•

• • • •
• • • I

atterns and Processes

~ ardon H. Orians and Martha J. Groom

· .._e most wonderful mystery of life may well be the means by which it created so much
~-_,· ersity from so little physical matter. The biosphere, all organisms combined, makes
.., only about one part in ten billion of the earth's mass. It is sparsely distributed
rough a kilometer-thick layer of soil, water, and air stretched over a half billion
- are kilometers of surface.
E. 0. Wilson, 1992

at Is Biodiversity and Why Is it Important?

3~ological diversity, or biodiversity, is the sum total of all living things - the im-
~:ense richness and variation of the living world. Biodiversity can be considered at
' -=-=-~any levels of biological variation, ranging from genetic variability within a species,
~ the biota of some selected region of the globe, to the number of evolutionary line-
ages and the degree of distinctness among them, to the diversity of ecosystems and
:i.omes on Earth. There is no one "correct" level at which to measure biodiversity be- ,
... . .--use
.. different scientific issues and practical problems find their focus at different
~"2\-els .
Few people are aware of the full spectrum of biodiversity, because our own expe-
~ence focuses on interactions with people and those species that interact directly .
-.-ith people, or a few others that attract our attention. Yet if we are to understand
·-hy it is important to preserve biodiversity, we must appreciate the richness that lies
a1!" each level from genes to biomes.
The various levels of biodiversity ar~ best understood from a hierarchical per-
S?ective, from genes, through populations and species, to communities, ecosystems,
and landscapes. Further, we also can describe biodiversity in terms of its variation in
:om position, structure, and functioning. This value of a hierarchical perspective is
:iescribed and illustrated by Reed Noss in Essay 2.1. The following discussion pro-
::eeds from lower to higher levels in the hierarchy of biodiversity.
/ "

28 Chapter 2

ESSAY2.1
Hierarchical Indicators for Monitoring Changes in Biodiversity
Reed F. Noss, University of Central Florida

· Biodiversity and how to save it is vegetation. Function includes the cli- magnitude with human activity. In
the subject matter of conservation biol- matic, geologic, hydrologic, ecological, order to have any chance of protecting
ogy. If you have read this far in this and evolutionary processes that gener- biodiversity against the onslaught of
book, or even skimmed its pages, two ate biodiversity and keep it forever these factors, we must have early warn-
things about biodiversity should be changing. ing of change; hence the need for moni-
clear: (1) it is complex, and (2) it is Change is universal, but some kinds toring. Because biodiversity is multifac-
always changing. How on earth can a of change threaten biodiversity. eted and hierarchical, those indicators
conservation biologist or land manager Changes in climate, changes in distur- that we select as targets for monitoring
deal with this mess? bance regime (such as fire suppression, should represent all of this complexity.
First, we need to make some sense or conversely, increases in ignitions), Otherwise, something might fall
of the complexity of nature. We can dis- introductions of novel chemicals into through the cracks.
sect the biodiversity concept into mean- the environment, and species introduc- Land managers are familiar with the
ingful components, and yet retain some tions or deletions are all changes likely use of indicator species, often selected to
idea of how they all fit together, by to degrade native biodiversity. These represent a suite of species with similar
appealing to hierarchy theory. There are kinds of changes happen naturally, but habitat requirements. As a well-known
several kinds of hierarchies in nature, often occur faster and are of greater example, the Northern Spotted Owl
including the familiar levels of biologi-
cal organization (such as genes, popula-
tions, species, communities), hierar-
chies of space and time, and hierarchies
of rates. There are also ethical hierar-
chies; for instance, many people care
about the suffering of individual ani-
mals, but some also care about the loss
of species, ecosystems, and biomes. All
of these hierarchies are nested; that is,
higher levels enclose lower levels and,
to a great extent, constrain their behav-
ior. A tree is part of a forest stand, the
stand is part of a landscape, the land-
scape is part of a physiographic region,
and so on. If the physiographic region
is inundated by a volcanic flow, every-
thing nested within it also goes.
Biodiversity is not just species
diversity. A comprehensive approach
to biodiversity conservation must
address multiple levels of organization
and many different spatial and tempo-
ral scales. Most definitions of biodiver-
sity recognize its hierarchical struc-
ture, with the genetic, population-
species, community-ecosystem, and
landscape levels considered most
often. Each of these levels can be fur-
ther divided into compositional, struc-
tural, and functional components.
Composition includes the genetic con-
stitution of populations, the identity
and relative abundances of species in a
natural community, and the kinds of
habitats and communities distributed
across the landscape. Structure
includes the sequence of pools and rif-
fles in a stream, downed logs and
snags in a forest, and the vertical lay- Figure A Compositional, structural, and functional attributes of biodiversity at four levels of
ering and horizontal patchiness of organization. (From Noss 1990.)
 Global Biodiversity 29

- · occidentalis caurina) was selected TABLE A Hierarchical Indicators for Monitoring Biodiversity
_ . . , managers of national forests in the .. ~·- . ",' ,.,.-
';.:.-. ~-
·~.

,, G~N·. ETI~· . . ,, ·.. . " .::,. . :· . ""'

~..,..,-~. ~-c Northwest as a surrogate for all
L
:~ \i. . ~ :z~,

~\"-~species associated with old-

"' .. . I!ei}c ,diyersjtY,: ~ · , ,: '1' •. _·: , foliage profiles in.staQd.. ~ _· : : :. .:·
·:· ··
.Q:tlJ~-\rth forests. However, the use of
". • ~resen~e/ absen.c~ of rar¢ ~leles . ·· ·., • Can0py density and size, dispersiQn "'.
:__.;.......-cator species has encountered B tructure , ,. . . . . .: ,·_ · ·· · · of-canopy openings. ,, , ,, .~ , , . :·
·-"'"""""""~ems, including biased selection "~ • Heterozygosity . . 1 :. . , . :. , .. 'Areal extent of each disturbance " . ....
,; •
...,....,;.:::::..._n· a, false assumptions about • Phenotypic: polymorphism . ,., ·~ ,. : . ~· 'event (e.g.,~ fires) · · · . . ;,: "" .;
S3aaJ-es habitat relationships, unwar- " ·Function" '· · · · · . · ... , , ! ., ,. , •• , ", Funct!on . ·, "' · · " -: , , " -~,.
""'"'!:"""·Eed extrapolations from one species " ~ Sy~ptoms ofinbr€ellin depression .. : ·. • Frequen9',:intepsity, rerum interval, '\
:· · .. " , pr rotation,period of fires anti other a
_ . - . . . ~..ers, and flawed design of moni-
~· or ...e~til_.lty, ·.a.' bn.o. .r mal snerm, redu.·ce ' " ; ' na~U:ral qp.d artf\rof?ogef:U~ distut- ' '" .,
"-----'..~..-~~..10..1_ programs. But species will con-
' >•

• r.es,.ts.t.anc~. t~ Stt~ease~ rh~rEJ:ologtcal

••• w · · .. bances, . . . ,, . - -· " ·~· .,
: "'t'e to be useful as indicators, partie- . ·~ <;tbrtqrmaUties, O! asynunetrtes) ~· · ~· ,, - : • Cycling rates. for yarioU:~ !Sex Iju~i: :.
'----'-~~ if we focus on those most . "" • Inbreeding/ou.t\)r~eding :rate~ • :; · ' · ,. ents (e.g., N, P} ., . , . .. , .
-
,."':: ·ve to human activities and those ·· • Rate of senetic intercliange qetweeh ' " • Intensity or sever,i ty of d..istu;barice ....
lay pivotal roles in their ecosys- · · populations (measured by.ra.Je.,o( d;is- "' ·· events ,,, · ,. * .~, · " , >. .,,

,, · persal and subsequent reprod uct!op ,. "' · : i Sej1S9rlality or periodi-city of,disturr- v

~. .-.. :.-· For example, change in the abun- ,_, · i! of migrants) · ,, ·~ ~, M • , ., " bances· · . · · · · ~ ~ < .: .
- ·- . re of woodpeckers may warn us of -~ POeULATION-SPECJES :• .. ·:. :: '" ' ,. Prediclal;)ility or variability of distur-,:
_ ..._:e changes in populations of ~ ~ bances · ~- · · · · ·· " ' ·· · "'
' Compositio~ ' ., · · . .· : , . ·:· ~ ,, . : : . Hu~~ intru~ion'~rates and interlsi; ·~·
• M •• • , • • ·
1

_---...Jj., species that use woodpecker cav- • Absolute aud r~lati~e abundance, · ·· "
.. -

ti' · ., "- · · , ,,. •..

~s. iBut we should not carry our ,: ·. densitfy basal area, cover, impqrt?n~e~ j( < es . ·.;.
> < • < ' y ; ; i ·.; ·., 1' '<

EC:!aJ)olations too far. The idea of one · ' value for ·,v arious ,species . .. , j • , . LANO.SCAPE. ,_. "'· · · · - ·. , . .~ .,.
.. ';: ~ ··,. .; . ' "

~ • ~St!~':a~o, age dis~il~un~,:and oth~r ' ~ ~ ~'%~~t6:, ~~~b~tio~, ~i;~e:s,. and ::

s::ea·es representing all others that 5

~a similar habitat is not ecologi-

, : • . aspects Of population stru9turefor ,. , , proportiqn$ Q.f pate~ !)rpes. (such·ag· ·
- · realistic. ~ sensitive species, keystone species, · . ~- forest type~ a~a serfil .st~ges) acrbss ·~
- cators for monitoring biodiver-
..........oiL&.·
·..·
,. .
.
: an~ ot}:ter special interest species . t • . . , ,,,, the laridscqp~ . .: :.. . ·. . . ·• · .
. . ;:::;:;;..-. ust consist of much more than :: ·.. • J.?isn;ibution ana dispersion ofspecial . . . ' • 'Total amount of late sp.ccessional for.:.'
setr•of indicator species. Because bio- ·' ,,, interest spe~ies~cross 'the region , ,.. , est interior· habi~t . -~ 1
, .· ·, . '" ""

·~
--- ~"'S . ity is distributed hierarchically,
should indicators be. A frame-
' """'""'"'·'""" ·or selecting biodiversity indica-
_____, tight follow a nested hierarchy
} >· ··.
:Function '" : . . ·.. · , " . . . ' . · , , }· ' • Total amount,of forest patchperime-.
• Population.growth ~n~ flucfYatiqn'~ ~, ,. ' ter' and edge zone , ., · L ,, . . , ,.
n trends of special interest species . · . Structure ' · ·; , · · ,. '· · ·
, • Fertility, fecundity; recruitmen\. r~te, . ." .· le-:>aac•·~~~se~r.za.,e sf..rtaeqgue'eann_.cy.· fdotr'setsrtibtyuptieo,nafnodr _.
-:· survivorship/ mortality rate,jndivid-.. .. ,.
ual growth Tate, and other~.indivi,dttal ... ~.
·j

. . :.
·!. ··
. 1 n d
?cross p.l! stage~ and types
· _, ·
. .
-...:,li.Lpositional, structural, and func-
__,_~~~ elements (Figure A). A monitor-
· • '· .· anti population-health parameters . . ., ,, . • Patctl size diversity index · i" ·~
, -~· • Trends';in' Habitat components fo:r. spe- · ,. • Size .frequency distribution qf !ate ,.: ·'"
__,....,., -rogram should select a broad , : _.. , ¢i~J ip.ter~st species (varies·by species~ ·: . · · successional interior forfst :patches,. :·
•
:::::::==~~-~of indicators that correspond to
-......._...... ~management questions, such : ' ~~~~!;eijJt ~~~;~~;=d : ; : .~. ', ~~~~~f~d;e~g~ ~o~e, ~vally: ,
--.....are populations of rare species •setJStti'lity qf ~Pec!es, 1.1} ~~lati~n.:to..1and
F '' ··· • Forest patch perimeter: area ratio.. " ,.
~...,~ _ maintained in sizes and distri- . us~ practic~s {Uld 9ffie~ iiffipe}l~es] : " ··. , :: .• Eqge zone:'interior zone ratio · ., &. ••

' COMMUNITY~ECOSYSTEM · .. , ~· .. • Fractar dirriensiort ' - ' . · ·!t ·r

~LJ~ that assure long-term demo-
_ . _ _ _ .....

