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• • • 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

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
:~ \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 •• • , • • ·

_---...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-
,. .
: 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
, .· ·, . '" ""

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

. . :.
·!. ··
. 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 ·•
' 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) . ,. " , ·,. · .· ~: . :

--~ 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 the evolutionary history of that population, and shape

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

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

_ 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
----. Endangered Species Act (ESA) and ·_p
-._e Convention on International Trade in ~
~ 250
~:;ecies (CITES). Therefore we will em~ ~
- 00
~~ .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
- -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 system. Kolata and his colleagues lethal yellow dwarf virus, which threat-
Van der Post remarked that the most worked with the Aymaras to dig a series ened an industry worth $160 million per
damning legacy of colonialism has been of raised beds surrounded by ditches year. The yield of cassava in Africa was
its relentless tendency to separate tropi- and the results were a crop yield seven increased ten-fold because of disease
cal peoples from their land and culture. times greater than the local average. resistance provided by cassava from the
"The great mistake people make is in Here we see three different fates for Amazon. And scientists have recently
thinking about the aboriginal races in three independently evolved agricul- found that a variety of sunflower being
South Africa the Bushmen and Hot- tural systems: The Mojeno system has cultivated by the Havasupai Indians in
tentots and the like as primitive peo- been lost, the chinampa system still the American southwest offers resistance
ples. They were actually very sophisti- exists, and the altiplano system would to a blight attacking sunflower crops in
cated societies and cultures with have remained lost if it had not been for the Old World. ·
immense awareness and very impor- an intrepid anthropologist and his The late economic botanist Edgar
tant values." indigenous colleagues. How many other Anderson once stumbled across an
The sophistication of many indige- equally valid and possibly even more Indian garden in Guatemala that ini-
nous cultures is most evident in their productive systems have completely dis- tially seemed more of a rat's nest than a
agricultural systems. Over 200 years ago, appeared without a trace? The superior- productive agricultural plot. It was .
a Jesuit priest in lowland Bolivia, Fran- ity complex that drives outsiders from only after careful study that he realized
cisco Eder, noted that an extraordinary Western societies to want to replace how much more sophisticated than the
irrigation system in the local savannas, "primitive" systems with an ill-suited botanist was the farmer:
which had been devised in pre- one developed on foreign soils continue In terms of our American and ...

Columbian times, still supported an to this day: The Indonesian government European equivalents the garden
enormous population of Mojeno Indtans. is currently trying to get tribal peoples was a vegetable garden, an or-
Thanks to the influence of the padre and in highland New Guinea to forgo their chard, a medicinal plant garden, a
his cohorts, as well as introduced dis- traditional and highly productive agri- rubbish heap, a compost heap, and
eases, this Amerindian agricultural sys- cultural system based on sweet potatoes a bee-yard. There was no problem
tem was abandoned and the savannas or sago palms, and replace it with rice, of erosion though it was at the top
that once fed the teeming local populace an inappropriate crop. This colonial atti- of a steep slope; the soil surface was
today are covered with a scrub vegeta- tude comes from cultural biases and a practically all covered and appar-
tion supporting a few mangy cows. lack of understanding. ently would be during most of the
The chinampa system, developed in Yet, the problem we face is not just year. Humidity would be kept up
pre-Columbian Mexico, remains in use, loss of agricultural systems themselves, during the dry season and plants
albeit in a much more restricted range but extinction of cultivar diversity as of the same sort were so isolated
than it once covered. Usually con- well. Indigenous agricultural systems from one another by intervening
structed in swampy or lacustrine envi- typically contain many different vari- vegetation that pests and diseases
ronments, chinampas consist of garden eties of a single crop, much as a farmer could not readily spread from plant
plots created by building small islands in the industrialized world may grow to plant. ... I suspect that if one
using layers of vegetation and mud. The several different types of corn: one to were to make a careful study of
surrounding waters seep in and provide feed his family, another to feed his ani- such an American Indian garden,
the necessary moisture, while mud mals, still.another to make popcorn, one would find it more productive
scooped up from canals around the plot and another to sell as a cash crop. The than ours in terms of pounds of
is periodically added as fertilizer to the Amerindian farmer in the Amazon usu- vegetables and fruit per man-hour
garden on top of the island. According ally cultivates distinct varieties of cas- per square foot of ground.
to Dr. Jim Nations of Co11servation Inter- sava for the production of bread, beer,
national, chinampas not only produce meal, porridge, and whatever other end At a time when there are ever more
food and other useful crops year-round, products he desires. The Tirio Indians mouths to feed on this planet, Ander-
they do not deplete the soil or require of northeastern Amazonia have at least son's admonition to look more closely
artificial fertilizers or pesticides.· 15 varieties of cassava in their gardens at the form and function of indigenous
A similar system was rediscovered in while other tribes like the Machiguenga systems seems ~ike advice worth fol-
the altiplano of northwestern Bolivia. of Peru may cultivate several dozen. lowing. Modern management tech-
Dr. Alan Kolata of the University of These crop varieties often harbor niques often overlook and disparage
Chicago had long been intrigued by a adaptations that enhance yields or these indigenous systems, which are
series of ridges and depressions in this increase resistance to pests and diseases. based on centuries of in situ sustainable
remote region that seemed to indicate ReceD.tly, crosses of indigenous and existence, in favor of high-tech but
some form of agricultural system that commercial varieties have led to tremen- often inappropriate and expensive sys-
involved raised planting beds sur- dous improvements. A barley plant from tems that fail. We can learn a great deal
rounded by canals. The local Aymara Ethiopia was crossbred with barley in about environmental management (and
Indians, however, employed no such California, providing resistance to the humility) from such cultures.
Global Biodiversity 33

