecuador (two localities: La Barquilla
Order, Family: Anura, Hylidae and El Reventador in Sucumbios Province and Napo Province) and Colombia (eastern slopes of the Cordillera Oriental
Country Distribution: Colombia, Ecuador in the departments of Caqueta and Putumayo). It might be more widespread than records suggest. Its altitudinal
© Natural History Museum and Biodiversity Research

Current Population Trend: Decreasing range is 740-1,700m asl.

Population It is a moderately common species.
Habitat and Ecology It inhabits cloud forests, as well as secondary forest. It is a stream-breeding species, particularly
associated with creeks, streams and swampy areas in forest, especially with vegetation overhanging streams.
Major Threats The major threats are habitat loss from agricultural development, planting of illegal crops, logging, and
human settlement, and pollution resulting from the spraying of illegal crops. The species lives in habitats where cata-
Center, University of Kansas

strophic extinctions have affected other frogs with stream-dwelling tadpoles, probably due to chytridiomycosis.
Conservation Measures It might be found in the Reserva Ecológica Cayambe-Coca in Ecuador, and might also occur
on Parque Nacional Natural Alto Fragua-Indi Wasi in Colombia. There is a need for close population monitoring of
this species, given the potential threat of chytridiomycosis.
Notes on taxonomy: This species was previously included in the genus Hyla but has recently been moved to the resurrected genus
Hyloscirtus (Faivovich et al. 2005).
Bibliography: Duellman, W.E. and Altig, R. (1978), Duellman, W.E. and Hillis, D.M. (1987), Faivovich, J. et al. (2005), Ruiz-Carranza,
P.M. and Lynch, J.D. (1982), Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and Lynch, J.D. (1996)
Data Providers: Mario Yánez-Muñoz, Wilmar Bolívar, Luis A. Coloma, Santiago Ron, Juan Manuel Renjifo

CR Hypsiboas cymbalum (Bokermann, 1963)

Critically Endangered B1ab(iii)+2ab(iii) Geographic Range This species is known from the type locality, Notes on taxonomy: This species was previously included in the genus Hyla but has recently been moved to the resurrected genus
Order, Family: Anura, Hylidae Campo Grande da Serra, and nearby Nova Manchester in São Paulo, Hypsiboas (Faivovich et al. 2005).
Country Distribution: Brazil in the state of São Paulo, Brazil. The type locality is between 600 Bibliography: Bokermann, W.C.A. (1963), Faivovich, J. et al. (2005), Garcia, P.C.A., Vinciprova, G. and Haddad, C.F.B. (2003), Lutz, B.
Current Population Trend: Decreasing and 800m asl. (1973a)
Population This species is only known from a few specimens, and Data Providers: Paulo Garcia, Miguel Trefaut Rodrigues
recent surveys in the area have not recorded it.
Habitat and Ecology Specimens were found on shrubs by the side
of a brook at the type locality. There is no information on breeding
habits, but reproduction might take place in streams.
Major Threats Habitat loss due to infrastructure development has
destroyed the species’ habitat at the type locality. Chytridiomycosis
also cannot be ruled out as a threat.
Conservation Measures This species does not occur within a pro-
tected area, and protection of any remaining habitat for the species
is urgently needed. Further survey work is required to determine the
biology and population status of this species, and whether or not it
still survives at, or outside the vicinity of, the type locality.
 Globally Threatened Amphibian Species 253

VU Hypsiboas heilprini (Noble, 1923)

Vulnerable B2ab(iii) Geographic Range This species occurs throughout much of Hispaniola, from sea level up to 1,856m asl, although
Order, Family: Anura, Hylidae populations are extremely fragmented.
Country Distribution: Dominican Republic, Haiti Population This species is believed to have undergone a decline in recent years, and during the course of recent
Current Population Trend: Decreasing

© Marcelino Hernández / Miguel A. Landestoy

surveys in Haiti was found to be absent from a number of streams (B. Hedges pers. comm.). In the Cordillera Central
of the Dominican Republic, it appears to persist in appropriate habitat at many localities (based on field observations
from 1998-2000; M. Hernandez pers. comm.).
Habitat and Ecology It is a stream-breeding amphibian found only in fast-flowing mountain streams in mesic
broadleaf forests. Males call from rocks or low vegetation near and in water.
Major Threats Due to the fact that it is a mountain stream-dweller, it is more sensitive to habitat destruction than
many other species. In Haiti, severe degradation of streams has already significantly altered its breeding habitat.
Mining is a threat in some areas in the Dominican Republic as is infrastructure development.
Conservation Measures It is known from several protected areas, including Reserva Científica Natural de Valle
Nuevo, although most are in need of improved management.
Notes on taxonomy: This species was previously included in the genus Hyla but has recently been moved to the resurrected genus
Hypsiboas (Faivovich et al. 2005). The populations on the Tiburon Peninsula probably represent an undescribed species (R. Thomas
pers. comm.).
Bibliography: Faivovich, J. et al. (2005), Hedges, S.B. (1993), Hedges, S.B. (1999), Hedges, S.B. (2001), Henderson, R.W. and Powell, R
(1999), Henderson, R.W. and Powell, R. (2001), Schwartz, A. and Henderson, R.W. (1991)
Data Providers: Blair Hedges, Sixto Inchaustegui, Marcelino Hernandez, Robert Powell

CR Isthmohyla angustilineata (Taylor, 1952)

Critically Endangered A2ae Population This is generally an uncommon species. The population is decreasing, but threats to the species remain
Order, Family: Anura, Hylidae unknown. In Costa Rica, the species has disappeared from Cerro Chompipe and Tapanti. It declined drastically in
Country Distribution: Costa Rica, Panama Monteverde, but a few individuals still hung on at least until the late 1990s, and the particular site where they were
Current Population Trend: Decreasing found has not been visited since; there is a chance that it still survives at Monteverde. In Panama there is a scarcity
of information on its distribution and abundance, though it appears to have declined, and the most recent record is
from Parque Internacional La Amistad, in Chiriquí Province, in 1991.
Habitat and Ecology It is a nocturnal treefrog of humid lower montane rainforest. Breeding usually takes place in
small puddles and water-filled depressions. Males call from leaves and twigs above the forest flood. Tadpoles are
benthic (Savage 2002).
Major Threats The reasons for the population decline of this species are not known with certainty, though it is likely
to be due to chytridiomycosis, perhaps in association with climate change.
Conservation Measures The current known range of this species falls entirely within protected areas (Parque
© Kenji Nishida

Geographic Range This species occurs in the Cordillera de Tilarán, Cordillera Central, and Cordillera de Talamanca
of Costa Rica and western Panama, from 1,500-2,040m asl (Savage 2002).
Internacional La Amistad, Panama, and at least three protected areas in Costa Rica). Further survey work is required
to determine the population status and trends of this species. In view of the threat of chytridiomycosis, ex-situ
populations might need to be established.
Notes on taxonomy: This species was previously included in the genus Hyla but has recently been moved to the new genus Isthmohyla
(Faivovich et al. 2005).
Bibliography: Arosemena, F.A. and Ibanez, D.R. (1991), Arosemena, F.A. and Ibanez, D.R. (1991), Duellman, W.E. (2001), Faivovich, J.
et al. (2005), Ibáñez, R. et al. (2000), Pounds, J.A. et al. (1997), Savage, J.M. (2002), Young, B. et al. (1999)
Data Providers: Frank Solís, Roberto Ibáñez, Alan Pounds, Federico Bolaños, Gerardo Chaves

CR Isthmohyla calypsa (Lips, 1996)

Critically Endangered A2ace Geographic Range This species is known from the southern Cordillera de Talamanca, on Cerro Pando on the
Order, Family: Anura, Hylidae Pacific slope in Costa Rica and Atlantic versant in Panama, and on the Pacific slope in south-western Panama, from
Country Distribution: Costa Rica, Panama 1,810-1,920m asl (Savage 2002).
Current Population Trend: Decreasing Population Extensive monitoring has shown that this species has disappeared from its former range. It was formerly
locally common in Tablas, Costa Rica, but has disappeared from all known sites since the early 1990s. There is no
information on the population size or abundance of this species in Panama, though this population has probably
also declined seriously.
Habitat and Ecology It is found along torrential stream courses in primary humid lower montane forest throughout the
year. Males are strongly territorial, with small home ranges. Eggs (clutches of 10-36) are deposited on leaf surfaces of
low vegetation above streams; hatching tadpoles fall or are washed by rain into the stream below (Savage 2002).
© Ignacio De La Riva

Major Threats The observed population decline is likely to be due to chytridiomycosis, since it has taken place within
pristine habitats. Habitat loss as a result of smallholder livestock farming is also a threat.
Conservation Measures This species has been recorded in Parque Internacional La Amistad and might occur in
Parque Nacional Volcán Barú. Further survey work is required to determine the population status and trends of this
species. In view of the threat of chytridiomycosis, ex-situ populations might need to be established.
Notes on taxonomy: This species was previously included in the genus Hyla but has recently been moved to the new genus Isthmohyla
(Faivovich et al. 2005).
Bibliography: Duellman, W.E. (2001), Faivovich, J. et al. (2005), Ibáñez, R. et al. (2000), Lips, K.R. (1996), Lips, K.R. (1998), Savage,
J.M. (2002), Young, B. et al. (1999)
Data Providers: Federico Bolaños, Gerardo Chaves, Frank Solís, Roberto Ibáñez, Jay Savage, César Jaramillo, Querube Fuenmayor

CR Isthmohyla debilis (Taylor, 1952)

Critically Endangered A2ace (at 1,200-1,400m asl) (Savage 2002).
Order, Family: Anura, Hylidae Population It is now a rare species (Savage 2002). The Costa Rican population has declined dramatically, and it
Country Distribution: Costa Rica, Panama might now be extinct in this country. A dramatic decline was documented in the Reserva Forestal Fortuna, Chiriquí,
Current Population Trend: Decreasing in Panama, and the most recent record from this country is from the Bosque Protector Palo Seco, on the highlands
of Bocas del Toro, in 1998.
Habitat and Ecology An arboreal species of the lower reaches of cloud forest. It is associated with low dense
vegetation overhanging small montane streams (less than 1m above the water). Tadpoles develop in these streams,
adhering to rocks (Savage 2002).
Major Threats The decline in Costa Rica and in the Reserva Forestal Fortuna, Panama, is probably due to chytrid-
iomycosis, and this is likely to be the most serious threat to the species. It is probably also impacted by habitat loss
© Ignacio De La Riva

as a result of agriculture, logging, and human settlement.

Geographic Range This species is known from the Atlantic slopes of the Cordillera Central and the Cordillera de
Talamanca in Costa Rica and western Panama (at 910-1,450m asl) and the Pacific slope of south-western Panama
Conservation Measures This species has been recorded from three protected areas in Panama, and two in Costa
Rica. Further survey work is required to determine the population status of this species. In view of the threat of
chytridiomycosis, ex-situ populations might need to be established.
Notes on taxonomy: This species was previously included in the genus Hyla but has recently been moved to the new genus Isthmohyla
(Faivovich et al. 2005).
Bibliography: Duellman, W.E. (1970), Duellman, W.E. (2001), Faivovich, J. et al. (2005), Hofer, U. and Bersier, L.-F. (2001), Ibáñez, R. et
al. (2000), Lips, K.R. (1999), Savage, J.M. (2002), Young, B. et al. (1999)
Data Providers: Frank Solís, Roberto Ibáñez, Gerardo Chaves, Jay Savage, César Jaramillo, Querube Fuenmayor
254 Threatened Amphibians of the World

CR Isthmohyla graceae (Myers and Duellman, 1982)

Critically Endangered A2ace Geographic Range This species is endemic to the western Cordillera Central of Panama at elevations of 1,120-
Order, Family: Anura, Hylidae 1,650m asl on both sides of the continental divide in the western part of the Serranía de Tabasará and on the Pacific
Country Distribution: Panama slopes of the eastern part of the Cordillera de Talamanca.
Current Population Trend: Decreasing Population The population of this species has declined seriously. It is not certain that any populations now survive,
and the species could be extinct.
Habitat and Ecology It is an arboreal species that inhabits humid montane forest, small ponds, puddles, and low-
gradient streams. Breeding occurs in these aquatic habitats.
Major Threats Infection by the chytrid pathogen was recorded by Lipps (1999) at the Reserva Forestal Fortuna,
in Chiriquí, and this is probably the most serious threat to the species. It is probably also impacted by habitat loss
© Karl-Heinz Jungfer caused by the destruction of natural forests.
Conservation Measures It is recorded as having been present in a number of protected areas including: Parque
Nacional Santa Fe, Parque Internacional La Amistad, and the Reserva Forestal Fortuna. This species is an urgent priority
for survey work, in order to determine whether or not this species still survives in the wild. In view of the threat of
chytridiomycosis, surviving individuals might need to form the basis for the establishment of an ex-situ population.
Notes on taxonomy: This species was previously included in the genus Hyla but has recently been moved to the new genus Isthmohyla
(Faivovich et al. 2005).
Bibliography: Duellman, W.E. (2001), Faivovich, J. et al. (2005), Ibáñez, R. et al. (2000), Lips, K.R. (1999), Myers, C.W. and Duellman,
W.E. (1982), Young, B. et al. (1999)
Data Providers: Frank Solís, Roberto Ibáñez, César Jaramillo, Querube Fuenmayor, Karen Lips

CR Isthmohyla insolita (McCranie, Wilson and Williams, 1993)

Critically Endangered B1ab(iii)+2ab(iii) Geographic Range This species is found in the extreme west of the Sierra Nombre de Dios, La Fortuna village,
Order, Family: Anura, Hylidae between the Departamentos of Yoro and Atlantica, north-central Honduras. The type locality is at 1,550m asl.
Country Distribution: Honduras Population This species is moderately common within its very restricted range.
Current Population Trend: Decreasing Habitat and Ecology This species is found on the ground and on low vegetation along streams, in lower montane
Society for the Study of Amphibians and Reptiles

wet forest. It lays its eggs in patches of moss in trees overhanging streams; when the eggs hatch the tadpoles fall
in to the stream.
© Hylid Frogs of Middle America, 2001,

Major Threats The major threat to this species is habitat loss due to agriculture and logging. This frog is very
dependent on pristine habitat as it depends on sufficient humidity for moss for breeding in the trees.
Conservation Measures The range of this species is known to be within Texiguat Wildlife Refuge. Maintenance
of existing tracts of remaining habitat is an important conservation measure for this species.
Notes on taxonomy: This species was previously included in the genus Hyla but has recently been moved to the new genus Isthmohyla
(Faivovich et al. 2005).
Bibliography: Faivovich, J. et al. (2005), McCranie, J.R. and Wilson, L.D. (2002b), McCranie, J.R., Wilson, L.D. and Williams, K.L. (1993c),
Wilson, L.D., McCranie, J.R. and Williams, K.L. (1994)
Data Providers: Gustavo Cruz, Larry David Wilson, Franklin Casteñeda

EN Isthmohyla picadoi (Dunn, 1937)

Endangered B1ab(iii) Geographic Range This species is known from three disjunct populations in the Cordillera Central and the Cordillera
Order, Family: Anura, Hylidae de Talamanca of Costa Rica and western Panama, from 1,920-2,770m asl (Savage 2002).
Country Distribution: Costa Rica, Panama Population The available evidence suggests that the Costa Rican populations are stable, and it is common at some
Current Population Trend: Decreasing sites in Panama. The species is difficult to observe and is mostly recorded by its call.
Habitat and Ecology It inhabits the canopy of humid upper montane forest, and is associated with the bromeliad
flora of this habitat. The larvae develop in bromeliad pools, and hence this is a phytotelmic species.
Major Threats The major threat is forest loss due to agricultural expansion, logging for timber, and human settle-
ment.
Conservation Measures It has been recorded from several protected areas, including: Parque Nacional Volcán Barú,
Costa Rica; Parque Internacional La Amistad, Panama; and the Reserva Forestal Fortuna, Panama.
Notes on taxonomy: This species was previously within the genus Hyla but has recently been moved to the new genus Isthmohyla
(Faivovich et al. 2005).
Bibliography: Duellman, W.E. (2001), Faivovich, J. et al. (2005), Ibáñez, R. et al. (2000), Lips, K.R. (1998), Savage, J.M. (2002), Young,
B. et al. (1999)
Data Providers: Frank Solís, Roberto Ibáñez, Gerardo Chaves, Jay Savage, César Jaramillo, Querube Fuenmayor
© Eric Lindquist

VU Isthmohyla pictipes (Cope, 1875)

Vulnerable B1ab(iii,v) Geographic Range This species occurs in the mountains of the Cordillera Central and Talamanca in Costa Rica, at
Order, Family: Anura, Hylidae 1,930-2,800m asl. It might occur slightly more widely.
Country Distribution: Costa Rica Population Recent searches in Cerro Chompipe, a known historical locality that has not suffered much habitat
Current Population Trend: Decreasing modification, have not found this species, although it continues to be seen at Cerro de la Muerte and several other
localities within its range.
Habitat and Ecology A nocturnal stream-breeding treefrog occurring in fast-flowing streams in lower montane and
montane rainforests. Males call from rocks in streams and along stream margins, and sometimes from low vegetation
near streams. Tadpoles typically are found adhering to large rocks in cold mountain streams (Savage 2002).
Major Threats Major threats include habitat destruction, pollution, and climate change. The decline witnessed at
Cerro Chompipe could be due to chytridiomycosis.
© Karl-Heinz Jungfer

Conservation Measures Much of the species’ range is included within national parks, including Parque Inter-
nacional La Amistad. There is a need for close population monitoring of this species, given the potential threat of
chytridiomycosis.
Notes on taxonomy: This species was previously included in the genus Hyla but has recently been moved to the new genus Isthmohyla
(Faivovich et al. 2005).
Bibliography: Duellman, W.E. (2001), Faivovich, J. et al. (2005), Savage, J.M. (2002), Savage, J.M. and Wake, M.H. (2001)
Data Providers: Federico Bolaños, Gerardo Chaves, Jay Savage
 Globally Threatened Amphibian Species 255

CR Isthmohyla rivularis (Taylor, 1952)

Critically Endangered A2ace Population In Costa Rica, it has disappeared from Monteverde, Tapantí, and Las Tablas where it once was common. It
Order, Family: Anura, Hylidae was last seen in 1993 at Las Tablas and had disappeared from Monteverde by 1989. There have been no other recent
Country Distribution: Costa Rica, Panama records from Costa Rica of this formerly common frog. In Panama, there are records from the Bajo Mono highlands
Current Population Trend: Decreasing of Chiriquí in 1982, and from Las Tablas in the early 1990s, but it had disappeared from the latter site by 1996. There
might not be any subsequent records, indicating a decline and possible disappearance in this country, too.
Habitat and Ecology This species occurs along or in clear streams in lower and premontane rainforest. Males call at
night from riparian bushes and herbaceous vegetation at the margin of or overhanging fast-moving mountain streams.
Amplexus and egg deposition have not been observed in this species; tadpoles have been collected from streams.
Major Threats The overall decline in this species is probably taking place as a result of infection of populations
with the chytrid pathogen. Within Panama, it is threatened by general habitat loss through agriculture and selective
logging.
Conservation Measures The species has been recorded from a number of protected areas in both Costa Rica
© Andrew Gray

Geographic Range This species occurs on the slopes of the cordilleras (Tilarán, Central and Talamanca) of Costa
Rica and adjacent western Panama, from 1,210-2,040m asl (Savage 2002).
and Panama. Further research is urgently needed into the population status of this species. Given the threat of
chytridiomycosis, recommended conservation measures likely should include the establishment of a captive-breed-
ing programme.
Notes on taxonomy: This species was previously included in the genus Hyla but has recently been moved to the new genus Isthmohyla
(Faivovich et al. 2005).
Bibliography: Duellman, W.E. (1970), Duellman, W.E. (2001), Faivovich, J. et al. (2005), Ibáñez, R. et al. (2000), Lips, K.R. (1998), Myers,
C.W. and Duellman, W.E. (1982), Pounds, J.A. et al. (1997), Savage, J.M. (2002), Young, B. et al. (1999)
Data Providers: Frank Solís, Roberto Ibáñez, Alan Pounds, Federico Bolaños, Gerardo Chaves, Karen Lips

CR Isthmohyla tica (Starrett, 1966)

Critically Endangered A2ace Population In Costa Rica, it has disappeared from Monteverde, Tapantí, and Las Tablas where it once was common.
Order, Family: Anura, Hylidae In Panama, there were records from the Cerro Horqueta highlands of Chiriquí in 1982. It used to be seen regularly
Country Distribution: Costa Rica, Panama at the Reserva Forestal Fortuna, Chiriquí, in Panama, but its population collapsed in January 1997, and it has not
Current Population Trend: Decreasing been seen there since. This is possibly the last record from Panama, and it seems that it might have disappeared
from this country as well.
Habitat and Ecology It is a nocturnal treefrog occurring in humid premontane and lower montane rainforest areas.
Males usually call from vegetation overhanging the fastest-flowing stretches of streams, 1-3m above the water.
Amplexus and oviposition are unknown for this species. Eggs are deposited under rocks in streams, and the tadpoles
attach themselves to rocks by means of the large oral funnel (Savage 2002).
Major Threats The observed population decline has taken place in pristine habitats, and might be due to chytridiomycosis,
© Ignacio De La Riva

Geographic Range This species occurs in the lowest portion of the humid lower montane zone in the Cordillera
de Tilarán, Cordillera Central, and Cordillera de Talamanca of Costa Rica and western Panama, from 1,100-1,650m
asl (Savage 2002).
and perhaps also the effects of climate change. It is generally also affected by habitat loss as a result of logging.
Conservation Measures This species has been recorded from a number of protected areas in Costa Rica. Further
research is urgently needed into the ecology and population status of this frog; in view of the possible threat of
chytridiomycosis, ex-situ populations might need to be established.
Notes on taxonomy: This species was previously included in the genus Hyla but has recently been moved to the new genus Isthmohyla
(Faivovich et al. 2005).
Bibliography: Duellman, W.E. (2001), Faivovich, J. et al. (2005), Ibáñez, R. et al. (2000), Myers, C.W. and Duellman, W.E. (1982), Pounds,
J.A. et al. (1997), Savage, J.M. (2002), Young, B. et al. (1999)
Data Providers: Frank Solís, Roberto Ibáñez, Alan Pounds, Federico Bolaños, Gerardo Chaves, Jay Savage, César Jaramillo, Querube
Fuenmayor

VU Isthmohyla zeteki (Gaige, 1929)

Vulnerable B1ab(iii) Geographic Range This species can be found in humid premontane areas or marginally in the lower montane
Order, Family: Anura, Hylidae zone of the Cordillera Central and Cordillera de Talamanca of Costa Rica and western Panama, from 1,200-1,804m
Country Distribution: Costa Rica, Panama asl (Savage 2002).
Current Population Trend: Decreasing Population The available evidence suggests that Costa Rican populations are stable in suitable habitat, and have
not exhibited the declines noted in many other montane species.
Habitat and Ecology It inhabits bromeliads in humid montane forest. Breeding and larval development takes place
in the bromeliads (phytotelmic species), with eggs laid on the outside leaves above the waterline in the bromeliad
cups (Savage 2002).
Major Threats The threats to this species are not well known, but likely include general habitat loss as a result of
logging, human settlement, and some agricultural activities.
© Robert Puschendorf

Conservation Measures This species is known from a number of protected areas, including Parque Nacional
Braulio Carrillo.
Notes on taxonomy: This species was previously included in the genus Hyla but has recently been moved to the new genus Isthmohyla
(Faivovich et al. 2005).
Bibliography: Duellman, W.E. (2001), Dunn, E.R. (1937), Faivovich, J. et al. (2005), Ibáñez, R. et al. (2000), Savage, J.M. (2002), Young,
B. et al. (1999)
Data Providers: Frank Solís, Roberto Ibáñez, Gerardo Chaves, Jay Savage, César Jaramillo, Querube Fuenmayor

VU Litoria andiirrmalin McDonald, 1997 CAPE MELVILLE TREEFROG

Vulnerable D2 Geographic Range This species is known only from four streams in boulder fields in northern Queensland, Australia,
Order, Family: Anura, Hylidae above 60m asl.
Country Distribution: Australia Population There is no information on the population status of this species, but it is believed to be stable.
Current Population Trend: Stable Habitat and Ecology This species is found associated with rocky streams with large granite boulders and granite
bedrock in rainforest and mesophyll vine forest with Melaleuca emergents. It is often seen sitting on rocks, vines and
twigs adjacent to cascades and riffles. It shelters under rocks and vegetation. It breeds in summer with the onset of
the wet season. Males can be heard calling in late November.
Major Threats There are no known major threats.
Conservation Measures All populations are found within Cape Melville National Park. There is a need for close
population monitoring of this species given its restricted range.
Bibliography: McDonald, K.R. (1997)
© Stephen Richards

Data Providers: Jean-Marc Hero, Keith McDonald, Ross Alford, Michael Cunningham, Richard Retallick
256 Threatened Amphibians of the World
VU Litoria aurea (Lesson, 1830)
Vulnerable A2ace
Order, Family: Anura, Hylidae
Country Distribution: Australia, New Caledonia
(Introduced), New Zealand (Introduced)
Current Population Trend: Decreasing
© Marion Anstis
Geographic Range This Australian species has been recorded along the south-east coast from East Gippsland in
Victoria, north to approximately Byron Bay in north-east New South Wales (Gillespie 1996; White and Pyke 1996).
Most records are from elevations below 100m asl. North of Sydney, there were a few high-elevation records of
the species and almost all records were east of the Great Divide (the only discrepant records are from Armidale
and Ebor) (White and Pyke 1996). On the Southern Tablelands, the species does not appear to occur above 800m
asl (Osborne, Littlejohn and Thomson 1996) and in Victoria the species does not appear to occur above about
670m asl (Gillespie 1996). Populations occur on two offshore islands in New South Wales, Bowen Island in Jervis
Bay (Osborne and McElhinney 1996) and Broughton Island north of Port Stephens (New South Wales NPWS Atlas
1998). It remains unknown whether or not these populations are relictual or the result of assisted translocation
(Mahony 1999). It has been introduced to New Zealand and is widespread across northern North Island, and pet
traders have moved it between the North and South Islands. It is also introduced to New Caledonia and New
Hebrides (Tyler 1979).
Population This species was formerly considered to be common throughout its range (Tyler 1992). Since about
1960, declines in the distribution and abundance of the species have been observed to the extent that it may now
be regarded as rare (White and Pyke 1996). In New South Wales, the species has disappeared completely from all
highland areas above 250m asl and coastal populations have been reduced in number and are more isolated from
other populations (White and Pyke 1996). Osborne, Littlejohn and Thomson (1996) observed that the species had
declined in the Australian Capital Territory and had not been recorded from the Southern Tablelands since 1980.
Declines in abundance have been observed in New South Wales and the Australian Capital Territory (White and
Pyke 1996; Osborne, Littlejohn and Thomson 1996; Lewis and Goldingay 1999; Goldingay and Lewis 1999), although
no similar decline in distribution and abundance in Victoria is so far apparent (Gillespie 1996). Recent censuses of
populations throughout the distribution of the species indicate that many are small, with most estimates being less
than 20 adults (White and Pyke 1996). In New Zealand, where the species has been introduced, there are many
thousands of individuals, but local declines have been observed with chytridiomycosis and introduced Gambusia
fish being implicated.
Habitat and Ecology The natural habitat requirements of the species have proved difficult to define because it has
been associated with almost every type of water body except fast-flowing streams (Pyke and White 1996). There
also appears to be some confusion over whether or not forested habitats are utilized by the species (Lemckert 1996;
Gillespie 1996). Pyke and White (1996) examined sites where L. aurea is known to have been present, and compared
VU Litoria becki (Loveridge, 1945)
Vulnerable D2
Order, Family: Anura, Hylidae
Country Distribution: Papua New Guinea
Current Population Trend: Unknown
© Stephen Richards
CR Litoria booroolongensis (Moore, 1961)
Critically Endangered B2ab(i,ii,iii,iv,v)
Order, Family: Anura, Hylidae
Country Distribution: Australia
Current Population Trend: Decreasing
© Marion Anstis
Geographic Range This Australian endemic ranges from the Queensland border south down the Great Dividing
Range almost to the Victorian border. It has not been recorded from the Northern Tablelands during the past 15 years
despite extensive surveys. The only extant population in Northern New South Wales is near Tamworth. The area of
GREEN AND GOLDEN BELL FROG
the habitat at sites where breeding was identified with that at locations where breeding was not identified, in New
South Wales. Sites which supported breeding populations were found to contain waterbodies which are still, shal-
low, ephemeral, unpolluted, unshaded, with aquatic plants and generally free of Gambusia and other predatory fish
(but not always); adjacent terrestrial habitats consisted of grassy areas and vegetation no higher than woodlands,
and a range of diurnal shelter sites. Breeding occurred in a significantly higher proportion of sites with ephemeral
ponds rather than sites with fluctuating or permanent ponds, and where predatory fish were absent. Mahony (1999)
commented on the limitations of the study and suggested that the results do not necessarily identify the requirements
of the species prior to declines. It is worthy to note that the use of ephemeral breeding sites was not a feature as-
sociated with members of the bell frog group in earlier habitat descriptions (Mahony 1999). Pyke and White (1996)
suggest that the habitat requirements of L. aurea in New South Wales and Victoria differ. In New South Wales the
species occupies disturbed habitats and breeding largely occurs in ephemeral ponds (Pyke and White 1996). However,
in Victoria it occupies habitats with little human disturbance and commonly breeds in permanent ponds as well as
ephemeral ponds (Pyke and White 1996). Goldingay (1996) argued that this is because most natural habitats are
degraded or lost in New South Wales. In Victoria the species is predominantly found on the coastal plains and low
foothills of the hinterland where it has been recorded in a range of lentic and terrestrial habitats (Gillespie 1996).
Breeding has been documented from dams in both forested and cleared areas, swamps in farmland, gravel pits,
billabongs, marshes, coastal lagoon wetlands, wet swale herb lands and isolated stream-side pools (Gillespie 1996).
These habitats are mostly permanent, but include some ephemeral waterbodies (Gillespie 1996). All habitats are
characterized by stationary water (Gillespie 1996). Virtually all isolated waterbodies are free of native fish species
and typically have dense emergent vegetation (Gillespie 1996). It can be found in a variety of terrestrial habitats
including lowland forest, banksia woodland, wet heath land, riparian scrub complex, riparian shrubland, riparian
forest, damp forest, shrubby dry forest and cleared pastoral lands (Gillespie 1996). It is seasonally active and has
been observed from September to early May (Daly 1995). Males call between September and March. Spawn is laid
amongst aquatic vegetation and has been observed in December, January and February (Daly 1995). Counts of eight
egg masses ranged from 2,463-11,682 eggs (van de Mortel and Goldingay 1998). Eggs hatch within three days and
metamorphosis can take from 2-11 months (Daly 1995); however, six weeks appears to be an average duration for
the field (R. Goldingay pers. comm.).
Major Threats The cause(s) of the apparent declines observed in populations of all taxa within the L. aurea complex
are unclear (Gillespie, Osborne and McElhinney 1995). Investigations of disappearances among the group have
primarily focused on L. aurea and L. castanea and two major directions in research have been pursued: the role of
increased ultraviolet radiation; and the impact of the introduced fish, Gambusia (Mahony 1999). It is also possible
that disease, such as a viral infection or chytrid fungus, might have contributed to the decline of this species (W.
Osborne pers. comm.). Chytrid fungus was detected in this species in Hoskinstown and Homebush Bay in Sydney,
New South Wales.
Conservation Measures There has been a lot of research into threats to the species and movement towards a
conservation strategy. Its range includes several protected areas. As an introduced species in New Zealand it has
the potential to spread chytridiomycosis to areas inhabited by native frogs. There is a cooperative program between
Taronga Zoo and a range of NSW agencies and NGO’s, involving breeding and release at a number of sites close to
Sydney. This program is currently under review.
Bibliography: Bell, B.D. (1982a), Bell, B.D. (1982b), Courtice, G.P. and Grigg, G.C. (1975), Daly, G. (1995), Gill, B.J., Whitaker, A.H. (1996),
Gillespie, G.R. (1996), Gillespie, G.R. and Hero, J.M. (1999), Gillespie, G.R., Osborne, W.S. and McElhinney, N.A. (1995), Goldingay, R. and
Lewis, B. (1999), Goldingay, R.L. (1996), Hero, J.-M., Littlejohn, M. and Marantelli, G. (1991), Lemckert, F. (1996), Lewis, B. and Goldingay,
R. (1999), Mahony, M. (1999), Morgan, L.A. and Buttermer, W.A. (1996), Murphy, M.J. (1995), NSW National Parks and Wildlife Service
(1998), Osborne, W.S. and McElhinney, N.A. (1996), Osborne, W.S., Littlejohn, M.J. and Thomson, S.A. (1996), Pyke, G.H. et al. (2002),
Pyke, G.H. and White, A.W. (1996), Tyler, M.J. (1979), Tyler, M.J. (1992), Tyler, M.J. (1997), van de Mortel, T.F. and Buttermer, W.A. (1996),
van de Mortel, T.F. and Goldingay, R. (1996), Webb, C.E. and Joss, J. (1997)
Data Providers: Jean-Marc Hero, Graeme Gillespie, Harold Cogger, Frank Lemckert, Peter Robertson
Geographic Range This species is currently known only from Mount Wilhelm in the central mountains of Papua
New Guinea, but it might have a broader distribution at high altitudes in New Guinea. Another population might
survive on Mount Giluwe and perhaps also on Mount Hagen. It has been recorded from above 3,000m asl, and so
the amount of habitat available within its potential range is probably limited.
Population There is no recent information available on the population status of this species. There are specimens
in a collection in Papua New Guinea from the 1970s and 1980s that might represent this species.
Habitat and Ecology There are no specific data available, but it is probably found in montane rainforest. It presum-
ably breeds in torrential high-altitude streams.
Major Threats There is no information on threats to this species, but it is known from an area where there is very
little human impact. A potential future threat is the spread of chytridiomycosis, which has been implicated in the
declines of high-altitude, torrent-dwelling treefrogs in Australia.
Conservation Measures It is not known from any protected areas. There is a need for further survey work to better de-
termine the distribution and population status of this species in the high mountains of central Papua New Guinea.
Notes on taxonomy: This species is part of a taxonomically poorly defined group of frogs (R. Zweifel pers. comm.).
Bibliography: Tyler, M.J. (1968)
Data Providers: Stephen Richards, Fred Parker
BOOROOLONG FROG
occupancy of this species is only approximately 10km² and is severely fragmented. It has been recorded between
200 and 1,000m asl.
Population There have been very few records of the species in the past five years and the species is believed to
have undergone massive declines over its entire range.
Habitat and Ecology This is a highland species associated with western-flowing rocky streams on the slopes and
tablelands of the Great Dividing Range. Streams are slow-flowing and bordered by grassy vegetation. Males begin
calling in August from rocks in or near the water. It is most active at night but also often found in daylight on rocks
on the waters edge.
Major Threats The widespread disappearance of this species across most of its range is probably due to chytridiomy-
cosis. In addition, introduced fish occur in many streams where the species has been recorded, which exert predatory
pressure upon tadpoles of this species. Land clearing, forest grazing and timber harvesting have occurred adjacent
to or in the headwaters of catchments in which the species has been recorded. Flow modification and weed invasion
(particularly by willows) has also occurred along many streams where the species occurs.
Conservation Measures Development of a management plan is under way, but much further research and survey
work is needed as is protection and rehabilitation of remaining habitat. The range of the species includes several
protected areas, and it is given protection where it occurs in state forests. Given the probable threat of chytridiomy-
cosis, recommended conservation measures will probably need to include the establishment of a captive-breeding
programme.
Bibliography: Barker, J., Grigg, G. and Tyler, M. (1995), Gillespie, G. and Hines, H.B. (1999), Moore, J.A. (1961)
Data Providers: Jean-Marc Hero, Graeme Gillespie, Frank Lemckert, Peter Robertson, Murray Littlejohn
 Globally Threatened Amphibian Species 257

EN Litoria brevipalmata Tyler, Martin and Watson, 1972 GREEN-THIGHED FROG

Endangered B2ab(iii,iv,v) Geographic Range This Australian endemic occurs from Cordalba State Forest in south-east Queensland south
Order, Family: Anura, Hylidae to Ourimbah, approximately 100km north of Sydney in New South Wales. The Darkes Forest records are erroneous.
Country Distribution: Australia They have been recorded from sea level up to at least 150m asl, and possibly a little higher. The area of occupancy
Current Population Trend: Decreasing is thought to be less than 500km2.
Population It has decreased at Ourimbah on the central coast of New South Wales, but elsewhere there have been no
reports of declines or disappearances. It is still found throughout the Ourimbah Valley. Additional populations have been
found in Queensland as a result of increased surveying. It is listed as rare in Queensland and New South Wales.
Habitat and Ecology It inhabits the leaf-litter and low vegetation of forests and prefers wetter forest types in the
southern half of its range, but extends also into open and drier forests in north-east New South Wales and south-east
Queensland (Lemckert et al. 2006). It breeds after heavy rains anywhere from September to May (spring to autumn
in the southern hemisphere), preferring larger temporary pools, and flooded areas for breeding (Lemckert and Slatyer
2002). Eggs (about 500-600) are laid in loose clumps among waterweed. The larvae are free swimming.
© Marion Anstis

Major Threats Recently, the development of an extensive highway system and rapidly expanding coastal lowland
development has increased threats to this species. Several populations have been directly affected by road construction
and fragmentation by roads and housing is probably significantly restricting movements between populations.
Conservation Measures Protection of breeding sites in state forests, national parks and other conservation parks
and reserves is in place. However, a management plan for the conservation of this species is needed.
Bibliography: Anstis, M. (1994), Barker, J., Grigg, G. and Tyler, M. (1995), Hines, H., Mahony, M. and McDonald, K. (1999), Lemckert,
F. et al. (2006), Lemckert, F.L. and Slatyer, C. (2002), Natrass, A.E.O. and Ingram, G.J. (1993)
Data Providers: Jean-Marc Hero, Harry Hines, Ed Meyer, Frank Lemckert, David Newell, John Clarke

CR Litoria castanea Steindachner, 1867 YELLOW-SPOTTED TREEFROG

Critically Endangered D Population There are no verified records of the northern population after 1975 (White and Ehmann 1997a) and the
Order, Family: Anura, Hylidae last specimen to be placed in the museum was collected in 1973 (Australian Museum Register in Mahony 1999).
Country Distribution: Australia The southern tablelands population suffered an extensive decline, with no confirmed records since 1980 (Osborne,
Current Population Trend: Unknown Littlejohn and Thomson 1996; Mahony 1999). It is not yet considered extinct because of the lack of surveys of potential
habitat, especially in areas between the northern and southern populations.
Habitat and Ecology Litoria castanea occupies similar habitat to L. aurea and L. raniformis which includes permanent
ponds, swamps, lagoons, farm dams and the still backwaters of rivers usually with tall reeds present (Courtice and Grigg
1975; White and Ehmann 1997a, b). The species was also found in ponds or slow moving streams with overhanging
grassy banks in the absence of reed beds (Courtice and Grigg 1975). Litoria castanea was found to over-winter in
the hollow centres of rotting logs and in the earth surrounding the roots of uprooted trees (Courtice and Grigg 1975),
and in the base of sedge tussocks (Humphries 1979). Little is known about the biology of this species; however, it is
© John Wombey

Geographic Range Some uncertainty surrounds the taxonomic status of the northern (L. flavipunctata of Courtice
and Grigg 1975) and southern populations of Litoria castanea. Thomson et al. (1996) suggest that the northern
and southern populations represent one species consisting of two disjunctive isolates separated by a distance of
about 500km (see also map in Osborne, Littlejohn and Thomson 1996). The northern population was known from
a relatively restricted distribution centred around the town of Guyra on the New England Tableland at altitudes
between 1,000 and 1,500m asl (White and Ehmann 1997a; Mahony 1999). It occupied the headwaters of the west
flowing Booroolong Creek and to a lesser extent those of the east flowing Anne River and Sarah River (Heatwole et
al. 1995). Near Armidale, the species has been recorded from Commissioners Waters, a tributary of the east flowing
Gara River (Heatwole et al. 1995). There are 13 known sites in the region (most above 1,000m asl) all of which have
been verified by examination of museum specimens or photographs (Mahony 1999). The southern population has
a restricted distribution between Canberra and Bombala on the Southern Tablelands at altitudes between 700 and
800m asl (Mahony 1999). The southern population was broadly sympatric with L. aurea in the north of its range and
with L. raniformis in the south-west of the region (Mahony 1999).
likely to be similar to that of L. aurea and L. raniformis (Gillespie, Osborne and McElhinney 1995). Individuals reach
sexual maturity at three years and live for six years.
Major Threats The cause(s) of the apparent declines observed in populations of all taxa within the L. aurea com-
plex are unclear (Gillespie, Osborne and McElhinney 1995), although chytridiomycosis must be strongly suspected.
Investigations of disappearances among the group have primarily focused on L. aurea and L. castanea and two major
directions in research have been pursued: the role of increased ultraviolet radiation; and the impact of the introduced
fish, Gambusia, which is native to southern and eastern USA (Mahony 1999). However, it is likely that disease, such
as chytrid fungus ora viral infection, contributed to the decline of this species (W. Osborne pers. comm.).
Conservation Measures Further survey work is required to determine whether or not this species still survives,
particularly in the southern tablelands, and in areas of suitable habitat between the northern and southern popula-
tions. Research into possible causes of this species’ disappearance and the decline of similar species (L. aurea and
L. raniformis) is urgently needed.
Bibliography: Barker, J., Grigg, G. and Tyler, M. (1995), Courtice, G.P. and Grigg, G.C. (1975), Gillespie, G.R. and Hero, J.M. (1999),
Gillespie, G.R., Osborne, W.S. and McElhinney, N.A. (1995), Goldingay, R. and Lewis, B. (1999), Heatwole, H. et al. (1995), Humphries,
R. (1979), Mahony, M. (1999), Morgan, L.A. and Buttermer, W.A. (1996), Osborne, W.S., Littlejohn, M.J. and Thomson, S.A. (1996),
Thompson, S.A. et al. (1996), Tyler, M.J. (1997), van de Mortel, T.F. and Buttermer, W.A. (1996), Webb, C.E. and Joss, J. (1997), White,
A.W. and Ehmann, H. (1997a), White, A.W. and Ehmann, H. (1997b)
Data Providers: Jean-Marc Hero, Harry Hines, Frank Lemckert, Peter Robertson

EN Litoria cooloolensis Liem, 1974 COOLOOLA TREEFROG

Endangered B1ab(iii) Geographic Range This Australian endemic is known from south-eastern Queensland in Cooloola (or Great Sandy)
Order, Family: Anura, Hylidae National Park on Fraser Island, and North Stradbroke Island.
Country Distribution: Australia Population It occurs in large numbers on Fraser Island.
Current Population Trend: Decreasing Habitat and Ecology The species is found in sandy coastal and island freshwater lakes and wallum creeks, where
it has a preference for dense reed beds. It is a spring and summer breeder, with males calling from reeds or trees
around freshwater lakes. Eggs are deposited on submerged vegetation; larvae are free-swimming.
Major Threats High human visitation to the freshwater lakes important for breeding is a major threat, as is the
trampling of reed beds and the pollution of water. Water extraction for sand mining and domestic use has had
significant impacts on populations on North Stradbroke Island through habitat loss, deterioration in water quality
and alterations in hydrology. More significantly, this species has virtually disappeared from Brown Lake on North
Stradbroke Island following the introduction of predatory Gambusia fish there in 2003. However, overall these threats
are relatively localized.
© Marion Anstis

Conservation Measures Most of the population is protected in the Cooloola or Great Sandy National Park. Fraser
Island was inscribed into the list of UNESCO World Heritage Areas in 1992. Measures need to be taken to prevent
the spread of Gambusia within its range, and to control the impacts of tourism.
Notes on taxonomy: Genetic, vocalization and morphological differences between the populations on Cooloola and Fraser Island
and those on North Stradbroke Island indicate that the the latter might represent a distinct species or subspecies (James 2000; E.
Meyer unpubl.).
Bibliography: Barker, J., Grigg, G. and Tyler, M. (1995), Hines, H., Mahony, M. and McDonald, K. (1999), James, C.H. (1998), Liem,
D.S. (1974a)
Data Providers: Harry Hines, Ed Meyer, Jean-Marc Hero, David Newell, John Clarke
258
VU Litoria daviesae Mahony, Knowles, Foster and Donnellan, 2001
Vulnerable B1ab(iii)
Order, Family: Anura, Hylidae
Country Distribution: Australia
Current Population Trend: Decreasing
© Ross Knowles
VU Litoria freycineti Tschudi, 1838
Vulnerable B2ab(ii,iii,iv,v)
Order, Family: Anura, Hylidae
Country Distribution: Australia
Current Population Trend: Decreasing
© Marion Anstis
CR Litoria lorica Davies and McDonald, 1979
Critically Endangered D
Order, Family: Anura, Hylidae
Country Distribution: Australia
Current Population Trend: Decreasing
est between 640 and 1,000m asl (McDonald 1992), although it might be present at lower altitudes (M. Cunningham
pers. comm.). It has been found on granite boulders in notophyll vine forest in the splash zone near turbulent fast
flowing water (Davies and McDonald 1979). There is no information on breeding season, timing of reproduction, or
VU Litoria lutea (Boulenger, 1887)
Vulnerable B1ab(iii)
Order, Family: Anura, Hylidae
Country Distribution: Papua New Guinea,
Solomon Islands
Current Population Trend: Decreasing
© Stephen Richards
Threatened Amphibians of the World
Geographic Range This species was first located in 1976 and
later described by Davies and McDonald (1979). It is known from
four localities-Alexandra Creek (TR165); Hilda Creek (near Thornton
Peak, Cape Tribulation National Park); Roaring Meg Cascades (via
Mount Sorrow TR165); and Bluff Creek (Mossman Gorge Daintree
National Park), north-east Queensland (Hero and Fickling 1994; M.
Cunningham pers. comm.).
Population It was once moderately common in suitable habitat, but
it has undergone a dramatic decline. Despite recent efforts to locate
the species, it has not been seen since 1991 (Ingram and McDonald
1993; Hero et al. 1998, 2002), and it might now be extinct. However,
few searches for the species have been undertaken at historical sites
(J-M. Hero and M. Cunningham pers. comm.).
Habitat and Ecology This species is known to be a rainforest
specialist endemic to the Wet Tropics Bioregion (Williams and Hero
1998, 2001). As a stream-dwelling/stream-breeding species (Hero
and Fickling 1994) it prefers fast-flowing streams in upland rainfor-
Geographic Range This species is known from 14 different locations in a narrow band on the eastern edge of the
tablelands and the great escarpment of the Great Dividing Range, within central-eastern to lower north-eastern New
South Wales, Australia. It is only found above 400m asl.
Population It is not abundant, and has generally been recorded as small populations.
Habitat and Ecology This species is associated with areas of unpolluted, upland streams in heath land or dry open
forest on the tablelands, or wet sclerophyll and rainforest vegetation on the edge of the escarpment and in deeper
gullies. The adults do not occur away from upland stream habitats. Nothing is known about habitat use of this species
outside the breeding season. The larvae develop in the streams.
Major Threats This species has a small geographic range and has suffered habitat loss and fragmentation through
clearance for agriculture. Many of the streams within the species’ range have been stocked with exotic fishes (carp,
trout, mosquito fish) that are likely to be predators of the tadpoles. The impact of forestry practices at many sites
where the species is known to occur requires assessment.
Conservation Measures It is known from a number of protected areas.
Bibliography: Mahony, M. et al. (2001)
Data Providers: Jean-Marc Hero, Michael Mahony
FREYCINET’S FROG
Geographic Range This Australian endemic occurs from Fraser Island in Queensland, south to Jervis Bay in mid-
eastern New South Wales.
Population There is no information on population status.
Habitat and Ecology This species occupies a variety of habitats, but is most commonly seen in the vicinity of
temporary coastal swamps (wallum). These areas typically have low nutrient soils and heath, sedge land, Melaleuca
swamp or Banksia woodland vegetation. It is mostly a nocturnal species, and is a spring and summer breeder. Eggs
are demersal and laid in shallow water, and the larvae are free-swimming.
Major Threats Clearing of habitat for agriculture, pine plantations, housing and other infrastructure is a threat; they
occur in an area with the highest rate of human population growth in Australia. Other threats include habitat degradation
through alterations in hydrology and increased nutrient loads, invading weeds, and inappropriate fire regimes.
Conservation Measures The range of the species includes several protected areas, including Fraser Island
National Park.
Bibliography: Barker, J., Grigg, G. and Tyler, M. (1995), Hines, H., Mahony, M. and McDonald, K. (1999), Moore, J.A. (1961)
Data Providers: Harry Hines, Jean-Marc Hero, Ed Meyer, David Newell
ARMOURED FROG
egg deposition sites. The tadpoles of the species have not been described but are thought to be torrent-adapted
and similar to that of a sympatric species, Litoria nannotis (Davies and McDonald 1979). Unpigmented eggs (M.
Cunningham unpubl.) are presumably laid in water under rocks in the stream.
Major Threats The reason(s) for the decline of this species are unknown, although chytridiomycosis must be strongly
suspected. The habitat of the species in the Wet Tropics has been protected since 1988; therefore, habitat destruction
is no longer a threat (McDonald and Alford 1999). Current research is examining the possibility that disease, such
as a viral and infection or chytrid fungus, might have contributed to the decline of this species (Berger, Speare and
Hyatt 1999). Feral pigs have been suggested as a potential cause of riparian habitat damage and adult frog mortal-
ity (Richards, McDonald and Alford 1993). However, feral pigs are unlikely to have a direct impact due to the close
association of this species with rocky cascades in streams (M. Cunningham pers. comm.).
Conservation Measures The habitat of the species in the Wet Tropics has been protected within protected areas
since 1988. This species is a priority for immediate survey work to determine whether or not it still survives at the
localities from which it has previously been recorded. Research is also needed into the possible reasons for the decline
of the species. Given the possible threat of chytridiomycosis, surviving individuals might need to form the basis for
the establishment of an ex-situ population.
Bibliography: Barker, J., Grigg, G. and Tyler, M. (1995), Berger, L., Speare, R. and Hyatt, A. (1999), Davies, M. and McDonald, K.R.
(1979), Hero, J.-M. et al. (1998), Hero, J.-M. et al. (2002), Hero, J.-M. and Fickling, S. (1994), Ingram, G.J. and McDonald, K.R. (1993),
McDonald, K. and Alford, R. (1999), McDonald, K.R. (1992), Richards, S.J., McDonald, K.R. and Alford, R.A. (1993), Williams, S.E. and
Hero, J.-M. (1998), Williams, S.E. and Hero, J.-M. (2001)
Data Providers: Jean-Marc Hero, Michael Cunningham, Ross Alford, Keith McDonald
Geographic Range This is a poorly known species occurring in the north-western islands of the Solomons archipelago,
from Isabel and New Georgia in the south, to Buka (in Papua New Guinea) in the north. Recent studies (S. Richards,
unpubl.) suggest that this species is more widespread in the Solomon Islands than previously documented. It is a
lowland species that probably occurs only up to about 50m asl.
Population There is no information on population status, but it most likely occurs at low densities.
Habitat and Ecology This species is arboreal, and found in tropical rainforest. It possibly breeds in swamps in
forests. In Bougainville it has been observed laying eggs on the vertical surface of trees above water-filled tree holes;
the tadpoles then fall into the water.
Major Threats The major threat to this species is habitat loss due to logging.
Conservation Measures It is not recorded from any protected areas, so there is a need for improved habitat
protection at the lowland sites where this species is known to occur. The distribution of this species requires further
documentation.
Bibliography: Brown, W.C. (1952), Tyler, M.J. (1968)
Data Providers: Stephen Richards, Fred Parker
 Globally Threatened Amphibian Species
EN Litoria nannotis (Andersson, 1916)
Endangered A2ae
Order, Family: Anura, Hylidae
Country Distribution: Australia
Current Population Trend: Stable
© Jodi Rowley
Geographic Range This species is endemic to the Wet Tropics Bioregion in north Queensland, from Paluma to
Cooktown (Hero and Fickling 1994; Williams and Hero 1998, 2001) at altitudes between 180 and 1,300m asl (McDonald
1992). It includes three deeply divergent mitochondrial DNA lineages, distributed from Paluma to Tully River, Tully
River to Lamb Range, and Mount Lewis to Big Tableland (Schneider, Cunningham and Moritz 1998).
Population This species was first noted to have declined in 1990 (Richards, McDonald and Alford 1993), since it
had apparently disappeared from most upland sites south of the Daintree River. The species occurred at all lowland
sites, and at upland sites north of the Daintree River during summer surveys in 1991-1992. In 1994 it was sighted at
several locations above 600m asl on Mount Father Clancy. Lowland populations surveyed in Tully Gorge appeared
to be relatively stable between 1995 and 1998 (J.-M. Hero pers. comm.). At the southern end of its range it was
last observed in Mount Spec State Forest in 1991 (Richards, McDonald and Alford 1993); however, adults occurred
at a lower elevation site in a different creek system (Crystal Creek Stone Bridge, 300m asl) from January 1994 to
September 1995 (J.-M. Hero pers. comm.). This species is currently known to have stable populations at lowland
sites (Hero et al. 1998; McDonald and Alford 1999).
CR Litoria nyakalensis Liem, 1974
Critically Endangered D
Order, Family: Anura, Hylidae
Country Distribution: Australia
Current Population Trend: Decreasing
© Stephen Richards
Geographic Range This species, an Australian endemic, formerly occurred across two thirds of the Wet Tropics from
Douglas Creek near Cardwell to Alexandra Creek, Thornton Peak north-east Queensland (Hero and Fickling 1994) at
altitudes between 380 and 1,020m asl (McDonald 1992).
Population It was once moderately common in suitable habitat, but it has undergone a catastrophic decline. Adults
were last recorded in April 1990, and tadpoles and metamorphs were last recorded in November 1990 on the Carbine
Tableland (Richards, McDonald and Alford 1993). However, this species had apparently disappeared from sites on
the Atherton Tableland much earlier (Richards, McDonald and Alford 1993). It was recorded from various sites on the
Atherton Tableland prior to 1973 (Liem 1974b), but was not encountered in Danbulla State Forest during 1989-1992
or at any Atherton Tableland site during surveys conducted between 1991 and 1992 (Richards, McDonald and Alford
1993). No information is available on population structure or genetic variation (M. Cunningham pers. comm.). The
VU Litoria olongburensis Liem and Ingram, 1977
Vulnerable B1ab(ii,iii,iv)+2ab(ii,iii,iv)
Order, Family: Anura, Hylidae
Country Distribution: Australia
Current Population Trend: Decreasing
© Marion Anstis
Geographic Range This species is distributed from Lake Woongeel, Fraser Island, south to near Woolgoolga including
Bribie, Moreton and Stradbroke Island, south-east Queensland, Australia (Hines, Mahony and McDonald 1999).
Population There is no information on population size, structure or dynamics (Hines, Mahony and McDonald 1999). The
species exhibits genetic structuring on a north-south gradient (James 1996). Populations in northern New South Wales,
North Stradbroke Island, Moreton Island and Cooloola-Fraser are each genetically different and significantly divergent
from each other, and should be considered demographically independent due to their insular nature (James 1996).
Habitat and Ecology This species inhabits coastal sand plains and dunes of “the wallum” (Hines, Mahony and Mc-
Donald 1999). Vegetation types typical of these environments include heath land, Melaleuca swamp, sedge land and
Banksia woodland (Hines, Mahony and McDonald 1999). It is associated with low pH waters characteristic of wallum
259
TORRENT TREEFROG
Habitat and Ecology Litoria nannotis is a habitat specialist, restricted to rocky stream habitats in rainforest or
wet sclerophyll forest where there is fast-flowing water, waterfalls and cascades (Liem 1974b; McDonald 1992). It
is a stream dwelling and breeding species (Hodgkison and Hero 2001), but, unlike most stream-breeding frog spe-
cies that live in the adjacent forest and use the stream habitat for breeding, the stream is the primary habitat for
both male and females throughout the year (Hodgkison and Hero 2001, 2002). On several occasions the adults and
juveniles were noted to form small aggregations (4-6 individuals) amongst large boulders behind waterfalls (Liem
1974b; J-M. Hero pers. obs.). Gravid females and males with nuptial pads are encountered all year round (Martin
and McDonald 1995). Unpigmented eggs numbering 136-216 (1.98-3.4mm in diameter) are laid in gelatinous egg
masses under rocks in water (Liem 1974b; Hero and Fickling 1996). Liem (1974b) described the larva and noted that
it is one of the few species of frog known to exhibit larval adaptations to torrent environments of Australia, such as
a streamlined body shape, large suctorial mouthparts and a muscular tail. Richards (1992) also provided information
on the larvae of the L. nannotis group.
Major Threats The reason(s) for the decline are unknown. Richards, McDonald and Alford (1993) reject drought,
floods, habitat destruction or pollution by pesticides, inorganic ions or heavy metals. The habitat of the species in
the Wet Tropics has been protected since 1988, and habitat destruction is no longer a major threat (McDonald and
Alford 1999). Research is examining the possibility that disease, such as a viral infection or chytrid fungus, may
have contributed to the decline of this species (Berger, Speare and Hyattt 1999). Feral pigs are a potential cause of
riparian habitat damage and adult frog mortality (Richards, McDonald and Alford 1993). The activity of feral pigs has
been recorded to have increased over the period 1989-1992 in an area previously inhabited by L. nannotis (Richards,
McDonald and Alford 1993). However, there has been very little research into the impact of feral pigs on native frog
populations (Richards, McDonald and Alford 1993).
Conservation Measures There is a need for continued and strengthened protection of habitat in the Wet Tropics,
including an improved management plan that involves dealing with the threat posed by feral pigs. Further research
into the reasons for the decline of this species in pristine habitat is necessary.
Bibliography: Barker, J., Grigg, G. and Tyler, M. (1995), Berger, L., Speare, R. and Hyatt, A. (1999), Cogger, H.G. (1992), Hero, J.-M. et al.
(1998), Hero, J.-M. and Fickling, S. (1996), Hodgkison, S.C. and Hero, J.-M. (2001), Hodgkison, S.C. and Hero, J.-M. (2002), Hodgkison,
S.C. and Hero, J.-M. (2003), Liem, D.S. (1974b), Martin, W.F. and McDonald, K.R. (1995), McDonald, K. and Alford, R. (1999), McDonald,
K.R. (1992), Richards, S.J. (1992), Richards, S.J., McDonald, K.R. and Alford, R.A. (1993), Schneider, C.J., Cunningham, M. and Moritz,
C. (1998), Tyler, M.J. (1997), Williams, S.E. and Hero, J.-M. (1998), Williams, S.E. and Hero, J.-M. (2001)
Data Providers: Jean-Marc Hero, Richard Retallick
MOUNTAIN MISTFROG
species might now be extinct.
Habitat and Ecology This species is a rainforest specialist, endemic to the Wet Tropics Bioregion (Williams and Hero
1998, 2001) found in upland rainforest and wet sclerophyll forest along fast-flowing streams where there is white
water from riffles and cascades (Liem 1974b; McDonald 1992). It is usually found perched on rocks or overhanging
vegetation adjacent to the water (Liem 1974b). Little is known about the life history of this species. Mating calls have
been heard from October to March (Liem 1974b). From 86-90 large unpigmented eggs (1.9-2.5mm diameter) are laid
under rocks in riffles (Richards 1993a; Hero and Fickling 1996). Richards (1992) described the tadpole and noted that
it is one of the few species of tadpole known to exhibit adaptations to torrent environments of Australia, including
a streamlined body shape, large suctorial mouthparts and muscular tail. Tadpoles commonly overwinter in upland
streams, although those hatching in early summer can metamorphose before the next autumn (Richards 1992).
Major Threats The reason(s) for the decline of this species are unknown, although chytridiomycosis must be strongly
suspected. Richards, McDonald and Alford (1993) reject drought, floods, habitat destruction, or pollution by pesticides,
inorganic ions or heavy metals. The habitat of the species in the Wet Tropics has been protected since 1988; therefore,
habitat destruction is no longer a threat (McDonald and Alford 1999). Current research is examining the possibility
that disease, such as a viral infection or chytrid fungus, might have contributed to the decline of this species (Berger,
Speare and Hyatt 1999). Feral pigs are a potential cause of riparian habitat damage and adult frog mortality (Richards,
McDonald and Alford 1993). The activity of feral pigs has been recorded to have increased over the period 1989-1992
in an area previously inhabited by this species (Richards, McDonald and Alford 1993).
Conservation Measures The habitat of the species in the Wet Tropics has been protected since 1988. This species
is a priority for immediate survey work to determine whether or not it still survives at the localities from which it has
previously been recorded. Research is also needed into the possible reasons for the decline of the species. Given
the possible threat of chytridiomycosis or some other disease, surviving individuals might need to form the basis for
the establishment of an ex-situ population.
Bibliography: Berger, L., Speare, R. and Hyatt, A. (1999), Hero, J.-M. and Fickling, S. (1994), Hero, J.-M. and Fickling, S. (1996), Liem, D.S.
(1974b), McDonald, K. and Alford, R. (1999), McDonald, K.R. (1992), Richards, S.J. (1992), Richards, S.J. (1993a), Richards, S.J., McDonald,
K.R. and Alford, R.A. (1993), Tyler, M.J. (1997), Williams, S.E. and Hero, J.-M. (1998), Williams, S.E. and Hero, J.-M. (2001)
Data Providers: Jean-Marc Hero, Michael Cunningham, Ross Alford, Keith McDonald, Richard Retallick
OLONGBURRA FROG
environments, usually found in sedge swamps, and less commonly in coastal lakes and creek environs (Liem and Ingram
1977; James 1996; Ehmann 1997). During wet periods these swamps are heavily inundated and the species can be found
clinging to emergent vegetation (grasses, reeds and Bungwall Fern; E. Meyers pers. comm.). During dry periods individuals
may be found at the base of sedges, grass clumps and/or Bungwall Fern in the same swamps (E. Meyer pers. comm.).
Ingram and Corben (1975) termed this species an ‘acid’ frog as it is confined to sandy heaths and their acidic water.
Non-breeding habitat is not known. Limited information is available on the breeding biology of the species. Ehmann
(1997) recorded males calling in spring, summer and early autumn at night and by day when swamps were rising or ample
water was available. It mainly breeds following heavy rain in perched swamps, amidst sedges, grasses and/or Bungwell
Fern (Blechnum indicum) in still water 0.5-1.5m deep (E. Meyer pers. comm.). The wallum waters in which the species
breeds are typically heavily tanin-stained, highly acidic and generally dilute (E. Meyer pers. comm.). Eggs are attached
to grasses and sedges and larvae are nektonic in form and dwell amongst reeds and grasses in water 0.5-1.5m deep (E.
Meyer pers. comm.). Fish are largely absent from habitat occupied by the species (E. Meyer pers. comm.).
Major Threats Populations appear to be relatively stable in protected habitat; however, this species is at risk from
continuing loss of habitat through clearing for agriculture, pine plantations, housing and infrastructure such as canal
development, drainage projects and transport corridors (Ingram and McDonald 1993; Hines, Mahony and McDonald
1999). Melaleuca forest and heathland are particularly threatened and have been extensively cleared since 1974 (Cat-
terall and Kingston 1993) suggesting an extensive loss of habitat. Other threats include habitat degradation through
changes in hydrological regimes, increased nutrients or sediments, weed invasion, inappropriate fire management,
competition from invading frog species and predation from introduced fish (Hines, Mahony and McDonald 1999). Weed
invasion is a potentially threatening process (Hines, Mahony and McDonald 1999). Competition from invading frog
species and predation from introduced fish (i.e., Gambusia holbrooki) have been identified as potentially threatening
processes (Ehmann 1997; Hines, Mahony and McDonald 1999) but further research is required.
Conservation Measures The range of the species includes several protected areas.
Bibliography: Catterall, C.P. and Kingston, M. (1993), Ehmann, H. (1997a), Hines, H., Mahony, M. and McDonald, K. (1999), Ingram, G.J. and
Corben, C.J. (1975), Ingram, G.J. and McDonald, K.R. (1993), James, C. (1996), Liem, D.S and Ingram, G.J. (1977), Tyler, M.J. (1997)
Data Providers: Harry Hines, David Newell, Ed Meyer, Jean-Marc Hero, John Clarke
260 Threatened Amphibians of the World

CR Litoria piperata Tyler and Davies, 1985 PEPPERED TREEFROG

Critically Endangered D Geographic Range This Australian endemic was formerly known
Order, Family: Anura, Hylidae from five streams draining the east of the Northern Tablelands, from
Country Distribution: Australia 800-1,120m asl, from Gibraltar Range to Armidale, northern New
Current Population Trend: Unknown South Wales (Tyler and Davies 1985a). The distribution map does
not include northern populations that might belong to this species
(see additional information under Notes on Taxonomy).
Population Despite searches of the historic localities and other
streams with similar habitat within the region (Mahony 1997), the
species has not been seen since 1973. If the northern populations
do not belong to this species (see Notes on Taxonomy), then it is
possible that it is extinct.
Habitat and Ecology This species is known to occupy open forest
and wet sclerophyll forest (Heatwole et al. 1995). Little is known
about the breeding biology of this species. However, morphological
similarity to Litoria pearsoniana and L. phyllochroa suggests that
ecological similarities are likely (Gillespie and Hines 1999).
Major Threats The causes of the apparent declines are unknown.
However, most of the historic sites and other streams in the region have undergone substantial alteration and suf-
fered significant habitat disturbance through clearance of vegetation, grazing and timber harvesting (H.B. Hines pers.
comm.). Introduced predatory fish species (Eastern Gambusia Gambusia holbrooki and salmonids) occur in streams
formerly occupied by the species and might have displaced frog populations by predation upon larvae (Gillespie and
Hines 1999). Given the vulnerability of other members of the L. citropa group to trout predation, these fish are likely
to have had a significant impact on populations of this species. In addition, chytridiomycosis cannot be ruled out as
a cause of the decline.
Conservation Measures It is recognized as Endangered in Queensland and New South Wales, and therefore
protected by state legislation. Its range includes Gara River Nature Reserve and Mount Mitchell State Forest. This
species is a priority for immediate further survey work to determine whether or not it might possibly still survive at the
localities from which it has previously been recorded, and both taxonomic and survey work is required to determine
the status of the possible northern populations. Research is also needed into the possible reasons for the decline of
the species. Given the possible threat of chytridiomycosis or some other disease, surviving individuals might need to
form the basis for the establishment of an ex-situ population.
Notes on taxonomy: This assessment is based only on the sites from which the species was described by Tyler and Davies (1985a), i.e.,
between Armidale and Glen Innes. However, in 1992 surveys outside the known range on the Northern Tablelands located populations
of frogs which closely resemble this species (NSW NPWS 1994). While the external morphology of the population closely resembles
L. piperata, the mating call is very similar to L. pearsoniana (M. Mahony pers. comm. in Tyler 1997). It is possible that this species
represents morphologically distinct outlying populations of L. pearsoniana. Considerable confusion exists over the systematics of the
Litoria barringtonensis, L. pearsoniana, L. phyllochroa, and L. piperata complex. Studies of the genetic variation in populations of this
complex revealed that the currently recognized species boundaries are in need of major review (Donellan et al. 1999). Further genetic
and morphometric studies are required to resolve the systematics of these northern populations.
Bibliography: Barker, J., Grigg, G. and Tyler, M. (1995), Donnellan, S.C. et al. (1999), Gillespie, G. and Hines, H.B. (1999), Heatwole, H.
et al. (1995), Mahony, M., Knowles, R. and Patterson, L. (1997b), Tyler, M.J. (1997), Tyler, M.J. and Davies, M. (1985a)
Data Providers: Jean-Marc Hero, Harry Hines, Frank Lemckert

VU Litoria quadrilineata Tyler and Parker, 1974

Vulnerable D2 Geographic Range This species is known only from the type locality, that it currently is known only from a single locality; further survey work is needed to determine whether or not it
Order, Family: Anura, Hylidae Merauke, in southern Papua, Indonesia, and there have been very might occur more widely.
Country Distribution: Indonesia few records since it was first discovered in the late 1960s. It occurs Bibliography: Tyler, M.J. and Parker, F. (1974)
Current Population Trend: Unknown close to sea level. Data Providers: Stephen Richards, Fred Parker, Djoko Iskandar
Population There is no information on the population status of
this species.
Habitat and Ecology It has only been found in a grassy, swampy
area in the town of Merauke, where it breeds. It presumably occurs
elsewhere, but its broader habitat requirements are not yet known,
and surveys have not been successful in locating it.
Major Threats There is no information on major threats, but given
that it occurs close to sea level it might be susceptible to habitat
loss due to development in coastal areas. It has been found once in
the pet trade in Jakarta.
Conservation Measures It is not known from any protected
areas. This species requires close population monitoring, given

EN Litoria raniformis (Keferstein, 1867) SOUTHERN BELL FROG

Endangered A2ae Habitat and Ecology This species is usually found in association with dams, ponds and marshes, either amongst
Order, Family: Anura, Hylidae sedges and other semi-aquatic vegetation, or sheltering under logs and rocks up to 1300m asl (Gillespie, Osborne and
Country Distribution: Australia, New Zealand McElhinney 1995). It appears to be associated with permanent waterbodies though it is unclear whether, like Litoria
(Introduced) aurea, the species also utilizes ephemeral pools (Mahony 1999). It occurs both in woodland and areas of improved
Current Population Trend: Decreasing pasture (Gillespie, Osborne and McElhinney 1995). Little is known about the biology of this species; however, it is
likely to be similar to that of L. aurea (Gillespie, Osborne and McElhinney 1995). Males call from August to April (Hero,
Littlejohn and Marantelli 1991). The species breeds in permanent ponds or swamps, usually with extensive areas
of sedges and rushes from which adults call (Gillespie, Osborne and McElhinney 1995). About 1,700 eggs are laid in
a loose clump (Hero, Littlejohn and Marantelli 1991, Hero and Warrell unpubl.). The larvae are free swimming and
develop over summer and autumn (Gillespie, Osborne and McElhinney 1995). Metamorphosis takes place between
late summer and autumn, although larvae may overwinter and metamorphose the following season (Gillespie,
Osborne and McElhinney 1995).
Major Threats The cause(s) of the apparent declines observed in populations of all taxa within the L. aurea complex
© Marion Anstis