..;;;;_.._,.;.v.~...- c and genetic viability? Are the

:,· . composilidn 'i ·· , -- • ·t ·.~ ·· , ., , ., . . " • Patch ~h~p~ mdices~ · ~ , :: . _:· ' ~ :;:·
.:. ··. ~ ld~~ti,ty, ~el~tive ab~dance, freq~en- · ., , · • Patch density. . , ,. . :· . .. ...·~ . :,
-~al structure and species compo- , . , cy, nc~ess, .ard eyenr1ess ~fspeetes . · ·· · ·• Fragmentation indices , , ; , . ·.
_____.._... . of the community being main- , . ansi gu.J.h:!s (m V~!oushab1tats) ' · - • Interpatch distanee (mean, median~.
~~: . Is the configuration of the .;J :t • Diversity;.of tree ag~s pr:siZes ih com.,; , ·· ·· · range)'for al11·forestpatches and fo~ .
~_....._-~~,ape adequate to permit normal ·•
1
' munity (stand) ·
J; , . , . . . .~ · .. ·~ ·: late successional forest patches "'·· .;i .,,
=:.:~m ents of organisms? Using the · i' • Ratio of exotic specie&to.native , . . . ,• Ju~tapositiqn mecrsures (percentage of
., ~· ~pec~~srin c~~mu:lity. (species xich"7, . . area witli.irl a''definoo·~distance from .;
~e of a managed forest land- ness, cover, and btomass) ·• ,, . . . ,, ; < p~tch oc~u£ied'by diffetertt habitat f
~~~ some measurable indicators
__ "'"'"'~-.ght help answer such ques- .• : ~ ~n~~~~e'!dn~:~:thr:~t~n~d, . : : . ~~~iJ~t~~~1h~:?t~~~~ia-: :
,____.~ ar:e listed in Table A. Only with .' .Btructure ' · ' · ~ , · " ' ·' · • Structurql contrast (~~grqtuae' of dif.:;;
~n, - comprehensive approach will .. :. ti_ Freqjie~cy (;i~trib¥tiort of seral' stages ·' ··· : ference between <!djac~rjt ljabi!afs, :· "
,. , (age ~lf1sses) for e~c!'t forest type artd ., ·· ,. · measured for; various structural · '
==.:sez·v.ation biologists be able to , 't" across all rypes . .· · · ,:·' .· ,, ·· ·~ , · ·· attributes) . ,. " , ·,. · .· ~: . :
1

--~ changes in biodiversity and • Average and range of tr~e age~· ~it!Urt ' ' • Road density (mi/ mH or km/knj. f ., 2

e information they need to ~ definea seral stages. , ~ ·. . . , ,. ..· · , · for different classes·2ot road and all .
~:sa~;~e it. ,. . • Ratio of area of Ratural forest ofaU ages '. ': ;, toad·classes combined , _ : 1

· ··· " to area in·clear...cuts and plantations " _ . :Function · ·< ·. · .., • ' :; ;;

· ' ~ ABundance and density,of snags; . ,, · · • DiStUrbance indicators (see above) · "~
:· ,. : aoWned logs,.-and other·definea ,. S· " .•• • Rat~~ of ~qtrferit/e~ergy, ana bio- .k ;\
~ · structural elements in various size , ~ logicaLtr~a.rtSfer .Pe.nyeen'differen! : ,
., : '., ~a <;iecay classes · : · · ;; · " , communiti~s; and,patch~ ·,in~,tJ!e ,. · · ·
'" " ~ $.p,9tia( disEersion of st~ctural ele-· t ·' landsc~pe ·' , . ~·. ,. .~ ·~
_ . ·' ments anfl pa.tcJ;tes . · : - · .., , · . ~ ··· , .. " , ::
_., t
" .
30 Chapter 2
Components of Biodiversity
Genetic diversity
Genetic variability is the ultimate source of biodiversity at
all levels. The number of genes found in different species
ranges over more than three orders of magnitude (from
about 500 in a parasitic bacterium to more than 20,000 in a
mouse or a mustard plant), although not all of those genes
code for products. In addition, most genes exist in several
forms, or alleles, so the possible number of combinations
of genes is enormous, much larger than the number of in-
dividuals present in any species. This genetic variability is
the material upon which the agents of evolution act.
Recent advances in molecular biology provide the
tools we need to measure the amount of genetic vari-
ability present in organisms. Measures of variability
within local populations provide important clues that
help us understand the nature of forces acting on genet-
ic variation. Such knowledge has important practical ap-
plications in, for example, designing captive breeding
programs for rare species to reduce deleterious effects of
inbreeding, or determining the best sources of individu-
als for reestablishing populations in areas from which
they have been exterminated. Measure.m ents of the
amount of genetic differences among species enable us
to determine rates of evolution and establish phyloge-
netic relationships among organisms.
Genes are the fundamental unit of biodiversity, with-
out which no evolutionary change occurs. Sp~cific genes
or cq.mbinations of genes allow individuals to tolerate
polluted conditions, exploit resources more efficiently, or
compete better with other species. Abilities to tolerate
temperature changes, or disperse great distances, which
are at least in part genetically based, may be crucial to
the persistence of species in the face of a rapidly chang-
ing climate. Preservation of genetic diversity is thus an
important goal of conservation biology.
Preservation of genetic diversity in crops and live-
stock breeds is also important. Distinct varieties of do-
mesticated species are adapted to a wide range of envi-
ronmental conditions. These varieties are especially
important for human welfare, particularly in the poorest
areas of the world where other supporting institutions
are weak or absent. Further, greater agricultural efficien-
cy means that less land must be cultivated or serve as
rangeland to meet the needs of a given population, leav-
ing more land available for preservation of biodiversity.
Population-level diversity
An extremely important component of biodiversity is
the variation found within members of a species or pop-
ulation. A population is a group of individuals of a par-
ticular species living in a specific area at the same time.
Genetic differences among individuals within a popula-
tion, and among different populations of a species reflect
the evolutionary history of that population, and shape
its future potential for adaptation. It is in local popula-
tions where responses to environmental challenges
occur, where adaptations arise, and where genetic diver-
sity is maintained and reshuffled each generation. We
are concerned here primarily with genetic differences,
because only heritable differences can be passed on to
future generations. The potential rate of evolutionary
change of a population is proportional to the amount of
available genetic variability. Variation among popula-
tions of a species is the result primarily of adaptations to
.local ecological conditions. Locally adapted populations
of a widespread species may have particular genes and
gerie combinations critical for survival and reprod.u ction
in those particular areas. The nature and extent of be-
tween-population genetic variability reveals much about
the evolutionary history of populations. The theoretical
and practical importance of population-level genetic di-
versity is treated in greater detail in Chapter 11.
Individual populations may have great conservation
value. Many wide-ranging species consist of a number
of genetically isolated or semi-isolated populations, each
of which is adapted to a different environment. For ex-
ample, populations of some plant species have evolved
high levels of tolerance to metals in the soil (Antonovics
. et al. 1971). If we were attempting to re-establish a pop-
ulation of a plant on metal-contaminated soil, we would
be successful only if we planted genotypes that could
tolerate those conditions.
Guppies (Poecilia reticulata) in Trinidad streams have
very different genetically based color patterns, clutch
sizes, and offspring sizes depending on whether they
occur in streams with or without predatory fishes
(Reznick and Bryga 1987; Reznick et al. 1990). If we were
trying to reintroduce an extirpated population of these
guppies into a stream with predators, we would need to
use individuals having the traits that promote coexis-
tence with predators. Some plant species have marked- .
ly different genetically based growth forms in low- and
high-elevation ecosystems (Clausen et al. 1940). Such ge- ·.
netically based plasticity in life history characters, which
is probably a pervasive feature of most organisms, is
under-appreciated by conservation biologists. . ~ ·
The ecological functioning of each population and the
services it provides may be largely independent of other
populations of the species. For example, if a population
of a bee is exterminated from a river basin due to pesti-
cides, the flowers that depend on that bee for pollination
have lost that service, even if the "species" still exists else-
where, unless some other species can assume that role.
Thus, the loss of local populations reduces the ability of
ecosystems to provide goods and services to people and
other species living there. For this reason, the massive de-
clines in abundances and ranges of many species are of
great conservation concern (Gaston and Spicer 2004).
 Global Biodiversity 31

uman cultural diversity
~.:te variety of human cultures is an important compo-
~-ent of population-level diversity. Cultural diversity em-
:: dies a reservoir of human knowledge, skills, values,
2·d management traditions that have evolved for thou-
::- ds of years. Different cultures interact with nature in
-.:.:-..any ways that reciprocally influence the development
:human societies and their impacts on nature. Cultural
-=-=actices represent "solutions" to the challenges of find-
:_ o or growing food and defeating infections and disease
::-- a given habitat. In Essay 2.2, Mark Plotkin and Adrian
: ::lrsyth describe some of the sophisticated solutions in-
.:.. aen ous cultures devised to maintain productive agri-
.::: .tural systems in difficult environmental condit.i ons.
There are 6526 distinct spoken languages, with the
~-eatest diversity concentrated in tropical regions, with
- :ew exceptions (e.g., Australia, United States) (Figure
- - J. To the extent that language reflects cultural differ-
=- ces/ this provides at least a lower bounds on the total
=-'!Ilb er of distinct cultures worldwide.
- cltural diversity can also be measured
(A)
""' _.- counting the number of localized, in-
1000
-·genou s human populations, which are
:..:. · - st numerous in tropical moist forest
800
_:- ;:~itats (see Figure 2.1).
The importance of cultural diversity is
-:dely unappreciated. Essentially, cultur-
~ diversity encompasses many of the ~ 400 Q)
ual process. Once a species becomes separated into two or
more populations/ the daughter populations may evolve
independently for a long time before they become repro-
ductively incompatible, or they may become reproduc-
tively incompatible before they evolve noticeable morpho-
logical, physiological or behavioral differences. Thus,
separate populations within a species may exist at various
stages in the process of becoming separate species. Con-
servation biologists are often concerned with preserving
evolving populations that have not yet become full
species. We do this because the genetic differences evolving
populations have accumulated may enable them to survive
in different future environments. We seek to preserve the
differing evolutionary lineages represented by each sub-
species, incipient species or full species, as well as the po-
tential for variable evolutionary outcomes that will unfold
as species or subspecies persist separately or interact.
Although species are generally defined in terms of re-
productive isolation, such information is often not avail-

_ echanisms that allow people to adapt to

-~anging conditions in their .environment. z 200 1
Diversity of species
_ ~ st people think of diversity of species
-~en they hear and use the term "biodi-
..--asity." Species are fundamental units of
.....,. . u tion, and species are the primary tar-
~:s of some of the most powerful pieces (B)
~ 350
: conservation legislation, such as the
0
----. Endangered Species Act (ESA) and ·_p
ro
300
-._e Convention on International Trade in ~
~ 250
~:;ecies (CITES). Therefore we will em~ ~
- 00
200
~~ .asize species diversity in this chapter ~

= throughout the book. Yet we should ~.......b.O 150

_ ~:J in mind that population-level diver- "d
.s 100
B. _.- can be as or more important for con- '-H
0
- -ation , and that communities, ecosys- 50
~.. . . , landscapes, or ecoregions may be
~--:e most appropriate level for s.ome con-
~-ation activities.
_-\ species is a group of actually or poten-
~;; ~~~y interbreeding natural populations
::----are reproductively isolated from other
=.::.h groups. Deciding whether two popu-
-.;;cons constitute different species may be Figure 2.1 Linguistic diversity (A) and numbers of indigenous cultures •

-:.;:=:1cult because speciation is often a grad- (B) across the world. (Modified from Mulder and Coppolillo 2005.)
32 Chapter 2
The Importance of Indigenous Knowledge Systems
Mark J. Plotkin, Amazon Conservation Team, and Adrian Forsyth, Gordon and Betty Moore Foundation
. The philosopher-naturalist Laurens
Van der Post remarked that the most
damning legacy of colonialism has been
its relentless tendency to separate tropi-
cal peoples from their land and culture.
"The great mistake people make is in
thinking about the aboriginal races in
South Africa the Bushmen and Hot-
tentots and the like as primitive peo-
ples. They were actually very sophisti-
cated societies and cultures with
immense awareness and very impor-
tant values."
The sophistication of many indige-
nous cultures is most evident in their
agricultural systems. Over 200 years ago,
a Jesuit priest in lowland Bolivia, Fran-
cisco Eder, noted that an extraordinary
irrigation system in the local savannas,
which had been devised in pre-
Columbian times, still supported an
enormous population of Mojeno Indtans.
Thanks to the influence of the padre and
his cohorts, as well as introduced dis-
eases, this Amerindian agricultural sys-
tem was abandoned and the savannas
that once fed the teeming local populace
today are covered with a scrub vegeta-
tion supporting a few mangy cows.
The chinampa system, developed in
pre-Columbian Mexico, remains in use,
albeit in a much more restricted range
than it once covered. Usually con-
structed in swampy or lacustrine envi-
ronments, chinampas consist of garden
plots created by building small islands
using layers of vegetation and mud. The
surrounding waters seep in and provide
the necessary moisture, while mud
scooped up from canals around the plot
is periodically added as fertilizer to the
garden on top of the island. According
to Dr. Jim Nations of Co11servation Inter-
national, chinampas not only produce
food and other useful crops year-round,
they do not deplete the soil or require
artificial fertilizers or pesticides.·
A similar system was rediscovered in
the altiplano of northwestern Bolivia.
Dr. Alan Kolata of the University of
Chicago had long been intrigued by a
series of ridges and depressions in this
remote region that seemed to indicate
some form of agricultural system that
involved raised planting beds sur-
rounded by canals. The local Aymara
Indians, however, employed no such
-..
system. Kolata and his colleagues
worked with the Aymaras to dig a series
of raised beds surrounded by ditches
and the results were a crop yield seven
times greater than the local average.
Here we see three different fates for
three independently evolved agricul-
tural systems: The Mojeno system has
been lost, the chinampa system still
exists, and the altiplano system would
have remained lost if it had not been for
an intrepid anthropologist and his
indigenous colleagues. How many other
equally valid and possibly even more
productive systems have completely dis-
appeared without a trace? The superior-
ity complex that drives outsiders from
Western societies to want to replace
"primitive" systems with an ill-suited
one developed on foreign soils continue
to this day: The Indonesian government
is currently trying to get tribal peoples
in highland New Guinea to forgo their
traditional and highly productive agri-
cultural system based on sweet potatoes
or sago palms, and replace it with rice,
an inappropriate crop. This colonial atti-
tude comes from cultural biases and a
lack of understanding.
Yet, the problem we face is not just
loss of agricultural systems themselves,
but extinction of cultivar diversity as
well. Indigenous agricultural systems
typically contain many different vari-
eties of a single crop, much as a farmer
in the industrialized world may grow
several different types of corn: one to
feed his family, another to feed his ani-
mals, still.another to make popcorn,
and another to sell as a cash crop. The
Amerindian farmer in the Amazon usu-
ally cultivates distinct varieties of cas-
sava for the production of bread, beer,
meal, porridge, and whatever other end
products he desires. The Tirio Indians
of northeastern Amazonia have at least
15 varieties of cassava in their gardens
while other tribes like the Machiguenga
of Peru may cultivate several dozen.
These crop varieties often harbor
adaptations that enhance yields or
increase resistance to pests and diseases.
ReceD.tly, crosses of indigenous and
commercial varieties have led to tremen-
dous improvements. A barley plant from
Ethiopia was crossbred with barley in
California, providing resistance to the
lethal yellow dwarf virus, which threat-
ened an industry worth $160 million per
year. The yield of cassava in Africa was
increased ten-fold because of disease
resistance provided by cassava from the
Amazon. And scientists have recently
found that a variety of sunflower being
cultivated by the Havasupai Indians in
the American southwest offers resistance
to a blight attacking sunflower crops in
the Old World. ·
The late economic botanist Edgar
Anderson once stumbled across an
Indian garden in Guatemala that ini-
tially seemed more of a rat's nest than a
productive agricultural plot. It was .
only after careful study that he realized
how much more sophisticated than the
botanist was the farmer:
In terms of our American and ...
European equivalents the garden
was a vegetable garden, an or-
chard, a medicinal plant garden, a
rubbish heap, a compost heap, and
a bee-yard. There was no problem
of erosion though it was at the top
of a steep slope; the soil surface was
practically all covered and appar-
ently would be during most of the
year. Humidity would be kept up
during the dry season and plants
of the same sort were so isolated
from one another by intervening
vegetation that pests and diseases
could not readily spread from plant
to plant. ... I suspect that if one
were to make a careful study of
such an American Indian garden,
one would find it more productive
than ours in terms of pounds of
vegetables and fruit per man-hour
per square foot of ground.
At a time when there are ever more
mouths to feed on this planet, Ander-
son's admonition to look more closely
at the form and function of indigenous
systems seems ~ike advice worth fol-
lowing. Modern management tech-
niques often overlook and disparage
these indigenous systems, which are
based on centuries of in situ sustainable
existence, in favor of high-tech but
often inappropriate and expensive sys-
tems that fail. We can learn a great deal
about environmental management (and
humility) from such cultures.
 Global Biodiversity 33

::= ""'le. For example, the concept cannot be applied to organ-
:.s::ns that reproduce asexually. Also, fossils, the source of
_ ..ost information about organisms that lived in the past,
_arely provide clues about breeding behavior. Finally, in-
.:c::rnation about reproductive behavior is not available for
_ ost species that live today. Therefore, many species are
::-ocognized on the basis of their morphology. For example,
:__ efforts to complete an inventory of the biota of Costa
-~~~ biologists have relied heavily on distinguishing mor-
,1-. .I.<L'ospecies, particularly among arthropods (INBIO 1995).
---:."""_e morphospecies concept is useful, because it can be ap-
_:::ed when other data are unavailable. Moreover, given
~..itt evolution is, to use Darwin's words, "descent with
.=-edification," organisms that look alike generally share
:::any alleles that code for their body structures. This is
~ -:)~ most species designations based on morphology
~;:-.--e been supported by subsequent genetic data.
-~ closely related species continue to evolve in genetic
: lation from one another, they diverge and accumulate
,;ettetic differences. Thus, the number of unique genes,
- d the morphological and physiological traits they en-
e, increases in a lineage of organisms over
-=:w.e. These unique traits are important compo-
:Le :lts of the rich fabric of life on Earth. To deter- TABLE 2.1 Number of Living Species in Major Phy/a
classify organisms hierarchically in ways that reflect
their evolutionary history, constructing classifications in
which the higher taxonomic categories contain all of the
descendants of their most recent ancestor, but no other
species; that is, categories that are monophyletic.
How Many Species Are There?
Approximately 1.75 mi1Hon living and 300,000 fossil species
have been described and given scientific names (Table
2.1). On average, about 300 new species are formally de-
scribed and named every day, and no slowdown is in
sight. New animal phyla, Cycliophora and Loricifera,
whose members live in interstitial spaces in ocean sedi-
ments, were first described within the past 25 years! Not
surprisingly, because they are based on incomplete and
indirect evidence, estimates of the actual number of living
species vary widely. Current estimates of the total number
of living species range from 10 million to as high as 50
million or more (May 1988; Wilson 1992; Gaston and
Spicer 2004). In other words we do not know within an
0

_._ methe degree and significance of these differ-

::.:·ces, biologists attempt to reconstruct
... ylogenies or cladograms. A phylogeny is a
~:.-pothesis that describes the history of descent
:a group of organisms from its common an-
.:estor. A lineage is represented as a branching
~-ee," each node of which represents a specia-
-= rr1 event. That is, the phylogeny shows the
:--der in which lineages are hypothesized to
~ Ye split. Systematists use a wide variety of in-
: .:n1ation morphological, developmental,
?: ysiological, behavioral, and molecular to
reconstruct phylogenies.· Computer programs
zre used to infer the most likely evolutionary re-
..::ionships among organisms in a lineage (e.g.,
_ HYLIP, Felsenstein 2004). Conservation biolo-
c_.:sts use phylogenies for many purposes, as we
Fill. describe in Chapter 11.
The biological classification system in use
_ day was first proposed by the Swedish biol-
. t, Carolus Linnaeus in 1758. Linnaeus gave
ach species two names, one identifying the
s~'ecies itself, and the other the genus (a group
:closely related species) to which it belongs
.:. . the Linnean system. Species and genera are
~_:rther grouped into a hierarchical system of
-i gher taxonomic categories families, orders, Source: Adapted from data in Margulis and Schwartz 2000; Lecointre et al. 2001;
.<zsses, phyla, kingdoms, and domains. This IUCN Red list 2004; Gaston and Spicer 2004; Maddison and Schultz 2004.
~-stem provides unique names for each aMajor phyla listed have >1000 described species.
5~'ecies and is an aid to memory and commu- bProtista lumps 5- 7 distinct kingdoms within the domain Eukaryota.
- .;cation. As much as possible, taxonomists cEstimates range from 6-30 million.
34 Chapter 2
order of magnitude the number of living species. Thus, a
large fraction of species likely to be exterminated during
the twenty-first century will disappear before they have
been named, much less understood ecologically.
The immense richness of viruses, bacteria, archaea,
protists, and unicellular algae is still largely uncata-
logued. Within the domain Archaea, new phylum-level
groups are discovered every year (Furhman and Camp-
bell 1998), and the richness of species within these
groups is unknown. There are few studies of viruses ex-
cept those that attack people, our domesticated plants
and animals, and the organisms we study scientifically.
How many types attack noncrop plants and insects is
unknown, as is the number of marine forms. Similarly,
bacteria and protists living in soils or that attack inverte-
brates have scarcely been examined. Terrestrial algae, es-
pecially those living on bark and rocks, have been little
studied. All estimates of the number of species in these
groups are crude guesses at best (Torsvik et al. 2002).
About 80,000 species of fungi have been described, but
the total number living today likely exceeds 1 million. For
example, the 12,000 fungal species described in Britain is
about six times the number of native vascular plants de-
scribed for the same area. If there were, on average, six
fungal species for each vascular plant species worldwide
(the ratio found in Britain), the number of fungal species
would be approximately 1.6 million (Hawksworth 1991).
The global ratio could be much higher or lower than the
British one, but we have little else to base estimates upon.
Taxonomists believe that the number of species of ne-
matodes is very large because millions of individuals
may be present in 1 square kilometer of soil or mud;
more than 200 species have been reported in samples of
just a few cubic centimeters of coastal mud (Poinar 1983).
Yet, almost nothing is known about species ranges and
rates of species turnover geographically, so global esti-
mates are very uncertain, although these data make a
compelling case for a vast diversity of nematode species.
Approximately 30,000 species of mites have been de-
scribed but, because knowledge of tropical mite faunas is
extremely poor, the actual number of living species could
easily exceed 1 million. Nearly 1 million species of insects,
the world's most speciose group of organisms, have been
described, but this is certainly a small fraction of the total.
Most (>55°/o) of the .insects collected by fogging the
canopies of tropical trees, for example, are members of un-
described species (Erwin 1991), and samples taken with-
in 70 km of one another at four sites in Manaus, Brazil had
only 1°/o of their species in common (Erwin 1983). These
samples hint at a vast unexplored diversity of arthropods,
just in the canopies of tropical trees.
Earth is a relatively unexplored planet biologically.
Not only are most living species still undescribed, but
very little is known about the life history and ecological
relations of most of the species that have been named.
The rate of description of new species today is higher
than it ever has been, but the rate is nonetheless inade-
quate to accomplish a reasonable inventory prior to the
likely extinction of many of them.
Diversity of Higher Taxa
Recent advances in molecular evolution-have painted a
radically different portrait of taxonomic diversity than
existed even 20 years ago. Until recently, taxonomists
classified all living organisms into five kingdoms: Ani-
malia, Plantae, Fungi, Protista, and Monera (all prokary-
otes). Today, we recognize that there are deep evolution-
ary divisions among the prokaryotes. They are now
divided into two large domains, the Archaea and the
Bacteria (Woese et al. 1990; Figure 2.2). The remaining
four kingdoms are grouped together in the third do-
. main, the Eukarya (see Figure 2.2). The genetic diversity
among the Bacteria (which contains all familiar prokary-
otes such as gram-positive bacteria, cyanobacteria, and
green sulfur bacteria) and Archaea (which are unfamil-
iar to most, but include organisms adapted to life in ex-
treme environments such as deep sea hydrothermal
vents, hot springs, the guts of animals, and hypersaline
environments) is as great as that among the long-recog-
nized kingdoms among the Eukaryotes. Two new king-
doms are now recognized among the Archaea (Crenar-
chaeota and Euryarchaeota) and at least six more are
recognized among the Bacteria. Similarly, the protists,
which now are known to be vastly more diverse than
previously recognized, have been divided into seven
more kingdoms. These discoveries are exciting because
they force us to consider the importance of organisms
whose size make them generally invisible to us, yet are
evolutionarily equal to the plant and animal kingdoms
that have a greater hold on our consciousness. To appre-
ciate this in another way, the diversity among all plants,
fungi, and animals in small subunit RNA, the molecule
used to create the modem phylogeny of all life on Earth,
is only 10°/o of the diversity spread across all the rest of
the Eukaryotes, Archaea, and Bacteria (Olsen and Woese
1996). Freeman (2002) describes their importance well:
Bacteria and archaea are ancient, abundant, ubiqui-
tous and diverse ... their abundance is well-docu-
mented. A teaspoon of good-quality soil contains
billions of microbes. In sheer numbers, the bacterium ·
Prochlorococcus found in the plankton of the world's
oceans may be the dominant life-form on the
planet ... a drop of seawater contains a population
equivalent to a large city. Yet Prochlorococcus was first
described and named only recently in 1998.
Bacteria and archaea are also found in almost
every conceivable habitat. On land, they live in envi-
ronments as unusual as oxygen-free mud, hot springs,
salt flats, the roots of plants, and the guts of animals.
They have been discovered living in bedrock to a
 Global Biodiversity 35

- Domain Bacteria Domain Eukarya

Bryophytes Seedless vascular plants Gymnosperms

~~"
'!o..,1>-
0.1>- ~1f
~1>- ·~ 0.1>-
~0 o.~'\-
-#
1>-
·0~
~...,_1>-
:\:'-0~
# '!o...o
~0~
0
»01>-
~~
~
~1>-
0
'!o..,1>-
o1f
~~~
~