::= ""'le. For example, the concept cannot be applied to organ- classify organisms hierarchically in ways that reflect
:.s::ns that reproduce asexually. Also, fossils, the source of their evolutionary history, constructing classifications in
_ ..ost information about organisms that lived in the past, which the higher taxonomic categories contain all of the
_arely provide clues about breeding behavior. Finally, in- descendants of their most recent ancestor, but no other
.:c::rnation about reproductive behavior is not available for species; that is, categories that are monophyletic.
_ 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
How Many Species Are There?
-~~~ biologists have relied heavily on distinguishing mor- Approximately 1.75 mi1Hon living and 300,000 fossil species
,1-. .I.<L'ospecies, particularly among arthropods (INBIO 1995). have been described and given scientific names (Table
---:."""_e morphospecies concept is useful, because it can be ap- 2.1). On average, about 300 new species are formally de-
_:::ed when other data are unavailable. Moreover, given scribed and named every day, and no slowdown is in
~..itt evolution is, to use Darwin's words, "descent with sight. New animal phyla, Cycliophora and Loricifera,
.=-edification," organisms that look alike generally share whose members live in interstitial spaces in ocean sedi-
:::any alleles that code for their body structures. This is ments, were first described within the past 25 years! Not
~ -:)~ most species designations based on morphology surprisingly, because they are based on incomplete and
~;:-.--e been supported by subsequent genetic data. indirect evidence, estimates of the actual number of living
-~ closely related species continue to evolve in genetic species vary widely. Current estimates of the total number
: lation from one another, they diverge and accumulate of living species range from 10 million to as high as 50
,;ettetic differences. Thus, the number of unique genes, million or more (May 1988; Wilson 1992; Gaston and
- d the morphological and physiological traits they en- Spicer 2004). In other words we do not know within an
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 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 The rate of description of new species today is higher
large fraction of species likely to be exterminated during than it ever has been, but the rate is nonetheless inade-
the twenty-first century will disappear before they have quate to accomplish a reasonable inventory prior to the
been named, much less understood ecologically. likely extinction of many of them.
The immense richness of viruses, bacteria, archaea,
protists, and unicellular algae is still largely uncata-
logued. Within the domain Archaea, new phylum-level
Diversity of Higher Taxa
groups are discovered every year (Furhman and Camp- Recent advances in molecular evolution-have painted a
bell 1998), and the richness of species within these radically different portrait of taxonomic diversity than
groups is unknown. There are few studies of viruses ex- existed even 20 years ago. Until recently, taxonomists
cept those that attack people, our domesticated plants classified all living organisms into five kingdoms: Ani-
and animals, and the organisms we study scientifically. malia, Plantae, Fungi, Protista, and Monera (all prokary-
How many types attack noncrop plants and insects is otes). Today, we recognize that there are deep evolution-
unknown, as is the number of marine forms. Similarly, ary divisions among the prokaryotes. They are now
bacteria and protists living in soils or that attack inverte- divided into two large domains, the Archaea and the
brates have scarcely been examined. Terrestrial algae, es- Bacteria (Woese et al. 1990; Figure 2.2). The remaining
pecially those living on bark and rocks, have been little four kingdoms are grouped together in the third do-
studied. All estimates of the number of species in these . main, the Eukarya (see Figure 2.2). The genetic diversity
groups are crude guesses at best (Torsvik et al. 2002). among the Bacteria (which contains all familiar prokary-