Geographic Range This species was formerly distributed across a large area of south-east Australia, including
Tasmania, from 0-1,300m asl (Osborne, Littlejohn and Thomson 1996). In New South Wales and the Australia Capital
Territory its range was centred on the Murray and Murrumbidgee River valleys and their tributaries. It occurred through-
out the Southern Tablelands and was also recorded on the Central Tablelands as far north as Bathurst (Ehmann and
White 1997). It was widespread across Victoria, absent only from the west desert regions and the east alpine regions
(Littlejohn 1963, 1982; Hero, Littlejohn and Marantelli 1991). In South Australia the species is known to occur along
the lower Murray River valley, the lower south-east to near Keith, and a small introduced population, in the Adelaide
Hills (Tyler 1978). In Tasmania, the species occurred broadly across the north and east of the island and on the Bass
Strait Island (Brook 1979), and in pockets in the south of the state. This species has also been introduced to New
Zealand, where it is widespread across North and South Islands and on Great Barrier and Stewart Islands.
Population It was once a common species, but serious declines have occurred in sections of the range (Mahony
1999). Ehmann and White (1997) noted that in New South Wales the species had disappeared from sites in the
central and southern highlands. It is currently widespread throughout the Murray River valley but has disappeared
from a number of sites along the Murrumbidgee River (Mahony 1999) and there are no recent records from the
Monaro District near the Victorian border (G. Gillespie pers. comm.). It persists in isolated populations in the greater
Melbourne area, and isolated populations are known from a few sites in central Victoria and Gippsland. A similar
decline has been noted in Tasmania, and it is now almost absent from the midlands of Tasmania. In New Zealand,
where the species is introduced, there are many thousands, although local declines due to chytridiomycosis and/or
introduced Gambusia fish have been observed.
are unclear (Gillespie, Osborne and McElhinney 1995). Investigations of disappearances among the group have
primarily focused on L. aurea and L. castanea and two major directions in research have been pursued: the role of
increased ultraviolet radiation; and the impact of the introduced fish, Gambusia (Mahony 1999). As for L. aurea, L.
raniformis has disappeared from sites where Gambusia is present (Mahony 1999; W. Osborne pers. comm.). The dates
of introduction of Gambusia to many regions are not well documented and this lack of information has hampered
research into declines (Mahony 1999). Introduced Gambusia fish are also a threat to the introduced populations in
New Zealand. It is also possible that disease, such as a viral infection or chytrid fungus, may have contributed to the
decline of some species (W. Osborne pers. comm.). Chytrid fungus was detected in this species in Mount Compass,
South Australia, but was first identified in New Zealand from populations of this species in Christchurch. The drainage
of wetlands in Tasmania is a particular threat.
Conservation Measures This species is protected by state legislation. As an introduced species in New Zealand
it has the potential to spread chytrid to areas inhabited by native frogs. Its range includes several protected areas.
In Tasmanian legislation the species is listed as “Vulnerable”. Werribee Open Range Zoo is supporting ongoing field
monitoring of a naturally occurring population, in conjunction with the Victorian Department of Sustainability and
Environment. Melbourne Zoo also maintains a breeding population in captivity.
Bibliography: Barker, J., Grigg, G. and Tyler, M. (1995), Bell, B.D. (1982a), Bell, B.D. (1982b), Brook, A.J. (1979), Courtice, G.P. and Grigg,
G.C. (1975), Ehmann, H. and White, A. (1997), Gill, B.J., Whitaker, A.H. (1996), Gillespie, G.R. and Hero, J.M. (1999), Gillespie, G.R.,
Osborne, W.S. and McElhinney, N.A. (1995), Goldingay, R. and Lewis, B. (1999), Hero, J.-M., Littlejohn, M. and Marantelli, G. (1991),
Littlejohn, M.J. (1982), Mahony, M. (1999), Morgan, L.A. and Buttermer, W.A. (1996), Osborne, W.S., Littlejohn, M.J. and Thomson, S.A.
(1996), Pyke, G. (2002), Tyler, M.J. (1978), Tyler, M.J. (1992), Tyler, M.J. (1997), van de Mortel, T.F. and Buttermer, W.A. (1996), Waldman,
B. et al. (2001), Webb, C.E. and Joss, J. (1997)
Data Providers: Jean-Marc Hero, Graeme Gillespie, Frank Lemckert, Murray Littlejohn, Peter Robertson, Raymond Brereton, Peter
Brown
 Globally Threatened Amphibian Species
EN Litoria rheocola Liem, 1974
Endangered A2ae
Order, Family: Anura, Hylidae
Country Distribution: Australia
Current Population Trend: Decreasing
© Stephen Richards
Geographic Range This species is endemic to the Wet Tropics Bioregion (Williams and Hero 1998, 2001) and occurs
from Broadwater Creek National Park to Amos Bay, north Queensland, at altitudes between sea level and 1,180m
asl (McDonald 1992). Three genetic lineages of Litoria rheocola have been identified, based on mitochondrial DNA,
distributed from Kirrama Range to Palmerston National Park; Bartle Frere to Harris Peak; and from Mt Lewis to Big
Tableland (Schneider, Cunningham and Moritz 1998). Each of these lineages carries substantial genetic variability
(Schneider, Cunningham and Moritz 1998).
Population This species was first noted to have declined in 1989 (Richards, McDonald and Alford 1993). In 1990
several sites were sampled between the Kirrama Range and Cooktown. L. rheocola was common at all foothill and
lowland sites and was recorded at some upland sites in the Kirrama Range in April and on the Carbine Tableland
in January of that year (Richards, McDonald and Alford 1993). It was abundant in Danbulla State Forest (at 700m
asl) in September 1982, but was not recorded there during monitoring between 1989 and 1992 (Richards, McDonald
and Alford 1993). It has since disappeared from most upland sites south of the Daintree River (Richards, McDonald
and Alford 1993). Richards, McDonald and Alford (1993) reported only two adults at Bobbin Falls on the Atherton
Tableland, although the species has been found regularly in that area between 1998 and February 2000 (R. Retallick
pers. comm.) and 2001-2002 (K.R. McDonald pers. comm.). At O’Keefe Creek, Big Tableland, this species has oc-
casionally reappeared near a 400m asl site, but has not established resident populations and is absent at another
monitoring site at600m asl (McDonald and Alford 1999). Adults and larvae remained common at upland sites north
of the Daintree River (Richards, McDonald and Alford 1993) but disappeared in 1993 (M. Cunningham pers. comm.).
Interestingly, the lowland populations still exist (McDonald and Alford 1999).
CR Litoria spenceri Dubois, 1984
Critically Endangered B2ab(ii,iii,iv,v)
Order, Family: Anura, Hylidae
Country Distribution: Australia
Current Population Trend: Decreasing
© Graeme Gillespie
Geographic Range This species, an Australian endemic, is restricted predominantly to the western fall of the Great
Divide, from Lake Eildon in the Central Highlands of Victoria to Mount Kosciuszko in New South Wales, at altitudes
of 200-1,100m asl (Gillespie and Hollis 1996; Victorian Wildlife Atlas unpublished data). The area of occupancy of
this species is believed to be less than 5km².
Population Despite extensive systematic surveys throughout eastern Victoria and southern New South Wales (Watson
et al. 1991; Gillespie 1992; Gillespie and Hollis 1996; Hunter and Gillespie 1999) this species has only ever been found
in 19 streams, and has always been considered rare (Watson et al. 1991). The species is now believed to be extinct in
four of these streams, and has declined substantially in distribution and abundance along most others (Gillespie and
Hollis 1996). The remaining streams comprise 12 discrete isolated populations (Gillespie and Hollis 1996; Hunter and
Gillespie 1999). Two populations are known to have died out over the past five years. Monitoring has been conducted at
eight populations since 1994 and has recently been expanded to include 15 streams (G. Gillespie pers. comm.). With the
exception of the historic site on Buffalo Creek, Mount Buffalo National Park, monitoring of sites where it is extinct ceased
in 1998 (G. Gillespie pers. comm.). Most extant adult populations have remained relatively stable since 1993; however, a
population at Bogong Creek, Mount Kosciuszko National Park suffered a precipitous decline in 1996 (Gillespie 1997). It is
at a higher elevation than other populations and was the only high-density population known prior to the decline (Gillespie
and Hollis 1996). Gillespie and Hines (1999) estimate the largest population to contain approximately 1,000-1,500 adults
in the Upper Goulburn River with the sizes of all other populations comprising less than 1,000 adults.
Habitat and Ecology This species is associated with a range of vegetation communities from montane forest at
high altitudes to wet and dry forest at moderate to low altitudes, respectively (Gillespie and Hollis 1996). The extent
of riparian forest at known localities ranges from virtually non-existent, with scattered riparian tree or shrub spe-
261
COMMON MIST FROG
Habitat and Ecology This is a rainforest specialist, restricted to fast-flowing rocky creeks and streams in rainforest
as well as wet sclerophyll forest (Liem 1974b; McDonald 1992). Within these streams they are often found in the
slower more open sections, away from waterfalls (Hodgkison and Hero 2002). Individuals can be found on rocks,
logs and vegetation in or adjacent to streams (Hero and Fickling 1994). There are differences in habitat use between
males and females of the species. Females and juveniles use streamside vegetation more frequently than males. In
contrast, males display strong fidelity to the rocky stream environment. Calling males and gravid females have been
observed throughout the year (Liem 1974b). Breeding has been observed in most months, except during cold winter
nights, and seems to reach a peak between November and March (Liem 1974b; Dennis and Trenerry 1984). Males call
from rocks or boulders in creeks or from vegetation overhanging water along streams and creeks (Liem 1974b). The
males also appear to display inter-male spacing, with males rarely found within 1m of each other, which is possibly a
territorial response to low availability of females (Hodgkison and Hero 2002). A number (46-63) of unpigmented eggs
(2.4-2.6mm in diameter) are laid in compact gelatinous clumps under rocks in water (Liem 1974b; Hero and Fickling
1996). Larvae can be found in fast-flowing sections of streams and adjacent pools in highly oxygenated water, clinging
to rocks and other substrates (Liem 1974b; Hero and Fickling 1994). Liem (1974b) described the larvae of this species
as torrent dwelling, having flattened bodies, large suctorial mouthparts and muscular tails. Richards (1992) and Hero
and Fickling (1994) also provided detailed information on the larvae of this species.
Major Threats The reason(s) for the decline of this species are unknown. Richards, McDonald and Alford (1993) reject
drought, floods, habitat destruction or pollution by pesticides, inorganic ions or heavy metals. The habitat of the species
in the Wet Tropics has been protected since 1988; therefore, habitat destruction is no longer a threat (McDonald and
Alford 1999). Current research is examining the possibility that disease, such as a viral infection or chytrid fungus,
may have contributed to the decline of this species (Berger, Speare and Hyatt 1999). In recent experiments involving
the translocation of larvae and adult frogs to sites previously occupied by the species, a clear pattern was evident
in the disease results, and nearly all of the animals found dead showed signs of chytridiomycosis (Retallick 1999,
2000, 2001). It is unknown as to whether this disease was solely responsible for the disappearance of L. rheocola at
these sites. Feral pigs are a potential cause of riparian habitat damage and adult frog mortality (Richards, McDonald
and Alford 1993). The activity of feral pigs has been recorded to have increased over the period 1989-1992 in an area
previously inhabited by L. rheocola (Richards, McDonald and Alford 1993). However, there has been very little research
into the impact of feral pigs on native frog populations (Richards, McDonald and Alford 1993).
Conservation Measures Listed as endangered in Queensland, and therefore protected by state legislation. The
habitat of the species in the Wet Tropics has been protected since 1988. A recovery plan is in place.
Bibliography: Berger, L., Speare, R. and Hyatt, A. (1999), Dennis, A. and Trenerry, M. (1984), Hero, J.-M. and Fickling, S. (1994),
Hero, J.-M. and Fickling, S. (1996), Hodgkison, S.C. and Hero, J.-M. (2002), Hodgkison, S.C. and Hero, J.-M. (2003), Liem, D.S. (1974b),
McDonald, K. and Alford, R. (1999), Retallick, R.W.R. (1999), Retallick, R.W.R. (2000), Retallick, R.W.R. (2001), Retallick, R.W.R. (2002),
Richards, S.J. (1992), Richards, S.J., McDonald, K.R. and Alford, R.A. (1993), Tyler, M.J. (1997), Williams, S.E. and Hero, J.-M. (1998),
Williams, S.E. and Hero, J.-M. (2001)
Data Providers: Jean-Marc Hero, Ross Alford, Michael Cunningham, Keith McDonald, Richard Retallick
SPOTTED TREEFROG
cies, to a dense canopy of Leptospermum spp. shading the stream (Gillespie and Hollis 1996). The species is found
almost exclusively in association with rock habitats along streams and occurs along sections of streams with steep
banks, invariably in steeply dissected country or gorges with numerous rapids and waterfalls (Gillespie and Hollis
1996). L. spenceri is restricted to riffle and cascade stream sections with exposed rock banks, resulting in a highly
patchy distribution along most streams (Gillespie and Hollis 1996). Adults and juveniles most likely remain in the
vicinity of the stream, rarely venturing far from riparian zone (Gillespie 1997). Frogs have been seen basking on rocks
mid-stream (Ehmann, Ehmann and Ehmann 1993). Adults have been found between September and May, and calling
males have been heard in October, November and early December (Hero 1990, 1991; Watson et al. 1991; Gillespie
1993) and February (G. Gillespie, in Hero, Watson and Gillespie 1995). Clutches consist of 200-1,000 eggs (Gillespie
1997). Oviposition sites are in narrow spaces beneath large river stones within the stream and eggs are hidden as they
adhere to the underside of the rock (Gillespie 2001b). The seasonal distribution of size classes of frogs and breeding
activity (Watson et al. 1991; Gillespie 1993) suggest that eggs are laid in late spring/early summer and tadpoles
reach metamorphosis in late summer/autumn (Hero, Watson and Gillespie 1995). Hero, Watson and Gillespie (1995)
described the tadpole. At lower altitudes, sexual maturity is reached 18 months and 3.5 years after metamorphosis
by males and females, respectively (Gillespie 2001a). At higher altitudes, sexual maturity is reached at least 3.5 and
4.5 years by males and females, respectively (Gillespie 1997).
Major Threats Declines occurred in the 1970s and early 1980s (Watson et al. 1991). Gillespie and Hollis (1996)
suggest that, based upon the known demography of the species, this species probably suffered population declines
over a wider area earlier in the 20th century, and possibly late in the 19th century. Human disturbances to streams,
such as gold dredging, forest roads and recreational pressures are correlated with the general pattern of decline of
this species (Gillespie and Hollis 1996). Trout species (Rainbow Trout Oncorhynchus mykiss and Brown Trout Salmo
trutta) occur throughout the geographic range and are able to exert significant predation pressure on larvae (Gil-
lespie 2001a). Trout are believed to be a major cause of population declines of this species (Gillespie 2001a). Several
moribund frogs have been located from four populations (Gillespie and Hines 1999) and recently identified as being
infected with the chytrid fungus (Berger et al. 1998; Gillespie and Hines 1999). However, the role of the fungus in the
population dynamics of this species remains to be resolved (Gillespie and Hines 1999).
Conservation Measures The range of the species is within a few protected areas. It is listed as Endangered in
Australian legislation. A monitoring programme for this species has been established, but further research is required
to determine the reasons for the observed declines. Given the possible threat of chytridiomycosis, the establishment
of a captive-breeding programme might be required, while in situ conservation measures should include control of
invasive trout species at least within protected areas.
Bibliography: Barker, J., Grigg, G. and Tyler, M. (1995), Berger, L. et al. (1998), Ehmann, H., Ehmann, J. and Ehmann, N. (1993), Gillespie,
G. and Hines, H.B. (1999), Gillespie, G.R and Hollis, G.J. (1996), Gillespie, G.R. (1992), Gillespie, G.R. (1993), Gillespie, G.R. (1997), Gillespie,
G.R. (2001a), Gillespie, G.R. (2001b), Gillespie, G.R. (2002), Gillespie, G.R. and Hero, J.M. (1999), Hero, J.-M. (1990), Hero, J.-M. (1991),
Hero, J.-M., Littlejohn, M. and Marantelli, G. (1991), Hero, J.-M., Watson, G. and Gillespie, G. (1995), Hunter, D. and Gillespie, G.R.
(1999), Tyler, M.J. (1997), Watson, G.F. et al. (1991)
Data Providers: Jean-Marc Hero, Graeme Gillespie, Peter Robertson, Murray Littlejohn, Frank Lemckert
262 Threatened Amphibians of the World

VU Litoria subglandulosa Tyler and Anstis, 1983 GLANDULAR FROG

Vulnerable B1ab(i,ii,iii)+2ab(i,ii,iii) Geographic Range This Australian endemic occurs on the eastern escarpment of the Great Dividing Range from
Order, Family: Anura, Hylidae “The Flags” near Walcha, New South Wales, in the south, to Girraween National Park, Queensland, in the north
Country Distribution: Australia (Mahoney et al. 2001). It occurs at altitudes above 600m, and possibly up to 1,400m asl.
Current Population Trend: Decreasing Population It appears that this species underwent historical declines. However, most populations were discovered
from 1990 onwards and so little historical basis exists for assessing the level of these declines. Noticeable declines
have been recorded in three populations on the coast of northern New South Wales. It is now absent from some
sites, but the status of populations at other sites is not well known.
Habitat and Ecology The species lives near slow-flowing and small streams in dry and wet sclerophyll forest,
rainforest, montane forest and heath land. It is also found in semi-cleared grazing lands. It breeds in spring and
tadpoles have tentacles around their mouths (rather than a horny beak).
Major Threats The removal of riparian vegetation, especially by grazing and timber harvesting, is a major threat.
Pollution of streams by agricultural and domestic by-products is a threat to this riparian species. Trout have also been
© Marion Anstis

released into streams that support small populations of the frog.

Conservation Measures The range of the species includes several protected areas.
Notes on taxonomy: Litoria daviesae, a newly described species, was recently separated from L. subglandulosa by Mahony et al. (2001).
Bibliography: Barker, J., Grigg, G. and Tyler, M. (1995), Gillespie, G. and Hines, H.B. (1999), Mahony, M. et al. (2001)
Data Providers: Jean-Marc Hero, Harry Hines, John Clarke, Peter Robertson

VU Litoria wisselensis (Tyler, 1968)

Vulnerable D2 Geographic Range This species is known only from several lakes in
Order, Family: Anura, Hylidae the Enarotali area of Papua, Indonesia, at about 1,700m asl.
Country Distribution: Indonesia Population It has been found to be very common within a small area,
Current Population Trend: Unknown and is very well represented in museum collections.
Habitat and Ecology It lives on the rocky shorelines of the lakes.
Major Threats No threats are known at present. However, since
it occurs in an area subject to heavy tourist activity, it might be
susceptible to disturbance and degradation of its habitat.
Conservation Measures It is not known from any protected areas.
The species requires close population monitoring given that it is
known only from around the Enarotali area. Further survey work is
necessary to establish whether or not the species occurs in habitats
away from the lake shores.
Bibliography: Tyler, M.J. (1968)
Data Providers: Stephen Richards, Hellen Kurniati

CR Megastomatohyla mixe (Duellman, 1965)

Critically Endangered B1ab(iii)+2ab(iii) Geographic Range This species is known only from Vista Her- Bibliography: Duellman, W.E. (1965), Duellman, W.E. (2001), Faivovich, J. et al. (2005)
Order, Family: Anura, Hylidae mosa town, Sierra de Juárez, north-central Oaxaca, Mexico, at Data Providers: Georgina Santos-Barrera, Luis Canseco-Márquez
Country Distribution: Mexico 1,800m asl.
Current Population Trend: Decreasing Population This is a rare species.
Habitat and Ecology It inhabits cloud forest, and occurs in mountain
streams and rocky habitats.
Major Threats The major threat is the loss of original, intact cloud
forest due to smallholder agricultural activities and logging.
Conservation Measures The range of this species does not include
any protected areas, and the protection of remaining forested habitat
in the Sierra de Juárez is urgent. This species is protected by Mexican
law under the “Special Protection” category (Pr).
Notes on taxonomy: This species was previously included in the genus Hyla
but has recently been moved to the new genus Megastomatohyla (Faivovich
et al. 2005).

EN Megastomatohyla mixomaculata (Taylor, 1950)

Endangered B1ab(iii) Geographic Range This species is known only from the Coscomatepec region in central Veracruz, Mexico, at
Order, Family: Anura, Hylidae altitudes of 900-1,500m asl.
Country Distribution: Mexico Population It is a rare species.
Current Population Trend: Decreasing Habitat and Ecology It is highly associated with epiphytic plants in cloud forest, but breeds in temporary
Society for the Study of Amphibians and Reptiles

streams.
Major Threats The major threat is habitat loss due to the conversion of original forest cover to agricultural land,
© Hylid Frogs of Middle America, 2001,

especially coffee plantations.

Conservation Measures The range of this species is not within any protected area, and protection and restoration
of cloud forest remnants is extremely urgent. Survey work is necessary to establish its current population status. It
is listed as “Threatened” (Amenazada) by the Mexican government.
Notes on taxonomy: This species was previously included in the genus Hyla but has recently been moved to the new genus
Megastomatohyla (Faivovich et al. 2005).
Bibliography: Duellman, W.E. (2001), Faivovich, J. et al. (2005)
Data Providers: Georgina Santos-Barrera, Luis Canseco-Márquez
 Globally Threatened Amphibian Species 263

EN Megastomatohyla nubicola (Duellman, 1964)

Endangered B1ab(iii) Geographic Range This species is known only from central Veracruz, Mexico, in the Huatusco region, at around
Order, Family: Anura, Hylidae 1,325m asl, although it might occur a little more widely.
Country Distribution: Mexico Population It is a rare species.
Current Population Trend: Decreasing Habitat and Ecology It inhabits cloud forest and breeds in streams.

Society for the Study of Amphibians and Reptiles

Major Threats The main threat is the transformation of cloud forest fragments to coffee plantations.
Conservation Measures It does not occur within any protected areas, and its cloud forest habitat is in urgent need

© Hylid Frogs of Middle America, 2001,

of protection. This species is listed as “Threatened” (Amenazada) by the Mexican government.
Notes on taxonomy: This species was previously included in the genus Hyla but has recently been moved to the new genus
Megastomatohyla (Faivovich et al. 2005).
Bibliography: Duellman, W.E. (2001), Faivovich, J. et al. (2005)
Data Providers: Georgina Santos-Barrera, Luis Canseco-Márquez

CR Megastomatohyla pellita (Duellman, 1968)

Critically Endangered A2ace Geographic Range This species occurs in the Sierra Madre del Sur in southern Oaxaca, north to about San Gabriel
Order, Family: Anura, Hylidae Mixtepec and Jalatengo, Mexico, at between 1,200 and 1,700m asl.
Country Distribution: Mexico Population This has always been a rare species, but it appears to have gone into serious decline, and has not seen
Current Population Trend: Decreasing recorded since the 1960s. Recent surveys to locate it have been unsuccessful, and it might now be extinct.
Society for the Study of Amphibians and Reptiles

Habitat and Ecology This species inhabits cloud forest. It generally dwells on low vegetation along streams, and
breeds in permanent streams.
© Hylid Frogs of Middle America, 2001,

Major Threats The destruction and degradation of its cloud forest habitat is a major threat to this species, and there
is high pressure from infrastructure development for human settlements, and other anthropogenic activities in this
region. However, this species has also disappeared in suitable habitat, probably due to chytridiomycosis.
Conservation Measures This species is not known to occur in any protected areas. There is a need for further
survey work in areas of suitable habiat in order to determine whether or not this species still survives in its natural
range, and to evaluate the present condition of the forested areas north to San Gabriel Mixtepec. The protection of
remaining intact habitat within this area is recommended.
Notes on taxonomy: This species was previously included in the genus Hyla but has recently been moved to the new genus
Megastomatohyla (Faivovich et al. 2005).
Bibliography: Duellman, W.E. (2001), Faivovich, J. et al. (2005), Lips, K.R. et al. (2004)
Data Providers: Georgina Santos-Barrera, Luis Canseco-Márquez

VU Nyctimystes avocalis Zweifel, 1958

Vulnerable D2 Geographic Range This species is known only from the type local- and particularly given its small range. Further survey work is also necessary to better understand the distribution
Order, Family: Anura, Hylidae ity on the east slope of Goodenough Island, in the D’Entrecasteaux (including its possible occurrence on other islands and on the mainland of New Guinea), habitat requirements, and
Country Distribution: Papua New Guinea Islands, Papua New Guinea, at about 900m asl. population status of this species.
Current Population Trend: Unknown Population The current population status is unknown. It has been Bibliography: Zweifel, R.G. (1958)
recorded on Goodenough Island at least once since it was first Data Providers: Stephen Richards, Allen Allison
described, but has not been found on neighbouring islands.
Habitat and Ecology It inhabits small streams in forest. As with
other species in the genus, this species presumably lays large eggs
in torrential, clear and rocky streams and the tadpoles presumably
have large suckers.
Major Threats The habitat on Goodenough Island is heavily impact-
ed by gardening and fires, as well as expanding human populations,
up to 300m asl, but is still intact at higher altitudes.
Conservation Measures It is not known from any protected areas.
There is a need for close population monitoring of this species, given
declines that have taken place in torrent-dwelling hylids in Australia
264 Threatened Amphibians of the World

EN Nyctimystes dayi (Günther, 1897) AUSTRALIAN LACE-LID

Endangered A2ac 1997; Hodgkinson and Hero 2002). Davies and Richards (1990) provide a detailed description of the developmental
Order, Family: Anura, Hylidae biology of the species. Males call at night from rocks and low foliage along rapidly flowing stretches of creek. The
Country Distribution: Australia competition for females may necessitate territorial behaviour in males, as they have never been found calling in a
Current Population Trend: Decreasing group or within 1m of another male (Hodgkinson and Hero 2002). Amplexus is axillary and eggs are laid in a cohesive
clump under rocks in rapidly flowing water (Czechura, Ingram and Liem 1987). One clutch collected contained 107
unpigmented eggs (egg diameter 2.3-2.6mm, capsule diameter 3.3-3.5mm, n=5) (Davies and Richards 1990). Larvae
can be found on or under rocks in fast-flowing sections of stream and show adaptations to living in torrents, such
as large suctorial mouthparts and muscular tails (Davies and Richards 1990; Fickling and Hero 1994). After hatching
they aggregate under a rock until their digestive tracts are fully formed (Davies and Richards 1990). After several
days they begin to graze on benthic algae and may drift downstream. Larvae from eggs laid in early summer complete
development in 3-4 months, while those eggs laid in late summer may overwinter and metamorphose the following
© Stephen Richards

summer (Davies and Richards 1990).

Geographic Range This species is endemic to the Wet Tropics Bioregion (Williams and Hero 1998, 2001) from
Paluma to Cooktown, north Queensland, at altitudes between sea level and 1,200m asl (McDonald 1992). The spe-
cies includes two divergent genetic lineages separated by Barron Creek-the diversity within each lineage is very low
(M. Cunningham pers. comm.).
Population Nyctimystes dayi has disappeared from upland sites throughout the Wet Tropics and was last recorded
from Mount Spec State Forest in 1990 and the Kirrama Range in 1989 (Richards, McDonald and Alford 1993; M.
Cunningham pers. comm.). Richards, McDonald and Alford (1993) noted that the species was still common at most
foothill and lowland sites and recorded adults and larvae from upland sites north of the Daintree River. These popula-
tions subsequently disappeared in 1992 and 1993 (M. Cunningham pers. comm.). At one monitoring site at O’Keefe
Creek, Big Tableland, this species has occasionally reappeared near a site at an altitude of 400m asl, but it has not
established resident populations and is absent from a monitoring site at 680m asl (McDonald and Alford 1999).
Interestingly, the lowland and foothill populations still exist (McDonald and Alford 1999).
Habitat and Ecology This species is a rainforest specialist (Czechura, Ingram and Liem 1987). In montane areas the
species prefers fast-flowing rocky streams although they also frequent slower watercourses where ample vegeta-
tion exists along the margins (Czechura, Ingram and Liem 1987). At low elevations, it favours rock soaks, narrow
ephemeral streams and rock outcrops in larger watercourses (Czechura, Ingram and Liem 1987). It may also be found
on rocks, boulders and vegetation in or adjacent to streams (Czechura, Ingram and Liem 1987). N. dayi is a spring
and summer breeder (Davies and Richards 1990) with peak breeding activity from October to April (Hero and Fickling
Major Threats The reason(s) for the decline of the species are largely unknown. Although in the past habitat
destruction may have been a factor, clearing or logging has not taken place in the Wet Tropics World Heritage Area,
where the species occurs, since 1988 (McDonald and Alford 1999). Richards, McDonald and Alford (1993) first noted
a decline of the species in pristine rainforest habitats in 1989. Richards, McDonald and Alford (1993) reject drought,
floods, habitat destruction or changes in water quality (pH, conductivity, temperature, dissolved oxygen and metal
ions) as the primary causes of the decline. However, in a study by Hodgkinson and Hero (2002) cold temperatures
were found to reduce the activity of N. dayi and it was suggested that prolonged exposure to cold, dry conditions
(particularly at high altitudes) may inhibit the breeding and survival of the frogs and influence recruitment in local
populations. Current research is examining the possibility that disease, possibly a virus or chytrid fungus, may have
contributed to the decline of this species (Berger,Speare and Hyatt 1999; McDonald and Alford 1999). Chytrid fungus
was detected in this species in Tully, Queensland. Feral pigs are a potential cause of riparian habitat damage and
adult frog mortality (Richards, McDonald and Alford 1993). The activity of feral pigs has been recorded to have
increased over the period 1989-1992 in an area previously inhabited by N. dayi (Richards, McDonald and Alford
1993). However, there has been very little research into the impact of feral pigs on native frog populations (Richards,
McDonald and Alford 1993).
Conservation Measures The range of the species includes several protected areas. Although a recovery plan is in
place for this species, further research into the cause of the decline of this species is necessary.
Bibliography: Berger, L., Speare, R. and Hyatt, A. (1999), Czechura, G.V., Ingram, G.J. and Liem, D.S. (1987), Davies, M. andRichards, S.J.
(1990), Hero, J.-M. and Fickling, S. (1994), Hodgkison, S.C. and Hero, J.-M. (2002), Hodgkison, S.C. and Hero, J.-M. (2003), McDonald,
K. and Alford, R. (1999), McDonald, K.R. (1992), Richards, S.J., McDonald, K.R. and Alford, R.A. (1993), Tyler, M.J. (1997), Williams, S.E.
and Hero, J.-M. (1998), Williams, S.E. and Hero, J.-M. (2001)
Data Providers: Jean-Marc Hero, Ross Alford, Michael Cunningham, Keith McDonald, Richard Retallick

VU Nyctimystes rueppelli (Boettger, 1895)

Vulnerable B1ab(iii) Geographic Range This species is known only from several loca- Bibliography: Zweifel, R.G. (1958)
Order, Family: Anura, Hylidae tions on Halmahera and Morotai Islands, in the Maluku Islands, Data Providers: Stephen Richards, David Price
Country Distribution: Indonesia Indonesia. Its elevational range is not known, but the highest point
Current Population Trend: Decreasing within its range is 1,200m asl.
Population The species was last collected on Halmahera Island
in 1989.
Habitat and Ecology There is no reliable information available,
but it probably occurs in streams in tropical rainforest. As with other
species in the genus, it presumably lays large eggs in torrential, clear
and rocky streams and the tadpoles have large suckers.
Major Threats The major threat to this species is habitat loss, fol-
lowing the widespread logging that has taken place on Halmahera
Island in the past few years.
Conservation Measures There are no protected areas within its
range. Improved habitat protection is required at sites at which this
species is known to occur. There is also a need for further survey work
to determine the current population status of this species.

EN Osteopilus crucialis (Harlan, 1826) JAMAICAN SNORING FROG

Endangered B1ab(iii) Geographic Range This species is endemic to central Jamaica. It increased management of these, and improved habitat protection, are necessary.
Order, Family: Anura, Hylidae formerly occurred on the south-western coast, at an altitudinal range Bibliography: Garrick, L.D., Sutton, R.L. and Lang, J.W. (1985), Hedges, S.B. (1993), Hedges, S.B. (1999), Hedges, S.B. (2001), Henderson,
Country Distribution: Jamaica of sea level to 1,200m asl. R.W. and Powell, R (1999), Henderson, R.W. and Powell, R. (2001), Schwartz, A. and Henderson, R.W. (1991)
Current Population Trend: Decreasing Population There has been a decline in its abundance, and this Data Providers: Blair Hedges, Susan Koenig, Byron Wilson
species is now rarely encountered or heard calling, except in pockets
of suitable habitat.
Habitat and Ecology This species is found in mesic broadleaf woods
and forests on tree trunks and in bromeliads; it requires large dead
trees. Males call from hollows in branches and bromeliads, and eggs
are laid in bromeliads. It is not found in significantly altered habitats
but can be found in regenerating forests.
Major Threats It is threatened by habitat degradation and deforesta-
tion due to agriculture, human settlements, development for tourist
activities, and logging.
Conservation Measures It occurs in several forest reserves,
but these do not guarantee the species’ long-term protection, and
 Globally Threatened Amphibian Species 265

EN Osteopilus marianae (Dunn, 1926) YELLOW BROMELIAD FROG

Endangered B1ab(iii) Geographic Range This species has a restricted and fragmented range Bibliography: Hedges, S.B. (1988b), Hedges, S.B. (1993), Hedges, S.B. (1999), Hedges, S.B. (2001), Henderson, R.W. and Powell, R
Order, Family: Anura, Hylidae in central Jamaica, where it has been recorded from 120-880m asl. (1999), Henderson, R.W. and Powell, R. (2001), Schwartz, A. and Henderson, R.W. (1991)
Country Distribution: Jamaica Population This species appears to be less common than Osteo- Data Providers: Blair Hedges, Susan Koenig, Byron Wilson
Current Population Trend: Decreasing pilus wilderi.
Habitat and Ecology It is found in bromeliads in pine and deciduous
forests. Eggs are laid in bromeliads. It was thought not to have a call
until recently as it does not have a vocal sac, but it actually does
vocalize by means of vibration. It is restricted to old growth forest,
and is not tolerant of disturbance of its forest habitat.
Major Threats Deforestation is a major threat throughout its range,
primarily for agriculture, human settlement, development for tourist
activities, and selective logging.
Conservation Measures It is known from a few forest reserves,
but these do not guarantee the species’ long-term protection, and
increased management of these reserves, and improved habitat
protection, are necessary.

EN Osteopilus pulchrilineatus (Cope, 1869) HISPANIOLAN YELLOW TREEFROG

Endangered B2ab(iii,v) Geographic Range This species has a highly fragmented distribution on Hispaniola, which suggests that it is declin-
Order, Family: Anura, Hylidae ing from a previously more uniform distribution. It has been recorded from sea level up to 1,091m asl. It is presently
Country Distribution: Dominican Republic, Haiti known to persist at only one location in the Dominican Republic.
Current Population Trend: Decreasing Population This species has always been hard to find, but even so it appears to be in decline, with many historic
populations that appear to have disappeared. It was not recorded during extensive searches of the Cordillera Central
in the Dominican Republic in suitable habitat from 1998 to 2000. It was last recorded in Haiti in 1985.
Habitat and Ecology It occurs in mesic broadleaf forests, including remnant forests. Males call in flooded pools
after heavy rain. Records from 40 years ago found the species in anthropogenic habitats such as rice fields, but recent
records are all from forest. Eggs are laid in still water where the larvae also develop.
© Eladio Fernandez 2006

Major Threats Major threats include deforestation due to agricultural expansion, infrastructure development, and
logging. Since it appears to have declined in suitable habitats, chytridiomycosis cannot be ruled out as a threat.
Conservation Measures Its range includes several protected areas in both Haiti and the Dominican Republic,
although these are not well managed for biodiversity conservation. Further survey work is necessary to determine
the current population status of this species in the wild, and to determine whether chytrid is a threat.
Bibliography: Hedges, S.B. (1993), Hedges, S.B. (1999), Hedges, S.B. (2001), Henderson, R.W. and Powell, R (1999), Henderson, R.W.
and Powell, R. (2001), Schwartz, A. and Henderson, R.W. (1991)
Data Providers: Blair Hedges, Sixto Inchaustegui, Marcelino Hernandez, Robert Powell

EN Osteopilus vastus (Cope, 1871) HISPANIOLAN GIANT TREEFROG

Endangered B2ab(iii,v) Geographic Range This species is widely, but very patchily, distributed on Hispaniola (Haiti and Dominican
Order, Family: Anura, Hylidae Republic) at an altitudinal range from sea level to 1,697m asl. It was also recently found (2001-2002) just outside
Country Distribution: Dominican Republic, Haiti Santo Domingo.
Current Population Trend: Decreasing Population This species is more common than Osteopilus pulchrilineata in the Cordillera Central. However, other than
the Cordillera Central and Santo Domingo localities, there have been very few records in the last two decades.
Habitat and Ecology It is found in mesic broadleaf forests, often along creeks and streams. Males call from trees
overhanging running water, in which eggs are deposited.
Major Threats The montane stream-dwelling habitat of this species is threatened by deforestation due to agriculture
and logging. M. Hernandez (pers. comm.) has found dead animals and animals with deformities (one animal with an
eye on its back), and declines in suitable habitat are suggestive of chytridiomycosis; indeed, chytrid was confirmed
in this species in La Vega, Arroyazo, in the Dominican Republic.
Conservation Measures The species occurs in several protected areas. Given the apparent risk of infection with
chytrid, some individuals should probably form the basis for the establishment of a captive-breeding colony.
Bibliography: Hedges, S.B. (1993), Hedges, S.B. (1999), Hedges, S.B. (2001), Henderson, R.W. (2003), Henderson, R.W. and Powell, R
(1999), Henderson, R.W. and Powell, R. (2001), Powell, R. et al. (2000), Schwartz, A. and Henderson, R.W. (1991)
Data Providers: Blair Hedges, Sixto Inchaustegui, Marcelino Hernandez, Robert Powell
© Twan Leenders

EN Osteopilus wilderi (Dunn, 1925) GREEN BROMELIAD FROG

Endangered B1ab(iii) Geographic Range This species is restricted to central Jamaica at Bibliography: Hedges, S.B. (1993), Hedges, S.B. (1999), Hedges, S.B. (2001), Henderson, R.W. and Powell, R (1999), Henderson, R.W.
Order, Family: Anura, Hylidae an altitudinal range of 120-880m asl. and Powell, R. (2001), Schwartz, A. and Henderson, R.W. (1991)
Country Distribution: Jamaica Population It is declining but is more common than Osteopilus Data Providers: Blair Hedges, Susan Koenig, Byron Wilson
Current Population Trend: Decreasing marianae.
Habitat and Ecology It is commonly found in terrestrial and arboreal
bromeliads in closed-canopy forests. The species can occur in second-
ary growth, as long as there are bromeliads (since the eggs are laid,
and larval development occurs, in bromeliads only).
Major Threats The major threat to this bromeliad-dependent
species is habitat loss due smallholder farming activites, wood
extraction, and human settlement.
Conservation Measures It occurs in several forest reserves, but
these do not guarantee the species’ long-term protection. Its range
overlaps with the Blue and John Crow Mountain National Park.
Given this species’ very specific habitat requirements, improved and
strengthened habitat protection and management is essential.
266 Threatened Amphibians of the World

CR Phyllodytes auratus (Boulenger, 1917)

Critically Endangered B1ab(iii)+2ab(iii) Geographic Range This species is known only from the type locality, the summit of Aripo, in the Northern Range, and
Order, Family: Anura, Hylidae probably also Morne Bleu Ridge, Trinidad Island, in Trinidad and Tobago. The total range of the species is estimated
Country Distribution: Trinidad and Tobago to be 10km² or less. It is known from around 940m asl.
Current Population Trend: Decreasing Population It is a rare species.
Habitat and Ecology It is found in montane rainforest and elfin woodland. It has a very specific microhabitat and
is associated with the giant epiphytic bromeliad, Glomeropitcairnia erectiflora. It breeds by larval development in
bromeliads. It is not adaptable to disturbance of its habitat.
Major Threats It is threatened by the collection of bromeliads from the forest, and the over collecting of speci-
mens.
Conservation Measures It is not known if the species is present within any protected areas, but maintenance and
protection of its habitat is urgently required.
© Dan Thornham

Bibliography: Kenny, J.S. (1969), Murphy, J.C. (1997), Parker, H.W. (1933a), Parker, H.W. (1933b)
Data Providers: Jerry Hardy

CR Phyllomedusa ayeaye (B. Lutz, 1966)

Critically Endangered B1ab(iii)+2ab(iii) Geographic Range This species is known only from the type locality (Poços de Caldas) in the state of Minas Gerais,
Order, Family: Anura, Hylidae Brazil. The type locality is around 1,400m asl.
Country Distribution: Brazil Population It is a rare species.
Current Population Trend: Decreasing Habitat and Ecology This species occurs in open areas on shrubs near pools. The egg clutch is deposited on leaves
above permanent flowing water, and when hatched the tadpoles fall into streams below.
Major Threats Habitat loss due to mining and fires, as well as pollution from mining activities, are the major
threats.
© Ana Cristina Monteiro Leonel

Conservation Measures The species is not known to occur within any protected areas, and protection of the
remaining habitat of this species is needed. Further survey work is required to determine the population status of
this species, as well as the limits of its range.
Bibliography: Cruz, C.A.G. (1982), Nascimento, L.B. (1998)
Data Providers: Ulisses Caramaschi, Carlos Alberto Gonçalves da Cruz, Raphael Lima

EN Phyllomedusa baltea Duellman and Toft, 1979

Endangered B1ab(iii)+2ab(iii) Geographic Range This species is known only from the west slope of the Serrania de Sira, Huánuco Department,
Order, Family: Anura, Hylidae Peru. It possibly occurs in the Cordillera Azul, but this requires further investigation. Its presumed altitudinal range
Country Distribution: Peru is around 1,280m asl.
Current Population Trend: Decreasing Population There is no information on the population status of this species.
Habitat and Ecology It has been recorded from low cloud forest at 1,280m asl. Otherwise, nothing further is known
about this species, though it may use temporary ponds for breeding.
Major Threats Although the restricted area that the species inhabits is relatively inaccessible, there is some habitat
disturbance due to farming and human settlement. In addition, the lower parts of the range are being selectively
logged.
Conservation Measures It is present in the Reserva Comunal El Sira (an indigenous people’s reserve). Further
© Karl-Heinz Jungfer

survey work is necessary to determine the precise limits of the distribution of this species, its current population
status in the wild, and its breeding biology.
Bibliography: Cannatella, D.C. (1982), Duellman, W.E. and Toft, C.A. (1979), Rodríguez, L.O., Cordova, J.H. and Icochea, J. (1993)
Data Providers: Ariadne Angulo, Karl-Heinz Jungfer, Robert Reynolds, Javier Icochea

EN Phyllomedusa ecuatoriana Cannatella, 1982

Endangered B1ab(iii) Geographic Range This species is known only from Agua Rica, and its habitat may be in need of formal protection to guard against encroachment. Further research is needed to
Order, Family: Anura, Hylidae south-west of Limón, on the Amazonian slopes of the Andes in determine the current population status of this species, and whether chytrid poses a genuine threat.
Country Distribution: Ecuador Morona-Santiago Province, Ecuador, at around 1,890m asl. It is Bibliography: Cannatella, D.C. (1982)
Current Population Trend: Decreasing likely to occur more widely. Data Providers: Luis A. Coloma, Santiago Ron
Population There is no information on the population status of
this species.
Habitat and Ecology It is apparently a species of montane cloud
forest, presumably breeding in slow-flowing streams, or perhaps
occasionally in still water.
Major Threats The habitat around the type locality is apparently not
very threatened at the moment, although there is some agricultural
encroachment of the forest. However, the species lives in montane
habitats where catastrophic declines have affected other frogs with
stream-dwelling larvae, probably as a result of chytridiomycosis, and
so this is a potential threat.
Conservation Measures It is not known from any protected areas,
 Globally Threatened Amphibian Species 267

CR Plectrohyla acanthodes Duellman and Campbell, 1992

Critically Endangered B1ab(iii,v) Geographic Range This species is found in the northern slopes of the Meseta Central de Chiapas, Mexico, and the
Order, Family: Anura, Hylidae Sierra de los Cuchumatanes, western Guatemala, from 1,540-2,200m asl.
Country Distribution: Guatemala, Mexico Population It was common according to Duellman (2001) but it might now be uncommon. Recent searches in
Current Population Trend: Decreasing Guatemala have failed to find the species, and it seems that this population is declining. It is known now from only

Society for the Study of Amphibians and Reptiles

one locality in Chiapas, in a fragile ecosystem supporting only a few individuals.
Habitat and Ecology This species inhabits cloud forests, and is associated with mountain streams where it

© Hylid Frogs of Middle America, 2001,

breeds.
Major Threats The expansion of agriculture and human settlements in northern Chiapas and Guatemala are a major
threat. However, the observed decrease might also be due to chytridiomycosis, which has seriously affected other
stream-breeding amphibians in the region.
Conservation Measures This species is not known to occur in any protected areas, and protection of the Chiapan
cloud forests is urgently needed to ensure the persistence of this, and other endemic, species in the region. Given the
possible threat of chytridiomycosis, recommended conservation measures might need to include the establishment of
a captive-breeding programme. Further research is needed to determine whether or not the species has been affected
by chytrid. This species is protected by Mexican law under the “Special Protection” category (Pr).
Bibliography: Duellman, W.E. (2001), Duellman, W.E. and Campbell, J.A. (1992)
Data Providers: Georgina Santos-Barrera, Luis Canseco-Márquez, Manuel Acevedo, Antonio Muñoz Alonso, Joseph Mendelson III

EN Plectrohyla arborescandens (Taylor, 1939)

Endangered B1ab(iii) Geographic Range This species is known from the Atlantic slopes of Mexico, from Tlaxcala and northern Puebla
Order, Family: Anura, Hylidae and east-central to southern Puebla, at elevations of 1,800-3,100m asl.
Country Distribution: Mexico Population It is a common species.
Current Population Trend: Decreasing Habitat and Ecology This stream-breeding amphibian requires highly humid habitats such as streams in high-eleva-
tion cloud and pine-oak forests. It also takes refuge in bromeliads.
Major Threats Disturbance and clearance of the primary forest and the desiccation of streams are the main threats
to this species. Chytridiomycosis may be a threat, as it is a high-elevation stream dweller, and declines due to the
disease have already been detected amongst other species of this genus elsewhere.
Conservation Measures The protection of forest fragments in eastern central Mexico is recommended since
no protected areas currently are included within the species’ range. Further survey work and research is needed
to determine whether chytrid poses a threat to this species. This species is protected by Mexican law under the
“Special Protection” category (Pr).
© Tim Burkhardt

Notes on taxonomy: This species was previously included in the genus Hyla but has recently been moved to the genus Plectrohyla
(Faivovich et al. 2005).
Bibliography: Duellman, W.E. (2001), Faivovich, J. et al. (2005)
Data Providers: Georgina Santos-Barrera, Luis Canseco-Márquez

CR Plectrohyla avia Stuart, 1952

Critically Endangered A3e Geographic Range This species can be found in extreme south-western Chiapas, Mexico, southward to northern
Order, Family: Anura, Hylidae Pacific Guatemala from 1,700-2,200m asl.
Country Distribution: Guatemala, Mexico Population It is uncommon in Guatemala and rare at one location in Chiapas.
Current Population Trend: Decreasing Habitat and Ecology It occurs in cloud forests and is commonly found in vegetation along mountain streams.
Society for the Study of Amphibians and Reptiles

Major Threats Deforestation and transformation of native forest to agricultural areas represents the main threat.
Chytridiomycosis is also a threat to this species, as it is a high-elevation, stream-breeder, and declines due to the
© Hylid Frogs of Middle America, 2001,

disease have already been detected amongst other species of this genus in this region.
Conservation Measures This species is not known to occur in any protected area, and protection of remaining
cloud forest habitat in southern Chiapas is urgently needed. Given the possible threat of chytridiomycosis, the es-
tablishment of a captive-breeding programme might be necessary. This species is protected by Mexican law under
the “Special Protection” category (Pr).
Bibliography: Duellman, W.E. (2001)
Data Providers: Georgina Santos-Barrera, Luis Canseco-Márquez, Manuel Acevedo, Antonio Muñoz Alonso, Joseph Mendelson III

CR Plectrohyla calthula (Ustach, Mendelson, McDiarmid and Campbell, 2000)

Critically Endangered B1ab(iii,v) were collected between 1,780 and 1,896m asl, and the specimens from the second locality were collected at 1,360m
Order, Family: Anura, Hylidae asl. The exact limits of its distribution are not known, but it is believed to have a restricted distribution.
Country Distribution: Mexico Population It is known only from the type series, and two other specimens recently collected from the second
Current Population Trend: Decreasing locality.
Society for the Study of Amphibians and Reptiles

Habitat and Ecology It inhabits tropical cloud forest. The adults have been found on vegetation along streams; larvae
have been found in a pool in a small stream. It is presumed to be sensitive to habitat modification.
© Hylid Frogs of Middle America, 2001,

Geographic Range This species has been recorded from the type locality of Totontepec, Sierra Mixes, and from one
other locality west of Zacatepec, very near the type locality in Oaxaca, Mexico. The specimens from the type locality
Major Threats The forest at the type locality has been cleared and no suitable habitat remains in the area. It is
unlikely that any animals now remain at the type locality (Ustach et al. 2000). The second locality is also suffering
from severe habitat fragmentation and the remaining habitat is likely to disappear in the near future. Chytridiomycosis
is also a threat to this species, as it is a high-elevation, stream-dweller, and declines due to the disease have already
been detected amongst other species of this genus.
Conservation Measures The species has not been recorded from any protected areas. Survey work is required
to determine the current population status and range of this species, and particularly to ascertain whether or not
it still survives at the type locality. In view of the threat of chytridiomycosis, ex-situ populations might need to be
established.
Notes on taxonomy: This species was previously included in the genus Hyla but has recently been moved to the genus Plectrohyla
(Faivovich et al. 2005).
Bibliography: Duellman, W.E. (2001), Faivovich, J. et al. (2005), Meik, J.M. et al. (2005), Ustach, P.C. et al. (2000)
Data Providers: Georgina Santos-Barrera, Joseph Mendelson III
268 Threatened Amphibians of the World

CR Plectrohyla calvicollina (Toal, 1994)

Critically Endangered B1ab(iii,v)+2ab(iii,v) Geographic Range This species is known only from the northern Survey work is needed to determine the current population status of this species and whether or not it still survives
Order, Family: Anura, Hylidae slope of the Cerro Pelón, Sierra de Juárez, north-central Oaxaca, in the wild. In view of the threat of chytridiomycosis, surviving individuals might need to form the basis for the
Country Distribution: Mexico Mexico at 2,500-2,700m asl. establishment of an ex-situ population.
Current Population Trend: Decreasing Population This has always been a rare species, but it appears to Notes on taxonomy: This species was previously included in the genus Hyla but has recently been moved to the genus Plectrohyla
have gone into serious decline, and has not been recorded since (Faivovich et al. 2005).
1984. Recent surveys to locate it have been unsuccessful, and it Bibliography: Duellman, W.E. (2001), Faivovich, J. et al. (2005), Lips, K.R. et al. (2004), Toal, K.R. (1994)
might now be extinct. Data Providers: Georgina Santos-Barrera, Luis Canseco-Márquez
Habitat and Ecology This stream-breeding amphibian seems to
occur in rocky streams in cloud forest areas.
Major Threats It has disappeared in suitable habitat, probably due
to chytridiomycosis. It occurs within an area in Sierra de Juárez,
Mexico, which is undergoing a rapid rate of land transformation at
present due to logging and agricultural activities.
Conservation Measures The range of this species is not within
any protected area; a plan for the protection and/or restoration of
the large forested areas of Sierra de Juárez is urgently required.

CR Plectrohyla celata (Toal III and Mendelson III, 1995)

Critically Endangered B2ab(iii,v) Geographic Range This species is known from the Sierra de Juárez, east of Oaxaca City, in south-eastern Mexico,
Order, Family: Anura, Hylidae at 2,640-2,890m asl.
Country Distribution: Mexico Population This has always been a rare species, but it appears to have gone into serious decline, and has not been
Current Population Trend: Decreasing recorded since 1984. Recent surveys to locate it have been unsuccessful, and it might now be extinct.
Society for the Study of Amphibians and Reptiles

Habitat and Ecology This species is associated exclusively with pristine streams in cloud forests.
Major Threats This species has disappeared in suitable habitat, probably due to chytridiomycosis. The fragmentation
© Hylid Frogs of Middle America, 2001,

and disturbance of the forest due to logging and other human activities, and the continuing desiccation of streams,
has probably also contributed to the disappearance of this species.
Conservation Measures The range of this species is not within any protected area. Urgent restoration and protection
of the remnants of cloud forest in the Sierra de Juárez is required. A field study is necessary to evaluate the popula-
tion status of this species, and whether or not it still survives in the wild. In view of the threat of chytridiomycosis,
surviving individuals might need to form the basis for the establishment of an ex-situ population.
Notes on taxonomy: This species was previously included in the genus Hyla but has recently been moved to the genus Plectrohyla
(Faivovich et al. 2005).
Bibliography: Duellman, W.E. (2001), Faivovich, J. et al. (2005), Lips, K.R. et al. (2004), Toal, K.R. and Mendelson III, J.R. (1995)
Data Providers: Georgina Santos-Barrera, Luis Canseco-Márquez

CR Plectrohyla cembra (Caldwell, 1974)

Critically Endangered A2ce Geographic Range This species occurs only in two locations, one on the Pacific slope of the Sierra Madre del
Order, Family: Anura, Hylidae Sur, Distrito de Pochutla, Oaxaca, Mexico (the type locality, at 2,160m asl), and one 7.5km south-east of Llano de
Country Distribution: Mexico Guadelupe, in north-west Oaxaca (2,850m asl).
Current Population Trend: Decreasing Population This species has only been collected twice, once in 1969 and again in 1993. Recent attempts to locate
Society for the Study of Amphibians and Reptiles

it have been unsuccessful.

Habitat and Ecology This species was collected in pine-oak forest. It is believed to breed in small streams.
© Hylid Frogs of Middle America, 2001,

Major Threats A major threat is the loss of pine-oak forest habitat to logging and agriculture. It might also have
been impacted by chytridiomycosis.
Conservation Measures The range of this species is not within any protected area, and the protection and/or
restoration of the original habitat of this species is required. This species is listed as “Threatened” (Amenazada) by
the Mexican government. Further survey work is required to determine the population status of this species, and
whether or not it survives in the wild; given the likely threat of chytridiomycosis, any surviving individuals might need
to form the basis for the establishment of an ex-situ population.
Notes on taxonomy: This species was previously included in the genus Hyla but has recently been moved to the genus Plectrohyla
(Faivovich et al. 2005).
Bibliography: Duellman, W.E. (2001), Faivovich, J. et al. (2005), Lips, K.R. et al. (2004), Mendelson III, J.R. and Canseco-Márquez,
L. (2002)
Data Providers: Georgina Santos-Barrera, Luis Canseco-Márquez, Joseph Mendelson III

EN Plectrohyla charadricola (Duellman, 1964)

Endangered B1ab(iii)+2ab(iii) Geographic Range This species occurs from north-western Hidalgo to the south and west to northern Puebla,
Order, Family: Anura, Hylidae Mexico. It is found from 2,000-2,300m asl.
Country Distribution: Mexico Population This species is relatively common within its restricted range.
Current Population Trend: Decreasing Habitat and Ecology It is specifically associated with epiphytic vegetation (bromeliads) in cloud and evergreen
Society for the Study of Amphibians and Reptiles

montane forest, and breeds in streams.

Major Threats The rapid rate of deforestation of the cloud and montane forests of Mexico is the main cause of the
© Hylid Frogs of Middle America, 2001,

decline of this species and others. The disappearance of the forest results in the disappearance of streams and other
water reservoirs where the species breeds. Chytridiomycosis is also a threat, as it is a high-elevation stream dweller,
and declines due to the disease have already been detected amongst other species of this genus elsewhere.
Conservation Measures Urgent protection of the cloud and montane forest habitat of this species is required. An
assessment of the population status of this species needs to be undertaken, particularly to determine whether chytrid
poses a threat. This species is listed as “Threatened” (Amenazada) by the Mexican government.
Notes on taxonomy: This species was previously included in the genus Hyla but has recently been moved to the genus Plectrohyla
(Faivovich et al. 2005).
Bibliography: Duellman, W.E. (2001), Faivovich, J. et al. (2005)
Data Providers: Georgina Santos-Barrera, Luis Canseco-Márquez
 Globally Threatened Amphibian Species 269

CR Plectrohyla chryses (Adler, 1965)

Critically Endangered A3ce Geographic Range This species occurs in Carrizal de Bravos, central-western Guerrero, Mexico. It is found from
Order, Family: Anura, Hylidae 2,340-2,600m asl.
Country Distribution: Mexico Population This is a very rare species.
Current Population Trend: Decreasing Habitat and Ecology It is associated with humid cool areas in wet pine-oak-fir cloud forest. Breeding takes place

Society for the Study of Amphibians and Reptiles

in streams.
Major Threats The disturbance and transformation of the species’ forest habitat is the major threat, mainly as a

© Hylid Frogs of Middle America, 2001,

result of logging and the continuing expansion of human settlements in the region. Chytridiomycosis might also be
a threat, since it is a high-elevation stream-dweller, and declines due to the disease have already been detected
amongst other species of this genus.
Conservation Measures Legal protection of the Natural Park in Guerrero is required since it is not in a biosphere
reserve and currently lacks any management plan. Further survey work is required to monitor the population status and
trends of this species; also, in view of the threat of chytridiomycosis, ex-situ populations might need to be established.
This species is protected by Mexican law under the “Special Protection” category (Pr).
Notes on taxonomy: This species was previously included in the genus Hyla but has recently been moved to the genus Plectrohyla
(Faivovich et al. 2005).
Bibliography: Duellman, W.E. (2001), Faivovich, J. et al. (2005), Mendelson III, J.R. and Toal III, K.R. (1996)
Data Providers: Georgina Santos-Barrera, Luis Canseco-Márquez, Joseph Mendelson III

CR Plectrohyla chrysopleura Wilson, McCranie and Cruz, 1994

Critically Endangered A2ace;B1ab(iii,v)+ Geographic Range This species occurs in only one locality, Cerro Bufalo, Quebrado de Oro, Sierra Nombre de Dios,
2ab(iii,v) Departemento Atlantida in north-central Honduras, between 930 and 1,550m asl.
Order, Family: Anura, Hylidae Population It is moderately uncommon and appears to be in decline.
Country Distribution: Honduras Habitat and Ecology A stream-breeding species, it has been found along streams in premontane wet forest.
Current Population Trend: Decreasing Major Threats Habitat destruction, due to smallholder farming activities and subsistence wood collection, and
landslides represent major threats to this species. Tadpoles have been found infected with the chytrid fungus, and
this is probably the cause of the recently observed decline.
Conservation Measures The only known locality is within Parque Nacional Pico Bonito. Further survey work is
required to monitor the population status and trends of this species; in view of the severe risk of chytridiomycosis,
ex-situ populations might need to be established.
Bibliography: McCranie, J.R. and Wilson, L.D. (2002b), Wilson, L.D., McCranie, J.R. and Cruz, G.A. (1994)
Data Providers: Gustavo Cruz, Larry David Wilson, Franklin Casteñeda
© James McCranie

CR Plectrohyla crassa (Brocchi, 1877)

Critically Endangered B1ab(iii,v) Geographic Range This species is known from Cerro San Felipe, northern Oaxaca and So la de Vega and La Cofradia,
Order, Family: Anura, Hylidae central Oaxaca, Mexico. It occurs at elevations from 1,540-2,650m asl and is currently known from only five localities.
Country Distribution: Mexico It might occur a little more widely.
Current Population Trend: Decreasing Population This is a rare species.
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Habitat and Ecology From the limited information available, it appears that this frog inhabits and breeds in lower
streams in montane cloud forest.
© Hylid Frogs of Middle America, 2001,

Major Threats Loss of the original montane forests on the slopes of northern Oaxaca due to agriculture, logging and
human settlement is a major threat. Chytridiomycosis might be a threat to this species, since it is a high-elevation
stream-dweller, and declines due to the disease have already been detected amongst other species of this genus.
Conservation Measures This species is not known to occur in any protected areas, but the protection of the Sierra
de Juárez and adjacent areas in northern Oaxaca is urgently required. In view of the risk of chytridiomycosis, the status
of this species should be closely monitored, and ex-situ populations might need to be established. It is protected by
Mexican law under the “Special Protection” category (Pr).
Notes on taxonomy: This species was previously included in the genus Hyla but has recently been moved to the genus Plectrohyla
(Faivovich et al. 2005).
Bibliography: Duellman, W.E. (2001), Faivovich, J. et al. (2005), Toal, K.R. and Mendelson III, J.R. (1995)
Data Providers: Georgina Santos-Barrera, Luis Canseco-Márquez, Joseph Mendelson III

CR Plectrohyla cyanomma (Caldwell, 1974)

Critically Endangered B1ab(iii,v)+2ab(iii,v) Geographic Range This species occurs in Cerro Pelón and Cerro Humo Chico in the Sierra de Juárez, north-eastern
Order, Family: Anura, Hylidae Oaxaca, Mexico. It is currently known from only three localities. It is found from 2,640-2,670m asl.
Country Distribution: Mexico Population This species used to be very conspicuous in its habitat. However, recent efforts to find individuals have
Current Population Trend: Decreasing been unsuccessful. It was last collected in 1980, and is possibly now extinct.
Society for the Study of Amphibians and Reptiles

Habitat and Ecology This species is known only from pristine forest, and requires the presence of low or moderate
streams at high elevations in areas covered by cloud forest. It is likely to breed in streams.
© Hylid Frogs of Middle America, 2001,

Major Threats This species has disappeared in suitable habitat, probably due to chytridiomycosis. The disturbance
and clearance of the forest in the Sierra de Juárez has probably also contributed to its disappearance.
Conservation Measures The protection and restoration of the original forest is necessary for the protection of
this species and others inhabiting the cloud forests of the Sierra de Juárez. This species is considered “Threatened”
by the Mexican government. In view of the risk of chytridiomycosis, it is a very high priority to conduct surveys to
relocate this species and determine its current population status; any surviving individuals might need to form the
basis for a captive-breeding programme.
Notes on taxonomy: This species was previously included in the genus Hyla but has recently been moved to the genus Plectrohyla
(Faivovich et al. 2005).
Bibliography: Duellman, W.E. (2001), Faivovich, J. et al. (2005), Lips, K.R. et al. (2004)
Data Providers: Georgina Santos-Barrera, Luis Canseco-Márquez, Oscar Flores-Villela
270 Threatened Amphibians of the World

EN Plectrohyla cyclada (Campbell and Duellman, 2000)

Endangered B1ab(iii) Geographic Range This species is known from the northern versant of the Sierra Juárez (at 1,600-2,180m asl), near
Order, Family: Anura, Hylidae the crest of the Sierra Juárez (Cerro Machín and Cerro San Felipe) (at 2,370 and 2,670m asl), and the Sierra Mixes
Country Distribution: Mexico (at 2,121-2,568m asl), Oaxaca, Mexico. It might occur a little more widely.
Current Population Trend: Decreasing Population The population status of this species is unknown.
Habitat and Ecology It inhabits cloud forest and pine-oak forest at high elevations. Adults may be found close to
forest streams or in arboreal bromeliads. The larvae have been found in pools in streams.
Major Threats It is presumably threatened by the conversion of forest habitat to agricultural land, and by logging.
Chytridiomycosis may also pose a threat, as it is a high-elevation stream-dwelling amphibian, and declines due to
the disease have already been detected amongst other species of this genus.
Conservation Measures It is not known if the species occurs in protected areas, but the montane pine-oak forest
habitats of Mexico in general are poorly protected. Further survey work is needed to determine the population status
of this species, and whether chytrid is a threat.
© Tim Burkhardt

Notes on taxonomy: This species was previously included in the genus Hyla but has recently been moved to the genus Plectrohyla
(Faivovich et al. 2005).
Bibliography: Campbell, J.A. and Duellman, W.E. (2000), Duellman, W.E. (2001), Faivovich, J. et al. (2005)
Data Providers: Georgina Santos-Barrera, Joseph Mendelson III

CR Plectrohyla dasypus McCranie and Wilson, 1981

Critically Endangered A2ace;B1ab(iii,v)+ Geographic Range This species is found in the Parque Nacional Cusuco on the Sierra de Omoa, Departamento de
2ab(iii,v) Cortes, north-western Honduras, between 1,410 and 1,990m asl.
Order, Family: Anura, Hylidae Population This species was previously moderately common, but populations are currently undergoing drastic
Country Distribution: Honduras declines.
Current Population Trend: Decreasing Habitat and Ecology This stream-breeding amphibian is found on low vegetation along streams, and in arboreal
bromeliads in lower montane wet forest.
Major Threats Tadpoles of this species have been found with deformed mouth parts, presumably as a result of
infection with the chytrid fungus, likely the cause of the observed decline.
Conservation Measures The entire range of this species is within Parque Nacional Cusuco. In view of the threat
of chytridiomycosis, the status of this species should be closely monitored, and ex-situ populations should be
established.
Bibliography: Duellman, W.E. (2001), McCranie, J.R. and Wilson, L.D. (1981), McCranie, J.R. and Wilson, L.D. (2002b)
© Silviu Petrovan

Data Providers: Gustavo Cruz, Larry David Wilson, Franklin Casteñeda

CR Plectrohyla ephemera (Meik, Canseco-Márquez, Smith and Campbell, 2005)

Critically Endangered A4e Geographic Range This species is known from 1,100-1,220m asl
Order, Family: Anura, Hylidae on Cerro Las Flores, a semi-isolated uplifted area that is part of the
Country Distribution: Mexico fragmented Sierra Mixe, in south-central Oaxaca, Mexico. Many
Current Population Trend: Decreasing hylids in this region have been found to be localized endemics.
Population No information is currently available on the species’
population status, but it is possible that is has undergone a serious
decline due to chytridiomycosis, and the only known population
might already be extinct.
Habitat and Ecology This species is recorded from cloud forest.
One individual was found on vegetation 1.8m above a small stream.
Tadpoles were found in pools in this stream, which contained little
flowing water at the time, and coursed through large boulders at
the bottom of a deep ravine. Tadpoles were found in several similar
streams nearby.
Major Threats The primary threat to the species is habitat loss, and
large expanses of forest are being cleared in this region. In addition,
many of the tadpoles found at Cerro Las Flores possessed deformed or missing mouthparts, which might indicate
infection with chytridiomycosis. Indeed, the specific name of this species is derived from the Greek ephemeros,
meaning shortlived, referring to the ominous observation that chytridiomycosis might be present.
Conservation Measures The species is not known from any protected areas, and protection and maintenace of
remaining habitat in the region is urgently required. It is a very high priority to conduct surveys to relocate this species
and determine its current population status and whether or not it is still extant; in view of the threat of chytridiomycosis,
any surviving individuals would need to form the basis for a captive-breeding programme.
Notes on taxonomy: This species was previously included in the genus Hyla but has recently been moved to the genus Plectrohyla
(Faivovich et al. 2005).
Bibliography: Faivovich, J. et al. (2005), Meik, J.M. et al. (2005)
Data Providers: Joseph Mendelson III

CR Plectrohyla exquisita McCranie and Wilson, 1998

Critically Endangered A3e Geographic Range This species is only found within the Parque Nacional Cusuco in the Sierra de Omoa, Departe-
Order, Family: Anura, Hylidae mento de Cortes, north-western Honduras, between 1,490 and 1,680m asl.
Country Distribution: Honduras Population It is considered moderately common within its very restricted range.
Current Population Trend: Stable Habitat and Ecology This stream-breeding frog is found on low vegetation along streams in the lower montane
Society for the Study of Amphibians and Reptiles

wet forest. It is restricted to streams throughout its life cycle.

Major Threats Natural disasters are most likely the major threats to this species. However, given the declines in some
© Hylid Frogs of Middle America, 2001,

other Plectrohyla species in Honduras due to chytridiomycosis, this must also be considered as a potential threat.
Conservation Measures The species occurs wholly within the Parque Nacional Cusuco that has been declared a
national park since 1987. In view of the severe risk of chytridiomycosis, the status of this species should be closely
monitored, and ex-situ populations might need to be established.
Bibliography: Duellman, W.E. (2001), Duellman, W.E. and Campbell, J.A. (1992), McCranie, J.R. and Wilson, L.D. (1998)
Data Providers: Gustavo Cruz, Larry David Wilson, Franklin Casteñeda, Joseph Mendelson III
 Globally Threatened Amphibian Species 271

EN Plectrohyla glandulosa (Boulenger, 1883)

Endangered B2ab(iii) Geographic Range This species is known only from the Sierra de los Cuchumatanes and the central and southern
Order, Family: Anura, Hylidae highlands of Guatemala at an altitudinal range of 2,400-3,500m asl.
Country Distribution: Guatemala Population There is no information on the population status of this species.
Current Population Trend: Decreasing Habitat and Ecology It is a stream-breeding amphibian, living in fir forest, pine-cypress forest, and montane

Society for the Study of Amphibians and Reptiles

grasslands. It is not known whether or not it can survive in lightly degraded habitats.
Major Threats The main threat is habitat destruction due to expanding agriculture and human settlement, and

© Hylid Frogs of Middle America, 2001,

extraction of wood. Chytridiomycosis is a potential threat to this species, as it is a high-elevation stream dweller,
and declines due to the disease have already been detected among congeners.
Conservation Measures It is not known from any protected areas. However, the range includes the proposed Parque
Nacional Cuchumatanes as well as the slopes of the southern Guatemalan volcanoes, which are also proposed for
protection. Further survey work is needed to determine the population status of this species, and whether chytrid
is a threat.
Notes on taxonomy: The form occurring in El Salvador and Honduras is considered a separate species, Plectrohyla psiloderma
(McCranie and Wilson 1999a).
Bibliography: Campbell, J.A. (2001), Duellman, W.E. (2001), McCranie, J.R. and Wilson, L.D. (1999a)
Data Providers: Manuel Acevedo, Eric Smith

CR Plectrohyla guatemalensis Brocchi, 1877

Critically Endangered A3e Geographic Range This species occurs on both versants from south-eastern Chiapas, Mexico, to east-central
Order, Family: Anura, Hylidae Honduras. Its altitudinal range is 950-2,600m asl.
Country Distribution: El Salvador, Guatemala, Population It is still relatively common in Honduras, but, while formerly common, it is now uncommon in Guatemala and
Honduras, Mexico El Salvador. It has not been recorded in Mexico since 1944, and recent surveys to locate it have been unsuccessful.
Current Population Trend: Decreasing Habitat and Ecology This stream-breeding species inhabits cloud forests and premontane and lower montane
Society for the Study of Amphibians and Reptiles

forest. It is associated with cascading mountain streams.

Major Threats Although it tolerates moderate habitat destruction, loss of original forest cover affects the microhabitat
© Hylid Frogs of Middle America, 2001,

of this species. In Honduras, tadpoles have been found with deformed mouthparts, indicating chytridiomycosis. This
is probably the explanation for the observed declines in the species in recent years.
Conservation Measures There is protection of the cloud forests in southern Mexico and it occurs in several parks in
Honduras, one in Guatemala, and one in El Salvador. However, given the threat of chytridiomycosis, a captive-breeding
programme might need to be established.
Bibliography: Duellman, W.E. (2001), Duellman, W.E. and Campbell, J.A. (1992), McCranie, J.R. and Wilson, L.D. (2002b)
Data Providers: Georgina Santos-Barrera, Luis Canseco-Márquez

CR Plectrohyla hartwegi Duellman, 1968

Critically Endangered A3e Geographic Range This species occurs in Chiapas, Mexico, and western Guatemala, in an altitudinal range of
Order, Family: Anura, Hylidae 1,000-2,400m asl.
Country Distribution: Guatemala, Mexico Population A recent survey in Chiapas indicated its disappearance, at least from some localities, and there have
Current Population Trend: Decreasing been no records from Mexico since 1993. A survey of Río Cafetal, Baja Verapaz, Guatemala in the Sierra de las Minas,
showed a dramatic decline of this species, most likely due to chytridiomycosis (confirmed in a tadpole of another
species at the same site). The survey also showed that the species no longer appears to be breeding at this site,
although tadpoles have been observed at nearby localities.
Habitat and Ecology This is a high-elevation cloud forest species, associated with rocky mountain streams where
it breeds.
Major Threats Although chytridiomycosis is the suspected cause of decline in Mexico and in Guatemala, deforesta-
© Ignacio De La Riva

tion and transformation of the original forested areas due to agriculture, logging and human settlement is also a
major threat to the species.
Conservation Measures This species occurs in the Reserva de la Biósfera El Triunfo, and also in Biotopo del Quetzal
and the Reserva de la Biósfera Sierra de las Minas. Protection of the cloud forested areas in southern Chiapas is
needed. In view of the severe risk of chytridiomycosis, ex-situ populations might need to be established. This species
is protected by Mexican law under the “Special Protection” category (Pr).
Bibliography: Duellman, W.E. (2001), Lips, K.R. et al. (2004), Mendelson III, J.R. et al. (2004)
Data Providers: Georgina Santos-Barrera, Luis Canseco-Márquez, Manuel Acevedo

CR Plectrohyla hazelae (Taylor, 1940)

Critically Endangered A2ace Geographic Range This species is found in Sierra de Juárez, Conservation Measures The range of this species does not include any protected area, and the implementation of
Order, Family: Anura, Hylidae north-central Oaxaca City, and Cerro San Felipe and Cerro Machin, a conservation strategy in the Sierra de Juárez is urgent. It is a very high priority to conduct surveys to relocate this
Country Distribution: Mexico Oaxaca, Mexico. It is also recorded from the Sierra Madre del Sur species and determine whether or not it survives in the wild; in view of the risk of chytridiomycosis, any surviving
Current Population Trend: Decreasing (Puerto del Gallo) in south-western Guerrero and Oaxaca. It is known individuals might need to form the basis for a captive-breeding programme. It is protected by Mexican law under
from elevations of 2,200-2,865m asl. With increased survey work, it the “Special Protection” category (Pr).
is quite likely to be found elsewhere, especially in intervening areas Notes on taxonomy: This species was previously included in the genus Hyla but has recently been moved to the genus Plectrohyla
between known sites. (Faivovich et al. 2005).
Population This was never a common species, but it appears to have Bibliography: Duellman, W.E. (2001), Faivovich, J. et al. (2005), Lips, K.R. et al. (2004)
gone into serious decline, and has not been recorded since 1984. Data Providers: Georgina Santos-Barrera, Luis Canseco-Márquez
Recent surveys at known sites to locate it have been unsuccessful.
Habitat and Ecology This stream-breeding species occurs only in
pine-oak forest at high elevations. It shows a close affinity to slow
streams and small bushes.
Major Threats This species has disappeared in suitable habitat,
probably due to chytridiomycosis. Logging of the original pine-oak
forest at Sierra de Juárez is also a major threat to this species.
272 Threatened Amphibians of the World

CR Plectrohyla ixil Stuart, 1942

Critically Endangered A3ce Geographic Range This species occurs in two disjunct populations on the Atlantic slopes of northern Chiapas,
Order, Family: Anura, Hylidae Mexico, and western Guatemala. Its altitudinal range is from 1,175-2,200m asl. Its distribution remains largely
Country Distribution: Guatemala, Mexico unknown, and the species could occur more widely.
Current Population Trend: Decreasing Population There is no information on its population status in Guatemala, but they are considered rare in Chi-
apas.
Habitat and Ecology The species occurs in montane cloud forests, and is associated with high-elevation streams
and adjacent low vegetation where it also breeds.
Major Threats The disturbance and deforestation due to human settlement and associated activities is a major
threat. Chytridiomycosis is also a threat to this species, since it is a high-elevation stream-dweller, and declines due
to the disease have already been detected amongst other species of this genus.
Conservation Measures The species is not known to occur in any protected areas, and protection of the remaining
forests in Chiapas, in particular, is urgent. In view of the threat of chytridiomycosis, the status of this species should
© Bruce Young

be closely monitored, and ex-situ populations should be established.

Bibliography: Duellman, W.E. (2001)
Data Providers: Georgina Santos-Barrera, Luis Canseco-Márquez, Manuel Acevedo, Antonio Muñoz Alonso, Joseph Mendelson III

EN Plectrohyla lacertosa Bumzahem and Smith, 1954

Endangered B1ab(iii)+2ab(iii) Geographic Range This species is known only from two localities survey work is needed to better understand the population status of this species, and whether chytrid poses a threat.
Order, Family: Anura, Hylidae in Cerro Tres Picos and Soconusco region, south-western Chiapas, This species is protected by Mexican law under the “Special Protection” category (Pr).
Country Distribution: Mexico Mexico. It is known from elevations at around 2,130m asl. Bibliography: Duellman, W.E. (2001)
Current Population Trend: Decreasing Population It is considered rare. Data Providers: Georgina Santos-Barrera, Luis Canseco-Márquez
Habitat and Ecology It is a stream-breeding amphibian that inhabits
cloud forest at high elevations.
Major Threats Transformation or deforestation of the cloud forests
in the Soconusco region is a major threat to this species. Chytridiomy-
cosis may also be a threat, as it is a high-elevation stream dweller,
and declines due to the disease have already been detected amongst
other species of this genus.
Conservation Measures The range of this species includes the
Reserva de la Biósfera La Sepultura, which still contains well-pre-
served forest areas. Regular surveillance of this area is required to
avoid illegal logging and other human activities from degrading the
forest. Protection of other forest areas in Chiapas is needed. Further

VU Plectrohyla matudai Hartweg, 1941

Vulnerable B1ab(iii) Geographic Range This species can be found on the Pacific slopes from Oaxaca and Chiapas, Mexico, to central
Order, Family: Anura, Hylidae Guatemala, at 1,500-2,000m asl.
Country Distribution: Guatemala, Mexico Population It is common to abundant in Chiapas and Guatemala.
Current Population Trend: Decreasing Habitat and Ecology This species inhabits pine-oak and cloud forests. It is closely associated with low vegetation
along small mountain streams. Reproduction occurs via larval development in streams.
Major Threats The major threat is habitat loss and disturbance due to agriculture, logging, and human settlement.
Chytridiomycosis is a potential future threat, given the impact that it has had on some other members of the genus
in the region.
Conservation Measures It is present in Reserva de la Biósfera El Triunfo in Mexico, and is presumably present in
several protected areas in Guatemala. Improved legal protection of the extensive forested areas in Los Chimalapas
is needed.
© James McCranie

Bibliography: Duellman, W.E. (2001)

Data Providers: Georgina Santos-Barrera, Luis Canseco-Márquez, Manuel Acevedo, Antonio Muñoz Alonso

EN Plectrohyla mykter (Adler and Dennis, 1972)

Endangered B1ab(iii) Geographic Range This species is known only from the Sierra Madre del Sur de Guerrero, in Omiltemi and Puerto
Order, Family: Anura, Hylidae del Gallo, Mexico, at elevations from 1,985-2,520m asl.
Country Distribution: Mexico Population This is a rare species.
Current Population Trend: Decreasing Habitat and Ecology It inhabits high-elevation cloud and pine-oak forest, in association with streams. Its breeding
Society for the Study of Amphibians and Reptiles

biology is unknown, but it might take place in streams.