~
0~
-~
~~
(;~ (}~ ~~~ o/~ ~0 '9# cY
~
~

Jellyfish, Comb Insects, spiders, Segmented Snails, clams, Sea stars, Vertebrates,
sea anemones jellies Flatworms Roundworms crustaceans worms squid sand dollars ascidians

~~are 2.2 (A) Domains of biodiversity: Bacteria, Archaea, and Eukarya. Note that
--'· fungi, and animals, evolved relatively recently within the Eukarya. (B) Major
~"'-'Tr":-""'4:.: of plants. (C) Major groups of animals. (Modified from Freeman 2002.)
36 Chapter 2
depth of 1500 m below Earth's surface. In the ocean
they are found from the surface to depths of 10,000 m,
and at temperatures ranging from ooc in Antarctic sea
ice to over 110°C near submarine volcanos.
Organisms so widespread and adaptable may not be
in danger of extinction. Nonetheless we should be con-
cerned that such incredible diversity be maintained be-
cause it may be of critical functional importance.
The distinctness of evolving lineages is an important
component of biodiversity. The higher taxonomic cate-
gories of the Linnaean biological classification system
provide rough measures of distinctness of lineages. For
example, phyla ill the animal kingdom are distinguished
by their developmental patterns and their adult body
plans (e.g., having two or three embryonic tissue types).
Measured by diversity of phyla, marine biodiversity is
much higher than terrestrial biodiversity even though
far fewer species of marine than of terrestrial organisms
have been described. Of the 96 phyla (most of them
prokaryotes) recognized by Margulis and Schwartz
(1998), about 69 have marine representatives and 55
have terrestrial ones. Of the 35 extant phyla of multicel-
lular animals, 34 have marine representatives, and 16 of
these are exclusively marine. From this perspective,
preservation of marine biodiversity is more important
than might be suggested simply by comparing the num-
bers of species in marine and terrestrial environments.
The preservation of evolutionarily distinct lineages
above the level of species is important for a number of
reasons. First, the evolutionary potential of life depends
upon the distinctness of evolving lineages, not just the
number of species. Lineages that have been evolving
separately for long periods of time have many unique
genes and gene combinations that would be lost were
those lineages to become extinct. Closely related species,
on the other hand, share nearly all of their alleles. For ex-
ample, we share an estimated 99°/o of our DNA with
chimpanzees. Second, evolutionary lineages are store-
houses of information about the history of life. Scientists
can read and interpret this information with increasing
accuracy using modern phylogenetic methods. Third,
the integrated functioning of ecosystems depends, in
part, on the variety of species in them. For example, mi-
croorganisms have more diverse mechanisms for ob-
taining energy from the environment and decompose a
wider variety of substances than multicellular organ-
isms. Without microorganisms, ecosystems would be
unable to provide, at high rates, such goods and servic-
es to humankind as absorbing and breaking down pol-
lutants, storing and cycling nutrients, forming soils, and
A
maintaining soil fertility. The numbers and kinds of
fuels, construction materials, and medicines we obtain
from nature depend upon the evolutioD.ary distinctness
of species. Finally, the aesthetic benefits we receive from
nat~re are strongly correlated with the variety of living
;
organisms with which we interact. No matter how many
species of beetles there may be, they cannot substitute
aesthetically for mammals, fishes, corals, or butterflies.
Diversity of Biological Communities
The distinctness of biological communities is defined by
the species within them and their interactions. The species
composition of ecological communities changes over
space because each species has unique adaptations to its
physical and biological environment. As we move across
a landscape, species will be present according to their tol-
erance for the physical conditions that vary spatially (tem-
perature, salinity, light levels, soil chemistry, etc.), and if
they are able to survive their interactions with other
species. Community membership is never absolute, al-
though suites of species may be commonly found togeth-
er under similar conditions.
The number of species ill a location can be determined
simply by counting them, but a simple tabulation does
not serve all purposes. The commonly used measures fall
into three major categories that emphasize, respectively,
numbers, evenness, and differences. The number of
species of organisms present in an area, habitat, or evolu-
tionary lineage, is called species richness. Measures of
evenness, or species diversity indices, weigh species by
some index of their importance, such as their abundance,
productivity, or size. For a given number of entities, the
highest value of evenness is obtained if all are equally
abundant. The degree of difference among species (or
populations, or biotas) is measured by indices of similar-
ity. These indices are also used to assess the degree of ge-
netic similarity among evolutionary lineages, or the di-
versity of habitat types across landscapes or ecosystems.
Indices of both diversity and species richness are
commonly used in ecological, biogeographical, and con-
servation studies because each type gives useful infor-
mation not provided by the others. Ecologists commonly
use diversity measures to assess the adverse effects of
pollution and other types of environmental distur-
bances. Typically a stressed ecological community expe-
riences losses of species and increases in abundance-
and hence dominance of a few species. Multimetric
biological indexes quantify diverse biological changes,
and an examination of the nature of the changes yields
clues as to their causes (for example, the Index of Biotic
Integrity, widely used to evaluate stream condition; Karr
1991). Understanding causes is essential for the design of
management plans that have the potential to counteract
negative effects of stresses and restore the systems to
their former states.
Conservation biologists usually use unweighted meas-
ures of species richness because the many rare species that
characterize most biotas are often of greater conservation
interest than the more common ones that dominate