About 80,000 species of fungi have been described, but otes such as gram-positive bacteria, cyanobacteria, and
the total number living today likely exceeds 1 million. For green sulfur bacteria) and Archaea (which are unfamil-
example, the 12,000 fungal species described in Britain is iar to most, but include organisms adapted to life in ex-
about six times the number of native vascular plants de- treme environments such as deep sea hydrothermal
scribed for the same area. If there were, on average, six vents, hot springs, the guts of animals, and hypersaline
fungal species for each vascular plant species worldwide environments) is as great as that among the long-recog-
(the ratio found in Britain), the number of fungal species nized kingdoms among the Eukaryotes. Two new king-
would be approximately 1.6 million (Hawksworth 1991). doms are now recognized among the Archaea (Crenar-
The global ratio could be much higher or lower than the chaeota and Euryarchaeota) and at least six more are
British one, but we have little else to base estimates upon. recognized among the Bacteria. Similarly, the protists,
Taxonomists believe that the number of species of ne- which now are known to be vastly more diverse than
matodes is very large because millions of individuals previously recognized, have been divided into seven
may be present in 1 square kilometer of soil or mud; more kingdoms. These discoveries are exciting because
more than 200 species have been reported in samples of they force us to consider the importance of organisms
just a few cubic centimeters of coastal mud (Poinar 1983). whose size make them generally invisible to us, yet are
Yet, almost nothing is known about species ranges and evolutionarily equal to the plant and animal kingdoms
rates of species turnover geographically, so global esti- that have a greater hold on our consciousness. To appre-
mates are very uncertain, although these data make a ciate this in another way, the diversity among all plants,
compelling case for a vast diversity of nematode species. fungi, and animals in small subunit RNA, the molecule
Approximately 30,000 species of mites have been de- used to create the modem phylogeny of all life on Earth,
scribed but, because knowledge of tropical mite faunas is is only 10°/o of the diversity spread across all the rest of
extremely poor, the actual number of living species could the Eukaryotes, Archaea, and Bacteria (Olsen and Woese
easily exceed 1 million. Nearly 1 million species of insects, 1996). Freeman (2002) describes their importance well:
the world's most speciose group of organisms, have been
Bacteria and archaea are ancient, abundant, ubiqui-
described, but this is certainly a small fraction of the total.
tous and diverse ... their abundance is well-docu-
Most (>55°/o) of the .insects collected by fogging the mented. A teaspoon of good-quality soil contains
canopies of tropical trees, for example, are members of un- billions of microbes. In sheer numbers, the bacterium ·
described species (Erwin 1991), and samples taken with- Prochlorococcus found in the plankton of the world's
in 70 km of one another at four sites in Manaus, Brazil had oceans may be the dominant life-form on the
only 1°/o of their species in common (Erwin 1983). These planet ... a drop of seawater contains a population
samples hint at a vast unexplored diversity of arthropods, equivalent to a large city. Yet Prochlorococcus was first
described and named only recently in 1998.
just in the canopies of tropical trees.
Bacteria and archaea are also found in almost
Earth is a relatively unexplored planet biologically. every conceivable habitat. On land, they live in envi-
Not only are most living species still undescribed, but ronments as unusual as oxygen-free mud, hot springs,
very little is known about the life history and ecological salt flats, the roots of plants, and the guts of animals.
relations of most of the species that have been named. They have been discovered living in bedrock to a
Global Biodiversity 35