Major Threats Degradation and destruction of the forest patches in Guerrero due to smallholder farming activities
© Hylid Frogs of Middle America, 2001,

and logging is a major threat to this species. Chytridiomycosis may also be a threat, since it occurs at high elevations
(and presumably breeds in streams), and declines due to the disease have already been detected amongst other
species of this genus.
Conservation Measures The range of this species does not include any protected areas, and effective formal
protection of the Omiltemi State Ecological Park is needed; the Puerto del Gallo also requires urgent protection.
Further survey work is needed to better understand the population status of this species, and whether chytrid poses
a threat. This species is listed as “Threatened” (Amenazada) by the Mexican government.
Notes on taxonomy: This species was previously included in the genus Hyla but has recently been moved to the genus Plectrohyla
(Faivovich et al. 2005).
Bibliography: Duellman, W.E. (2001), Faivovich, J. et al. (2005), Mendelson III, J.R. and Toal III, K.R. (1996)
Data Providers: Georgina Santos-Barrera, Luis Canseco-Márquez
 Globally Threatened Amphibian Species 273

CR Plectrohyla pachyderma (Taylor, 1942)

Critically Endangered B1ab(iii) Geographic Range This species is known only from the Atlantic
Order, Family: Anura, Hylidae slopes of the Sierra Madre Oriental in central Veracruz close to
Country Distribution: Mexico Teziutlán, at the border with the state of Puebla, Mexico. The type
Current Population Trend: Decreasing locality is at 1,600m asl.
Population This species has not been collected since it was first
described, and is most likely a rare species.
Habitat and Ecology It inhabits streams and bushes in cloud forest,
and presumably breeds in streams.
Major Threats There is high pressure from infrastructure develop-
ment for human settlements, and other anthropogenic activities in
this region, and alteration of the original forest cover is resulting in
the disappearance of streams and other water systems required for
the survival of this species. Chytridiomycosis might also be a threat
to this species, since it is a high-elevation stream-dweller, and
declines due to the disease have already been detected amongst
other species of this genus.
Conservation Measures This species is not known to occur within any protected area. A specific survey to evaluate
the conservation status of the cloud forest areas in the Huasteca region in Mexico is required, and a programme to
restore and protect this area is important to conserve this and other amphibian species. It is a very high priority to
relocate this species and determine its current population status; in view of the threat of chytridiomycosis, surviving
individuals might need to form the basis for the establishment of a captive-breeding programme. This species is
protected by Mexican law under the “Special Protection” category (Pr).
Notes on taxonomy: This species was previously included in the genus Hyla but has recently been moved to the genus Plectrohyla
(Faivovich et al. 2005).
Bibliography: Duellman, W.E. (2001), Faivovich, J. et al. (2005)
Data Providers: Georgina Santos-Barrera, Luis Canseco-Márquez

EN Plectrohyla pentheter (Adler, 1965)

Endangered B1ab(iii,v) Geographic Range This species is found on the Pacific versant of the Sierra Madre del Sur in Guerrero and Oaxaca,
Order, Family: Anura, Hylidae and in central Guerrero and south-western Oaxaca in fragmented areas, at elevations of 1,280-2,000m asl.
Country Distribution: Mexico Population Recent surveys suggest that this formerly common species has declined and is now rare.
Current Population Trend: Decreasing Habitat and Ecology It is a stream-breeding amphibian that inhabits cloud forest, close to low streams.
Major Threats The recently recorded declines are probably due to chytridiomycosis, although habitat loss and
disturbance due to human activities (agriculture and logging) also represents a major threat to this species.
Conservation Measures The range of this species does not include any protected areas, and protection of the
original cloud forest habitat is required. Given the apparent risk posed by chytrid, a captive-breeding programme
may need to be established.
Notes on taxonomy: This species was previously included in the genus Hyla but has recently been moved to the genus Plectrohyla
(Faivovich et al. 2005).
Bibliography: Duellman, W.E. (2001), Faivovich, J. et al. (2005), Lips, K.R. et al. (2004), Parra-Olea, G. et al. (2003)
© Tim Burkhardt

Data Providers: Georgina Santos-Barrera, Luis Canseco-Márquez, Joseph Mendelson III

CR Plectrohyla pokomchi Duellman and Campbell, 1984

Critically Endangered A4ce Geographic Range This species is known only from the Sierra de las Minas and the Sierra de Xucaneb, central and
Order, Family: Anura, Hylidae eastern Guatemala, from 1,400-1,900m asl.
Country Distribution: Guatemala Population It is an uncommon species. There is evidence of a recent dramatic decline and possible extirpation in Río
Current Population Trend: Decreasing Cafetal, Baja Verapaz, Guatemala, in the Sierra de las Minas, possibly due to chytrid (a tadpole of another species
Society for the Study of Amphibians and Reptiles

at this site was confirmed to have chytridiomycosis).

Habitat and Ecology This species inhabits cloud forests, living in vegetation near cascading streams, and breeds
© Hylid Frogs of Middle America, 2001,

in clear streams. It can survive in old secondary forest.

Major Threats The main threat is habitat loss, especially due to encroaching agriculture (in particular leather-leaf
plantations). There is also the risk of contamination or change in flow-regime of streams. As noted, chytridiomycosis is
also a threat, given the impact that it has had on some other members of this genus in Central America, and due to the
apparent disappearance of this species at a site where chytridiomycosis in another species has been confirmed.
Conservation Measures It occurs in the Reserva de la Biósfera Sierra de las Minas and Biotopo del Quetzal. In
view of the threat of chytrid, the status of this species should be very closely monitored, and ex-situ populations
should be established.
Bibliography: Campbell, J.A. (2001), Duellman, W.E. (2001), Duellman, W.E. and Campbell, J.A. (1984), Mendelson III, J.R. et al. (2004)
Data Providers: Manuel Acevedo, Eric Smith, Joseph Mendelson III

CR Plectrohyla psarosema (Campbell and Duellman, 2000)

Critically Endangered B1ab(iii) Geographic Range This species is known only from the type locality: 5.8km (by road) west of Totontepec (17° 14’
Order, Family: Anura, Hylidae 24’’N; 96° 03’ 36”W), at an elevation of 2,103m asl, on the Sierra Mixes in northern Oaxaca, Mexico. It might occur
Country Distribution: Mexico more widely, but it is believed to have a small distribution.
Current Population Trend: Decreasing Population The population status of this recently described species is not known.
Society for the Study of Amphibians and Reptiles

Habitat and Ecology It was collected in pristine cloud forest, and was found in crevices in boulders in a small
stream and on leaves of vegetation surrounding the same stream. It is presumed to reproduce by larval development
© Hylid Frogs of Middle America, 2001,

in the water.
Major Threats The type locality of this species was totally cleared and burned as far back as 1983, and the stream
was badly silted and littered with burned logs. Even if the species does occur elsewhere, chytridiomycosis is a very
real threat, since it is a high-elevation stream-dweller, and declines due to the disease have been detected amongst
other species of this genus.
Conservation Measures The species is not known to occur in any protected areas. In view of the risk of chytrid-
iomycosis, it is a very high priority to conduct surveys to relocate this species and determine whether or not it still
survives in the wild.
Notes on taxonomy: This species was previously included in the genus Hyla but has recently been moved to the genus Plectrohyla
(Faivovich et al. 2005).
Bibliography: Campbell, J.A. and Duellman, W.E. (2000), Duellman, W.E. (2001), Faivovich, J. et al. (2005)
Data Providers: Georgina Santos-Barrera
274 Threatened Amphibians of the World

EN Plectrohyla psiloderma McCranie and Wilson, 1999

Endangered B1ab(iii) Geographic Range This species is known from Parque Nacional Celaque, Montana Celaque, in the department of
Order, Family: Anura, Hylidae De Lempira, western Honduras, and from El Pital in northern El Salvador near the Honduran border. It might occur in
Country Distribution: El Salvador, Honduras Parque Nacional Pital in Honduras (on the Salvadorian border), where larvae thought to belong to this species have
Current Population Trend: Decreasing been found. Its altitudinal range is from 2,450-2,530m asl.
Population It is moderately common in Honduras; there are no recent records in El Salvador.
Habitat and Ecology It lives on low vegetation and boulders along pristine streams in lower montane moist forest.
It breeds in streams, and is not found in disturbed habitats.
Major Threats The main threat is habitat loss, mainly due to agricultural encroachment and wood extraction. Pollution
from pesticides is also suspected to be a problem. Chytridiomycosis is a potential future threat, given the impact that
it has had on some other high-elevation stream-breeding members of this genus in Central America.
Conservation Measures It occurs in Parque Nacional Celaque, and possibly also Parque Nacional Pital, both in
© James McCranie

Honduras; the Salvadorian population is not protected. Further survey work is needed to determine the status of this
species in the Salvadorian part of the range.
Bibliography: Duellman, W.E. (2001), McCranie, J.R. and Wilson, L.D. (1999a), McCranie, J.R. and Wilson, L.D. (2002b)
Data Providers: Gustavo Cruz, Larry David Wilson, Randy McCranie, Gunther Köhler

CR Plectrohyla pycnochila Rabb, 1959

Critically Endangered B1ab(iii) Geographic Range This species is known only from the Meseta should be closely monitored, and ex-situ populations might need to be established.
Order, Family: Anura, Hylidae Central of Chiapas, in northern San Cristobal de las Casas, Mexico, Bibliography: Duellman, W.E. (2001)
Country Distribution: Mexico at 2,400m asl. Data Providers: Georgina Santos-Barrera, Luis Canseco-Márquez
Current Population Trend: Decreasing Population This is a rare species.
Habitat and Ecology It occurs in pine-oak forests, presumably
inhabiting and breeding in mountain streams.
Major Threats The major threat to the species is clear-cut logging
of its forest habitat, and transformation of the forest to agricultural
land. Chytridiomycosis might also be a threat, since this is a high-
elevation stream-dwelling species, and declines due to the disease
have already been detected in other species of this genus.
Conservation Measures The range of this species does not include
any protected areas, and protection of the forested areas surround-
ing San Cristobal de las Casas is needed. This species is listed as
“Threatened” (Amenazada) by the Mexican government. In view of
the threat of chytridiomycosis, the population status of this species

CR Plectrohyla quecchi Stuart, 1942

Critically Endangered A4ce Geographic Range This species is known from the slopes of the northern highlands in Guatemala from the Sierra
Order, Family: Anura, Hylidae de los Cuchumatanes to the highlands of Alta Verapaz and the Sierra de las Minas, from 615-1,850m asl.
Country Distribution: Guatemala Population It is a common species, but there is evidence of a recent dramatic decline and possible extirpation in
Current Population Trend: Decreasing Río Cafetal, Baja Verapaz, Guatemala, in the Sierra de las Minas, possibly due to chytridiomycosis (a tadpole of
Society for the Study of Amphibians and Reptiles

another species at this site was confirmed to have the disease). However, P. quecchi is reportedly still breeding at
nearby sites.
© Hylid Frogs of Middle America, 2001,

Habitat and Ecology This stream-breeding species inhabits cloud forest, and individuals have been found among
rocks alongside streams. It is not found in degraded habitats, but does occur in old secondary forest.
Major Threats The main threat is habitat loss, especially due to encroaching agriculture (in particular leather-leaf
plantations). There is also the risk of contamination or change in flow-regime of streams. As noted, chytridiomycosis is
also a threat, given the impact that this has had on some other members of this genus in Central America, and due to the
apparent disappearance of this species at a site where chytridiomycosis in another species has been confirmed.
Conservation Measures This species occurs in the Reserva de la Biósfera Sierra de las Minas, Biotopo del Quetzal,
and the Reserva de la Biósfera Bisis Caba. In view of the threat of chytridiomycosis, the population status of this
species requires close monitoring, and animals might need to be maintained in captivity. The taxonomic status of
this species, relative to Plectrohyla sagorum, requires investigation.
Notes on taxonomy: This species could be conspecific with Plectrohyla sagorum (M. Acevedo pers. comm.).
Bibliography: Campbell, J.A. (2001), Duellman, W.E. (2001), Duellman, W.E. and Campbell, J.A. (1992), Mendelson III, J.R. et al.
(2004), Stuart, L.C. (1942)
Data Providers: Manuel Acevedo, Eric Smith, Joseph Mendelson III

EN Plectrohyla robertsorum (Taylor, 1940)

Endangered B1ab(iii)+2ab(iii) Geographic Range This species is known from the Sierra Madre Oriental, in extreme east-central Hidalgo, up to the
Order, Family: Anura, Hylidae border with the states of Puebla and Tlaxcala, Mexico. It has been recorded at 2,250-3,050m asl. It probably occurs
Country Distribution: Mexico more widely than current records suggest, especially in areas between known sites.
Current Population Trend: Decreasing Population This is thought to be a rare species. Surveys conducted in 2001 within the species’ range collected
Society for the Study of Amphibians and Reptiles

only a single specimen.

Habitat and Ecology It occurs in pine and fir forests and montane meadows, and is commonly found close to
© Hylid Frogs of Middle America, 2001,

streams, in which it breeds.

Major Threats The main threat to this species is habitat loss and degradation due to the activities of smallholder
farmers, logging, and human settlement. Chytridiomycosis may also be athreat, as it is a high-elevation stream-breeding
amphibian, and declines due to the disease have already been detected amongst other species of this genus.
Conservation Measures The range of this species includes Parque Nacional El Chico, although this was logged in
the last 10-15 years and is subject to human settlement. Expanded and strengthened protection of the high-elevation
pine and fir forests in the Sierra Madre Oriental of eastern Hidalgo is necessary, and an extensive survey to establish
the current population status of this species (and the threat posed by chytrid) is required. This species is listed as
“Threatened” (Amenazada) by the Mexican government.
Notes on taxonomy: This species was previously included in the genus Hyla but has recently been moved to the genus Plectrohyla
(Faivovich et al. 2005).
Bibliography: Duellman, W.E. (2001), Faivovich, J. et al. (2005)
Data Providers: Georgina Santos-Barrera, Luis Canseco-Márquez
 Globally Threatened Amphibian Species 275

CR Plectrohyla sabrina (Caldwell, 1974)

Critically Endangered A3ce Geographic Range This species occurs on the northern slopes of Sierra de Juárez, northern Oaxaca, Mexico, at
Order, Family: Anura, Hylidae 1,580-2,020m asl.
Country Distribution: Mexico Population There is no information on the population status of this species; however, it is probably quite rare, and
Current Population Trend: Decreasing there have only been a few individuals collected since the 1970s.

Society for the Study of Amphibians and Reptiles

Habitat and Ecology This stream-breeding species exclusively inhabits cloud forest in the highlands of the Sierra
de Juárez, and is found in moist and rocky habitats with abundant aquatic vegetation along streams.

© Hylid Frogs of Middle America, 2001,

Major Threats The transformation and clearance of original cloud forest in the region is the main threat to this species.
In southern Mexico, tadpoles have been found with damaged mouthparts, an indication of chytrid infection.
Conservation Measures The protection of the Sierra de Juárez forest fragments is extremely urgent since there
are currently no protected areas in this region. Further surveys to establish the species’ population status are also
required, and in view of the severe risk of chytridiomycosis, ex-situ populations might need to be established. This
species is listed as “Threatened” (Amenazada) by the Mexican government.
Notes on taxonomy: This species was previously included in the genus Hyla but has recently been moved to the genus Plectrohyla
(Faivovich et al. 2005).
Bibliography: Duellman, W.E. (2001), Faivovich, J. et al. (2005)
Data Providers: Georgina Santos-Barrera, Luis Canseco-Márquez, Joseph Mendelson III

EN Plectrohyla sagorum Hartweg, 1941

Endangered B1ab(iii) Geographic Range This species occurs on the Pacific slopes of the Sierra Madre del Sur de Chiapas, south-central
Order, Family: Anura, Hylidae Chiapas, Mexico, south-west to Guatemala and again in northern El Salvador (not mapped here), from 1,000-2,050m
Country Distribution: El Salvador, Guatemala, asl.
Mexico Population It is abundant in two localities in the Cuchumatanes, Guatemala. It is very rare in Mexico, where it is
Current Population Trend: Decreasing known from a single locality. Only one specimen is known from El Salvador.
Society for the Study of Amphibians and Reptiles

Habitat and Ecology This stream-breeding amphibian exists only in cloud forests.
Major Threats The main threat to the species is habitat loss due to deforestation and transformation of the original
© Hylid Frogs of Middle America, 2001,

forest habitats in both southern Chiapas and Guatemala for agriculture, logging, timber plantations, and human
settlement.Since this is a relatively high-elevation, stream-breeding species, chytridiomycosis is a potential future
threat, and has already led to declines in other species in this genus in Central America.
Conservation Measures It is present in the Reserva de la Biósfera El Triunfo (Mexico) where a well-preserved forest
still exists, but is not known from any protected areas in Guatemala, and there is a need for improved protection of
suitable habitats for this species. Further survey work is needed to determine the status of populations in Mexico
and El Salvador, and to assess the risk posed by chytrid.
Data Providers: Georgina Santos-Barrera, Luis Canseco-Márquez, Manuel Acevedo

CR Plectrohyla siopela (Duellman, 1968)

Critically Endangered B1ab(iii,v)+2ab(iii,v) Geographic Range This species is known only from the west slope of the Cofre de Perote Mountain, central
Order, Family: Anura, Hylidae Veracruz, Mexico, at 2,500-2,550m asl.
Country Distribution: Mexico Population This species used to be abundant, but it has not been collected in over 10 years despite numerous surveys
Current Population Trend: Decreasing in suitable habitats in the area from which it has been recorded.
Society for the Study of Amphibians and Reptiles

Habitat and Ecology It inhabits dry pine forest, and is found only in mountain streams in crevices and rocks.
Major Threats The major threat to this species is the disappearance of suitable stream habitat. Its apparent disap-
© Hylid Frogs of Middle America, 2001,

pearance in areas of remaining suitable habitat is suggestive of a threat such as chytridiomycosis, so this requires
investigation.
Conservation Measures The range of this species falls entirely in the Parque Nacional Cofre de Perote. In view of
the risk of chytridiomycosis, it is a very high priority to conduct surveys to determine whether or not this species still
survives; surviving individuals might need to form the basis for the establishment of an ex-situ population.
Notes on taxonomy: This species was previously included in the genus Hyla but has recently been moved to the genus Plectrohyla
(Faivovich et al. 2005).
Bibliography: Duellman, W.E. (2001), Faivovich, J. et al. (2005)
Data Providers: Georgina Santos-Barrera, Luis Canseco-Márquez

CR Plectrohyla tecunumani Duellman and Campbell, 1984

Critically Endangered B1ab(iii)+2ab(iii) Geographic Range This species is known from two localities in the Sierra de los Cuchumatanes in western
Order, Family: Anura, Hylidae Guatemala, at 3,200-3,395m asl.
Country Distribution: Guatemala Population The type series was collected in 1968. A visit to the cave where the series was collected in June 1989
Current Population Trend: Decreasing turned up no individuals. The forest around the cave had been cleared, the area was heavily grazed, and the water
Society for the Study of Amphibians and Reptiles

in the stream in the cave was heavily silted. In 1995, a tadpole was found very close to the type locality in a stream
that has subsequently dried up after the water was piped to the local community. In 2002, tadpoles were found at
© Hylid Frogs of Middle America, 2001,

another locality (Magdalena). There are few other streams in the region and most natural habitat has been severely
degraded.
Habitat and Ecology All records have been from small caves, with streams flowing out of them. It is presumed that
the adults live in the surrounding montane pine forest, which is now somewhat degraded.
Major Threats The main threats to the species, as already indicated, include forest destruction (due to expanding
farming and human settlements, as well as wood extraction) and deterioration of water quality. Chytridiomycosis is
a potential threat, given the impact that it has had on some other members of this genus in Central America.
Conservation Measures It occurs in the proposed Parque Nacional Cuchumatanes. Further survey work is required
to determine the biology and population status and trends of this species and the limits of its range. In view of the
threat of chytridiomycosis, ex-situ populations might need to be established.
Bibliography: Campbell, J.A. (2001), Duellman, W.E. (2001), Duellman, W.E. and Campbell, J.A. (1984), Duellman, W.E. and Campbell,
J.A. (1992)
Data Providers: Manuel Acevedo, Eric Smith
276 Threatened Amphibians of the World

CR Plectrohyla teuchestes Duellman and Campbell, 1992

Critically Endangered B1ab(iii)+2ab(iii) Geographic Range As currently understood, this species is known private land. In view of the threat of chytridiomycosis, it is necessary to conduct surveys to relocate this species and
Order, Family: Anura, Hylidae only from the type locality on the southern slopes of the Sierra de determine its current population status and the limits of its distribution, and possibly to initiate a captive-breeding
Country Distribution: Guatemala Xucaneb, north of the Río Polochic, Alta Verapaz, in central Guatemala programme.
Current Population Trend: Decreasing (15° 22’N; 90° 01’W), at 1,000m asl. Notes on taxonomy: The forms that occur in Honduras have been separated to form the species Plectrohyla exquisita (McCranie and
Population When discovered, a handful of adults and tadpoles were Wilson 1998).
observed at the type locality. However, recent attempts to locate the Bibliography: Campbell, J.A. (2001), Duellman, W.E. (2001), Duellman, W.E. and Campbell, J.A. (1992), McCranie, J.R. and Wilson,
species have not been successful. L.D. (1998)
Habitat and Ecology It apparently lives near, and breeds in, streams Data Providers: Manuel Acevedo, Eric Smith
and waterfalls in montane forest. It has only been recorded from
undisturbed habitats.
Major Threats The only known site is facing habitat loss as a result
of farming and wood extraction. Chytridiomycosis is a potential
threat, given the impact that it has had on some other members of
this genus in Central America.
Conservation Measures This species is not known to occur in
any protected area; the site from which it has been recorded is on

CR Plectrohyla thorectes (Adler, 1965)

Critically Endangered A2ace Geographic Range This species is known from the Sierra Madre del Sur in Oaxaca and Guerrero, Mexico, in the vicinity
Order, Family: Anura, Hylidae of San Gabriel Mixtepec, south-western Oaxaca and Puerto del Gallo, Guerrero, Mexico, at elevations of 1,600-1,900m
Country Distribution: Mexico asl. It probably occurs more widely than current records suggest, especially in areas between known sites.
Current Population Trend: Decreasing Population This was never a common species, but it appears to have gone into serious decline, and has not been
recorded since 1984. Recent surveys to locate it within known localities have been unsuccessful.
Habitat and Ecology A stream-breeding species, it occurs in pine-oak forest, and individuals are often found on
low vegetation such as ferns and bushes along mountain streams.
Major Threats Clearance of the original forest cover for conversion to agricultural land in the region is a threat to
this species. However, the species has also disappeared in suitable habitat, probably due to chytridiomycosis.
Conservation Measures The range of this species does not include any protected areas, and urgent protection of
the forested areas surrounding San Gabriel Mixtepec and Puerto del Gallo is recommended. It is a very high priority
to conduct surveys to relocate this species and determine its current population status; if the threat of chytrid proves
© Tim Burkhardt

to be real, then surviving individuals might need to form the basis for a captive-breeding programme. This species is
protected by Mexican law under the “Special Protection” category (Pr).
Notes on taxonomy: This species was previously included in the genus Hyla but has recently been moved to the genus Plectrohyla
(Faivovich et al. 2005).
Bibliography: Duellman, W.E. (2001), Faivovich, J. et al. (2005), Lips, K.R. et al. (2004)
Data Providers: Georgina Santos-Barrera, Luis Canseco-Márquez

CR Ptychohyla dendrophasma (Campbell, Smith and Acevedo, 2000)

Critically Endangered B1ab(iii) Geographic Range This species is known from a single female specimen collected in 1998 from the type locality:
Order, Family: Anura, Hylidae Quebrada Sancapech, Finca San Francisco, Huehuetenango department, Guatemala, at 1,270m asl. This locality is
Country Distribution: Guatemala in the north-western portion of the Sierra de Los Cuchumatanes.
Current Population Trend: Decreasing Population There is no information on current population status, and three subsequent trips to the type locality
Society for the Study of Amphibians and Reptiles

have not turned up any more individuals.

Habitat and Ecology It is recorded from cloud forest. There is no information on breeding, although closely related
© Hylid Frogs of Middle America, 2001,

species breed by larval development in tree cavities and in bromeliads.

Major Threats Habitat loss is a threat to this species: it was found in the only small forest fragment in the area,
which is surrounded by extensively cleared agricultural lands. Chytridiomycosis is also a potential threat, as other
species of the genus have also declined in Guatemala due to the impacts of the disease.
Conservation Measures It does not occur in any protected area. Further survey work is required to determine the
population status of this species and the limits of its range, particularly to ascertain whether or not it might survive
in forested areas outside the type locality. Given the possible threat of chytridiomycosis, surviving individuals might
need to form the basis for the establishment of an ex-situ population.
Notes on taxonomy: This species was previously included in the genus Hyla but has recently been moved to the genus Ptychohyla
(Faivovich et al. 2005).
Bibliography: Campbell, J.A., Smith, E.N. and Acevedo, M.E. (2000), Faivovich, J. et al. (2005)
Data Providers: Manuel Acevedo, Bruce Young

EN Ptychohyla erythromma (Taylor, 1937)

Endangered B1ab(iii)+2ab(iii) Geographic Range This species occurs on the Pacific slopes of the Sierra Madre del Sur de Guerrero, central Guer-
Order, Family: Anura, Hylidae rero, Mexico, at elevations from 700-950m asl.
Country Distribution: Mexico Population It is not a common species.
Current Population Trend: Decreasing Habitat and Ecology It inhabits pine-oak forest and tropical deciduous forest. It is often found on low vegetation,
Society for the Study of Amphibians and Reptiles

under logs, rocks and vegetation along streams where it breeds.

Major Threats Habitat loss as a result of smallholder farming activities and logging is the main threat to this species.
© Hylid Frogs of Middle America, 2001,

A larva has been found with keratinised mouthparts, indicative of chytridiomycosis.

Conservation Measures The range of this species does not include any protected areas, and protection and restora-
tion of forested areas in Guerrero is urgently needed. Further research and survey work is needed to determine the extent
to which chytrid poses a threat; there may beed a need to establish a captive assurance colony of this species.
Bibliography: Duellman, W.E. (2001), Lips, K.R. et al. (2004)
Data Providers: Georgina Santos-Barrera, Luis Canseco-Márquez
 Globally Threatened Amphibian Species 277

CR Ptychohyla hypomykter McCranie and Wilson, 1993

Critically Endangered A3e Geographic Range This species ranges from western Guatemala, through most of Honduras to central Nicaragua,
Order, Family: Anura, Hylidae from 620-2,070m asl.
Country Distribution: Guatemala, Honduras, Population It is still relatively common in some areas, but is in decline.
Nicaragua Habitat and Ecology It is a forest species, living on low vegetation along mountain streams where it breeds, but
Current Population Trend: Decreasing is tolerant of some habitat disturbance. However, it does require some woody vegetation cover, such as trees along
steams in pastureland.
Major Threats The main threats are habitat loss due to expanding agriculture and pasture, forest fires, and water
pollution. The recent emergence of chytridiomycosis is also a threat. Tadpoles have been found with deformed
mouthparts, and one was confirmed with chytridiomycosis in Río Cafetal, Baja Verapaz, Guatemala in the Sierra de
las Minas, where there has been an observed dramatic population decline (although the species is still present at
this site) (Mendelson et al. 2004).
© Ignacio De La Riva

Conservation Measures It occurs in a number of protected areas in its range. In view of the threat of chytridiomycosis,
the status of this species should be closely monitored, and ex-situ populations should be established.
Notes on taxonomy: Much of the scientific literature uses the name Ptychohyla spinipollex for this species. McCranie and Wilson
(1993b) explain the correct usage of the names.
Bibliography: Campbell, J.A. (2001), Duellman, W.E. (2001), Köhler, G. (2001), McCranie, J.R. and Wilson, L.D. (1993b), McCranie, J.R.
and Wilson, L.D. (2002b), Mendelson III, J.R. et al. (2004)
Data Providers: Manuel Acevedo, Gustavo Cruz, Larry David Wilson, Randy McCranie, Gunther Köhler, Joseph Mendelson III

EN Ptychohyla legleri (Taylor, 1958)

Endangered B1ab(iii) Geographic Range This species occurs on the Pacific slopes of the Cordillera de Talamanca of Costa Rica and
Order, Family: Anura, Hylidae extreme western Panama, from 880-1,524m asl (Savage 2002).
Country Distribution: Costa Rica, Panama Population It is moderately common in appropriate habitat.
Current Population Trend: Decreasing Habitat and Ecology It is a nocturnal, stream-breeding species, present in humid montane forest and usually found
along (and occasionally in) small streams and dense vegetation no more than one metre above the water.
Major Threats The major threats to the species are the alteration of water flow in streams (for instance through
hydroelectric projects), and general habitat destruction due to smallholder livestock ranching and more extensive
agriculture (crops). Chytridiomycosis is also a potential threat to this species.
Conservation Measures Although present in protected areas such as Parque Internacional La Amistad and the
Las Cruces Biological Station (the latter is part of the Reserva de la Biósfera La Amistad), the habitat in the range of
© Karl-Heinz Jungfer

this species is in need of further protection.

Bibliography: Duellman, W.E. (2001), Ibáñez, R. et al. (2000), Savage, J.M. (2002), Young, B. et al. (1999)
Data Providers: Frank Solís, Roberto Ibáñez, Gerardo Chaves, Jay Savage, César Jaramillo, Querube Fuenmayor, Brian Kubicki,
Federico Bolaños

EN Ptychohyla leonhardschultzei (Ahl, 1934)

Endangered B1ab(v) Geographic Range This species is known from the Pacific slopes of Guerrero and Oaxaca, Mexico, at elevations
Order, Family: Anura, Hylidae between 700 and 2,000m asl. It probably occurs more widely than current records suggest.
Country Distribution: Mexico Population This is not a common species.
Current Population Trend: Decreasing Habitat and Ecology It inhabits pine-oak and cloud forest, and is found in, or on, low vegetation along mountain
streams. It breeds in streams and some populations have been found surviving in coffee plantations.
Major Threats The main threat is habitat loss mainly as a result of deforestation and the planting of coffee and
other non-timber plantations. Larvae have been found in southern Mexico with keratinised mouthparts, indicative
of chytridiomycosis.
Conservation Measures The range of this species does not include any protected areas, and the protection of the
original forested areas in Guerrero and Oaxaca is urgently needed. Further research and survey work is needed to
determine the extent to which chytrid poses a threat; there may beed a need to establish a captive assurance colony
of this species. This species is protected by Mexican law under the “Special Protection” category (Pr).
© Tim Burkhardt

Bibliography: Duellman, W.E. (2001), Lips, K.R. et al. (2004)

Data Providers: Georgina Santos-Barrera, Luis Canseco-Márquez, Joseph Mendelson III

CR Ptychohyla macrotympanum Campbell and Smith, 1992

Critically Endangered B1ab(iii) Geographic Range This species is known only from the road to La Cumbre, Municipio de la Democracia, Huehu-
Order, Family: Anura, Hylidae etenango, Guatemala, from approximately 600-1,000m asl.
Country Distribution: Guatemala Population It was found to be common in 1996 within this restricted range.
Current Population Trend: Decreasing Habitat and Ecology It occurs along, and reproduces in, streams in dry forest.
Society for the Study of Amphibians and Reptiles

Major Threats Although this species does tolerate moderate levels of habitat destruction, wholesale habitat
loss caused by agricultural activities is a major threat to the species. Chytridiomycosis is also a major threat, and
© Hylid Frogs of Middle America, 2001,

declines due to the disease have already been detected amongst species of this genus in Guatemala (Mendelson
et al. 2004).
Conservation Measures It is not known to occur in any protected areas, and habitat protection is urgently needed.
In view of the severe risk of chytridiomycosis, the status of this species should be closely monitored, and ex-situ
populations might need to be established.
Notes on taxonomy: This species was removed from synonymy of Ptychohyla euthysanota by Campbell and Smith (1992).
Bibliography: Campbell, J.A. and Smith, E.N. (1992), Mendelson III, J.R. et al. (2004)
Data Providers: Manuel Acevedo, Bruce Young
278 Threatened Amphibians of the World

EN Ptychohyla panchoi Duellman and Campbell, 1982

Endangered B1ab(iii) Geographic Range This species is known from the Sierra de las Minas, the Montañas del Mico, and the Sierra
Order, Family: Anura, Hylidae Xucaneb in eastern and central Guatemala, from 100-895m asl. It probably occurs more widely.
Country Distribution: Guatemala Population It is an uncommon species.
Current Population Trend: Decreasing Habitat and Ecology This is a stream-breeding species that lives in moist tropical forest, often along streams, and

Society for the Study of Amphibians and Reptiles

is not found in degraded habitats.
Major Threats Habitat loss, due to expanding agriculture, wood extraction, and human settlement, as well as water

© Hylid Frogs of Middle America, 2001,

pollution, are the major threats to the species. Chytridiomycosis is also a major threat, and declines due to the disease
have already been detected amongst species of this genus in Guatemala (Mendelson et al. 2004).
Conservation Measures It occurs in the Reserva de la Biosfera Sierra de las Minas and Montanos del Mico Catch-
ment Reserve. Given the risk of chytridiomycosis, there may be a need to establish a captive-breeding programme
for this species.
Bibliography: Campbell, J.A. (2001), Duellman, W.E. (2001), Duellman, W.E. and Campbell, J.A. (1982), Mendelson III, J.R. et al.
(2004)
Data Providers: Manuel Acevedo, Eric Smith

EN Ptychohyla salvadorensis (Mertens,1952)

Endangered B1ab(iii) Geographic Range This species occurs in extreme south-eastern Guatemala, northern and central El Salvador,
Order, Family: Anura, Hylidae and south-western to south-central Honduras, from 1,440-2,050m asl. It probably occurs more widely than current
Country Distribution: El Salvador, Guatemala, records suggest.
Honduras Population It is relatively common.
Current Population Trend: Decreasing Habitat and Ecology This stream-breeding species lives on low vegetation along streams in premontane to lower
montane moist forest. It is tolerant of some habitat disturbance; however, it requires some woody vegetation, such
as trees along streams in pastureland.
Major Threats The main threats are habitat loss due to expanding agriculture, creation of pasturelands, and
forest fires. Chytridiomycosis is also a major threat, and declines due to the disease have already been detected
amongst species of this genus in Guatemala (Mendelson et al. 2004), and similar declines have also been observed
in Honduras.
Conservation Measures It occurs in protected areas in all three countries. Given the risk of chytridiomycosis, there
may be a need to establish a captive-breeding programme for this species.
Bibliography: Campbell, J.A. (2001), Campbell, J.A. and Smith, E.N. (1992), Duellman, W.E. (2001), McCranie, J.R. and Wilson, L.D.
(2002b), Mendelson III, J.R. et al. (2004), Mertens, R. (1952)
Data Providers: Gustavo Cruz, Larry David Wilson, Randy McCranie, Gunther Köhler, Manuel Acevedo, Joseph Mendelson III
© Twan Leenders

CR Ptychohyla sanctaecrucis Campbell and Smith, 1992

Critically Endangered B1ab(iii,v) Geographic Range This species is known only from the Sierra de Santa Cruz, eastern Guatemala, from 366-1,150m,
Order, Family: Anura, Hylidae though it might occur a little more widely.
Country Distribution: Guatemala Population It is a common species within its range.
Current Population Trend: Decreasing Habitat and Ecology It inhabits undisturbed subtropical moist forest and is found near, and breeds in, streams.
Society for the Study of Amphibians and Reptiles

Major Threats The major threats to this species include habitat loss due to expanding agriculture, wood extraction,
and human settlement, and water pollution. Chytridiomycosis is also a potential threat, and declines due to the disease
© Hylid Frogs of Middle America, 2001,

have already been detected amongst other species of this genus in Guatemala (Mendelson et al. 2004).
Conservation Measures This species has not been found in any protected area, though one has been proposed
within its range. In view of the severe risk of chytridiomycosis, the status of this species should be closely monitored,
and ex-situ populations might need to be established.
Bibliography: Campbell, J.A. (2001), Campbell, J.A. and Smith, E.N. (1992), Duellman, W.E. (2001), Mendelson III, J.R. et al. (2004)
Data Providers: Manuel Acevedo, Eric Smith

EN Ptychohyla spinipollex (K. Schmidt, 1936)

Endangered B1ab(iii)+2ab(iii) Geographic Range This species occurs in the Cordillera Nombre de Dios in the departments of Atlantida, Colon
Order, Family: Anura, Hylidae and Yoro in northern Honduras, at 160-1,580m asl.
Country Distribution: Honduras Population It is relatively common.
Current Population Trend: Decreasing Habitat and Ecology A stream-breeding species, it lives on vegetation and boulders along streams, in lowland moist
Society for the Study of Amphibians and Reptiles

forest and premontane and lower montane wet forest. It can survive in somewhat degraded forest.
Major Threats The main threats are loss of habitat due to subsistence agriculture, creation of pastureland, and
© Hylid Frogs of Middle America, 2001,

forest fires. Chytridiomycosis is also a plausible potential threat.

Conservation Measures It is found in three protected areas, namely Pico Bonito, Texiguat, and Capiro y Calentura
National Parks. Further survey work is necessary to determine the degree to which chytrid poses a threat to this
species.
Notes on taxonomy: The use of this name has been clarified by McCranie and Wilson (1993b).
Bibliography: Duellman, W.E. (2001), McCranie, J.R. and Wilson, L.D. (1993b), McCranie, J.R. and Wilson, L.D. (2002b), Schmidt, K.P.
(1936b), Stuart, L.C. (1954), Wilson, L.D. and McCranie, J.R. (1989)
Data Providers: Gustavo Cruz, Larry David Wilson, Randy McCranie
 Globally Threatened Amphibian Species 279

CR Scinax alcatraz (B. Lutz, 1973)

Critically Endangered B1ab(iii)+2ab(iii) Geographic Range This species is known only from the type and population status of this species.
Order, Family: Anura, Hylidae locality: Ilha de Alcatrazes, an island off the coast of the state of Notes on taxonomy: This species was removed from synonymy (as a subspecies) of Scinax catharinae by Peixoto (1988).
Country Distribution: Brazil São Paulo, Brazil. The maximum elevation of the island is no more Bibliography: Lutz, B. (1973a), Lutz, B. (1973b), Peixoto, O.L. (1981)
Current Population Trend: Decreasing than 100m asl. Data Providers: Miguel Trefaut Rodrigues, Carlos Alberto Gonçalves da Cruz
Population The population status of this species is unknown.
Habitat and Ecology It occurs in primary and secondary forest,
and has also been recorded from degraded forest. The breeding
habits are unknown, but it presumably breeds in water by larval
development.
Major Threats Ilha de Alcatrazes was used as a military base in
the past, but this is no longer the case. Although there currently is
no human settlement on the island, direct disturbance of the spe-
cies, and degradation of its habitat from touristic activities and fire
are ongoing threats.
Conservation Measures There are no protected areas established
on the island. Further survey work is required to determine the biology

EN Smilisca dentata (Smith, 1957)

Endangered B1ab(iii)+2ab(iii) Geographic Range This species occurs in southern Agusacalientes and extreme north-western Jalisco, Mexico, at
Order, Family: Anura, Hylidae elevations between 1,800 and 1,900m asl.
Country Distribution: Mexico Population It is a rare species.
Current Population Trend: Decreasing Habitat and Ecology It occurs only in flooded grasslands, and breeds in temporary and permanent pools.
Society for the Study of Amphibians and Reptiles

Major Threats The transformation of the original grassland habitat into agricultural land, and the subsequent pol-
lution from pesticides, are major threats to this species.
© Hylid Frogs of Middle America, 2001,

Conservation Measures It does not occur within any protected areas, and urgent protection of the original habitat
of this species is required (especially in Agusacalientes where only a small area of suitable habitat remains). This
species is listed as “Threatened” (Amenazada) by the Mexican government.
Notes on taxonomy: This species was previously included in the genus Pternohyla which was recently synonymized with Smilisca
(Faivovich et al. 2005).
Bibliography: Duellman, W.E. (2001), Faivovich, J. et al. (2005)
Data Providers: Georgina Santos-Barrera, Luis Canseco-Márquez

VU Smilisca puma (Cope, 1884)

Vulnerable B1ab(iii) Geographic Range This species can be found from the Caribbean lowlands of Costa Rica and adjacent Nicaragua,
Order, Family: Anura, Hylidae at 15-520m asl.
Country Distribution: Costa Rica, Nicaragua Population It is uncommon, but regularly recorded in Costa Rica.
Current Population Trend: Decreasing Habitat and Ecology It lives in lowland tropical rainforest, and has also been found breeding in somewhat disturbed
habitats, such as secondary growth. Males call throughout the rainy season, from shallow water and low bushes,
usually hidden in vegetation. Breeding takes place in small, shallow temporary pools or ponds (Savage 2002).
Major Threats The major threat is habitat loss and degradation due to small- and large-scale agriculture and logging.
The forests of south-eastern Nicaragua have been less altered than those on the Costa Rican side of the border.
Conservation Measures The range of this species includes several national parks and other protected areas.
Bibliography: Savage, J.M. (2002)
Data Providers: Federico Bolaños, Gerardo Chaves
© Brian Kubicki

VU Tepuihyla rimarum Ayarzaguena, Señaris and Gorzula, 1992

Vulnerable D2 Geographic Range This species is restricted to the Ptari-tepui (5° (2004), Duellman, W.E. and Yoshpa, M. (1996), Gorzula, S. and Señaris, J.C. (1998), La Marca, E. (1997)
Order, Family: Anura, Hylidae 47’N; 61° 47’W), at 2,400m asl, in Bolívar state, Venezuela. It is likely Data Providers: Celsa Señaris, Enrique La Marca
Country Distribution: Venezuela to have a genuinely restricted range.
Current Population Trend: Stable Population It is a common species.
Habitat and Ecology It is found in crevices and under slabs in
rocky areas adjacent to shrublands on the bare sandstone summit
of Ptari-tepui. Reproductive habits are unknown, but it presumably
breeds in water like other species of the genus.
Major Threats Although there are no major threats, its restricted
range makes it more susceptible to stochastic threatening pro-
cesses.
Conservation Measures It occurs in Parque Nacional Canaima.
There is a need for close population monitoring of this species, given
that it is known only from a single location.
Bibliography: Ayarzaguena, J., Señaris, J.C. and Gorzula, S. (1992a),
Ayarzaguena, J., Señaris, J.C. and Gorzula, S. (1992b), Barrio Amorós, C.L.
280 Threatened Amphibians of the World

VU Tlalocohyla godmani (Günther, 1901)

Vulnerable B1ab(iii) Geographic Range This species is known from north-eastern Puebla and adjacent Veracruz, south to south-central
Order, Family: Anura, Hylidae Veracruz, in Mexico.
Country Distribution: Mexico Population It is not common.
Current Population Trend: Decreasing Habitat and Ecology It inhabits montane forest, and breeding takes place in temporary streams.

Society for the Study of Amphibians and Reptiles

Major Threats Conversion of forest to agricultural land has resulted in the disappearance of the original habitat of
this species. The mesic montane forest is one of the most heavily impacted environments in Mexico. Pollution from

© Hylid Frogs of Middle America, 2001,

mining is also degrading the quality of the species’ habitat.
Conservation Measures The range of this species does not include any protected areas, and habitat preservation
and restoration is required. It is listed as “Threatened” by the Mexican government.
Notes on taxonomy: This species was previously included in the genus Hyla but has recently been moved to the new genus Tlalocohyla
(Faivovich et al. 2005).
Bibliography: Duellman, W.E. (2001), Faivovich, J. et al. (2005)
Data Providers: Georgina Santos-Barrera, Luis Canseco-Márquez

HYPEROLIIDAE

VU Afrixalus clarkeorum Largen, 1974

Vulnerable B1ab(iii) Geographic Range This species is endemic to Ethiopia, where it is currently known from only two areas (the vicinity
Order, Family: Anura, Hyperoliidae of Bonga, and Godare) at 820-1,800m asl, in the south-western region of the country.
Country Distribution: Ethiopia Population It appeared to be relatively common at the type locality in 1971, but its status is otherwise unknown
Current Population Trend: Decreasing due to lack of recent survey work.
Habitat and Ecology It is found in tropical deciduous forest and forest edge, breeding amongst herbaceous
vegetation surrounding pools in forest clearings. It has also been found in coffee plantations and other secondary
habitats. For breeding it requires emergent vegetation in marshy pools, at least some of which are likely to become
dry for part of the year.
Major Threats The major threat is habitat loss due to selective logging, human settlement, and agricultural encroach-
ment (including coffee plantations).
Conservation Measures It is not known from any protected areas. There is a need for improved habitat protection
at sites where this species is known to occur.
© M.J. Largen

Notes on taxonomy: Largen (2001) emended the name from Afrixalus clarkei to Afrixalus clarkeorum.
Bibliography: Largen, M.J. (1974), Largen, M.J. (2001), Schiøtz, A. (1999)
Data Providers: Malcolm Largen, Arne Schiøtz

VU Afrixalus enseticola Largen, 1974 ETHIOPIAN BANANA FROG

Vulnerable B2ab(iii) Geographic Range This species is endemic to high-altitude areas of southern Ethiopia on both sides of the Rift
Order, Family: Anura, Hyperoliidae Valley at altitudes of 1,700-2,750m asl.
Country Distribution: Ethiopia Population It is locally common within its relatively restricted range.
Current Population Trend: Decreasing Habitat and Ecology It is found in montane forest at altitudes, and rarely in montane grassland after forest clearance,
typically breeding amongst herbaceous vegetation surrounding pools in forest clearings. It can survive in suitable
situations in plantations, rural gardens, and even urban areas. For breeding it requires emergent vegetation in marshy
pools, at least some of which are likely to become dry for part of the year.
Major Threats Threats include forest clearance, human settlement, and agricultural encroachment.
Conservation Measures It is known to occur in the Bale Mountains National Park.
Bibliography: Largen, M.J. (1974), Largen, M.J. (2001), Schiøtz, A. (1999)
Data Providers: Malcolm Largen, Arne Schiøtz
© M.J. Largen

EN Afrixalus knysnae (Loveridge, 1954) KNYSNA BANANA FROG

Endangered B1ab(ii,iii,iv,v)+2ab(ii,iii,iv,v) Geographic Range This species is known from a few localities at low altitude on the south coast of South Africa
Order, Family: Anura, Hyperoliidae on either side of the border between the Eastern Cape and Western Cape Provinces.
Country Distribution: South Africa Population It is a rare species, with only a few individuals known in each locality.
Current Population Trend: Decreasing Habitat and Ecology It lives in a coastal mosaic of vegetation types, including mountain fynbos heathland, forest,
and to some extent on agricultural land. It breeds in small dams and shallow semi-permanent water with much
emergent vegetation.
Major Threats The main threat is habitat loss due to urban and recreational development, afforestation, and
© Frank Glaw and Miguel Vences

agricultural expansion.
Conservation Measures It occurs in Tsitsikamma National Park, Goukamma Nature Reserve, and Diepwalle
Forestry Area.
Notes on taxonomy: This species is closely related to Afrixalus spinifrons (Schiøtz 1999).
Bibliography: Channing, A. (2001), Minter, L.R. et al. (2004), Passmore, N.I. and Carruthers, V.C. (1995), Pickersgill, M. (1996), Pickersgill,
M. (2000), Schiøtz, A. (1999)
Data Providers: Leslie Minter, Arne Schiøtz
 Globally Threatened Amphibian Species 281

EN Afrixalus lacteus Perret, 1976

Endangered B1ab(i,iii) Geographic Range This species is known only from a few mountains in western Cameroon, specifically the western
Order, Family: Anura, Hyperoliidae and southern slopes of Mount Manenguba, Mount Nlonako, and the southern portion of the Bamileke Plateau (at
Country Distribution: Cameroon Mount Bana, Foto and Batie), at an altitude of 1,200-1,900m asl.
Current Population Trend: Decreasing Population It is generally an uncommon species.
Habitat and Ecology It lives in montane forest (including secondary forest and forest edges), and raffia palm swamps,
and is often associated with streams in which it breeds (which is unusual for a species in this genus).
Major Threats It is threatened by forest loss as a result of agricultural encroachment, expanding human settle-
ments, and removal of wood.
Conservation Measures It does not occur in any protected areas. The protected area network in western Camer-
oon urgently needs to be expanded to include the remaining montane forest habitats, particularly those on Mount
Manenguba.
© Jean-Louis Amiet

Bibliography: Amiet, J.-L. (1972d), Amiet, J.-L. (1975), Herrmann, H.-W. et al. (2005), Perret, J.-L. (1976a), Schiøtz, A. (1999)
Data Providers: Jean-Louis Amiet, Arne Schiøtz

VU Afrixalus morerei Dubois, 1986 “1985”

Vulnerable B1ab(iii) Geographic Range This species is endemic to the Udzungwa Mountains in southern Tanzania, occurring between
Order, Family: Anura, Hyperoliidae 1,300 and 2,050m asl.
Country Distribution: Tanzania Population It is moderately abundant in suitable habitats.
Current Population Trend: Decreasing Habitat and Ecology It is a species of marshy areas in open montane grasslands, and from grassy glades in for-
est/grassland mosaic. It has been found in pastureland, but it is not clear that in can survive in croplands.
Major Threats Its montane grassland habitat is threatened by afforestation, agricultural expansion, fire, and hu-
man settlement.
Conservation Measures There are no records from Udzungwa National Park, but it is likely to occur there.
Bibliography: Dubois, A. (1985), Pickersgill, M. (1992), Pickersgill, M. (1995), Poynton, J.C. (2003b), Schiøtz, A. (1974), Schiøtz, A.
(1975), Schiøtz, A. (1999)
© Michele Menegon

Data Providers: Martin Pickersgill, Arne Schiøtz, Kim Howell

VU Afrixalus orophilus (Laurent, 1947)

Vulnerable B1ab(iii) Geographic Range This species occurs in Kivu Province in eastern Democratic Republic of Congo, western
Order, Family: Anura, Hyperoliidae Rwanda, western Burundi, and south-western Uganda, probably north to Mount Ruwenzori. It is normally found
Country Distribution: Burundi, Congo, D.R., above 1,500m asl.
Rwanda, Uganda Population It is not a very abundant species.
Current Population Trend: Decreasing Habitat and Ecology Its habitat is not well known, but it has been found in montane grassland, bamboo forests,
and in wetland areas in reeds and papyrus. It breeds in marshy areas.
Major Threats Little information is available, though it is likely to be impacted by loss of habitat for agriculture
and human settlements.
Conservation Measures It has been recorded from Virungas National Park in the Democratic Republic of Congo
and Bwindi National Park in Uganda.
Bibliography: Drewes, R.C. and Vindum, J.V. (1994), Laurent, R.F. (1972), Laurent, R.F. (1982), Schiøtz, A. (1999), Vonesh, J. (2001)
Data Providers: Robert Drewes, Arne Schiøtz
© R.C. Drewes

VU Afrixalus spinifrons (Cope, 1862) NATAL BANANA FROG

Vulnerable B2ab(ii,iii,iv,v) Geographic Range This species, which is endemic to South Africa, occurs as two subspecies: A. s. spinifrons oc-
Order, Family: Anura, Hyperoliidae curs in the KwaZulu-Natal lowlands, and eastern Cape coast of South Africa at low to intermediate altitudes; A.s.
Country Distribution: South Africa intermedius occurs at altitudes above 1,000m asl in western KwaZulu-Natal.
Current Population Trend: Decreasing Population There have been relatively few records in recent years, perhaps because of a decline, but this species is
somewhat hard to detect. It certainly seems to have disappeared from some localities, and many surviving populations
are small and isolated. However, in other places, high-density choruses are common.
Habitat and Ecology It is associated with low vegetation in shrubland, dry forest and to some extent agricultural
land. It breeds in vleis and temporary pools and pans (including roadside pools).
Major Threats Certain populations are affected by urban and recreational development, afforestation, agricultural
expansion, pesticides, and overgrazing by livestock.
Conservation Measures Afrixalus spinifrons intermedius occurs in the Khahlamba-Drakensberg National Park.
© Richard Boycott

Afrixalus s. spinifrons occurs in a number of coastal protected areas.

Notes on taxonomy: We follow Pickersgill (1996) in considering what were formerly regarded as the eastern populations of Afrixalus
knysnae to be a subspecies of A. spinifrons (A. s. intermedius).
Bibliography: Alexander, G.J. (1990), Armstrong, A. (2001), Backwell, P.R.Y. and Passmore, N.I. (1991), Channing, A. (2001), Lambiris,
A.J.L. (1989a), Minter, L.R. et al. (2004), Passmore, N.I. and Carruthers, V.C. (1995), Pickersgill, M. (1984), Pickersgill, M. (1996), Pickersgill,
M. (2000), Poynton, J.C. (1964b), Schiøtz, A. (1999)
Data Providers: Leslie Minter, Arne Schiøtz, Martin Pickersgill
282 Threatened Amphibians of the World

EN Afrixalus sylvaticus Schiøtz, 1974

Endangered B2ab(iii) Geographic Range This species ranges from the Shimba Hills in southern coastal Kenya south through the East
Order, Family: Anura, Hyperoliidae Usambara foothills in north-eastern Tanzania, as far south as the Kazizumbwi Forest in central coastal Tanzania. There
Country Distribution: Kenya, Tanzania is an unconfirmed record from further north along the Tana River in eastern Kenya. It occurs only very patchily within
Current Population Trend: Decreasing the mapped range due to limited suitable habitat.
Population It is fairly abundant where it occurs.
Habitat and Ecology It is a species of lowland forest that can survive in secondary growth and plantations, but not
in completely degraded habitats. It breeds in temporary pools and water-filled depressions in forest.
Major Threats Although it can tolerate degraded forest and secondary growth, these habitats are also known to
be suitable for A. stuhlmanni with which this species hybridizes, and this is probably the most serious threat to the
species.
Conservation Measures It occurs in the Shimba Hills National Park, but the maintenance and protection of other
tracts of lowland coastal forest habitat is essential to ensure the persistence of this species.
© Arne Schiøtz

Notes on taxonomy: Pickersgill (2000, 2005) considers this form to be a subspecies of Afrixalus stuhlmanni, with which it hybridizes
as forests are cleared. We follow Schiøtz (1999, and pers. comm.) in considering it to be a valid species. However, we follow Pickersgill
(2005) and K. Howell (pers. comm.) in considering it to occur in the coastal areas of Tanzania, as well as in the Shimba Hills in Kenya.
Bibliography: Howell, K.M. (1993), Pickersgill, M. (2005), Schiøtz, A. (1974), Schiøtz, A. (1975), Schiøtz, A. (1999)
Data Providers: Martin Pickersgill, Arne Schiøtz, Kim Howell

VU Afrixalus uluguruensis (Barbour and Loveridge, 1928)

Vulnerable B1ab(iii) Geographic Range This species occurs in the Usambara, Nguu, Uluguru and Udzungwa mountains in Tanzania. Its
Order, Family: Anura, Hyperoliidae altitudinal range is not fully known, but it is a species of medium to high altitudes. Records from the coastal plain
Country Distribution: Tanzania of Tanzania require confirmation.
Current Population Trend: Decreasing Population It is abundant where it occurs.
Habitat and Ecology It is a forest-dependent species, and does not survive in degraded habitats. It breeds in swampy
valley bottoms and temporary pools in closed-canopy forest.
Major Threats The main threat is habitat loss due to agricultural encroachment, logging, and expanding human
settlements. Its habitat in the East Usambara Mountains has recently come under serious threat as a result of the
activities of illegal gold miners.
Conservation Measures It occurs in the Amani Nature Reserve. It might occur in the Udzungwa National Park, but
there have not yet been any confirmed records.
© Elizabeth Harper

Bibliography: Drewes, R.C., Altig, R. and Howell, K.M. (1989), Harper, E. and Vonesh, J.R. (2003), Howell, K.M. (1993), Schiøtz,
A. (1975), Schiøtz, A. (1999)
Data Providers: Arne Schiøtz, Kim Howell, Martin Pickersgill, Michele Menegon

CR Alexteroon jynx Amiet, 2000

Critically Endangered B1ab(iii)+2ab(iii) Geographic Range This recently described species is known only from two localities at 800-1,050m asl, 6km apart
Order, Family: Anura, Hyperoliidae near the villages of Betenge and Mofako on the eastern slopes of the Rumpi Hills in south-western Cameroon. It might
Country Distribution: Cameroon occur a little more widely, but appears to be absent from other sites that have been explored nearby.
Current Population Trend: Decreasing Population There is no available information on its population status.
Habitat and Ecology It is associated with narrow streams in forests, over which there is a continuous canopy. It
favours well aerated, flowing water, but avoids torrents. Males call close to streams in dense vegetation, and they
presumably breed in a similar manner to Alesteroon obstetricans (i.e., small clumps of eggs are laid on leaves above
streams, into which the larvae fall, and develop).
Major Threats Severe forest loss is taking place in the Rumpi Hills due to smallholder farming activities, logging,
and human settlement.
Conservation Measures It is not known whether or not this species occurs in any protected areas, though it might
© Jean-Louis Amiet

well occur in the Rumpi Hills Forest Reserve; habitat protection is a matter of urgent priority for this species. Survey
work is also required to determine the current population status of this species and the limits of its range.
Bibliography: Amiet, J.-L. (2000)
Data Providers: Jean-Louis Amiet, Arne Schiøtz

EN Arlequinus krebsi (Mertens, 1938)

Endangered B1ab(iii) Geographic Range This species is known from only two localities in western Cameroon: Mubenge on the southern
Order, Family: Anura, Hyperoliidae slopes of Mount Cameroon, where it is known only from the type specimen; and Nkondjock at 750m asl on the
Country Distribution: Cameroon southern slopes of the Bamileke Plateau, where it is known from a small number of specimens and a handful of field
Current Population Trend: Decreasing observations. It presumably occurs more widely, but it has not been found elsewhere despite extensive fieldwork
in the region.
Population This species is poorly known, but it seems to be rare. It is probably mute, and this makes it very hard
to find.
Habitat and Ecology It has been found in dense forest where the eggs are placed above small pools with stagnant
or slowly flowing water, and the larvae develop in these pools. It is not known whether or not it can survive in
secondary habitats.
Major Threats The major threat is likely to be ongoing forest loss due to subsistence wood cutting and agriculture,
© Jean-Louis Amiet

and human settlement.

Conservation Measures It is not known to occur in any protected areas. The protected area network in western
Cameroon urgently needs to be expanded to include remaining montane forest habitats, such as those on Mount
Cameroon.
Bibliography: Amiet, J.-L. (1972d), Amiet, J.-L. (1976a), Perret, J.-L. (1988b), Schiøtz, A. (1999)
Data Providers: Jean-Louis Amiet, Arne Schiøtz

VU Callixalus pictus Laurent, 1950
Vulnerable B1ab(iii)
Order, Family: Anura, Hyperoliidae
Country Distribution: Congo, D.R., Rwanda
Current Population Trend: Decreasing
EN Hyperolius bobirensis Schiøtz, 1967
Endangered B1ab(iii)
Order, Family: Anura, Hyperoliidae
Country Distribution: Ghana
Current Population Trend: Decreasing
© Annika Hillers
VU Hyperolius castaneus Ahl, 1931
Vulnerable B1ab(iii)
Order, Family: Anura, Hyperoliidae
Country Distribution: Burundi, Congo, D.R.,
Rwanda, Uganda
Current Population Trend: Decreasing
© R.C. Drewes
VU Hyperolius chrysogaster Laurent, 1950
Vulnerable B1ab(iii)
Order, Family: Anura, Hyperoliidae
Country Distribution: Congo, D.R.
Current Population Trend: Decreasing
Globally Threatened Amphibian Species 283
Geographic Range This species is known only from the Itombwe Conservation Measures It is not known from any protected areas. There is a need for improved habitat protection
and Kabobo highlands of eastern Democratic Republic of Congo, and at sites where this species is known to occur, such as on the Itombwe massif. Further survey work is required to
from the mountains of western Rwanda. It has never been found determine the current population status of the species.
below 2,100m, and is rare below 2,400m asl. There have been no Bibliography: Laurent, R.F. (1950), Laurent, R.F. (1964), Schiøtz, A. (1999)
recent records of this species, presumably due to lack of fieldwork Data Providers: Arne Schiøtz
within its range.
Population There is no recent information on the population status
of this species.
Habitat and Ecology It is a species of high-altitude forests, espe-
cially bamboo forest. By day, it hides in broken bamboo stems at 2-4m
above the ground, or, where bamboo is scarce, it hides between bark
and moss, which covers the tree trunks. There is no information on
its breeding, but it is presumed to breed in water.
Major Threats There is no direct information, but it is restricted to
unprotected habitats that are probably being lost due to agriculture,
wood extraction, and expanding human settlements.
Geographic Range This species is known from the Bobiri Forest Reserve in south-western Ghana and from the
Ankasa Conservation Area (ACA), a twin wildlife protected area comprised of Nini-Suhien National Park to the north
and the Ankasa Forest Reserve to the south, also in Ghana. It is presumed to occur in other forests in Ghana, but it
cannot range much further to the west, where Hyperolius zonatus occurs.
Population It is very common at the type locality (Bobiri Forest Reserve).
Habitat and Ecology It is confined to lowland forest, where it is associated with temporary, stagnant pools in which
it breeds. The eggs are laid on leaves above water, into which the larvae fall and develop.
Major Threats It is presumably threatened by ongoing habitat loss due to logging, agriculture and human settle-
ments.
Conservation Measures The species occurs in the Bobiri Forest Reserve (although it is not clear how well protected
this is) as well as the Ankasa Conservation Area.
Bibliography: Rödel, M.-O. et al. (2005), Schiøtz, A. (1967), Schiøtz, A. (1999)
Data Providers: Arne Schiøtz, Mark-Oliver Rödel
Geographic Range This species occurs in eastern Democratic Republic of Congo, western Rwanda, western Burundi,
and south-western Uganda. It occurs between 1,600 and 2,850m asl.
Population It is an abundant species.
Habitat and Ecology It inhabits and breeds in swamps in montane grassland and forest.
Major Threats Little information is available, though it is likely to be impacted by loss of habitat for agriculture
(crops and livestock) and human settlements.
Conservation Measures It is present in Bwindi National Park (Uganda) and the Virungas National Park (Democratic
Republic of Congo).
Bibliography: Drewes, R.C. and Vindum, J.V. (1994), Laurent, R.F. (1951), Laurent, R.F. (1972), Schiøtz, A. (1975), Schiøtz, A. (1999)
Data Providers: Arne Schiøtz, Robert Drewes
Geographic Range This species is known only from Kivu Province Bibliography: Laurent, R.F. (1950), Laurent, R.F. (1972), Schiøtz, A. (1999)
in eastern Democratic Republic of Congo, from Mount Ruwenzori Data Providers: Arne Schiøtz
south to the northern slopes of the Itombwe Mountains. It occurs at
intermediate altitudes, and the type locality is at 1,750m asl. There
have been no recent records, presumably due to the lack of survey
work within its range.
Population The population status of this species is unknown.
Habitat and Ecology It is a species of transition forests, and its
adaptability to degraded habitats is not known. Its breeding biology
is unknown, though it presumably takes place in water.
Major Threats It is likely to be threatened by ongoing habitat loss
due to logging, agriculture and human settlements.
Conservation Measures It occurs in the Virungas National Park.
Further survey work is required to determine the current population
status of the species.
284 Threatened Amphibians of the World

VU Hyperolius cystocandicans Richards and Schiøtz, 1977

Vulnerable B1ab(v) Geographic Range This species is endemic to the Kenyan Highlands, east of the Rift Valley, from Limuru to Thompson’s
Order, Family: Anura, Hyperoliidae Falls, and east to the Nyambeni Mountains. It is a montane species and the type locality is at 2,200m asl.
Country Distribution: Kenya Population It said to be an abundant species, but there has apparently been a general decline in its population.
Current Population Trend: Decreasing Habitat and Ecology It is a species of open farmland and high-altitude grassland. It breeds in temporary, and
sometimes permanent, pools. It is often found together with Hyperolius montanus.
Major Threats Its montane grassland and pastureland habitats are probably not seriously threatened (except
perhaps by afforestation), but there has apparently been a general population decline in areas of suitable habitat
for reasons that are not understood.
Conservation Measures It occurs in the Mount Kenya and Aberdares National Parks. Research is needed to
investigate the reasons for the decline of this species in apparently suitable habitat.
Bibliography: Lötters, S. et al. (2004), Richards, C.M. and Schiøtz, A. (1977), Schiøtz, A. (1999)
Data Providers: Arne Schiøtz, Kim Howell, Stefan Lötters
© R.C. Drewes

EN Hyperolius dintelmanni Lötters and Schmitz, 2004

Endangered B1ab(iii) Geographic Range This recently described species is known only from the Edib Hills in the Bakossi Mountains in
Order, Family: Anura, Hyperoliidae south-western Cameroon at an altitude of 1,100-1,250m asl. Surveys in nearby areas, such as Mount Kupe and Mount
Country Distribution: Cameroon Nlonako, have not located this species, and it is probably endemic to the Bakossi Mountains.
Current Population Trend: Decreasing Population There is little information on its current population status.
Habitat and Ecology All specimens have been found on the outer margins of patches of primary forest. Most
specimens were found sitting at night in grassy vegetation in areas with ferns, less than one metre above the ground.
It is presumed to breed in water by larval development.
Major Threats Although it appears to be somewhat adaptable to habitat disturbance (since it has been found on
the edges of forest), it occurs in an area where extensive forest clearance for smallholder farming is taking place
and it is unlikely to persist in the face of such rampant habitat loss.
Conservation Measures It does not occur in any protected areas, and its habitat in the Bakossi Mountains should
© Andreas Schmitz

be protected. Further survey work is needed to determine the population status of this species.
Bibliography: Lötters, S. and Schmitz, A. (2004)
Data Providers: Stefan Lötters

VU Hyperolius discodactylus Ahl, 1931

Vulnerable B1ab(iii) Geographic Range This species occurs in western Rwanda, western Burundi, south-western Uganda and eastern
Order, Family: Anura, Hyperoliidae Democratic Republic of Congo (west of Lake Edward and in Mount Ruwenzori). It is a montane species, but its exact
Country Distribution: Burundi, Congo, D.R., altitudinal range is unclear.
Rwanda, Uganda Population It is rare and seldom found.
Current Population Trend: Decreasing Habitat and Ecology It is a species of montane forests, associated with rivers, streams and moving waters where
it presumably breeds. Its adaptability to secondary habitats is not known.
Major Threats Little information is available, though it is probably impacted by the loss of habitat for agriculture
(crops and livestock), wood extraction and human settlements.
Conservation Measures It is present in the Bwindi National Park (Uganda), Ruwenzori Mountains National Park
(Uganda), and Virungas National Park (Democratic Republic of Congo).
Notes on taxonomy: There is some question as to the correct name for this species. Hyperolius alticola is usually considered to be
a synonym of H. discodactylus, and the name discodactylus has priority over alticola. However, Schiøtz (1999) found the type of H.
© Alan Channing

discodactylus to be indeterminable, and therefore preferred to use the name alticola. Here we use the older name, discodactylus,
pending resolution of this problem. There appears to be some question as to whether or not this species is specifically distinct from
Hyperolius frontalis (Schiøtz 1999).
Bibliography: Drewes, R.C. and Vindum, J.V. (1994), Laurent, R.F. (1972), Schiøtz, A. (1975), Schiøtz, A. (1999)
Data Providers: Arne Schiøtz, Robert Drewes

VU Hyperolius endjami Amiet, 1980

Vulnerable B1ab(iii) Geographic Range This species is known only from south-western Cameroon at 100-1,100m asl. It occurs as two
Order, Family: Anura, Hyperoliidae separate populations: a northern range from Mboassoum and Mankouat (near Nkongsamba) to Fopouanga (near
Country Distribution: Cameroon Nkondjock); and a southern population from near Kribi to the general vicinity of Yaounde. It probably occurs a little
Current Population Trend: Decreasing more widely than current records suggest.
Population It appears to be an uncommon species.
Habitat and Ecology It inhabits dense primary and secondary forest, and does not survive outside forest. It breeds
in flooded swamps with shallow water and in small pools.
Major Threats It is likely to be threatened by ongoing habitat loss for logging, agriculture, and human settle-
ments.
Conservation Measures It is not confirmed from any protected areas, though it probably does occur in at least one.
Nonetheless, there is a need for improved habitat protection at sites where this species is known to occur.
© Jean-Louis Amiet

Bibliography: Amiet, J.-L. (1975), Amiet, J.-L. (1980b), Amiet, J.-L. (1986), Schiøtz, A. (1999)
Data Providers: Jean-Louis Amiet, Arne Schiøtz
 Globally Threatened Amphibian Species 285

VU Hyperolius frontalis Laurent, 1950

Vulnerable B1ab(iii) Geographic Range This species occurs in eastern Democratic Republic of Congo from north-west of Mount Ru-
Order, Family: Anura, Hyperoliidae wenzori, south to west of Lake Kivu, extending into extreme south-western Uganda in the Bwindi Forest. Its recorded
Country Distribution: Congo, D.R., Uganda altitudinal range is 700-2,000m asl.
Current Population Trend: Decreasing Population It is reasonably abundant where it occurs.
Habitat and Ecology It is a species of transition and montane forest. It has been found in dense secondary vegetation
overhanging a small stream and in a small marsh in forest. Its breeding biology is unknown, though it presumably
takes place in aquatic habitats.
Major Threats Little information is available, though it is likely to be impacted by loss of habitat for agriculture
(crops and livestock), wood extraction, and human settlements.
Conservation Measures It is present in Bwindi National Park (Uganda) and Virungas National Park (Democratic
Republic of Congo).
Notes on taxonomy: There appears to be some question as to whether or not this species is specifically distinct from Hyperolius
© R.C. Drewes

discodactylus (Schiøtz 1999).