weighted indices of diversity. In addition, because accurate
estimates of population densities on geographic scales sel-
dom are available, species lists are the only available infor-
mation for most areas.
However, species richness is a poor indicator of dif-
ferences among biological communities in the degree to
w hich they retain significant species interactions. Nei-
ther the abundance nor the functional roles in a commu-
nity ate indicated through a total species count, and thus
w e do not have an indication of which species play sig-
nificant roles or those that play lesser ones. Further,
species richness does not differentiate between native
and nonnative species, and the species richness of de-
graded communities may be increased initially by an in-
flux of nonnative species. Finally, we may most wish to
know the role and importance of variability in commu-
nities, the dynamical responses of communities to dis-
turbance, or several other factors that influence both the
current status of a community or its ability to recover fol-
~owing disturbances. The extent to which communities
can recover from disturbance on their own, or may re-
quire intervention such as ecological restoration, is im-
portant for conservation biologists to understand. Yet,
other measures of species diversity, though they might
indicate differences in the degree of dominance of the
community by one or more species, also can not distin-
guish among these ecologically important differences.
-nfortunately, at present we do not have many good in-
dicators of the functioning of these more complex as-
pects of biological communities (see Chapter 18).
Biological communities are of conservation interest
largely because all species are influenced by their interac-
:ions with other species. Evolutionary change takes place
:n the context of ecological communities, and thus alter-
ations to communities will change the course of evolution.
_ lore immediately, changes in the abundance or presence
of individual species can have strong effects on the re-
:::taining species in a community, which can cause species
~ tinctions or other disruption (see Chapter 3). Thus, con-
servation of biological communities may be a necessary
means to conserving species and evolutionary processes.
Ecosystem and Biome Diversity, and the
orld's Ecoregions
I
Oassification of terrestrial ecological systems typically has
:..een based on the shapes and life-forms of the plants that
~ ominate the structure of those communities (von Hum-
~oldt 1806; Raunkaier 1934; Dansereau 1957; Halle et al.
:978). Koppen (1884) even used plant life-form distribu-
__;on to define climates. Holdridge's (1967) widely used life
zone system, on the other hand, is based entirely upon cli-
~..atic variables. The diversity of schemes reflects the var-
:ed goals of biologists who classify large ecological units.
I
Global Biodiversity 37
Although ecological communities typically gradually
grade into one another, recognition of major divisions is
useful for analysis. Based on the composition of the bio-
tas of different regions, Earth has been divided into eight
major biogeographic realms: the Palearctic, Afrotropic,
Indo-Malay, Australasia, Oceania, Nearctic, Neotropic,
and Antarctic (see Plate 1). Each of these biogeographic
realms contains a number of biomes, large ecological
units that are identified on land on the basis of the dom-
inant type of vegetation (see Plate 1), and, in the sea, on
ocean currents and spatial patterns of primary produc-
tivity (Figure 2.3).
Terrestrial biomes change along environmental (pre-
cipitation, elevation or temperature, and latitudinal) gra-
dients (Figure 2.4). At temperate latitudes, commonly
recognized biomes change along a precipitation gradient
from mesophytic forest, to woodland, tallgrass prairie,
shortgrass prairie, and finally, desert. With increasing
latitude, mesophytic forests change into conifer-domi-
nated boreal forests, and eventually tundra. In tropical
regions, biomes change along precipitation gradients
(rainforest, evergreen seasonal forests, dry forests, thorn
woodland, desert scrub, and desert) and elevational gra-
dients (rainforest, montane rainforest, cloud forest, elfin
woodland, paramo). Finer divisions of biomes into
ecosystem types are based on drainage, soil type, slope,
and species composition. Classifications of ecosystems
and biomes are inevitably somewhat arbitrary, but they
identify ecological units that are useful for a variety of
purposes. Moreover, the richness of human experience
depends upon the richness of biomes as well as the rich-
ness of species.
Recently, a group of scientists led by the Conservation
Science Program at the WWF-US reclassified Earth's bio-
mes into 867 terrestrial ecoregions (see Plate 2; Olson et
al. 2001). An ecoregion is a relatively large area contain-
ing a distinct assemblage of natural communities and
ecological conditions, characterized by a dominant and
widespread assemblage of species. With contributions
from over 1000 scientists, the WWF map of ecoregions
has become the most comprehensive description of
higher-level landscape diversity available, and is a wide-
ly accepted base map for conservation planning.
Preservation of representatives of all habitat types in
all ecoregions is necessary for .preservation of species,
because species will not survive without sufficient quan-
tities of their natural habitats. Captive propagation can,
and does, serve a role in keeping species alive for short
periods until they can be reintroduced into the wild (see
Chapter 15). But captive propagation is of little ultimate
use if there are no suitable habitats into which to reintro-
duce the species. Managing species in zoos and botani-
cal gardens is expensive, and an animal in a cage or a
plant in a garden lives isolated from its natural physical
and biological environment.
 38 Chapter 2

@) ·.

1. East Bering Sea 14. Patagonian Shelf 27. Canary Current 40. Northeast Australia 53. West Bering Sea
2. Gulf of Alaska 15. South Brazil Shelf 28. Guinea Current 41. East-Central Australia 54. Chukchi Sea
3. California Current 16. East Brazil Shelf 29. Benguela Current 42. Southeast Australia 55. Beaufort Sea
4. Gulf of California 17. North Brazil Shelf 30. Agulhas Current 43. Southwest Australia 56. East Siberian Sea
5. Gulf of Mexico 18. West Greenland Shelf 31. Somali Coastal Current 44. West-Central Australia 57. Laptev Sea
6. Southeast U.S. Continental Shelf 19. East Greenland Shelf 32. Arabian Sea 45. Northwest Australia 58. Kara Sea
7. Northeast U.S. Continental Shelf 20. Barents Sea 33. Red Sea 46. New Zealand Shelf 59. Iceland Shelf
8. Scotian Shelf 21. Norwegian Sea 34. Bay of Bengal 47. East China Sea 60. Faroe Plateau
9. Newfoundland-Labrador Shelf 22. North Sea 35. Gulf of Tailand 48. Yellow Sea 61. Antartic
10. Insular Pacific-Hawaiian 23. Baltic Sea 36. South China Sea 49. Kuroshio Current 62. Black Sea
11. Pacific Central-American 24. Celtic-Biscay Shelf 37. Sulu-Celebes Sea SO. Sea of Japan 63. Hudson Bay
12. Caribbean Sea 25. Iberian Coastal 38. Indonesian Sea 51. Oyashio Current 64. Artie Ocean
'
13. Humboldt Current 26. Mediterranean 39. North Australia 52. Sea of Okhotsk

Figure 2.3 Large marine ecosystems of the world and their associated major water-
sheds. These ecosystems account for the majority of ocean productivity due to their
proximity to land masses. However, this view neglects other important marine bio-
mes, particularly pelagic zones making up the majority of the ocean basins. (Modified
from map created by NOAA and the University of Rhode Island, 2002.)
-15
Arctic-
alpine
-10 Tundra

-... -5
u
...._
0

Q)
Cold ::l
~
0
temperate ro .f.j

~
Q)
0... 5
s
Q)

-.f.j

ro
;:$ 10
Figure 2.4 Biomes and cli-
§
ro mate. Distributions of the
~ 15 major biomes are plotted on
ro
Q)

~ axes of mean annual precipi-

Warm 20 tation. Within the region
temperate
bounded by the dashed line,
25 factors such as seasonality of
Tropical drought, fire, and grazing
30 strongly affect which type of ·
0 so 100 150 200 250 300 350· 400 450 vegetation is present (Modi-
Mean annual precipitation (em) fied from Whittaker 1970.)

Ecosystems are often a conservation target because
--.:tical functions that support human populations are
Ellilintained by this level of biodiversity. As ecosystems
-:e degraded by human activities, their ability to provide
,...__:tical ecosystem services and to support a broad array
: iological functions is compromised, as we discuss in
.: e:ail later in this chapter. Biomes across the world are
-~g modified at different rates, and by different forces
see Chapter 6). Our focus at the biome level is often to
.. . . ._. erstand the drivers of change in these areas, and how
u:r options for mitigating these changes may be used for
... ~rvation (Millennium Ecosystem Assessment 2005).
Species Richness over Geological Time
_-:3 number of species on Earth at any moment in time
~ ~ e result of the difference between the rate of produc-
LE 2@2 Earth's Geological History
Era
Cenozoi
Global Biodiversity 3.
tion of new species and the rate of loss of existing
species. The rate of species formation depends both on
events that generate barriers to gene flow (vicariance
events), and on the number of species the more
species, the more ranges there are to be separated, the
more species available to cross existing barriers, and the
more opportunities for divergence. Rates of loss of
species depend on the kinds and severity of distur-
bances, and the strength of factors that permit species to
coexist. Many such factors exist but their operation de-
pends fundamentally on the existence of trade-offs in the
abilities of species to deal with the factors that influence
their abundance and distributions.
The fossil record, though very incomplete, provides a
rough measure of trends in species richness during the
history of life on Earth (Table 2.2). Cellular life in the form
of bacteria evolved about 3.8 billion years ago (bya); eu-
 40 Chapter 2

karyotic organisms probably evolved about 2 bya. Al-
though many species must have lived in ancient times,
species richness appears to have been low during the first
2 billion years of Earth's existence. During the late Pre-
cambrian, the richer Ediacaran fauna, consisting of
strange frond- and disc-shaped, soft-bodied animals and
some forms that appear to be arthropods and echino-
derms, evolved. The first explosion of biodiversity took
place during the early Cambrian period. Some Cambrian
species may be members of phyla that left no surviving
descendants. As measured by the number of phyla, ani-
mallife may have been more diverse during the Cambri-
an period than at any time since (Gould 1989).
The fossil record for marine invertebrates with hard
skeletons is good enough to provide a general picture of
the number of evolutionary lineages present at different
times in the past. The Cambrian explosion was followed,
about 60 million years later, by the extensive radiation of
proximately 40,000 species of marine invertebrates have
been described from the Paleozoic and Mesozoic eras, a
number that increased to about 250,000 in the late Ceno-
zoic era. The fossil record of terrestrial animals is much
poorer, especially among such species-rich groups as in-
sects, which fossilize poorly. The fossil record of verte-
brates, particularly mammals, is much better. It indicates
th at richness, as measured by the number of orders, is
slightly higher today than earlier in the Cenozoic.
Terrestrial vascular plants appeared by the early Sil-
urian and their richness increased rapidly during the De-
vonian when seed-bearing plants first appeared. Species
richness has continued to increase overall but the num-
bers of species of ferns and gymnosperms has decreased,
whereas the number of species of angiosperms has in-
creased dramatically (Figure 2.6).
Rates of species formation
•

the Paleozoic fauna. Following the Permian mass extinc- Speciation rates have varied greatly over evolutionary
tion, the modem fauna evolved, and overall family rich- time. Large numbers of new evolutionary lineages origi-
ness has increased steadily throughout the Mesozoic and nated three times during the history of life. The first event,
Cenozoic eras to a maximum today (Figure 2.5). Ap- known as the Cambrian explosion, took place about half
a billion years ago. The second, about 60 mi11ion years
later, resulted in the Paleozoic fauna. Biodiversity was
Present
greatly reduced 300 million years later by the great Per-
()
@ mian mass extinction, which was followed by the Triassic
0
N
0
explosion that led to our modem biota. Although all three
~·
(')
of these explosions resulted in many new species, they
were qualitatively very different. Virtually all the major
100

~
ro
00
0 700
N
0
~·
Jurassic (')

200 600
0
00
ro Triassic
rJ)
~
ro
(!)
::>... Permian
* 500 Angiosperms
'-4ool rJ)
0 Q)
.u"""
~ 300 . '
(!)
0 0.. 400
•
."""
...-4
Carboniferous
rJ)

."""
...-4 '-4ool
0
~ ~
(!)

D evon1an
. ~
~::s 300

400
~
ro
0
z
N 200 Gymnosperms
Silurian 9.
(') •

100
Ordovician
500
Cambrian S D c P Tr K T
0 200 400 600 800 1000 439 409 360 296 251 206 144 65 2
Number of marine families Millions of years before present

Figure 2.5 Diversity of marine families from the Cambrian Figure 2.6 Terrestrial plant species richness. Ferns, gym-
to the present. The asterisks mark the five major mass extinc- nosperms, and angiosperms have, in turn, dominated the
tion events. world's flora. (Modified from Signor 1990.)
 Global Biodiversity 41

groups of living organisms appeared in the Cambrian pe-

riod, along with some phyla that subsequently became
extinct. The Paleozoic and Triassic explosions greatly in- 60
creased the number of families, genera, and species, but *
:10 new phyla of multicellular organisms evolved.
High rates of speciation have been favored by three .....u 40
.5
.....
:5actors: mass extinctions, increasing separation of land ><
Q)

:nasses across Earth, and evolution of new life forms and .....
~
Q)
20
* * '