- Domain Bacteria Domain Eukarya

Bryophytes Seedless vascular plants Gymnosperms

0.1>- ~1f
~1>- ·~ 0.1>-
~0 o.~'\-
# '!o...o

(;~ (}~ ~~~ 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 organisms with which we interact. No matter how many
they are found from the surface to depths of 10,000 m, species of beetles there may be, they cannot substitute
and at temperatures ranging from ooc in Antarctic sea
ice to over 110°C near submarine volcanos. aesthetically for mammals, fishes, corals, or butterflies.

Organisms so widespread and adaptable may not be

in danger of extinction. Nonetheless we should be con-
Diversity of Biological Communities
cerned that such incredible diversity be maintained be- The distinctness of biological communities is defined by
cause it may be of critical functional importance. the species within them and their interactions. The species
The distinctness of evolving lineages is an important composition of ecological communities changes over
component of biodiversity. The higher taxonomic cate- space because each species has unique adaptations to its
gories of the Linnaean biological classification system physical and biological environment. As we move across
provide rough measures of distinctness of lineages. For a landscape, species will be present according to their tol-
example, phyla ill the animal kingdom are distinguished erance for the physical conditions that vary spatially (tem-
by their developmental patterns and their adult body perature, salinity, light levels, soil chemistry, etc.), and if
plans (e.g., having two or three embryonic tissue types). they are able to survive their interactions with other
Measured by diversity of phyla, marine biodiversity is species. Community membership is never absolute, al-
much higher than terrestrial biodiversity even though though suites of species may be commonly found togeth-
far fewer species of marine than of terrestrial organisms er under similar conditions.
have been described. Of the 96 phyla (most of them The number of species ill a location can be determined
prokaryotes) recognized by Margulis and Schwartz simply by counting them, but a simple tabulation does
(1998), about 69 have marine representatives and 55 not serve all purposes. The commonly used measures fall
have terrestrial ones. Of the 35 extant phyla of multicel- into three major categories that emphasize, respectively,
lular animals, 34 have marine representatives, and 16 of numbers, evenness, and differences. The number of
these are exclusively marine. From this perspective, species of organisms present in an area, habitat, or evolu-
preservation of marine biodiversity is more important tionary lineage, is called species richness. Measures of
than might be suggested simply by comparing the num- evenness, or species diversity indices, weigh species by
bers of species in marine and terrestrial environments. some index of their importance, such as their abundance,
The preservation of evolutionarily distinct lineages productivity, or size. For a given number of entities, the
above the level of species is important for a number of highest value of evenness is obtained if all are equally
reasons. First, the evolutionary potential of life depends abundant. The degree of difference among species (or
upon the distinctness of evolving lineages, not just the populations, or biotas) is measured by indices of similar-
number of species. Lineages that have been evolving ity. These indices are also used to assess the degree of ge-
separately for long periods of time have many unique netic similarity among evolutionary lineages, or the di-
genes and gene combinations that would be lost were versity of habitat types across landscapes or ecosystems.
those lineages to become extinct. Closely related species, Indices of both diversity and species richness are
on the other hand, share nearly all of their alleles. For ex- commonly used in ecological, biogeographical, and con-
ample, we share an estimated 99°/o of our DNA with servation studies because each type gives useful infor-
chimpanzees. Second, evolutionary lineages are store- mation not provided by the others. Ecologists commonly
houses of information about the history of life. Scientists use diversity measures to assess the adverse effects of
can read and interpret this information with increasing pollution and other types of environmental distur-
accuracy using modern phylogenetic methods. Third, bances. Typically a stressed ecological community expe-
the integrated functioning of ecosystems depends, in riences losses of species and increases in abundance-
part, on the variety of species in them. For example, mi- and hence dominance of a few species. Multimetric
croorganisms have more diverse mechanisms for ob- biological indexes quantify diverse biological changes,
taining energy from the environment and decompose a and an examination of the nature of the changes yields
wider variety of substances than multicellular organ- clues as to their causes (for example, the Index of Biotic
isms. Without microorganisms, ecosystems would be Integrity, widely used to evaluate stream condition; Karr
unable to provide, at high rates, such goods and servic- 1991). Understanding causes is essential for the design of
es to humankind as absorbing and breaking down pol- management plans that have the potential to counteract
lutants, storing and cycling nutrients, forming soils, and
negative effects of stresses and restore the systems to
maintaining soil fertility. The numbers and kinds of their former states.
fuels, construction materials, and medicines we obtain Conservation biologists usually use unweighted meas-
from nature depend upon the evolutioD.ary distinctness ures of species richness because the many rare species that
of species. Finally, the aesthetic benefits we receive from characterize most biotas are often of greater conservation
nat~re are strongly correlated with the variety of living interest than the more common ones that dominate
Global Biodiversity 37