Bibliography: Drewes, R.C. and Vindum, J.V. (1994), Laurent, R.F. (1950), Laurent, R.F. (1972), Schiøtz, A. (1975), Schiøtz, A. (1999)
Data Providers: Arne Schiøtz, Robert Drewes

VU Hyperolius horstockii (Schlegel, 1837)

Vulnerable B1ab(iii) Geographic Range This is a coastal species occurring at low elevations along the southern coast of the Western Cape
Order, Family: Anura, Hyperoliidae Province (including the Cape Peninsula) in South Africa, east into the western part of the Eastern Cape Province.
Country Distribution: South Africa Population It is a locally common species, though it has declined at the western extremities of its range.
Current Population Trend: Decreasing Habitat and Ecology It lives in wetlands in coastal fynbos heathland. It breeds in large and small pans, dams, vleis,
and even slow-flowing streams. It needs emergent vegetation, and therefore requires relatively permanent water,
though it seems to avoid very deep water.
Major Threats Spreading alien vegetation can lead to drying out of its breeding habitats. It is probably also adversely
affected by fires. Several populations have disappeared due to the impacts of agricultural and urban expansion on
its native habitat.
Conservation Measures It occurs in several protected areas, including Cape Peninsula National Park and De Hoop
© Atherton de Villiers

Nature Reserve.
Bibliography: Channing, A. (2001), Minter, L.R. et al. (2004), Passmore, N.I. and Carruthers, V.C. (1995), Poynton, J.C. (1963), Schiøtz,
A. (1999), Visser, J. (1979a), Wager, V.A. (1986)
Data Providers: Leslie Minter, Alan Channing, James Harrison

EN Hyperolius kihangensis Schiøtz and Westergaard In Schiøtz, 1999

Endangered B1ab(iii) Geographic Range This species is known only from the Luhega Forest in the Udzungwa Scap Forest Reserve at 1,740m
Order, Family: Anura, Hyperoliidae asl in the Udzungwa Mountains in eastern Tanzania. It is probably more widespread in the Udzungwa Mountains,
Country Distribution: Tanzania though it appears to be absent from apparently suitable sites close to the type locality.
Current Population Trend: Decreasing Population It is common in its only known locality.
Habitat and Ecology It has been found in swamps in dense montane forest. Its breeding biology is unknown, though
it is likely to take place in swamps. Its adaptability to altered habitats is unknown.
Major Threats The main threat to the species is likely to be the loss of habitats for agriculture, livestock, wood
extraction, and human settlements.
Conservation Measures It occurs in the Udzungwa Scap Forest Reserve; it could also occur in the Udzungwa
National Park, although its presence there has not yet been confirmed.
© Michele Menegon

Bibliography: Poynton, J.C. (2003b), Schiøtz, A. (1999), Schiøtz, A. and Westergaard, M. (2000)
Data Providers: Arne Schiøtz, Kim Howell

VU Hyperolius laurenti Schiøtz, 1967

Vulnerable B1ab(iii) Geographic Range This species has been recorded from Lakota in south-central Côte d’Ivoire, and in Ghana from
Order, Family: Anura, Hyperoliidae Bobiri Forest Reserve in the south-west and Boi-Tano Forest Reserve, Draw River Forest Reserve, and the Ankasa
Country Distribution: Côte d’Ivoire, Ghana Conservation Area. Surveys in the forests of south-western Ghana in recent years have not succeeded in locating
Current Population Trend: Decreasing this species.
Population It could be a rare species, but it is very cryptic, and so might be under-recorded.
Habitat and Ecology It has only been found in lowland wet evergreen tropical rainforest, and it probably cannot
survive in altered habitats. The males call at dusk, from branches above small, swiftly flowing streams in which the
species breeds (it can sometimes be found up to 4m above water in the vegetation).
Major Threats It is threatened by ongoing habitat loss for logging, agriculture, and human settlements.
Conservation Measures It occurs in the Bobiri Forest Reserve in Ghana, but it is not known how well protected
this area is, as well as the Boi-Tano Forest Reserve, Draw River Forest Reserve, and the Ankasa Conservation Area,
a twin wildlife protected area comprised of Nini-Suhien National Park to the north, and the Ankasa Forest Reserve
© Annika Hillers

to the south.
Bibliography: Rödel, M.-O. et al. (2005), Schiøtz, A. (1967), Schiøtz, A. (1999)
Data Providers: Arne Schiøtz, Mark-Oliver Rödel
286
EN Hyperolius leleupi Laurent, 1951
Endangered B1ab(iii)
Order, Family: Anura, Hyperoliidae
Country Distribution: Congo, D.R.
Current Population Trend: Decreasing
EN Hyperolius leucotaenius Laurent, 1950
Endangered B1ab(iii)
Order, Family: Anura, Hyperoliidae
Country Distribution: Congo, D.R.
Current Population Trend: Decreasing
VU Hyperolius minutissimus Schiøtz, 1975
Vulnerable B1ab(iii)
Order, Family: Anura, Hyperoliidae
Country Distribution: Tanzania
Current Population Trend: Decreasing
© Michele Menegon
EN Hyperolius nienokouensis Rödel, 1999
Endangered B1ab(iii)
Order, Family: Anura, Hyperoliidae
Country Distribution: Côte d’Ivoire
Current Population Trend: Decreasing
© Mark-Oliver Rödel
Threatened Amphibians of the World
Geographic Range This species is known only from the Itombwe Bibliography: Laurent, R.F. (1951), Schiøtz, A. (1999)
Mountains in eastern Democratic Republic of Congo. The type local- Data Providers: Arne Schiøtz
ity is at 2,550m asl. A mapped occurrence in Burundi (Schiøtz 1999)
appears not to be correct.
Population It is poorly known, but it is apparently a rare species.
There have been no recent records, presumably due to lack of recent
herpetological work within its range.
Habitat and Ecology It is a species of high-altitude bamboo forests.
Its breeding biology is unknown, but if it is similar to the closely
related Hyperolius castaneus, then it is likely to breed in swamps.
Major Threats The major threat is likely to be habitat loss for
agriculture, livestock and human settlements.
Conservation Measures It is not known from any protected
areas, and the Itombwe Mountains are in urgent need of formal
protection.
Geographic Range This species is known only from the area to the
west of the southern part of the Itombwe Highlands and northern
slopes of the Kabobo Highlands, in eastern Democratic Republic of
Congo. It has been recorded at 1,850-2,000m asl.
Population The population status of this species is unknown.
There have been no recent records, presumably due to lack of recent
herpetological work within its range.
Habitat and Ecology It is a species of transition forests, and its
adaptability to altered habitats is not known. Its breeding biology is
also unknown, though it presumably takes place in water.
Major Threats It is likely to be threatened by habitat loss for
agriculture, livestock and human settlements.
Conservation Measures It is not known from any protected areas,
and there is a need for urgent protection of its habitat.
Bibliography: Laurent, R.F. (1950), Schiøtz, A. (1999)
Data Providers: Arne Schiøtz
Geographic Range This species is known only from southern Tanzania, where it has been recorded from the Udzungwa
Mountains and from the Njombe area in the western part of the Southern Highlands. It has been suggested that it
might occur in northern Malawi, but recent surveys on Mount Rungwe, in southern Tanzania, close to the Malawi
border, did not find this species. Its altitudinal range is not recorded, but it is likely to be mainly above 1,800m asl.
Population It is rarely seen, but easily overlooked, and so perhaps not as rare as records suggest.
Habitat and Ecology It lives in open, windswept, montane grasslands. It has been found in pastureland, but it is
not clear that it can survive in croplands. It breeds in shallow, temporary swamps.
Major Threats Its montane grassland habitat is threatened by afforestation, agricultural expansion, fire, and hu-
man settlement.
Conservation Measures It has not been found in the Udzungwa National Park, though it might occur there.
Bibliography: Poynton, J.C. (2003b), Schiøtz, A. (1975), Schiøtz, A. (1999), Schiøtz, A. and Westergaard, M. (2000)
Data Providers: Arne Schiøtz, Kim Howell
Geographic Range This species is known only from two localities in south-western Côte d’Ivoire: the foot of Mont
Niénokoué (approximately 15km west of Guiroutou) on the south-western edge of Taï National Park; and a nearby
locality 30km to the north of this.
Population It is a very uncommon species. A visit to the type locality in 2002 was unsuccessful in locating the
species.
Habitat and Ecology It is found only in primary rainforest, and probably attaches eggs to leaves above shallow
temporary ponds and swamps, where the larvae develop.
Major Threats It is presumably threatened by ongoing habitat loss for logging, agriculture and human settle-
ments.
Conservation Measures It occurs in Taï National Park.
Bibliography: Rödel, M.-O. (1999), Rödel, M.-O. (2000b), Schiøtz, A. (1999)
Data Providers: Arne Schiøtz, Mark-Oliver Rödel

EN Hyperolius nimbae Laurent, 1958
Endangered B1ab(iii)
Order, Family: Anura, Hyperoliidae
Country Distribution: Côte d’Ivoire
Current Population Trend: Decreasing
© Arne Schiøtz
EN Hyperolius pickersgilli Raw, 1982
Endangered B1ab(ii,iii,iv)+2ab(ii,iii,iv)
Order, Family: Anura, Hyperoliidae
Country Distribution: South Africa
Current Population Trend: Decreasing
© Richard Boycott
VU Hyperolius polystictus Laurent, 1943
Vulnerable D2
Order, Family: Anura, Hyperoliidae
Country Distribution: Congo, D.R.
Current Population Trend: Unknown
VU Hyperolius riggenbachi Nieden, 1910
Vulnerable B1ab(iii)
Order, Family: Anura, Hyperoliidae
Country Distribution: Cameroon, Nigeria
Current Population Trend: Decreasing
© Václav Gvoždík
Globally Threatened Amphibian Species
Geographic Range This species is probably restricted to the sur-
roundings of the falls of the Lofoi River, in south-eastern Democratic
Republic of Congo. There have been no recent records, presumably
due to a lack of survey work within its range.
Population The population status of this species is unknown.
Habitat and Ecology It is only known from the area around the Lofoi
River waterfalls. It could be a specialized species restricted to the
spray zone of the waterfalls, or to humid forest fed by the spray.
Major Threats There is no information on threats to this species.
However, any developments upstream of the falls could have a
devastating impact on the micro-habitat on which this species
depends, as happened with the Kihansi Spray Toad Nectophrynoides
asperginis.
Conservation Measures It is not known from any protected areas.
There is a need for further survey work to determine, and monitor,
the current population status of this species.
287
Geographic Range This species is known only from lowlands on the south-eastern foot of Mount Nimba in Côte
d’Ivoire, though it presumably also occurs in Guinea and Liberia.
Population It is apparently extremely common in its restricted range.
Habitat and Ecology It lives in clearings in lowland forest, where it calls at the edge of large swamps. It presum-
ably breeds in swamps.
Major Threats The threats are hard to determine, since it is not known to what extent the species can adapt to
opening up of its habitat. It appears to have a genuinely small range, since it is a conspicuous species and probably
would have been found elsewhere if it is more widespread. It might be adversely affected by expanding agriculture,
logging and human settlement.
Conservation Measures It is not known from any protected areas. Although part of Mount Nimba is protected
as the Mount Nimba Strict Nature Reserve (which was added to the list of World Heritage Sites in 1981) the site is
urgently in need of stricter protection and improved management, particularly given that it represents the only known
site where several highly threatened species are known to persist.
Notes on taxonomy: This species is is part of the Hyperolius tuberculatus Group within the H. viridiflavus complex (Schiøtz 1999).
Bibliography: Laurent, R.F. (1958a), Laurent, R.F. (1976a), Schiøtz, A. (1967), Schiøtz, A. (1971), Schiøtz, A. (1999)
Data Providers: Arne Schiøtz, Mark-Oliver Rödel
PICKERSGILL’S REED FROG
Geographic Range This species is endemic to the coast of Kwa-Zulu Natal Province, South Africa, ranging from
Kingsburgh in the south to Bonamanzi Game Park in the north. It occurs close to sea level.
Population It is secretive and so is under-recorded, but appears to be a rare species.
Habitat and Ecology It is a species of coastal mosaic bushland and grassland, breeding in stagnant, usually temporary
to semi-permanent, water, rarely exceeding 50cm in depth, surrounded by dense sedges. It is seldom found at the
same breeding sites as the abundant Hyperolius marmoratus.
Major Threats It is confined to a small area subject to urbanization, habitat fragmentation, afforestation, and
drainage for agricultural and urban development. Some breeding sites are being polluted by DDT, which is used
for controlling malarial mosquitoes. The spread of alien vegetation, in particular eucalyptus, is responsible for the
drying out of some breeding sites.
Conservation Measures It occurs in the Greater St Lucia Wetland Park, the Ulalazi Game Reserve, and the Twin-
streams-Mtunzini Natural Heritage Site.
Bibliography: Branch, W.R. (1988), Channing, A. (2001), Lambiris, A.J.L. (1989a), Minter, L.R. et al. (2004), Passmore, N.I. and Carruthers,
V.C. (1995), Poynton, J.C. and Broadley, D.G. (1987), Raw, L.R.G. (1982), Schiøtz, A. (1999)
Data Providers: Leslie Minter, James Harrison, Arne Schiøtz
Bibliography: Laurent, R.F. (1943), Schiøtz, A. (1999)
Data Providers: Arne Schiøtz
Geographic Range This species is known from the Obudu and Mambilla Plateaus of eastern Nigeria, and the
Bamenda Highlands, and the Adamawa Plateau (at Ngaouyanga and Banguve), western and central Cameroon. Two
subspecies occur: H. r. riggenbachi and H. r. hieroglyphicus, the latter occurring in the Bamenda Highlands and on the
Obudu Plateau, in an enclave surrounded by the former subspecies. Its altitudinal range is 900-1,800m asl.
Population It is a very common species.
Habitat and Ecology It lives in wetlands and small wooded watercourses in montane grassland. Males call at dusk
from the stems of waterside plants. It breeds in still water near streams.
Major Threats It is probably affected by the ongoing loss of its montane grassland, wetland and stream habitat due
to small-scale agricultural activities and wood collection, as well as human settlement.
Conservation Measures It is not known to occur in any protected areas. Improved habitat protection at sites where
this species is known to occur is needed.
Bibliography: Amiet, J.-L. (1973b), Böhme, W. and Schneider, B. (1987), Gartshore, M.E. (1986), Joger, U. (1982), Laurent, R.F. (1961),
Perret, J.-L. (1962), Perret, J.-L. (1966), Schiøtz, A. (1967), Schiøtz, A. (1999)
Data Providers: Arne Schiøtz, Jean-Louis Amiet
288 Threatened Amphibians of the World

EN Hyperolius rubrovermiculatus Schiøtz, 1975

Endangered B1ab(iii) Geographic Range This species is known only from the Shimba Hills, south of Mombassa, in coastal Kenya, being
Order, Family: Anura, Hyperoliidae replaced further south by Hyperolius mitchelli.
Country Distribution: Kenya Population It is common in its tiny range.
Current Population Trend: Decreasing Habitat and Ecology It lives in dry forest, dense humid savannah, and heavily degraded former forest (farm bush).
It breeds in marshes and temporary pools.
Major Threats Although somewhat adaptable, there is a limit to the amount of opening up of its habitat that it can

© De Saix, courtesy of Arne Schiøtz

tolerate, and so it is probably impacted by agricultural expansion and increasing human settlements.
Conservation Measures It probably occurs in the Shimba Hills National Reserve, which is in need of strengthened
management.
Notes on taxonomy: This species is possibly a subspecies of Hyperolius mitchelli (Schiøtz 1999).
Bibliography: Howell, K.M. (1993), Schiøtz, A. (1975), Schiøtz, A. (1999)
Data Providers: Arne Schiøtz, Robert Drewes

EN Hyperolius tannerorum Schiøtz, 1982

Endangered B1ab(iii)+2ab(iii) Geographic Range This species is known only from just outside the Mazumbai Forest Reserve at 1,410m asl in the
Order, Family: Anura, Hyperoliidae West Usambara Mountains, in north-eastern Tanzania. It might occur more widely in the West Usambara Mountains,
Country Distribution: Tanzania although there is little remaining undisturbed forest at this altitude.
Current Population Trend: Decreasing Population There is very little information on its population status, and only one small population is known. It pos-
sibly has no advertisement call, so it might be easily overlooked.
Habitat and Ecology It is known only from undisturbed montane forest. The only known breeding site is a small
swamp in forest, bordering a small stream.
Major Threats The forest habitat outside the Mazumbai Forest Reserve is being impacted by agricultural expansion,
logging and expanding human settlements.
Conservation Measures It probably occurs in the University of Dar es Salaam’s forest reserve at Mazumbai.
Further survey work is necessary to obtain a better understanding of the population status and distribution range
of this species.
© Arne Schiøtz

Notes on taxonomy: Schiøtz (1999) emended the name from Hyperolius tanneri to H. tannerorum.
Bibliography: Howell, K.M. (1993), Schiøtz, A. (1982), Schiøtz, A. (1999)
Data Providers: Arne Schiøtz, Kim Howell

EN Hyperolius thomensis Bocage, 1886

Endangered B1ab(iii) Geographic Range This species is endemic to São Tomé Island in São Tomé and Príncipe. It is restricted to primary
Order, Family: Anura, Hyperoliidae forest remnants at elevations usually above 800m asl.
Country Distribution: São Tomé and Príncipe Population There is little information on the population status of this species, since it is notoriously difficult to
Current Population Trend: Decreasing find.
Habitat and Ecology It is restricted to primary rainforest, and is known to breed by larval development in tree holes.
The same tree hole can be utilized by many individuals for breeding at different times.It has previously been reported
from towns, but there is now significant doubt about these records.
Major Threats Little information is available on threats, though it is likely to be impacted by habitat loss for agri-
culture, livestock, wood extraction, and human settlements.
Conservation Measures A large portion of its range falls within the proposed 295km² Parc Naturel Obo.
Notes on taxonomy: This species has previously been assigned to the genus Nesionixalus.
Bibliography: Drewes, R.C. (2002), Drewes, R.C. and Stoelting, R.E. (2004), Drewes, R.C. and Wilkinson, J.A. (2004), Loumont, C. (1992),
© R.C. Drewes

Perret, J.-L. (1988b), Schiøtz, A. (1999)

Data Providers: Robert Drewes, Arne Schiøtz

EN Hyperolius torrentis Schiøtz, 1967

Endangered B1ab(iii)+2ab(iii) Geographic Range This species occurs only from the Akwapim-Togo Range in eastern Ghana and western Togo.
Order, Family: Anura, Hyperoliidae Population It is an uncommon species, and only a very few localities are known.
Country Distribution: Ghana, Togo Habitat and Ecology It is a forest species that is restricted to an area where forest is severely degraded (though
Current Population Trend: Decreasing it does not occur in degraded habitats outside forest). A stream-breeding species, it is closely associated with very
fast-flowing streams and waterfalls; the eggs are laid on leaves above water. It has been found on or near vertical
cliffs in forest, in gullies in open forest, and in trees and on vegetation close to streams.
Major Threats Its forest habitat is being impacted by agricultural expansion, logging, and increasing human
settlements.
Conservation Measures It occurs in Kyabobo National Park in Ghana.
Bibliography: Rödel, M.-O. and Agyei, A.C. (2003), Schiøtz, A. (1967), Schiøtz, A. (1999)
© Mark-Oliver Rödel

Data Providers: Arne Schiøtz, Mark-Oliver Rödel

 Globally Threatened Amphibian Species 289

VU Hyperolius viridigulosus Schiøtz, 1967

Vulnerable B2ab(iii) Geographic Range This species is known only from south-central and western Ghana and southern Côte d’Ivoire.
Order, Family: Anura, Hyperoliidae Population It appears to be a rare species. It is easy to overlook, since it is a canopy species, but it has a distinctive,
Country Distribution: Côte d’Ivoire, Ghana though rarely heard, call.
Current Population Trend: Decreasing Habitat and Ecology It occurs in closed-canopy lowland rainforest, where it calls from high up in the trees, sometimes
far from water, but usually beside streams. Its breeding habits are unknown.
Major Threats Its forest habitat is being impacted by agricultural expansion, logging, and increasing human
settlements.
Conservation Measures It occurs in Bobiri Forest Reserve, but it is not clear how well protected this site is, as well
as Boi-Tano Forest Reserve, Draw River Forest Reserve and the Ankasa Conservation Area, a twin wildlife protected
area comprising Nini-Suhien National Park to the north and the Ankasa Forest Reserve to the south, in Ghana.
Bibliography: Rödel, M.-O. et al. (2005), Schiøtz, A. (1967), Schiøtz, A. (1999)
Data Providers: Arne Schiøtz, Mark-Oliver Rödel
© Raffael Ernst

VU Kassina arboricola Perret, 1985

Vulnerable B2ab(iii) Geographic Range This species ranges from south-western Côte d’Ivoire to south-central Ghana, though within
Order, Family: Anura, Hyperoliidae this general range it is known only from 4-5 localities.
Country Distribution: Côte d’Ivoire, Ghana Population There have only been a few records, but it is abundant where it has been found.
Current Population Trend: Decreasing Habitat and Ecology It is a species of secondary forest and forest edges, rather than undisturbed primary forest.
It only occurs marginally in heavily degraded former forest (farm bush). It breeds in both temporary and permanent
water, favouring large, well-vegetated pools.
Major Threats It is probably suffering from severe deforestation as a result of agricultural expansion, logging, and
growing human settlements.
Conservation Measures It occurs in Banco National Park and Mont Peko National Park in Côte d’Ivoire, and in
Bobiri Forest Reserve in Ghana.
© Mark-Oliver Rödel

Notes on taxonomy: We follow Perret (1985) and Rödel et al. (2002) in considering this to be a species distinct from Kassina
cochranae.
Bibliography: Perret, J.-L. (1985), Rödel, M.-O. (2003), Rödel, M.-O. et al. (2002), Rödel, M.-O. and Ernst, R. (2003), Schiøtz, A. (1999)
Data Providers: Mark-Oliver Rödel, Arne Schiøtz

VU Kassina lamottei Schiøtz, 1967

Vulnerable B1ab(iii)+2ab(iii) Geographic Range This species is only known from western and south-western Côte d’Ivoire, from the Mount
Order, Family: Anura, Hyperoliidae Nimba area, south to the Taï National Park. It is likely to occur in Liberia and Guinea, but there have not so far been
Country Distribution: Côte d’Ivoire any records. Records from Sierra Leone are in error.
Current Population Trend: Decreasing Population It is a very localized species, but nonetheless abundant where it occurs.
Habitat and Ecology It is a terrestrial species inhabiting lowland rainforest, and living only in mature and closed-
canopy forest. It breeds in large temporary forest pools.
Major Threats It is probably suffering from severe deforestation as a result of agricultural expansion, logging, and
growing human settlements. The trend towards decreasing rainfall seriously affects this species in particular, since
it lives in a sandy area, and low rainfall leads to drying out of its breeding ponds.
Conservation Measures It occurs in Taï National Park and Cavally Classified Forest.
© Mark-Oliver Rödel

Bibliography: Rödel, M.-O. (2000b), Rödel, M.-O. et al. (2002), Rödel, M.-O. and Branch, W.R. (2002), Rödel, M.-O. and Ernst, R. (2001),
Schiøtz, A. (1967), Schiøtz, A. (1999)
Data Providers: Mark-Oliver Rödel, Arne Schiøtz

VU Leptopelis barbouri Ahl, 1929

Vulnerable B1ab(iii) Geographic Range This species is endemic to eastern and southern Tanzania, and occurs in the East Usambara
Order, Family: Anura, Hyperoliidae Mountains, the Uzungwa Mountains, and on Mount Rungwe. It probably also occurs on other mountain ranges in
Country Distribution: Tanzania the Eastern Arc Mountain chain. It ranges from 700-2,100m asl.
Current Population Trend: Decreasing Population It is common along watercourses in the Udzungwa Mountains, but is apparently less common in the
East Usambara Mountains.
Habitat and Ecology It is a species of transition and montane rainforest, and can tolerate limited habitat disturbance,
although it requires forest in reasonable condition nearby. The eggs are laid in an underground nest near streams and
pools (sometimes as much as 10m away), into which tadpoles emerge.
Major Threats Its forest habitat is being impacted by agricultural expansion, logging, and increasing human
© Tim Davenport / WCS

settlements. Its habitat in the East Usambara Mountains has recently come under serious threat as a result of the
activities of illegal gold miners.
Conservation Measures It occurs in the Amani Nature Reserve, and is likely to occur in the Udzungwa National
Park, though there have not yet been any records.
Bibliography: Harper, E. and Vonesh, J.R. (2003), Howell, K.M. (1993), Poynton, J.C. (2003b), Schiøtz, A. (1975), Schiøtz, A. (1999)
Data Providers: Arne Schiøtz, Kim Howell, John Poynton, Tim Davenport
290 Threatened Amphibians of the World

EN Leptopelis karissimbensis Ahl, 1929

Endangered B1ab(iii) Geographic Range This species is known only from high altitudes in north-western Rwanda, south-western Uganda
Order, Family: Anura, Hyperoliidae and adjacent eastern Democratic Republic of Congo. Its range is centred on the Virunga volcanoes, with an altitudinal
Country Distribution: Congo, D.R., Rwanda, range of 2,000-2,800m asl and occasionally down to 1,500m asl. The map in Schiotz (1999) is in error, since it is, by
Uganda mistake, a duplicate of that for Leptopelis oryi.
Current Population Trend: Decreasing Population There is no recent information on the population status of this species.
Habitat and Ecology It is known from montane meadows in steep forest on volcanic soils, and also from highland
savannahs and heathland. Its adaptability to secondary habitats is also unknown. There is no information on its
breeding. However, assuming that it breeds in a similar manner to other species in the genus, it lays its eggs in a
nest buried in mud near water, into which the larvae emerge and develop.
Major Threats There is no direct information on threats, but most of its range is in protected areas. However, political
instability and high human population pressure continue to threaten the remaining habitat in its range.
Conservation Measures It occurs in the Virunga National Park in the Democratic Republic of Congo, the Volcanoes
National Park in Rwanda, and the Mgahinga Gorilla National Park in Uganda. Further survey work is needed to determine
© Ronalda Keith

the current population status of this species and to investigate aspects of its breeding biology.
Notes on taxonomy: This species is often confused with Leptopelis kivuensis (see Schiøtz 1999).
Bibliography: Laurent, R.F. (1972), Laurent, R.F. (1973), Schiøtz, A. (1975), Schiøtz, A. (1999)
Data Providers: Arne Schiøtz, Robert Drewes

VU Leptopelis palmatus (Peters, 1868)

Vulnerable D2 Geographic Range This species is endemic to Príncipe Island, in São Tomé and Príncipe, in the Gulf of Guinea. It
Order, Family: Anura, Hyperoliidae is present through most of the island, up to at least 700m, and possibly up to 1,000m asl.
Country Distribution: São Tomé and Príncipe Population It is a reasonably abundant species.
Current Population Trend: Stable Habitat and Ecology It generally inhabits wet forest along the edges of creeks and streams. It can also be found
in forest remnants and possibly in towns. It presumably buries its eggs close to water, with the tadpoles moving into
pools or streams where they develop further.
Major Threats There are no immediate threats, but increasing development of the island, leading to severe opening
up of the habitat, is a potential future threat.
Conservation Measures It is not known from any protected areas. This species requires close population monitoring
given that it is restricted entirely to Príncipe.
Notes on taxonomy: Populations from mainland Africa assigned to this species are now considered to be Leptopelis rufus.
Bibliography: Drewes, R.C. (2002), Drewes, R.C. and Stoelting, R.E. (2004), Loumont, C. (1992), Perret, J.-L. (1973), Schiøtz, A. (1999)
© R.C. Drewes

Data Providers: Robert Drewes, Arne Schiøtz

VU Leptopelis parkeri Barbour and Loveridge, 1928

Vulnerable B1ab(iii) Geographic Range This species is endemic to several mountains in the Eastern Arc chain of Tanzania, namely:
Order, Family: Anura, Hyperoliidae Uluguru, Udzungwa, East and West Usambara, and South Pare. It probably occurs a little more widely than these
Country Distribution: Tanzania records suggest. Although mainly a montane species, it lives in the foothills down at least to 300m asl (perhaps
Current Population Trend: Decreasing lower), and up to at least 2,000m asl.
Population In some places it appears to be easy to find and even common, whereas in other places it is rare.
Habitat and Ecology It inhabits closed, intact rainforest, not surviving in seriously disturbed habitats outside forest.
It breeds in slow-flowing streams, although the eggs are laid in a nest on land not far from water.
Major Threats It is affected by habitat loss as a result of expanding human settlements and agriculture, as well
as the harvesting of wood. Its habitat in the East Usambara Mountains has recently come under serious threat as a
result of the activities of illegal gold miners.
Conservation Measures It occurs in the Amani Nature Reserve, and in the University of Dar es Salaam’s reserve
at Mazumbai. It has not yet been recorded from Udzungwa National Park, but with further survey work probably will
© Don Church

be shown to occur there.

Bibliography: Harper, E. and Vonesh, J.R. (2003), Howell, K.M. (1993), Poynton, J.C. (2003b), Schiøtz, A. (1975), Schiøtz, A. (1999)
Data Providers: Kim Howell, John Poynton, Arne Schiøtz

VU Leptopelis ragazzii (Boulenger, 1896)

Vulnerable B2ab(iii) Geographic Range This species is endemic to the Ethiopian plateau, on both sides of the Rift Valley, at altitudes
Order, Family: Anura, Hyperoliidae of 1,930-3,010m asl.
Country Distribution: Ethiopia Population It is still locally common at suitable sites within its relatively limited range.
Current Population Trend: Decreasing Habitat and Ecology It is strictly associated with montane forest, not surviving in degraded habitats. It breeds by
larval development in pools and small streams, both permanent and temporary. The eggs are laid in nests on land
not far from water.
Major Threats The main threat is habitat loss from forest clearance, human settlement, and agricultural encroach-
ment. In some areas, this species seems only to survive in isolated forest remnants, such as often occur along the
slopes of steep-sided valleys.
Conservation Measures It is present in the Bale Mountains National Park.
Bibliography: Largen, M.J. (1977), Largen, M.J. (2001), Schiøtz, A. (1999)
Data Providers: Malcolm Largen, Arne Schiøtz
© M.J. Largen
 Globally Threatened Amphibian Species 291

EN Leptopelis susanae Largen, 1977

Endangered B1ab(iii)+2ab(iii) Geographic Range This species is endemic to the Gughe Mountains, at 2,600-3,000m asl, in southern Ethiopia.
Order, Family: Anura, Hyperoliidae Population It was still reasonably numerous in at least two localities in 1975, but the absence of subsequent records
Country Distribution: Ethiopia is due to a lack of fieldwork.
Current Population Trend: Decreasing Habitat and Ecology It is confined to montane forest remnants, and has not been found in degraded habitats. Calling
males have been found along the banks of streams, in which larval development presumably occurs. The eggs are
presumably laid in nests on land, near the water.
Major Threats The main threats are forest clearance, human settlement, and agricultural encroachment. The Gughe
Mountains are heavily populated and intensively cultivated, and only small patches of montane forest remain.
Conservation Measures It is not currently known to be present in any protected areas, and the protection of
forest remnants in the Gughe Mountains, and the search for further sites where the species might be found, are of
the highest priority.
Bibliography: Largen, M.J. (1977), Largen, M.J. (2001), Schiøtz, A. (1999)
© M.J. Largen

Data Providers: Malcolm Largen, Arne Schiøtz

VU Leptopelis uluguruensis Barbour and Loveridge, 1928

Vulnerable B1ab(iii) Geographic Range This species is endemic to several mountains in the Eastern Arc chain of Tanzania, namely:
Order, Family: Anura, Hyperoliidae East Usambara, Nguu, Nguru, Uluguru, and Udzungwa (Kihansi Gorge and Mwanihana Forest). It probably occurs a
Country Distribution: Tanzania little more widely than records suggest. It occurs mainly above 900m asl, and it is probably sub-montane, perhaps
Current Population Trend: Decreasing not ranging much above 1,650m asl.
Population It is a common species.
Habitat and Ecology It inhabits wet submontane rainforest, but can tolerate slight disturbance and is sometimes
found in banana patches in forest clearings. However, it is never found in open areas away from mature forest. Its
breeding biology is unknown, but it presumably breeds in water, and males call near streams. The eggs are probably
laid in nests on land, not far from water (either streams or temporary pools).
Major Threats It is affected by habitat loss as a result of expanding human settlements and agriculture, as well
© Michele Menegon

as the harvesting of wood. Its habitat in the East Usambara Mountains has recently come under serious threat from
the activities of illegal gold miners.
Conservation Measures It occurs in the Udzungwa National Park and the Amani Nature Reserve.
Bibliography: Harper, E. and Vonesh, J.R. (2003), Howell, K.M. (1993), Poynton, J.C. (2003b), Schiøtz, A. (1975), Schiøtz, A. (1999)
Data Providers: Kim Howell, John Poynton, Arne Schiøtz, Michele Menegon

VU Leptopelis vannutellii (Boulenger, 1898)

Vulnerable B2ab(iii) Geographic Range This species is known only from the highlands of south-western Ethiopia, at altitudes of
Order, Family: Anura, Hyperoliidae 1,500-2,200m asl.
Country Distribution: Ethiopia Population It is still locally common at suitable sites within its relatively limited range.
Current Population Trend: Decreasing Habitat and Ecology It is confined to dense, tropical deciduous forest, and does not survive in degraded habitats.
It breeds by larval development in pools and small streams, both permanent and temporary. The eggs are laid in
nests on land, not far from water.
Major Threats The main threat is habitat loss due to forest clearance, human settlement, and agricultural en-
croachment.
Conservation Measures It is not currently known from any protected areas. The most obvious requirement is habitat
conservation at selected sites, within the species range, where still undamaged lowland forest survives.
Bibliography: Largen, M.J. (1977), Largen, M.J. (2001), Schiøtz, A. (1999)
Data Providers: Malcolm Largen, Arne Schiøtz
© M.J. Largen

VU Leptopelis vermiculatus (Boulenger, 1909)

Vulnerable B2ab(iii) Geographic Range This species is endemic to several mountains in the Eastern Arc chain of Tanzania, namely:
Order, Family: Anura, Hyperoliidae East and West Usambara, Nguu, Udzungwa, Poroto and Mount Rungwe. It occurs above 900m asl, up to elevations
Country Distribution: Tanzania of at least 1,800m asl in southern Tanzania.
Current Population Trend: Decreasing Population It is a very common species through much of its range.
Habitat and Ecology It inhabits submontane and montane forest, and requires mature, closed-canopy forest. Its
breeding biology is unknown, but it presumably breeds in water, and males call near streams. The eggs are probably
laid in nests on land, not far from water (either streams or temporary pools).
Major Threats It is affected by habitat loss as a result of expanding human settlements and agriculture, and the
harvesting of wood. Its habitat in the East Usambara Mountains has recently come under serious threat from the
activities of illegal gold miners. It is sometimes found in the international pet trade although this is not currently
believed to constitute a major threat.
© Elizabeth Harper

Conservation Measures It occurs in the Amani Nature Reserve and the Udzungwa National Park.
Bibliography: Drewes, R.C., Altig, R. and Howell, K.M. (1989), Harper, E. and Vonesh, J.R. (2003), Howell, K.M. (1993), Poynton, J.C.
(2003b), Schiøtz, A. (1975), Schiøtz, A. (1999)
Data Providers: Kim Howell, John Poynton, Arne Schiøtz, Michele Menegon
292 Threatened Amphibians of the World

EN Leptopelis xenodactylus Poynton, 1963

Endangered B2ab(ii,iii,iv) Geographic Range This species is endemic to south-eastern South Africa in the southern KwaZulu-Natal Province
Order, Family: Anura, Hyperoliidae highlands, and marginally in Eastern Cape Province. It is not found on the steep slopes of the escarpment, and is usually
Country Distribution: South Africa found between 1,000 and 1,830m asl. Only a few localities are known, but it might be more widespread.
Current Population Trend: Decreasing Population It is a rare species.
Habitat and Ecology It occurs in grassland, keeping to areas away from trees, and breeds in upland bogs, grassy
wetlands and marshes, generally using semi-permanent water. Eggs are presumably laid in a nest on the ground
near water.
Major Threats It has a small range, and lives in a habitat that is threatened by afforestation, fire, and the spread
of alien plants that lower the water table (leading to drying out of breeding sites).
Conservation Measures It is found in the uKhahlamba-Drakensberg Park, which is at least well managed for
biodiversity conservation.
© Richard Boycott

Bibliography: Armstrong, A. (2001), Branch, W.R. (1988), Channing, A. (2001), Lambiris, A.J.L. (1989a), Minter, L.R. et al. (2004), Passmore,
N.I. and Carruthers, V.C. (1995), Poynton, J.C. (1963), Poynton, J.C. (1964b), Schiøtz, A. (1999)
Data Providers: Leslie Minter, Alan Channing, Arne Schiøtz, James Harrison

VU Phlyctimantis keithae Schiøtz, 1975

Vulnerable B1ab(iii) Geographic Range This species is known only from the Udzungwa Mountains in eastern Tanzania. It has generally
Order, Family: Anura, Hyperoliidae been recorded at 1,800-2,000m asl, but probably has a wider altitudinal range than this.
Country Distribution: Tanzania Population It is common within its restricted range.
Current Population Trend: Decreasing Habitat and Ecology It lives in forest, montane grassland, and open farmland near forest. It breeds in shallow pools,
including artificial ponds, with emergent vegetation. It is probably a forest species that moves into open areas when
it is breeding. It exhibits a rather unusual and cryptic defensive posture.
Major Threats Its montane forest and grassland habitat is threatened by afforestation, agricultural expansion, fire,
and human settlement.
Conservation Measures There are no records from the Udzungwa National Park, but with further survey work it
probably will be shown to occur there.
© Michele Menegon

Bibliography: Channing, A. and Howell, K. (2003), Perret, J.-L. (1986), Poynton, J.C. (2003b), Schiøtz, A. (1975), Schiøtz, A. (1999)
Data Providers: Arne Schiøtz, Kim Howell

LEIOPELMATIDAE

CR Leiopelma archeyi Turbott, 1942 ARCHEY’S FROG

Critically Endangered A2ae Geographic Range This species occurs in the Whareorino range in the west and Coromandel ranges in the east on
Order, Family: Anura, Leiopelmatidae North Island, New Zealand. Ranges from 400-1,000m asl.
Country Distribution: New Zealand Population Formerly, this species was recorded in the tens of thousands, but declines since 1996 have reduced the
Current Population Trend: Decreasing numbers throughout their range. The decline was first noted in 1996; one study population on the Tapu Ridge declined
by 88% (433 frogs down to 53 frogs) over the period 1996-2002.
Habitat and Ecology A terrestrial and nocturnal species, it occurs mostly at higher altitudes in forested ranges and
more open sub-alpine scrub. It lays large unpigmented eggs in damp situations on the ground, which undergo direct
development without a larval stage.
Major Threats The definitive cause of the decline of this species is still not known, though disease of some kind
is the most likely cause, in particular chytridiomycosis. Chytrid fungus infection was first identified in this species in
September 2001 (specimen collected June 2001 at Te Moehu). Declines of species were documented through most
of its range-central Coromandel (1996-1998), northern Coromandel (2000-2001) and Whareorino (2001).
© Avi Holzapfel

Conservation Measures The NZ Department of Conservation (DOC) through its Native Frog Recovery Group and
Native Frog Recovery Plan administers conservation management of the species and permits appropriate research,
including a collaborative program with Auckland Zoo which has recently established a new facility for breeding and
maintaining the species. Recent declines and the positive identification of chytrid fungus has stimulated urgent
research and management of the species, including pathology, population monitoring, captive management and
molecular research. This involves DOC and Canterbury, Massey, Otago and Victoria Universities.
Bibliography: Archey, G. (1922), Bell, B.D. (1978), Bell, B.D. (1982b), Bell, B.D. (1985), Bell, B.D. (1996), Bell, B.D. (1999), Bell, B.D. et
al. (2004), Bell, B.D., Daugherty, C.H. and Hitchmough, R.A. (1998), Gill, B.J., Whitaker, A.H. (1996), Newman, D.G. (1996), Stephenson,
E.M., Stephenson, N.G. (1957), Turbott, E.G. (1942)
Data Providers: Ben Bell

EN Leiopelma hamiltoni McCulloch, 1919 HAMILTON’S FROG

Endangered D Geographic Range This species is confined to a single rock stack on Stephens Island in New Zealand. Translocation
Order, Family: Anura, Leiopelmatidae to an adjoining man-made rock stack on Stephens Island has been attempted but with limited success.
Country Distribution: New Zealand Population The total population numbers less than 300 individuals.
Current Population Trend: Stable Habitat and Ecology It is found on boulder banks near the summit of Stephens Island. It lays large unpigmented
eggs in damp situations on the ground, which undergo direct development without a larval stage.
Major Threats The major threats to the species are introduced mammalian predators (e.g. Rattus rattus) and pos-
sibly disease (chytridiomycosis has been recently identified in the closely related L. archeyi). Tuatara (Sphenodon
punctatus) also prey on the species.
Conservation Measures A tuatara-proof fence has been constructed around the frog’s habitat. Regular population
monitoring is also in place, and an island habitat restoration programme is under way. The prevention of accidental
colonization by introduced mammalian predators is necessary. Translocation of some frogs to a second island is under
consideration. The NZ Department of Conservation (DOC) through its Native Frog Recovery Group and Native Frog
© Paddy Ryan

Recovery Plan administers conservation management of this species and permits appropriate species research.
Notes on taxonomy: Populations on Maud Island were previously considered to belong to this species, but were described as a new
species, Leiopelma pakeka, in 1998.
Bibliography: Bell, B.D. (1978), Bell, B.D. (1985), Bell, B.D. (1996), Bell, B.D., Daugherty, C.H. and Hay, J.M. (1998), Brown, D. (1994),
Gill, B.J., Whitaker, A.H. (1996), Holyoake, A., Waldman, B. and Gemmell, N.J. (2001), McCulloch, A.R. (1919), Newman, D.G. (1990),
Newman, D.G. (1996)
Data Providers: Mandy Tocher, Derek Brown, Ben Bell

VU Leiopelma hochstetteri Fitzinger, 1861
Vulnerable A3e; B1ab(iii)
Order, Family: Anura, Leiopelmatidae
Country Distribution: New Zealand
Current Population Trend: Decreasing
© Paddy Ryan
Geographic Range This species is known from numerous localities in northern North Island, New Zealand.
Population It is a common species.
VU Leiopelma pakeka Bell, Daugherty and Hay, 1998
Vulnerable D2
Order, Family: Anura, Leiopelmatidae
Country Distribution: New Zealand
Current Population Trend: Stable
© Paddy Ryan
VU Adelophryne baturitensis Hoogmoed, Borges and Cascon, 1994
Vulnerable B1ab(iii)
Order, Family: Anura, Leptodactylidae
Country Distribution: Brazil
Current Population Trend: Decreasing
EN Adelophryne maranguapensis Hoogmoed, Borges and Cascon, 1994
Endangered B1ab(iii)
Order, Family: Anura, Leptodactylidae
Country Distribution: Brazil
Current Population Trend: Decreasing
Globally Threatened Amphibian Species
LEPTODACTYLIDAE
Geographic Range This species is known only from the Serra (or
Maciço) de Baturité, in the State of Ceará, north-eastern Brazil, at
600-1,000m asl.
Population It was very common within its small range, and often
found at numerous localities since it was first described up until
1993; however, from 1994 to 2003 there were 34 surveys of the
same sites and not a single individual was found, until in July 2004
three individuals were recorded.
Habitat and Ecology Most specimens have been found in dry or
moist leaf-litter on the ground, in bromeliads, and stream margins
in reasonably well-preserved closed forests. However, it can also
survive in shaded coffee plantations. It is a diurnal species that
breeds by direct development, and the eggs are probably deposited
in wet spots on the ground.
Major Threats The major threat is habitat loss due to logging,
agriculture, and human settlement. The very good soils and favour-
Geographic Range This species is known only from the Serra
de Maranguape, in the State of Ceará, north-eastern Brazil, at
800-900m asl.
Population It is abundant in certain parts of its small range and
is regularly recorded.
Habitat and Ecology This is a diurnal species that lives in the
leaf-litter of primary and secondary forest, and has also been found
in bamboo and banana plantations, as well as forest fragments (but
not open habitats). It is a direct developer, and the eggs are probably
laid on the ground.
Major Threats The Serra de Maranguape is being very heavily
deforested, due to logging, agriculture, and human settlement.
Conservation Measures It is not found in any protected areas,
and the formal protection of remaining forest and forest fragments
in the Serra de Maranguape is urgently needed.
293
HOCHSTETTER’S FROG
Habitat and Ecology This semi-aquatic species can be found alongside forested creeks and watercourses, mostly in
native forest, but surviving to some extent in modified habitats such as farmland and exotic forest. It lays large unpig-
mented eggs in damp situations on the ground, which then undergo direct development without a larval stage.
Major Threats Disease (chytridiomycosis) will most likely be a major threat to this species in the near future, as it
has recently been identified in the related L. archeyi. This species co-exists with introduced mammalian predators
(such as Rattus rattus, Mustela erminea), but their long-term impact on the species is uncertain. Habitat loss, due to
the conversion of native habitat to pine plantations, is certainly a threat, as is the siltation of breeding streams that
takes place as a result of logging and road building activities.
Conservation Measures The NZ Department of Conservation (DOC), through its Native Frog Recovery Group and
Native Frog Recovery Plan, administers conservation management of this species and permits appropriate species
research. Possible declines, and the identity of chytrid fungus in Leiopelma archeyi, have stimulated urgent research
and management of the two species, including studies on pathology, population monitoring, captive management
and molecular research. This involves DOC and Canterbury, Massey, Otago, and Victoria Universities. Recent studies
have suggested that this species might comprise cryptic species, and hence taxonomic research is recommended
to resolve this taxonomic issue.
Notes on taxonomy: Recent genetic studies suggest that this species, as currently recognized, represents a species complex.
Bibliography: Bell, B.D. (1978), Bell, B.D. (1985), Bell, B.D. (1996), Bowsher, J.H. (2000), Fitzinger, L.J. (1861), Gill, B.J., Whitaker, A.H.
(1996), Newman, D.G. (1996), Stephenson, E.M., Stephenson, N.G. (1957)
Data Providers: Ben Bell, Mandy Tocher, Philip Bishop, Bruce Waldman
MAUD ISLAND FROG
Geographic Range This species is confined to Maud Island, New Zealand. In 1997, 300 individuals were translocated
to Motuara Island, where it seems to have been established successfully.
Population This species’ population is estimated at between 27,500 and 39,500 individuals in a 16-ha forest remnant
on Maud Island. There are approximately 200 individuals estimated on Motuara Island.
Habitat and Ecology It can be found in coastal forest, mostly in deep boulder banks on lower slopes. It lays large un-
pigmented eggs in damp situations on the ground, which then undergo direct development without a larval stage.
Major Threats The introduction of mammalian predators, such as Rattus rattus and Mustela erminea, represents
a major potential threat to this species. Disease (chytridiomycosis) also represents a possible threat, since it has
recently been identified in the closely related L. archeyi.
Conservation Measures NZ Department of Conservation (DOC), through its Native Frog Recovery Group and Native
Frog Recovery Plan, administers conservation management of the species and permits appropriate species research.
Regular population monitoring and a habitat restoration programme are needed. The colonization of the islands by
introduced mammalian predators must be prevented.
Notes on taxonomy: The frogs now described as Leiopelma pakeka were previously considered to belong to L. hamiltoni.
Bibliography: Bell, B.D. (1978), Bell, B.D. (1985), Bell, B.D. (1996), Bell, B.D., Daugherty, C.H. and Hay, J.M. (1998), Gill, B.J., Whitaker,
A.H. (1996), Holyoake, A., Waldman, B. and Gemmell, N.J. (2001), Newman, D.G. (1990), Newman, D.G. (1996), Tocher, M. and Newman,
D.G. (1997)
Data Providers: Ben Bell, Mandy Tocher, Philip Bishop, Bruce Waldman
able climate of the area are encouraging agricultural expansion, and the species is unable to survive in the banana
plantations that are rapidly taking over the area.
Conservation Measures It is not known from any protected areas, and there is clearly a need for improved habitat
protection at sites where this species is known to occur. Further survey work is needed to determine whether or not
this species is experiencing a decline, or whether or not it is just undergoing natural population fluctuations.
Bibliography: Eterovick, P.C. et al. (2005), Hoogmoed, M.S., Borges, D.M. and Cascon, P. (1994)
Data Providers: Débora Silvano, Diva Borges-Najosa
Bibliography: Borges-Nojosa, D.M. and Lima, D.C. (2001), Eterovick, P.C. et al. (2005), Hoogmoed, M.S., Borges, D.M. and Cascon,
P. (1994)
Data Providers: Débora Silvano, Diva Borges-Najosa
294 Threatened Amphibians of the World

VU Alsodes barrioi Veloso, Díaz, Iturra, Penna, 1981

Vulnerable A3c Geographic Range This species is known only from the Cordillera de Nahuelbuta, Estero Cabrería, Provincia de
Order, Family: Anura, Leptodactylidae Malleco, Chile (37° 50’S; 73° 00’ W). It has an altitudinal range of 1,000-1,500m asl.
Country Distribution: Chile Population The species is relatively abundant at the type locality. During breeding periods, it is possible to find
Current Population Trend: Decreasing large numbers of tadpoles in streams.
Habitat and Ecology It occurs under logs or stones close to mountain streams, and the surrounding vegetation
mainly comprises Nothofagus dombeyi and Araucaria araucana. Breeding takes place in streams.
Major Threats The main threat is the afforestation of the watershed where this species occurs with an introduced
pine. When the pine is harvested, the effect on the surrounding breeding streams could be catastrophic.
Conservation Measures It occurs in Parque Nacional Nahuelbuta.
Bibliography: Formas, J.R. (1995), Glade, A. (1993), Servicio Agrícola Ganadero (1998), Veloso, A. et al. (1981), Veloso, A. and Navarro,
J. (1988)
© Andrés Morya

Data Providers: Alberto Veloso, Herman Núñez, Ramón Formas

CR Alsodes montanus (Lataste, 1902)

Critically Endangered B1ab(iii,v)+2ab(iii,v) Geographic Range This species is currently known only from Farellones (33° 32’S; 70° 19’W), in central Chile,
Order, Family: Anura, Leptodactylidae 35km north-east of Santiago, at 2,300-3,000m asl. The exact location of the type locality is uncertain, though it is
Country Distribution: Chile somewhere on the western slopes of the Andes in central Chile.
Current Population Trend: Decreasing Population The current known population comprises only a few adults. It was last collected in 2003, and the popula-
tion is known to have declined substantially in recent years. Areas in which the species has been known to reproduce
were found to have been completely destroyed in February 2002.
Habitat and Ecology This stream-breeding species has been found under stones, close to mountain streams, where
the surrounding vegetation comprises grasses and shrubs of cold Andean areas. It has free-swimming tadpoles.
Major Threats Currently, the primary threat to this species is from recreational skiing and tourist infrastructure
development, since the only known locality where the species is found is in a ski resort.
Conservation Measures Protection and maintenance of its existing habitat is a priority for this species, since it is
not currently known from any protected area.
© Andrés Morya

Bibliography: Díaz, N. (1989), Formas, J.R. (1995), Glade, A. (1993), Servicio Agrícola Ganadero (1998), Veloso, A. and Navarro, J.
(1988)
Data Providers: Alberto Veloso, Herman Núñez

CR Alsodes tumultuosus Veloso, Iturra, Galleguillos, 1979

Critically Endangered B1ab(iii,v)+2ab(iii,v) Geographic Range This species is known only from Farellones (33° 32’S; 70°19’W), Central Chile, 35km north-east
Order, Family: Anura, Leptodactylidae of Santiago, at an altitudinal range of 2,300-3,000m asl, though it is possibly more widely distributed.
Country Distribution: Chile Population The population is currently believed to comprise only a few adults. Like its congener, A. montanus, it
Current Population Trend: Decreasing was last collected in 2003, and the population has declined substantially in recent years. Areas where the species
is known to breed were found to have been completely destroyed by February 2002.
Habitat and Ecology This stream-breeding species has been found under stones, close to mountain streams,
where the surrounding vegetation comprises grasses and shrubs characteristic of cold Andean areas. It has free-
swimming tadpoles.
Major Threats Currently, the primary threat to this species is from recreational skiing and tourist infrastructure
development, since the only known locality where the species is found is in a ski resort.
Conservation Measures Protection and maintenance of its existing habitat is a priority for this species, as it is
currently not known from any protected area.
© Andrés Morya

Bibliography: Formas, J.R. (1995), Glade, A. (1993), Servicio Agrícola Ganadero (1998), Veloso, A. and Navarro, J. (1988), Veloso, A.,
Iturra, P. and Galleguillos, R. (1979)
Data Providers: Alberto Veloso, Herman Núñez

CR Alsodes vanzolinii (Donoso-Barros, 1974)

Critically Endangered A2ac; B2ab(iii) Geographic Range This species is endemic to the western slopes of the Nahuelbuta Range, Arauco Province (37°
Order, Family: Anura, Leptodactylidae 59’S; 73° 22’W), Chile, at 50m asl.
Country Distribution: Chile Population The most recent collection was of tadpoles in 2002 which were found in the only pool in the area.
Current Population Trend: Decreasing Habitat and Ecology This species can be found in small patches of Nothofagus forest, and is not tolerant of habitat
modification. It reproduces in pools within permanent streams and has free-swimming tadpoles.
Major Threats It is highly threatened by habitat destruction due to expanding pine plantations and human settle-
ment. In the last 10 years, in particular, the range of this species has been nearly completely destroyed as a result
of afforestation with pine plantations.
Conservation Measures The species is not known to occur in any protected area, and protection and maintenance of
its remaining native habitat is an urgent priority. Further survey work is necessary to determine the current population
status of this species and the limits of its range.
© Andrés Morya

Bibliography: Donoso-Barros, R. (1974), Formas, J.R. (1981), Formas, J.R. (1995), Formas, J.R. and Brieva, L. (2003), Servicio Agrícola
Ganadero (1998), Veloso, A. and Navarro, J. (1988)
Data Providers: Alberto Veloso, Herman Núñez, Juan Carlos Ortiz
 Globally Threatened Amphibian Species 295

VU Atelognathus nitoi (Barrio, 1973)

Vulnerable D2 Geographic Range This species occurs only at the type locality (Laguna Verde on Cerro Challhuanco) and in neigh-
Order, Family: Anura, Leptodactylidae bouring areas in Argentina. Its altitudinal range is between 1,300 and 1,550m asl.
Country Distribution: Argentina Population It is common within its restricted range. The minimum population size is estimated at around 1,000
Current Population Trend: Stable individuals (Ubeda et al. 1999).
Habitat and Ecology This species can be found in permanent and temporary ponds and humid areas of caducifolius
forest (Nothofagus pumilio); breeding takes place in permanent ponds. It is not known from modified habitats.
Major Threats Potential major threats include fires (both natural and human initiated) and increasing touristic
activities.
Conservation Measures The entire known population is in the reserve area of the Parque Nacional Nahuel Huapi.
There is a need for close population monitoring of this species given that it is known from only a single location.
Bibliography: Barrio, A. (1973), Basso, N.G. and Ubeda, C.A. (1997), Echeverria, D.D., Ubeda, C.A. and Basso, N.G. (2001), Lavilla, E.O.
et al. (2000), Lynch, J.D. (1978), Ubeda, C. et al. (1999)
© Boris Blotto

Data Providers: Carmen Úbeda, Esteban Lavilla

EN Atelognathus patagonicus (Gallardo, 1962)

Endangered A2ace; B1ab(i,ii,v) Geographic Range This species is endemic to north-western Argentinean Patagonia where it has been recorded
Order, Family: Anura, Leptodactylidae from a system of endorreic and isolated lagoons scattered in the volcanic tablelands of Neuquen (between 38° 55’
Country Distribution: Argentina and 39° 32’S and 70° 20’ to 70° 39’W). Its altitudinal range is 1,265-1,410m asl.
Current Population Trend: Decreasing Population It is common within its preferred habitat. New subpopulations have recently been discovered (at 14
satellite ponds), but the major subpopulation (Laguna Blanca) is believed to have been extirpated.
Habitat and Ecology The species inhabits permanent lagoons that are surrounded by steppe and/or semi-desert.
Individuals are only found in areas adjacent to the ponds, and breeding takes place within these ponds. It is not
known from degraded habitats.
Major Threats The main subpopulation (Laguna Blanca) is believed to have been extirpated through predation by
introduced fishes (perch and salmonids). The remaining subpopulations are isolated from this lake and mostly appear
to be stable, although eutrophication of two ponds due to livestock may be leading to a decline in two subpopulations.
Introduction of predatory fishes to the remaining lakes is a potential major threat to all subpopulations.
© Boris Blotto

Conservation Measures While the species was previously recorded from Parque Nacional Laguna Blanca it is
now known only from the ponds within the buffer zone of the park. Measures need to be put in place to prevent the
introduction of predatory fishes into the habitat of the remaining subpopulations.
Notes on taxonomy: This is a polymorphic species that might consist of several species or subspecies.
Bibliography: Cei, J.M. (1980), Cei, J.M. and Roig, V.G. (1968), Gallardo, J.M. (1962b), Lynch, J.D. (1978)
Data Providers: Carmen Úbeda, Esteban Lavilla, Néstor Basso

EN Atelognathus praebasalticus (Cei and Roig, 1968)

Endangered A2ace; B1ab(i,ii,v) Geographic Range This species is endemic to north-western Argentinean Patagonia, and has been recorded from
Order, Family: Anura, Leptodactylidae a number of localities in Neuquen Province. The altitudinal range is from 1,000-1,500m asl.
Country Distribution: Argentina Population It is a rare species.
Current Population Trend: Decreasing Habitat and Ecology It lives on the ground under stones and in humid ravines surrounding small-scattered lagoons,
in volcanic tablelands, with steppe and semi-desert landscapes; it is presumed to breed in the lagoons. The species
does not occur in modified habitats.
Major Threats The main threat is the introduction of predatory fishes (perch and salmonids) in Laguna Blanca, but it
is also threatened by the loss of pond side vegetation through overgrazing and trampling by livestock (sheep).
Conservation Measures The species was previously protected in Parque Nacional Laguna Blanca; however, it is
now extirpated in this park because of the introduction of predatory fishes and habitat loss. Three of the subspecies
lack any protection, and habitat protection is urgently needed as are measures to prevent the future introduction of
predatory fish species.
© Boris Blotto

Notes on taxonomy: The Atelognathus praebasalticus complex needs to be re-examined taxonomically. There are four known subspecies,
some of which may constitute new species. Some of these subspecies may be extinct.
Bibliography: Cei, J.M. (1972), Cei, J.M. (1980), Cei, J.M. and Roig, V.G. (1968), Lavilla, E.O. et al. (2000), Lynch, J.D. (1978)
Data Providers: Carmen Úbeda, Néstor Basso, Esteban Lavilla, Boris Blotto

EN Atelognathus reverberii (Cei, 1969)

Endangered B1ab(i,ii,iii)+2ab(i,ii,iii) Geographic Range This species is known from five lagoons on the Somuncura Plateau, an isolated basaltic plateau
Order, Family: Anura, Leptodactylidae in Río Negro Province, Argentinean Patagonia, at an altitude of 1,000-1,200m asl.
Country Distribution: Argentina Population It is a rare species.
Current Population Trend: Decreasing Habitat and Ecology The species inhabits steppe tablelands that contain a number of temporary shallow lagoons
scattered over the volcanic plateau. Breeding takes place in permanent pools. It is not present in modified habitats.
Major Threats It is threatened by eutrophication of its aquatic habitat, and the trampling of the shallow ponds by
livestock (sheep and goats).
Conservation Measures The entire population of this species is included within the Somuncura Provincial Reserve;
however, this Reserve requires better management, since it affords no real protection for the area. Further surveys are
needed throughout the Somuncura Plateau area to better establish the population status of this species.
Bibliography: Cei, J.M. (1969a), Cei, J.M. (1969b), Cei, J.M. (1980), Lavilla, E.O. et al. (2000), Lynch, J.D. (1978)
Data Providers: Carmen Úbeda, Esteban Lavilla, Néstor Basso
© Néstor Basso
296
VU Atelognathus salai Cei, 1984
Vulnerable D2
Order, Family: Anura, Leptodactylidae
Country Distribution: Argentina
Current Population Trend: Stable
© Carmen Ubeda
VU Atelognathus solitarius (Cei, 1970)
Vulnerable D2
Order, Family: Anura, Leptodactylidae
Country Distribution: Argentina
Current Population Trend: Stable
CR Atopophrynus syntomopus Lynch and Ruíz-Carranza, 1982
Critically Endangered B1ab(iii)+2ab(iii)
Order, Family: Anura, Leptodactylidae
Country Distribution: Colombia
Current Population Trend: Decreasing
© John D. Lynch
EN Batrachophrynus brachydactylus Peters, 1873
Endangered A2d
Order, Family: Anura, Leptodactylidae
Country Distribution: Peru
Current Population Trend: Decreasing
© Mikael Lundberg
Threatened Amphibians of the World
Geographic Range This species is known only from the type
locality, Las Bayas Creek, 48km south of Pilcaniyeu, Río Negro
Province, Argentina. It is believed to have a restricted range, but
further surveys are needed to confirm this. It has been collected at
an altitude of 900m asl.
Population The current population status of this species is
unknown.
Habitat and Ecology The species’ habitat includes rocky ravines
along creeks in the wasted volcanic landscape of the Pilcaniyeu
steppe and tableland. The breeding habitat is not known.
Major Threats There is no information on the threats to this species.
There is only some grazing of sheep in the area, though it is unclear
what impact this might be having on the species.
Conservation Measures It is not recorded from any protected
areas. Further surveys are needed to obtain more information on the
population status and range of this poorly known species.
Geographic Range This species is known only from its type locality pond, Laguna de los Gendarmes (46° 6’S; 71°
41’W; ca 1,050m asl), and other nearby ponds in south-western Argentinean Patagonia (north-west of Santa Cruz
Province). It is known only from a single plateau.
Population It is a common species.
Habitat and Ecology Its habitat comprises permanent and endorreic lagoons at approximately 1,000m asl in the
hilly landscape of the Patagonian steppe, and the temperate deciduous austral beech forest (Nothofagus pumilio).
Breeding takes place in permanent ponds. It is not known if the species can occur in modified habitats.
Major Threats The region where this species occurs is relatively remote, and is not currently experiencing any
threats.
Conservation Measures It is not known from any protected areas. Further surveys are necessary to determine the
limits of the range of this species in this unexplored and remote area.
Bibliography: Cei, J.M. (1984), Ubeda, C.A. and Basso, N.G. (1998)
Data Providers: Carmen Úbeda, Esteban Lavilla, Néstor Basso
Bibliography: Cei, J.M. (1970b), Lavilla, E.O. et al. (2000), Lynch, J.D. (1978)
Data Providers: Carmen Úbeda, Esteban Lavilla, Néstor Basso, Boris Blotto
Geographic Range This species is known only from the region of the type locality, 8km east of Sonsón, in the
Cordillera Central, in the department of Antioquia, Colombia, at 2,780m asl. It might be more widespread, but it is
difficult at present to survey in surrounding areas due to security problems.
Population It is known only from three specimens, and while there have been at least a dozen surveys of the type
locality since its first description they have all failed to find the species. It has not been seen again since it was
first described.
Habitat and Ecology The species was recorded on wet rocks very close to the surface of the stream in primary
forest.
Major Threats Unfortunately, the habitat of the type locality has been quite degraded, largely as a result of guerilla
activities.
Conservation Measures Further survey work is needed to confirm whether or not the species still survives at the
type locality or if it occurs outside the vicinity of the type locality (although it is not a safe area to conduct fieldwork
at present). If the species still survives, restoration and protection of its habitat will be necessary for its survival,
since the type locality is not within any protected area.
Bibliography: Lynch, J.D. and Ruiz-Carranza, P.M. (1982), Myers, C.W. and Ford, L.S. (1986)
Data Providers: Fernando Castro, John Lynch
Geographic Range This species’ geographic range comprises the small tributaries of Lake Junin, and highlands and
neighbouring cordilleras of Junin and Pasco Departments in central Peru. It occurs at an altitude of approximately
4,000m asl.
Population It can be abundant where it occurs, but appears to have declined.
Habitat and Ecology This is a semi-aquatic riparian species, found in small tributaries of Lake Junin but not in
the lake itself.
Major Threats The main threat to the species is exploitation for food and medicine.
Conservation Measures It occurs in the Reserva Nacional de Junin. Measures are urgently required to control and
manage the offtake of this species from the wild.
Bibliography: Czopek, J. (1983), Instituto Nacional de Recursos Naturales (INRENA) (2000), Salas, A.W. and Sinsch, U. (1996), Sinsch,
U. (1986), Sinsch, U. (1990), Sinsch, U., Salas, A.W. and Canales, V. (1995)
Data Providers: Ariadne Angulo, Ulrich Sinsch, Edgar Lehr
 Globally Threatened Amphibian Species 297

EN Batrachophrynus macrostomus Peters, 1873

Endangered A2cde Geographic Range This species occurs in Junin and Pasco Departments in Peru; it has also been introduced into the
Order, Family: Anura, Leptodactylidae upper valley of the Río Mantaro, in the central Andes of Peru. It has an altitudinal range of 3,200-4,300m asl.
Country Distribution: Peru (Native and Introduced) Population Previously common, this species has undergone a noticeable decline over the last twenty years.
Current Population Trend: Decreasing Habitat and Ecology It is an exclusively aquatic species and can be found in marshy lakes such as Lake Junin and
smaller, deep, isolated lakes.
Major Threats The species is threatened by introduction of trout into lakes in the high Andes, by uncontrolled
exploitation as a food source, and potentially through the loss of habitat quality due to water pollution.
Conservation Measures Based on its known habitat preferences and distribution, this species occurs in the Reserva
Nacional de Junin and may also occur in the Santuario Nacional Huayllay and the Santuario Historico Chacamarca.
Measures are urgently required to control and manage the offtake of this species from the wild.
Bibliography: Czopek, J. (1983), Czopek, J. (1984), Dubois, A. (1984a), Instituto Nacional de Recursos Naturales (INRENA) (2000), Leja,
© Mikael Lundberg

M. (1991), Rodríguez, L.O., Cordova, J.H. and Icochea, J. (1993), Sinsch, U. (1986), Sinsch, U. (1990), Szarski, H. (1979), Tantalean, M.
and Garcia, L. (1989), Tantalean, M. and Garcia, L. (1993)
Data Providers: Ariadne Angulo, Ulrich Sinsch, César Aguilar Puntriano

VU Batrachyla fitzroya Basso, 1994

Vulnerable D2 Geographic Range This species is known only from the type locality (Isla Grande in Menendez Lake, 42° 40’S;
Order, Family: Anura, Leptodactylidae 71° 51’W), an island completely protected in Parque Nacional Los Alerces, Chubut Province, Argentina. It occurs
Country Distribution: Argentina at 500m asl.
Current Population Trend: Stable Population This species is locally common within its restricted range.
Habitat and Ecology It can be found in very humid temperate forest dominated by Nothofagus. Its breeding habitat
is not known, but it may breed within small temporary pools. It is not known whether or not it can occur in modified
habitats.
Major Threats There are no current threats at the type locality, but its restricted range renders it susceptible to
stochastic threatening processes.
Conservation Measures The population occurs in a highly protected area in Parque Nacional Los Alerces. There
is a need for close population monitoring of this species given its restricted range.
© Carmen Ubeda

Bibliography: Basso, N.G. (1994), Lavilla, E.O. et al. (2000)

Data Providers: Carmen Úbeda, Néstor Basso, Esteban Lavilla, Boris Blotto

VU Caudiverbera caudiverbera (Linnaeus, 1758)

Vulnerable A2ad Geographic Range This species occurs from Coquimbo (at approx. 29°S) to Puerto Montt (40°S), in Chile. It has
Order, Family: Anura, Leptodactylidae an altitudinal range of 0-500m asl.
Country Distribution: Chile Population Populations of this species are currently declining in central Chile; it is practically absent in ponds and
Current Population Trend: Decreasing lagoons close to towns where it was very abundant only a few years ago. However, populations are apparently
common and stable in southern Chile.
Habitat and Ecology It occurs in large, deep ponds, and small reservoirs, and breeding takes place in ponds. It
tolerates some minor disturbance of its habitat, but requires permanent ponds to persist.
Major Threats The main threats include harvesting as an exotic food source, and water pollution due to agriculture.
Introduced trout and the drainage of ponds for development and agriculture are additional threats.
Conservation Measures It occurs in several protected areas. The harvesting of this species from the wild needs
to be managed sustainably.
© Andrés Morya

Bibliography: Formas, J.R. (1995), Glade, A. (1993), Servicio Agrícola Ganadero (1998), Veloso, A. and Navarro, J. (1988)
Data Providers: Alberto Veloso, Ramón Formas

VU Ceratophrys stolzmanni Steindachner, 1882 PACIFIC HORNED FROG

Vulnerable B1ab(iii) Geographic Range This species is found in xeric environments of north-western Peru (Departamento Tumbes) and
Order, Family: Anura, Leptodactylidae the Gulf of Guayaquil, Ecuador (Guayas and Manabi Provinces). Two subspecies are recognized: C. s. scaphiopeza in
Country Distribution: Ecuador, Peru Ecuador, and C. s. stolzmanni in Peru. It has an altitudinal range of 0-100m asl.
Current Population Trend: Decreasing Population It is a very rare species, but has been recorded in the last few years in Ecuador. There is no information
on its population status in Peru.
Habitat and Ecology The habitat where some individuals have been collected can be described as tropical scrub,
which is usually fairly open with a few trees, but remains green for about half the year. Other specimens have been
collected in an open and sandy desert, with scattered, low, desert vegetation, and trees that are absent except in
or near temporary streambeds (Peters 1967). It lives below ground when not breeding, but is an explosive breeder
during good rains (with breeding taking place in water).
Major Threats The major threat is habitat loss due to agriculture (especially cattle ranching), logging, and human
© Ana Almendáriz

settlement. Soil pollution, due to agrochemicals, also represents a threat.

Conservation Measures It occurs in Parque Nacional Machililla and Reserva Ecológica Manglares Churute in
Ecuador, and the Zona Reservada de Tumbes in Peru. Further survey work is needed to determine the population
status of this species in Peru.
Bibliography: Lynch, J.D. (1982a), Parker III, T.A. and Carr, J.L. (1992), Peters, J.A. (1967), Rodríguez, L.O., Cordova, J.H. and Icochea,
J. (1993), Steindachner, F. (1882), Tello, G. (1998)
Data Providers: Ariadne Angulo, Luis A. Coloma, Santiago Ron, Diego Cisneros-Heredia
298
VU Craugastor alfredi (Boulenger, 1898)
Vulnerable B2ab(iii)
Order, Family: Anura, Leptodactylidae
Country Distribution: Guatemala, Mexico
Current Population Trend: Decreasing
VU Craugastor anatipes (Lynch and Myers, 1983)
Vulnerable B1ab(iii)
Order, Family: Anura, Leptodactylidae
Country Distribution: Colombia, Ecuador
Current Population Trend: Decreasing
© John D. Lynch
CR Craugastor anciano (Savage, McCranie and Wilson, 1988)
Critically Endangered B1ab(iii,v)+2ab(iii,v)
Order, Family: Anura, Leptodactylidae
Country Distribution: Honduras
Current Population Trend: Decreasing
CR Craugastor andi (Savage, 1974)
Critically Endangered A2ace
Order, Family: Anura, Leptodactylidae
Country Distribution: Costa Rica
Current Population Trend: Decreasing
© Federico Bolaños
Threatened Amphibians of the World
Geographic Range This species can be found from central Veracruz
in Mexico to western El Peten, Guatemala, at 0-600m asl.
Population This is a poorly known species that has not often been
collected; little is known about its biology and ecology. It is locally
common in Chiapas and is known from a single locality from El
Peten, Guatemala.
Habitat and Ecology It is an inhabitant of lowland moist forest,
and occurs in low vegetation, including under leaf sheaths of banana
plants. Reproduction occurs by direct development, probably on the
ground in moist leaf-litter.
Major Threats The main threat to this species is the loss of the
original forest cover, in particular due to logging and agriculture.
Conservation Measures It has been recorded in the protected ar-
eas at Yaxchilan, and the El Triunfo and El Ocote Biosphere Reserves
(where extensive forest remains) in Mexico. Additional fieldwork is
needed to evaluate the status of the populations in the field.
Geographic Range This species is known only from two sites in
the Montana de Celaque, Departments of Lempira and Ocopepeque,
western Honduras, at 1,400-1,840m asl.
Population It has always been extremely rare, and might now
be extinct.
Habitat and Ecology It lives on the ground along mountain streams
in premontane and lower montane moist forest. It presumably breeds
by direct development, with eggs laid on land.
Major Threats All streamside Craugastor known to occur above
900m asl have disappeared in Honduras for reasons that remain
unclear, although infection with chytridiomycosis is clearly a possibil-
ity. Other threats include agricultural encroachment, wood extraction,
forest fires, water pollution, and landslides.
Conservation Measures One of the two recorded sites is in Parque
Nacional Celaque. In view of the likely threat of chytridiomycosis,
it is a very high priority to conduct surveys to relocate this species
Notes on taxonomy: This species was previously within the genus Eleutherodactylus (Crawford and Smith 2005).
Bibliography: Campbell, J.A. (2001), Campbell, J.A., Lamar, W.W. and Hillis, D.M. (1989), Crawford, A.J. and Smith, E.N. (2005)
Data Providers: Georgina Santos-Barrera, Luis Canseco-Márquez, Gunther Köhler, Antonio Muñoz Alonso
Geographic Range This species occurs mainly in southern Colombia in the Pacific lowlands and slopes of the
Cordillera Occidental in the Departments of Nariño and Valle del Cauca, south to the Río Mira in extreme northern
Ecuador. Its altitudinal range is 100-1,600m asl.
Population It is a very uncommon species.
Habitat and Ecology It is a forest inhabitant, occurring from tropical lowland forest to lower cloud forest. The
ability of this species to adapt to disturbed habitats is unknown, although it is probably restricted to closed forest.
It is closely associated with small streams, and only rarely strays from the immediate vicinity of these; they are not
found along larger rivers. It is active at night, sitting on rocks or in rocky crevices, or on very steep cliffs alongside
streams. It is presumed to breed by direct development.
Major Threats The major threat is probably habitat loss, due to agricultural development (including plantations,
and the planting of illegal crops), logging, and human settlement. Pollution, resulting from the spraying of illegal
crops, is also a significant threat.
Conservation Measures It is present in several protected areas in Colombia, including Parque Nacional Natural
Farallones de Cali.
Notes on taxonomy: This species was previously included in the genus Eleutherodactylus (Crawford and Smith 2005).
Bibliography: Acosta-Galvis, A.R. (2000), Crawford, A.J. and Smith, E.N. (2005), Hedges, S.B. (1989), Lynch, J.D. (1998b), Lynch, J.D. (1999),
Lynch, J.D. and Ardila-Robayo, M.C. (1993), Lynch, J.D. and Duellman, W.E. (1997), Lynch, J.D. and Myers, C.W. (1983), Lynch, J.D., Ruiz-
Carranza, P.M. and Ardila-Robayo, M.C. (1997), Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and Lynch, J.D. (1996), Savage, J.M. (1987)
Data Providers: Fernando Castro, Santiago Ron, Luis A. Coloma
and determine its current population status; surviving individuals might need to form the basis for the establishment
of an ex-situ population.
Notes on taxonomy: This species was previously included in the genus Eleutherodactylus (Crawford and Smith 2005).
Bibliography: Campbell, J.A. and Savage, J.M. (2000), Crawford, A.J. and Smith, E.N. (2005), McCranie, J.R. and Wilson, L.D. (2002b),
Savage, J.M., McCranie, J.R. and Wilson, L.D. (1988)
Data Providers: Gustavo Cruz, Larry David Wilson, Randy McCranie
Geographic Range This species occurs on the Atlantic slopes of the northern and central portions of the Costa
Rican cordilleras at an altitude of 1,000-1,200m asl (Savage 2002).
Population It was once common, or at least regularly seen, in Monteverde and the San Ramon Reserve, but it has
not been recorded since 1990, despite searches, thus suggesting that a serious decline has taken place.
Habitat and Ecology This nocturnal arboreal frog inhabits premontane wet forest and rainforest. It usually is found
close to streams, but has occasionally been collected in bromeliads as much as 5m above ground. Males call from
stream banks, and females descend from trees for mating (Savage 2002). Eggs are laid out of the water on the ground,
and breeding occurs by direct development.
Major Threats This species has disappeared from apparently suitable habitats, with the implication that the
major threat is likely to be chytridiomycosis (perhaps in combination with the effects of climate change), leading to
a catastrophic population decline, as has occurred in many other montane amphibian species that are associated
with streams.
Conservation Measures Much of the known range of this species is in protected areas. However, further surveys
are urgently needed to relocate this species and determine its current population status; surviving individuals might
need to form the basis for the establishment of an ex-situ population.
Notes on taxonomy: This species was previously included in the genus Eleutherodactylus (Crawford and Smith 2005).
Bibliography: Crawford, A.J. and Smith, E.N. (2005), Pounds, J.A. et al. (1997), Savage, J.M. (2002)
Data Providers: Alan Pounds, Federico Bolaños

CR Craugastor angelicus (Savage, 1975)
Critically Endangered A2ace
Order, Family: Anura, Leptodactylidae
Country Distribution: Costa Rica
Current Population Trend: Decreasing
© Wayne Van Devender
VU Craugastor aphanus (Campbell, 1994)
Vulnerable D2
Order, Family: Anura, Leptodactylidae
Country Distribution: Guatemala
Current Population Trend: Stable
EN Craugastor aurilegulus (Savage, McCranie and Wilson, 1988)
Endangered B1ab(iii,v)+2ab(iii,v)
Order, Family: Anura, Leptodactylidae
Country Distribution: Honduras
Current Population Trend: Decreasing
EN Craugastor azueroensis (Savage, 1975)
Endangered B1ab(iii)
Order, Family: Anura, Leptodactylidae
Country Distribution: Panama
Current Population Trend: Decreasing
Globally Threatened Amphibian Species 299
Geographic Range This species occurs in the Cerro Cacao in the Cordillera de Guanacaste, Cordillera de Tilarán
(1,200-1,600m asl), and on the northern (600-1,700m asl) and eastern (900-1,400m asl) slopes of Volcán Poas (Cordil-
lera Central) in Costa Rica (Savage 2002).
Population The species has disappeared from Monteverde where it was abundant until the 1980s. There was one
sighting in the northern part of the Cordillera de Tilaran in 1994, but the site has not been resurveyed since (J.A.
Pounds pers. comm.). Recent surveys in Cerro Cacao (2001) and the Cordillera Central where the species was previ-
ously recorded have not turned up any individuals (F. Bolaños and G. Chaves pers. comm.).
Habitat and Ecology It inhabits tropical premontane and lower montane forest. It is nocturnal, foraging along steep
sloping banks of mountain streams (Savage 2002). A female has been observed to lay her clutch of eggs in a hole
and then cover it with sand (Hayes 1985).
Major Threats Since this species has disappeared from apparently suitable habitats-and some parts of its original
habitat remain pristine-the major threat to the species is likely to be chytridiomycosis (perhaps in synergy with climate
change), which has lead to catastrophic population declines in many other amphibian species that are associated
with high-elevation streams.
Conservation Measures Much of the known range of this species is in protected areas. However, further surveys
are urgently needed to relocate this species and determine its current population status; surviving individuals might
need to form the basis for the establishment of an ex-situ population.
Notes on taxonomy: This species was previously within the genus Eleutherodactylus (Crawford and Smith 2005).
Bibliography: Crawford, A.J. and Smith, E.N. (2005), Hayes, M.P. (1985), Pounds, J.A. et al. (1997), Savage, J.M. (2002)
Data Providers: Alan Pounds, Federico Bolaños
Geographic Range This species occurs in the Montañas del Notes on taxonomy: This species was previously included in the genus Eleutherodactylus (Crawford and Smith 2005).
Mico and the northern slopes of the eastern portion of the Sierra Bibliography: Campbell, J.A. (1994a), Campbell, J.A. (2001), Crawford, A.J. and Smith, E.N. (2005), Savage, J.M. and Myers, C.W.
de las Minas, in the Departamento de Izabal, eastern Guatemala, (2002)
at 591-786m asl. Data Providers: Manuel Acevedo, Eric Smith
Population It is a rare species.
Habitat and Ecology Although it is found at relatively low eleva-
tions, it lives in typical cloud forest conditions with frequent fog and
drizzling rain. It is found in undisturbed forest only. It breeds by direct
development and is not dependent on water.
Major Threats Small-scale wood extraction is taking place, but
much of its habitat is included in a protected area.
Conservation Measures It occurs in the Montanos del Mico
Catchment Reserve, and might also occur in Parque Nacional Sierra
de las Minas. Close population monitoring is required given that this
species is known from only a few sites.
Geographic Range This species occurs in the Sierra Nombre de Conservation Measures It occurs in several protected areas, including Parque Nacional Pico Bonito and Parque
Dios, the Montana La Muralla, and the Montana Texiguat, in the Nacional Texiguat. Further research is necessary to establish the cause of declines at high elevations, particularly
departments of Atlantida, Colon, and Yoro y Olancho in north-central to ascertain whether chytrid is a threat.
Honduras, at 50-1,550m asl. Notes on taxonomy: This species was previously included in the genus Eleutherodactylus (Crawford and Smith 2005).
Population The lowland populations appear to be stable and Bibliography: Campbell, J.A. and Savage, J.M. (2000), Crawford, A.J. and Smith, E.N. (2005), McCranie, J.R. and Wilson, L.D. (2002b),
common, but higher elevation populations are severely reduced Savage, J.M., McCranie, J.R. and Wilson, L.D. (1988)
above 900m asl. Data Providers: Gustavo Cruz, Larry David Wilson, Randy McCranie
Habitat and Ecology It lives on the ground, along small and me-
dium-sized streams in lowland moist forest, and premontane and
lower montane wet forest. It does not tend to survive in degraded
habitats. It is a species that breeds by direct development.
Major Threats The major threats include deforestation as a result
of agricultural and livestock encroachment, human settlements,
logging, fires and landslides. However, these threats do not account
for the inexplicable declines at high elevations; chytridiomycosis is
clearly a possibility.
Geographic Range This species is endemic to the Azuero Peninsula, Notes on taxonomy: This species was previously included in the genus Eleutherodactylus (Crawford and Smith 2005).
Panama, where it has been recorded at altitudes of 61-940m asl. Bibliography: Campbell, J.A. and Savage, J.M. (2000), Crawford, A.J. and Smith, E.N. (2005), Ibáñez, R. et al. (2000), Young, B. et
Population It is apparently locally abundant. al. (1999)
Habitat and Ecology This is a terrestrial species associated with Data Providers: Frank Solís, Roberto Ibáñez, César Jaramillo, Querube Fuenmayor
streams in dry lowland and montane forest. It breeds by direct
development and eggs are deposited on the ground.
Major Threats General habitat loss through deforestation is a major
threat. Some other species of the genus that are associated with
streams have undergone dramatic declines and disappearances,
possibly due to chytridiomycosis, so the status of this species should
be monitored carefully.
Conservation Measures The species has been recorded from
Parque Nacional Cerro Hoya, but expanded protection of habitat on
the Azuero Peninsula is needed. Further research is also required to
determine whether chytrid poses a real threat to this species.
300 Threatened Amphibians of the World

VU Craugastor biporcatus (Boettger, 1893)

Vulnerable B1ab(iii) Geographic Range This species is known from the Venezuelan Coastal Ranges in the States of Aragua, Carabobo,
Order, Family: Anura, Leptodactylidae Miranda, Sucre, and Yaracuy. It has been recorded from 250-1,600m asl.
Country Distribution: Venezuela Population It is a common species.
Current Population Trend: Decreasing Habitat and Ecology A terrestrial and nocturnal frog that inhabits lowland humid and montane cloud forests. It
breeds by direct development.
Major Threats The major threat is habitat destruction and degradation due to agriculture.
Conservation Measures Its range includes several protected areas, including Parque Nacional Guatopo, Parque
Nacional Henri Pittier, and Parque Nacional San Esteban.