*
types of species interactions. Mass extinctions tend to be u

::allowed by increases in rates of speciation because ~

:hose lineages that survive inherit a biologically depau-
perate Earth, an ecological setting favorable for the evo-
~ution of many different life-forms and body plans. In-
8"easing provinciality, such as occurred during the
breakup of Pangaea and, later, Gondwana, stimulated
speciation, because foll~wing their separation, evolution
on each of the new continents produced many new en-
demic species. Much of the Cenozoic era increase in
:1umbers of species appears to be due to such provin-
d alization. South America, Africa, and Southeast Asia
are all rich in species, but they share few species or gen-
era because they have been separated from one another
ior many millions of years. Increasing complexity of eco-
-rogical interactions also has stimulated rates of specia-
tion. On land, the richness of vascular plants increased
dramatically with the evolution of angiosperms and
:heir complicated interactions with animals during re-
?roduction and dispersal of seeds. The evolution of
flight undoubtedly also contributed to the great diversi-
ty of both insects and birds by allowing better exploita-
tion of the third dimension on land. In the oceans, diver-
sity was stimulated by evolution of organisms' ability to
burrow into sediments and to swim in open water. The
number of species living together also increased as a re-
sult of finer adaptations to particular environmental con-
ditions and ways of exploiting the environment made
possible by minor variations in morphology, physiology,
and behavior.
Rates of extinction
Extinctions have occurred at all times throughout the
history of life, but rates have changed dramatically. Pa-
leontologists distinguish between "normal" or "back-
ground" extinction rates and the much higher rates as-
sociated with mass extinctions (Sepkoski and Raup 1986;
Figure 2.7). The first of six mass extinctions, at the end of
the Cambrian period, destroyed about half of the known
animal species. About 75°/o of species became extinct in
the second event at the end of the Devonian period. The
most extreme of the mass extinctions, at the close of the
Permian period eliminated about 95o/o of both marine
and terrestrial organisms. In the oceans, trilobites de-
clined to extinction and brachiopods almost became ex-
tinct. On land, the trees that formed the great coal forests
~
Q) *
C 0 S D Crb P Tr Jur Cret Tert
Paleozoic Mesozoic Cen
500 400 200 0
Time (mya)
Figure 2. 7 Extinctions of families through geologic time.
The five historical mass extinction events are marked with an
asterisk.
became extinct, as did most lineages of amphibians. At
the end of the Triassic period, nearly all ammonites and
approximately 80°/o of reptile species vanished, resulting
in an overall loss of about 65°/o of species. The fifth ex-
tinction event at the end of the Cretaceous is most famous
because dinosaurs and other large reptiles became extinct
during that period, along with most marine lineages and
about 75o/o of all species. Discussed in greater detail in
Chapter 3, the current and sixth mass extinction event,
promulgated by human expansion over the planet, ini-
tially exterminated large mammals and island species,
but if current trends continue, organisms of all sizes and
lineages will be seriously affected.
Following each of the previous mass extinctions, bio-
diversity expanded, but species richness did not reach its
previous value until after an average lag of about 10 mil-
lion years, and in some cases much longer (Kirchner and
Weil2000; see Figure 2.5). However, even with such long
lags, species richness has not been directly affected by
mass extinction events during most of geological histo-
ry. Their major effect has been to eliminate some lineag-
es, thereby making ecological room for other lineages
that proliferated (over geological time intervals) follow-
ing episodes of mass extinction.
Current patterns of species richness
Earth is not uniform and neither is the distribution of or-
ganisms across its surface. Some important general pat-
terns in the geographical distribution of species richness
have been discovered, but much remains unknown, in
large part because the inventory of living organisms is so
incomplete. Also, the distributions of species are best
known for temperate regions where most taxonomists
and ecologists live and work. Tropical regions, where
 ,
42 Chapter 2

most of the world's species live, are poorly known bio-
logically. Nonetheless, some places on Earth clearly have
exceptionally high biodiversity (tropical rainforests,
coral reefs) whereas others are virtually devoid of life
(extremely dry tropical and polar deserts) (Figure 2.8).
These patterns are discussed in greater detail in the later
section, "Latitudinal Gradients in Species Richness."
Much of the interest in describing and interpreting
current patterns of distribution of biodiversity has been
stimulated by a growing concern about the future of bio-
diversity on Earth (Gaston 2000). If efforts to preserve
Earth's biodiversity are to be successful, they must be
based on accurate information on where species are
found, why they are there, and what is threatening their
continued existence. Recently, there has been a revolu-
tion in the quantity of new data on distributions and in
the development of new technical tools, such as distri-
bution-mapping schemes (i.e., geographic information
systems or GIS) and remote-sensing technology, with
which to analyze complex data sets.
To describe spatial patterns of biodiversity, ecologists
and biogeographers have found it useful to divide
species richness into three major components: alpha (a)-
richness, beta(~) -richness, and gamma (y) -richness (Fig-
ure 2.9). Alpha-richness refers to the number of species
found in a small, homogeneous area; ~-richness refers to

(A) 22,000

20,000

18,000 Amphibians
lr --. , I Birds
16,000
U)
UiiU Mammals
Q)
• !""'( I I Reptiles
u
Q)
14,000
~
U)
......... 12,000
0
H
Q)

~;j 10,000

z 8000

6000

4000

2000

(B) 9000

8000 Figure 2.8 Species richness (A) and endemism (B) of mammals, birds, amphib-
ians, and reptiles is greatest in tropical moist forests, and lowest in deserts. TMF,
7000 tropical and subtropical moist broadleaf forests; TDF, tropical and subtropical dry
U)
Q)
broadleaf forests; TCF, tropical and subtropical coniferous forests; TeBF, temperate
• !""'(
u
Q) 6000 broadleaf and mixed forests; TeCF, temperate coniferous forests; BF, boreal
~
U)
forests/taiga; TG, tropical and subtropical grasslands, savannas and shrublands;
u TeG, temperate grasslands, savannas, and shrublands; FG, flooded grasslands and
s
•!""'(
5000
Q) savannas; MG, montane grasslands and shrublands; T, Tundra; MF, Mediterranean
'\:)
c: forests, woodlands, and scrub; D, deserts and xeric shrublands; M, mangroves.
Q)
.........
4000 (Modified from Millennium Ecosystem Assessment 2005.)
0
H
Q)
3000
~;j
z 2000

1000

~ ~"'- "'-6 4f ""§- <);)"'- "'-0 ""p "'-0 .$' "" $ ~

Biome

40 I I I 4
----
~
X;- 30
--
-
! ;...
~
-§
_::)
20
--
:;
> (l)
------
:-:
----
10
o~~--~----~------~----~----~----~
0 0.4 0.8 1.2 1.6
. Habitat gradient
· figure 2.9 Species turnovers along habitat gradients. (A) Species accumulation
· ·.. :urves for birds across habitat gradients in Mediterranean vegetation (from dry scrub
:o ¥voodland) in southern Africa, California, and Chile. (B) The differentials of the
curves in (A) give the rate of species turnover across the gradient. (Modified from
Cody 1975.)
.
the rate of change in species composition across habitats
r among communities; and y-richness refers to changes
across larger landscape gradients. A high B-richness
m eans that the cumulative number of species recorded
r apidly increases as additional areas are censused along
some environmental gradient. Species may also drop out
rapidly along such gradients, resulting in a high rate of
species turnover. Susan Harrison and Jim Quinn discuss
some of the ways in which studies of B-richness enhance
our understanding of regional ecology, and inform con-
servation planning in Box 2.3.
The rate at which the species composition of commu-
nities changes across environmental gradients is deter-
nlined by the sizes of species ranges and the degree to
\-hich species are habitat specialists. The ranges of trop-
~cal species are much less known than the ranges of their
ternperate counterparts, but on average terrestrial tropi-
cal species have smaller ranges than species of higher
. . atitudes. The altitudinal ranges of species on the slopes
of tropical mountains appear to be narrower than ranges
of species on temperate mountains, but this impression
m ay be an artifact of inadequate sampling of rich tropi-
cal biotas (Colwell and Hurtt 1994).
An inevitable and important corollary of patterns of
species richness is that areas with high a-richness in-
evitably have many rare species. A tropical wet forest in
South America or Southeast Asia, for example, may har-
bor between 300 and 400 species of trees per square kilo-
m eter, whereas a temperate forest harbors an order of
magnitude less. However, the number of trees per hectare
is roughly the same in tropical and temperate forests. It
follows that most of the tree species in tropical forests
Global Biodiversity 43
•
(B)
Africa •
I
I 1
Africa
California
\•
Chile
California
.u
..-(
~ Chile
cJ)
c 10
~
(l)
1
z o~----~----~----~------~----~----~
2.0 0 0.4 0.8 1.2 1.6 2.0
Habitat gradient
must be present at very low densities. Many of those
species are more abundant elsewhere, but some species
evidently are present only at low densities throughout
their ranges.
The following sections describe some major patterns
in the distribution of species and discuss mechanisms
that may generate these patterns. An understanding of
both the patterns and the mechanisms that generate and
maintain them can be used to develop strategies to pro-
tect or recover biodiversity.
Patterns of Endemism
A species that is found in a particular region but nowhere
else is said to be endemic to that region. However, what is
an appropriate spatial scale for assessing endemism varies
greatly. All species are, as far as we know, endemic to
Earth. At the opposite extreme, some species are restricted
to single desert springs, small islands, or isolated moun-
tain tops. Regions with many endemic species are the re-
sult of one or more major events that caused the ranges of
many taxa to separate at approximately the same place.
Causes of such geographical isolation include continental
drift, mountain building, climate change, and sea level
rises. Following such isolation, many taxa may undergo
evolutionary radiations in the same general area. Vicari-
ance due to continental drift has been extremely important
in generating the high degrees of endemism found in the
biotas of Madagascar, Australia, New Guinea, and New
Caledonia. Owing to their isolation, islands often have
high proportions of endemic species, but, given that is-
lands often have relatively impoverished biotas, high
,:100
 •
lo a iod· rs1 45

a
A
wh
due

demism ofte ·s not ssociated w·th · spe ies rich- Patt r s o end mi m i fe reatly mong t xa
Less. Cor 1 efs assoc·ate with ·sl s, howeve, ar an ( ble 2. ). th h a e r i n of ou h Africa d
""". . . .""~ti n. e island- tudde west m ac · ·c e is . . ,. .-'\ sou este Aus a ·a a e x ely hig n ers of
exampl ofar gion fm · p · ri s d n- ndemi plant sp cies bu ver few end ic a s
--.. . . ·sm for corals, reef · es, snails, and 1 sters ( i o birds ( owlin 92; y eta . 200 ). These differ-
2.10; Bri gs 1996; Ro erts et 1. 2 02). Fin y, con · ent 1 ence aris e au pl t c s eciate vi ol loi y ·
"-'LL'--"~
• e Tropi of n c ntain both highs eci s ·ch- uc s lie areas t can e e rates, a ong w ich
n and high degr fend · m (s i e 2.8). polyploidy is r re. o ever, t ere are ron cor la-
46 Chapter 2

Figure 2.10 The Indo-West Pacific

is a marine diversity hotspot. The
distribution of species richness for
damselfish (Pomacentridae) shown
here is typical of many marine taxa
in the region. (From Briggs 1996.)

68 30

Scale
0 1000 2000 Miles
I, , , , J,, , ,I,,,,J , , , ,J
III I I II II I pI IIJI II I III II jilill
0 1500 3000 Kilometers
(True distances on mid-meridians and parallels 0° to 40°)

TABLE 2.3 Numbers of Endemic Species of Plants, Mammals, Reptiles and Amphibians in Regions
0
of Exceptional Degrees of Endemism

Source: Data from Meyers et al. 2000.

a1°/o or greater of global total for endemic plants or vertebrates.
bSize (0/o) refers to remaining primary vegetation in km2 and to percent remaining of original extent.
 Global Biodiversity 47

tions among patterns of endemism in mammals, birds, 1000

and reptiles, all of which require relatively large areas for
geographic speciation. s

Latitudinal Gradients in Species Richness

In both marine and terrestrial environments tropical re- • • • •
gions harbor many more species in most higher taxo-
n omic categories than higher-latitude regions. For exam- ~
Q)
• G
-~---Q
ple, Arctic waters have about 100 species of tunicates, 400 ~
H 100 ----
0
u 'C\
q., .... ' 0 0 "a
species are known from temperate waters, and more than z •
4
o/ "'~
0
°
°
F
',
Q\
600 species inhabit tropical seas. The richness of species,
e
. I
/ . \
\
genera, and families of bivalve mollusks peaks in tropical • I
1 0
~ ® Gte
•
• •
\
\
regions and declines rapidly with increasing latitude I \
foo b
(Figure 2.11). A similar pattern is found among the fauna o,a
I ~
living on hard substrates (Thorson 1957) and benthic liv- •
ing and fossil foraminiferans (Buzas and Gibson 1969;
Stehli et al. 1969). The number of ant species found in
local regions increases from about 10 at 60°N latitude, to 10~----~----~----~----~----~----~
90N 0 90S
as many as 2000 species in equatorial regions. Greenland
Latitude
h osts 56 species of breeding birds, New York, 105,
Guatemala 469, and Colombia 1395. Latitudinal gradi- Figure 2.11 Latitudinal species richness in bivalve mollusks.
ents in species richness of birds and mammals in North Points are average numbers of species (S), genera (G), and
and Central America are illustrated in Figure 2.12. families (F). (Modified from Stehli et al. 1969.)
There are relatively few exceptions to these latitudinal
p atterns, in which highest richness occurs in mid- or
high latitudes, or there is no correlation with latitude. positional variable, cannot by itself be a determinant of
However, most of these cases occur when the scale of species richness patterns. Because of their differing re-
analysis is smaller (<20° latitude), or a lower taxonomic quirements, it seems likely that different combinations of
category is examined (e.g., orders). But parasites, ich- factors may determine the distributions of bacteria, fungi,
neumonid parasitoid wasps, and aquatic plants (re- vascular plants, amphibians, and mammals, yet some
viewed in Willig et al. 2003) all appear to be exceptions suite of factors that co-varies with latitude must operate
even at large scales of analysis. Seaweeds show no con- to increase species richness at low relative to high lati-
sistent pattern in relation to latitude (Bolton 1994). Also, tudes. Differences in species richness must be generated
the richness of marine birds and mammals is greater at by differences in the rates of speciation, extinction, immi-
high latitudes (Willig et al. 2003). gration and/or emigration among locations. Because lati-
Although latitudinal gradients in species richness tude cannot directly influence speciation, extinction, or
have been known for a long time, determining their caus- movement rates, it must be a surrogate for some combi-
es has proven difficult. For a starter, latitude or any other nation of variables that can directly influence species

Breeding bird species Mammal species

120 ~ so

150
190 . 80
200 100
210
200 110
Figure 2.12 Latitudinal gradients of
. 220 110
species richness of birds and mam-
~ 280 140 mals in North and Central America.
360 150 Species richness corresponds to lati-
480 150 tude map at left. The numbers for
600 150 birds are from breeding species only.
660 160 (Modified from Briggs 1996.)
48 Chapter 2

(A ) (B)
1000 1000

Hispanola
• •
r.f)

.~ 100 Puerto •Cuba •• • •

u
Q)

~
Rico
100 • •• •• • ••
........
Jamaica Sunda Islands
~
0
•
Q)

~;J 10
• •
z
Montserrat 10 •
Saba
Redonda

1 ~----~----~----~----~----~ 1~----~------~------~------~----~
10 100 1000 10,000 100,000 100 1000 10,000 100,000

Figure 2.13 Relationship between area and number of species on islands of various
sizes. (A) The number of species of amphibians and reptiles found on selected islands
in the West Indies. (B) The number of land and freshwater bird species on the Sunda
Islands, the Philippines, and New Guinea. These islands are close to the Asian conti-
nent, and many were connected to the mainland during glacial periods. Therefore,
many of the larger islands are relatively species-rich. (Modified from MacArthur and
Wilson 1967.)

richness and that are correlated with position. One likely
correlated factor is area, which can influence both speci-
ation and extinction rates.
One of the first ecological relationships to be estab-
lished empirically was the relationship between area and
number of species (Arrhenius 1921). This relationship~,
which is true for both continental and island biotas
(Rosenzweig 1995), is commonly expressed using a
power function of the form
2 and McKay 2004). Therefore, area is probably a contrib-
S = cA
where S is the number of species, A is the area, and c and
z are constants fitted to the data. On a logarithmic scale,
this relationship plots as a straight line where c is the y-in-
tercept and z is the slope of the line (Figure 2.13). Analy-
ses of species-area relationships in many groups of or-
ganisms revealed that values of z range from 0.15 for
continents, to between 0.25 and 0.45 for islands associat-
ed with a continent, to 0.19 for comparisons among con-
tinents or biogeographic provinces (Rosenzweig 1995).
The regular relationship between the size of an area and
the number of species it supports, which is most readily
observed using island data, was a key empirical general-
ization in the development of the theory of island bio-
geography (MacArthur and Wilson 1967).
The Tropics have a larger climatically similar area
than any other ecological zone because the surface area
of latitudinal bands decreases toward the poles and be-
cause temperatures change relatively slowly between
the equator and 20°N and 20°5 latitudes. Larger area is
correlated with higher rates of speciation and lower
rates of extinction, and thus, also is correlated with high
species richness. Large areas can facilitate the long-term
survival of populations, which in tum, allows the accu-
mulation of the genetic differences that may prevent hy-
bridization should the populations come together at
some time in the future. This favors greater speciation
rates. Populations of vertebrates in lower latitudes show
greater genetic differentiation, which is consistent with
greater capacity for speciation in the Tropics (Martin
utor to high species richness in the Tropics, but so are
productivity and available energy, as discussed in the
next section.
Species Richness-Energy Relationships
Available energy affects local species richness because the
more energy that is available, the more biomass per unit
area that can be supported. More biomass enables more
individuals to coexist in an area, potentially resulting in
more individuals per species and, hence, lower extinction
rates. If high productivity is combined with moist tropi-
cal conditions, organisms expend relatively little energy
maintaining their temperatures and moistures at appro-
priate levels, so even more energy is potentially available
for reproduction (Connell and Orias 1964).
More available energy might also enable species to
persist on relatively specialized diets (Gaston 2000).
There is some evidence that tropical animals may, on av-
erage, have more specialized diets than their temperate
counterparts, but much more research is needed to es-
'\

:ablish the degree to which this is true and to identify the
~-:-:mensions along which dietary specialization has
eYolved (Beaver 1979; Marquis and Braker 1994).
Primary production,-the major source of available en-
=::g)" is determined primarily by temperature, moisture,
~1d nutrient availability. Highest terrestrial production
:S found in regions with high rainfall and year-round
...-ar m temperatures. Production drops with elevation
~ecause of lowered temperatures. In many areas of the
.·.-orld, the amount of precipitation determines total pro-
iuction. In arid and semiarid regions, annual production
:an be predicted fairly accurately from the amount and
~.; tribution of precipitation.
Marine production is also limited by temperature, but
nore often by shortage of nutrients in surface waters
Longhurst 1998). Organisms and nutrients sink in the
- -ater column. Therefore, processes that keep organisms
'-Lose to the surface where photosynthesis is possible, and
:hose that bring nutrients and organisms back to the sur-
:=ace, are major determinants of marine productivity. Pro-
. uctivity is highest in areas where vertical turbulence
a p welling) is produced by friction between tidal streams
and the seabed and at the confluence of two major ocean
currents; it is lowest in the open ocean (Koblentz-Mishke
et al.1970; Bunt 1975; Longhurst 1998). Primary produc-
:i\i ty also is higher near continental margins due to the
:runoff of nutrients from the land. Finally, primary pro-
uction decreases with depth because light levels de-
crease rapidly with depth even in clear water.
The hypothesis that available energy contributes to
species richness is supported by a correlation between
___ roductivity and species richness along both latitudinal
and altitudinal gradients. A strong correlation exists be-
uveen realized annual evapotranspiration, a measure of
10
20
20
40
20
soJ 40
50
Acacia Eucalyptus
Global Biodiversity 49
available energy to plants, and tree species richness in
North America (Currie and Paquin 1987). Similarly, Gen-
try (1988) demonstrated a strong correlation between
plant species richness in Neotropical forests and ab-
solute annual precipitation, a variable strongly positive-
ly correlated with productivity.
However, the relationship between productivity and
species richness is not simple. Many of the world's most
productive systems, such as estuaries, seagrass beds,
and hot springs, are species-poor. Conversely, plant
species richness is higher in semiarid regions with nutri-
ent-deficient soils than in similar areas with richer soils.
The remarkably rich plant communities of the Cape re-
gion of South Africa (fynbos) and the extreme southwest
corner of Australia (Figure 2.14) are found on highly in-
fertile soils (Kruger and Taylor 1979; Bond 1983; Rice and
Westoby 1983). Nitrogen levels in those soils are an order
of magnitude lower than in lowland California and
Chile (Specht and Moll1983), where species richness is
lower. Evidence suggests that soil infertility itself is a
major contributor to the inverse correlation between soil
fertility and plant species richness because low fertility
favors abilities to exploit slightly different microhabitats
more efficiently (Tilman 1982, 1985; Cowling et al. 1992;
Willis et al. 1996). Fynbos plants are remarkably variable
in leaf morphology (Cody 1986) and growth phenologies
(Kruger 1981; Cowling 1992). These differences are prob-
ably correlated with a corresponding diversity of car-
bon-fixing strategies. In addition, the low palatability of
leaves of plants growing on nutrient-poor soils results in
a rapid accumulation of flammable biomass, leading to
high frequencies of fires in these summer-dry climates.
Soil infertility also may favor high species richness in-
directly via interactions with seed dispersers. Compared
Figure 2.14 Numbers of species of
Eucalyptus and Acacia in southwestern
Australia. Notice how many species in
these two genera can be found living
55 in proximity. Similar patterns are
found among many other genera of
plants in southwestern Australia.
(Modified from Lamont et al. 1984.)
50 Chapter 2
to birds, ants move seeds relatively short distances, but
they usually bury them. Also, ants pick up and move
seeds that offer much smaller rewards than those that at-
tract birds. Australia, a continent with notoriously poor
soils, has the highest proportion of plants with ant-dis-
persed seeds of any continent (Berg 1975; Westoby et al.
1991) and it has unusually high ant species richness (An-
dersen 1983; Greenslade and Greenslade 1984). Ant-dis-
persed seeds are also common in South African fynbos
(Milewski and Bond 1982). In combination, these factors
could result in very high a-species richness and unusu-
ally high B-species richness. Species richness on scales of
one hectare is higher in southwest Australia than in rain-
forests even though the latter accumulate more species
at slightly larger scales (Figure 2.15).
There is a consistent unimodal relationship between
species richness and depth for many marine taxa. Maxi-
mum richness of most groups is found at depths be-
tween 2000 and 4000 meters (Rex 1973, 1983). In contrast,
coral species richness peaks at depths between 15 and 30
meters because corals, which obtain much of their ener-
gy from photosynthetic algae embedded in their tissues,
are confined to the photic zone (Huston 1994).
In addition, in many ecosystems, species richness in-
creases with primary production at low production lev-
els, but declines at high levels of primary production
(Huston 1994). This phenomenon has been called "the
paradox of enrichment" (Rosenzweig 1971) because ad-
dition of fertilizer to aquatic and terrestrial plant com-
munities often results in sharp decreases in species rich-
ness (Huston 1980).
1 terrestrial environments, plants provide most of the phys-
400
300
(/)
Q)
• ....C 2 plex communities have a greater variety of microclimates,
u
Q)
~
(/)
~
0 3
200
J.-1
Q)
s ~ 4
z lated with structural complexity. Structurally simple habi-
100 5 tats, such as the open ocean, grasslands, and cold deserts,
o~ -~------------~----------------~
0 0.1 1 2
Area (ha)
Figure 2.15 Plant species richness in Borneo and in south-
western Australia. Curves 1, 2, and 3 represent trees greater
than 10 em in diameter mmixed forests in Borneo. Curves 4
and 5 are for plots in southwestern Australia. (Modified from
Lamont et al. 1984.)
(/)
Q)
•....C
u
Q)
~
(/)
20 40 60 80 100
Salinity (0/o)
Figure 2.16 Estimated species richness of aquatic inverte-
brates living in waters of different salinity throughout the
world. The two peaks correspond to fresh water (<2°/o salinity)
and seawater (30°/o-40°/o salinity). (Modified from Kinne 1971.)
Most highly productive but species-poor systems are
distributed as relatively small, fragmented patches
whose physical environments differ strikingly from those
of the surrounding, more extensive ecosystems. Evolu-
tionary biologists believe that the combination of major
physical environmental differences and isolation of the
patches results in fewer species of organisms evolving
adaptations to those unusual environments. For the same
reason, species adapted to the more common surround-
ing environments are less likely to survive well in those
rare habitat types. Both fresh water and seawater are
widely distributed, but waters of intermediate salinity
and waters that are more saline than the oceans are rare.
The species richness of aquatic invertebrates throughout
the world parallels this pattern (Figure 2.16).
Available energy may influence species richness indi-
rectly via vegetation structure. The structural complexity
of vegetation is strongly correlated with potential evapo-
transpiration and, hence, primary productivity. In most
ical structure within which the activities of all other or-
ganisms are carried out. Coral reefs create complex struc-
tures in tropical marine environments that influence fish
species richness (Holbrook et al. 2002). Structurally com-
a greater variety of resources, more ways in which to ex-
ploit those resources, and more places in which to find
shelter from predators and the physical enviro~ent.
The richness of species in most taxa is positively corre-
generally support fewer species of organisms than struc-
turally more complex communities, such as forests and
coral reefs. Animal groups that exploit the environment in
three dimensions are most sensitive to plant community
structure. A positive correlation between foliage height di-
versity and bird species richness exists in many plant com-
munities on all continents (MacArthur and MacArthur
1961; MacArthur 1964). Similarly, the richness of species of
web-building spiders is positively correlated with the het-

erogeneity in heights of the tips of vegetation to which
such spiders attach their webs (Greenstone 1984).
In contrast, there is no consistent relationship between
v egetation structure and lizard species richness in hot
deserts in North and South America, Africa, and Australia
(Pianka 1986). The number of lizard species in similar
h abitats in African deserts averages twice that in North
.i\merica; Australian deserts are about twice as rich as
African ones. The main differences are due to the presence
m Australia of non-lizardlike lizards and nocturnal
species. Only one nocturnal lizard is found at the North
American sites, whereas there are four nocturnal species
at the African and eight at the Australian desert sites.
Mammal-like species (the monitors) and worm-like
species add to the richness of Australian desert lizard fau-
nas. These differences relate primarily to the long-term
evolutionary history of deserts on the three continents,
not to differences in today's vegetation (Pianka 1986).
Disturbance and Species Richness
•
One explanation for the latitudinal patterns of species
richness comes from considerations of the extreme dis-
turbances caused by climate change over Earth's history
(Dynesius and Jansson 2000). During the Pleistocene and
before, periods of rapid climatic change (influenced by
orbital oscillations of Earth), which were more intense
toward the poles, clearly had enormous influence on
worldwide patterns of species riclmess. Rapid climate
fluctuations caused increases in extinction rate as species
were unable to adapt to new conditions, or migrate to
more hospitable ones, and the more extreme fluctuations
toward the poles resulted in more extinctions at high lat-
itudes. Importantly, these fluctuations also should have
changed patterns of speciation, favoring characteristics
that lead to lower speciation rates among the survivors
0
Global Biodiversity 51
(less specialization, greatervagility,larger ranges), par-
ticularly in higher latitudes that were most affected by
climate change (Dynesius and Jansson 2000). This com-
bination of selective factors in response to climate cycles
could contribute to the latitudinal gradient of species
riclmess we see today (Figure 2.17). At a finer spatial
scale, physical and biological disturbances clearly influ-
ence the number of species present in many ecological
communities. Small disturbances occur much more fre-
quently than large disturbances. Physical perturbations
include heavy rains, strong winds, landslides, earth-
quakes, and fires. Biological disturbances, most of which
are small scale, include activities of predators and para-
sites, tree-falls, activities of competitors, and trampling.
Physical disturbances influence species riclmess by
destroying habitat structure, selectively killing individ-
uals of different species, and sterilizing soils. Physical
disturbances typically also have important indirect ef-
fects on biodiversity by altering the biological interac-
tions that affect species riclmess. For example, fire, one
of the most important terrestrial physical disturbances,
kills plants and animals, destroys soil organic matter,
and redistributes nutrients, resulting in a new ecological
community with a dramatically different physical struc-
ture and altered biological interactions.
Theoretical and empirical studies both suggest that
disturbances do not have consistent effects on species
richness. The hypothesis that lack of disturbances might
favor high species richness was originally advanced by
Sanders (1968), and Sanders and Hessler (1969), who
noted that productive estuaries and continental shelves
in most latitudes have few species of benthic animals,
but the cold, dark, unproductive floor of the deep sea is
very species-rich. Sanders suggested that the deep sea
supported many species because its environments had
relatively constant levels of temperature, salinity, and
Figure 2.17 Sudden climate shifts
influence species richness by chang-
ing speciation and extinction rates.
Selective pressure for species to be
less specialized, more vagile, and
have larger range sizes result in slow-
er speciation rates. Direct effects of
climate change increase extinction .
rates. The net result of these forces is
a decrease in species richness. (Modi-
fied from Dynesius and Jansson
2000.)
52 Chapter 2
oxygen concentration and had been so for millennia.
However, food resources in the deep sea are patchily dis-
tributed, and loc.a l disturbances are caused by the feed-
ing, burrowing, and mound-building activities of ani-
mals (Grassle 1989, 1991). These activities create a
patchiness similar to that produced by tree-falls inter-
restrial environments. They should create conditions
suitable for many different species.
Biological interactions can also influence species rich-
ness. For example, competition could reduce a-richness
if some species exclude others from ecosystems. How-
ever, competition could increase a-richness by favoring
finer habitat segregation among species (Rosenzweig
1995). Predation can increase species richness if preda-
tors prey preferentially upon competitive dominants,
thereby preventing competitive exclusion. A well-known
example is the increase in species richness in rocky in-
tertidal communities of the Pacific Coast of North Amer-
ica. This increase is a result of selective predation by the
sea star Pisaster ochraceus on the competitively dominant
mussel Mytilus californianus (Paine 1974). On the other
hand, predation can reduce species richness by prevent-
ing vulnerable species from living in an area. Experi-
mental evidence exists for all of these outcomes, but how
often and where these forces exert their influence are un-
known (Orians and Kunin 1991). Analyzers of the litera-
ture have reached different conclusions, in part because
they used different criteria for including studies in their
samples (Connell1975, 1983; Schoener 1983).
Some researchers have hypothesized that many organ-
isms are held at sufficiently low population densities such
that competition rarely occurs (Connell1975). Yet other re-
searchers have suggested that competition occurs prima-
rily during unusually hard times, "competitive crunches,"
when resources are scarce (Wiens 1977). Which of these
two scenarios applies depends upon whether low popu-
lation densities are caused by harsh physical conditions or
by scarcity of consumable resources.
Often species richness is highest at intermediate lev-
els of disturbance (Figure 2.18). The Intermediate Dis-
turbance Hypothesis states that physical disturbances
or predation should augment species richness where
other sources of disturbance are few, but should reduce
species richness when exogenous disturbance is high
(Connell 1978; Abugov 1982). If disturbances are rare,
species richness may decline owing to competitive ex-
clusion. If disturbances are common, many species may
be unable to complete their life cycles during the short
intervals between disturbances.
Interactions between local and
regional species richness
The structure of local ecological communities may be de-
termined by loc~ interactions among species and their en-
vironments, and by processes operating at larger spatial
Low High
Disturbance frequency or intensity
Figure 2.18 Model of the intermediate disturbance hypothe-
sis. Species diversity is lowest at high and low levels (fre-
quencies or intensities) of disturbance, and highest at an in-
termediate level. (Modified from Connell1978.)
and temporal scales (MacArthur 1972; Ricklefs 1987). In
other words, to understand why a particular set of species
exists in a specific place, ecologists must study more than
local competition, predation, and mutualistic interactions.
Larger-scale processes, such as dispersal, speciation, and
historical biogeography cannot be ignored.
The ecological interactions that appear to influence
current patterns of species richness, in combination with
physical environmental factors, could set absolute limits
to species richness. In other words, we can ask if there
are limits to the number of species that can live in a
given ecological community. If so, are those limits regu-
larly reached? Is there a critical limit to overlap in re-
source use that prevents more species from being ac-
commodated unless dramatic changes occur in the
nature of available resources? Does the chemical warfare
between plants and herbivores set limits to species rich-
ness or does it offer possibilities for more species? Are
there limits to the size of mimicry systems, and do mim-
icry systems allow more species to persist in ecosystems
than would be possible without them? Have the rich-
ness-generating interactions between plants and their
pollinators and between plants and their seed dispersers
been exhausted? To answer these questions research
must be carried out at many different spatial scales. .
For many diverse taxa, including gall wasps (Cornell
1985), birds (Ricklefs 1987), tiger beetles (Pearson and Ju-
liano 1993), fishes (Griffiths 1997), and primates (Eeley
and Lawes 1999), local species richness appears to in-
crease along with regional species richness, but at a slow-
er rate. This pattern suggests that rigid limits to local
species richness are not set by competition, predation,
and parasitism (Comell1999). Thus, although ecological
interactions are typically very intense in local communi-
ties they apparently influence abundances more strongly
than numbers of species. Moreover, their influence on the
numbers of coexisting species appears to be weaker than
the influence of regional processes. A major challenge in
ecological research is to determine how regional process-
es act to determine local species richness.
 Global ~iodiversity 53