© Francisco José López-López

Notes on taxonomy: This species was previously included in the genus Eleutherodactylus (Crawford and Smith 2005).
Bibliography: Barrio Amorós, C.L. (2004), Cordero, G. (1987), Crawford, A.J. and Smith, E.N. (2005), Frost, D.R. (1985), Heatwole,
H. (1962), La Marca, E. (1992), Lynch, J.D. (1975b), Manzanilla, J. et al. (1995), Manzanilla, J., Rivero, R. and Schmid, M. (1996), Savage,
J.M. and Myers, C.W. (2002)
Data Providers: Jesús Manzanilla, Enrique La Marca

VU Craugastor bocourti (Brocchi, 1877)

Vulnerable B1ab(iii) Geographic Range This species occurs in the mountains of Alta Bibliography: Campbell, J.A. (2001), Crawford, A.J. and Smith, E.N. (2005), Stuart, L.C. (1948)
Order, Family: Anura, Leptodactylidae Verapaz and the Sierra de las Minas, Guatemala, at 1,300-1,700m Data Providers: Manuel Acevedo, Eric Smith
Country Distribution: Guatemala asl.
Current Population Trend: Decreasing Population It is an uncommon species.
Habitat and Ecology It occurs on the forest floor in undisturbed
cloud forest. It breeds by direct development, and is not dependent
on water for breeding.
Major Threats The major threat is habitat loss due to agriculture
(in particular leather-leaf plantations), wood extraction, and human
settlement.
Conservation Measures It occurs in the Parque Nacional Sierra
de las Minas and Biotopo del Quetzal.
Notes on taxonomy: This species was previously included in the genus
Eleutherodactylus (Crawford and Smith 2005).

VU Craugastor brocchi (Boulenger, 1882)

Vulnerable B1ab(iii) Geographic Range This species is found on the Atlantic versant species of Craugastor that are associated with streams have undergone dramatic declines and disappearances,
Order, Family: Anura, Leptodactylidae slopes of the northern highlands of Guatemala from Sierra de las possibly due to chytridiomycosis, so the status of this species should be monitored carefully.
Country Distribution: Guatemala, Mexico Minas, Alta Verapaz, and Baja Verapaz, westward through Sierra de Conservation Measures It occurs in a single protected area in Mexico, Parque Nacional Lagunas de Montebello,
Current Population Trend: Decreasing los Cuchumatanes to north-western Huehuetenango, Guatemala, and is present in Biotopo del Quetzal and the Bisis-Cabá and Sierra de las Minas Biosphere Reserves in Guatemala.
then to Chiapas, Mexico, in Parque Nacional Lagunas de Montebello, Survey work is needed to evaluate the status of this species in Chiapas.
at altitudes of 1,200-2,000m asl. Notes on taxonomy: This species was previously included in the genus Eleutherodactylus (Crawford and Smith 2005).
Population It is uncommon throughout its range; there is no recent Bibliography: Campbell, J.A. and Savage, J.M. (2000), Crawford, A.J. and Smith, E.N. (2005), Lynch, J.D. (2000b), Savage, J.M.
information from Chiapas. (1987)
Habitat and Ecology This species prefers riparian habitats along Data Providers: Georgina Santos-Barrera, Manuel Acevedo, Antonio Muñoz Alonso
streams flowing through premontane and montane wet forests,
cloud forest and humid pine-oak forest. It is commonly found on the
ground or beneath leaf-litter along streams, where it also breeds
by direct development.
Major Threats A major threat to this species is habitat loss and
disturbance due to agriculture and selective logging, particularly
since it clearly does not adapt well to disturbed areas. Some other

CR Craugastor catalinae (Campbell and Savage, 2000)

Critically Endangered A2ace Geographic Range This species is found on the extreme south-western Pacific slope of Costa Rica and adjacent
Order, Family: Anura, Leptodactylidae western Panama, at elevations of 1,219-1,800m asl (Savage 2002).
Country Distribution: Costa Rica, Panama Population This species used to be common in Costa Rica; however, like many other frogs in the genus, it has
Current Population Trend: Decreasing disappeared from much of its known range. Likewise, populations are known to have declined in the uplands of
western Panama.
Habitat and Ecology This terrestrial species occurs in streams in premontane and lower montane humid forests,
and breeds by direct development.
Major Threats While the destruction of natural forests is presumably a threat to this species, the large population
© Michael and Patricia Fogden

decline has taken place within pristine habitats, and this might well be due to chytridiomycosis.
Conservation Measures The status of this species should be closely monitored, and, given the possible threat of
chytridiomycosis, some individuals might need to form the basis for the establishment of an ex-situ population. The
species has been recorded from a few protected areas including Parque Internacional La Amistad.
Notes on taxonomy: This species was previously included in the genus Eleutherodactylus (Crawford and Smith 2005).
Bibliography: Campbell, J.A. and Savage, J.M. (2000), Crawford, A.J. and Smith, E.N. (2005), Savage, J.M. (2002)
Data Providers: Frank Solís, Roberto Ibáñez, Gerardo Chaves, Jay Savage, César Jaramillo, Querube Fuenmayor

EN Craugastor charadra (Campbell and Savage, 2000)
Endangered B1ab(iii,v)
Order, Family: Anura, Leptodactylidae
Country Distribution: Guatemala, Honduras
Current Population Trend: Decreasing
© Silviu Petrovan
VU Craugastor cheiroplethus (Lynch, 1990)
Vulnerable B1ab(iii)
Order, Family: Anura, Leptodactylidae
Country Distribution: Colombia
Current Population Trend: Decreasing
CR Craugastor coffeus (McCranie and Köhler, 1999)
Critically Endangered B1ab(iii)+2ab(iii)
Order, Family: Anura, Leptodactylidae
Country Distribution: Honduras
Current Population Trend: Decreasing
CR Craugastor cruzi (McCranie, Savage and Wilson, 1989)
Critically Endangered A2ace;
B1ab(iii,v)+2ab(iii,v)
Order, Family: Anura, Leptodactylidae
Country Distribution: Honduras
Current Population Trend: Decreasing
Globally Threatened Amphibian Species 301
Geographic Range This species is known from the highlands between the Río Motagua in Guatemala and the Río
Chamelecón in Honduras, at 30-1,370m asl.
Population It has disappeared from above 900m asl in Honduras and much of the Honduran portion of the range is
badly deforested; it was still common in 1998 in streams at 30-330m asl in extreme north-western Cortés Department
in Honduras. It is uncommon in Guatemala.
Habitat and Ecology This species inhabits lowland and premontane wet forest, living along small tributaries, espe-
cially of the Río Motagua. It is often found sitting on low vegetation, sometimes on the banks of streams. It breeds
by direct development and is not dependent on water for breeding. It probably tolerates some habitat disturbance
because one individual was found in a coffee plantation.
Major Threats Serious habitat degradation is taking place due to agriculture, livestock, logging, and human settle-
ment. However, these threats do not account for the high-altitude declines seen in this species, which could be due
to chytridiomycosis.
Conservation Measures One locality in Honduras is on the edge of Parque Nacional Cerro Azul. Surveys are needed
to determine where other remnant populations survive and how these could be better protected. Further research is
necessary to determine whether chytrid is a real threat.
Notes on taxonomy: This species was previously included in the genus Eleutherodactylus (Crawford and Smith 2005).
Bibliography: Campbell, J.A. (2001), Campbell, J.A. and Savage, J.M. (2000), Crawford, A.J. and Smith, E.N. (2005), McCranie, J.R.
and Wilson, L.D. (2002b)
Data Providers: Manuel Acevedo, Larry David Wilson, Gustavo Cruz, Randy McCranie
Geographic Range This species is known from the departments to chytridiomycosis, so the status of this species should be monitored carefully.
of Antioquia, Choco, Risaralda and Valle del Cauca, on the western Conservation Measures There are several protected areas within the range of the species, including Parque
slopes of the northern half of the Cordillera Occidental in Colombia, Nacional Natural Farallones de Cali.
at moderate elevations (800-1,540m asl). Notes on taxonomy: This species was previously included in the genus Eleutherodactylus (Crawford and Smith 2005).
Population It is a relatively abundant species. Bibliography: Acosta-Galvis, A.R. (2000), Crawford, A.J. and Smith, E.N. (2005), Lynch, J.D. (1990a), Lynch, J.D. (1998b), Lynch, J.D.
Habitat and Ecology An inhabitant of rainforest, it can be found and Duellman, W.E. (1997), Lynch, J.D., Ruiz-Carranza, P.M. and Ardila-Robayo, M.C. (1997), Paez, V.P. et al. (2002), Ruiz-Carranza, P.M.,
along streams, on medium to low perches on vegetation, although Ardila-Robayo, M.C. and Lynch, J.D. (1996), Ruiz-Carranza, P.M., Lynch, J.D. and Ardila-Robayo, M.C. (1997)
adults are usually too heavy to be supported by the leaves so they Data Providers: Fernando Castro, Maria Isabel Herrera, John Lynch
are commonly found on rocks. Provided there is gallery forest present
over streams, it is able to tolerate considerable habitat modification.
Breeding takes place by direct development.
Major Threats Although there are no major threats to the species
overall at present, some localized habitat loss is taking place due
to agricultural development (including the planting of illegal crops).
Some other species of Craugastor that are associated with streams
have undergone dramatic declines and disappearances, possibly due
Geographic Range This species is known only from between occur within the protected area although there are no records to date. Further survey work is required to determine
Laguna del Cerro and Quebrada Grande in the Department of the biology and population status of this species and to determine whether or not it is indeed present in Parque
Copan, in the extreme west of Honduras, at 1,000m asl, close to Nacional Cerro Azul.
the Guatemala border. Notes on taxonomy: This species was previously included in the genus Eleutherodactylus (Crawford and Smith 2005).
Population It is known only from a single specimen. Bibliography: Crawford, A.J. and Smith, E.N. (2005), McCranie, J.R. and Köhler, G. (1999a), McCranie, J.R. and Wilson, L.D. (2002b)
Habitat and Ecology Very little is known of the habitat and ecology Data Providers: Gustavo Cruz, Larry David Wilson, Randy McCranie
of this species, but it is believed to occur in premontane wet forest,
although the holotype was found on the ground in a coffee plantation,
which might suggest some ability to adapt to secondary habitats.
It presumably breeds by direct development and is not dependent
on water for breeding.
Major Threats Primary threats probably include deforestation as a
result of agricultural and livestock encroachment, human settlements,
logging, fires and landslides.
Conservation Measures The only known locality is right on
edge of Parque Nacional Cerro Azul, so the species might well
Geographic Range This species is known only from the type locality, given the threat of chytridiomycosis, surviving individuals might need to form the basis for the establishment of an
the Cordillera Nombre de Dios on the Atlantic slope south of La Ceiba ex-situ population.
in northern Honduras at 1,520m asl. Notes on taxonomy: This species was previously included in the genus Eleutherodactylus (Crawford and Smith 2005).
Population A landslide destroyed the stream where the type speci- Bibliography: Crawford, A.J. and Smith, E.N. (2005), McCranie, J.R. and Wilson, L.D. (2002b), McCranie, J.R., Savage, J.M. and Wilson,
men was collected in 1988. Visits to the area in 1995 and 1996 turned L.D. (1989), Wilson, L.D. and McCranie, J.R. (1998)
up no additional specimens. Data Providers: Larry David Wilson, Randy McCranie, Gustavo Cruz
Habitat and Ecology The two known specimens were collected
along a very small stream in lower montane wet forest.
Major Threats The major threat is likely to be chytridiomycosis,
leading to a catastrophic population decline, as has occurred in
many other high-elevation amphibian species that are associated
with streams. Considering that all streamside Craugastor above
900m asl in Honduras have disappeared recently, this species might
well be extinct.
Conservation Measures The type locality of this species is
within Parque Nacional Pico Bonito. Surveys to determine whether
or not there are surviving populations of this species are needed;
302
EN Craugastor daryi (Ford and Savage, 1984)
Endangered B1ab(iii)
Order, Family: Anura, Leptodactylidae
Country Distribution: Guatemala
Current Population Trend: Decreasing
VU Craugastor decoratus (Taylor, 1942)
Vulnerable B1ab(iii)
Order, Family: Anura, Leptodactylidae
Country Distribution: Mexico
Current Population Trend: Decreasing
© Tim Burkhardt
CR Craugastor emcelae (Lynch, 1985)
Critically Endangered A3ce
Order, Family: Anura, Leptodactylidae
Country Distribution: Panama
Current Population Trend: Decreasing
© Ignacio De La Riva
CR Craugastor epochthidius (McCranie and Wilson, 1997)
Critically Endangered A3ce
Order, Family: Anura, Leptodactylidae
Country Distribution: Honduras
Current Population Trend: Decreasing
Threatened Amphibians of the World
Geographic Range This species can be found in the cloud forest of
the Sierra Xucaneb and the Sierra de las Minas, central Guatemala,
at 1,500-2,290m asl.
Population It is uncommon but still regularly seen in appropriate
habitat.
Habitat and Ecology This species can be found along streams
in cloud forest. Reproduction occurs via direct development. It can
persist in reduced numbers in moderately disturbed areas.
Major Threats A major threat to this species is habitat destruc-
tion, and more than 50% of its habitat has been lost in the last 10
years due to conversion of forest to ornamental plant farms. Some
other species of the genus that are associated with high-elevation
streams have undergone dramatic declines and disappearances,
possibly due to chytridiomycosis, so the status of this species should
be monitored carefully.
Conservation Measures This species has been collected within
Geographic Range This species can be found in Sierra de Agalta
and Sierra Punta Piedra in the departments of Olancho and Colon
east-central and north-eastern Honduras, at elevations of 150-
1,459m asl.
Population Formerly, this species was relatively common, but now
is apparently in decline. Substantial habitat has been lost in recent
years, and a recent survey conducted in appropriate habitat turned
up no records of this species.
Habitat and Ecology It occurs and breeds (by direct development)
in leaf-litter on the ground in lowland and premontane moist forest.
It is often found near streams.
Major Threats The major threat is habitat loss due to agriculture
(crops and cattle ranching), logging, human settlement, and forest
fires. However, since the observed decline has also taken place in
suitable habitat this could be suggestive of chytridiomycosis as a
significant threat (at least to higher-altitude populations).
Biotopo del Quetzal and the Reserva de la Biósfera Sierra de las Minas. Further research is necessary to determine
whether chytrid poses a real threat.
Notes on taxonomy: This species was previously included in the genus Eleutherodactylus (Crawford and Smith 2005).
Bibliography: Campbell, J.A. (2001), Crawford, A.J. and Smith, E.N. (2005), Ford, L.S. and Savage, J.M. (1984)
Data Providers: Manuel Acevedo, Bruce Young
Geographic Range This species is found in southern Tamaulipas, southern San Luis Potosi and adjacent north
Querétaro, and northern Hidalgo to central Veracruz and adjacent northern Puebla, Mexico.
Population This is a rare species.
Habitat and Ecology It inhabits montane cloud forest and several individuals have been found on trees and bushes in
cultivated land, suggesting that it is tolerant of a degree of habitat modification. It breeds by direct development.
Major Threats The major threat is habitat loss and degradation due to logging, as well as some agricultural
activities.
Conservation Measures The range of this species includes the El Cido Biosphere Reserve and Parque Nacional
Cofre de Perote. Improved protection of the montane forests along the Huasteca region in Mexico is needed. It is
protected by Mexican law under the “Special Protection” category (Pr).
Notes on taxonomy: This species was previously included in the genus Eleutherodactylus (Crawford and Smith 2005).
Bibliography: Campbell, J.A., Lamar, W.W. and Hillis, D.M. (1989), Crawford, A.J. and Smith, E.N. (2005)
Data Providers: Georgina Santos-Barrera, Luis Canseco-Márquez
Geographic Range This species is endemic to the western central cordillera of Panama. It has been recorded from
altitudes of between 910 and 1,450m asl.
Population While there is little information available on the population status of this species, it is known to have
declined in the Reserva Forestal Fortuna, Chiriquí.
Habitat and Ecology This is a terrestrial species of humid lowland and montane forest. It breeds by direct develop-
ment, with eggs deposited on the forest floor.
Major Threats The major threat to the species is general habitat loss as a result of smallholder farming activities,
logging and human settlement, although the current observed population decline is thought to be due to chytridio-
mycosis (Lips 1999).
Conservation Measures The species has been recorded from the protected areas of Parque Internacional La Amistad,
Bosque Protector Palo Seco, and Reserva Forestal Fortuna. In view of the threat of chytridiomycosis, the status of this
species should be closely monitored, and ex-situ populations should be established.
Notes on taxonomy: This species was previously included in the genus Eleutherodactylus (Crawford and Smith 2005).
Bibliography: Crawford, A.J. and Smith, E.N. (2005), Ibáñez, R. et al. (2000), Lips, K.R. (1999), Lynch, J.D. (1985), Young, B. et al.
(1999)
Data Providers: Frank Solís, Roberto Ibáñez, César Jaramillo, Querube Fuenmayor
Conservation Measures It has been collected within the Reserve de la Biosfera Río Platano and on the edge
of Parque Nacional Agalta. In view of the threat of chytridiomycosis, the status of this species should be closely
monitored, and ex-situ populations should be established.
Notes on taxonomy: This species was previously included in the genus Eleutherodactylus (Crawford and Smith 2005).
Bibliography: Crawford, A.J. and Smith, E.N. (2005), McCranie, J.R. and Wilson, L.D. (1997a), McCranie, J.R. and Wilson, L.D.
(2002b)
Data Providers: Gustavo Cruz, Larry David Wilson

CR Craugastor escoces (Savage, 1975)
Critically Endangered A2ace
Order, Family: Anura, Leptodactylidae
Country Distribution: Costa Rica
Current Population Trend: Decreasing
© Wayne Van Devender
CR Craugastor fecundus (McCranie and Wilson, 1997)
Critically Endangered A2ace
Order, Family: Anura, Leptodactylidae
Country Distribution: Honduras
Current Population Trend: Decreasing
CR Craugastor fleischmanni (Boettger, 1892)
Critically Endangered A2ace
Order, Family: Anura, Leptodactylidae
Country Distribution: Costa Rica
Current Population Trend: Decreasing
CR Craugastor glaucus (Lynch, 1967)
Critically Endangered B1ab(iii)
Order, Family: Anura, Leptodactylidae
Country Distribution: Mexico
Current Population Trend: Decreasing
© Thomas Bille
Globally Threatened Amphibian Species
Geographic Range This species can be found in Quebrada de Oro
and Cerro Calentura in the Sierra Nombre de Dios in the Depart-
ments of Atlantica and Colon, northern Honduras, at elevations of
200-1,260m asl.
Population Although once relatively common, it is now in decline: it
was not recorded once during a total of 20 days of searching during
1995 and 1996 in suitable habitat.
Habitat and Ecology The species is known to occur in leaf-litter in
undisturbed lowland moist forest and premontane wet forest, and
breeds by direct development.
Major Threats The most likely cause of the apparent disappearance
of this species is chytridiomycosis.
Conservation Measures There are records of this species from
Parque Nacional Pico Bonito, Parque Nacional Capiro and Parque
Nacional Calentura. Surveys to relocate this species and to deter-
mine its current population status are urgently required; surviving
Geographic Range This species is found in Costa Rica on the
Meseta Central Oriental and Occidental, on the Pacific slopes of the
volcanoes Poás and Barva, on the Atlantic slopes of the volcanoes
Irazú, Turrialba, and on the slopes of the Cordillera de Talamanca,
at elevations of 1,050-2,286m asl.
Population It has not been seen since the 1980s despite extensive
searches in sites from which it was known. It appears that a serious
decline has taken place.
Habitat and Ecology It is a nocturnal species that has been found
along streams in premontane and lower montane wet forest at both
pristine and moderately disturbed sites. It reproduces in stream
margins by direct development.
Major Threats The most likely cause of the apparent disappear-
ance of this species is chytridiomycosis, possibly in synergy with the
effects of climate change.
303
Geographic Range This species occurs on the volcanoes Barva, Irazú, and Turrialba, of the Cordillera Central of
Costa Rica, at elevations of 1,100-2,100m asl (Savage 2002).
Population Although it has been well studied and collected extensively, and was formerly abundant throughout its
range, it has not been recorded since 1986. A serious decline appears to have taken place.
Habitat and Ecology It has been found along streams in premontane and lower montane rainforest; breedin takes
place on land by direct development along stream margins.
Major Threats The most likely cause of its disappearance in suitable habitats is chytridiomycosis, perhaps in
combination with climate change.
Conservation Measures Further survey work is required to determine whether or not this species still survives in
the wild and the reasons for its decline. It is well protected in Parque Nacional Braulio Carrillo, but in view of the
likely threat of chytridiomycosis, any surviving individuals might need to form the basis for the establishment of a
captive-breeding programme.
Notes on taxonomy: This species was previously included in the genus Eleutherodactylus (Crawford and Smith 2005).
Bibliography: Campbell, J.A. and Savage, J.M. (2000), Crawford, A.J. and Smith, E.N. (2005), Savage, J.M. (2002)
Data Providers: Federico Bolaños, Gerardo Chaves
individuals might need to form the basis for the establishment of an ex-situ population.
Notes on taxonomy: This species was previously included in the genus Eleutherodactylus (Crawford and Smith 2005).
Bibliography: Crawford, A.J. and Smith, E.N. (2005), McCranie, J.R. and Wilson, L.D. (1997a), McCranie, J.R. and Wilson, L.D.
(2002b)
Data Providers: Gustavo Cruz, Larry David Wilson
Conservation Measures Its range includes Parque Nacional Tapantí as well as other protected areas. It is a high
priority to conduct surveys to relocate this species and determine whether or not it survives in the wild; in view of
the threat of chytridiomycosis, surviving individuals might need to form the basis for the establishment of an ex-situ
population.
Notes on taxonomy: This species was previously included in the genus Eleutherodactylus (Crawford and Smith 2005).
Bibliography: Campbell, J.A. and Savage, J.M. (2000), Crawford, A.J. and Smith, E.N. (2005), Savage, J.M. (2002)
Data Providers: Federico Bolaños, Gerardo Chaves, Jay Savage
Geographic Range This species is known from the areas surrounding San Cristobal de las Casas, Chiapas, Mexico,
at 2,100m asl.
Population It is a rare species.
Habitat and Ecology The primary habitat is pine and pine-oak forests at intermediate elevations. It breeds by
direct development.
Major Threats Most of its range has been dramatically transformed in the past few years by an increased rate of
logging and urbanization.
Conservation Measures It is not known to occur in any protected areas, and restoration and protection of the
remaining habitat of this species is urgently required. This species is protected by Mexican law under the “Special
Protection” category (Pr).
Notes on taxonomy: This species was previously included in the genus Eleutherodactylus (Crawford and Smith 2005).
Bibliography: Campbell, J.A., Lamar, W.W. and Hillis, D.M. (1989), Crawford, A.J. and Smith, E.N. (2005)
Data Providers: Georgina Santos-Barrera, Luis Canseco-Márquez
304
CR Craugastor greggi (Bumzahem, 1955)
Critically Endangered A3e
Order, Family: Anura, Leptodactylidae
Country Distribution: Guatemala, Mexico
Current Population Trend: Decreasing
CR Craugastor guerreroensis (Lynch, 1967)
Critically Endangered A2ace
Order, Family: Anura, Leptodactylidae
Country Distribution: Mexico
Current Population Trend: Decreasing
© Tim Burkhardt
EN Craugastor gulosus (Cope, 1875)
Endangered B1ab(iii)
Order, Family: Anura, Leptodactylidae
Country Distribution: Costa Rica, Panama
Current Population Trend: Decreasing
EN Craugastor hobartsmithi (Taylor, 1936)
Endangered B1ab(iii)
Order, Family: Anura, Leptodactylidae
Country Distribution: Mexico
Current Population Trend: Decreasing
Threatened Amphibians of the World
Geographic Range This species occurs in the Soconusco region Conservation Measures It is not found within any protected areas, and protection of the forested areas along the
and slopes of Volcán Tacana, southern Chiapas, Mexico, and Soconusco region in Mexico is clearly necessary. In view of the threat of chytridiomycosis, the status of this species
adjacent western Guatemala at Volcán Tejumulco, at elevations of should be closely monitored, and ex-situ populations should be established. This species is protected by Mexican
1,500-2,700m asl. law under the “Special Protection” category (Pr).
Population It is uncommon and is known only from few localities. Notes on taxonomy: This species was previously included in the genus Eleutherodactylus (Crawford and Smith 2005).
The population in Guatemala is known to be declining, and there Bibliography: Campbell, J.A. (2001), Crawford, A.J. and Smith, E.N. (2005), Lips, K.R. et al. (2004)
have been no records from Mexico since 1997 (recent surveys in Data Providers: Georgina Santos-Barrera, Manuel Acevedo, Antonio Muñoz Alonso
Chiapas have not been successful in locating it).
Habitat and Ecology The primary habitat is montane cloud forests,
and it breeds near streams and does so via direct development.
Major Threats It is likely that this species is declining because of
chytridiomycosis, which has heavily impacted some other streamside
Craugastor species. Habitat loss is also a threat to the species,
particularly in Mexico where the population occurs in a continu-
ously altered forest.
Geographic Range This species is known only from Agua del Obispo at 980m asl, central Guerrero, Mexico, although
it is likely to occur a little more widely.
Population This has always been a rare species, but it appears to have gone into serious decline, and has not seen
recorded since 1984. Recent surveys to locate it have been unsuccessful, and it might now be extinct.
Habitat and Ecology It has been recorded inhabiting pine-oak forests, and breeds by direct development.
Major Threats Loss of original forest cover due to conversion to agricultural land, and as a result of excessive
logging, has had an impact of this species. However, it has also disappeared in suitable habitat, probably due to
chytridiomycosis.
Conservation Measures This species is a priority for further survey work to determine whether or not it might still
survive in the wild, and surviving populations might need to form the basis for a captive-breeding programme. The
range of this species does not include any protected areas, and protection of the Agua del Obispo forests is urgent
since there are many threatened amphibians known to occur in this area. It is protected by Mexican law under the
“Special Protection” category (Pr).
Notes on taxonomy: This species was previously included in the genus Eleutherodactylus (Crawford and Smith 2005).
Bibliography: Campbell, J.A., Lamar, W.W. and Hillis, D.M. (1989), Crawford, A.J. and Smith, E.N. (2005), Lips, K.R. et al. (2004)
Data Providers: Georgina Santos-Barrera, Luis Canseco-Márquez
Geographic Range This species can be found in the southern the borderland of Costa Rica and Panama (Savage and Myers 2002).
Cordillera de Talamanca of Costa Rica and western Panama, at Bibliography: Crawford, A.J. and Smith, E.N. (2005), Savage, J.M. (2002), Savage, J.M. and Myers, C.W. (2002)
elevations of 1,000-1,873m asl (Savage 2002). Data Providers: Frank Solís, Roberto Ibáñez, Gerardo Chaves, Jay Savage, César Jaramillo, Querube Fuenmayor
Population It is generally a rare species.
Habitat and Ecology It inhabits humid premontane forest. Little
is known about the species’ biology, but it is presumed to breed by
direct development.
Major Threats The main threat is general habitat loss primarily due
to agriculture and logging.
Conservation Measures The species has been recorded from
the protected areas of Parque Internacional La Amistad and the
Reserva Forestal Fortuna. Further research is needed into the range
and breeding ecology of this species.
Notes on taxonomy: This species was previously included in the genus
Eleutherodactylus (Crawford and Smith 2005). The rediscovery of this species
showed it to be a largely montane frog, occupying an apparently small range in
Geographic Range This Mexican endemic is known from the Notes on taxonomy: This species was previously included in the genus Eleutherodactylus (Crawford and Smith 2005).
south-western portion of the Mexican plateau, from Nayarit and Bibliography: Crawford, A.J. and Smith, E.N. (2005), Flores-Villela, O.A., Mendoza-Quijano, F. and Gonzalez-Porter, G. (1995), Taylor,
Jalisco westward into Michoacán and the adjacent eastern state E.H. (1939)
of Mexico. Data Providers: Georgina Santos-Barrera, Oscar Flores-Villela
Population There is no information on its population status, but it
is believed to be declining.
Habitat and Ecology This species occurs in lowland deciduous
seasonal forest and even in lowland dry mixed forests. It breeds by
direct development.
Major Threats The major threat is habitat loss and disturbance due
to smallholder farming, logging, and human settlement.
Conservation Measures There is a need for improved and
expanded protection of the seasonal forest areas along Pacific
coastal Mexico as well as the inland mixed forests. Further survey
work is also necessary to determine the current population status
of this species.

EN Craugastor inachus (Campbell and Savage, 2000)
Endangered B1ab(iii)
Order, Family: Anura, Leptodactylidae
Country Distribution: Guatemala
Current Population Trend: Decreasing
EN Craugastor lauraster (Savage, McCranie and Espinal, 1996)
Endangered B1ab(iii,v)
Order, Family: Anura, Leptodactylidae
Country Distribution: Honduras, Nicaragua
Current Population Trend: Decreasing
© Twan Leenders
CR Craugastor lineatus (Brocchi, 1879)
Critically Endangered A3e
Order, Family: Anura, Leptodactylidae
Country Distribution: Guatemala, Mexico
Current Population Trend: Decreasing
© Tim Burkhardt
VU Craugastor matudai (Taylor, 1941)
Vulnerable D2
Order, Family: Anura, Leptodactylidae
Country Distribution: Guatemala, Mexico
Current Population Trend: Stable
Globally Threatened Amphibian Species
Geographic Range This species can be found in mid Motagua
Valley and associated areas in central and eastern Guatemala and
the slopes of the Salamá Basin in Baja Verapaz, at elevations of
500-1,400m asl. It is likely to occur somewhat more widely.
Population It is an uncommon species, but it is still seen regu-
larly.
Habitat and Ecology It is restricted to narrow riparian corridors
in premontane dry forest. No other member of the genus occurs in
such dry areas in this part of Central America. Reproduction occurs
via direct development.
Major Threats The main threat to this species is habitat loss due
to agricultural activities and extraction of wood. Some other spe-
cies of the genus that are associated with high-elevation streams
have undergone dramatic declines and disappearances, possibly
due to chytridiomycosis, so the status of this species should be
monitored carefully.
Geographic Range This species is known from Cerro Ovando,
south-western Chiapas, Mexico, and from one location in the Pacific
versant at Aldea Fraternidad, northern Guatemala. It occurs from
1,500-2,000m asl.
Population This is a rare species.
Habitat and Ecology It is a terrestrial species inhabiting pine-oak
forest at moderate elevations. It breeds by direct development.
Major Threats At present, the pine-oak habitat in the only two
locations where this species occurs is still intact. However, logging
represents a potential threat.
Conservation Measures The range of this species does not include
any protected areas, and protection of the montane forests at Cerro
Ovando is needed. It is protected by Mexican law under the “Special
Protection” category (Pr).
Notes on taxonomy: This species was previously included in the genus
Eleutherodactylus (Crawford and Smith 2005).
305
Conservation Measures It occurs outside any protected areas although the Motagua Valley has been proposed for
a new national park. There is an urgent need to expand the protected areas network to include additional tracts of
suitable remaining habitat in the range of this species. Further research is necessary to determine whether chytrid
poses a threat.
Notes on taxonomy: This species was previously included in the genus Eleutherodactylus (Crawford and Smith 2005).
Bibliography: Campbell, J.A. and Savage, J.M. (2000), Crawford, A.J. and Smith, E.N. (2005)
Data Providers: Manuel Acevedo, Bruce Young
Geographic Range This species occurs from central and eastern Honduras on the Atlantic versant to northern
Nicaragua, at 40-1,200m asl.
Population It can be very common, but it is undergoing severe declines in Nicaragua at higher elevations.
Habitat and Ecology It lives on the ground of lowland moist forest and premontane wet forest, and can survive in
somewhat degraded habitats providing there is shade over leaf-litter. It presumably breeds by direct development
and is not dependent on water for breeding.
Major Threats The main threats to its habitat come from agriculture, cattle ranching, logging, forest fires, and severe
dry seasons. The causes of the declines at high altitudes are not understood.
Conservation Measures It occurs in several protected areas in both countries, although improved and strengthened
management of these is needed. Further research is needed to investigate the reason for this species’ decline at
higher elevations.
Notes on taxonomy: This species was previously included in the genus Eleutherodactylus (Crawford and Smith 2005).
Bibliography: Campbell, J.A. (2001), Crawford, A.J. and Smith, E.N. (2005), Köhler, G. (1998), Köhler, G. (2001), McCranie, J.R. and
Wilson, L.D. (2002b), Savage, J.M., McCranie, J.R. and Espinal, M. (1996)
Data Providers: Gustavo Cruz, Larry David Wilson, Randy McCranie, Gunther Köhler
Geographic Range This species can be found on the Atlantic versant of Guerrero, Oaxaca and Chiapas, Mexico,
southeast to Guatemala. It also occurs on the Pacific versant, from eastern Oaxaca through Chiapas to the south-
western highlands of Guatemala, at elevations of 300-2,000m asl.
Population There are very few records of this species and it appears to be uncommon. In Chiapas, it is currently
known only from a single site. Recent surveys in Guerrero and Oaxaca indicate that it has disappeared from all sites
surveyed, suggesting a serious decrease.
Habitat and Ecology It can be found in lower montane evergreen forest, and it occurs and breeds by direct develop-
ment near streams.
Major Threats The declines witnessed in Mexico are probably due to chytridiomycosis, since some other species of
Craugastor that are associated with streams have undergone dramatic declines and disappearances, possibly due to
chytridiomycosis. Habitat loss due to agriculture, logging, and human settlement is also a threat.
Conservation Measures This species is known to occur in Biotopo del Quetzal and the Reserva de la Biósfera Bisis
Cabá. In view of the severe risk of chytridiomycosis, the status of this species should be closely monitored, and ex-situ
populations should be established. It is protected by Mexican law under the “Special Protection” category (Pr).
Notes on taxonomy: This species was previously included in the genus Eleutherodactylus (Crawford and Smith 2005).
Bibliography: Crawford, A.J. and Smith, E.N. (2005), Hoogmoed, M.S. (1986), Lips, K.R. et al. (2004), Regos, J. and Schluter, A. (1984),
Savage, J.M. (1987), Schluter, A. and Regos, J. (1981)
Data Providers: Georgina Santos-Barrera, Manuel Acevedo, Antonio Muñoz Alonso
Bibliography: Crawford, A.J. and Smith, E.N. (2005), Taylor, E.H. (1941)
Data Providers: Georgina Santos-Barrera, Luis Canseco-Márquez
306
CR Craugastor megalotympanum (Shannon and Werler, 1955)
Critically Endangered B1ab(iii)
Order, Family: Anura, Leptodactylidae
Country Distribution: Mexico
Current Population Trend: Decreasing
VU Craugastor melanostictus (Cope, 1875)
Vulnerable B1ab(iii)
Order, Family: Anura, Leptodactylidae
Country Distribution: Costa Rica, Panama
Current Population Trend: Decreasing
© Wayne Van Devender
CR Craugastor merendonensis (K. Schmidt, 1933)
Critically Endangered A2ace; B1ab(v)+2ab(v)
Order, Family: Anura, Leptodactylidae
Country Distribution: Honduras
Current Population Trend: Decreasing
VU Craugastor necerus (Lynch, 1975)
Vulnerable B1ab(iii)
Order, Family: Anura, Leptodactylidae
Country Distribution: Ecuador
Current Population Trend: Decreasing
Threatened Amphibians of the World
Geographic Range This species is known only from Sierra de Los Notes on taxonomy: This species was previously included in the genus Eleutherodactylus (Crawford and Smith 2005).
Tuxtlas, southern Veracruz, Mexico. The type locality is recorded Bibliography: Campbell, J.A., Lamar, W.W. and Hillis, D.M. (1989), Crawford, A.J. and Smith, E.N. (2005)
as 900-1,200m asl. Data Providers: Georgina Santos-Barrera, Oscar Flores-Villela
Population It is an uncommon species.
Habitat and Ecology The primary habitat is lowland tropical humid
rainforest; it seems to live on the ground and under leaf-litter. Breed-
ing takes place by direct development.
Major Threats The high rate of transformation of original forest
habitat in this area is notorious, and deforestation for the cultivation
of coffee, fruits and tobacco is rapidly increasing.
Conservation Measures The range of this species is wholly within
Reserva de la Biósfera Sierra de Los Tuxtlas, although logging is taking
place within this area. A survey to evaluate the population status of this
population in Los Tuxtlas is needed, and real protection of the Sierra
de Los Tuxtlas and the surrounding areas is urgent. This species is pro-
tected by Mexican law under the “Special Protection” category (Pr).
Geographic Range This species is found in the cordilleras of Costa Rica and the highlands of central Panama, at
elevations of 1,150-2,700m asl (Savage 2002).
Population In Costa Rica, this species is rarely seen and there is little information on its population status. Lips
(1998) reported a sharp decline at Las Tablas (Panama), but populations have persisted at Monteverde (Costa Rica)
(Pounds et al. 1997).
Habitat and Ecology It inhabits relatively undisturbed, dense, primary humid montane forest. Breeding takes place
by direct development.
Major Threats In Panama, the major threat is logging. The cause of the decline at Las Tablas has not been determined,
but chytridiomycosis is probable, as it was detected in this species at this location.
Conservation Measures It has been recorded from a number of protected areas in Costa Rica (including Parque
Nacional Braulio Carrillo, and Parque Internacional La Amistad) and Panama (Parque Nacional Chagres and Parque
Nacional Volcán Barú).
Notes on taxonomy: This species was previously included in the genus Eleutherodactylus (Crawford and Smith 2005). Populations in
Panama might belong to a different species.
Bibliography: Crawford, A.J. and Smith, E.N. (2005), Ibáñez, R. et al. (2000), Lips, K.R. (1998), Lips, K.R., Green, D.E. and Papendick, R.
(2003), Lips, K.R., Reeve, J.D. and Witters, L.R. (2003), Miyamoto, M.M. and Tennant, M.R. (1984), Pounds, J.A. et al. (1997), Savage,
J.M. (2002), Savage, J.M. and DeWeese, J.E. (1981)
Data Providers: Frank Solís, Roberto Ibáñez, Gerardo Chaves, Jay Savage, César Jaramillo, Querube Fuenmayor
Geographic Range This species is known only from Montañas del be a major threat to this species. On the other hand, this species occurs at a much lower altitude than most frog
Merendon on the Sierra de Omoa, west of San Pedro Sula in the species that have been affected by such unexplained declines.
Department of Cortes, Honduras, at elevations of 150-200m asl. Conservation Measures Further survey work is required to determine the current population status of this species
Population It was formerly relatively common but is now in pre- (including whether or not it even still survives in the wild), and particularly to establish the reasons for its decline.
cipitous decline. It was not recorded in the course of visits to the Notes on taxonomy: This species was previously included in the genus Eleutherodactylus (Crawford and Smith 2005).
known range of the species in 1996 and 1998 where the habitat Bibliography: Campbell, J.A. and Savage, J.M. (2000), Crawford, A.J. and Smith, E.N. (2005), McCranie, J.R. and Wilson, L.D. (2002b),
remains relatively intact. Schmidt, K.P. (1933)
Habitat and Ecology It occurs and reproduces by direct develop- Data Providers: Gustavo Cruz, Larry David Wilson
ment in leaf-litter on the ground of lowland moist forest, and is often
found in the vicinity of streams.
Major Threats Since the range of this species is well protected
from obvious human disturbance (the area from which it is known
has been managed and protected for the last 30 years), the cause
of its decline is unclear. Some other species of Craugastor that are
associated with streams have undergone dramatic declines and
disappearances, possibly due to chytridiomycosis, and so this might
Geographic Range This species is known from more than ten chytridiomycosis, so the status of this species should be monitored carefully.
localities on the lower Pacific slopes of the Andes, from the province Conservation Measures Its range overlaps slightly with Reserva Ecológica Cotacachi-Cayapas and Reserva
of Cotopaxi northwards to the province of Carchi in Ecuador. It is also Ecológica Los Illinizas.
likely to occur in adjacent Colombia, although there are, as yet, no Notes on taxonomy: This species was previously included in the genus Eleutherodactylus (Crawford and Smith 2005).
confirmed records. It ranges from 600-1,540m asl. Bibliography: Crawford, A.J. and Smith, E.N. (2005), Hedges, S.B. (1989), Lynch, J.D. (1975b), Lynch, J.D. and Duellman, W.E. (1997)
Population It is a rare species. Data Providers: Luis A. Coloma, Santiago Ron, Diego Cisneros-Heredia, Ana Almandáriz
Habitat and Ecology An inhabitant of humid premontane forest;
most individuals have been observed in the immediate vicinity of
streams (Lynch and Duellman 1997). Breeding takes place by direct
development.
Major Threats The major threat to this species is habitat loss and
degradation, which has been particularly severe in the last few years
due to agriculture (both crops and livestock) and logging; pollution
due to agriculture is also a threat. It should be noted that some
other species of Craugastor that are associated with streams have
undergone dramatic declines and disappearances, possibly due to

EN Craugastor obesus (Barbour, 1928)
Endangered B1ab(iii,v)
Order, Family: Anura, Leptodactylidae
Country Distribution: Costa Rica, Panama
Current Population Trend: Decreasing
© Karl-Heinz Jungfer
EN Craugastor omiltemanus (Günther, 1901)
Endangered B1ab(iii)
Order, Family: Anura, Leptodactylidae
Country Distribution: Mexico
Current Population Trend: Decreasing
CR Craugastor omoaensis (McCranie and Wilson, 1997)
Critically Endangered A2ace; B1ab(iii)
Order, Family: Anura, Leptodactylidae
Country Distribution: Honduras
Current Population Trend: Decreasing
VU Craugastor persimilis (Barbour, 1926)
Vulnerable B1ab(iii)
Order, Family: Anura, Leptodactylidae
Country Distribution: Costa Rica
Current Population Trend: Decreasing
© Brian Kubicki
Globally Threatened Amphibian Species
Geographic Range This species is known only from the Sierra
Madre del Sur, central Guerrero, Mexico, at altitudes of 2,200-
2,600m asl.
Population This is a relatively common species.
Habitat and Ecology It inhabits pine, oak and pine-oak forests,
and requires an abundance of leaf-litter on the ground. It breeds
by direct development.
Major Threats Habitat loss and disturbance due to smallholder
farming activities, livestock and removal of wood through logging
and subsistence collection, is having a negative impact on the
microhabitat of this species.
Conservation Measures A survey to determine the presence of
this species in the only protected area within its range, Omiltemi
Park, is required. No records have been reported recently, even
when the park was under state protection. Recently, the park has
been subject to human activities that jeopardize the survival of the
Geographic Range This species can be found at moderate eleva-
tions of the Sierra de Omoa west-south-west of San Pedro Sula
on the Atlantic versant of north-western Honduras, at elevations
of 760-1,150m asl.
Population It is known from 24 specimens. Formerly, it was moder-
ately common, but is now undergoing a precipitous decline. A visit
to the area of the type locality in 1996 failed to turn up individuals
of this or any other amphibian.
Habitat and Ecology The species occurs in, and breeds by direct
development along, streams in premontane wet forest.
Major Threats Although habitat loss is certainly affecting this
species, this does not explain the dramatic decline that has taken
place. Some other species of Craugastor that are associated with
streams have undergone dramatic declines and disappearances at
higher elevations, possibly due to chytridiomycosis, and so this might
be a major threat to this species.
307
Geographic Range This species occurs on the Atlantic slopes of south-eastern Costa Rica and north-western
Panama at an altitude of 400-1,450m asl.
Population It is rare in Costa Rica and is believed to have declined in the Reserva Forestal Fortuna, Chiriquí,
Panama.
Habitat and Ecology This species can be found in the spray zone on rocks, boulders, and cliff faces in the middle of
moderate-sized cascading streams in lowland moist forest, premontane and lower montane wet forest and rainforest.
It does not occur in modified habitats. It presumably breeds by direct development.
Major Threats The main threat to this species is believed to be chytridiomycosis, and it has already disappeared
from one site where chytrid fungus was found.
Conservation Measures It has been recorded from Parque Internacional La Amistad (Costa Rica and Panama) and
two protected areas in Panama, and subpopulations within these seem relatively secure. Further research is needed
into the range and ecology of this species, as well as the impact of chytridiomycosis.
Notes on taxonomy: This species was previously included in the genus Eleutherodactylus (Crawford and Smith 2005). This species
was removed from synonymy of Craugastor punctariolus (Campbell and Savage 2000).
Bibliography: Campbell, J.A. and Savage, J.M. (2000), Crawford, A.J. and Smith, E.N. (2005), Savage, J.M. (2002)
Data Providers: Frank Solís, Roberto Ibáñez, Gerardo Chaves, Jay Savage, César Jaramillo, Querube Fuenmayor, Federico Bolaños
native fauna and flora. This species is in urgent need of habitat protection. It is protected by Mexican law under the
“Special Protection” category (Pr).
Notes on taxonomy: This species was previously included in the genus Eleutherodactylus (Crawford and Smith 2005).
Bibliography: Crawford, A.J. and Smith, E.N. (2005), Lynch, J.D. (2000b)
Data Providers: Georgina Santos-Barrera, Oscar Flores-Villela
Conservation Measures It is not known to occur in any protected areas. It is a very high priority to conduct surveys
to relocate this species and determine its current population status; in view of the risk of chytridiomycosis, surviving
individuals might need to form the basis for the establishment of an ex-situ population.
Notes on taxonomy: This species was previously included in the genus Eleutherodactylus (Crawford and Smith 2005).
Bibliography: Crawford, A.J. and Smith, E.N. (2005), McCranie, J.R. and Wilson, L.D. (1997a)
Data Providers: Gustavo Cruz, Larry David Wilson
Geographic Range This species ranges from central to south-eastern Costa Rica, on the Atlantic slopes at eleva-
tions of 39-1,200m asl.
Population It is collected occasionally. It can experience marked population fluctuations from year to year.
Habitat and Ecology It occurs in leaf-litter of lowland and premontane moist rainforest. It persists in moderately
disturbed areas and breeds on the forest floor.
Major Threats Habitat destruction, due to agriculture and logging, is the major threat to this species.
Conservation Measures It is present in several protected areas, including Parque Nacional Barbilla.
Notes on taxonomy: This species was previously included in the genus Eleutherodactylus (Crawford and Smith 2005). It was recently
separated from Eleutherodactylus bransfordii.
Bibliography: Crawford, A.J. (2003), Crawford, A.J. and Smith, E.N. (2005), Savage, J.M. (2002)
Data Providers: Federico Bolaños, Gerardo Chaves, Jay Savage
308
VU Craugastor podiciferus (Cope, 1876)
Vulnerable B1ab(iii)
Order, Family: Anura, Leptodactylidae
Country Distribution: Costa Rica, Panama
Current Population Trend: Decreasing
© Andrew Crawford
Geographic Range This species is fairly widespread on both slopes of the cordilleras of Costa Rica and adjacent
western Panama, at elevations of 1,089-2,650m asl (Savage 2002).
Population It is generally common and regularly recorded in Costa Rica (Pounds et al. 1997; Lips 1998), although
there are indications that it has undergone declines at several sites at which it was formerly common, such as the
Reserva Forestal Fortuna, Chiriquí, in Panama.
CR Craugastor polymniae (Campbell, Lamar and Hillis, 1989)
Critically Endangered A2ace
Order, Family: Anura, Leptodactylidae
Country Distribution: Mexico
Current Population Trend: Decreasing
CR Craugastor pozo (Johnson and Savage, 1995)
Critically Endangered B1ab(iii)
Order, Family: Anura, Leptodactylidae
Country Distribution: Mexico
Current Population Trend: Decreasing
VU Craugastor psephosypharus (Campbell, Savage and Meyer, 1994)
Vulnerable B1ab(iii)
Order, Family: Anura, Leptodactylidae
Country Distribution: Belize, Guatemala
Current Population Trend: Decreasing
Threatened Amphibians of the World
Habitat and Ecology It is a diurnal species inhabiting leaf-litter in primary humid premontane and montane forest.
It presumably breeds by direct development.
Major Threats Declines of this species have been recorded in both Panama and Costa Rica. For example, in Panama
a massive die-off of amphibians has been observed at the Reserva Forestal Fortuna, with chytridiomycosis suspected
as the major causal factor (Lips 1999), and they are now no longer common at Las Tablas in Costa Rica where they
were formerly abundant (K. Lips pers. comm.). On the other hand, museum specimens of this species collected from
La Selva-Volcán Barva (now in the Parque Nacional Braulio Carrillo) in 1986 have been found infected with chytrid
fungi (Puschendorf, Bolaños and Chaves 2006), yet the species appears to have persisted in this area, where strong
declines have taken place amongst other species (R. Puschendorf pers. comm.).
Conservation Measures The species has been recorded from many protected areas in Costa Rica and Panama,
including Parque Internacional La Amistad and Parque Nacional Braulio Carrillo. Further research is needed into the
possible decline of the species at Reserva Forestal Fortuna, and the potential impacts of the chytrid pathogen on
this species.
Notes on taxonomy: This species was previously included in the genus Eleutherodactylus (Crawford and Smith 2005). This form
might represent a complex of several species.
Bibliography: Crawford, A.J. and Smith, E.N. (2005), Ibáñez, R. et al. (2000), Lips, K.R. (1998), Lips, K.R. (1999), Pounds, J.A. et al. (1997),
Puschendorf, R., Bolaños, F. and Chaves, G. (2006), Savage, J.M. (2002), Schlaepfer, M.A. (1998), Schlaepfer, M.A. and Figeroa-Sandi,
R. (1998), Schlaepfer, M.A. and Gavin, T.A. (2001)
Data Providers: Frank Solís, Roberto Ibáñez, Gerardo Chaves, Jay Savage, César Jaramillo, Querube Fuenmayor
Geographic Range This species is known only from Vista Hermosa ity if the risk of chytridiomycosis proves genuine. The range of this species does not include any protected areas, and
town, Sierra de Juárez, north-central Oaxaca, Mexico, at about protection and restoration of the remaining cloud forest is urgently needed in the Sierra de Juárez where no protected
1,500m asl. areas currently exist. This species is protected by Mexican law under the “Special Protection” category (Pr).
Population This has always been a rare species, but it appears to Notes on taxonomy: This species was previously included in the genus Eleutherodactylus (Crawford and Smith 2005).
have gone into serious decline, and has not been recorded since Bibliography: Campbell, J.A., Lamar, W.W. and Hillis, D.M. (1989), Crawford, A.J. and Smith, E.N. (2005), Lips, K.R. et al. (2004)
1983. Recent surveys to locate it have been unsuccessful, and it Data Providers: Georgina Santos-Barrera, Gabriela Parra-Olea
might now be extinct.
Habitat and Ecology This species inhabits cloud forest; it appears
to have arboreal habits and breeds by direct development.
Major Threats Although disturbance and clear cutting of montane
cloud forest habitats in Sierra de Juárez (primarily for agriculture)
represent a threat to this species, it has disappeared even in suitable
habitat, probably due to chytridiomycosis.
Conservation Measures This species is an urgent priority for survey
work in order to determine whether or not it still survives in the wild,
and any surviving populations might need to be maintained in captiv-
Geographic Range This species is known only from the western Notes on taxonomy: This species was previously included in the genus Eleutherodactylus (Crawford and Smith 2005).
foothills and highlands of Chiapas, Mexico, at 760-1,100m asl, in an Bibliography: Campbell, J.A. and Savage, J.M. (2000), Crawford, A.J. and Smith, E.N. (2005), Johnson, J.D. and Savage, J.M. (1995)
area bordered by Cintalapa, Berriozabal and Presa Nezahualcoyotl. Data Providers: Georgina Santos-Barrera, Gabriela Parra-Olea
Population This is a rare species.
Habitat and Ecology It is associated with wet forest areas, and
inhabits the forest floor under leaf-litter among limestone boulders
and limestone caves. It breeds by direct development.
Major Threats The primary threat to this species is reduction of
original forest cover, since it occurs in an area of high human-induced
disturbance, and its only known habitat covers a very small area.
Conservation Measures Currently, the species is not known to oc-
cur in any protected area, and so protection of the only highland forest
where this species is found is essential to ensure its persistence.
Geographic Range This species is known only from central and Notes on taxonomy: This species was previously included in the genus Eleutherodactylus (Crawford and Smith 2005).
eastern Guatemala, and adjacent southern Belize, at 150-1,170m asl. Bibliography: Campbell, J.A. (1998), Campbell, J.A. (2001), Campbell, J.A. and Savage, J.M. (2000), Campbell, J.A., Savage, J.M. and
It might occur in Honduras, but has not yet been recorded. Meyer, J.R. (1994), Crawford, A.J. and Smith, E.N. (2005), Lee, J.C. (1996), Lee, J.C. (2000)
Population It is rarely encountered, but is sometimes locally Data Providers: Julian Lee, Paul Walker, Manuel Acevedo
common.
Habitat and Ecology It inhabits undisturbed lowland and premon-
tane moist and wet forests, and is often found far from streams. It
is sometimes found in limestone outcrops and caves. It presumably
breeds by direct development and is not dependent on water for
breeding.
Major Threats The major threat is habitat loss and degradation
due to agriculture and logging.
Conservation Measures It occurs in the Columbia River Forest Re-
serve, and possibly in the Bladen Nature Preserve and the Chiquipul
Forest Reserve, in Belize. It has not been recorded from any protected
areas in Guatemala, but might occur in Parque Nacional Río Dulce.
 Globally Threatened Amphibian Species 309

EN Craugastor punctariolus (Peters, 1863)

Endangered B1ab(iii) Geographic Range This species is endemic to the western and central cordilleras of Panama, and is present at
Order, Family: Anura, Leptodactylidae altitudes of between 560 and 1,000m asl.
Country Distribution: Panama Population It is reported to be common where it occurs.
Current Population Trend: Decreasing Habitat and Ecology It is a semi-aquatic species associated with humid montane forest streams. It breeds by direct
development, although the site of egg deposition is not known.
Major Threats Although general habitat loss through deforestation is a recognized threat, pathogens, possibly
including the chytrid fungus, are also considered to be a threat.
Conservation Measures It has been recorded from several protected areas; further research is urgently required
into the possible ongoing impact of pathogens on populations of this species.
Notes on taxonomy: This species was previously included in the genus Eleutherodactylus (Crawford and Smith 2005).
Bibliography: Campbell, J.A. and Savage, J.M. (2000), Crawford, A.J. and Smith, E.N. (2005), Ibáñez, R. et al. (2000)
Data Providers: Frank Solís, Roberto Ibáñez, César Jaramillo, Querube Fuenmayor
© Brian Kubicki

VU Craugastor pygmaeus (Taylor, 1937)

Vulnerable B1ab(iii) Geographic Range This species is known from extreme southern Michoacán and México (state) in Mexico, south
Order, Family: Anura, Leptodactylidae and east along the Pacific slope to western Guatemala. It also occurs in Veracruz and eastern Oaxaca, Mexico, on
Country Distribution: Guatemala, Mexico the Atlantic slope. In Guatemala it is known from above 2,000m asl.
Current Population Trend: Decreasing Population It is a common species.
Habitat and Ecology It occurs in a great variety of habitats, from lowland forests (tropical rainforest, tropical semi-
deciduous forest) to montane pine and pine-oak forests, and cloud forests. It can survive in secondary forests, but
not in open habitats. It breeds by direct development.
Major Threats The major threat is the clearance of forest habitat for agriculture and as a result of logging, which
is probably leading to the disappearance of shade, humidity and leaf-litter microhabitats on which the species
depends.
Conservation Measures This species occurs in Reserva de la Biósfera El Triunfo in Mexico.
Notes on taxonomy: This species was previously included in the genus Eleutherodactylus (Crawford and Smith 2005).
© Tim Burkhardt

Bibliography: Crawford, A.J. and Smith, E.N. (2005), Taylor, E.H. (1939)
Data Providers: Georgina Santos-Barrera, Luis Canseco-Márquez

CR Craugastor ranoides (Cope, 1886)

Critically Endangered A2ace where it was formerly common, such as Volcán Cacao, have been searched repeatedly in the last decade without
Order, Family: Anura, Leptodactylidae success. A small population persists in the Río Murciélago, in the Guanacaste Conservation Area, where several
Country Distribution: Costa Rica, Nicaragua, individuals were recorded in 1994, 1995, and 2003 (Sasa and Solórzano 1995; M. Sasa unpubl.; Puschendorf et al.
Panama 2005). There is nothing known about the populations in Panama, and only a few specimens are known from Nicaragua
Current Population Trend: Decreasing (though this might well be due to a lack of survey effort).
Habitat and Ecology The species is associated with small streams principally in lowland and premontane wet
forests, although also dry forest with perenial streams, and is often found on rocks and under boulders in the
stream. The only known remaining site in Costa Rica is dry forest that has a constant flowing stream (even through
the dry season) and has a water temperature higher than other sights in Costa Rica from which it has disappeared.
It presumably breeds by direct development.
Major Threats It is generally affected by the destruction of natural forests for agriculture and timber, but this does
© Robert Puschendorf

Geographic Range This species can be found from the lowlands and premontane slopes from eastern Nicaragua
on the Atlantic slope and north-western Costa Rica on the Pacific versant to extreme western Panama (including Isla
Escudo de Veraguas), exclusive of the Golfo Dulce region of south-western Costa Rica, at elevations from sea level
up to 1,220m asl (Savage 2002).
Population It was formerly widespread in Costa Rica but has not been seen in much of its range since 1986. Sites
not explain its disappearance from pristine habitats. Some other species of Craugastor that are associated with
streams have undergone dramatic declines and disappearances, possibly due to chytridiomycosis, and so this might
be a threat to this species.
Conservation Measures In view of the threat of chytridiomycosis, the status of this species, and particularly
the surviving population at Río Murciélago, should be closely monitored, and ex-situ populations might need to be
established. The species has been recorded from several protected areas in the past.
Notes on taxonomy: This species was previously included in the genus Eleutherodactylus (Crawford and Smith 2005). It was removed
from the synonymy of Eleutherodactylus rugulosus (Campbell and Savage 2000).
Bibliography: Campbell, J.A. and Savage, J.M. (2000), Crawford, A.J. and Smith, E.N. (2005), Puschendorf, R. et al. (2005), Sasa, M.
and Solorzano, A. (1995), Savage, J.M. (2002)
Data Providers: Frank Solís, Roberto Ibáñez, Alan Pounds, Federico Bolaños, Gerardo Chaves, César Jaramillo, Jay Savage, Gunther
Köhler

VU Craugastor rhodopis (Cope, 1867)

Vulnerable B1ab(iii) Geographic Range This species is largely restricted to western Veracruz and adjacent Hidalgo and Puebla in Mexico
Order, Family: Anura, Leptodactylidae between 1,200 and 2,100m asl. There is an apparently disjunct population in central and south-eastern Chiapas (and
Country Distribution: Mexico adjacent Oaxaca) (Lynch 2000b). It probably occurs more widely than current records suggest.
Current Population Trend: Decreasing Population There is no information on the population status of this species.
Habitat and Ecology This species inhabits tropical montane forest, and it is not known if it can survive in disturbed
habitats. Breeding takes place by direct development.
Major Threats The main threat is forest loss due to logging, human settlement, and some agricultural activities.
Conservation Measures It occurs in Reserva de la Biósfera La Sepultura.
Notes on taxonomy: This species was previously included in the genus Eleutherodactylus (Crawford and Smith 2005). We follow Lynch
(2000b) in separating Eleutherodactylus loki from E. rhodopis.
Bibliography: Crawford, A.J. and Smith, E.N. (2005), Goldberg, S.R. et al. (2002), Lynch, J.D. (2000b), Savage, J.M., McCranie, J.R.
© Twan Leenders

and Espinal, M. (1996)

Data Providers: John Lynch, Georgina Santos Barrera
310
EN Craugastor rhyacobatrachus (Campbell and Savage, 2000)
Endangered B1ab(iii,v)
Order, Family: Anura, Leptodactylidae
Country Distribution: Costa Rica, Panama
Current Population Trend: Decreasing
EN Craugastor sabrinus (Campbell and Savage, 2000)
Endangered A2ace
Order, Family: Anura, Leptodactylidae
Country Distribution: Belize, Guatemala
Current Population Trend: Decreasing
CR Craugastor saltuarius (McCranie and Wilson, 1997)
Critically Endangered A2ace
Order, Family: Anura, Leptodactylidae
Country Distribution: Honduras
Current Population Trend: Decreasing
EN Craugastor sartori (Lynch, 1965)
Endangered B1ab(iii)
Order, Family: Anura, Leptodactylidae
Country Distribution: Mexico
Current Population Trend: Decreasing
Threatened Amphibians of the World
Geographic Range This species can be found in premontane and Conservation Measures The species has been recorded from at least two parks in Panama, but there is clearly a
lower montane southern slopes of the Cordillera Talamanca-Barú need for additional protection of its montane forest habitat. Further research is needed into the potential impacts of
of Costa Rica and western Panama, at elevations of 950-1,800m the chytrid pathogen on this species.
asl (Savage 2002). Notes on taxonomy: This species was previously included in the genus Eleutherodactylus (Crawford and Smith 2005).
Population It is generally quite common where it occurs. However, Bibliography: Campbell, J.A. and Savage, J.M. (2000), Crawford, A.J. and Smith, E.N. (2005), Lips, K.R. (1998), Savage, J.M. (2002)
it is believed to have declined in upland western Panama, though Data Providers: Frank Solís, Roberto Ibáñez, Gerardo Chaves, Jay Savage, César Jaramillo, Querube Fuenmayor
this requires further investigation.
Habitat and Ecology It is associated with rocks and waterfalls
in streams within humid montane forest. It presumably breeds by
direct development.
Major Threats It is generally affected by the destruction of natural
forests for agriculture and timber, but this does not explain its appar-
ent decline in pristine habitats in western Panama. Other species of
the genus that are associated with streams have undergone dramatic
declines and disappearances, possibly due to chytridiomycosis, and
so this might be a major threat to this species.
Geographic Range This species is known from the foothills of disappearances, possibly due to chytridiomycosis, so the status of this species should be monitored carefully.
eastern Guatemala, including the Montañas del Mico, the eastern Conservation Measures It occurs in several protected areas, including the Columbia River Forest Reserve and
portion of the Sierra de las Minas, and the northern portion of the probably in the Bladen Forest Reserve in Belize; and in the Montanas del Mico Catchment Reserve and the Reserva de
Sierra de Merendón. It is also known from the Maya Mountains of la Biosfera Sierra de las Minas in Guatemala. Further research is necessary to establish the reasons for the declines
Belize. It ranges from near sea level to about 900m asl. witnessed in this species in undisturbed habitat.
Population Although still common in Belize, it has suffered serious Notes on taxonomy: This species was previously included in the genus Eleutherodactylus (Crawford and Smith 2005).
population reductions since the 1990s in Guatemala, where it was Bibliography: Campbell, J.A. (1998), Campbell, J.A. (2001), Campbell, J.A. and Savage, J.M. (2000), Crawford, A.J. and Smith, E.N.
formerly common. (2005), Lee, J.C. (1996), Lee, J.C. (2000)
Habitat and Ecology It inhabits lowland and premontane wet Data Providers: Julian Lee, Paul Walker, Manuel Acevedo
and moist forest, living in and alongside streams. It is not found
in degraded forest. It breeds by direct development and the eggs
are laid on land.
Major Threats Habitat loss is taking place due to agriculture, wood
extraction, and human settlement. However, it is declining even in
suitable habitat in Guatemala. Other species of the genus that are
associated with streams have undergone dramatic declines and
Geographic Range This species is found in Cerro Bufalo in the survey work is required to determine the population status of this species and the reasons for its decline in pristine
Dept. of Atlantida and Montaña La Fortuna in the Department of habitat. If disease is shown to be a major threat, then surviving individuals might need to form the basis for the
Yoro, Honduras, at elevations of 1,550-1,800m asl. establishment of an ex-situ population.
Population Although always rare, the species is now in precipitous Notes on taxonomy: This species was previously included in the genus Eleutherodactylus (Crawford and Smith 2005).
decline. Visits to the sites, both in 1995 and 1996, failed to turn up Bibliography: Crawford, A.J. and Smith, E.N. (2005), McCranie, J.R. and Wilson, L.D. (1997a), McCranie, J.R. and Wilson, L.D. (2002b),
any individuals. Wilson, L.D. and McCranie, J.R. (1998)
Habitat and Ecology It occurs in leaf-litter in lower montane wet Data Providers: Gustavo Cruz, Larry David Wilson
forest. Breeding takes place by direct development.
Major Threats Habitat destruction caused by subsistence agricul-
ture and landslides are the major threats. However, these threats do
not account for its decline in pristine habitats. Rather, its disappear-
ance is likely to be as a result of disease (such as chytridiomycosis),
although it is perhaps surprising for this to be the case in such a
species since it is not associated with streams.
Conservation Measures The species occurs in Parque Nacional
Pico Bonito and in the Reserva de Vida Silvestre Texiguat. Further
Geographic Range This species is known only from Cerro Ovando
and surrounding areas, in the Sierra Madre de Chiapas, Mexico, at
1,200-1,900 m asl.
Population The current population status is unknown.
Habitat and Ecology It inhabits montane cloud and mixed forests,
and breeds by direct development.
Major Threats The main threat to this species is habitat loss and
degradation due, in particular, to logging.
Conservation Measures The range of this species includes the
Reserva de la Biósfera El Triunfo. Further survey work is necessary
to evaluate the current population status of this species. This spe-
cies is protected by Mexican law under the “Special Protection”
category (Pr).
Notes on taxonomy: This species was previously included in the genus
Eleutherodactylus (Crawford and Smith 2005).
Bibliography: Crawford, A.J. and Smith, E.N. (2005), Lynch, J.D. (2000b)
Data Providers: Georgina Santos-Barrera, Oscar Flores-Villela
 Globally Threatened Amphibian Species 311

EN Craugastor silvicola (Lynch, 1967)

Endangered B1ab(iii)
Order, Family: Anura, Leptodactylidae
Country Distribution: Mexico
Current Population Trend: Decreasing
Geographic Range This species is known only from Zanatepec, Notes on taxonomy: This species was previously included in the genus Eleutherodactylus (Crawford and Smith 2005).
Isthmus of Tehuantepec, in southern Oaxaca, Mexico, at 1,450- Bibliography: Campbell, J.A., Lamar, W.W. and Hillis, D.M. (1989), Crawford, A.J. and Smith, E.N. (2005)
1,600m asl. Data Providers: Georgina Santos-Barrera, Luis Canseco-Márquez
Population It is a very rare species.
Habitat and Ecology It inhabits pine-oak forest and breeds by
direct development.
Major Threats The major threat is habitat disturbance and degrada-
tion due to logging and the creation of agricultural land. The area
where this species occurs is still relatively undisturbed, due to the
lack of roads and towns; however, logging is expected to increase
in the next few years.
Conservation Measures The range of this species does not include
any protected areas, and there is an urgent need for protection of
montane pine-oak forest habitat in this part of the state of Oaxaca.
This species is protected by Mexican law under the “Special Protec-
tion” category (Pr).