The importance of biodiversity

100
Ve asserted in the first chapter that a normative postu- --- --- --- --- I

late of conservation biology is that biodiversity is worthy .. .... ..

A ......
.... --- ...
I

80 .... .. .... I
I

of being conserved for its own sake. Yet, many people do .... .... I

:1ot find this normative argument compelling, and seek , .. .. I

I

, , I
,, I
anthropocentric justifications, primarily economic ones, B
,,
I
~ I
ior the conservation of biodiversity, as discussed in 0 ,,
I I
I
Chapters 4 and 5. In the past 15 years, conservationists I
I
I
I
I
I
have focused on describing the nature of the links be- I
I
I
I I
n¥een biodiversity and ecosystem processing (Chapin et I
I
I
/
/
/
al. 1997). Ecosystems provide a large array of goods and 20 I
1
1
c // /
services vital to human existence, including water and I
I
,.. .... /

I ,.. ....
air purification, cycling of critical elements and water, I
..... ..........
development and retention of rich soils, regulation of
0
I

l( ater flow, carbon sequestration, decomposition, and
p rimary and secondary production (Table 2.4). Claire
Kremen discusses the importance of one of these servic-
es, pollination, in Box 2.2.
If clean air and water, fertile soils, pollination of crops,
and other "services" depend upoh the maintenance of a
b road diversity of native species, this becomes a power-
ful motivator for their conservation. For example, in an
experiment manipulating species richness in a prairie
ecosystem in Minnesota, Tilman and Downing (1994)
showed that drought resistance was greater in species-
rich plots than in species-poor plots. Many observation-
al, experimental and theoretical studies have found that
species-rich communities have higher values of some
metric of ecosystem functioning than species-poor com-
munities. If generally true, then preserving as many
species as possible should be a primary goal.
Yet, ecologists still know little about how attributes of
communities including species richness, relative abun-
·d ances, and functional traits of individual species (such
as ability to fix nitrogen or seed dispersal effectiveness)-
TABLE 2.4 Examples of Ecosystem Services
()
Source: Costanza et al. 1997 and Millenium Ecosystem Assessment
2005.
0 20
--- 40 60 80 100
Richness (0/o)
Figure 2.19 Ecosystem function could increase quickly with
species richness (A), linearly (B), or as an accelerating func-
tion (C). Increases in ecosystem function with species richness
could occur due to increases in ecological functionality either
because of changes in the number, abundance, or functional
traits held by the species in the community. (Modified from
Kremen 2005.)
interact to influence the magnitude and stability of
ecosystem processes (Figure 2.19). For example, are rela-
tively few or many species required for high values of
some ecosystem process (curve A versus C in Figure
2.19)? Or is the relative abundance of key species, or the
diversity of functional groups, most strongly related to
ecosystem processes?
Considerable debate has sprung up regarding the inter-
pretation of experimental investigations of relationships
between species richness and ecosystem processes. The
importance of rare species to ecosystem functioning is
poorly known, and many studies show that a particular
ecosystem process, such as primary productivity, may
reach a plateau at only modest species richness (e.g.,
20°/o-30°/o of maximum richness) (Schwartz et al. 2000).
Only a few studies show a linear increase of ecosystem
functioning with species richness (e.g., Naeem and Li
1997), although several show that a more diverse commu-
nity yields the least variability in biomass (e.g., Mc-
Naughton 1977; Tilman 1996). Further, most experimental
investigations to date used synthetic communities, where
less than 40 species are brought together in different mix-
tures in lab or field plots at small scales. The results of these
studies may not be good predictors of processes and pat-
terns in natural ecosystems where the assembled species
have long histories of interaction. Yet, these experiments
point to interesting principles to test in the field, and the
scale of some experiments is appropriate for the process of
interest. For example, microcosm experiments of soil com-
munities are complex enough to show that functional di-
versity of species (i.e., differences in the roles they play in
5 Chapt r 2
 '
the ecosystem), and not species richness, influences the
rate of decomposition (Heemsbergen et al. 2004).
Increasingly researchers are using large-scale removal
experiments and comparative studies of communities
1vith varying species membership and abundances to
gauge the role of individual species as well as diversity
on ecosystem functioning (e.g., Kremen et al. 2002a,b;
Diaz et al. 2003). A number of biologists believe that eco-
logical differences among species should create the con-
ditions for an increase in ecosystem process with diver-
sity, while others believe that only the attributes of a few
d ominant species will control that relationship (Figure
2.20). A growing consensus focuses on the importance of
differences in the ecological roles of species, and the joint
contributions these differences make (Diaz and Cabido
2001; Lareau et al. 2001). However, measuring the signif-
icance of a species for a particular ecosystem process is
m uch harder than counting numbers of species or esti-
m ating population densities. Full understanding of how
biodiversity affects the ability of an ecosystem to provide
goods and services over time awaits much further work.
Species pool
Low richness
Low functional diversity
Dominance of species Complimentarity among
\vith particular traits functional groups of species
Lower Functioning ECOSYSTEM PROCESSES
Global Biodiversity 55
The Future of Biodiversity Studies
The incomplete state of our knowledge of the identities,
taxonomic relationships, and distributions of the vast ma-
jority of the world's organisms means that the primary
work of cataloging biodiversity is yet to be done. Today
relatively few scientists are being trained as taxonomists.
Therefore, increasing the cadre of competent taxonomists,
particularly in tropical nations, is an important goal
(Mikkelsen and Cracraft 2001; Gotelli 2004). How this in-
ventory should be carried out is the subject of much de-
bate. Some biologists have committed themselves to an
intense global surve~ aimed at the discovery and classifi-
cation of all species (e.g., the Species 2000 Project). Others,
pointing to the shortage of people, funds, and time, be-
lieve that the only realistic hope lies in the rapid recogni-
tion and preservation of those threatened habitats that
contain the largest number of endemic species (Conserva-
tion International strategy), or to conserve representatives
of the habitats of every ecoregion (World Wildlife Fund
and The Nature Conservancy strategies). They give the in-
ventory task a lower immediate priority.
Figure 2.20 Ecosystem function can
be influenced by the functional diver-
sity among species in a community
via several mechanisms. Representa-
tion within a community of more
than a single or few functional
groups, or of many species with dis-
tinct functional roles should lead to
higher functioning of various ecosys-
tem processes. (Modified from Lore-
au et al. 2001.)
Complimentarity among species
with different functional roles
Higher Functioning
56 Chapter 2

E. 0. Wilson (1992) strongly advocated a strategy that
combines global surveys designed to achieve a complete
biodiversity inventory in 50 years with special attention
to areas of unusually high species richness ...a nd en-
demism. His strategy has three components. The first is
a Rapid Assessment Program (RAP) that would investi-
gate, within a few years, poorly known ecosystems. RAP
teams would be formed, consisting of experts on groups
such as flowering plants, reptiles, mammals, birds, fish-
es, and butterflies that are well enough known to be in-
ventoried quickly and accurately. These groups would
then serve as proxies for the entire biota (see Essay 2.3 by
Lily Rodriguez; for a discussion of applications to prior-
ity setting in biodiversity conservation see Chapter 14).
The next stage would be to establish research stations in
areas believed to be major hotspots of diversity. Invento-
ries and ecological studies would then be carried out
there and in surrounding regions. The third stage, with
a time frame of 50 years, would combine the inventories
from RAP and the intensive studies at a small number of
research stations with monographic studies of many
groups of organisms to provide a more complete picture
of global biodiversity and its distribution.
.
While a reasonably complete inventory of the biota of a
region, or significant components of it, is being achieved, a
monitoring program also needs to be established. Such a
program should be designed to detect trends in biodiver-
sity and to identify impending problems to which atten-
tion should be directed (National Resource Council2000).
Without a monitoring program society cannot know if its
efforts to preserve biodiversity are succeeding or why
and where they are failing.
Human societies will need to greatly increase their in-
vestments in biodiversity studies during future decades
if we are to understand the patterns of biodiversity more
fully. The scope of the untapped wealth residing in bio-
diversity, as well as the importance of inventorying bio-
diversity and understanding ecological relationships
among species, is unappreciated. It is doubtful that hu-
mans can devise a sustainable future without a more
complete knowledge of biodiversity.
The relationship between area and species richness
has major practical implications for the location, design,
and management of parks and reserves that are estab-
lished and maintained to preserve biodiversity. There is
increasing evidence that even the largest parks and re-

ESSAY2.3 • • .. . ..
-:-

Rapid Inventories for Conservation

Lily 0 . Rodriguez, CIMA, Cordillera Azul

Developed under the Rapid Assess-
ment Program (RAP, of Conservation
International, CI) or as Rapid Biological
Inventories (RBI, by The Field Museum
of Chicago), rapid inventories are a rel-
atively new conservation tool designed
to assess the biological importance of
priority sites for conservation. Using
short field trips, usually of no more
than four weeks, these inventories eval-
uate the biodiversity values of sites of
global conservation interest, providing
scientifically based recommendations
for prompt establishment of protected
areas and conservation action plans.
Identifying conservation targets and
their threats, and generating recom-
mendations for immediate conservation
action are the main goals of these
inventories.
RAP or RBI results rely highly on
the expertise of the team undertaking
the inventory, as there is no single
methodology that is or can be used in
all places with all taxa. Here, I provide
some insights and recommendations
based on my experience after partici-
pating in inventories over the last
decade.
Multidisciplinary teams are formed
for each expedition, usually with a core
group of well-recognized scientists (tax-
onomists and ecologists), combined
with resident field biologists, who are
both specialists and specialists-in-train-
ing. Each specialist selects the methods
and groups that will be the focus of its
work, based on abundance, rarity or
uniqueness, and capacity to document
the group in a short period of time.
None of these inventories are meant to
yield complete information on the site.
Planning the trip and identifying
2-4 campsites are crucial steps to
ensure exciting results. Satellite
imagery is used first to assess the gen-
eral area and locate points of main
interest. The preliminary selection of
sites to inventory is made based on this
first step, combined with a review of
any previous research in the area or
nearby locations, as well as local
knowledge. Overflights to the area are
then made by a few team members.
During those overflights, team mem-
bers select points for easy access to as
many different types of habitats as pos-
sible, verify any unique characteristics
seen in the images, and assess impacts
of human activities in the area to be
surveyed. Final decisions on sites are
made based on accessibility and trans-
portation. Although relatively expen-
sive, helicopters are the best way to
reach remote areas, and to relocate the
team between main campsites. To maxi-
mize time for the inventory, camps and
trails should be prepared prior to
arrival of the team. While preparing the
trails (usually loop trails), satellite
campsites can be prepared and used by
the team according to distances, inten-
sity of work in different habitats, taxa,
and technique requirements.
Different approaches are needed for
each of the principle taxonomic groups
addressed in rapid inventories. Plants
provide the best characterization of
habitat types and communities associ-
ated with the area and are therefore a
must in any rapid inventory. Transects,