EN Craugastor spatulatus (Smith, 1939)

Endangered B1ab(iii,v)
Order, Family: Anura, Leptodactylidae
Country Distribution: Mexico
Current Population Trend: Decreasing
Geographic Range This species is known from Cuautlapan, central or whether some other factors might also be at play.
Veracruz, and Vista Hermosa in the Sierra de Juárez, north-central Conservation Measures Urgent protection of the Sierra de Juárez forests is needed. There are no federal protected
Oaxaca, Mexico, above 1,800m asl. areas in this region, and those that do exist are “ejidal”, meaning they belong to several local owners with no legal
Population Formerly an abundant species (E. Smith pers. comm.), and continuous protection. This species is protected by Mexican law under the “Special Protection” category (Pr).
at present it is hard to find more than one or two specimens during Notes on taxonomy: This species was previously included in the genus Eleutherodactylus (Crawford and Smith 2005).
the course of an evening’s fieldwork suggesting a recent popula- Bibliography: Campbell, J.A., Lamar, W.W. and Hillis, D.M. (1989), Crawford, A.J. and Smith, E.N. (2005)
tion decline. Data Providers: Georgina Santos-Barrera, Gabriela Parra-Olea
Habitat and Ecology Individuals of this species have been found in
rainforest (cloud forest). It breeds by direct development.
Major Threats The accelerated rate of transformation of the original
forests in northern Oaxaca is certainly the cause of the decline of
amphibian populations in the Sierra de Juárez region. Both regions
where the species is found are under extreme pressure from local
human populations with some parts completely transformed by
agriculture and logging. However, it is not clear whether the recent
decline reported in this species is solely due to habitat degradation,

CR Craugastor stadelmani (K. Schmidt,1936)

Critically Endangered A2ace
Order, Family: Anura, Leptodactylidae
Country Distribution: Honduras
Current Population Trend: Decreasing
Geographic Range This species is known from three localities
in north-central Honduras: Parque Nacional La Muralla, in north-
western Olancho Department; Montanas La Fortuna, in Parque
Nacional Pico Pijol, near Portillo Grande in Yoro Department; and
Cerro San Francisco, in Atlantida Department. It has been recorded
at 1,125-1,900m asl.
Population Although formerly relatively common, this species is now
in precipitous decline. Recent surveys indicate that it has disappeared
from at least two of its three known localities (last seen in Texiguat
in 1993), with no recent information from the third locality (Parque
Nacional Pico Pijol).
Habitat and Ecology It occurs and reproduces by direct devel-
opment on the ground along streams in premontane and lower
montane wet forest.
Major Threats Although habitat destruction caused by subsistence
agriculture is a major threat, the species’ decline in pristine areas is
not entirely understood. However, it should be noted that many other species of Craugastor that are associated with
streams have undergone dramatic declines and disappearances at higher elevations due to chytridiomycosis, and so
this is presumably a major threat to this species.
Conservation Measures Further survey work is required to determine the population status of this species and the
reasons for its decline within pristine habitats. If disease is shown to be a major threat, then surviving individuals
might need to form the basis for the establishment of an ex-situ population. It occurs in Parque Nacional La Muralla
and Parque Nacional Pico Pijol and the Reserva de Vida Silvestre Texiguat.
Notes on taxonomy: This species was previously included in the genus Eleutherodactylus (Crawford and Smith 2005).
Bibliography: Campbell, J.A. (1994b), Crawford, A.J. and Smith, E.N. (2005), Holm, P.A. and Cruz, G.A. (1994), McCranie, J.R. and
Wilson, L.D. (1997a), McCranie, J.R. and Wilson, L.D. (2002b), Schmidt, K.P. (1936b)
Data Providers: Gustavo Cruz, Larry David Wilson

EN Craugastor stuarti (Lynch, 1967)

Endangered B1ab(iii)
Order, Family: Anura, Leptodactylidae
Country Distribution: Guatemala, Mexico
Current Population Trend: Decreasing
Geographic Range This species can be found from San Marcos Bibliography: Campbell, J.A., Lamar, W.W. and Hillis, D.M. (1989), Crawford, A.J. and Smith, E.N. (2005)
and Sololá, Guatemala, and adjacent eastern Chiapas, Mexico, at Data Providers: Georgina Santos-Barrera, Manuel Acevedo
elevations of 1,300-2,200m asl.
Population There are no recent records from Chiapas and the spe-
cies is uncommon in its small range in Guatemala.
Habitat and Ecology It inhabits tropical humid cloud forests. It is a
terrestrial species that can also be found in small bushes. Reproduc-
tion occurs by direct development.
Major Threats Loss and disturbance of the original forest habitat,
primarily due to agriculture and logging, is the major threat.
Conservation Measures This species is not found in any protected
area, and protection of the remaining montane forest habitat is
urgently needed. It is protected by Mexican law under the “Special
Protection” category (Pr).
Notes on taxonomy: This species was previously included in the genus
Eleutherodactylus (Crawford and Smith 2005).
312
CR Craugastor tabasarae (Savage, Hollingsworth, Lips and Jaslow, 2004)
Critically Endangered A4ae
Order, Family: Anura, Leptodactylidae
Country Distribution: Panama
Current Population Trend: Decreasing
© Karen Lips
VU Craugastor tarahumaraensis (Taylor, 1940)
Vulnerable B1ab(iii)
Order, Family: Anura, Leptodactylidae
Country Distribution: Mexico
Current Population Trend: Decreasing
VU Craugastor taurus (Taylor, 1958)
Vulnerable B1ab(iii)
Order, Family: Anura, Leptodactylidae
Country Distribution: Costa Rica, Panama
Current Population Trend: Decreasing
© Twan Leenders
CR Craugastor trachydermus (Campbell, 1994)
Critically Endangered A3ce
Order, Family: Anura, Leptodactylidae
Country Distribution: Guatemala
Current Population Trend: Decreasing
Threatened Amphibians of the World
Geographic Range This species is known from eastern Sonora and
western Chihuahua (Tarahumara mountains), to Jalisco, Mexico, in
the Sierra Madre Occidental, at elevations of around 2,400m asl. It
probably occurs more widely than current records suggest.
Population This is a rare species.
Habitat and Ecology It inhabits pine forest and lives under bark and
leaves on the ground. Breeding is by direct development.
Major Threats The main causes of habitat loss and disturbance in
the Tarahumara forests are logging and agriculture (both for crops
and livestock).
Conservation Measures The range of this species does not include
any protected areas, and urgent protection and restoration of the
original native forests in the Tarahumara Mountains is necessary.
This species is protected by Mexican law under the “Special Protec-
tion” category (Pr).
Geographic Range This species is known only from the type
locality in the Sierra de Santa Cruz, Izabal, Guatemala, at 900m
asl. Recent specimens collected 10km from the type locality might
belong to this species.
Population There is no information on population status, but
congenerics inhabiting similar habitats and elevations have declined
significantly.
Habitat and Ecology It is known to inhabit old-growth premon-
tane wet forest, and is unlikely to be able to adapt to degraded
habitats. Individuals have been seen at night along streams. They
presumably breed by direct development and are not dependent on
water for breeding.
Major Threats Habitat destruction is taking place due to recent
human settlement, as well as agricultural encroachment and wood
extraction. Some other species of Craugastor that are associated
with streams have undergone dramatic declines and disappearances,
Geographic Range This species is known from Coclé Province, west-central Panama (at 600-800 m asl in the
Serranía de Tabasará above El Copé), and Colon and Panama Provinces, east-central Panama (730-910 m asl) near
Cerro Bruja in the Parque Nacional G. D. Omar Torríjos H. and the Serranía Piedras-Pacora in the Parque Nacional
Chagres, respectively.
Population There is little information available on population status, but it is quite rare. A population crash took
place at El Copé in 2004, and it might now be extinct there.
Habitat and Ecology It has been recorded from premontane forest, and is not likely to be tolerant of significant
habitat alteration. Most individuals have been found at night up to 2m above the ground on vegetation overhanging
or near streams, and it is presumed to breed by direct development.
Major Threats Selective logging has occurred in the Serranía de Tabasará. It appears that chytrid fungus has caused
the loss of this species from El Copé, and it is likely to be similarly lost from other sites as the fungus continues to
spread (K. Lips in litt. 2005). The long-term survival of the species in the wild is, therefore, very much in doubt.
Conservation Measures Further survey work is required to determine the population status and, given the threat of
chytridiomycosis, some surviving individuals might need to form the basis for the establishment of an ex-situ captive
breeding programme. This species is known from two national parks: G. D. Omar Torríjos H. and Chagres.
Notes on taxonomy: This species was previously included in the genus Eleutherodactylus (Crawford and Smith 2005).
Bibliography: Crawford, A.J. and Smith, E.N. (2005), Savage, J.M. et al. (2004)
Data Providers: Roberto Ibanez, Karen Lips
Notes on taxonomy: This species was previously included in the genus Eleutherodactylus (Crawford and Smith 2005).
Bibliography: Crawford, A.J. and Smith, E.N. (2005), Taylor, E.H. (1940)
Data Providers: Georgina Santos-Barrera, Luis Canseco-Márquez
Geographic Range This species is known from south-western Costa Rica and extreme south-western Panama, at
elevations of 25-525m asl (Savage 2002).
Population It is a common species.
Habitat and Ecology A nocturnal species associated with rocky areas of streams in humid lowland forest. It breeds
via direct development.
Major Threats Outside protected areas, the main threat is habitat loss due to logging. A number of other species of
the genus Craugastor have suffered serious recent declines, perhaps due to chytridiomycosis; as this species occurs
at lower elevations than some of these affected congeners, it might be at lower risk of infection.
Conservation Measures Within Costa Rica this species has been recorded from Parque Nacional Corcovado and
the Reserva Biólogica el Naranjal; it is not recorded from any protected areas in Panama. Close population monitoring
is required given the potential threat of chytridiomycosis.
Notes on taxonomy: This species was previously included in the genus Eleutherodactylus (Crawford and Smith 2005).
Bibliography: Campbell, J.A. and Savage, J.M. (2000), Crawford, A.J. and Smith, E.N. (2005), Ibáñez, R. et al. (2000), Savage, J.M.
(2002)
Data Providers: Frank Solís, Roberto Ibáñez, Gerardo Chaves, Jay Savage, César Jaramillo, Querube Fuenmayor
possibly due to chytridiomycosis, so the status of this species should be monitored carefully.
Conservation Measures There are no protected areas near where this species has been found, but one has been
proposed for the Sierra de Danta Cruz. Given the likely threat of chytridiomycosis, recommended conservation measures
likely should include the establishment of a captive-breeding programme.
Notes on taxonomy: This species was previously included in the genus Eleutherodactylus (Crawford and Smith 2005).
Bibliography: Campbell, J.A. (1994b), Campbell, J.A. (2001), Crawford, A.J. and Smith, E.N. (2005)
Data Providers: Manuel Acevedo, Eric Smith
 Globally Threatened Amphibian Species 313

EN Craugastor uno (Savage, 1984)

Endangered B1ab(iii) Geographic Range This species is known from Puerto del Gallo, north-central Guerrero, and the Sierra del Sur de
Order, Family: Anura, Leptodactylidae Oaxaca, Mexico. The recent record from Oaxaca is probably in error and is currently under taxonomic revision. The
Country Distribution: Mexico type locality in Guerrero is at 2,034m asl.
Current Population Trend: Decreasing Population It is a rare species.
Habitat and Ecology It inhabits pine-oak and montane cloud forest remnants, and breeds by direct development.
Major Threats The main threat to this species is deforestation due to logging and the cultivation of crops.
Conservation Measures The range of this species does not include any protected areas, and protection of the
fragments of cloud forest and pine-oak forests that still remain in the Guerrero area is urgently needed. This species
is protected by Mexican law under the “Special Protection” category (Pr).
Notes on taxonomy: This species was previously included in the genus Eleutherodactylus (Crawford and Smith 2005).
Bibliography: Crawford, A.J. and Smith, E.N. (2005), Savage, J.M. (1984), Smith, H. and Chiszar, D. (2000)
Data Providers: Georgina Santos-Barrera, Luis Canseco-Márquez
© Tim Burkhardt

VU Craugastor xucanebi (Stuart, 1941)

Vulnerable B1ab(iii) Geographic Range This species occurs in the central highlands of Guatemala, including the Sierra de Los Cuchu-
Order, Family: Anura, Leptodactylidae matanes, the Sierra de Xucaneb, and the Sierra de las Minas. It lives at 600-1,300m asl.
Country Distribution: Guatemala Population It is an uncommon species.
Current Population Trend: Decreasing Habitat and Ecology It lives in cloud forest, including slightly degraded forest, and is usually found on bushes and
undergrowth. Breeding takes place by direct development, and it is not dependent on water for breeding.
Major Threats The major threat is habitat loss due to agricultural encroachment, extraction of wood, and human
settlement.
Conservation Measures It occurs in Parque Nacional Sierra de las Minas, Biotopo del Quetzal, and the proposed
Parque Nacional Cuchumatanes.
Notes on taxonomy: This species was previously included in the genus Eleutherodactylus (Crawford and Smith 2005).
Bibliography: Campbell, J.A. (2001), Crawford, A.J. and Smith, E.N. (2005), Stuart, L.C. (1941)
© Sean M. Rovito

Data Providers: Manuel Acevedo, Eric Smith

EN Cryptobatrachus boulengeri Ruthven, 1916

Endangered B1ab(iii) Geographic Range This species is known from the northern slopes of the Sierra Nevada de Santa Marta, in the
Order, Family: Anura, Leptodactylidae department of Magdalena, in northern Colombia, where it has been recorded from 1,230-2,700m asl.
Country Distribution: Colombia Population It is a common species.
Current Population Trend: Decreasing Habitat and Ecology It is found on rocks within fast-flowing streams near waterfalls in montane forest. The female
puts her eggs on her back and the young develop directly.
Major Threats Agriculture, as well as logging and infrastructure development for human settlement, are major
threats to the species’ habitat. In particular, deforestation around streams is a major threat.
Conservation Measures Its range includes Parque Nacional Natural Sierra Nevada de Santa Marta. ProAves, a
Colombian NGO, in partnership with international conservation NGOs, has recently purchased an additional 1,560
acres of land to the north-west of the park, thereby expanding the current area under protection.
Notes on taxonomy: This genus has recently been moved from the family Hylidae (Faivovich et al. 2005).
Bibliography: Cochran, D.M. and Goin, C.J. (1970), de Perez, G.R., Ruiz-Carranza, P.M. and Ramirez-Pinilla, M.P. (1992), Faivovich, J.
© Taran Grant

et al. (2005), Ramirez-Pinilla, M.P. and Jerez, A. (2000), Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and Lynch, J.D. (1996), Ruthven, A.G.
(1922)
Data Providers: Martha Patricia Ramírez Pinilla, Mariela Osorno-Muñoz, Jose Vicente Rueda, Adolfo Amézquita, María Cristina
Ardila-Robayo

VU Cryptobatrachus fuhrmanni (Peracca, 1914)

Vulnerable B1ab(iii) Geographic Range This species is known from the western flank of the central Andes, in south Antioquia and Caldas
Order, Family: Anura, Leptodactylidae departments, the north and south-eastern flank of the central Andes (in Antioquia and southern Tolima departments),
Country Distribution: Colombia and the western slope of the eastern Andes, in Santander and Cundinamarca departments, in Colombia. It has been
Current Population Trend: Decreasing recorded from 900-2,000m asl.
Population It is a common species.
© Conservacion Internacional-Colombia

Habitat and Ecology It is found on rocks in fast-flowing mountain streams in transition and montane forest. The
eggs are carried on the back of the female, and the young develop directly.
Major Threats Major threats to the species’ habitat include agriculture (both crops and livestock) and logging;
agricultural pollution is also a threat.
Conservation Measures Its range includes Reserva Regional Bosque de Florencia in Caldas department.
Notes on taxonomy: This genus has recently been removed from the family Hylidae (Faivovich et al. 2005).
Bibliography: de Perez, G.R., Ruiz-Carranza, P.M. and Ramirez-Pinilla, M.P. (1992), Faivovich, J. et al. (2005), Ramirez-Pinilla, M.P. and
Jerez, A. (2000), Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and Lynch, J.D. (1996)
Data Providers: Martha Patricia Ramírez Pinilla, Mariela Osorno-Muñoz, Jose Vicente Rueda, Adolfo Amézquita, María Cristina
Ardila-Robayo
314 Threatened Amphibians of the World

CR Cryptobatrachus nicefori Cochran and Goin, 1970

Critically Endangered B1ab(iii)+2ab(iii) Geographic Range This species is known only from the type local- although further taxonomic research is also needed to ascertain whether or not it is a valid species or not.
Order, Family: Anura, Leptodactylidae ity: La Salina, Boyacá Department, Colombia, at 1,450m asl. Notes on taxonomy: Many researchers doubt the validity of this species. The single specimen might belong in the genus Hyla. This
Country Distribution: Colombia Population There is no information on the population status of this genus has recently been moved from the family Hylidae (Faivovich et al. 2005).
Current Population Trend: Decreasing species; it is known only from the single type specimen. Surveys in the Bibliography: Cochran, D.M. and Goin, C.J. (1970), Faivovich, J. et al. (2005), Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and Lynch,
region of the type locality have not produced any individuals. J.D. (1996)
Habitat and Ecology There is no information on the habitat and Data Providers: María Cristina Ardila-Robayo, Jose Vicente Rueda
ecology of this species. However, like other members of its genus,
it is probably associated with streams and waterfalls in montane
forest. The eggs are carried on the back of the female, where they
develop directly without a larval stage.
Major Threats The region where this species occurs has been
severely affected by mining for salt production, and all possible
suitable remaining habitat for this taxon has been destroyed at
the type locality.
Conservation Measures Additional survey work is urgently
required to determine whether or not it still survives in the wild,

VU Cycloramphus acangatan Verdade and Rodrigues, 2003

Vulnerable B1ab(iii) Geographic Range This species is known from Cotia, Ibiúna, Juquitiba, Piedade, Pilar do Sul, Ribeirão Grande, and São
Order, Family: Anura, Leptodactylidae Bernardo do Campo Municipalities, Reserva Biológica de Paranapiacaba, Santo André municipality, and Parque Estadual
Country Distribution: Brazil Intervales, in São Paulo State, south-east Brazil. It is likely to occur throughout the Serra de Paranapiacaba.
Current Population Trend: Decreasing Population This species is common in suitable habitat, being the second most abundantly collected species in
Reserva Biológica do Morro Grande (Dixo and Verdade 2006).
Habitat and Ecology This species is found in Atlantic Forest remnants. Individuals are usually found on the forest
floor, far from water bodies. Breeding takes place by larval development, with terrestrial larvae. It seems to prefer
pristine areas, but is also present in forest fragments (Dixo and Verdade 2006).
Major Threats The Atlantic Forest has been subject to substantial deforestation and fragmentation due to historical
logging and ongoing large-scale clearance for cattle pasture, and crops such as sugar cane, coffee, and exotic trees,
© Francisco L. Franco

as well as for smallholder agriculture.

Conservation Measures It is known from Parque Estadual Intervales and Reservas Biológica de Paranapiacaba
and Morro Grande.
Bibliography: Dixo, M. and Verdade, V.K. (2006), Verdade, V.K. and Rodrigues, M.T. (2003)
Data Providers: Vanessa Verdade

EN Eleutherodactylus acerus Lynch and Duellman, 1980

Endangered B1ab(iii) Geographic Range This species is known only from the type locality and its vicinity between Papallacta and
Order, Family: Anura, Leptodactylidae Cuyuja in Napo Province, Ecuador, at 2,660-2,750m asl. It probably occurs elsewhere, though it is unlikely to be a
Country Distribution: Ecuador widespread species.
© Natural History Museum and Biodiversity Research

Current Population Trend: Decreasing Population There is no recent information on the population status of this species.
Habitat and Ecology It occurs in upper montane cloud forest, and presumably breeds by direct development. There
is no information on its adaptability to disturbed habitats.
Major Threats The major threats are likely to be deforestation due to agricultural development, creation of planta-
tions, logging, and human settlement.
Conservation Measures It is not known to occur in any protected areas, and habitat protection is urgently
Center, University of Kansas

needed.
Bibliography: Lynch, J.D. and Duellman, W.E. (1980), Lynch, J.D. and Duellman, W.E. (1997)
Data Providers: John Lynch, Luis A. Coloma, Santiago Ron

EN Eleutherodactylus acmonis Schwartz, 1960

Endangered B1ab(iii) Geographic Range This species occurs in Guantanamo Province, Bibliography: Centro Nacional de Areas Protegidas (CNAP) (2002), Hedges, S.B. (1993), Hedges, S.B. (1999), Hedges, S.B. (2001),
Order, Family: Anura, Leptodactylidae Cuba, with an altitudinal range of 30-1,150m asl. Henderson, R.W. and Powell, R (1999), Henderson, R.W. and Powell, R. (2001), Schwartz, A. (1960a), Schwartz, A. and Henderson,
Country Distribution: Cuba Population It is uncommon even in suitable habitat. R.W. (1991)
Current Population Trend: Decreasing Habitat and Ecology It is found in mesic hardwood forests on the Data Providers: Blair Hedges, Luis Díaz
ground and in rock crevices. Eggs are laid on the ground and it breeds
by direct development.
Major Threats It is threatened by habitat degradation and de-
forestation due to agriculture, subsistence farming, charcoaling,
and tourism (both because of infrastructure development and the
impact from tourists).
Conservation Measures It occurs in several protected areas,
although these are in need of improved and strengthened manage-
ment.

EN Eleutherodactylus actinolaimus Lynch and Rueda-Almonacid, 1998
Endangered B1ab(iii)
Order, Family: Anura, Leptodactylidae
Country Distribution: Colombia
Current Population Trend: Decreasing
VU Eleutherodactylus actites Lynch, 1979
Vulnerable D2
Order, Family: Anura, Leptodactylidae
Country Distribution: Ecuador
Current Population Trend: Stable
© Luis A. Coloma
EN Eleutherodactylus acutirostris Lynch, 1984
Endangered B1ab(iii,v)
Order, Family: Anura, Leptodactylidae
Country Distribution: Colombia
Current Population Trend: Decreasing
© John D. Lynch
EN Eleutherodactylus adelus Diaz, Cadiz and Hedges, 2003
Endangered B1ab(iii)
Order, Family: Anura, Leptodactylidae
Country Distribution: Cuba
Current Population Trend: Decreasing
Globally Threatened Amphibian Species 315
Geographic Range This species is known only from the vicinity Bibliography: Lynch, J.D. and Rueda-Almonacid, J.V. (1998a), Lynch, J.D. and Rueda-Almonacid, J.V. (1998b)
of the type locality, El Estadero, in Caldas Department, Colombia, Data Providers: John Lynch, Fernando Castro
from 2,000m asl.
Population It is not thought to be a rare species, but it is a difficult
and dangerous area in which to conduct survey work.
Habitat and Ecology It is an inhabitant of the understorey of very
wet primary forest, and it breeds by direct development with eggs
laid on the ground. It has not been recorded outside primary forest,
suggesting that it is very sensitive to habitat disturbance.
Major Threats The main threat is habitat loss and destruction due
to logging, agriculture and the planting of illegal crops.
Conservation Measures The range of the species is not within any
protected area., and there is an urgent need for the establishment
of a protected area to safeguard the primary forest remaining at the
type locality. Further survey work is needed to determine the popula-
tion status of this species, and to determine if it occurs outside the
vicinity of the type locality.
Geographic Range This species is known only from the vicinity of Pilalo on the Pacific slope of the Cordillera Oc-
cidental in central Ecuador, at 760-2,486m asl. It is not likely to occur anywhere else.
Population It is extremely common in its tiny area of distribution.
Habitat and Ecology It inhabits a region of cool cloud forest, but it is rarely observed within the forest, and is
more often seen in exceedingly moist microhabitats in cleared areas near forest. It presumably breeds by direct
development.
Major Threats Ongoing habitat destruction and degradation might impact this species, though it seems to adapt
relatively well to degraded areas. However, its restricted range means that it might not be able to withstand extremely
severe habitat alteration that leads to a drying out of its moist microhabitats.
Conservation Measures It is not known from any protected areas, but might occur in the Reserva Ecológica Los
Illinizas. This species is in need of close population monitoring given its very restricted range.
Bibliography: Lynch, J.D. (1979a), Lynch, J.D. and Duellman, W.E. (1997)
Data Providers: John Lynch, Luis A. Coloma, Santiago Ron
Geographic Range This species is known only from a tiny area on the western slopes of the Cordillera Oriental of
the Colombian Andes, near Virolín, in the department of Santander, Colombia, at 1,740-2,400m asl.
Population It is a very uncommon species that has declined dramatically since it was first discovered. In 2001, only
two individuals were recorded in the entire year.
Habitat and Ecology It occurs on vegetation in the undergrowth of cloud forests, and can only be found in old-growth
forest. It reproduces by direct development.
Major Threats The major threat to the species is habitat loss and degradation due to agriculture and livestock
ranching.
Conservation Measures It occurs in the Santuario de Fauna y Flora Guanentá Alto Río Fonce and the Cachalú
private reserve.
Bibliography: Lynch, J.D. (1984b), Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and Lynch, J.D. (1996)
Data Providers: Martha Patricia Ramírez Pinilla, Jose Vicente Rueda
Geographic Range This species is currently known only from Bibliography: Díaz, L.M., Cadiz, A. and Hedges, S.B. (2003)
the type locality: Loma del Espejo, Alturas de Pizarras del Sur, Data Providers: Blair Hedges, Luis Díaz
Sabanas Llanas, Pinar del Río, in western Cuba. It was collected
at 130m asl.
Population It is moderately common but is a very cryptic species.
Habitat and Ecology It occurs in open, secondary pine forest, with
a herbaceous vegetation layer primarily comprising Lycopodiella sp.,
ferns (Pteris sp.), and grass (Eleocharis sp.). The soil is acidic and is
derived from sandstone. It is found on the ground under leaf-litter and
other cover. It has not been recorded outside forest habitat. Eggs are
laid on the ground, and it breeds by direct development.
Major Threats The major threat to this species is habitat loss due
to fires and clear-cut logging of the forest.
Conservation Measures Its range includes a forest reserve, but
there is no management of this area for conservation. There is an
urgent need for effective and expanded protection of the pine forest
habitat of this species.
316 Threatened Amphibians of the World

VU Eleutherodactylus affinis (Werner, 1899)

Vulnerable D2 Geographic Range This species’ geographic range is from the Paramos de Chingaza, Guerrero, and
Order, Family: Anura, Leptodactylidae Sumapaz,Cundinamarca Department, Colombia. Its altitudinal range is 2,600-3,300m asl.
Country Distribution: Colombia Population It was recorded as abundant in appropriate habitat in 2001.
Current Population Trend: Stable Habitat and Ecology It occurs in low shrubs in páramo habitat, and is able to tolerate some habitat disturbance.
It breeds by direct development.
Major Threats Habitat loss caused by fires and cattle ranching is a major threat.
Conservation Measures It occurs in Parque Nacional Natural Chingaza and Parque Nacional Natural Sumapaz.
Bibliography: Acosta-Galvis, A.R. (2000), Cochran, D.M. and Goin, C.J. (1970), Duellman, W.E. (1972), Lynch, J.D. and Duellman,
W.E. (1997), Lynch, J.D., Ruiz-Carranza, P.M. and Ardila-Robayo, M.C. (1997), Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and Lynch, J.D.
(1996)
Data Providers: Fernando Castro, Maria Isabel Herrera, Mariela Osorno-Muñoz, Jose Vicente Rueda
© John D. Lynch

CR Eleutherodactylus albericoi Lynch and Ruíz-Carranza, 1996

Critically Endangered B1ab(iii)+2ab(iii) Geographic Range This species is known from only one stream Conservation Measures The only known locality is not within a protected area. Further survey work is needed to
Order, Family: Anura, Leptodactylidae in Alta de Oso, on the western flank of the Cordillera Occidental, in confirm the persistence of the species at the type locality, and any surviving individuals might need to form the basis
Country Distribution: Colombia Chocó department, Colombia, at 950m asl. for the establishment of an ex-situ population.
Current Population Trend: Decreasing Population It has only been collected once and was observed to be Bibliography: Acosta-Galvis, A.R. (2000), Lynch, J.D. (1998b), Lynch, J.D. and Ruiz-Carranza, P.M. (1996b), Lynch, J.D., Ruiz-Carranza,
uncommon. There have been numerous surveys of the type locality P.M. and Ardila-Robayo, M.C. (1997), Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and Lynch, J.D. (1996), Ruiz-Carranza, P.M., Lynch, J.D.
and surrounding areas since its original discovery, but the species and Ardila-Robayo, M.C. (1997)
has not been rediscovered. Data Providers: Fernando Castro, Maria Isabel Herrera, John Lynch
Habitat and Ecology This species occurs next to waterfalls and in
channelled crevices, or on vegetation on the sides of the waterfalls.
It breeds by direct development.
Major Threats Habitat loss and degradation caused by subsistence
wood collecting, agriculture (illegal crops), as well as pollution from
the spraying of illegal crops, are all potential threats. Other species of
Eleutherodactylus that are associated with streams have undergone
dramatic declines and disappearances, possibly due to chytridiomyco-
sis, so the status of this species should be monitored carefully.

CR Eleutherodactylus albipes Barbour and Shreve, 1937

Critically Endangered B1ab(iii) Geographic Range This species occurs on Turquino Peak, Sierra Maestra, in Cuba, at 1,700-1,974m asl.
Order, Family: Anura, Leptodactylidae Population It is an uncommon species.
Country Distribution: Cuba Habitat and Ecology It occurs in high-elevation closed-canopy moist forest, and has not been recorded from disturbed
Current Population Trend: Decreasing forest. Eggs are laid on the ground, and it breeds by direct development.
Major Threats Severe habitat destruction is taking place as a result of clear-cutting, charcoaling, agricultural expan-
sion, disturbance from touristic activities, and infrastructure development for human settlement.
Conservation Measures Although its range is wholly within Parque Nacional Turquino and Parque Nacional la
Bayamesa, these protected areas are in need of improved management.
© S. Blair Hedges, Penn State

Bibliography: Centro Nacional de Areas Protegidas (CNAP) (2002), Hedges, S.B. (1993), Hedges, S.B. (1999), Hedges, S.B. (2001),
Henderson, R.W. and Powell, R (1999), Henderson, R.W. and Powell, R. (2001), Schwartz, A. and Henderson, R.W. (1991)
Data Providers: Blair Hedges, Luis Díaz

EN Eleutherodactylus alcoae Schwartz, 1971

Endangered B1ab(iii) Geographic Range This species occurs on the Peninsula de Barahona, in Hispaniola, from sea level up to 600m
Order, Family: Anura, Leptodactylidae asl.
Country Distribution: Dominican Republic, Haiti Population It is a very common species in suitable habitat.
Current Population Trend: Decreasing Habitat and Ecology It is found in dry scrub forest, and retreats by day into caves and rock crevices. Eggs are laid
on the ground, and it breeds by direct development.
Major Threats The main threat is habitat loss from charcoaling and infrastructure development for human settlement.
There is also a cement mine within the range of the species.
Conservation Measures It occurs in Parque Nacional Jaragua, but this park is not well managed and significant
habitat destruction is ongoing within the park’s limits; this protected area is in need of improved and strengthened
© Eladio Fernandez 2006

management for biodiversity conservation. The species is also present in the Reserva Cientifica Via Panomámica
Aceitillar.
Bibliography: Hedges, S.B. (1993), Hedges, S.B. (1999), Hedges, S.B. (2001), Henderson, R.W. and Powell, R (1999), Henderson, R.W.
and Powell, R. (2001), Schwartz, A. (1971), Schwartz, A. (1977a), Schwartz, A. and Henderson, R.W. (1991)
Data Providers: Blair Hedges, Sixto Inchaustegui, Richard Thomas, Robert Powell
 Globally Threatened Amphibian Species 317

VU Eleutherodactylus altae Dunn, 1942

Vulnerable B1ab(iii) Geographic Range This species occurs on the Atlantic slope of northern and central Costa Rica, south to extreme
Order, Family: Anura, Leptodactylidae north-western Panama. Its altitudinal range is 60-1,245m asl (Savage 2002).
Country Distribution: Costa Rica, Panama Population This species is rarely seen and continues to appear occasionally on censuses at the San Ramon Reserve
Current Population Trend: Decreasing and at Monteverde, Costa Rica. It has always been a rare species for which there is little information on population
trends.
Habitat and Ecology This species is found in lowland and premontane wet forest and rainforest areas. It has been
found active on low vegetation at night, and in bromeliads and leaf-litter during the day (Savage 2002). Breeding
takes place by direct development, with the eggs possibly deposited on vegetation. It does not adapt well to modi-
fied habitats.
Major Threats The major threat to this species is habitat loss due to agriculture and logging.
Conservation Measures The species has been recorded from many protected areas including the San Ramon,
© Andrew Crawford

Monteverde, and Braulio Carrillo protected areas of Costa Rica, and Parque Internacional La Amistad (Costa Rica
and Panama).
Bibliography: Savage, J.M. (2002)
Data Providers: Alan Pounds, Federico Bolaños, Frank Solís, Roberto Ibáñez, Gerardo Chaves, Jay Savage, César Jaramillo, Querube
Fuenmayor

CR Eleutherodactylus alticola Lynn, 1937

Critically Endangered B1ab(iii) Geographic Range This species has a very small range in the Blue Mountain Peak, Jamaica, at 1,650-2,250m asl.
Order, Family: Anura, Leptodactylidae Population It was once a common species within its restricted range. However, it was last collected in the 1980s,
Country Distribution: Jamaica and there have been no surveys of the area since then.
Current Population Trend: Decreasing Habitat and Ecology It is a terrestrial species, found hiding under rocks in closed-canopy forest, and is not tolerant
of habitat disturbance. Eggs are laid on the ground and it breeds by direct development.
Major Threats Although this species occurs in the Blue and John Crow Mountain National Park, the primary threat
remains habitat destruction (even within the National Park where there are coffee plantations). Disturbance as a
result of touristic activities is also a threat, since visitors to the park have a marked influence on the habitat and
© S. Blair Hedges, Penn State

leave a great deal of litter behind.

Conservation Measures Recommended conservation measures include improved management of the existing
protected area and habitat maintenace within the national park.
Bibliography: Hedges, S.B. (1993), Hedges, S.B. (1999), Hedges, S.B. (2001), Henderson, R.W. and Powell, R (1999), Henderson, R.W.
and Powell, R. (2001), Schwartz, A. and Henderson, R.W. (1991)
Data Providers: Blair Hedges, Susan Koenig, Byron Wilson

CR Eleutherodactylus amadeus Hedges, Thomas and Franz, 1987

Critically Endangered A3c Geographic Range This species has a small range in the Massif de la Hotte, Haiti, where it has been recorded
Order, Family: Anura, Leptodactylidae from 1,000-2,340m asl.
Country Distribution: Haiti Population It is a very common species within its restricted range.
Current Population Trend: Decreasing Habitat and Ecology It is found in closed-canopy forest under rocks and logs in the daytime, and calls in the
early evening from low herbaceous vegetation. It has been recorded from disturbed forest, but only in very recently
disturbed forest suggesting that it might only be hanging on and not persisting in this habitat. Eggs are laid on the
ground, and it breeds by direct development.
Major Threats Severe habitat destruction is taking place in its range, primarily due to logging by local people
(charcoaling) and slash-and-burn agriculture.
© Eladio Fernandez 2006

Conservation Measures The species occurs in the Parc National Macaya, but there is no management of this area
for conservation, and the habitat continues to be destroyed. Urgent site-based action is required in the Massif de
la Hotte to conserve the remaining habitat in the area, to ensure the persistence of this species, as well as other
threatened amphibians known only from this area.
Bibliography: Hedges, S.B. (1993), Hedges, S.B. (1999), Hedges, S.B. (2001), Hedges, S.B. and Powell, R. (1998a), Hedges, S.B.,
Thomas, R. and Franz, R. (1987), Henderson, R.W. and Powell, R (1999), Henderson, R.W. and Powell, R. (2001), Schwartz, A. and
Henderson, R.W. (1991)
Data Providers: Blair Hedges, Richard Thomas, Robert Powell

EN Eleutherodactylus amplinympha Kaiser, Green and Schmidt, 1994

Endangered B1ab(iii) Geographic Range This species has a small range in the interior of Dominica, where it has been recorded from
Order, Family: Anura, Leptodactylidae 300-1,200m asl.
Country Distribution: Dominica Population It is generally abundant, and fills the ecological niche occupied by Eleutherodactylus martinicensis at
Current Population Trend: Decreasing lower elevations.
Habitat and Ecology Although a habitat generalist, it is an arboreal species with a preference for higher elevation
rainforests, where the forest becomes a little more open. The eggs develop directly without a larval stage.It has
been found in disturbed areas, such as along access roads created to support the hydroelectric project in Parque
Nacional Morne Trois Pitons.
Major Threats On Dominica, the montane forests are largely inaccessible and secure due to the rugged topography
of the area. However, there is some degradation taking place in the south of the island due to small-scale farming
© Alejandro Sanchez

and selective logging.

Conservation Measures This species occurs in several protected areas, but improved protection of habitat at lower
elevations, particularly in the south, is needed.
Bibliography: Hedges, S.B. (1999), Hedges, S.B. (2001), Kaiser, H. and Henderson, R.W. (1994), Kaiser, H., Green, D.M. and Schmid,
M. (1994)
Data Providers: Blair Hedges, Robert Powell
318 Threatened Amphibians of the World

EN Eleutherodactylus andrewsi Lynn, 1937

Endangered B1ab(iii) Geographic Range This species occurs in the Blue and John Crow Schwartz, A. and Henderson, R.W. (1991)
Order, Family: Anura, Leptodactylidae Mountains, Jamaica, at an altitude of 545-1,970m asl. Data Providers: Blair Hedges, Susan Koenig, Byron Wilson
Country Distribution: Jamaica Population It is common in old-growth forests.
Current Population Trend: Decreasing Habitat and Ecology It is found in deep woods on streamside rocks
and among ground litter; the males call from concealed terrestrial
sites. It can only tolerate very temporary habitat disturbance. It breeds
by direct development and the eggs are laid on the ground.
Major Threats The main threat to the species is habitat loss and de-
struction, primarily due to smallholder farming activities, the removal
of woody vegetation, and the impact of tourist activities.
Conservation Measures It occurs in the Blue and John Crow
Mountains National Park, although coffee plantations have been
expanding within the park’s limits. Improved management of this
important protected area is needed.
Bibliography: Hedges, S.B. (1993), Hedges, S.B. (1999), Hedges, S.B. (2001),
Henderson, R.W. and Powell, R (1999), Henderson, R.W. and Powell, R. (2001),

VU Eleutherodactylus angustidigitorum (Taylor, 1940)

Vulnerable B1ab(iii) Geographic Range Historically, this species ranged in a wide area in Michoacán, Mexico, but it is now restricted
Order, Family: Anura, Leptodactylidae to north-western and central Michoacán at around 1,500m asl.
Country Distribution: Mexico Population There is no information regarding the population status of this species.
Current Population Trend: Decreasing Habitat and Ecology This species inhabits pine-oak forest. It is clearly a terrestrial species that prefers to live under
stones and on the ground. It breeds by direct development.
Major Threats The major threat is habitat loss due to logging and infrastructure development for human settle-
ments.
Conservation Measures It is not recorded from any protected areas, but might occur in Parque Nacional Pico de
Tancitaro. There is a need for improved habitat protection at sites where the species is known to occur. Further survey
work is also needed to determine the current population status of the species. This species is protected by Mexican
law under the “Special Protection” category (Pr).
Bibliography: Dixon, J.R. (1957)
© Thomas Bille

Data Providers: Georgina Santos-Barrera, Luis Canseco-Márquez

EN Eleutherodactylus angustilineata Lynch, 1998

Endangered B1ab(iii) Geographic Range This species is known from Chicoral in Valle del Cauca Department, northwards to Pueblo Rico
Order, Family: Anura, Leptodactylidae in Risaralda Department, in the Cordillera Occidental, Colombia, from 1,880-2,500m asl.
Country Distribution: Colombia Population It is a locally common species.
Current Population Trend: Decreasing Habitat and Ecology It is found on low herbaceous vegetation or on epiphytic vegetation in forest habitats, and
has not been recorded outside primary forest. It breeds by direct development.
Major Threats The major threat to this species is habitat loss due to expanding agricultural activities, including
the planting of illegal crops.
Conservation Measures The range of the species includes Parque Nacional Natural Tatama. Continued protection
of primary forest habitat is essential to ensure the survival of this species.
Bibliography: Acosta-Galvis, A.R. (2000), Lynch, J.D. (1998b), Lynch, J.D. (1999)
Data Providers: Fernando Castro, Maria Isabel Herrera, John Lynch
© John D. Lynch

CR Eleutherodactylus apostates Schwartz, 1973

Critically Endangered A3c Geographic Range This species is confined to the Massif de la Hotte, Haiti. Its altitudinal range is from 333-1,640m
Order, Family: Anura, Leptodactylidae asl, although it occurs at lower altitudes only on the northern slope.
Country Distribution: Haiti Population This species was formerly common in good habitat, which probably no longer exists. It was last recorded
Current Population Trend: Decreasing in 1991, although no one has looked for the species since.
Habitat and Ecology It lives in closed-canopy forest. Males call near water, and it breeds by direct development
with the eggs laid on the ground.
Major Threats Severe habitat destruction is taking place in its range, primarily due to logging by local people
(charcoaling) and slash-and-burn agriculture.
Conservation Measures The species occurs in the Parc National Macaya, but there is no management of this area
© Eladio Fernandez 2006

for conservation, and the habitat continues to be destroyed. Urgent site-based action is required in the Massif de
la Hotte to conserve the remaining habitat in the area, to ensure the persistence of this species, as well as other
threatened amphibians known only from this area. Survey work in the region is required to determine the population
status of this species.
Bibliography: Hedges, S.B. (1993), Hedges, S.B. (1999), Hedges, S.B. (2001), Henderson, R.W. and Powell, R (1999), Henderson, R.W.
and Powell, R. (2001), Schwartz, A. (1973), Schwartz, A. (1977b), Schwartz, A. and Henderson, R.W. (1991)
Data Providers: Blair Hedges, Richard Thomas, Robert Powell
 Globally Threatened Amphibian Species 319

EN Eleutherodactylus armstrongi Noble and Hassler, 1933

Endangered B1ab(iii) Geographic Range This species occurs on the Massif de la Selle Bibliography: Hedges, S.B. (1993), Hedges, S.B. (1999), Hedges, S.B. (2001), Henderson, R.W. and Powell, R (1999), Henderson, R.W.
Order, Family: Anura, Leptodactylidae in Haiti and the Sierra de Baoruco in the Dominican Republic, and Powell, R. (2001), Schwartz, A. (1978), Schwartz, A. and Henderson, R.W. (1991)
Country Distribution: Dominican Republic, Haiti Hispaniola, at an altitude of 152-1,697m asl. Data Providers: Blair Hedges, Sixto Inchaustegui, Marcelino Hernandez, Robert Powell
Current Population Trend: Decreasing Population It is common within suitable habitat.
Habitat and Ecology It is arboreal in closed pine and hardwood
forests. Eggs are laid in bromeliads and on the ground under rocks.
Habitat quality is declining and the species has been found in
degraded habitats.
Major Threats Habitat destruction due to small-scale agriculture,
cattle grazing, mining and charcoal collecting is taking place at a rapid
rate in both Massif de la Selle and the Sierra de Baoruco.
Conservation Measures In the Dominican Republic, this species
occurs in Parque Nacional Sierra Baoruco Oriental. There is an urgent
need for strengthened and expanded protection of forest habitats in
the Massif de la Selle in Haiti.

VU Eleutherodactylus ashkapara Köhler, 2000

Vulnerable D2 Geographic Range This species is known from the type locality: the province of Chapare, in the department of
Order, Family: Anura, Leptodactylidae Cochabamba, Bolivia, at 2,100m asl, and from a very nearby locality at 1,700m asl.
Country Distribution: Bolivia Population It is a common species at the type locality.
Current Population Trend: Stable Habitat and Ecology This is an arboreal species that inhabits moist montane forest. It appears to be dependent
on the presence of epiphytes. There is no information known about breeding habits, but it probably breeds by direct
development, like other species of the genus.
Major Threats Logging might be a threat in the future, but at present there are no major threats, and the popula-
tion is stable.
Conservation Measures The type locality is within Parque Nacional Carrasco. This species requires close population
monitoring given that it has a very restricted range.
Bibliography: Köhler, J. (2000a), Köhler, J. (2000b)
Data Providers: Claudia Cortez, Steffen Reichle, Ignacio De la Riva, Jörn Köhler
© Jörn Köhler

EN Eleutherodactylus atratus Lynch, 1979

Endangered B1ab(iii) Geographic Range This species is known from the Amazonian slopes of the eastern Andean Cordillera (Cordillera de
Order, Family: Anura, Leptodactylidae Matanga to the Abra de Zamora) in southern Ecuador from 2,195-2,850m asl. It has also been reported from 3,400m
Country Distribution: Ecuador asl at Río Mazan, Azuay. It is recorded from six localities.
Current Population Trend: Decreasing Population There is very little information available on the current population status of this species. Four individu-
als were collected in 1985, but there appear to be no subsequent records, though this is probably due to a lack of
survey work within its range.
Habitat and Ecology It inhabits páramo grassland, sub-páramo shrubland and cloud forests (Lynch and Duellman
© Mario Humberto Yánez-Muñoz

1980). Breeding is by direct development.

Major Threats The habitat of this species is under threat from agricultural activities and dam construction.
Conservation Measures Its range overlaps Parque Nacional Sangay, and also Parque Nacional Cajas.
Bibliography: Lynch, J.D. (1979c), Lynch, J.D. and Duellman, W.E. (1980), Lynch, J.D. and Duellman, W.E. (1997)
Data Providers: Luis A. Coloma, Santiago Ron, Mario Yánez-Muñoz, Diego Almeida

VU Eleutherodactylus audanti Cochran, 1934

Vulnerable B1ab(iii) Geographic Range This species occurs on the Massif de la Hotte, Massif de la Selle, Sierra de Baoruco, Cordillera
Order, Family: Anura, Leptodactylidae Central, and Sierra de Neiba, in Hispaniola. It has been recorded from 800-2,500m asl.
Country Distribution: Dominican Republic, Haiti Population It is moderately common in suitable habitat.
Current Population Trend: Decreasing Habitat and Ecology It is found in upland closed-canopy forest and forest edge, under rocks and debris. Eggs are
laid under rocks, and it breeds by direct development. It has not been recorded outside forest habitat.
Major Threats The cloud forest habitat of this species is being destroyed at a rapid rate, even in protected areas,
as a result of logging by local people (charcoaling) and slash-and-burn agriculture.
Conservation Measures It occurs in the protected areas of Parques Nacionales Sierra de Bahoruco, Parques Sierra
de Bahoruco Oriental and in the Reserva Cientifica Laguna Rincón in the Dominican Republic, and also in protected
areas in Haiti, but there is a need for improved management of these areas for conservation.
Bibliography: Hedges, S.B. (1993), Hedges, S.B. (1999), Hedges, S.B. (2001), Henderson, R.W. and Powell, R (1999), Henderson, R.W.
© Pedro Rodriguez

and Powell, R. (2001), Schwartz, A. (1979b), Schwartz, A. and Henderson, R.W. (1991)
Data Providers: Blair Hedges, Sixto Inchaustegui, Marcelino Hernandez, Robert Powell
320 Threatened Amphibians of the World

EN Eleutherodactylus auriculatoides Noble, 1923

Endangered B1ab(iii) Geographic Range This species has a patchy distribution in the
Order, Family: Anura, Leptodactylidae Cordillera Central, Dominican Republic, from 788-1,879m asl.
Country Distribution: Dominican Republic Population It is relatively uncommon even within suitable
Current Population Trend: Decreasing habitat.
Habitat and Ecology It is an arboreal species, inhabiting montane
closed forest where it is often found in bromeliads. The males guard
eggs laid in bromeliads.
Major Threats The major threat to the species is habitat destruction
due to agriculture and charcoal production.
Conservation Measures It occurs in several protected areas,
incuding the Valle Nuevo Natural Scientific Reserve.
Bibliography: Hedges, S.B. (1993), Hedges, S.B. (1999), Hedges, S.B. (2001),
Henderson, R.W. and Powell, R (1999), Henderson, R.W. and Powell, R. (2001),
Schwartz, A. (1980a), Schwartz, A. and Henderson, R.W. (1991)
Data Providers: Blair Hedges, Sixto Inchaustegui, Marcelino Hernandez,
Robert Powell

EN Eleutherodactylus bacchus Lynch, 1984

Endangered B1ab(iii) Geographic Range This species can be found on the western slopes of the Cordillera Oriental in the Department
Order, Family: Anura, Leptodactylidae of Santander, Colombia, 1,740-2,300m asl.
Country Distribution: Colombia Population This is a rare species that was last recorded in 2002.
Current Population Trend: Decreasing Habitat and Ecology It occurs on vegetation in the undergrowth of cloud forests. It can also be found at forest
edges and shrubs in pastures, so it can tolerate some level of disturbance, but probably cannot tolerate the complete
© Conservacion Internacional-Colombia

opening up of its habitat. It breeds by direct development.

Major Threats Major threats include habitat loss and degradation due to agriculture and cattle ranching.
Conservation Measures It occurs in the Santuario de Fauna y Flora Guanentá Alto Río Fonce. Additional survey
work is needed to determine the current population status of this species.
Bibliography: Lynch, J.D. (1984b), Lynch, J.D. and Duellman, W.E. (1997), Lynch, J.D., Ruiz-Carranza, P.M. and Ardila-Robayo, M.C.
(1997), Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and Lynch, J.D. (1996)
Data Providers: Fernando Castro, Maria Isabel Herrera, Martha Patricia Ramírez Pinilla, Jose Vicente Rueda

CR Eleutherodactylus bakeri Cochran, 1935

Critically Endangered A3c Geographic Range This species is restricted to the Massif de la Hotte, Haiti, at an altitude of 890-2,325m asl.
Order, Family: Anura, Leptodactylidae Population This species was common within its original habitat, which has now largely been destroyed.
Country Distribution: Haiti Habitat and Ecology It is an arboreal species dependent on closed-canopy forest, retreating by day into bromeliads.
Current Population Trend: Decreasing Eggs are laid inside the bromeliads and the species breeds by direct development.
Major Threats The habitat of this species is being destroyed at a high rate throughout the Tiburon Peninsula, as
a result of logging (charcoal collection) by local people and slash-and-burn agriculture. Only small pockets of intact
habitat remain.
Conservation Measures Although it occurs in the Parc National Macaya, there is no active management of this
area for conservation, and the habitat continues to be destroyed. Urgent site-based action is required in the Massif
© Eladio Fernandez 2006

de la Hotte to conserve the remaining habitat in the area, to ensure the persistence of this species, as well as other
threatened amphibians known only from this area.
Bibliography: Hedges, S.B. (1993), Hedges, S.B. (1999), Hedges, S.B. (2001), Henderson, R.W. and Powell, R (1999), Henderson, R.W.
and Powell, R. (2001), Schwartz, A. (1980b), Schwartz, A. and Henderson, R.W. (1991)
Data Providers: Blair Hedges, Richard Thomas, Robert Powell

EN Eleutherodactylus balionotus Lynch, 1979

Endangered B1ab(iii) Geographic Range This species is known only from the type locality, 13.5km east of Loja, at the crest of the Cordil-
Order, Family: Anura, Leptodactylidae lera (Abra de Zamora) between Loja Province and Zamora-Chinchipe Province, Ecuador, at 2,800m asl. It might occur
Country Distribution: Ecuador elsewhere, though it is unlikely to have a very wide distribution.
© Natural History Museum and Biodiversity Research

Current Population Trend: Decreasing Population It is a moderately common species.

Habitat and Ecology It lives in high-altitude sub-páramo bushland habitats. Specimens have been collected during
the day from terrestrial bromeliads. It is presumed to breed by direct development.
Major Threats Habitat destruction and degradation is ongoing within its range, with the expansion of both agricultural
activities and pine plantations.
Conservation Measures It is not known to occur in any protected areas, but it is found close to the northern limit
Center, University of Kansas

of Parque Nacional Podocarpus. There is a definite need for expanded and improved protection of the sub-páramo
habitat of this species.
Bibliography: Lynch, J.D. (1979c), Lynch, J.D. and Duellman, W.E. (1997)
Data Providers: John Lynch, Luis A. Coloma, Santiago Ron
 Globally Threatened Amphibian Species 321

EN Eleutherodactylus barlagnei Lynch, 1965

Endangered B1ab(iii)+2ab(iii) Habitat and Ecology It is aquatic, living in large and small streams, preferring boulders in reasonably fast-moving
Order, Family: Anura, Leptodactylidae waters. It is usually found in rainforest, and also grassy savannahs. It only occurs in disturbed habitats where there
Country Distribution: Guadeloupe are trees along rivers. The eggs are laid on vegetation and in rock crevices, where they develop directly without a
Current Population Trend: Decreasing larval stage.
Major Threats The stream habitat on Guadeloupe is fragile, small in extent, and the habitat continues to decline.
The forest is being cleared for agriculture, especially banana plantations, and also for the construction of human
settlements; the activities of local fishermen also lead to habitat disturbance. Another major threat is pollution from
pesticides used in banana plantations, as well as from domestic sources. Predation from introduced predators, such as
rats, cats and mongooses, may also be having an impact on numbers, while the introduced congener Eleutherodactylus
johnstonei might be a competitor. Chytridiomycosis has also been recorded as present in some amphibian species on
Dominica, and is a serious potential threat to this species.
© Renaud Boistel

Conservation Measures It occurs in Parc National de la Guadeloupe, but the area is not well managed for biodi-
versity conservation. There is a need to expand and strengthen the existing protected area coverage of Guadeloupe.
Furthermore, the species requires careful population monitoring, particularly given the combined likely impact of
introduced predators and the potential threat of chytridiomycosis; it may be necessary to establish a captive-breed-
ing programme for this species.
Bibliography: Breuil, M. (2002), Breuil, M. (2004), Hedges, S.B. (1993), Hedges, S.B. (1999), Hedges, S.B. (2001), Henderson, R.W. and
Geographic Range This species occurs on the Basse-Terre portion of Guadeloupe, from sea level up to Powell, R (1999), Henderson, R.W. and Powell, R. (2001), Kaiser, H. and Henderson, R.W. (1994), Kaiser, H., Green, D.M. and Schmid,
1,400m asl. M. (1994), Lynch, J.D. (1965), Schwartz, A. (1967), Schwartz, A. and Henderson, R.W. (1991)
Population It is moderately common in suitable habitats within its small range. Data Providers: Blair Hedges, Beatrice Ibéné, Michel Breuil, Robert Powell

CR Eleutherodactylus bartonsmithi Schwartz, 1960

Critically Endangered B1ab(iii) Geographic Range This species has an extremely restricted range in two locations in eastern Cuba: the mouth and
Order, Family: Anura, Leptodactylidae lower reaches of the Río Yumuri; and the Parque Nacional Alejandro de Humboldt (though the taxonomic status of
Country Distribution: Cuba this population has been questioned). The altitudinal range is from 30-212m asl.
Current Population Trend: Decreasing Population There are only a few records of this species, despite extensive survey efforts to locate it. However, the
species appears to be locally common when heard calling.
Habitat and Ecology It is arboreal, found at low elevations in closed forest. They call from shrubs, and are also
© Nils Navarro, courtesy of Luis Diaz

found in rock crevices. Breeding takes place by direct development. Although they can occur in coffee plantations,
they are not found in open habitats.
Major Threats This species is threatened by habitat degradation and deforestation due to subsistence wood col-
lecting, and smallholder farming activities, as well as moderate impacts from touristic activities.
Conservation Measures Besides the Parque Nacional Alejandro de Humboldt, this species also occurs in the Pro-
tected Landscape Maisi-Yumuri. Habitat maintenance within the range of this species is an essential requirement.
Notes on taxonomy: The taxonomic allocation of the western population to this species has been questioned (S.B. Hedges and
L. Diaz pers. comm.).
Bibliography: Centro Nacional de Areas Protegidas (CNAP) (2002), Hedges, S.B. (1993), Hedges, S.B. (1999), Hedges, S.B. (2001),
Henderson, R.W. and Powell, R (1999), Henderson, R.W. and Powell, R. (2001), Schwartz, A. (1960a), Schwartz, A. and Henderson,
R.W. (1991)
Data Providers: Blair Hedges, Luis Díaz

EN Eleutherodactylus baryecuus Lynch, 1979

Endangered B1ab(iii) Geographic Range This species is known only from eight localities
Order, Family: Anura, Leptodactylidae on the eastern face of the Cordillera Oriental, east of Cuenca, in
Country Distribution: Ecuador Morono-Santiago Province, Ecuador, at 2,195-2,988m asl.
Current Population Trend: Decreasing Population It is a reasonably common species.
Habitat and Ecology It inhabits high-altitude sub-páramo bushland
and cloud forests, but can survive in somewhat degraded forest. It
is presumed to breed by direct development.
Major Threats Habitat destruction and degradation is ongoing
within its range, with the expansion of both agricultural activities
and pine plantations.
Conservation Measures Its geographic range overlaps Parque
Nacional Sangay. There is a need for expanded and improved protec-
tion of the sub-páramo habitat of this species.
Bibliography: Lynch, J.D. (1979c), Lynch, J.D. and Duellman, W.E. (1980), Lynch,
J.D. and Duellman, W.E. (1997)
Data Providers: John Lynch, Luis A. Coloma, Santiago Ron

EN Eleutherodactylus bellona Lynch, 1992

Endangered B1ab(iii)+2ab(iii) Geographic Range This species is known from the type locality (Murri) and from Parque Nacional Natural Las
Order, Family: Anura, Leptodactylidae Orquideas, both in the municipality of Frontino, in Antioquia Department, on the western flank of the Cordillera
Country Distribution: Colombia Occidental, in Colombia, from 1,100-2,000m asl.
Current Population Trend: Decreasing Population It is more common at Murri than at Parque Nacional Natural Las Orquideas.
Habitat and Ecology It is usually found on low vegetation and in leaf-litter, inside cloud forests and very humid
forest; it has not been recorded outside forest habitat. It breeds by direct development.
Major Threats Outside the national park, there is increasing pressure on the remaining habitat from agricultural
development, including the planting of illegal crops.
Conservation Measures Although this species occurs in Parque Nacional Natural Las Orquideas, the type locality
remains unprotected. Further research is needed to determine whether this species occurs outside the two known
localities.
© John D. Lynch

Bibliography: Acosta-Galvis, A.R. (2000), Lynch, J.D. (1992a), Lynch, J.D. (1998b), Lynch, J.D. and Duellman, W.E. (1997), Lynch, J.D.,
Ruiz-Carranza, P.M. and Ardila-Robayo, M.C. (1997), Paez, V.P. et al. (2002), Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and Lynch, J.D.
(1996), Ruiz-Carranza, P.M., Lynch, J.D. and Ardila-Robayo, M.C. (1997)
Data Providers: Fernando Castro, Maria Isabel Herrera, John Lynch
322 Threatened Amphibians of the World

CR Eleutherodactylus bernali Lynch, 1986

Critically Endangered B2ab(iii) Geographic Range This species is only known from the type locality: 4km south of the junction of the Sonsón-Dorada
Order, Family: Anura, Leptodactylidae and Argelia roads, Municipality of Sonsón, Departamento of Antioquia, in the northern part of the Central Cordillera
Country Distribution: Colombia of the Colombian Andes, at 2,350m asl.
Current Population Trend: Decreasing Population It is uncommon; two specimens were collected in 1978 and a further four in 1981. Since then, repeated
searches of the type locality have not found any individuals, and surveys in the surrounding areas have not recorded
this species.
Habitat and Ecology It has been recorded from primary cloud forest on herbaceous vegetation and fallen leaves,
and breeds by direct development.
Major Threats Habitat loss caused by logging and agricultural development (illegal crops) is the major threat at
the type locality.
Conservation Measures The type locality does not fall within a protected area, and the habitat has been so badly
disturbed that it might not even be suitable for protection. Further survey work is required to determine whether
© John D. Lynch

or not the species still survives at the type locality; any surviving individuals might need to form the basis for the
establishment of an ex-situ population.
Bibliography: Acosta-Galvis, A.R. (2000), Lynch, J.D. (1986a), Lynch, J.D. and Duellman, W.E. (1997), Lynch, J.D., Ruiz-Carranza, P.M.
and Ardila-Robayo, M.C. (1997), Paez, V.P. et al. (2002), Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and Lynch, J.D. (1996)
Data Providers: Fernando Castro, Maria Isabel Herrera, John Lynch

VU Eleutherodactylus bicolor Rueda-Almonacid and Lynch, 1983

Vulnerable B1ab(iii) Geographic Range This species can be found from the western slope of the Cordillera Oriental in Colombia (depart-
Order, Family: Anura, Leptodactylidae ments of Cundinamarca, Boyaca, and Santander) at elevations of 1,750-2,400m asl.
Country Distribution: Colombia Population This is a common species.
Current Population Trend: Decreasing Habitat and Ecology It occurs in under storey vegetation near watercourses in cloud forest, and also occurs in
secondary forest. Reproduction is by direct development.
Major Threats Habitat loss and degradation due to agriculture, both crops and cattle ranching, is a major threat.
Some other species of Eleutherodactylus that are associated with streams at high elevations have undergone
dramatic declines and disappearances, possibly due to chytridiomycosis, so the status of this species should be
monitored carefully.
Conservation Measures It occurs in Santuario de Fauna y Flora Guanentá Alto Río Fonce and the Estacion Dem-
ostrativa El Rasgon.
© John D. Lynch

Bibliography: Lynch, J.D. and Duellman, W.E. (1997), Lynch, J.D., Ruiz-Carranza, P.M. and Ardila-Robayo, M.C. (1997), Rueda-Almonacid,
J.V. and Lynch, J.D. (1983), Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and Lynch, J.D. (1996)
Data Providers: Fernando Castro, Maria Isabel Herrera, Mariela Osorno-Muñoz, Jose Vicente Rueda

VU Eleutherodactylus bicumulus (Peters, 1863)

Vulnerable B1ab(iii) Geographic Range This species is known from the Venezuelan Coastal Range in the states of Aragua, Distrito
Order, Family: Anura, Leptodactylidae Federal and Sucre. It has been recorded from 577-2,060m asl.
Country Distribution: Venezuela Population It is a common species.
Current Population Trend: Decreasing Habitat and Ecology It is an inhabitant of upland montane cloud forests; its ability to adapt to secondary habitats
is not known. It presumably breeds by direct development.
Major Threats Habitat loss, due to agriculture and logging, is a major threat.
© Javier Mesa / Fundación AndígenA

Conservation Measures This species occurs in several protected areas, including the national parks of Henri Pittier,
San Esteban, Macarao, and El Avila.
Bibliography: Barrio Amorós, C.L. (2004), Frost, D.R. (1985), La Marca, E. (1992), La Marca, E. (1995b), La Marca, E. (1997), Lynch, J.D.
and La Marca, E. (1993), Manzanilla, J. (2001), Manzanilla, J. et al. (1995)
Data Providers: Jesús Manzanilla, Enrique La Marca

EN Eleutherodactylus bisignatus (Werner, 1899)

Endangered B1ab(iii) Geographic Range This species is known only from the type locality, Notes on taxonomy: This species was formerly considered a synonym of Eleutherodactylus fenstratus by Lynch (1980), but it was
Order, Family: Anura, Leptodactylidae Chaco, Bolivia, and from the nearby surroundings. The type locality elevated to species status by Kohler (2000a).
Country Distribution: Bolivia might actually be “Chaco in the Unduavi valley, Yungass de La Paz, Bibliography: Cortez, C. (2000), Köhler, J. (2000a), Reichle, S. (1999)
Current Population Trend: Decreasing 16° 21’S; 67° 49’W, at approximately 1,850m.” Its altitudinal range Data Providers: Claudia Cortez, Steffen Reichle, Ignacio De la Riva, Jörn Köhler
is around 1,850-2,700m asl.
Population It is an abundant species at the type locality. However,
although it is currently common, it is feared that the species will
undergo a steep population decline in the near future, especially as
the quality of its habitat declines.
Habitat and Ecology This species is a diurnal inhabitant of montane
cloud forest. There is no information on breeding, but it probably
breeds by direct development like other species of the genus.
Major Threats Agriculture and logging are the major threats to
its habitat.
Conservation Measures It occurs in Parque Nacional Cotapata,
and the ecological needs of this species should be incorporated as
part of the management plan of this protected area.

CR Eleutherodactylus blairhedgesi Estrada, Diaz and Rodriguez, 1997
Critically Endangered B1ab(iii)+2ab(iii)
Order, Family: Anura, Leptodactylidae
Country Distribution: Cuba
Current Population Trend: Decreasing
CR Eleutherodactylus boconoensis Rivero and Mayorga, 1973
Critically Endangered B1ab(iii)
Order, Family: Anura, Leptodactylidae
Country Distribution: Venezuela
Current Population Trend: Decreasing
© Juan Elías García-Pérez
CR Eleutherodactylus bresslerae Schwartz, 1960
Critically Endangered B1ab(iii)
Order, Family: Anura, Leptodactylidae
Country Distribution: Cuba
Current Population Trend: Decreasing
© Luis M. Diaz
CR Eleutherodactylus brevirostris Shreve, 1936
Critically Endangered A3c
Order, Family: Anura, Leptodactylidae
Country Distribution: Haiti
Current Population Trend: Decreasing
© Eladio Fernandez 2006
Globally Threatened Amphibian Species
Geographic Range This species is only known from two small
nearby sites, near Santa Cruz del Norte, La Havana Province, in Cuba.
It has been recorded from around sea level.
Population It is a very common species within its restricted
range.
Habitat and Ecology It is found on limestone rocks and cliffs in
open areas just beside the coast. The eggs are laid on the ground,
and it breeds by direct development.
Major Threats The species is restricted to an unprotected site
close to an area of high human population density. Canadian tourist
companies are planning the development of this part of the coast
in the near future, and oil extraction activities are also taking place
in this area.
Conservation Measures The species is not currently found within
any protected area and, although a proposed local protected area
(Managed Flora Reserve of Canasí) will include its habitat, it is
323
unclear whether or not this area, if designated, will be well managed.
Bibliography: Estrada, A.R., Díaz, L.M. and Rodríguez, A. (1997), Hedges, S.B. (1999), Hedges, S.B. (2001), Henderson, R.W. and Powell,
R (1999), Henderson, R.W. and Powell, R. (2001), Schwartz, A. and Henderson, R.W. (1991)
Data Providers: Blair Hedges, Luis Díaz
Geographic Range This species is known from the Páramo de Guaramacal, at an elevation of 2,700-3,150m asl,
near the town of Boconó, in Trujillo State, Venezuela.
Population It is an uncommon species.
Habitat and Ecology It occurs in páramo grassland, cloud forests, and high-altitude forest. It breeds by direct
development.
Major Threats The primary threat to the species is habitat loss due to infrastructure development (primarily the
construction of power lines) and fire.
Conservation Measures The species’ range includes Parque Nacional Guaramacal, but this area is not sufficiently
managed to counter the threats to this species.
Bibliography: Barrio Amorós, C.L. (2004), Frost, D.R. (1985), García-Pérez, J.E. (1999), La Marca, E. (1992), La Marca, E. (1997), Rivero,
J.A. and Mayorga, H. (1973)
Data Providers: Enrique La Marca, Juan Elías García-Pérez
Geographic Range This species is known only from two isolated locations on the Maisi Peninsular in extreme
eastern Cuba, including from the mouth of the Río Yumuri. Its altitudinal range is from 30-221m asl.
Population There are only a few records of this species, despite extensive efforts to locate it.
Habitat and Ecology It is found on rocks and cliffs in closed hardwood forest, habitat within which it is moderately
common. It breeds by direct development.
Major Threats The primary threats to the species include habitat degradation and loss due to subsistence agriculture
and wood collecting, and moderate impacts from touristic activities.
Conservation Measures This species occurs in the Protected Landscape Maisi-Yumuri, but maintenance of other
existing habitat within the range of this species is required.
Bibliography: Centro Nacional de Areas Protegidas (CNAP) (2002), Hedges, S.B. (1993), Hedges, S.B. (1999), Hedges, S.B. (2001),
Henderson, R.W. and Powell, R (1999), Henderson, R.W. and Powell, R. (2001), Schwartz, A. (1960a), Schwartz, A. and Henderson,
R.W. (1991)
Data Providers: Blair Hedges, Luis Díaz
Geographic Range This species is restricted to a very small area on the Massif de la Hotte, in Haiti, where it has
been recorded at elevations of 575-2,375m asl.
Population It was formerly common in good habitat in the past, but due to habitat loss the species is now consid-
ered to be an extremely rare species. It has not been recorded since 1991, although there have been no subsequent
surveys in the region for the species.
Habitat and Ecology It is found under ground cover in dwarf cloud forest, and also in clearings (although the latter
is not its main habitat, and it is not known whether or not the species can persist outside forest). Eggs are laid on
the ground and it breeds by direct development.
Major Threats Severe habitat destruction is taking place in its range, primarily due to logging by local people
(charcoaling) and slash-and-burn agriculture.
Conservation Measures Although it occurs in the Parc National Macaya, there is no active management of this
area for conservation, and the habitat continues to be destroyed. Urgent site-based action is required in the Massif
de la Hotte to conserve the remaining habitat in the area, in order to ensure the persistence of this species as well
as other threatened amphibians known only from this area. Survey work in the region is required to determine the
population status of this species, and whether or not it still survives in the wild.
Bibliography: Hedges, S.B. (1993), Hedges, S.B. (1999), Hedges, S.B. (2001), Henderson, R.W. and Powell, R (1999), Henderson, R.W.
and Powell, R. (2001), Schwartz, A. (1980c), Schwartz, A. and Henderson, R.W. (1991)
Data Providers: Blair Hedges, Richard Thomas, Robert Powell
324 Threatened Amphibians of the World

VU Eleutherodactylus briceni Boulenger, 1903

Vulnerable D2 Geographic Range This species is known from the mountains near Mérida City, in the Cordillera de Mérida,
Order, Family: Anura, Leptodactylidae Venezuela. It has been recorded from 1,600-3,300m asl.

© César Luis Barrio-Amorós / Fundación AndígenA

Country Distribution: Venezuela Population It is an uncommon species.
Current Population Trend: Stable Habitat and Ecology It is terrestrial and normally found under rocks in páramo and the ecotone of páramo/cloud
forest. It breeds by direct development.
Major Threats There are no known threats to the species’ habitat at present, but its restricted range renders it
susceptible to stochastic threatening processes.
Conservation Measures Populations are found within Parque Nacional Sierra de la Culata and possibly Parque
Nacional Sierra Nevada.
Bibliography: Barrio Amorós, C.L. (2004), Frost, D.R. (1985), La Marca, E. (1992), La Marca, E. (1997)
Data Providers: Enrique La Marca, Juan Elías García-Pérez

VU Eleutherodactylus bromeliaceus Lynch, 1979

Vulnerable B1ab(iii) Geographic Range This species is known from the Amazonian slopes of the Cordillera Oriental (1,707-2,622m
Order, Family: Anura, Leptodactylidae asl), northern (1,500-1,600m asl) and western (1,830m asl) slopes of the Cordillera del Cóndor, province of Zamora-
Country Distribution: Ecuador, Peru Chinchipe, Loja, and the Cordillera de Cutucù (1,700m asl), province of Morona-Santiago in southern Ecuador. Also
Current Population Trend: Decreasing recorded from Abra Pardo Miguel (2,180m asl) in the northern part of the Cordillera Central, in the department of
San Martin, Peru (05° 46’S; 77°42’W).
Population This is an uncommon species. In Peru, only two specimens have ever been recorded.
Habitat and Ecology It is present within humid montane forests. Individuals occur on low vegetation at night and
in bromeliads by day (Duellman and Pramuk 1999). Breeding takes place by direct development although the egg
deposition site is not known. The ability of this species to adapt to modified habitats is unknown.
Major Threats The major threat to this species is habitat loss and degradation due to agriculture and human
settlement.
Conservation Measures It is recorded from the Bosque de Proteccion Alto Mayo, Peru, and the Parque Nacional
© Edgar Lehr

Podocarpus in Ecuador.
Bibliography: Almendariz, A. (1997), Duellman, W.E. and Pramuk, J.B. (1999), Lynch, J.D. (1979c), Lynch, J.D. and Duellman, W.E.
(1980), Lynch, J.D. and Duellman, W.E. (1997)
Data Providers: Lily Rodríguez, Jorge Luis Martinez, Luis A. Coloma, Santiago Ron, Diego Almeida

EN Eleutherodactylus cabrerai Cochran and Goin, 1970

Endangered B1ab(iii) Geographic Range This species is known from the departments of Antioquia and Caldas on the Cordillera Occidental
Order, Family: Anura, Leptodactylidae of the Colombian Andes from 1,140 -1,940m asl. Cochran (1970) recorded one individual from Valle de Cauca Cisneros
Country Distribution: Colombia below 500m asl, but Lynch (1997, 1998b) and Ruiz-Carranza, Ardila-Robayo, and Lynch (1996) did not consider this to
Current Population Trend: Decreasing be a valid record. Probably by mistake, Acosta-Galvis (2000) included Valle del Cauca in its distribution.
Population It is not a common species.
Habitat and Ecology It is found among fallen leaves, and branches and leaves of low herbaceous vegetation in
secondary montane forest and mature forests. It breeds by direct development. It has not been recorded outside
forest habitat.
Major Threats Habitat loss due to agricultural development (including the planting of illegal crops) is the main
threat to the species outside protected areas.
Conservation Measures It occurs in Parque Nacional Natural Orquideas, but there is a need to strengthen and
expand the existing coverage of protected areas in the Colombian Andes.
© John D. Lynch

Notes on taxonomy: Mention of this species by Kattan (1984) refers to E. capitonis.