q:uadrat plots, collections, and photo-
graphs are some of the techniques that
can be used to characterize the flora,
providing quantitative information,
::eld observations and general collec-
::ons of the diversity of plants. Focal
groups, such as ferns or understory
~egetation, are important as they pro-
-tide better resolution of habitat differ-
ences than do trees. Groups of special
-.--alue such as orchids may also add to
~.,_e richness of the results.
Birds are one. of the best-documented
groups around the world. Because of
:heir high species numbers, and the fact
that the range, distribution, and biology
is reasonably well known for most
species or groups of species, birds are
used as habitat indicators. Avian data
can be used to provide quick compar-
isons among sites, and to guide consid-
erations such as connectivity, size of the
nrotected area, and conservation status
~
and perturbation of the site. Most com-
monly used methods involve auditory
surveys along trails, including song
recording and point counts. Assessing
the status of local populations of hunted
species is an important step for manage-
ment recommendations.
Fishes are the main community
associated with aquatic habitats. Often
easy to collect, they can be captured
1-vith small nets that are for scientific
use only. A high percentage of the fish
community can thus be surveyed.
Interviewing local people with photo-
graphs in hand provides rich informa-
tion on consumed species and seasonal
variations. Usually fishes contribute to
Figure A Members of a RAP team examining fish specimens collected
that day. (Photograph courtesy of L. Rodriguez.)
the inventory with rare records or new
species and the presence of suites of
species can help relate basins and
aquatic communities (Figure A).
Insects, because of their immense
species richness, can rarely be used as a
group for rapid inventories. The best
choice is to take experts on specific tax-
onomic groups that are best known
and easily seen in few days. Butterflies,
beetles, dung beetles, and spiders are
some candidates for rapid inventories,
but, as always, this will depend on the
availability of experts and the scope of
knowledge of the group.
Mammals (especially large mam-
mals) experience heavy human pres-
sures and are important to document,
providing a primary indica tor of the
conservation quality of the area. Tracks
and visual encounters are used most
frequently to document mammalian
species. However, small mammals such
as rodents and bats are speciose groups
that if specifically surveyed can pro-
vide a more detailed evaluation of the
site. A trade-off between time and
value is usually applied here, with bats
(which are easily censused with mist
nets) being one of the best choices.
Because of their sensitivity to distur-
bance, frogs are usually included in
rapid inventories. Easily located by
their calls, auditory transects and
visual encounters are the most com-
mon techniques employed to survey
for frogs. Snakes, because of their low
population densities and secretive
habits, are difficult to include; however,
they can add value as indicators of
Global Biodiversity 57
regional faunas. Because aquatic and
terrestrial turtles as well as crocodilians
are frequently hunted they are always
included and evaluated as target
species for conservation actions. Tissue
collection of amphibians (especially in
mountain ecosystems, where popula-
tions declines due to disease appear to
occur more often) is now recom-
mended to be able to evaluate possible
causes of decline in amphibian popula-
tions.
Geology, soil diversity, freshwater
characteristics, and substratum are sur-
veyed to understand and document the
uniqueness of the area and to indicate
species distributions and plant commu-
nity types. An additional but extremely
important aspect is to include human
dimensions in the rapid inventory
through rapid social asset inventories.
This consists of visiting the communi-
ties surrounding the site surveyed, and
is achieved through a diversity of tech-
niques (interviews of local leaders,
authorities, key people of the commu-
nity, and focal groups; meetings with
the whole community; and visits to
their crop fields), the team explores the
history, demography, economy, social
organization and structure, and the use
of natural resources. While identifying
patterns of social organization and
opportunities for capacity building,
these· inventories contribute to hone the
recommendations, and to engage local
participation for future conservation
action.
Rapid inventories should not be
taken as the sole source of information
on biodiversity. While they are a quick
and effective way to document biodi-
versity, they do not provide complete
species lists, and abundance informa-
tion is limited and depends on the
areas visited and the seasons in which
inventory occurs. Furthermore, it is
highly expensive, and requires a core of
highly trained taxonomists who can
quickly recognize what is new to sci-
ence and relevant to the area, and who
are able to compare findings with the
widest possible range of other sites.
It is also important that local people,
as well as local scientists participate in
the inventory. Being part of the field-
work will give the residents a first-
hand insight of the global importance
and will increase their support and
interest in the area, as well as con-
tribute to developing pride in the site.
Recognizing species that are of no use
to them but that can be of high scien-
tific value (rare or unique species)
58 Chapter 2

usually gives them a new perspective

on the value of protected areas and
their potential to attract new and com-
patible activities such as research and Brazil
tourism.
Student participation is usually lim-
ited because the time devoted in the
field to the learning process is expen-
sive. However, it is a wonderful oppor- Bolivia
tunity for training and increasing local
capacities.
Presenting results to local, regional,
and national key players is another
important issue. Immediately after gatoni
completing fieldwork and before leav- Tambopata
Vileabamba
ing the area, a meeting with local com- complex
munities to present preliminary results
will inform them of the value of their
area and enhance relationships. After
some processing and analysis, a more
elaborate report should be given to
regional authorities, academics, and
conservation, tourism, and other inter-
ests. A similar presentation in the capi-
tal will enable the results to be high-
lighted to proper decision makers. But Figure B Locations of RAP expeditions to Tambopata and Megantoni (light gray), areas adja-
the most important step is to involve cent to globally important national parks (Manu Biosphere Reserve, Otishi, and Bahuaja-Sonene
National Parks are in gray) in southeastern Peru.
any local institutions that will be in
charge of the next steps for conserva-
tion management and oversight of the only 500 m Epipedobates simulans, a Rapid biological inventories are an
protected area. poison arrow frog. Thus, RAP invento- efficient tool for conservation. They
In 1992, I was the herpetologist for ries can add new species, even in areas provide quick results and recommenda-
the RAP to Tambopata, a large expanse that have been well studied previously. tions that can be used immediately by ·
of Andean forest in southeastern Peru, Eleven species of reptiles and decision makers. For example, the Tam-
from lowland rainforest at 200 m to amphibians, including a new snake, bopata reserved zone was created adja-
cloud forest at 2400 m (Figure B). three lizards, and seven frogs were dis- cent to Bahuaja-Sonene National Park,
Importantly, Tambopata also included covered in a more recent inventory and the boundaries of this park were
the only piece of lowland Amazonian (May 2004) of the Megantoni Reserved expanded based on the Tambopata
grasslands (pa~tanal) in Peru. Because Zone, the strip of land connecting inventory. Despite the fact that
Roy McDiarmid and others had studied Manu Biosphere Reserve and the Vil- RAP /RBI is expensive and reliant on
amphibians in Tambopata in the low- eabamba complex. Data in this case the expertise of the team, this modern
lands, I assumed any new species I supported the categorization of this tool is a good solution to fill gaps of
might find would be in the highlands. area as a national sanctuary and has information needed to complete pro-
As expected, I added new species and provided very valuable recommenda- tected area systems, and to enable local
occurrence records at a high elevation tions for management and zoning of governments to take further steps for
site, yet I also found a new species at the area. effective conservation.

serves are too small to maintain viable populations of
those species with the largest aerial requirements (dis-
cussed in Chapters 7 and 14). Many of the national parks
of the United States have already lost their largest mam-
mal species, and the trend continues (Newmark 1987).
Therefore, additional studies of the species-area rela-
tionship and its causes are essential to inform manage-
ment of protected areas. Meanwhile, existing knowledge
of species-area relationships is being used to guide es-
tablishment of protected areas. A notable example is the
megareserve system of Costa Rica, which is designed to
preserve about 80°/o of the biodiversity of the country
over the long term (Figure 2.21). Each megareserve in-
cludes natural areas and areas managed for economical-
ly valuable products. Some conservation areas remain
the homes of indigenous people who continue to use the
environment in their traditional ways. The largest of the
Costa Rican reserves, La Amistad Biosphere Reserve, is
a mosaic of more than 500,000 ha and includes three na-
tional or international parks, a large biological reserve,
five Indian reservations, and two large forest reserves.
On a theoretical level, biodiversity studies continue to
be needed to resolve the many uncertainties surround-
ing the historical and present-day ecological processes

Nicaragua
Conservation areas
1. Guanacaste
2. Arenal Pacific Ocean
3. Tempisque
4. Cordillera Volcanica Central
5. Pacifico Central
6. Llanuras del Tortuguero
7. Amistad Caribe
8. Amistad Pacifico
9. Osa
10. Isla del Coco 0 10 20 30 40 50 60 70
Kilometers
that determine today' s patterns of biodiversity. Modern
phylogenetic techniques that enable systematists to de-
velop soundly based phylogenies are being combined
w ith biogeographical studies to provide a more com-
plete picture of the history of the distribution of life on
Earth. Processes operating over ecological time frames
are increasingly being studied using manipulative ex-
periments in which some or all species are removed
from restricted areas, or species are introduced. Many
.unatural experiments," are being studied to gain ecolog-
ical insights from them volcanic eruptions that elimi-
nate the biotas of islands, the massive inadvertent move-
ment of species around the world by human travel;
deliberate introductions for agricultural, aesthetic, or
pest-control purposes; and habitat fragmentation by
conversion of natural .landscapes to ones dominated by
highly modified communities that are managed to chan-
nel most of their productivity to human uses. Such sce-
narios provide opportunities to examine the results of
manipulations over longer time frames than is possible
with investigator-initiated experiments.
Studies of the influences of human activities on species
distributions and species richness are adding rapidly to
our understanding of the roles of the varied processes that
interact to cause the patterns in the distribution of biodi-
Global Biodiversity 59
Figure 2.21 The extent of Costa
Rica's conservation areas as of 1996.
Shaded areas indicate protected areas,
which are organized into 10 distinct
conservation areas.
Caribbean Sea
Panama
®
versity on Earth that so fascinate us today. This knowl-
edge is also being used to attempt to reduce the rates of
species extinction and to restore landscapes so that they
can continue to support the array of species originally
found in them and the evolutionary processes that gener-
ate new species. For example, to promote survival of ex-
isting species and the evolutionary processes that gener-
ate new species, devoting large areas to biodiversity
preservation will be critical.
Recently, the Millennium Ecosystem Assessment
(MA), a large coalition of international development and
conservation organizations, governments, and scientists
has come together to assess the status of Earth's ecosys-
tems, the goods and services they provide, and the like-
ly effects of potential pathways of human economic de-
velopment on the future provisioning of these services
and human well-being (Figure 2.22). The MA focuses
both globally, and on subglobal regions of particular
concern due to the difficulty of human existence or po-
tential for serious declines in human welfare in these re-
gions. Published in 2005, this is the first comprehensive
assessment of the condition and status of global ecosys-
tem services. The information summarized in the MA
will be used to guide development policy both regional-
ly and globally. In addition, the MA will help focus re-

common species typically dominate ecosystem
processes.
_ 1.- \bout 1.75 million living and 300,000 fossil species
have been described, but estimates of the total num-
ber of living species range from 10 mi11ion to as high
as 50 million or more. Although species richness is
higher on land than in the oceans, 34 of the 35 extant
animal phyla have marine representatives, and 16
are exclusively marine.
.i. Cellular life in the form of bacteria evolved about 3.8
billion years ago; eukaryotic organisms evolved
about 2 billion years ago. The first major explosion
of biodiversity took place in the early Cambrian pe-
riod, and except during times of mass extinctions,
the number of species has increased since then.
ore species are probably alive today than at any
other time in the history of life, even though some
taxa had more species in the past than they do today.
The number of species present at any moment is the
result of the difference in the rate of formation of
new species and the rate of extinction of existing
species. High rates of speciation have been favored
by mass extinctions, the breakup of the continents,
and the evolution of more diverse body plans that
enabled animals to burrow, swim, and fly.
~· Several broad patterns characterize the current dis-
tribution of species. Areas that have experienced
long geographical isolation and that have great
topographic relief often support many endemic
species. Tropical regions often have both high
species richness and endemism, and islands have
high endemism of both marine and terrestrial
•
species.
- . Among most taxa, more species live in tropical re-
gions than at higher latitudes. The large land area of
the Tropics contributes to this pattern. Species rich-
ness is also positively correlated with available ener-
gy, which enables more individuals to be supported
per unit area, and with structural complexity, which
is positively correlated with available energy. On
land, structure is provided primarily by vascular
plants, whereas in many marine communities, ani-
mals such as corals generate most structure. There is
a positive correlation between productivity and
species richness at low levels of productivity, but
species richness typically declines at higher produc-
tivities. Some highly productive systems, such as salt
marshes, seagrass beds, and hot springs are species-
poor. Plant species richness is extremely high in
some unproductive semiarid regions with poor soils.
Island communities are poorer in species than com-
parable mainland communities at all latitudes.
Global Biodiversity 61
6. Climatic oscillations producing swift, radical changes
in climate increased extinction rates and likely de-
creased speciation rates toward the poles, thus acting
as a strong driver of latitudinal patterns in species
richness.
7. Both physical (heavy rains, strong winds, landslides,
earthquakes, and fires) and biological disturbances
(tree-falls and activities of predators and competi-
tors) influence species richness but their effects are
highly varied. Most disturbances are small and di-
rectly affect only small areas, but they may influence
more distant regions via indirect effects. Strong com-
petition and predation may eliminate species from
particular areas but intermediate levels of predation,
by preventing competitive exclusion, may increase
local species richness.
8. Despite the demonstrated importance of local bio-
logical interactions, local species richness in many
taxa appears to increase without apparent limit along
with increasing regional species richness. This sur-
prising result suggests either that local interactions
exert strong effects on abundances of species but not
on their numbers, or that the influence of regional
processes on numbers of species in local communi-
ties is strong enough to override local influences.
9. The state of knowledge of Earth's biodiversity is so
poor that the primary work of cataloging biodiver-
sity is yet to be done. Resources currently devoted to
this task are inadequate, especially given the rate at
which species are becoming extinct. Better informa-
tion on biodiversity, its distribution, at1d its causes is
needed for wise management of Earth's biotic re-
sources. Further, research that allows us to under-
stand the contributions of biodiversity to human
welfare, and the effects of development on ecosys-
tem services is needed to help us preserve the rich-
ness of biodiversity and human existence.
. -
Pleases refer te the;~e}fsite~www.s:Un.au~r.c(i)Pl/~oq;p:l. s~¥
fqr Stlggesjed ~ea~~' ~eb l~s,_,,adqitiol).al ques¥ons,
aha supplementary resouices.~w -,~'{' !'7 ~- -~i ' ,,ifl't Yf~ '
.
*
.
. . . ·,1S,;~ -~-1>:' . _,Jf~ . ;·~~t- -~_;;i; '<·..... .;o
Questions for Discussion
1. The history of life has been punctuated by five
episodes during which extinction rates were very
high. If extinction is a normal process, and if life has
rediversified after each mass extinction, why should
we be worried about the prospects of high extinction
rates during this century? How does the current ex-
tinction spasm differ from previous ones?
 62 Chapter 2
2. Given that millions of species are yet to be described
and named, how should the limited human and fi-
nancial resources available for taxonomic research
be allocated? Should attention be concentrated on
poorly known taxa? Should efforts be directed to-
ward areas threatened with habitat destruction so
that species can be collected before they are elimi-
nated? Should major efforts be directed to obtain
complete "all taxa" surveys of selected areas? How
and by whom should these decisions be made?
3. Indices of species diversity that are weighted by
abundance, biomass, or productivity are used fre-
quently by ecologists, but seldom by conservation
biologists. For what purposes might conservation
biologists wish to use weighted indices instead of
simple lists of species?
4. Many conservation efforts are directed at particular
local areas harboring rare species or having high
species richness. Why is concentrating only on local
problems insufficient as an effective conservation
strategy?
5. For which animal taxa would you expect species
richness to be most positively correlated with plant
community structure? Mammals? Amphibians? In-
sects? Why?