Bibliography: Acosta-Galvis, A.R. (2000), Cochran, D.M. and Goin, C.J. (1970), Kattan, G. (1984), Lynch, J.D. (1998b), Lynch, J.D. and
Duellman, W.E. (1997), Lynch, J.D., Ruiz-Carranza, P.M. and Ardila-Robayo, M.C. (1997), Paez, V.P. et al. (2002), Rueda-Almonacid, J.V. (2000),
Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and Lynch, J.D. (1996), Ruiz-Carranza, P.M., Lynch, J.D. and Ardila-Robayo, M.C. (1997)
Data Providers: Fernando Castro, Maria Isabel Herrera, John Lynch

EN Eleutherodactylus cacao Lynch, 1992

Endangered B1ab(iii) Geographic Range This species is known only from a few localities within Parque Nacional Natural Munchique in
Order, Family: Anura, Leptodactylidae the department of Cauca, in the cloud forests on the western slopes of the western Andes in south-western Colombia,
Country Distribution: Colombia from 2,200-2,440m asl. However, it may be more widely distributed than is currently known.
Current Population Trend: Decreasing Population It is not a common species. Recent fieldwork has failed to find the species at any of the known localities
(but the species has always been rare so no conclusions can be drawn yet).
Habitat and Ecology It occurs on the ground among logs and rocks inside cloud forests, and has not thus far been
recorded outside forest habitat. It breeds by direct development.
Major Threats A major threat to the species is habitat loss within the park, due to the planting of illegal crops.
© Francisco José López-López

Conservation Measures Although the known distribution of the species is wholly within Parque Nacional Natural
Munchique, this protected area is in need of improved and strengthened management. Further survey work is neces-
sary to determine the current population status of the species.
Bibliography: Acosta-Galvis, A.R. (2000), Lynch, J.D. (1992b), Lynch, J.D. (1998b), Lynch, J.D. and Duellman, W.E. (1997), Lynch, J.D.
and Ruiz-Carranza, P.M. (1996b), Lynch, J.D., Ruiz-Carranza, P.M. and Ardila-Robayo, M.C. (1997), Ruiz-Carranza, P.M., Ardila-Robayo,
M.C. and Lynch, J.D. (1996)
Data Providers: Fernando Castro, Maria Isabel Herrera, John Lynch

EN Eleutherodactylus calcaratus (Boulenger, 1908)
Endangered B1ab(iii)
Order, Family: Anura, Leptodactylidae
Country Distribution: Colombia
Current Population Trend: Decreasing
© John D. Lynch
VU Eleutherodactylus calcarulatus Lynch, 1976
Vulnerable B1ab(iii)
Order, Family: Anura, Leptodactylidae
Country Distribution: Colombia, Ecuador
Current Population Trend: Decreasing
© John D. Lynch
EN Eleutherodactylus capitonis Lynch, 1998
Endangered B1ab(iii)
Order, Family: Anura, Leptodactylidae
Country Distribution: Colombia
Current Population Trend: Decreasing
© Martha Silva-Velasco
CR Eleutherodactylus caribe Hedges and Thomas, 1992
Critically Endangered A3c; B2ab(iii)
Order, Family: Anura, Leptodactylidae
Country Distribution: Haiti
Current Population Trend: Decreasing
Globally Threatened Amphibian Species
Geographic Range This species is only known from one site (1ha),
south-west of Dame-Marie, Département de la Grand ‘Anse, on the
Tiburon Peninsula, Haiti. It occurs at sea level.
Population There is very little information available on the popula-
tion status of this species.
Habitat and Ecology This amphibian is confined entirely to coastal
mangrove marshes. Its breeding habits and requirements are un-
known, although it presumably breeds by direct development.
Major Threats The fragile mangrove marsh in which it occurs is
in an area of severe habitat destruction, and the mangroves are
probably being cut for charcoal.
Conservation Measures It is not known to occur in any protected
areas. Protection and maintenance of the existing mangrove habitat
of this species is essential, and further survey work is needed to
determine whether or not the species might occur in other marshes
nearby.
325
Geographic Range This species occurs in the cloud forests of the western flank of the Cordillera Occidental, from
the department of Risaralda to the department of Valle del Cauca, in Colombia, from 1,400-2,700m asl.
Population It is a common species.
Habitat and Ecology It is found on the ground in primary and secondary cloud forests, but has not been recorded
outside forest. It breeds by direct development.
Major Threats The main threat to the species is habitat loss caused by agricultural development, including the
planting of illegal crops.
Conservation Measures It occurs in Parque Nacional Natural Farallones de Cali, but there is an urgent need for the
expansion and strengthening of the existing protected areas network in Colombia’s Cordillera Occidental.
Notes on taxonomy: Lynch (1996b) notes misidentifications in earlier accounts.
Bibliography: Acosta-Galvis, A.R. (2000), Cochran, D.M. and Goin, C.J. (1970), Lynch, J.D. (1996b), Lynch, J.D. (1998b), Lynch, J.D.
(1999), Lynch, J.D. and Duellman, W.E. (1997), Lynch, J.D., Ruiz-Carranza, P.M. and Ardila-Robayo, M.C. (1997), Ruiz-Carranza, P.M.,
Ardila-Robayo, M.C. and Lynch, J.D. (1996), Ruiz-Carranza, P.M., Lynch, J.D. and Ardila-Robayo, M.C. (1997)
Data Providers: Fernando Castro, Maria Isabel Herrera, John Lynch
Geographic Range This Andean species ranges along the western flank of the Cordillera Occidental from La Planada
in the department of Nariño in extreme southern Colombia, south to Cotopaxi Province in Ecuador. Its altitudinal range
is 1,140-3,000m asl. It might occur a little more widely.
Population It is a very common species.
Habitat and Ecology It inhabits primary and secondary forest, and does not occur in open areas. It is arboreal
in bushes and trees, and is associated with epiphytes in high vegetation. It is especially abundant along trails
through cloud forest, particularly in the proximity of small streams. It breeds by direct development, and the eggs
are deposited in leaf-litter.
Major Threats The major threats are likely to be deforestation for agricultural development (including plantations,
and the planting of illegal crops), logging, and human settlement, and pollution resulting from the spraying of illegal
crops. Some other species of Eleutherodactylus that are associated with streams at high elevations have undergone
dramatic declines and disappearances, possibly due to chytridiomycosis, so the status of this species should be
monitored carefully.
Conservation Measures In Ecuador, its geographic range overlaps with the Reserva Ecológica Cotacachi-Cayapas
and the Reserva Ecológica Los Illinizas. In Colombia it occurs in the La Planda private reserve.
Bibliography: Acosta-Galvis, A.R. (2000), Lynch, J.D. (1976a), Lynch, J.D. (1998b), Lynch, J.D. and Duellman, W.E. (1997), Lynch, J.D.,
Ruiz-Carranza, P.M. and Ardila-Robayo, M.C. (1997), Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and Lynch, J.D. (1996)
Data Providers: Fernando Castro, Santiago Ron, Luis A. Coloma, Mario Yánez-Muñoz, Diego Cisneros-Heredia
Geographic Range This species is known from three localities in the departments of Cauca and Valle del Cauca,
on the western flank of the Cordillera Occidental, in Colombia, and most likely occurs in suitable habitat in between.
It has been recorded from 2,440-2,800m asl.
Population It is a reasonably common species, and is very common in Munchique.
Habitat and Ecology It is found on low vegetation or in leaf-litter in primary and secondary forest or forest edge,
but has not been recorded outside forest habitat. It breeds by direct development.
Major Threats Habitat loss caused by deforestation for agricultural development (including the planting of illegal
crops) are the main threats to the species.
Conservation Measures The range of this species includes Parque Nacional Natural Munchique and Parque
Nacional Natural Farallones de Cali.
Notes on taxonomy: This species was mistakenly reported as E. cabrerai by Kattan (1984).
Bibliography: Acosta-Galvis, A.R. (2000), Kattan, G. (1984), Lynch, J.D. (1998b), Lynch, J.D. (1999)
Data Providers: Fernando Castro, Maria Isabel Herrera, John Lynch
Bibliography: Hedges, S.B. (1993), Hedges, S.B. (1999), Hedges, S.B. (2001), Hedges, S.B. and Thomas, R. (1992a), Henderson, R.W.
and Powell, R (1999), Henderson, R.W. and Powell, R. (2001), Schwartz, A. and Henderson, R.W. (1991)
Data Providers: Blair Hedges, Richard Thomas
326 Threatened Amphibians of the World

EN Eleutherodactylus casparii Dunn, 1926

Endangered B1ab(iii) Geographic Range This species is restricted to Cienfuegos species, and are in need of expansion and improved management.
Order, Family: Anura, Leptodactylidae Province, in Cuba, where it has been recorded from sea level up Bibliography: Centro Nacional de Areas Protegidas (CNAP) (2002), Hedges, S.B. (1993), Hedges, S.B. (1999), Hedges, S.B. (2001),
Country Distribution: Cuba to 830m asl. Henderson, R.W. and Powell, R (1999), Henderson, R.W. and Powell, R. (2001), Schwartz, A. and Henderson, R.W. (1991)
Current Population Trend: Decreasing Population It is a very common species. Data Providers: Blair Hedges, Luis Díaz
Habitat and Ecology It inhabits a wide variety of mesic situations,
including natural forest and some anthropogenic habitats such as
plantations, as well as introduced vegetation. This suggests that it
is able to tolerate some alteration of its habitat, although it does
require large trees. Eggs are laid on the ground, and it breeds by
direct development.
Major Threats The main threat to this species is habitat loss and
degradation as a result of deforestation for agriculture, the develop-
ment of tourism infrastructure, and the subsequent disturbance of
the habitat due to tourist activities.
Conservation Measures It is known from a few protected areas,
although these areas do not provide sufficient protection for the

CR Eleutherodactylus cavernicola Lynn, 1954

Critically Endangered B1ab(iii)+2ab(iii) Geographic Range This species is only known from two caves near Jackson’s Bay, on the Portland Ridge Peninsula,
Order, Family: Anura, Leptodactylidae very close to sea level (10-15m asl), in Jamaica.
Country Distribution: Jamaica Population It is a very rare species and is difficult to find. It was last seen in the 1990s, but there have been no
Current Population Trend: Decreasing surveys for the species since.
Habitat and Ecology It has only been recorded inhabiting caves in moderately well-forested areas. Eggs are laid
among rocks and it breeds by direct development.
Major Threats The caves in which it occurs on the Portland Ridge Peninsula are being disturbed by humans, including
by people occupying caves illegally. There is also significant disturbance of the habitat from agriculture, the nearby
© S. Blair Hedges, Penn State

lighthouse, and from touristic activities.

Conservation Measures It is not known to occur in any protected areas, but both caves are currently within an
area designated as a national park for the future (although this is not yet confirmed). One cave is currently within
the Jackson Bay Hunting Club.
Bibliography: Hedges, S.B. (1993), Hedges, S.B. (1999), Hedges, S.B. (2001), Henderson, R.W. and Powell, R (1999), Henderson, R.W.
and Powell, R. (2001), Schwartz, A. and Henderson, R.W. (1991)
Data Providers: Blair Hedges, Susan Koenig, Byron Wilson

VU Eleutherodactylus ceuthospilus Duellman and Wild, 1993

Vulnerable B1ab(iii) Geographic Range This species is known from elevations of 1,735-1,840m asl on the western slopes of the Cordillera
Order, Family: Anura, Leptodactylidae de Huancabamba (the Huancambamba depression is a complex system of relatively low ridges, basins, and valleys),
Country Distribution: Peru and at 1,500m asl on the Pacific slope of the Cordillera Occidental in the province of Piura, northern Peru. The range
Current Population Trend: Decreasing of this species is not completely known, and it could occur more widely.
Population It is a common species where it occurs.
Habitat and Ecology An inhabitant of montane forest. Males have been found calling from leaves and stems of
herbaceous vegetation 14-40cm above the ground along a road cut in cloud forest. Adults of both sexes were found
in bromeliads by day. Breeding occurs by direct development.
Major Threats The major threat is the destruction and loss of habitat for livestock ranching and as a result of
selective logging.
Conservation Measures It does not occur in any protected areas, and there is a need for improved habitat protection
© German Chávez

at sites where the species is known to occur.

Bibliography: Duellman, W.E. and Pramuk, J.B. (1999), Duellman, W.E. and Wild, E.R. (1993), Rodríguez, L.O., Cordova, J.H. and
Icochea, J. (1993)
Data Providers: Lily Rodríguez, Jorge Luis Martinez, Erik Wild

VU Eleutherodactylus charlottevillensis Kaiser, Dwyer, Feichtinger and Schmidt, 1995

Vulnerable D2 Geographic Range This species is restricted to north-eastern Tobago Island, Trinidad and Tobago, from sea level
Order, Family: Anura, Leptodactylidae to approximately 550m asl.
Country Distribution: Trinidad and Tobago Population This species is quite common.
Current Population Trend: Stable Habitat and Ecology It is often found in forest leaf-litter and is more rarely encountered close to forest streams.
The males call from low bushes. It is a species that breeds by direct development, and the eggs are deposited in
leaf-litter.
Major Threats There are no known threats to this species other than its rather restricted range which renders it
susceptible to stochastic threatening processes.
Conservation Measures The forests on Tobago in which the species is found are protected in the Little Tobago
Wildlife Sanctuary. This species requires close population monitoring given its very restricted range.
Bibliography: Goldberg, S.R., Bursey, C.R. and Kaiser, H. (1998), Hardy, Jr, J.D. (1982), Kaiser, H. et al. (1995), Murphy, J.C. (1997)
Data Providers: Jerry Hardy
© Adrian Hailey
 Globally Threatened Amphibian Species 327

CR Eleutherodactylus chlorophenax Schwartz, 1976

Critically Endangered A3c Geographic Range This species has a very restricted range on in the area, in order to ensure the persistence of this species as well as other threatened amphibians known only from
Order, Family: Anura, Leptodactylidae the Massif de la Hotte, in Haiti, where it has been recorded from this area. Survey work in the region is required to determine the population status of this species.
Country Distribution: Haiti 990-1,290m asl. Bibliography: Cunningham, C.A., Powell, R. and Hedges, S.B. (1998), Hedges, S.B. (1993), Hedges, S.B. (1999), Hedges, S.B. (2001),
Current Population Trend: Decreasing Population Relatively few specimens of this species are known, and Henderson, R.W. and Powell, R (1999), Henderson, R.W. and Powell, R. (2001), Schwartz, A. (1976a), Schwartz, A. and Henderson,
there is no information on its current population status. R.W. (1991)
Habitat and Ecology It has been found in upland hardwood forests Data Providers: Blair Hedges, Richard Thomas, Robert Powell
and ravine pinelands. It prefers to hide under rocks, in sinkholes or
in vertical rock crevices. Eggs are laid on the ground, and it breeds
by direct development.
Major Threats Severe habitat destruction is taking place in its
range, primarily due to logging by local people (charcoaling) and
slash-and-burn agriculture.
Conservation Measures Although it occurs in the Parc National
Macaya, there is no management of this area for conservation, and
the habitat continues to be destroyed. Urgent site-based action is
required in the Massif de la Hotte to conserve the remaining habitat

EN Eleutherodactylus chrysops Lynch and Ruíz-Carranza, 1996

Endangered B1ab(iii) Geographic Range This species is known from Valle del Cauca and Choco Departments in the cloud forests of the
Order, Family: Anura, Leptodactylidae western flank of the Cordillera Occidental in Colombia, from 900-2,200m asl.
Country Distribution: Colombia Population It is a rare species.
Current Population Trend: Decreasing Habitat and Ecology The species is found mainly in the forest canopy and is mostly active in vegetation 1.5m above the
ground in primary and secondary forest; it has not been recorded outside forest. It breeds by direct development.
Major Threats Forest clearance for agricultural development and cattle ranching has already destroyed some of
the habitat within the range of the species.
Conservation Measures It occurs in Parque Nacional Natural Los Farallones de Cali, but there remains a need for
improved protection of its remaining forest habitat in the Cordillera Occidental. Further survey work is necessary to
establish the current population status of this species.
Bibliography: Acosta-Galvis, A.R. (2000), Lynch, J.D. (1998b), Lynch, J.D. (1999), Lynch, J.D. and Ruiz-Carranza, P.M. (1996b), Ruiz-
© John D. Lynch

Carranza, P.M., Ardila-Robayo, M.C. and Lynch, J.D. (1996)

Data Providers: Fernando Castro, Maria Isabel Herrera, John Lynch

VU Eleutherodactylus colodactylus Lynch, 1979

Vulnerable B1ab(iii) Geographic Range This species is known from two disjunct regions: the Amazonian slopes (2,195-3,140m asl) of
Order, Family: Anura, Leptodactylidae the Cordillera Oriental in the province of Morona-Santiago and Amazonian slopes (2,710-2,800m asl) of the Abra
Country Distribution: Ecuador, Peru Zamora, Zamora-Chinchipe Province in the southern Cordillera Oriental, Ecuador (in southern Ecuador it is known
Current Population Trend: Decreasing from at least 12 localities); and the crest and upper eastern slopes of the Cordillera de Huancabamba in northern
Peru (around 05° 28’S; 79°17’W). It probably occurs in intervening areas between the currently known Ecuadorian
and Peruvian distributions.
Population It is a common species.
Habitat and Ecology This species can be found in the cloud forest and sub-páramo of the high Amazonian slopes
of the Andes (Lynch and Duellman 1980). The Huancambamba depression is a complex system of relatively low
ridges, basins, and valleys. All individuals for which habitat data are available were found in terrestrial and arboreal
bromeliads by day (Duellman and Pramuk 1999). Breeding takes place by direct development, but the site of egg
deposition is not known. The ability of this species to adapt to modified habitats is unknown.
© John D. Lynch

Major Threats In Peru, this species is threatened by habitat loss as a result of livestock farming and selective
logging.
Conservation Measures In Peru, this species might occur in one small-protected area in the north-west, the Taba-
conas Namballe National Sanctuary. In Ecuador, its geographic range overlaps with Parque Nacional Sangay.
Bibliography: Duellman, W.E. and Pramuk, J.B. (1999), Duellman, W.E. and Wild, E.R. (1993), Lynch, J.D. (1979c), Lynch, J.D. and
Duellman, W.E. (1980), Lynch, J.D. and Duellman, W.E. (1997), Rodríguez, L.O., Cordova, J.H. and Icochea, J. (1993)
Data Providers: Lily Rodríguez, Jorge Luis Martinez, Luis A. Coloma, Santiago Ron

EN Eleutherodactylus colomai Lynch and Duellman, 1997

Endangered B1ab(iii) Geographic Range This species is known from three localities in the foothills of the Andes in north-western Ecua-
Order, Family: Anura, Leptodactylidae dor and south-western Colombia: Pote, Río Santiago, in Esmeraldas Province, Ecuador; Alto Tambo, in Esmeraldas
Country Distribution: Colombia, Ecuador Province, Ecuador; and the Altaquer-Tumaco Road, department of Nariño, Colombia. Its known altitudinal range is
Current Population Trend: Decreasing 360-1,200m asl.
Population It is an extremely rare species.
Habitat and Ecology It inhabits mature and secondary moist forest, but it is not found in open areas. Specimens have
been found on leaves of bushes 50-60cm above the ground at night. It is presumed to breed by direct development,
© Mario Humberto Yánez-Muñoz

although the site of egg deposition is not known.

Major Threats The major threats are habitat loss -- most likely due to deforestation for agricultural development, plan-
tations, illegal crops, logging, and human settlement -- and pollution (resulting from the spraying of illegal crops).
Conservation Measures It is not known from any protected areas, but it occurs on the border of the Reserva
Ecológica Cotacachi-Cayapas in Ecuador. It may be worth investigating the possible expansion of this reserve to
include part of the range of this species.
Bibliography: Acosta-Galvis, A.R. (2000), Lynch, J.D. (1998b), Lynch, J.D. and Duellman, W.E. (1997), Morales, M. et al. (2002), Ruiz-
Carranza, P.M., Ardila-Robayo, M.C. and Lynch, J.D. (1996)
Data Providers: Fernando Castro, Santiago Ron, Luis A. Coloma, Manuel Morales
328 Threatened Amphibians of the World

VU Eleutherodactylus colostichos La Marca and Smith, 1982

Vulnerable D2 Geographic Range This species is known only from a single, very Bibliography: Barrio Amorós, C.L. (2004), Frost, D.R. (1985), La Marca, E. (1992), La Marca, E. (1995b), La Marca, E. (1997), La Marca,
Order, Family: Anura, Leptodactylidae restricted locality in the Páramo de Los Conejos, in the Sierra de E. and Smith, H.M. (1982)
Country Distribution: Venezuela la Culata, in the state of Mérida, Venezuela. It has been recorded Data Providers: Enrique La Marca, Juan Elías García-Pérez
Current Population Trend: Stable from 3,000-3,600m asl.
Population It is an uncommon species.
Habitat and Ecology It is a terrestrial species that inhabits páramo
grassland. It breeds by direct development.
Major Threats Agriculture and habitat disturbance by tourists are
major threats.
Conservation Measures The population is protected within the
Parque Nacional Sierra de la Culata. Close population monitoring
of this species is required given that it is known only from a single
locality.

VU Eleutherodactylus condor Lynch and Duellman, 1980

Vulnerable B1ab(iii) Geographic Range This species is known from 1,975m asl in the Major Threats The major threat is habitat loss and degradation through agriculture (especially livestock ranching),
Order, Family: Anura, Leptodactylidae Cordillera de Cutucù and Coangos at 1,500-1,600m asl on the western logging, and mining. The Cordillera del Cóndor region is not currently threatened because of land mines remaining
Country Distribution: Ecuador, Peru slope of Cordillera del Cóndor, Morona-Santiago Province, Ecuador, there from the 1995 border war.
Current Population Trend: Decreasing and from 1,750m asl on the eastern slope of the Cordillera del Cóndor, Conservation Measures The range of this species overlaps with Tumbes Reserve Zone in the department of Tumbes
in the department of Amazonas, Peru. This species is also observed and Santiago Comainas Reserve Zone in the department of Amazonas, both in northern Peru. In Ecuador, it occurs
in the department of Tumbes. It might occur more widely, especially in Parque Nacional Podocarpus.
in areas between known locations. Bibliography: Almendariz, A. (1997), Duellman, W.E. and Pramuk, J.B. (1999), Lynch, J.D. and Duellman, W.E. (1980), Lynch, J.D. and
Population There is limited information on the population status of Duellman, W.E. (1997), Morales, M. (2003), Reynolds, R. and Icochea, J. (1997), Tello, G. (1998), Wust, W.H. (1998)
this species. In Ecuador, this species is rare although it was recorded Data Providers: Lily Rodríguez, Jorge Luis Martinez, Luis A. Coloma, Santiago Ron, Diego Almeida
in 2003 in two localities in the Cordillera del Cóndor. The Peruvian
locality is the first record, and based on an adult female.
Habitat and Ecology All individuals have been collected in humid
montane cloud forest. Adults have been found perched on low vegeta-
tion at night; juveniles and adults were on the forest floor by day. This
species breeds by direct development, but the site of egg deposition
is not known. It has not been found outside old growth forest.

EN Eleutherodactylus cooki Grant, 1932

Endangered B1ab(iii) Geographic Range This species has a restricted range at Cuchilla de Pandura, Puerto Rico. Its altitudinal range
Order, Family: Anura, Leptodactylidae is 91-303m asl.
Country Distribution: Puerto Rico Population The current population status is not clear, but it is not currently believed to be declining.
Current Population Trend: Decreasing Habitat and Ecology It lives in humid forests where it is found among large boulder clusters; the eggs are laid on
damp boulder surfaces in grottoes and develop directly.
Major Threats Major threats include habitat destruction and disturbance from agriculture and urbanization (since
the boulder areas in which it occurs are close to human habitation).
Conservation Measures It is not known to occur in any protected areas, and protection of the habitat of this
species is urgently needed.
Bibliography: Burrowes, P.A. (2000), Burrowes, P.A. and Joglar, R.L. (1999), Hedges, S.B. (1993), Henderson, R.W. and Powell, R
© Alejandro Sanchez

(1999), Henderson, R.W. and Powell, R. (2001), Joglar, R.L. (1999), Joglar, R.L., Burrowes, P.A. and Rios, N. (1996), Rivero, J.A. (1998),
Vega-Castillo, S.I. (2000)
Data Providers: Blair Hedges, Rafael Joglar, Richard Thomas, Luis J. Villanueva-Rivera, Neftalí Ríos-Lopez

VU Eleutherodactylus cornutus (Jiménez dela Espada, 1871)

Vulnerable B1ab(iii) Geographic Range This species can be found at moderate elevations (1,150-1,800m asl) along the eastern face of
Order, Family: Anura, Leptodactylidae the Cordillera Oriental from southern Colombia (Caquetá) to southern Ecuador.
Country Distribution: Colombia, Ecuador Population This is a rare species across its range; it has not been seen since 1990 in Colombia, although there has
© Natural History Museum and Biodiversity Research

Current Population Trend: Unknown been little survey work.

Habitat and Ecology This species occurs in cloud forest. One individual was found on the forest floor by day, while
others were found on the ground at night; some individuals have been found on stream banks (Lynch and Duellman
1980). Reproduction occurs by direct development.
Major Threats Habitat destruction and degradation due to agriculture (including the planting of illegal crops), hu-
man settlement, and logging is the main threat. Some other species of Eleutherodactylus that are associated with
Center, University of Kansas

streams have undergone dramatic declines and disappearances, possibly due to chytridiomycosis, so the status of
this species should be monitored carefully.
Conservation Measures In Ecuador, it occurs in Parque Nacional Podocarpus, and its range overlaps with Reserva
Ecológica Cayamabe-Coca, and Parque Nacional Sumaco Napo-Galeras. In Colombia, it occurs in Parque Nacional
Natural Alto Fragua-Indi Wasa.
Bibliography: Jiménez de la Espada, M. (1871), Lynch, J.D. (1975b), Lynch, J.D. (1997), Lynch, J.D. and Duellman, W.E. (1980), Lynch,
J.D. and Duellman, W.E. (1997)
Data Providers: Luis A. Coloma, Santiago Ron, John Lynch, Diego Almeida
 Globally Threatened Amphibian Species 329

CR Eleutherodactylus corona Hedges and Thomas, 1992

Critically Endangered A3c; B1ab(iii)+2ab(iii) Geographic Range This species is known from only one site on the Massif de la Hotte, in Haiti, at 1,120m asl.
Order, Family: Anura, Leptodactylidae Population It is known from fewer than 10 specimens, and is probably very rare.
Country Distribution: Haiti Habitat and Ecology It is an arboreal species, occurring in high-elevation cloud forest. It has been recorded only
Current Population Trend: Decreasing from forest edge but this is probably not suitable habitat. Males call from bromeliads or orchids, which they appear
to require for reproduction. The species breeds by direct development.
Major Threats Severe habitat destruction is taking place in its range, primarily due to logging by local people
(charcoaling) and slash-and-burn agriculture.
Conservation Measures The species occurs only in the Parc National Macaya, but there is no management of this

© S. Blair Hedges, Penn State

area for conservation, and the habitat continues to be destroyed. Urgent site-based action is required in the Massif
de la Hotte to conserve the remaining habitat in the area, in order to ensure the persistence of this species as well
as other threatened amphibians known only from this area. Survey work is required to determine the population
status of this species.
Bibliography: Hedges, S.B. (1993), Hedges, S.B. (1999), Hedges, S.B. (2001), Hedges, S.B. and Thomas, R. (1992b), Henderson, R.W.
and Powell, R (1999), Henderson, R.W. and Powell, R. (2001), Schwartz, A. and Henderson, R.W. (1991)
Data Providers: Blair Hedges, Richard Thomas

EN Eleutherodactylus cosnipatae Duellman, 1978

Endangered B1ab(iii) Geographic Range This species is believed to be entirely restricted to the Cosñipata Valley (1,580-1,700m asl), on
Order, Family: Anura, Leptodactylidae the north-eastern slopes of the Cadena de Paucartambo, a frontal range of the Andean Cordillera Oriental in Cuzco
Country Distribution: Peru Department, Peru.
Current Population Trend: Decreasing Population It is a rare species.
Habitat and Ecology It can be found in tall forest with some tree ferns and bromeliads and luxuriant undergrowth
of mosses and ferns. Except for a single individual that was collected under a rock, all specimens have been recorded
calling from low vegetation in cloud forest at night. It is not known from modified habitats. This species breeds by
direct development.
Major Threats The major threat is small-scale agriculture (involving both crops and livestock) taking place within
the valley.
© Alessandro Catenazzi

Conservation Measures Part of the range is well protected within Parque Nacional Manu. Further survey work
is necessary to determine the species’ current population status and whether it is indeed confined to the Cosñipata
Valley.
Bibliography: Duellman, W.E. (1978), Rodríguez, L.O., Cordova, J.H. and Icochea, J. (1993)
Data Providers: Lily Rodríguez, Jorge Luis Martinez, Wilfredo Arizabal

EN Eleutherodactylus counouspeus Schwartz, 1964

Endangered B1ab(iii) Geographic Range This species is restricted to the Massif de la Bibliography: Hedges, S.B. (1993), Hedges, S.B. (1999), Hedges, S.B. (2001), Henderson, R.W. and Powell, R (1999), Henderson, R.W.
Order, Family: Anura, Leptodactylidae Hotte, Haiti, from 303-760m asl. and Powell, R. (2001), Schwartz, A. (1964a), Schwartz, A. (1981), Schwartz, A. and Henderson, R.W. (1991)
Country Distribution: Haiti Population It is moderately common. Data Providers: Blair Hedges, Richard Thomas, Robert Powell
Current Population Trend: Decreasing Habitat and Ecology It inhabits limestone caves and crevices in
closed, humid forest; males call from well within the caves, where
the eggs are laid. It breeds by direct development.
Major Threats Severe habitat destruction is taking place in the
Formon-Macaya region, including throughout Parc National Ma-
caya, due to logging (charcoal collection) by locals and slash and
burn agriculture.
Conservation Measures It occurs in Parc National Macaya, but
there is no management of this area for conservation, and the habitat
continues to be destroyed. Urgent site-based action is required in
the Massif de la Hotte to conserve the remaining existing habitat in
the area, to ensure the persistence of this species, as well as other
threatened amphibians known only from this area.

EN Eleutherodactylus cremnobates Lynch and Duellman, 1980

Endangered B1ab(iii) Geographic Range This species is only known from five localities on the Amazonian slopes of the Andes in Ecuador,
Order, Family: Anura, Leptodactylidae at 1,410-1,700m asl, in the Río Quijos Valley. It may have a slightly wider distribution.
Country Distribution: Ecuador Population It is reasonably common in its small range.
© Natural History Museum and Biodiversity Research

Current Population Trend: Decreasing Habitat and Ecology It is a nocturnal cloud forest species, found among moss, ferns or herbaceous plants on cliffs
over streams. It is not known whether or not it can adapt to disturbed habitats. It is presumed to breed by direct
development, but the site of egg deposition is not known.
Major Threats Habitat destruction and degradation are taking place due to agricultural development, plantations,
logging, and human settlement. About 60% of the natural vegetation had been removed from within its known range
by 1996. Some other species of Eleutherodactylus that are associated with high-elevation streams have undergone
Center, University of Kansas

dramatic declines and disappearances, possibly due to chytridiomycosis, so the status of this species should be
monitored carefully.
Conservation Measures Its range possibly overlaps with the Reserva Ecológica Cayambe-Coca. There is a need
for improved protection of the cloud forest habitat of this species.
Bibliography: Lynch, J.D. and Duellman, W.E. (1980), Lynch, J.D. and Duellman, W.E. (1997)
Data Providers: John Lynch, Luis A. Coloma, Santiago Ron
330 Threatened Amphibians of the World

EN Eleutherodactylus crenunguis Lynch, 1976

Endangered B1ab(iii) Geographic Range This species is known from more than ten locations in the vicinity of Pilalo on the Pacific slope
Order, Family: Anura, Leptodactylidae of the Cordillera Occidental in central Ecuador, where it has been recorded from 760-2,486m asl.
Country Distribution: Ecuador Population It is not an uncommon species.

© Natural History Museum and Biodiversity Research

Current Population Trend: Decreasing Habitat and Ecology Individuals have been found on low vegetation along, or on banks of, streams at night (Lynch
and Duellman 1997) in montane forest. It breeds by direct development, and the eggs are laid in leaf-litter.
Major Threats Agriculture, both crops and livestock, as well as logging, are major threats to the species’ habitat.
Some other species of Eleutherodactylus that are associated with montane streams have undergone dramatic declines
and disappearances, possibly due to chytridiomycosis, so the status of this species should be monitored carefully.
Conservation Measures Its range is not included in any protected areas, and there is an urgent need for the
Center, University of Kansas protection of the montane forest habitat of this species.
Bibliography: Lynch, J.D. (1976a), Lynch, J.D. and Duellman, W.E. (1997)
Data Providers: Luis A. Coloma, Santiago Ron, Diego Cisneros-Heredia, Ana Almandáriz

VU Eleutherodactylus crucifer (Boulenger, 1899)

Vulnerable B1ab(iii) Geographic Range This species occurs at 1,200-1,800m asl on the Pacific slopes of the Cordillera Occidental in Ecuador,
Order, Family: Anura, Leptodactylidae from the provinces of Imbabura and Esmeraldas, south to Bolivar. It has been recorded from around ten localities.
Country Distribution: Ecuador Population It appears to be an uncommon species, but it is hard to find, and so it might not be as rare as it seems.
Current Population Trend: Decreasing Habitat and Ecology It is a cloud forest species that lives in bromeliads by day. At night, individuals have been
found on ferns and leaves up to 2m above the ground, in dense forest, and along streams. It is not known whether or
not it can adapt to disturbed habitats. It is presumed to breed by direct development, but the site of egg deposition
is not known.
Major Threats Habitat destruction and degradation are taking place due to agricultural development, plantations,
logging, and human settlement. Some other species of Eleutherodactylus that are associated with streams have
undergone dramatic declines and disappearances, possibly due to chytridiomycosis, so the status of this species
should be monitored carefully.
Conservation Measures Its geographic range overlaps the Reservas Ecológica Cotacachi-Cayapas and the Reserva
© John D. Lynch

Ecológica Los Illinizas.

Bibliography: Boulenger, G.A. (1899), Lynch, J.D. and Duellman, W.E. (1997)
Data Providers: John Lynch, Luis A. Coloma, Santiago Ron

EN Eleutherodactylus cryophilius Lynch, 1979

Endangered B1ab(iii) Geographic Range This species is known from five localities east and west of Cuenca, Morona-Santiago Province, Ec-
Order, Family: Anura, Leptodactylidae uador, from 2,835-3,384m asl. It is also known from the Protected Forest of Mazan, within Parque Nacional Cajas.
Country Distribution: Ecuador Population There is no information on the current population status of this species.
Current Population Trend: Decreasing Habitat and Ecology Individuals have been found beneath rocks and logs in sub-páramo shrubland and páramo
grassland habitats. It breeds by direct development.
Major Threats Habitat destruction and degradation, primarily due to agriculture, is the major threat.
Conservation Measures Its range overlaps Parque Nacional Cajas and Parque Nacional Sangay. Further survey
work is needed to determine the current population status of this species.
Bibliography: Lynch, J.D. (1976a), Lynch, J.D. (1979c), Lynch, J.D. and Duellman, W.E. (1997)
Data Providers: Luis A. Coloma, Santiago Ron, Mario Yánez-Muñoz
© Eduardo Toral

EN Eleutherodactylus cryptomelas Lynch, 1979

Endangered B1ab(iii) Geographic Range This Andean species is known from elevations of 2,470-2,710m asl on the Amazonian slopes of
Order, Family: Anura, Leptodactylidae the Cordillera Oriental and at 3,000-3,100m asl on the the ridges north of the Cuenca de Loja, in southern Ecuador,
Country Distribution: Ecuador, Peru and at elevations of 2,770-2,820m asl on the western slope of the Cordillera de Huancabamba in Piura Province in
Current Population Trend: Decreasing northern Peru.
Population It is a rare species.
Habitat and Ecology It inhabits páramo, sub-páramo and humid cloud forests (Lynch 1979c; Duellman and Pramuk
1999). Specimens have been found in terrestrial bromeliads or under rocks; one individual from Peru was found in a
tree at night (Duellman and Pramuk 1999). The ability of this species to adapt to modified habitats is unknown. It is
presumed to breed by direct development, although the site of egg deposition is not known.
Major Threats General habitat loss through deforestation for livestock ranching and selective wood extraction is
the main threat to this species, including even in protected areas.
© German Chávez

Conservation Measures In Peru, this species might occur in one small-protected area (Tabaconas Namballe National
Sanctuary). In Ecuador, its geographic range overlaps with Parque Nacional Podocarpus. Improved and expanded
protection of the habitat of this species is needed.
Bibliography: Duellman, W.E. and Pramuk, J.B. (1999), Duellman, W.E. and Wild, E.R. (1993), Lynch, J.D. (1979c), Lynch, J.D. and
Duellman, W.E. (1980), Lynch, J.D. and Duellman, W.E. (1997), Rodríguez, L.O., Cordova, J.H. and Icochea, J. (1993)
Data Providers: Lily Rodríguez, Jorge Luis Martinez, Luis A. Coloma, Santiago Ron

CR Eleutherodactylus cubanus Barbour and Shreve, 1937
Critically Endangered B2ab(iii)
Order, Family: Anura, Leptodactylidae
Country Distribution: Cuba
Current Population Trend: Decreasing
VU Eleutherodactylus cundalli Dunn, 1926
Vulnerable B1ab(iii)
Order, Family: Anura, Leptodactylidae
Country Distribution: Jamaica
Current Population Trend: Decreasing
CR Eleutherodactylus darlingtoni Cochran, 1935
Critically Endangered A3c; B1ab(iii)+2ab(iii)
Order, Family: Anura, Leptodactylidae
Country Distribution: Haiti
Current Population Trend: Decreasing
EN Eleutherodactylus degener Lynch and Duellman, 1997
Endangered B1ab(iii)
Order, Family: Anura, Leptodactylidae
Country Distribution: Colombia, Ecuador
Current Population Trend: Decreasing
Globally Threatened Amphibian Species 331
Geographic Range This species is restricted to the Sierra Maestra, Bibliography: Centro Nacional de Areas Protegidas (CNAP) (2002), Díaz, L.M. (1998b), Hedges, S.B. (1993), Hedges, S.B. (1999), Hedges,
Cuba, at an altitude of 800-1,400m asl. S.B. (2001), Hedges, S.B., González, L. and Estrada, A.R. (1995), Henderson, R.W. and Powell, R (1999), Henderson, R.W. and Powell, R.
Population It is a common species within suitable habitat. (2001), Schwartz, A. and Henderson, R.W. (1991)
Habitat and Ecology It is restricted to closed humid forest at high Data Providers: Blair Hedges, Luis Díaz
elevations, and breeds by direct development.
Major Threats The primary threat to this species is habitat
destruction due to agriculture, woodcutting, and disturbance from
touristic activities, as well as infrastructure development for human
settlement.
Conservation Measures The range of this species is wholly within
Parque Nacional Turquino and Parque Nacional La Bayamesa, but
there is no management of these specific areas for conservation,
and the habitat continues to be destroyed. Improved management
of these areas, and maintenance of the existing habitat, is urgently
needed.
Geographic Range This species is widely, but patchily, distributed
in western Jamaica from sea level to 635m asl.
Population It is more abundant than many other species of Eleu-
therodactylus on Jamaica and is moderately common.
Habitat and Ecology It is found in association with rocks and
caves and requires closed-canopy moist forest. This species breeds
by direct development, and eggs are laid on the ground. It does not
occur in open habitats.
Major Threats The main threat is habitat degradation and loss due to
agriculture, human settlements, tourism development, and logging.
Conservation Measures This species occurs in several forest
reserves, but many of these are in need of improved management.
Bibliography: Diesel, R., Baurle, G. and Vogel, P (1995), Hedges, S.B. (1993),
Hedges, S.B. (1999), Hedges, S.B. (2001), Henderson, R.W. and Powell, R
(1999), Henderson, R.W. and Powell, R. (2001), Schwartz, A. and Henderson,
R.W. (1991)
Data Providers: Blair Hedges, Susan Koenig, Byron Wilson
Geographic Range This species is restricted to the Massif de la Bibliography: Hedges, S.B. (1992), Hedges, S.B. (1993), Hedges, S.B. (1999), Hedges, S.B. (2001), Henderson, R.W. and Powell, R (1999),
Selle, in Haiti, where it has been recorded from 1,720-2,200m asl. Henderson, R.W. and Powell, R. (2001), Schwartz, A. (1982a), Schwartz, A. and Henderson, R.W. (1991)
Population It is very rare, and only a few specimens have ever been Data Providers: Blair Hedges, Richard Thomas, Robert Powell
found. It was last recorded in the mid 1980s, but there have been
few subsequent surveys.
Habitat and Ecology The species is known to occur in high-eleva-
tion pine forest, and has not been recorded from disturbed habitats.
Eggs are laid on the ground, and it breeds by direct development.
Major Threats Severe habitat destruction is the primary threat to
this species, largely as a result of logging by local people (charcoal-
ing) and slash-and-burn agriculture.
Conservation Measures The species occurs in the Parc National
Morne La Visite, but there is no active management of this area for
conservation, and the habitat continues to be destroyed. Improved
management of this protected area, and maintenance of the existing
habitat, is urgently needed.
Geographic Range This species is known from three localities Cotacachi-Cayapas in Ecuador and in La Planada private reserve in Colombia.
in the foothills of the Andes in north-western Ecuador and south- Bibliography: Acosta-Galvis, A.R. (2000), Lynch, J.D. and Duellman, W.E. (1997), Lynch, J.D. and Duellman, W.E. (1997)
western Colombia: El Cristal, in Esmeraldas Province, Ecuador; Alto Data Providers: Fernando Castro, Santiago Ron, Luis A. Coloma
Tambo, in Esmeraldas Province, Ecuador; and the Altaquer-Tumaco
Road, department of Nariño, Colombia. Its known altitudinal range
is 830-1,200m asl.
Population It is a very rare species.
Habitat and Ecology It lives in humid montane and submontane
forests, and also in disturbed forest, but not in open areas. It is
generally active on low vegetation at night. It is presumed to breed
by direct development, but the site of egg deposition is not known.
Major Threats The major threats are habitat loss -- most likely due
to deforestation for agricultural development, plantations, illegal
crops, logging, and human settlement -- and pollution (resulting
from the spraying of illegal crops).
Conservation Measures It occurs in the Reserva Ecológica
332 Threatened Amphibians of the World

EN Eleutherodactylus deinops Lynch, 1996

Endangered B1ab(iii) Geographic Range This species is known from Valle del Cauca Department on the western flank of the Cordillera
Order, Family: Anura, Leptodactylidae Occidental in Colombia, from 1,750-2,600m asl.
Country Distribution: Colombia Population It is not a rare species, but it is not common either.
Current Population Trend: Decreasing Habitat and Ecology It is usually active in the medium-high canopy inside primary and good quality secondary
forest. It breeds by direct development.
Major Threats The major threat to the species is habitat loss caused by deforestation for agricultural development
(including the planting of illegal crops).
Conservation Measures It occurs in Parque Nacional Natural Los Farallones de Cali.
Bibliography: Acosta-Galvis, A.R. (2000), Lynch, J.D. (1996b), Lynch, J.D. (1998b), Lynch, J.D. (1999), Lynch, J.D., Ruiz-Carranza, P.M.
and Ardila-Robayo, M.C. (1997), Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and Lynch, J.D. (1996), Ruiz-Carranza, P.M., Lynch, J.D. and
Ardila-Robayo, M.C. (1997)
© John D. Lynch

Data Providers: Fernando Castro, Maria Isabel Herrera, John Lynch

EN Eleutherodactylus dennisi (Lynch, 1970)

Endangered B1ab(iii) Geographic Range This species is known only from the type locality in Antiguo Morelos, southern Tamaulipas,
Order, Family: Anura, Leptodactylidae Mexico at 250m asl.
Country Distribution: Mexico Population It is a rare species.
Current Population Trend: Unknown Habitat and Ecology It has been collected in caves and lowland forest. Breeding is by direct development.
Major Threats The major threat to this species is habitat loss, particularly due to logging.
Conservation Measures Its range does not include any protected areas. A systematic survey to corroborate the
presence of this species in the type locality is required. This species is protected by Mexican law under the “Special
Protection” category (Pr).
Bibliography: Lynch, J.D. (1970)
Data Providers: Georgina Santos-Barrera, Luis Canseco-Márquez
© Tim Burkhardt

EN Eleutherodactylus devillei (Boulenger, 1880)

Endangered B1ab(iii) Geographic Range This species occurs on the eastern slopes of the Andes in Napo Province, Ecuador, at 2,350-
Order, Family: Anura, Leptodactylidae 3,155m asl. It has been recorded from nine localities and probably occurs somewhat more widely.
Country Distribution: Ecuador Population It is a common species within its small known range.
© Natural History Museum and Biodiversity Research

Current Population Trend: Decreasing Habitat and Ecology It inhabits upper montane forests and clearings and is tolerant of some limited habitat deg-
radation. It is found beneath rocks and logs by day and on low vegetation at night. It is presumed to breed by direct
development, but the site of egg deposition is not known.
Major Threats The major threat is habitat destruction and degradation due to deforestation for agricultural develop-
ment, logging, and human settlement.
Conservation Measures Its geographic range overlaps with Parque Nacional Sumaco, and slightly with the Reserva
Center, University of Kansas

Ecológica Antisana. Improved protection of the montane forest habitat of this species is needed.
Bibliography: Boulenger, G.A. (1880), Flores, G. (1987), Flores, G. (1988a), Lynch, J.D. (1983), Lynch, J.D. and Duellman, W.E. (1980),
Lynch, J.D. and Duellman, W.E. (1997)
Data Providers: John Lynch, Luis A. Coloma, Santiago Ron

VU Eleutherodactylus diaphonus Lynch, 1986

Vulnerable D2 Geographic Range This species is only known from the type locality (over about 1km) of the Alto Río Calima in
Order, Family: Anura, Leptodactylidae the Valle del Cauca department on the western slopes of the Cordillera Occidental, Colombia, from 1,230-1,250m
Country Distribution: Colombia asl. Surveys of the surrounding area have not recorded the species indicating that it is most likely a very restricted
© Natural History Museum and Biodiversity Research

Current Population Trend: Stable range species.

Population It is quite common at the type locality.
Habitat and Ecology It is found in vegetation along the stream and on rocks inside and immediately adjacent to
the stream in low cloud forest. It breeds by direct development. It is only known from this one stream and is thought
to be a microhabitat specialist, like other species of the group.
Major Threats The main threat to this species is potentially the hydroelectric power dam only a few kilometres
Center, University of Kansas

upstream, but there is no access to the species’ habitat as it is part of the dam property. Its habitat does not ap-
pear to be otherwise threatened. Some other species of Eleutherodactylus that are associated with streams have
undergone dramatic declines and disappearances, possibly due to chytridiomycosis, so the status of this species
should be monitored carefully.
Conservation Measures The type locality is not within a protected area, although it lies just outside the Parque
Nacional Natural Farallones de Cali. This locality is within a “protected” area where the Corporación del Valle del
Cauca (CVC) generates electricity as waters leave Lago Calima.
Bibliography: Acosta-Galvis, A.R. (2000), Lynch, J.D. (1986a), Lynch, J.D. (1998b), Lynch, J.D. (1999), Lynch, J.D. and Duellman, W.E.
(1997), Lynch, J.D. and Ruiz-Carranza, P.M. (1996b), Lynch, J.D., Ruiz-Carranza, P.M. and Ardila-Robayo, M.C. (1997), Ruiz-Carranza, P.M.,
Ardila-Robayo, M.C. and Lynch, J.D. (1996)
Data Providers: Fernando Castro, Maria Isabel Herrera, John Lynch

EN Eleutherodactylus dilatus (Davis and Dixon, 1955)
Endangered B1ab(iii)+2ab(iii)
Order, Family: Anura, Leptodactylidae
Country Distribution: Mexico
Current Population Trend: Decreasing
VU Eleutherodactylus diogenes Lynch and Ruíz-Carranza, 1996
Vulnerable D2
Order, Family: Anura, Leptodactylidae
Country Distribution: Colombia
Current Population Trend: Stable
EN Eleutherodactylus dissimulatus Lynch and Duellman, 1997
Endangered B1ab(iii)
Order, Family: Anura, Leptodactylidae
Country Distribution: Ecuador
Current Population Trend: Decreasing
© John D. Lynch
CR Eleutherodactylus dixoni Lynch, 1991
Critically Endangered B1ab(iii)+2ab(iii)
Order, Family: Anura, Leptodactylidae
Country Distribution: Mexico
Current Population Trend: Decreasing
Globally Threatened Amphibian Species 333
Geographic Range This species is found in Omiltemi, Mazatlan Bibliography: Dixon, J.R. (1957)
and the surrounding areas of Chilpancingo, in north-central Guerrero, Data Providers: Georgina Santos-Barrera, Luis Canseco-Márquez
Mexico. The type locality is at about 2,250m asl. It might occur a
little more widely.
Population This is not a common species.
Habitat and Ecology It inhabits pine-oak forest and crops in the
forest, and is commonly found under logs and rocks during the day.
During the mating season males climb on rocks and bushes, and
breeding is by direct development.
Major Threats Disturbance and/or loss of the forests north of
Chilpancingo are the main threat for this species since it is restricted
to high-elevation forests in the Omiltemi area.
Conservation Measures The Omiltemi Park is under protection
by the local people, although it no longer has any formal protection
status and is subject to increasing anthropogenic disturbance. There
is a need to protect the montane forest habitat of this species. A
new survey to evaluate the current status of the populations in
Guerrero is required.
Geographic Range This species is known from two localities close well-protected primary forest. The maintenance of these protected areas is essential for the long-term survival of
together; one in Parque Nacional Natural Munchique and the other this restricted-range, microhabitat specialist.
just outside the park, both in Cauca department, on the western slope Bibliography: Acosta-Galvis, A.R. (2000), Lynch, J.D. (1998b), Lynch, J.D. and Ruiz-Carranza, P.M. (1996b), Lynch, J.D., Ruiz-Carranza,
of the Cordillera Occidental in Colombia, from 1,470-1,600m asl. P.M. and Ardila-Robayo, M.C. (1997), Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and Lynch, J.D. (1996)
Population It is not thought to be a rare species. Data Providers: Fernando Castro, Maria Isabel Herrera, John Lynch
Habitat and Ecology This species is a microhabitat specialist, found
on rocks near waterfalls in streams in primary forests. It has not been
recorded outside primary forest. It breeds by direct development.
Major Threats There are no major threats to the species’ habitat
at present, as its habitat is relatively well protected. Some other
species of Eleutherodactylus that are associated with streams at high
elevations have undergone dramatic declines and disappearances,
possibly due to chytridiomycosis, so the status of this species should
be monitored carefully.
Conservation Measures One locality is within Parque Nacional
Natural Munchique, and the other is in a private reserve that has
Geographic Range This species is known from two localities on the western slopes of the Andes in Pichincha
Province, north-western Ecuador: Quebrada Zapadores, near Chiriboga, at 2,020m asl; and from San Ignacio, at
1,920m asl. Reports from Colombia are in error (J. Lynch pers. comm.).
Population It is not a common species.
Habitat and Ecology It inhabits montane cloud forest, although its ability to adapt to modified habitats is unknown.
Specimens have been found on low vegetation along streams at night. It is presumed to breed by direct development,
but the site of egg deposition is not known.
Major Threats Habitat destruction and degradation is ongoing within its range, due to expanding agricultural activi-
ties (including the rearing of livestock) and human settlements.
Conservation Measures It is not known from any protected areas, and there is an urgent need for the protection
of the montane cloud forest habitat of this species.
Bibliography: Lynch, J.D. and Duellman, W.E. (1997)
Data Providers: John Lynch, Luis A. Coloma, Santiago Ron
Geographic Range This species is known only from Agua del Bibliography: Hedges, S.B. (1989), Lynch, J.D. (1991b)
Obispo, central Guerrero, Mexico. Data Providers: Georgina Santos-Barrera, Luis Canseco-Márquez
Population This is a rare species.
Habitat and Ecology It inhabits a small area of pine-oak forest
at moderate elevations. Individuals are frequently found perched
on the branches of trees from which they usually call. It breeds by
direct development.
Major Threats The primary threat is habitat loss and degradation, as
the forested areas surrounding Agua del Obispo have been severely
disturbed or transformed by human activities.
Conservation Measures The range of this species does not include
any protected areas, and an urgent programme of protection and
restoration of the Agua del Obispo area is required. Further survey
work to evaluate the population status of this species is also neces-
sary. It is protected by Mexican law under the “Special Protection”
category (Pr).
334 Threatened Amphibians of the World

CR Eleutherodactylus dolomedes Hedges and Thomas, 1992

Critically Endangered A3c; B1ab(iii)+2ab(iii) Geographic Range This species is restricted to one site on the Massif de la Hotte, in Haiti, at 1,120m asl.
Order, Family: Anura, Leptodactylidae Population It is known only from a few specimens.
Country Distribution: Haiti Habitat and Ecology It is an arboreal species, occurring in high-elevation cloud forest. While it has been recorded from
Current Population Trend: Decreasing forest edge, this is probably not suitable habitat. Eggs are laid on the ground, and it breeds by direct development.
Major Threats The primary threat to the species is severe habitat destruction, which is taking place as a result of
logging by local people (charcoaling) and slash-and-burn agriculture.
Conservation Measures This species is known only from the Parc National Macaya; there is no active manage-
ment of this area for conservation, and the habitat continues to be destroyed. Urgent site-based action is required

© S. Blair Hedges, Penn State

in the Massif de la Hotte to conserve the remaining habitat in the area, in order to ensure the persistence of this
species as well as other threatened amphibians known only from this area. Survey work is required to determine
the population status of this species.
Bibliography: Hedges, S.B. (1993), Hedges, S.B. (1999), Hedges, S.B. (2001), Hedges, S.B. and Thomas, R. (1992b), Henderson, R.W.
and Powell, R (1999), Henderson, R.W. and Powell, R. (2001), Schwartz, A. and Henderson, R.W. (1991)
Data Providers: Blair Hedges, Richard Thomas

VU Eleutherodactylus dolops Lynch and Duellman, 1980

Vulnerable B1ab(iii) Geographic Range This species occurs at an elevation of 1,440-1,950m asl on the eastern face of the Andes in
Order, Family: Anura, Leptodactylidae Putumayo and Caqueta (Colombia) and Napo (Ecuador). In Ecuador it is only known from three nearby localities.
Country Distribution: Colombia, Ecuador Population Generally, this is not a common frog anywhere in its range, although a good-sized series was collected
© Natural History Museum and Biodiversity Research

Current Population Trend: Unknown in Colombia in 1990 in Parque Nacional Natural Alto Fragua.
Habitat and Ecology This species occurs in cloud forest, and is associated with stream habitats (Lynch and Duellman
1980). It is presumed to breed by direct development, but the site of egg deposition is not known. The ability of this
species to adapt to modified habitats is unknown.
Major Threats The major threat is habitat loss due to agriculture (including the planting of illegal crops) and logging.
Some other species of Eleutherodactylus that are associated with streams have undergone dramatic declines and
Center, University of Kansas

disappearances, possibly due to chytridiomycosis, so the status of this species should be monitored carefully.
Conservation Measures This species occurs in Parque Nacional Natural Alto Fragua-Indi Wasa, in Colombia. Its
range overlaps with the Reserva Ecológica Cayambe-Coca in Ecuador.
Bibliography: Acosta-Galvis, A.R. (2000), Lynch, J.D. (1989), Lynch, J.D. and Duellman, W.E. (1980), Lynch, J.D. and Duellman, W.E.
(1997), Lynch, J.D., Ruiz-Carranza, P.M. and Ardila-Robayo, M.C. (1997), Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and Lynch, J.D. (1996),
Suarez, A.M. (1999)
Data Providers: Fernando Castro, Maria Isabel Herrera, Luis A. Coloma, Santiago Ron, John Lynch

EN Eleutherodactylus dorsopictus Rivero and Serna, 1988 “1987”

Endangered B1ab(iii) Geographic Range This species is known from Páramo Sonsón, in Bibliography: Acosta-Galvis, A.R. (2000), Ardila-Robayo, M.C. and Acosta-Galvis, A. (2000a), Lynch, J.D. and Duellman, W.E. (1997),
Order, Family: Anura, Leptodactylidae southern Antioquia and eastern Caldas Departments, in the northern Lynch, J.D., Ruiz-Carranza, P.M. and Ardila-Robayo, M.C. (1997), Paez, V.P. et al. (2002), Rivero, J.A. and Serna, M.A. (1987), Rueda-
Country Distribution: Colombia part of the Cordillera Central in Colombia, from 2,400-3,000m asl. Almonacid, J.V. (2000), Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and Lynch, J.D. (1996)
Current Population Trend: Decreasing Population It is a reasonably common species. Data Providers: Fernando Castro, Maria Isabel Herrera, John Lynch
Habitat and Ecology An inhabitant of sub-páramo, it is usually
active among vegetation in the low stratum and is often found
sheltering in epiphytes, in forested areas, but it can also be found
in open areas. It has not been recorded from disturbed forest or
from anthropogenic habitats, although it may occur in these areas.
It breeds by direct development.
Major Threats The major threat to this species is habitat loss
caused by deforestation for agricultural development (including the
planting of illegal crops).
Conservation Measures It is not known to occur in any protected
areas, and formal habitat protection is urgently needed to secure
the future survival of this species.

VU Eleutherodactylus douglasi Lynch, 1996

Vulnerable B1ab(iii) Geographic Range This species can be found on the eastern and western flanks of the Cordillera Oriental in Santander
Order, Family: Anura, Leptodactylidae and Norte de Santander, Colombia, at 1,800-2,550m asl. It might occur a little more widely.
Country Distribution: Colombia Population It is a common species.
Current Population Trend: Decreasing Habitat and Ecology It occurs in the understorey of cloud forests, sometimes near streams. It also occurs in secondary
forest, but has not been found outside forested areas. Reproduction takes place by direct development.
Major Threats Habitat loss due to agricultural crops and cattle ranching is the main threat to this species.
Conservation Measures It occurs in the Estación El Rasgón.
Bibliography: Acosta-Galvis, A.R. (2000), Lynch, J.D. (1996a), Lynch, J.D. and Duellman, W.E. (1997), Lynch, J.D., Ruiz-Carranza, P.M.
and Ardila-Robayo, M.C. (1997), Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and Lynch, J.D. (1996)
Data Providers: Fernando Castro, Maria Isabel Herrera, Martha Patricia Ramírez Pinilla
© John D. Lynch
 Globally Threatened Amphibian Species 335

VU Eleutherodactylus duellmani Lynch, 1980

Vulnerable B1ab(iii) Geographic Range This species occurs on the Pacific slopes of the western Andes in northern Ecuador (Carchi,
Order, Family: Anura, Leptodactylidae Imabura, Pichincha Province) and southern Colombia (departments of Cauca and Nariño), from 1,780-2,700m asl.
Country Distribution: Colombia, Ecuador Population In 1977, this species was recorded as abundant in Carchi and Imbabura in Ecuador. However, during

© Natural History Museum and Biodiversity Research

Current Population Trend: Decreasing surveys conducted between 1997 and 2003, it was found to be rare at many localities.
Habitat and Ecology This is a montane tropical cloud forest species, and is present in humid microhabitats. At
night,it perches on branches and herbaceous vegetation immediately above small streams and sits on rock faces
in the spray zones of waterfalls; some have been found in crevices or between rocks in small streams (Lynch and
Duellman 1997). It is presumed to be a species that breeds by direct development, but the site of egg deposition is
not known. The ability of this species to adapt to modified habitats is unknown.
Center, University of Kansas Major Threats Habitat loss due to logging and agricultural development (including the planting of illegal crops) is
a major threat. Some other species of Eleutherodactylus that are associated with streams have undergone dramatic
declines and disappearances, possibly due to chytridiomycosis, so the status of this species should be monitored
carefully.
Conservation Measures In Colombia, it is recorded from Parque Nacional Natural Munchique (Cauca) and La
Planada (Nariño). In Ecuador, its range overlaps with Reserva Geobotánica Pululahua.
Bibliography: Acosta-Galvis, A.R. (2000), Lynch, J.D. (1980d), Lynch, J.D. (1998b), Lynch, J.D. and Burrowes, P.A. (1990), Lynch, J.D.
and Duellman, W.E. (1997), Lynch, J.D., Ruiz-Carranza, P.M. and Ardila-Robayo, M.C. (1997), Ruiz-Carranza, P.M., Ardila-Robayo, M.C.
and Lynch, J.D. (1996)
Data Providers: Fernando Castro, Maria Isabel Herrera, Santiago Ron, Luis A. Coloma, Diego Almeida, John Lynch

EN Eleutherodactylus elassodiscus Lynch, 1973

Endangered B1ab(iii) Geographic Range This species is found at intermediate elevations (from 2,300-2,900m asl) on the Amazonian
Order, Family: Anura, Leptodactylidae slopes of the Andes in northern Ecuador (Sucumbios and Napo Provinces) and southern Colombia (Putumayo Province,
Country Distribution: Colombia, Ecuador eastern flank of the Nudo de Pasto). It is known from relatively few localities.
© Natural History Museum and Biodiversity Research

Current Population Trend: Decreasing Population It has never been a common species but is regularly encountered in appropriate habitat. The last records
might be from the 1970s in Ecuador, but there have been no known recent attempts to find it.
Habitat and Ecology This terrestrial species occurs in sloping areas of upper humid montane forest, but can also
be found in forest edges. Specimens were found under rocks and logs, and on areas of humid grass by day (Lynch
and Duellman 1980). Reproduction occurs by direct development.
Major Threats The major threats to this species are general habitat loss, caused by agricultural development
Center, University of Kansas

(including the planting of illegal crops), and pollution (from the spraying of illegal crops).
Conservation Measures In Ecuador, its geographic range overlaps with the Reserva Ecológica Cayambe-Coca;
however, it does not occur in any protected areas in Colombia. There is a need for improved protection of the
montane forest habitat of this species. Further survey work is also necessary to determine the current population
status of this species.
Bibliography: Acosta-Galvis, A.R. (2000), Lynch, J.D. (1973a), Lynch, J.D. (1989), Lynch, J.D. and Duellman, W.E. (1980), Lynch, J.D.
and Duellman, W.E. (1997), Lynch, J.D., Ruiz-Carranza, P.M. and Ardila-Robayo, M.C. (1997), Ruiz-Carranza, P.M., Ardila-Robayo, M.C.
and Lynch, J.D. (1996)
Data Providers: Fernando Castro, Maria Isabel Herrera, Luis A. Coloma, Santiago Ron, John Lynch

VU Eleutherodactylus elegans (Peters, 1863)

Vulnerable B1ab(iii) Geographic Range This species can be found on the eastern slopes of the Cordillera Oriental in the páramos of Chin-
Order, Family: Anura, Leptodactylidae gaza, Sumapaz, and La Rusia, in the departments of Cundinamarca and Boyacá, Colombia, from 2,600-3,300m asl.
Country Distribution: Colombia Population It is an abundant species.
Current Population Trend: Decreasing Habitat and Ecology This species occurs in páramo areas, and also in cloud forest. It is associated with herbaceous
vegetation, and very small bushes.
Major Threats The major threat is habitat loss and degradation due to crop agriculture and cattle ranching.
Conservation Measures It occurs in Parque Nacional Natural Chingaza and Parque Nacional Natural Sumapaz.
Bibliography: Acosta-Galvis, A.R. (2000), Bernal, X. and Guzman, F. (2000), Cochran, D.M. and Goin, C.J. (1970), Luddecke, H. et al.
(1997), Luddecke, H. et al. (2000), Lynch, J.D. and Duellman, W.E. (1997), Lynch, J.D., Ruiz-Carranza, P.M. and Ardila-Robayo, M.C. (1997),
Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and Lynch, J.D. (1996)
Data Providers: Fernando Castro, Maria Isabel Herrera, Adolfo Amézquita, Jose Vicente Rueda
© Taran Grant

EN Eleutherodactylus emiliae Dunn, 1926

Endangered B1ab(iii)+2ab(iii) Geographic Range This species is restricted to the Sierra de Bibliography: Díaz, L.M. (1999), Hedges, S.B. (1993), Hedges, S.B. (1999), Hedges, S.B. (2001), Henderson, R.W. and Powell, R (1999),
Order, Family: Anura, Leptodactylidae Trinidad, Cuba, from 350-830m asl. Henderson, R.W. and Powell, R. (2001), Schwartz, A. and Henderson, R.W. (1991)
Country Distribution: Cuba Population It is an uncommon species. Data Providers: Blair Hedges, Luis Díaz
Current Population Trend: Decreasing Habitat and Ecology It is a terrestrial inhabitant of closed-canopy,
humid forests, and has not been recorded outside forest habitat. Eggs
are laid on the ground, and it breeds by direct development.
Major Threats The major threat to this species is habitat loss due
to agricultural expansion, the development of tourism infrastructure,
and the disturbance of the habitat from tourist activities.
Conservation Measures It is known from a few protected areas,
although these areas do not provide sufficient protection for the
species, and there is a need to provide more adequate protection of
the remaining forest habitat.
336 Threatened Amphibians of the World

CR Eleutherodactylus emleni Dunn and Emlen, 1932

Critically Endangered A2ace; B2ab(v) Geographic Range This species is known from the Río Choluteca decline. It is known to occur in the Parque Nacional La Tigra and the Reserva Biológica Cerro Uyuca.
Order, Family: Anura, Leptodactylidae watershed from Cerro La Tigra and Cerro Uyuca to El Zamorano in Bibliography: Campbell, J.A. and Savage, J.M. (2000), McCranie, J.R. and Wilson, L.D. (2002b)
Country Distribution: Honduras south-central Honduras at elevations of 800-2,000m asl. Data Providers: Larry David Wilson, Randy McCranie
Current Population Trend: Decreasing Population There have been no sightings of this species since 1985
despite one targeted effort to look for this species. Considering
that all streamside Eleutherodactylus occurring above 900m asl in
Honduras have disappeared recently, this species might be extinct.
Habitat and Ecology The preferred habitat of this species is pre-
montane and lower montane moist forest. It occurs in leaf-litter, and
has been collected along streams and in a citrus grove away from
streams. Reproduction occurs by direct development.
Major Threats Although the precise reasons for the declines seen
in this species are unclear, it is possible that the apparent disap-
pearance might be due to chytridiomycosis.
Conservation Measures Survey work is urgently needed to deter-
mine the population status of this species, and the reasons for its

CR Eleutherodactylus eneidae Rivero, 1959 ELEGANT COQUI

Critically Endangered A2ae Geographic Range This species formerly occurred in the interior uplands of Puerto Rico, at an altitude of 300-
Order, Family: Anura, Leptodactylidae 1,150m asl.
Country Distribution: Puerto Rico Population Formerly uncommon even in the 1980s, this species was last recorded in 1990 and subsequent extensive
Current Population Trend: Decreasing searches have failed to locate this species. It is now believed to be most probably extinct.
Habitat and Ecology A terrestrial species that breeds by direct development, it is known from extremely humid
closed-canopy forest.
Major Threats The cause of its disappearance is most likely chytridiomycosis linked to climate change, although
invasive rats and other predators have also been suggested.
© S. Blair Hedges, Penn State

Conservation Measures This species was known to occur in Luquillo National Forest in the El Yunque area. In
view of the risk of chytridiomycosis, it is a very high priority to conduct surveys to determine whether or not this
species could still survive in the wild; surviving individuals might need to form the basis for the establishment of
an ex-situ population.
Bibliography: Burrowes, P.A., Joglar, R.L. and Green, D.E. (2004), Hedges, S.B. (1993), Henderson, R.W. and Powell, R (1999), Henderson,
R.W. and Powell, R. (2001), Joglar, R.L. (1999), Rivero, J.A. (1998)
Data Providers: Blair Hedges, Rafael Joglar, Luis J. Villanueva-Rivera, Neftalí Ríos-Lopez

VU Eleutherodactylus eremitus Lynch, 1980

Vulnerable B1ab(iii) Geographic Range This species is found at elevations of 1,540-2,100m asl on the Pacific slopes of the western
Order, Family: Anura, Leptodactylidae Andes in Ecuador (Saloya Valley, Pichincha Province) and extreme south-western Colombia (La Planada, Nariño
Country Distribution: Colombia, Ecuador Department). It probably occurs more widely than current records suggest.
Current Population Trend: Decreasing Population It was abundant in suitable habitat in La Planada, Colombia, in the late 1980s, and has been recorded
during the course of recent surveys in Ecuador.
Habitat and Ecology It is a montane cloud forest species. It occurs in old growth and secondary forest, but not
outside forest habitats. During the day, it can be found in epiphytic bromeliads as much as 7m off the ground. It is
presumed to breed by direct development, but the site of egg deposition is not known.
Major Threats General habitat loss due to logging and agricultural development (including the planting of illegal
crops) is a major threat.
Conservation Measures In Colombia, the species occurs in Reserva La Planada (department of Nariño), and in
Ecuador its range overlaps with Reserva Ecológica Los Illinizas.
© John D. Lynch

Bibliography: Acosta-Galvis, A.R. (2000), Lynch, J.D. (1980a), Lynch, J.D. (1998b), Lynch, J.D. and Burrowes, P.A. (1990), Lynch, J.D.
and Duellman, W.E. (1997), Lynch, J.D., Ruiz-Carranza, P.M. and Ardila-Robayo, M.C. (1997), Ruiz-Carranza, P.M., Ardila-Robayo, M.C.
and Lynch, J.D. (1996)
Data Providers: Fernando Castro, Maria Isabel Herrera, Santiago Ron, Luis A. Coloma, Diego Cisneros-Heredia, John Lynch, Mario
Yánez-Muñoz

VU Eleutherodactylus eriphus Lynch and Duellman, 1980

Vulnerable B1ab(iii) Geographic Range This species can be found in the valley of the Río Papallacta on the eastern face of the Andes
Order, Family: Anura, Leptodactylidae in Napo Province, Ecuador, and Putumayo Department, Colombia, at elevations of 2,160-2,630m asl. In Ecuador, the
Country Distribution: Colombia, Ecuador species is only known from four localities. It probably occurs more widely.
© Natural History Museum and Biodiversity Research

Current Population Trend: Decreasing Population This species was recorded as common in Guacamayos, La Sofia, Cosanga, and Reventador, Ecuador,
in 2003.
Habitat and Ecology This species has been found in partially cleared and old growth cloud forest. It is active
at night on top of low herbaceous vegetation. It is presumed to breed by direct development, but the site of egg
deposition is not known.
Major Threats General habitat loss due to logging and agricultural development (including the planting of illegal
Center, University of Kansas

crops) is a major threat.

Conservation Measures It occurs in Reserva Ecológica Cayambe-Coca and Reserva Ecológica Antisana in
Ecuador.
Bibliography: Acosta-Galvis, A.R. (2000), Funk, C.W. et al. (2003), Lynch, J.D. and Duellman, W.E. (1980), Lynch, J.D. and Duellman, W.E.
(1997), Lynch, J.D., Ruiz-Carranza, P.M. and Ardila-Robayo, M.C. (1997), Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and Lynch, J.D. (1996)
Data Providers: Fernando Castro, Maria Isabel Herrera, Santiago Ron, Luis A. Coloma, Diego Almeida, John Lynch, Mario Yánez-
Muñoz, Ana Almandáriz
 Globally Threatened Amphibian Species 337

VU Eleutherodactylus ernesti Flores, 1987

Vulnerable D2 Geographic Range This species is restricted to the peak of Volcán Bibliography: Flores, G. (1987), Lynch, J.D. and Duellman, W.E. (1997)
Order, Family: Anura, Leptodactylidae Sumaco, in Napo Province, Ecuador, at around 3,900m asl. It is not Data Providers: John Lynch, Luis A. Coloma, Santiago Ron
Country Distribution: Ecuador likely to occur elsewhere.
Current Population Trend: Stable Population The population status of this species is unknown.
Habitat and Ecology It lives in high-elevation grassland and
bushland habitats on the Volcán Sumaco. It is presumed to breed
by direct development, but the site of egg deposition is not known.
Major Threats There is no recent information on threats in its only
known locality. Its highly restricted range renders it susceptible to
stochastic threatening processes.
Conservation Measures It is not known from any protected areas.
There is a need for further survey work to investigate the current
population status of this species, and to determine whether or not
there are any immediate threats at present. Close population monitor-
ing is required given that it is known only from the Volcán Sumaco.

EN Eleutherodactylus etheridgei Schwartz, 1958

Endangered B1ab(iii)+2ab(iii) Geographic Range This species occurs in Santiago de Cuba and Bibliography: Hedges, S.B. (1993), Hedges, S.B. (1999), Hedges, S.B. (2001), Henderson, R.W. and Powell, R (1999), Henderson, R.W.
Order, Family: Anura, Leptodactylidae Guantanamo Province, in Cuba, from sea level up to 151m asl. It and Powell, R. (2001), Schwartz, A. (1958c), Schwartz, A. and Henderson, R.W. (1991)
Country Distribution: Cuba occurs on Guantanamo Naval Station and has only rarely been Data Providers: Blair Hedges, Luis Díaz
Current Population Trend: Decreasing encountered outside this site.
Population It is moderately common in suitable habitat.
Habitat and Ecology This species is found in xeric rocky regions at
low elevations in mesic tropical forest, and has not been recorded
outside forest habitat. It breeds by direct development.
Major Threats Infrastructure development for human settlement
and tourist facilities, as well as the disturbance of the habitat by
tourists, is the major threat to this species.
Conservation Measures It is known from two protected areas, and
the naval station affords the species some level of protection.

EN Eleutherodactylus eugeniae Lynch and Duellman, 1997

Endangered B1ab(iii) Geographic Range This species occurs in four localities in a limited area of cloud forest in the upper valley of the
Order, Family: Anura, Leptodactylidae Río Pilatón in the province of Pichincha, Ecuador, at elevations of 1,700-2,010m asl.
Country Distribution: Ecuador Population It is an uncommon species. It has been recorded as recently as December 2002 in Pichincha.
Current Population Trend: Decreasing Habitat and Ecology It is an inhabitant of cloud forest, in humid subtropical and humid temperate regimes, and is
usually found away from streams. It occurs in bromeliads in the sub canopy. It breeds by direct development, which
takes place in bromeliads.
Major Threats Habitat loss, due to logging, is the major threat to the species.
© Mario Humberto Yánez-Muñoz

Conservation Measures It is not known to occur in any protected areas, and protection of the cloud forest habitat
of this species is urgently needed.
Bibliography: Guayasamin, J.M. et al. (2004), Lynch, J.D. and Duellman, W.E. (1997)
Data Providers: Luis A. Coloma, Santiago Ron, Mario Yánez-Muñoz

CR Eleutherodactylus eunaster Schwartz, 1973

Critically Endangered A3c Geographic Range This species is restricted to the Massif de la Hotte, in Haiti, at 575-1,300m asl.
Order, Family: Anura, Leptodactylidae Population It is known to be a moderately common species in suitable habitat; however, it was last recorded in
Country Distribution: Haiti 1991, but there have been no surveys for the species in subsequent years.
Current Population Trend: Decreasing Habitat and Ecology It is an arboreal species, occurring in mesic hardwood closed-canopy forests, and has not been
recorded from disturbed forest. Eggs are laid on the ground, and it breeds by direct development.
Major Threats The primary threat to the species is severe habitat destruction, which is taking place as a result of
logging by local people (charcoaling) and slash-and-burn agriculture.
Conservation Measures Although it occurs in the Parc National Macaya, there is no management of this area
for conservation, and the habitat continues to be destroyed. Urgent site-based action is required in the Massif de
© Eladio Fernandez 2006

la Hotte to conserve the remaining habitat in the area, in order to ensure the persistence of this species as well as
other threatened amphibians known only from this area. Survey work is also required to determine the population
status of this species.
Bibliography: Hedges, S.B. (1993), Hedges, S.B. (1999), Hedges, S.B. (2001), Henderson, R.W. and Powell, R (1999), Henderson, R.W.
and Powell, R. (2001), Schwartz, A. (1973), Schwartz, A. (1982b), Schwartz, A. and Henderson, R.W. (1991)
Data Providers: Blair Hedges, Richard Thomas, Robert Powell
338 Threatened Amphibians of the World

EN Eleutherodactylus euphronides Schwartz, 1967

Endangered B1ab(iii) Geographic Range This species is restricted to central and south- Bibliography: Germano, J.M. et al. (2003), Hedges, S.B. (1993), Hedges, S.B. (1999), Hedges, S.B. (2001), Henderson, R.W. and Powell,
Order, Family: Anura, Leptodactylidae eastern Grenada, Lesser Antilles, where it has been recorded from R (1999), Henderson, R.W. and Powell, R. (2001), Kaiser, H., Green, D.M. and Schmid, M. (1994), Kaiser, H., Hardy, Jr., J.D. and Green,
Country Distribution: Grenada 300-840m asl. D.M. (1994), Sander, J.M., Kaiser, H. and Powell, R. (2003), Schwartz, A. (1967), Schwartz, A. and Henderson, R.W. (1991)
Current Population Trend: Decreasing Population This species is poorly known, but it appears to be Data Providers: Blair Hedges, Robert Powell
moderately common in suitable habitat.
Habitat and Ecology It inhabits rainforests and forest edge and
montane meadows surrounded by agriculture. It lives on the ground
and on vegetation. The eggs are laid on the ground, and it breeds
by direct development.
Major Threats Habitat loss and destruction is the major threat to
this species, primarily due to urbanization and tourism development,
as well as agriculture.
Conservation Measures This species is present in the Grand
Etang Forest Reserve. Increased protection of its remaining rainfor-
est habitat is needed.

EN Eleutherodactylus fallax Lynch and Rueda, 1999

Endangered B1ab(iii) Geographic Range This species is found along forested streams in Antioquia, Tolima, and Caldas Departments,
Order, Family: Anura, Leptodactylidae on the eastern flank of the Cordillera Central, in Colombia, from 1,100-1,850m asl. The type locality is in San Carlos
Country Distribution: Colombia (Lynch and Rueda-Almonacid 1999).
Current Population Trend: Decreasing Population There is no information on its current population status.
Habitat and Ecology This is a forest species usually found along the edge of streams that is associated with naturally
© Fundación ProAves / www.proaves.org

occurring open areas in primary and secondary forests. Gallery forest cover is needed to keep humidity high; hence,
it is very sensitive to any disturbance of its habitat. It breeds by direct development.
Major Threats The main threat to this species is habitat loss due to deforestation for agricultural development and
the planting of illegal crops, although these activities are mainly localized at present. Some other species of Eleu-
therodactylus that are associated with high-elevation streams have undergone dramatic declines and disappearances,
possibly due to chytridiomycosis, so the status of this species should be monitored carefully.
Conservation Measures There are no protected areas within the range of the species, and its habitat is in need
of urgent protection (particularly given its dependence on gallery forest to maintain humidity levels). Further survey
work is needed to determine the current population status of this species.
Bibliography: Acosta-Galvis, A.R. (2000), Kaiser, H. (1995), Lescure, J. (2000), Lynch, J.D. and Rueda-Almonacid, J.V. (1999), Paez, V.P.
et al. (2002), Rueda-Almonacid, J.V. (2000)
Data Providers: Fernando Castro, Maria Isabel Herrera

EN Eleutherodactylus fetosus Lynch and Rueda-Almonacid, 1998

Endangered B1ab(iii) Geographic Range This species is known only from a few Conservation Measures There are no protected areas within the range of the species, and there is a need for the
Order, Family: Anura, Leptodactylidae localities within the vicinity of the type locality, Puerto Suárez, on protection of remaining tracts of montane forest in this region.
Country Distribution: Colombia the Cordillera Occidental in Caldas Department, Colombia, from Bibliography: Acosta-Galvis, A.R. (2000), Lynch, J.D. and Rueda-Almonacid, J.V. (1998a), Rueda-Almonacid, J.V. (2000)
Current Population Trend: Decreasing 1,800-2,650m asl. Data Providers: Fernando Castro, Maria Isabel Herrera, John Lynch
Population It is not a common species.
Habitat and Ecology It is found in primary and secondary forests,
and is very dependent on humid conditions and streamside condi-
tions. It can be found on rocks and low vegetation. It needs gallery
forest to maintain the humid conditions, and hence is not tolerant of
any disturbance of its habitat. It breeds by direct development.
Major Threats Habitat loss, as a result of deforestation for agri-
cultural development (including the planting of illegal crops), is the
major threat to this species. Some other species of Eleutherodactylus
associated with montane streams have undergone dramatic declines
and disappearances, possibly due to chytridiomycosis, so the status
of this species should be monitored carefully.

VU Eleutherodactylus floridus Lynch and Duellman, 1997

Vulnerable B1ab(iii) Geographic Range This species occurs on the western flanks of
Order, Family: Anura, Leptodactylidae the Andes in the Ecuadorian provinces of Cotopaxi, Imbabura, and
Country Distribution: Ecuador Pichincha, at 700-2,000m asl.
Current Population Trend: Decreasing Population It is a reasonably common species.
Habitat and Ecology It inhabits humid lowland tropical forest and
cloud forest. Its ability to adapt to modified habitats is unknown.
It is presumed to breed by direct development, but the site of egg
deposition is not known.
Major Threats Habitat destruction and degradation are taking place
due to agricultural development, logging, and human settlement.
Conservation Measures Its range overlaps with the Reserva
Ecológica Cotacachi-Cayapas and Reserva Ecológica Los Illinizas.
Bibliography: Lynch, J.D. and Duellman, W.E. (1997)
Data Providers: John Lynch, Luis A. Coloma, Santiago Ron
 Globally Threatened Amphibian Species 339

CR Eleutherodactylus fowleri Schwartz, 1973

Critically Endangered B1ab(iii)+2ab(iii) Geographic Range This species has a restricted range on the Massif de la Selle, Hispaniola (Haiti and Dominican
Order, Family: Anura, Leptodactylidae Republic) where it is known from only two isolated sites, one in Haiti, and one in the Dominican Republic. Its altitu-
Country Distribution: Dominican Republic, Haiti dinal range is 1,045-1,303m asl.
Current Population Trend: Decreasing Population This species is probably very rare; very few specimens have ever been collected and few individuals
have been heard calling.
Habitat and Ecology It inhabits high-elevation mesic forests, occurring in bromeliads in tall trees (which it uses as
diurnal retreats, and for calling sites and breeding). It breeds by direct development.
Major Threats The primary threat to the species is habitat destruction due to small-scale agriculture and charcoaling,

© S. Blair Hedges, Penn State

a particular threat to the tall trees that are its favoured habitat.
Conservation Measures The species occurs in the Parque Nacional Sierra de Baoruco in the Dominican Republic,
but is not known to occur in a protected area in Haiti. Maintenance of the existing habitat is urgently needed, and
further survey work is required to determine the current population status of the species.
Bibliography: Hedges, S.B. (1993), Hedges, S.B. (1999), Hedges, S.B. (2001), Henderson, R.W. and Powell, R (1999), Henderson, R.W.
and Powell, R. (2001), Schwartz, A. (1973), Schwartz, A. (1982d), Schwartz, A. and Henderson, R.W. (1991)
Data Providers: Blair Hedges, Sixto Inchaustegui, Marcelino Hernandez, Robert Powell

VU Eleutherodactylus frater (Werner, 1899)

Vulnerable B1ab(iii) Geographic Range This species can be found on the eastern slopes of the Cordillera Oriental in the departments
Order, Family: Anura, Leptodactylidae of Cundinamarca and Meta, and from the Serranía de la Macarena, in Colombia. Its altitudinal range is from 1,000-
Country Distribution: Colombia 1,600m asl. It might be more widely distributed than existing records indicate.
Current Population Trend: Decreasing Population It is a common species, and was last collected in 1993 when the last surveys took place.
Habitat and Ecology This species inhabits premontane humid forests, and is found on low vegetation. It has been
recorded from secondary forest. Reproduction occurs by direct development.
Major Threats Habitat loss and degradation due to agriculture (crops and cattle ranching) is the main threat to
this species.
Conservation Measures It has been recorded from Parque Nacional Natural Serrania de la Macarena.
Bibliography: Acosta-Galvis, A.R. (2000), Cochran, D.M. and Goin, C.J. (1970), Flores, G. and Vigle, G.O. (1994), Lynch, J.D. and Duellman,
W.E. (1997), Lynch, J.D., Ruiz-Carranza, P.M. and Ardila-Robayo, M.C. (1997), Pyburn, W.F. and Lynch, J.D. (1981), Ruiz-Carranza, P.M.,
© John D. Lynch

Ardila-Robayo, M.C. and Lynch, J.D. (1996)

Data Providers: Fernando Castro, Maria Isabel Herrera, Jose Vicente Rueda

CR Eleutherodactylus furcyensis Shreve and Williams, 1963

Critically Endangered A3c Geographic Range This species is restricted to the Massif de la Morne La Visite in Haiti. However, there is little or no management of this area for conservation, and the habitat is
Order, Family: Anura, Leptodactylidae Selle in Haiti, and the Sierra de Bahoruco in the Dominican Republic, being destroyed. Improved management of these protected areas, and maintenance of the existing habitat is urgently
Country Distribution: Dominican Republic, Haiti in Hispaniola. It has been recorded from 803-2,100m asl. needed, while further survey work is required to determine the current population status of the species.
Current Population Trend: Decreasing Population Known to be moderately common in suitable habitat, it Bibliography: Hedges, S.B. (1993), Hedges, S.B. (1999), Hedges, S.B. (2001), Henderson, R.W. and Powell, R (1999), Henderson, R.W.
was last collected in 1990 in the Dominican Republic, although there and Powell, R. (2001), Schwartz, A. (1983a), Schwartz, A. and Henderson, R.W. (1991)
have been no surveys of the region since then. Data Providers: Blair Hedges, Sixto Inchaustegui, Robert Powell
Habitat and Ecology It occurs in upland mesic pine forests, usually
under rocks and logs. It has been recorded from recently disturbed
forest, although this is probably not suitable habitat. Eggs are laid
on the ground, and it breeds by direct development.
Major Threats Severe habitat destruction, as a result of charcoal-
ing and slash-and-burn agriculture, is the primary threat to this
species, and is taking place even in the protected areas of La Selle
and Bahoruco.
Conservation Measures This species occurs in the Parque Nacional
Sierra de Bahoruco in the Dominican Republic and the Parc National

CR Eleutherodactylus fuscus Lynn and Dent, 1943

Critically Endangered B2ab(iii) Geographic Range This species has a restricted range in western Jamaica, and has been recorded from 120-680m
Order, Family: Anura, Leptodactylidae asl.
Country Distribution: Jamaica Population It can be locally abundant within its restricted range, but it has a very fragmented and restricted distribu-
Current Population Trend: Decreasing tion. It has not been recorded for many decades in the eastern part of the range.
Habitat and Ecology This is a terrestrial species found in mesic closed-canopy forests, where it calls from bare
ground or rocks. Eggs are laid on the ground, and it breeds by direct development.
Major Threats The primary threat to the species is habitat loss and degradation, in particular due to agriculture,
selective logging, and infrastructure development for ecotourism. Invasive bamboo is also becoming a problem in
© S. Blair Hedges, Penn State

regenerating areas.
Conservation Measures There are at least two forest reserves in the area where this species occurs, as well as
a proposed National Park (Dolphin Head National Park). Ensuring the gazettment of this new protected area, and
maintenance of the existing habitat is essential, and further survey work is necessary to determine the current
population status of the species.
Bibliography: Hedges, S.B. (1993), Hedges, S.B. (1999), Hedges, S.B. (2001), Henderson, R.W. and Powell, R (1999), Henderson, R.W.
and Powell, R. (2001), Koenig, S.E. (2001), Schwartz, A. and Henderson, R.W. (1991)
Data Providers: Blair Hedges, Susan Koenig, Byron Wilson
340 Threatened Amphibians of the World

EN Eleutherodactylus gentryi Lynch and Duellman, 1997

Endangered B1ab(iii) Geographic Range This species occurs at elevations of 2,850-3,380m asl in a small part of the Ecuadorian Andes,
Order, Family: Anura, Leptodactylidae west of the Páramo de Apagua, in the province of Cotopaxi.
Country Distribution: Ecuador Population It is a common species within its small known range.
Current Population Trend: Decreasing Habitat and Ecology It is a high-altitude cloud forest species, where individuals have been found under rocks by day.
It has apparently also been found in high-altitude bush land and grassland. Its ability to adapt to modified habitats is
unknown. It is presumed to breed by direct development, but the site of egg deposition is not known.
Major Threats Habitat destruction and degradation are taking place due to deforestation for agricultural develop-
ment and human settlement.
Conservation Measures It is not known from any protected areas, and there is therefore a need for urgent habitat
protection in the range of this species.
Bibliography: Lynch, J.D. and Duellman, W.E. (1997)
© John D. Lynch

Data Providers: John Lynch, Luis A. Coloma, Santiago Ron

EN Eleutherodactylus ginesi Rivero, 1964

Endangered B1ab(v) Geographic Range This species has a very restricted distribution near and within the Páramo de Mucubají, in the
Order, Family: Anura, Leptodactylidae Venezuelan Andes, in the state of Mérida. It has been recorded from 2,800-4,100m asl. Reports from the tepui region
© César Luis Barrio-Amorós / Fundación AndígenA

Country Distribution: Venezuela in Bolívar and Amazonas States are in error.

Current Population Trend: Decreasing Population It is rare and in decline. Populations of this frog are undergoing declines in areas where it was previously
abundant, such as the Páramo de Mucubají, although it is still common in less surveyed areas.
Habitat and Ecology It is a terrestrial species found in páramo grassland. It breeds by direct development.
Major Threats Although most of the populations are in protected areas, agriculture (involving both crops and live-
stock), as well as fires, are threats to the species’ habitat. However, the cause of the current declines is not known,
and chytridiomycosis cannot be ruled out.
Conservation Measures Part of the species’ range is protected within Parque Nacional Sierra Nevada. Further
research is needed to establish the reasons for the decline of this species, even within protected areas; if chytridio-
mycosis is proved to be a genuine threat, then a captive assurance colony may need to be established.
Bibliography: Barrio Amorós, C.L. (2001), Barrio Amorós, C.L. (2004), La Marca, E. (1992), Péfaur, J.E. and Rivero, J.A. (2000), Rivero,
J.A. (1964), Vial, J.L. and Saylor, L. (1993)
Data Providers: Enrique La Marca, Juan Elías García-Pérez

EN Eleutherodactylus gladiator Lynch, 1976

Endangered B1ab(iii) Geographic Range This species is known from two localities: Papallacta Valley near Cuyuja, in Napo Province,
Order, Family: Anura, Leptodactylidae and Monte Ollivo, in Imbabura Province, Ecuador, from 2,350-2,910m asl, on the Amazonian slopes of the Andes. It
Country Distribution: Ecuador probably occurs more widely but this requires confirmation.
© Natural History Museum and Biodiversity Research

Current Population Trend: Decreasing Population This species was rare at Monte Ollivo, where it was last collected in the year 2000.
Habitat and Ecology This is a fossorial species of páramo grassland, cloud forest, and clearings (Lynch and Duellman
1980). It breeds by direct development.
Major Threats Habitat loss and degradation, due to agriculture development and agricultural pollution, are the
major threats.
Conservation Measures It is not known to occur in any protected areas, making the protection of habitat within
Center, University of Kansas

its range an urgent priority.

Bibliography: Frolich, L.M. et al. (2003), Lynch, J.D. (1976c), Lynch, J.D. and Duellman, W.E. (1980), Lynch, J.D. and Duellman, W.E.
(1997)
Data Providers: Luis A. Coloma, Santiago Ron, Diego Almeida, Fernando Nogales

EN Eleutherodactylus glamyrus Estrada and Hedges, 1997

Endangered B1ab(iii) Geographic Range This species has a restricted range on the Sierra del Turquino in Cuba, from 800-1,974m asl.
Order, Family: Anura, Leptodactylidae Population It is common in suitable habitat.
Country Distribution: Cuba Habitat and Ecology It is arboreal and requires closed mesic cloud forests. It breeds by direct development, and
Current Population Trend: Decreasing the eggs are laid on the ground.
Major Threats The major threat to the species is habitat loss and degradation, as a result of deforestation due to
agriculture, woodcutting, disturbance from tourist activities, and infrastructure development for human settlement.
Although it may occur in the core of the Sierra Maestra, it is restricted to high-elevation cloud forest, which is
declining rapidly.
Conservation Measures It occurs in Parque Nacional Turquino and Parque Nacional La Bayamesa, but there is no
management of these areas for conservation, and the habitat continues to be destroyed. Improved management and
protection of these areas is necessary.
Bibliography: Centro Nacional de Areas Protegidas (CNAP) (2002), Estrada, A.R. and Hedges, S.B. (1997b), Hedges, S.B. (1993),
© Luis M. Diaz

Hedges, S.B. (1999), Hedges, S.B. (2001), Henderson, R.W. and Powell, R (1999), Henderson, R.W. and Powell, R. (2001), Schwartz, A.
and Henderson, R.W. (1991)
Data Providers: Blair Hedges, Luis Díaz
 Globally Threatened Amphibian Species 341

CR Eleutherodactylus glandulifer Cochran, 1935

Critically Endangered A3c Geographic Range This species is restricted to the Massif de la Hotte, Haiti, at an altitude of 300-1,886m asl.
Order, Family: Anura, Leptodactylidae Population At the time of the last survey in 1991 it was known to be moderately common.
Country Distribution: Haiti Habitat and Ecology It is a terrestrial species, found in closed-canopy forests, and appears to be associated with
Current Population Trend: Decreasing streamside habitats. Eggs are laid on the ground and it breeds by direct development.
Major Threats Severe habitat destruction is taking place in its range, primarily due to logging by local people
(charcoaling) and slash-and-burn agriculture.
Conservation Measures It is known to occur in the Parc National Macaya, but there is no management of this area
for conservation, and the habitat continues to be destroyed. Urgent site-based action is required in the Massif de

© S. Blair Hedges, Penn State

la Hotte to conserve the remaining habitat in the area, in order to ensure the persistence of this species as well as
other threatened amphibians known only from this area. Survey work is also necessary to determine the population
status of this species.
Bibliography: Hedges, S.B. (1993), Hedges, S.B. (1999), Hedges, S.B. (2001), Henderson, R.W. and Powell, R (1999), Henderson, R.W.
and Powell, R. (2001), Schwartz, A. (1983b), Schwartz, A. and Henderson, R.W. (1991)
Data Providers: Blair Hedges, Richard Thomas

CR Eleutherodactylus glanduliferoides Shreve, 1936

Critically Endangered B1ab(iii)+2ab(iii) Geographic Range This species is restricted to one small locality in the Massif de la Selle, in Haiti, where it has
Order, Family: Anura, Leptodactylidae been recorded from 1,515-2,121m asl.
Country Distribution: Haiti Population It is very rare, and possibly already extinct. It has been seen only a few times, most recently in 1985
Current Population Trend: Decreasing although there have been few herpetological surveys in the region since.
Habitat and Ecology It occurs only at high elevations in moist forest, calling from the ground or from low grasses
in the forest. Eggs are laid on the ground, and it breeds by direct development.
Major Threats The primary threat to the species is severe habitat destruction, due to charcoaling and slash-and-
burn agriculture.
© S. Blair Hedges, Penn State

Conservation Measures The known range of the species is just outside Parc National Morne La Visite, although
there is no management of this area for conservation, and the habitat within the park is being destroyed. This is
clearly a species requiring urgent site-based conservation action specifically to conserve the remaining habitat in the
area. Survey work is also needed to determine the current population status of this species.
Bibliography: Hedges, S.B. (1993), Hedges, S.B. (1999), Hedges, S.B. (2001), Henderson, R.W. and Powell, R (1999), Henderson, R.W.
and Powell, R. (2001), Schwartz, A. (1983c), Schwartz, A. and Henderson, R.W. (1991)
Data Providers: Blair Hedges, Richard Thomas, Robert Powell

EN Eleutherodactylus glandulosus (Boulenger, 1880)

Endangered B1ab(iii) Geographic Range This species has been recorded from five localities on the eastern slopes of the Andes in the
Order, Family: Anura, Leptodactylidae upper Río Papallacta Valley, in Napo Province, and Playon de San Francisco, in Sucumbios Province, Ecuador. It has
Country Distribution: Ecuador been recorded from 2,105-2,890m asl.
© Natural History Museum and Biodiversity Research

Current Population Trend: Decreasing Population The species is rare at Playon de San Francisco, where one individual was collected in the year 2000.
Habitat and Ecology This is an inhabitant of páramo, cloud forest, and clearings by day, and at night it has been
found on rocks, on the ground and on low vegetation (Lynch and Duellman 1980). It breeds by direct development.
Major Threats The major threats to the species are habitat loss and degradation, due to livestock farming, fires
and agricultural pollution.
Conservation Measures Its range includes the Reserva Ecológica Antisana. Increased protection of the habitat of
Center, University of Kansas

this species in the Ecuadorian Andes is necessary.

Bibliography: Boulenger, G.A. (1880), Frolich, L.M. et al. (2003), Lynch, J.D. and Duellman, W.E. (1980), Lynch, J.D. and Duellman,
W.E. (1997)
Data Providers: Luis A. Coloma, Santiago Ron, Diego Almeida, Fernando Nogales

EN Eleutherodactylus glaphycompus Schwartz, 1973

Endangered B1ab(iii) Geographic Range This species is restricted to the Tiburon Peninsula, Haiti, at an altitudinal range of 576-1,480m
Order, Family: Anura, Leptodactylidae asl.
Country Distribution: Haiti Population It is moderately common in suitable habitat.
Current Population Trend: Decreasing Habitat and Ecology It is found in crevices of exposed limestone in closed moist forest, and calls from low vegeta-
tion. Eggs are laid on the ground and it breeds by direct development.
Major Threats This species is sensitive to habitat destruction (which is ongoing), although perhaps less so than other
frog species. Logging (charcoal collection) by locals and slash and burn agriculture are major threats.
Conservation Measures It occurs in Parc National Macaya, but there is no management of this area for conserva-
tion, and the habitat continues to be destroyed. There is a need for urgent site-based conservation action to ensure
© Eladio Fernandez 2006

the formal protection of remaining habitat on the Tiburon Peninsula and thereby the persistence of this and several
other species of threatened amphibians in the region.
Bibliography: Hedges, S.B. (1991), Hedges, S.B. (1993), Hedges, S.B. (1999), Hedges, S.B. (2001), Henderson, R.W. and Powell, R (1999),
Henderson, R.W. and Powell, R. (2001), Schwartz, A. (1973), Schwartz, A. (1983d), Schwartz, A. and Henderson, R.W. (1991)
Data Providers: Blair Hedges, Richard Thomas
342
VU Eleutherodactylus goini Schwartz, 1960
Vulnerable B1ab(iii)
Order, Family: Anura, Leptodactylidae
Country Distribution: Cuba
Current Population Trend: Decreasing
EN Eleutherodactylus grabhami Dunn, 1926
Endangered B1ab(iii)
Order, Family: Anura, Leptodactylidae
Country Distribution: Jamaica
Current Population Trend: Decreasing
VU Eleutherodactylus gracilis Lynch, 1986
Vulnerable B1ab(iii)
Order, Family: Anura, Leptodactylidae
Country Distribution: Colombia
Current Population Trend: Decreasing
© John D. Lynch
EN Eleutherodactylus grahami Schwartz, 1979
Endangered B1ab(iii)
Order, Family: Anura, Leptodactylidae
Country Distribution: Haiti
Current Population Trend: Decreasing
Threatened Amphibians of the World
Geographic Range This species is restricted to Pinar del Río Hedges, S.B. (1999), Hedges, S.B. (2001), Henderson, R.W. and Powell, R (1999), Henderson, R.W. and Powell, R. (2001), Schwartz, A.
Province, in western Cuba. It has been recorded from sea level up (1960a), Schwartz, A. and Henderson, R.W. (1991)
to at least 700m asl. Data Providers: Blair Hedges, Luis Díaz
Population It is common and abundant.
Habitat and Ecology It is found in rocky habitats and in the leaf-lit-
ter of moist forest, but has not been recorded outside forest habitat.
It breeds by direct development.
Major Threats Major threats include habitat loss due to agriculture
and infrastructure development for human settlement and tourism,
and agricultural pollution.
Conservation Measures It is known from several protected areas,
including Parque Nacional Viñales, but most are in need of improved
management and protection.
Notes on taxonomy: This species was elevated from subspecies status under
Eleutherodactylus planirostris by Estrada and Hedges (1997a).
Bibliography: Estrada, A.R. and Hedges, S.B. (1997a), Hedges, S.B. (1993),
Geographic Range This species has a restricted range in western Bibliography: Hedges, S.B. (1993), Hedges, S.B. (1999), Hedges, S.B. (2001), Henderson, R.W. and Powell, R (1999), Henderson, R.W.
and central Jamaica, with an altitudinal range of 150-670m asl. and Powell, R. (2001), Schwartz, A. and Henderson, R.W. (1991)
Much of the mapped range includes many historical localities, and Data Providers: Blair Hedges, Susan Koenig, Byron Wilson
it is unclear whether the species still occurs throughout this range.
Population It is very rare, with only a few records in the last
decade.
Habitat and Ecology It is found in terrestrial and arboreal bromeli-
ads or on rocks in wet limestone forests, and appears to be intolerant
of any disturbance to its habitat. Eggs are laid on the ground and it
breeds by direct development.
Major Threats It is threatened by habitat degradation and de-
forestation due to agriculture, human settlements, logging, and
bauxite mining.
Conservation Measures Although recorded from several forest
reserves, these do not guarantee the species’ long-term protection,
and there is a need to strengthen and improve the management of
these reserves for biodiversity.
Geographic Range This species is known from the eastern slopes of the Cordillera Occidental in the departments of
Valle del Cauca, Risaralda and Antioquia, and from the western slopes north of the Cordillera Central in the depart-
ments of Caldas and Antioquia, Colombia. It has been recorded from 1,680-2,320m asl.
Population It is a common species.
Habitat and Ecology It occurs mostly on leaves or branches along streams in cloud forest. It can also survive in
secondary forests, but not in open areas outside forest. It breeds by direct development.
Major Threats Major threats included habitat loss due to agriculture (both crops and livestock), and pollution as a
result of the fumigation of crops. Some other species of Eleutherodactylus that are associated with streams at high-
elevations have undergone dramatic declines and disappearances, possibly due to chytridiomycosis, so the status of
this species should be monitored carefully.
Conservation Measures Its range includes Parque Nacional Natural Tatama, Parque Nacional Natural Farallones
de Cali and Regional Reserve Ucumari.
Bibliography: Acosta-Galvis, A.R. (2000), Lynch, J.D. (1986a), Lynch, J.D. (1998b), Lynch, J.D. (1999), Lynch, J.D. and Duellman, W.E.
(1997), Lynch, J.D., Ruiz-Carranza, P.M. and Ardila-Robayo, M.C. (1997), Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and Lynch, J.D. (1996),
Ruiz-Carranza, P.M., Lynch, J.D. and Ardila-Robayo, M.C. (1997)
Data Providers: Martha Patricia Ramírez Pinilla, Mariela Osorno-Muñoz, Jose Vicente Rueda, Adolfo Amézquita, María Cristina
Ardila-Robayo
Geographic Range This species is restricted to the north-west of Bibliography: Hedges, S.B. (1993), Hedges, S.B. (1999), Hedges, S.B. (2001), Henderson, R.W. and Powell, R (1999), Henderson, R.W.
Haiti, ranging from 20-330m asl. and Powell, R. (2001), Schwartz, A. (1979a), Schwartz, A. and Henderson, R.W. (1991)
Population It is relatively common in suitable habitats. Data Providers: Blair Hedges, Richard Thomas
Habitat and Ecology It is found on limestone ridges with boulders
and xerophytic vegetation. Eggs are laid on the ground and it breeds
by direct development.
Major Threats Extremely severe habitat destruction due to charcoal-
ing and small-scale agriculture is taking place on the north-western
peninsula of Haiti, which now looks like a lunar landscape, devoid
of vegetation, although some pockets remain where this species
might occur.
Conservation Measures It is not known to occur in any protected
areas. There is an urgent need for effective protection of remaining
suitable habitat in the range of this species.
 Globally Threatened Amphibian Species 343

CR Eleutherodactylus grandis (Dixon, 1957)

Critically Endangered B1ab(iii)+2ab(iii) Geographic Range This species is known from a single locality (El which a recovery program and strict protection of the species in the Pedregal Reserve and surroundings areas might
Order, Family: Anura, Leptodactylidae Pedregal Reserve) in the Southern Distrito Federal, central Mexico, in need to be implemented. It is protected by Mexican law under the “Special Protection” category (Pr).
Country Distribution: Mexico the lava fields of Volcán Xitle, in the southern part of Mexico City. Notes on taxonomy: The taxonomic status of this species is uncertain.
Current Population Trend: Decreasing Population This is not a very common species and it is difficult Bibliography: Dixon, J.R. (1957)
to collect. Data Providers: Georgina Santos-Barrera, Oscar Flores-Villela
Habitat and Ecology It inhabits dry shrubland, and breeds by
direct development.
Major Threats The extension of Mexico City has resulted in the
disappearance of most of the available suitable habitat for this
species, with the result that it is now believed to be restricted to a
very small protected area, the Pedregal Reserve. The Pedregal has
been subject to long-term disturbance (fragmentation and isolation),
and is now little more than an isolated vegetation fragment of 160
ha with no natural corridors.
Conservation Measures Survey work is urgently needed to es-
tablish the current population status of this species, subsequent to

EN Eleutherodactylus greyi Dunn, 1926

Endangered B1ab(iii) Geographic Range This species has a restricted range in central Bibliography: Hedges, S.B. (1993), Hedges, S.B. (1999), Hedges, S.B. (2001), Henderson, R.W. and Powell, R (1999), Henderson, R.W.
Order, Family: Anura, Leptodactylidae Cuba, where it has been recorded from sea level up to 820m asl. and Powell, R. (2001), Schwartz, A. and Henderson, R.W. (1991)
Country Distribution: Cuba Population It is common in suitable habitat, but is patchily Data Providers: Blair Hedges, Luis Díaz
Current Population Trend: Decreasing distributed.
Habitat and Ecology It is associated with rocky areas in mesic,
closed-canopy forests. It is restricted to these forest habitats and
breeds by direct development.
Major Threats The major threat to this species is habitat loss
and degradation, as a result of deforestation for agriculture, the
development of tourism infrastructure, and the disturbance of habitat
by tourist activities.
Conservation Measures It occurs in a few protected areas, but
these are not well managed for biodiversity conservation and the
habitat within them continues to be degraded. The existing pro-
tected areas network is in need of more effective and strengthened
management.

CR Eleutherodactylus griphus Crombie, 1986

Critically Endangered B2ab(iii) Geographic Range This species is known presently from only three localities in Cockpit Country, central-western
Order, Family: Anura, Leptodactylidae Jamaica. It has been recorded from 250-640m asl.
Country Distribution: Jamaica Population It is a rare species.
Current Population Trend: Decreasing Habitat and Ecology It is terrestrial, and is found only in pristine closed-canopy forests, and does not tolerate
habitat disturbance. Eggs are laid on the ground, and it breeds by direct development.
Major Threats The primary threat to the species is habitat destruction, due to agriculture development, expansion of
human settlements, cutting of timber for firewood, and selective logging. Improved access to previously inaccesible
parts of the interior by means of the building of roads has led to increased deforestation and selective logging.
© S. Blair Hedges, Penn State

Conservation Measures The majority of the Cockpit Country is government-owned Forest Reserve, although it has
not yet been formally declared a protected area. Maintenance of the existing habitat is urgently required.
Bibliography: Crombie, R.I. (1986), Hedges, S.B. (1993), Hedges, S.B. (1999), Hedges, S.B. (2001), Henderson, R.W. and Powell, R (1999),
Henderson, R.W. and Powell, R. (2001), Schwartz, A. and Henderson, R.W. (1991)
Data Providers: Blair Hedges

EN Eleutherodactylus gryllus Schmidt, 1920 GREEN COQUI

Endangered B1ab(v) Geographic Range This species has a restricted range in the interior uplands in Puerto Rico, having been recorded
Order, Family: Anura, Leptodactylidae from 300-1,182m asl.
Country Distribution: Puerto Rico Population It is known to have undergone recent population declines, and is now known only from cloud forest
Current Population Trend: Decreasing above 700m asl.
Habitat and Ecology It occurs in forest and along forest edges or openings, retreating by day into bromeliads or
under moss and rocks. It has not been recorded outside forest habitats. Eggs are laid in bromeliads, and it breeds
by direct development.
Major Threats There is a high abundance of introduced predators such as Rattus rattus, even in the protected areas
on El Yunque, which may pose a threat. It is also possible that chytridiomycosis, in combination with climate change,
may be a threat to this species.
© Alejandro Sanchez

Conservation Measures It occurs in a few protected areas, which are well managed. However, there is a need for
the control of invasive predators within these protected areas, and the species requires careful population monitoring,
particularly in light of the potential threat posed by chytridiomycosis.
Bibliography: Burrowes, P.A., Joglar, R.L. and Green, D.E. (2004), Hedges, S.B. (1993), Henderson, R.W. and Powell, R (1999), Henderson,
R.W. and Powell, R. (2001), Joglar, R.L. (1999), Joglar, R.L. and Burrowes, P.A. (1996), Rivero, J.A. (1998)
Data Providers: Blair Hedges, Rafael Joglar, Luis J. Villanueva-Rivera, Neftalí Ríos-Lopez
344 Threatened Amphibians of the World

EN Eleutherodactylus guanahacabibes Estrada and Rodriguez, 1985

Endangered B1ab(iii) Geographic Range This species has a restricted range in the Bibliography: Estrada, A.R. and Rodríguez, J.N. (1985b), Hedges, S.B. (1993), Hedges, S.B. (1999), Hedges, S.B. (2001), Henderson, R.W.
Order, Family: Anura, Leptodactylidae Peninsula de Guanahacabibes, in extreme western Cuba. It has been and Powell, R (1999), Henderson, R.W. and Powell, R. (2001), Schwartz, A. and Henderson, R.W. (1991)
Country Distribution: Cuba recorded from sea level up to 20m asl. Data Providers: Blair Hedges, Luis Díaz
Current Population Trend: Decreasing Population It is moderately common in suitable habitat.
Habitat and Ecology It occurs in rocky areas and caves with high
humidity in moist forest areas; it has not been recorded outside
forest habitat. Eggs are laid on the ground, and it breeds by direct
development.
Major Threats The major threat to this species is habitat loss and
degradation due to infrastructure development; a road has recently
been built directly through the habitat of the species, and the penin-
sula may become a landing point for cruise ships in the future.
Conservation Measures It occurs in Parque Nacional Guana-
hacabibes, which is a well-protected area in Cuba. Protection of
other tracts of suitable forest habitat where the species occurs is
recommended.

VU Eleutherodactylus guantanamera Hedges, Estrada and Thomas, 1992

Vulnerable B1ab(iii) Geographic Range This species has a restricted range in the upland
Order, Family: Anura, Leptodactylidae areas (60-1,150m asl) of south-eastern Cuba.
Country Distribution: Cuba Population This species is moderately common in suitable
Current Population Trend: Decreasing habitat.
Habitat and Ecology It is arboreal in closed moist forest. Eggs
are laid in bromeliads and breeding takes place by direct develop-
ment.
Major Threats The main threat is habitat loss and degradation
due to agriculture, woodcutting, disturbance from tourists, and
human settlement.
Conservation Measures It occurs in several protected areas, but
these are in need of improved protection and management.
Bibliography: Centro Nacional de Areas Protegidas (CNAP) (2002), Hedges,
S.B. (1993), Hedges, S.B. (1999), Hedges, S.B. (2001), Hedges, S.B., Estrada, A.R.
and Thomas, R. (1992), Henderson, R.W. and Powell, R (1999), Henderson, R.W.
and Powell, R. (2001), Schwartz, A. and Henderson, R.W. (1991)
Data Providers: Blair Hedges, Luis Díaz

EN Eleutherodactylus gundlachi Schmidt, 1920

Endangered B1ab(iii) Geographic Range This species has a restricted range in the Sierra Bibliography: Centro Nacional de Areas Protegidas (CNAP) (2002), Hedges, S.B. (1993), Hedges, S.B. (1999), Hedges, S.B. (2001),
Order, Family: Anura, Leptodactylidae Maestra, the Sierra del Cobre, and the Sierra de la Gran Piedra, Cuba. Henderson, R.W. and Powell, R (1999), Henderson, R.W. and Powell, R. (2001), Schwartz, A. and Henderson, R.W. (1991)
Country Distribution: Cuba The altitudinal range is 650-1,375m asl. Data Providers: Blair Hedges, Luis Díaz
Current Population Trend: Decreasing Population It is common in suitable habitat.
Habitat and Ecology It is terrestrial and requires closed mesic
forest. The species calls from the ground, and breeds by direct
development.
Major Threats The major threat is habitat destruction and loss due
to agriculture, woodcutting, disturbance from tourist activities, and
infrastructure development for human settlement.
Conservation Measures Although occurring in several protected
areas, several of these are not well managed for biodiversity con-
servation and habitat loss is proceeding within the boundaries of
these reserves. There is a clear need to improve and strengthen the
management of these existing protected areas.

EN Eleutherodactylus haitianus Barbour, 1942

Endangered B1ab(iii)+2ab(iii) Geographic Range This species has a restricted range in the Cordillera Central, Dominican Republic, where it has
Order, Family: Anura, Leptodactylidae been recorded from 1,545-2,455m asl.
Country Distribution: Dominican Republic Population It can be locally common, but it is patchily distributed.
Current Population Trend: Decreasing
© Marcelino Hernández / Miguel A. Landestoy

Habitat and Ecology It is terrestrial in high-elevation pine forests, and has not been recorded outside forested
regions. It calls from low vegetation, the eggs are laid on the ground, and it breeds by direct development.
Major Threats Disturbance from ecotourism is a major threat, as is habitat destruction due to agricultural activi-
ties.
Conservation Measures Its range includes several protected areas, although these are often not well managed
and habitat disturbance is ongoing within the parks’ limits. Improved and strengthened management of these existing
protected areas is recommended.
Bibliography: Hedges, S.B. (1993), Hedges, S.B. (1999), Hedges, S.B. (2001), Henderson, R.W. and Powell, R (1999), Henderson, R.W.
and Powell, R. (2001), Schwartz, A. (1983e), Schwartz, A. and Henderson, R.W. (1991)
Data Providers: Blair Hedges, Sixto Inchaustegui, Robert Powell

CR Eleutherodactylus hamiotae Flores, 1993
Critically Endangered B1ab(iii)
Order, Family: Anura, Leptodactylidae
Country Distribution: Ecuador
Current Population Trend: Decreasing
EN Eleutherodactylus hedricki Rivero, 1963
Endangered B1ab(v)
Order, Family: Anura, Leptodactylidae
Country Distribution: Puerto Rico
Current Population Trend: Decreasing
© Alejandro Sanchez
CR Eleutherodactylus helonotus (Lynch, 1975)
Critically Endangered B2ab(iii)
Order, Family: Anura, Leptodactylidae
Country Distribution: Ecuador
Current Population Trend: Decreasing
EN Eleutherodactylus helvolus Lynch and Rueda-Almonacid, 1998
Endangered B1ab(iii)
Order, Family: Anura, Leptodactylidae
Country Distribution: Colombia
Current Population Trend: Decreasing
© John D. Lynch
Globally Threatened Amphibian Species 345
Geographic Range This species is known only from a very small population status of this species.
area north-west of Nono at 2,140m asl on the western slope of the Bibliography: Flores, G. (1993), Lynch, J.D. and Duellman, W.E. (1997)
Andes in Pichincha Province, Ecuador. Data Providers: John Lynch, Luis A. Coloma, Santiago Ron
Population It is a very rare species, and only five specimens have
ever been collected.
Habitat and Ecology It is found in cloud forest, where it is a highly
specialized species, living on wet rock faces. It is unlikely to be very
adaptable to modified habitats in view of its specialized microhabitat
requirements. It is presumed to breed by direct development, but the
site of egg deposition is not known.
Major Threats The primary threat to the species is deforestation
as a result of agricultural development (crops and livestock), and
human settlement.
Conservation Measures This species is not known from any
protected areas, and maintenance of the existing habitat is urgently
needed. Further survey work is required to determine the biology and
TREEHOLE COQUI
Geographic Range This species has a patchy distribution in the interior uplands of Puerto Rico, and has an altitudinal
range of 455-1,152m asl.
Population There are indications that populations have declined over the last three decades, even in suitable
habitat (R. Joglar pers. comm.).
Habitat and Ecology It is arboreal in mesic broadleaf forests. Males call from tree trunks and the canopy, eggs are
laid in tree trunk cavities, and breeding is by direct development.
Major Threats The major threat to this species is chytridiomycosis (linked to climate change), which is suspected
to be causing the observed declines in this species.
Conservation Measures It occurs in several protected areas. In view of the threat of chytridiomycosis, ex-situ
populations may need to be established.
Bibliography: Burrowes, P.A., Joglar, R.L. and Green, D.E. (2004), Hedges, S.B. (1993), Henderson, R.W. and Powell, R (1999), Henderson,
R.W. and Powell, R. (2001), Joglar, R.L. (1999), Joglar, R.L. and Burrowes, P.A. (1996), Rivero, J.A. (1963a), Rivero, J.A. (1998)
Data Providers: Blair Hedges, Rafael Joglar, Richard Thomas, Luis J. Villanueva-Rivera, Neftalí Ríos-Lopez
Geographic Range This species is known only from two localities Bibliography: Lynch, J.D. (1975b), Lynch, J.D. (1981c), Lynch, J.D. and Duellman, W.E. (1997)
in the vicinity of Río Pitzara, Pichincha Province, on the Pacific slopes Data Providers: John Lynch, Luis A. Coloma, Santiago Ron
of the Andes in Ecuador, at 1,000-2,000m asl.
Population It is a very rare species.
Habitat and Ecology The species is terrestrial, restricted to primary
cloud forest, and cannot tolerate any disturbance of its habitat. It
is presumed to breed by direct development, but the site of egg
deposition is not known.
Major Threats The major threat to the species is habitat loss
due to agricultural development (crops and livestock), logging, and
human settlement.
Conservation Measures This species is not known from any
protected areas, and maintenance of the existing habitat is urgently
needed. Further survey work is required to determine the biology and
population status of this species.
Geographic Range This species is known from cloud forest in the municipalities of Pensilvania and Samaná, in the
department of Caldas, and from the municipalities of Guatepé and Anorí, in Antioquia Department, on the eastern
flank of the Cordillera Central, in Colombia. It has been recorded from 1,800-2,000m asl.
Population It is not a common species.
Habitat and Ecology It is restricted to primary forest and breeds by direct development and is not associated with
streams. It is very small (at most 23mm long) and is very sensitive to any variation in humidity. It is not tolerant of
any habitat disturbance.
Major Threats The major threat to this species is habitat loss caused by agricultural development (including the
planting of illegal crops).
Conservation Measures Its range does not include any protected areas; given the species’ sensitivity to any habitat
disturbance, protection of remaining primary forest habitat within the range of the species is essential.
Bibliography: Acosta-Galvis, A.R. (2000), Lynch, J.D. and Rueda-Almonacid, J.V. (1998b), Paez, V.P. et al. (2002), Rueda-Almonacid,
J.V. (2000)
Data Providers: Fernando Castro, Maria Isabel Herrera, John Lynch
346
EN Eleutherodactylus heminota Shreve and Williams, 1963
Endangered B2ab(iii)
Order, Family: Anura, Leptodactylidae
Country Distribution: Dominican Republic, Haiti
Current Population Trend: Decreasing
EN Eleutherodactylus hernandezi Lynch and Ruíz-Carranza, 1983
Endangered B1ab(iii)
Order, Family: Anura, Leptodactylidae
Country Distribution: Colombia
Current Population Trend: Decreasing
© John D. Lynch
VU Eleutherodactylus hybotragus Lynch, 1992
Vulnerable B1ab(iii)
Order, Family: Anura, Leptodactylidae
Country Distribution: Colombia
Current Population Trend: Decreasing
© John D. Lynch
EN Eleutherodactylus hypostenor Schwartz, 1965
Endangered B1ab(iii)
Order, Family: Anura, Leptodactylidae
Country Distribution: Dominican Republic, Haiti
Current Population Trend: Decreasing
Threatened Amphibians of the World
Geographic Range This species is restricted to the Tiburon Penin-
sula and eastern Sierra de Baoruca, Hispaniola (Haiti and Dominican
Republic). Its altitudinal range is from sea level up to 1,697m asl.
Population Although this species can be moderately common in
suitable habitat, it is declining and rarely encountered in much
of its range.
Habitat and Ecology This species requires closed-canopy forest
and frequents bromeliads. The eggs are laid in bromeliads and it
breeds by direct development.
Major Threats It is rapidly declining throughout its range due to
severe habitat destruction, even in protected areas. In the Dominican
Republic much habitat destruction has taken place in the last 10
years due to agriculture and cattle grazing. In Haiti, severe habitat
destruction has occurred in the Formon-Macaya region, including
throughout Parc National Macaya, mainly due to logging (charcoal
collection) by locals and slash and burn agriculture.
Geographic Range This species has a restricted range in the
western Massif de la Selle in Haiti and the Sierra de Baoruco in the
Dominican Republic, Hispaniola. Its altitudinal range is from sea
level up to 1,061m asl.
Population It can be locally uncommon and is patchily distrib-
uted.
Habitat and Ecology An inhabitant of closed mesic broadleaf
forests, as well as shade-grown coffee and cacao plantations,
this is a very specialized burrowing species, with males calling
from constructed underground chambers and the eggs being laid
underground.
Major Threats The main threat to this species is habitat loss: the
forest of Baoruco is severely threatened by expanding agriculture and
cattle grazing, even within Parque Nacional Sierra de Baoruco.
Conservation Measures Although it occurs in Parque Nacional
Sierra de Bahoruca and Parque Nacional Sierra Bahoruca Oriental
Conservation Measures It occurs in Parc Naturel Morne La Visite and Parc National Macaya in Haiti, and also
in Parque Nacional Sierra de Bahoruca and Parque Nacional Sierra Bahoruca Oriental in the Dominican Republic.
There is an urgent need for strengthened management and protection of these reserves, particularly on the Tiburon
Peninsula.
Bibliography: Hedges, S.B. (1993), Hedges, S.B. (1999), Hedges, S.B. (2001), Henderson, R.W. and Powell, R (1999), Henderson, R.W.
and Powell, R. (2001), Schwartz, A. (1983f), Schwartz, A. and Henderson, R.W. (1991)
Data Providers: Blair Hedges, Sixto Inchaustegui, Richard Thomas, Robert Powell
Geographic Range This species is known from two localities: one on the Oriental slope from the Cordillera Central
in Huila Department, and the other on the eastern slope of the Cordillera Central in Caquetá Department, Colombia,
from 2,300-2,600m asl.
Population It is an uncommon species, and is known only from six specimens.
Habitat and Ecology This species inhabits intermediate strata in primary forest. As only a few individuals have been
recorded, not much information is known regarding its exact habitat requirements, but it is believed to be extremely
moisture dependent. It breeds by direct development.
Major Threats Habitat loss caused by agricultural development (including the planting of illegal crops) is the main
threat to the species.
Conservation Measures One known locality is within Parque Nacional Natural Purace, and the second locality
will soon be protected within the newly created Parque Nacional Natural Fragua. Further survey work is necessary
to establish the current population status of this species, its ecological requirements, and to determine whether it
occurs outside the two known localities.
Bibliography: Acosta-Galvis, A.R. (2000), Lynch, J.D. and Duellman, W.E. (1997), Lynch, J.D. and Ruiz-Carranza, P.M. (1983), Lynch, J.D.,
Ruiz-Carranza, P.M. and Ardila-Robayo, M.C. (1997), Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and Lynch, J.D. (1996)
Data Providers: Fernando Castro, Maria Isabel Herrera, John Lynch
Geographic Range This species is known from Valle del Cauca department in the Pacific lowlands of Colombia,
from 10-920m asl. It probably occurs more widely than records suggest.
Population It is an uncommon species.
Habitat and Ecology An arboreal species found on middle and low strata on sloping terrain, inside or alongside
small streams wherever there is suitable forest cover. It breeds by direct development.
Major Threats Although this species occurs in an area of low human population density, its habitat remains at
risk from logging.
Conservation Measures The range includes Parque Nacional Natural Farallones de Cali.
Bibliography: Acosta-Galvis, A.R. (2000), Lynch, J.D. (1992c), Lynch, J.D. (1998b), Lynch, J.D. (1999), Lynch, J.D. and Duellman, W.E. (1997),
Lynch, J.D., Ruiz-Carranza, P.M. and Ardila-Robayo, M.C. (1997), Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and Lynch, J.D. (1996)
Data Providers: Fernando Castro, Maria Isabel Herrera, John Lynch
in the Dominican Republic, there is a need to greatly strengthen and improve the management of these protected
areas to ensure more effective protection of the broad-leaf forest habitat within these areas.
Bibliography: Hedges, S.B. (1993), Hedges, S.B. (1999), Hedges, S.B. (2001), Henderson, R.W. and Powell, R (1999), Henderson, R.W.
and Powell, R. (2001), Schwartz, A. (1965b), Schwartz, A. (1984), Schwartz, A. and Henderson, R.W. (1991)
Data Providers: Blair Hedges, Sixto Inchaustegui, Richard Thomas, Robert Powell
 Globally Threatened Amphibian Species 347

CR Eleutherodactylus iberia Estrada and Hedges, 1996

Critically Endangered B1ab(iii) Geographic Range The smallest frog in world, this species occurs only in Holguin Province in eastern Cuba from
Order, Family: Anura, Leptodactylidae sea level up to 600m asl.
Country Distribution: Cuba Population The species is common in suitable habitat.
Current Population Trend: Decreasing Habitat and Ecology It is found in closed rainforest on poorly drained soil, and maintenance of local humidity is
critically important for its survival. It breeds by direct development.
Major Threats This species is threatened mainly by habitat loss as a result of the deforestation taking place from
subsistence farming and wood collecting.
Conservation Measures It occurs in the Parque Nacional Alejandro de Humboldt, but even this protected area
is subject to the effects of habitat loss. Maintenance of the remaining suitable habitat and improved protection of
© Alberto R. Estrada existing protected areas is essential.
Bibliography: Centro Nacional de Areas Protegidas (CNAP) (2002), Estrada, A.R. and Hedges, S.B. (1996b), Hedges, S.B. (1993),
Hedges, S.B. (1999), Hedges, S.B. (2001), Henderson, R.W. and Powell, R (1999), Henderson, R.W. and Powell, R. (2001), Schwartz, A.
and Henderson, R.W. (1991)
Data Providers: Blair Hedges, Luis Díaz

EN Eleutherodactylus ignicolor Lynch and Duellman, 1980

Endangered B1ab(iii) Geographic Range This species is known from only three localities in the upper Río Papallacta Valley, in Napo
Order, Family: Anura, Leptodactylidae Province, Ecuador, from 2,160-2,750m asl. It probably occurs a little more widely.
Country Distribution: Ecuador Population Very little is known of this species’ current population status, but it is a rare species at the Estacion
© Natural History Museum and Biodiversity Research

Current Population Trend: Decreasing Biólogica Yanayacu (where it was collected as recently as 2003).
Habitat and Ecology Individuals have been found on bushes, and in bamboo at the edges of small cascading
streams at night in upper humid montane forest. It has not been recorded from secondary forest. It breeds by direct
development.
Major Threats The major threat to the species is habitat loss due to agricultural activities and logging. Some other
species of Eleutherodactylus that are associated with streams have undergone dramatic declines and disappearances,
Center, University of Kansas

possibly due to chytridiomycosis.

Conservation Measures It is not known from any protected areas in Ecuador, although it has recently been collected
from the Estacion Biólogica Yanayacu near Cusunga. Protection of montane forest habitat in the range of this species
is recommended, and further survey work is needed to monitor the population status of this species, particularly
given the potential of chytrid.
Bibliography: Lynch, J.D. and Duellman, W.E. (1980), Lynch, J.D. and Duellman, W.E. (1997)
Data Providers: Luis A. Coloma, Santiago Ron, Diego Almeida, Fernando Nogales

EN Eleutherodactylus incanus Lynch and Duellman, 1980

Endangered B1ab(iii) Geographic Range This species is known from only three localities on the eastern face of the Andes in Cocodrillos,
Order, Family: Anura, Leptodactylidae Napo Province, Ecuador. It has been recorded from 1,700-2,200m asl.
Country Distribution: Ecuador Population It is a rare species (only three specimens were collected in 2001).
Current Population Trend: Decreasing Habitat and Ecology It is found in well-shaded cloud forest, and most individuals have been recorded perched on
Biodiversity Research Center, University of Kansas

bushes or low trees at night (Lynch and Duellman 1980). It has also been found above small streams. Breeding is
by direct development.
Major Threats The major threat is habitat loss due to agricultural activities, involving both crops and livestock,
as well as logging. Some other species of Eleutherodactylus that are associated with streams have undergone
© Natural History Museum and

dramatic declines and disappearances, possibly due to chytridiomycosis, so the status of this species should be
monitored carefully.
Conservation Measures It has been recorded from the Reserva Ecológica Cayambe-Coca, the Reserva Ecológica
Antisana, and Parque Nacional Sumaco Napo-Galeras.
Bibliography: Lynch, J.D. and Duellman, W.E. (1980), Lynch, J.D. and Duellman, W.E. (1997)
Data Providers: Luis A. Coloma, Santiago Ron, Diego Almeida, Fernando Nogales

VU Eleutherodactylus incomptus Lynch and Duellman, 1980

Vulnerable B1ab(iii) Geographic Range This species has been reported from elevations of 1,370-1,910m asl on the Amazonian slopes of
Order, Family: Anura, Leptodactylidae the Cordillera Oriental in Napo and Sucumbios Provinces, Ecuador, and from about 1,300m asl on the southern slopes
Country Distribution: Ecuador, Peru of the Cordillera del Cóndor in Departmento Cajamarca, Peru (06° 05’S; 78° 43’W). It is likely to occur more widely.
Current Population Trend: Decreasing Population In Ecuador, it is an uncommon species. In Peru, it has only been reported from two specimens (both
Biodiversity Research Center, University of Kansas

sub-adult females).
Habitat and Ecology This species occurs in cloud forest. They are most frequently encountered at night on leaves of
herbaceous vegetation or low bushes in clearings or at the forest edge (Lynch and Duellman 1980). It breeds by direct
development, but the site of egg deposition is not known. It occurs in old and second growth forest.
© Natural History Museum and

Major Threats The major threat is habitat loss due to livestock ranching, agro-industry, and selective logging.
Conservation Measures In Ecuador, its distribution range overlaps with Reserva Ecológica Antisana, and Reserva
Ecológica Cayambe-Coca; it might also overlap with Parque Nacional Llanganatis and Parque Nacional Sumaco
Napo-Galeras. It is not recorded from any protected areas in Peru. Further survey work is needed to determine if the
species is more continuously distributed between records in Ecuador and Peru.
Notes on taxonomy: We follow Lynch and Duellman (1997) in regarding Eleutherodactylus kirklandi as a synonym of E. incomptus.
Bibliography: Duellman, W.E. and Pramuk, J.B. (1999), Flores, G. (1985b), Lynch, J.D. and Duellman, W.E. (1980), Lynch, J.D. and
Duellman, W.E. (1997), Ortiz, A. and Morales, M. (2000)
Data Providers: Lily Rodríguez, Jorge Luis Martinez, Luis A. Coloma, Santiago Ron, Diego Almeida
348 Threatened Amphibians of the World

VU Eleutherodactylus ingeri (Cochran and Goin, 1961)

Vulnerable B1ab(iii) Geographic Range This species has a discontinuous distribution on both slopes of the Cordillera Oriental of Colombia,
Order, Family: Anura, Leptodactylidae where it is found from 1,550-2,350m asl, in the departments of Cundinamarca (Laguna Pedro Palo), Boyacá, Santander,
Country Distribution: Colombia and Norte de Santander. It probably occurs more widely, in particular between known localities.
Current Population Trend: Decreasing Population This is a rare species that was last collected in 1991, but there has been no survey work since.
Habitat and Ecology This species is a cloud forest inhabitant, and occurs in leaf-litter and disturbed areas provided
that small patches of forest remain nearby. Reproduction is by direct development.
Major Threats Habitat loss caused by logging and agricultural development (including planting of illegal crops) is
the main threat.
Conservation Measures It occurs in Santuario de Fauna y Flora Guanentá Alto Río Fonce and in the Parque
Nacional Natural Tamá.
Bibliography: Acosta-Galvis, A.R. (2000), Cochran, D.M. and Goin, C.J. (1970), Lynch, J.D. (1981c), Lynch, J.D. (1997), Lynch, J.D. and
© John D. Lynch

Duellman, W.E. (1997), Lynch, J.D., Ruiz-Carranza, P.M. and Ardila-Robayo, M.C. (1997), Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and
Lynch, J.D. (1996)
Data Providers: Fernando Castro, Maria Isabel Herrera, Jose Vicente Rueda

EN Eleutherodactylus insignitus Ruthven, 1917

Endangered B1ab(iii) Geographic Range This species is known from the northern and north-western flank of the Sierra Nevada de Santa
Order, Family: Anura, Leptodactylidae Marta, in the departments of Magdalena and Guajira, Colombia. It has been recorded from 1,530-2,130m asl.
Country Distribution: Colombia Population It is an uncommon species.
Current Population Trend: Decreasing Habitat and Ecology A terrestrial frog, individuals have been found under logs or rocks on roadsides. When active,
they occur on top of rocks, logs or low vegetation, beside streams. It breeds by direct development.
Major Threats Major threats include habitat loss, due to agricultural activities (involving both crops and livestock),
as well as pollution due to the fumigation of crops.
Conservation Measures Its range includes Parque Nacional Natural Sierra Nevada de Santa Marta, and the
adjacent, recently established El Dorado Nature Reserve.
Bibliography: Acosta-Galvis, A.R. (2000), Cochran, D.M. and Goin, C.J. (1970), Lynch, J.D. and Duellman, W.E. (1997), Lynch, J.D. and
Ruiz-Carranza, P.M. (1985a), Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and Lynch, J.D. (1996), Ruthven, A.G. (1917a)
© John D. Lynch

Data Providers: Martha Patricia Ramírez Pinilla, Mariela Osorno-Muñoz, Jose Vicente Rueda, Adolfo Amézquita, María Cristina
Ardila-Robayo

EN Eleutherodactylus intermedius Barbour and Shreve, 1937

Endangered B1ab(iii) Geographic Range This species has a restricted range in the Schwartz, A. and Henderson, R.W. (1991)
Order, Family: Anura, Leptodactylidae Sierra Maestra and the Sierra del Cobre, Cuba. Its altitudinal range Data Providers: Blair Hedges, Luis Díaz
Country Distribution: Cuba is 454-1,880m asl.
Current Population Trend: Decreasing Population The species is uncommon even in suitable habitat.
Habitat and Ecology It occurs only in moist closed forest, and
breeds by direct development.
Major Threats The major threat is habitat loss, due to agriculture,
woodcutting, disturbance from tourists, and infrastructure develop-
ment for human settlement.
Conservation Measures It occurs in Parque Nacional Turquino and
Parque Nacional La Bayamesa, but there is no management of these
areas for conservation, and the habitat continues to be destroyed
even within the park boundaries. Improved and strengthened man-
agement of these existing protected areas is necessary.
Bibliography: Hedges, S.B. (1993), Hedges, S.B. (1999), Hedges, S.B. (2001),
Henderson, R.W. and Powell, R (1999), Henderson, R.W. and Powell, R. (2001),

VU Eleutherodactylus inusitatus Lynch and Duellman, 1980

Vulnerable B1ab(iii) Geographic Range This species is known from scattered localities along the eastern face of the Andes in Ecuador,
Order, Family: Anura, Leptodactylidae at low to moderate elevations from 1,300-2,160m asl. It is currently known from only five localities, but probably
Country Distribution: Ecuador occurs more widely.
Current Population Trend: Decreasing Population It is fairly common in some areas, such as Estación Biólogica Yanayacu, where it has been seen repeat-
edly from 2000-2003.
Habitat and Ecology Specimens have been found in cloud forest in low vegetation (Lynch and Duellman 1980). It
does not occur outside old growth forest. It breeds by direct development.
Major Threats The main threat is habitat loss due to agriculture (both crops and livestock), logging, and human
© H. Mauricio Ortega-Andrade

settlement.
Conservation Measures Its distribution range overlaps with Parque Nacional Sumaco Napo-Galeras, Reserva
Biólogica Limoncocha, Reserva de Produccion Faunistica Cuyabeno, Reserva Ecológica Cayambe-Coca and Reserva
Ecológica Antisana.
Bibliography: Funk, C.W. et al. (2003), Lynch, J.D. and Duellman, W.E. (1980), Lynch, J.D. and Duellman, W.E. (1997)
Data Providers: Luis A. Coloma, Santiago Ron, Diego Almeida, Fernando Nogales
 Globally Threatened Amphibian Species 349

EN Eleutherodactylus ionthus Schwartz, 1960

Endangered B1ab(iii) Geographic Range This species has a restricted range in the S.B. (1993), Hedges, S.B. (1999), Hedges, S.B. (2001), Henderson, R.W. and Powell, R (1999), Henderson, R.W. and Powell, R. (2001),
Order, Family: Anura, Leptodactylidae Sierra Maestra, Cuba. Its altitudinal range is from sea level up to Schwartz, A. (1960a), Schwartz, A. and Henderson, R.W. (1991)
Country Distribution: Cuba 1,230m asl. Data Providers: Blair Hedges, Luis Díaz
Current Population Trend: Decreasing Population It is moderately common in suitable habitat.
Habitat and Ecology It is arboreal in closed mesic forest, but
appears to be less sensitive to habitat alteration than some other
species in the genus. Breeding is by direct development.
Major Threats The major threat is habitat loss, due to agriculture,
woodcutting, disturbance from tourists, and infrastructure develop-
ment for human settlement.
Conservation Measures It occurs in Parque Nacional Turquino and
Parque Nacional La Bayamesa, but there is no management of these
areas for conservation, and habitat continues to be destroyed even
within the park boundaries. Improved and strengthened management
of these existing protected areas is necessary.
Bibliography: Centro Nacional de Areas Protegidas (CNAP) (2002), Hedges,

EN Eleutherodactylus jamaicensis Barbour, 1910

Endangered B1ab(iii) Geographic Range This species is widely, but very patchily, distrib- issue. There is clearly a need for more effective and strengthened management of the existing protected areas in
Order, Family: Anura, Leptodactylidae uted in the Jamaican interior, ranging from 120-1,290m asl. the region.
Country Distribution: Jamaica Population It is uncommon. Bibliography: Hedges, S.B. (1993), Hedges, S.B. (1999), Hedges, S.B. (2001), Henderson, R.W. and Powell, R (1999), Henderson, R.W.
Current Population Trend: Decreasing Habitat and Ecology It occurs only in lowland and montane moist and Powell, R. (2001), Schwartz, A. and Henderson, R.W. (1991)
old growth forests, and is almost always associated with arboreal Data Providers: Blair Hedges, Susan Koenig, Byron Wilson
or terrestrial bromeliads. Males call from, and eggs are laid in,
bromeliads.
Major Threats Its dependence upon bromeliads in mostly un-
disturbed forest makes it especially susceptible to habitat loss.
Unfortunately, habitat degradation and deforestation is taking place
in much of its range (such as the Blue Mountains) due to agricultural
activities, human settlements, and logging.
Conservation Measures It occurs in the Blue and John Crow
Mountains National Park and several forest reserves; however,
there are coffee plantations even within the park’s limits and habitat
disturbance resulting from tourist activities in the park remains an

CR Eleutherodactylus jasperi Drewry and Jones, 1976 GOLDEN COQUI

Critically Endangered A2ae; B2ab(i,ii,iv,v) Geographic Range This species is found in the Sierra de Cayey, Puerto Rico, where it has been recorded from
Order, Family: Anura, Leptodactylidae 647-785m asl.
Country Distribution: Puerto Rico Population This species has not been seen since 1981, and it is now possibly extinct. Extensive surveys of suitable
Current Population Trend: Decreasing habitat have failed to find any individuals. It has clearly undergone a catastrophic decline.
Habitat and Ecology This species has been recorded from bromeliads (terrestrial and arboreal) in forests and
open and rocky areas. It is known to be a live-bearing species, giving birth to 1-3 young, unique amongst members
of the genus.
Major Threats The primary reason for the decline of this species is thought to be chytridiomycosis in combination
with climate change. Introduced predators might also be a contributing factor.
© Erik Wild, Proyecto Coquí

Conservation Measures The range of this species includes Carite Forest Reserve, which is a well-managed pro-
tected area. Further surveys are needed to relocate this species and determine whether or not it might still survive
in the wild. In view of the severe risk of chytridiomycosis, surviving individuals might need to form the basis for the
establishment of an ex-situ population.
Bibliography: Burrowes, P.A., Joglar, R.L. and Green, D.E. (2004), Drewry, G.E. and Jones, K. (1976), Hedges, S.B. (1993), Henderson,
R.W. and Powell, R (1999), Henderson, R.W. and Powell, R. (2001), Joglar, R.L. (1999), Joglar, R.L. and Burrowes, P.A. (1996), Rivero,
J.A. (1998)
Data Providers: Blair Hedges, Rafael Joglar, Luis J. Villanueva-Rivera, Neftalí Ríos-Lopez

CR Eleutherodactylus jaumei Estrada and Alonso, 1997

Critically Endangered B1ab(iii) Geographic Range This species is known from only a single locality in the Sierra Maestra, Cuba, at an altitudinal
Order, Family: Anura, Leptodactylidae range of 200-950m asl.
Country Distribution: Cuba Population This is a common species in suitable habitat.
Current Population Trend: Decreasing Habitat and Ecology It occurs in closed mesic forest, but also occurs in secondary forests. It is usually found in
leaf-litter and it breeds by direct development.
Major Threats The main threat to this species is habitat destruction due to agricultural expansion, woodcutting,
© Nils Navarro, courtesy of Luis Diaz

disturbance from tourists, and infrastructure development.

Conservation Measures The species does occur in the lower elevations of the Parque Nacional La Bayamesa, but
there is no management of this area for conservation, and the habitat continues to be destroyed. Maintenance of the
remaining suitable habitat and improved protection of existing protected areas is essential.
Bibliography: Estrada, A.R. and Alonso, R. (1997), Hedges, S.B. (1993), Hedges, S.B. (1999), Hedges, S.B. (2001), Henderson, R.W. and
Powell, R (1999), Henderson, R.W. and Powell, R. (2001), Schwartz, A. and Henderson, R.W. (1991)
Data Providers: Blair Hedges, Luis Díaz
350 Threatened Amphibians of the World

EN Eleutherodactylus johannesdei Rivero and Serna, 1988 “1987”

Endangered B1ab(iii) Geographic Range This species is known from Antioquia to Ri- Bibliography: Acosta-Galvis, A.R. (2000), Lynch, J.D. (1998b), Lynch, J.D. and Duellman, W.E. (1997), Lynch, J.D., Ruiz-Carranza, P.M.
Order, Family: Anura, Leptodactylidae saralda Departments on the western flank of the Cordillera Occidental and Ardila-Robayo, M.C. (1997), Paez, V.P. et al. (2002), Rivero, J.A. and Serna, M.A. (1987), Ruiz-Carranza, P.M., Ardila-Robayo, M.C.
Country Distribution: Colombia in Colombia, at elevations of 1,500-1,800m asl. and Lynch, J.D. (1996), Ruiz-Carranza, P.M., Lynch, J.D. and Ardila-Robayo, M.C. (1997)
Current Population Trend: Decreasing Population It is an uncommon species. Data Providers: Fernando Castro, Maria Isabel Herrera, John Lynch
Habitat and Ecology It is active on low vegetation in primary
montane forests, and has not been recorded outside forest habitat.
Breeding is by direct development.
Major Threats The major threat to the species is habitat loss,
as a result of the expansion of agriculture and livestock farming
in the region.
Conservation Measures Populations of this species occur within
three national parks, including Parque Nacional Natural Paramillo
and Parque Nacional de Las Orquídeas; the maintenance of these
protected areas is essential to ensure the long-term survival of this
species, particularly given its association with primary forest.

EN Eleutherodactylus jorgevelosai Lynch, 1994

Endangered B1ab(iii,v) Geographic Range This species is known from the region of the type locality on the western slope of the Cordillera
Order, Family: Anura, Leptodactylidae Occidental, in the department of Santander, Colombia. It has been recorded from 1,900-2,400m asl.
Country Distribution: Colombia Population It is a rare species, and appears to be in decline.
Current Population Trend: Decreasing Habitat and Ecology It lives on rocks and vegetation up to3m above the ground along streams in mature forests
(and is found only in the forest interior). It breeds by direct development.
Major Threats The major threat to this species is habitat loss, due to agriculture, involving both crops and live-
stock, as well as wood plantations. Some other species of Eleutherodactylus that are associated with streams have
undergone dramatic declines and disappearances, possibly due to chytridiomycosis, so the population status of this
species should be monitored carefully.
Conservation Measures Its range includes the Estacion Demostrativa El Rasgón. Additional protection of forest
habitat in the range of this species is necessary.
© John D. Lynch

Bibliography: Acosta-Galvis, A.R. (2000), Lynch, J.D. (1994a), Lynch, J.D. and Duellman, W.E. (1997), Lynch, J.D., Ruiz-Carranza, P.M.
and Ardila-Robayo, M.C. (1997), Ruiz-Carranza, P.M., Ardila-Robayo, M.C. and Lynch, J.D. (1996)
Data Providers: Martha Patricia Ramírez Pinilla, María Cristina Ardila-Robayo, Fernando Castro, Maria Isabel Herrera

CR Eleutherodactylus jugans (Cochran, 1937)

Critically Endangered A3c Geographic Range This species has a restricted range on the minican Republic, which is better-managed, but is nonetheless still suffering from significant habitat degradation.
Order, Family: Anura, Leptodactylidae Massif de la Selle, in Hispaniola (Haiti and Dominican Republic). It The maintenance of remaining suitable habitat across the range of the species and improved protection of existing
Country Distribution: Dominican Republic, Haiti has been recorded from 1,242-2,146m asl. protected areas is essential.
Current Population Trend: Decreasing Population It is moderately common in suitable habitat, which is Bibliography: Hedges, S.B. (1993), Hedges, S.B. (1999), Hedges, S.B. (2001), Henderson, R.W. and Powell, R (1999), Henderson, R.W.
rapidly disappearing. and Powell, R. (2001), Schwartz, A. and Henderson, R.W. (1991)
Habitat and Ecology It is a terrestrial species, occurring in mesic Data Providers: Blair Hedges, Sixto Inchaustegui, Robert Powell
pine and broadleaf forests, and does not tolerate habitat disturbance.
Eggs are laid on the ground, and it breeds by direct development.
Major Threats The primary threat to this species is severe habitat
destruction (even in the protected areas of Parc National Morne La
Visite and Parque Nacional Sierra de Baoruco) as a result of logging
by local people (charcoaling), and slash-and-burn agriculture.
Conservation Measures Although the species does occur in the
Parc National Morne La Visite in Haiti, there is no management of
this area for conservation, and the habitat continues to be lost. It
is also known from Parque Nacional Sierra de Baoruco in the Do-

CR Eleutherodactylus junori Dunn, 1926

Critically Endangered B2ab(iii,v) Geographic Range This species is known from only four localities in the central uplands of Jamaica. It has been
Order, Family: Anura, Leptodactylidae recorded from 600-835m asl.
Country Distribution: Jamaica Population It is known to be extremely rare and has only been seen or heard on a few occasions since it was first
Current Population Trend: Decreasing described in 1926. Two specimens were collected in the mid 1980s, and one individual was heard in the late 1990s. A
series of specimens was collected in the 1960s, suggesting that it might have been more common in the past. Although
there has been no recent survey work for this species, it does appear to have declined significantly.
Habitat and Ecology This is a terrestrial species that has only been found on the edge of forest in banana groves and
pastures. It calls from rock and earthen crevices, but can it be quite a difficult species to locate, mainly because its call
© S. Blair Hedges, Penn State

is drowned out by that of the invasive Eleutherodactylus johnstonei. It presumably breeds by direct development.
Major Threats The primary threat is habitat destruction, due to agriculture, logging, and infrastructure development.
Their range has now been completely invaded by the introduced Eleutherodactylus johnstonei, which has a much
louder call (and is more locally abundant), and which is quite possibly out-competing this species ecologically, and
resulting in noise interference in mating calls (S.B. Hedges pers. obs.).
Conservation Measures This species is not known to occur in any formally protected areas. Although its range
includes Cockpit Country Forest Reserve, this does not guarantee long-term protection for the remaining habitat. The
maintenance and protection of remaining suitable habitat across the range of the species is essential.
Bibliography: Hedges, S.B. (1993), Hedges, S.B. (1999), Hedges, S.B. (2001), Henderson, R.W. and Powell, R (1999), Henderson, R.W.
and Powell, R. (2001), Schwartz, A. and Fowler, D.C. (1973), Schwartz, A. and Henderson, R.W. (1991)
Data Providers: Blair Hedges
 Globally Threatened Amphibian Species 351

CR Eleutherodactylus karlschmidti Grant, 1931 STREAM COQUI

Critically Endangered A2ae Geographic Range This species is known from eastern Puerto Rico as well as the western mountains, at an
Order, Family: Anura, Leptodactylidae altitudinal range of 45-630m asl.
Country Distribution: Puerto Rico Population Formerly known to be abundant in eastern Puerto Rico as well as the western mountains, it has not been
Current Population Trend: Decreasing seen since 1976. Habitats where it once occurred have been surveyed repeatedly by experts at different times of year
(including the rainy season), without success, and it is now considered to be almost certainly extinct.
Habitat and Ecology It is an aquatic species, restricted to mountain streams, with a preference for rocky torrents in
closed mesic forest. It is known to call from boulders, banks, and waterfalls, and to breed by direct development.
Major Threats The main cause of the decline of this species appears to be related to the synergistic effects of
chytridiomycosis and climate change, although invasive predators might also have had an impact.
© Erik Wild, Proyecto Coquí Conservation Measures This species formerly occurred in El Yunque National Forest, but it has not been recorded
recently from this area. Further survey work is required to determine the population status of this species and whether
or not it survives in the wild. In view of the risk of chytridiomycosis, any surviving individuals might need to form the
basis for the establishment of an ex-situ population.
Bibliography: Burrowes, P.A., Joglar, R.L. and Green, D.E. (2004), Hedges, S.B. (1993), Hedges, S.B. (1999), Hedges, S.B. (2001),
Henderson, R.W. and Powell, R (1999), Henderson, R.W. and Powell, R. (2001), Joglar, R.L. (1999), Joglar, R.L. and Burrowes, P.A. (1996),
Schwartz, A. and Henderson, R.W. (1991)
Data Providers: Blair Hedges, Rafael Joglar, Richard Thomas

EN Eleutherodactylus katoptroides Flores, 1988

Endangered B1ab(iii) Geographic Range This species is known from the type locality near Parque Nacional Podocarpus, although the population at the type locality remains unprotected. Further survey work
Order, Family: Anura, Leptodactylidae Puyo, in the province of Pastaza, in Ecuador, and also from the upper is necessary to better determine the current population status of this species, and to determine whether it might
Country Distribution: Ecuador watershed of Curintza in Parque Nacional Podocarpus at 2,700m asl. occur more widely than currently known.
Current Population Trend: Decreasing It is likely to occur more widely, at least in the area between the Bibliography: Almeida, D. and Nogales, F. (2003), Flores, G. (1988b), Lynch, J.D. and Duellman, W.E. (1997)
two currently known sites. Data Providers: Luis A. Coloma, Santiago Ron, Diego Almeida, Fernando Nogales
Population It is a very rare species. In two years of study in Parque
Nacional Podocarpus, only two individuals were collected.
Habitat and Ecology This species is an inhabitant of montane
cloud forest, occurring in the understorey herbaceous layer. The type
locality consists of a mixture of heavily disturbed forest, a cutover
and cultivated hillside, and less-disturbed, secondary growth forest.
It breeds by direct development.
Major Threats The major threat is habitat loss due to agriculture
(involving both crops and livestock), as well as logging and infra-
structure development for human settlement.
Conservation Measures It is known to occur in the 145,000-ha

VU Eleutherodactylus kelephas Lynch, 1998

Vulnerable D2 Geographic Range This species is only known from the region of the type locality in the municipality of El Cairo,
Order, Family: Anura, Leptodactylidae in Valle del Cauca and Chocó departments, Colombia, from 1,900-2,250m asl. It is unlikely to have a much wider
Country Distribution: Colombia distribution, as surveys of nearby areas have not found the species.
Current Population Trend: Stable Population It is a common species.
Habitat and Ecology It is usually abundant on vegetation along small streams in primary and secondary cloud
forests; it has not been recorded outside forest. It breeds by direct development.
Major Threats There are no known threats to the species at present, as its known distribution is currently conserved
within a protected area.
Conservation Measures The type locality is within Regional Reserve Serrania de los Paraguas. Maintenance of
the species’ habitat is essential to its long-term survival given its restriction to forest habitats. There is a need for
close population monitoring of this species.
© John D. Lynch

Notes on taxonomy: This taxon has previously been confused with Eleutherodactylus calcaratus (Lynch 1998).
Bibliography: Acosta-Galvis, A.R. (2000), Lynch, J.D. (1996b), Lynch, J.D. (1998b)
Data Providers: Fernando Castro, Maria Isabel Herrera, John Lynch

EN Eleutherodactylus klinikowskii Schwartz, 1959

Endangered B1ab(iii) Geographic Range This species has a restricted range in the Sierra de los Organos and the Sierra del Rosario,
Order, Family: Anura, Leptodactylidae Cuba. It has been recorded from 75-182m asl.
Country Distribution: Cuba