weighted indices of diversity. In addition, because accurate Although ecological communities typically gradually
estimates of population densities on geographic scales sel- grade into one another, recognition of major divisions is
dom are available, species lists are the only available infor- useful for analysis. Based on the composition of the bio-
mation for most areas. tas of different regions, Earth has been divided into eight
However, species richness is a poor indicator of dif- major biogeographic realms: the Palearctic, Afrotropic,
ferences among biological communities in the degree to Indo-Malay, Australasia, Oceania, Nearctic, Neotropic,
w hich they retain significant species interactions. Nei- and Antarctic (see Plate 1). Each of these biogeographic
ther the abundance nor the functional roles in a commu- realms contains a number of biomes, large ecological
nity ate indicated through a total species count, and thus units that are identified on land on the basis of the dom-
w e do not have an indication of which species play sig- inant type of vegetation (see Plate 1), and, in the sea, on
nificant roles or those that play lesser ones. Further, ocean currents and spatial patterns of primary produc-
species richness does not differentiate between native tivity (Figure 2.3).
and nonnative species, and the species richness of de- Terrestrial biomes change along environmental (pre-
graded communities may be increased initially by an in- cipitation, elevation or temperature, and latitudinal) gra-
flux of nonnative species. Finally, we may most wish to dients (Figure 2.4). At temperate latitudes, commonly
know the role and importance of variability in commu- recognized biomes change along a precipitation gradient
nities, the dynamical responses of communities to dis- from mesophytic forest, to woodland, tallgrass prairie,
turbance, or several other factors that influence both the shortgrass prairie, and finally, desert. With increasing
current status of a community or its ability to recover fol- latitude, mesophytic forests change into conifer-domi-
~owing disturbances. The extent to which communities nated boreal forests, and eventually tundra. In tropical
can recover from disturbance on their own, or may re- regions, biomes change along precipitation gradients
quire intervention such as ecological restoration, is im- (rainforest, evergreen seasonal forests, dry forests, thorn
portant for conservation biologists to understand. Yet, woodland, desert scrub, and desert) and elevational gra-
other measures of species diversity, though they might dients (rainforest, montane rainforest, cloud forest, elfin
indicate differences in the degree of dominance of the woodland, paramo). Finer divisions of biomes into
community by one or more species, also can not distin- ecosystem types are based on drainage, soil type, slope,
guish among these ecologically important differences. and species composition. Classifications of ecosystems
-nfortunately, at present we do not have many good in- and biomes are inevitably somewhat arbitrary, but they
dicators of the functioning of these more complex as- identify ecological units that are useful for a variety of
pects of biological communities (see Chapter 18). purposes. Moreover, the richness of human experience
Biological communities are of conservation interest depends upon the richness of biomes as well as the rich-
largely because all species are influenced by their interac- ness of species.
:ions with other species. Evolutionary change takes place Recently, a group of scientists led by the Conservation
:n the context of ecological communities, and thus alter- Science Program at the WWF-US reclassified Earth's bio-
ations to communities will change the course of evolution. mes into 867 terrestrial ecoregions (see Plate 2; Olson et
_ lore immediately, changes in the abundance or presence al. 2001). An ecoregion is a relatively large area contain-
of individual species can have strong effects on the re- ing a distinct assemblage of natural communities and
:::taining species in a community, which can cause species ecological conditions, characterized by a dominant and
~ tinctions or other disruption (see Chapter 3). Thus, con- widespread assemblage of species. With contributions
servation of biological communities may be a necessary from over 1000 scientists, the WWF map of ecoregions
means to conserving species and evolutionary processes. has become the most comprehensive description of
higher-level landscape diversity available, and is a wide-
ly accepted base map for conservation planning.
Ecosystem and Biome Diversity, and the Preservation of representatives of all habitat types in
all ecoregions is necessary for .preservation of species,
orld's Ecoregions

because species will not survive without sufficient quan-

Oassification of terrestrial ecological systems typically has tities of their natural habitats. Captive propagation can,
:..een based on the shapes and life-forms of the plants that and does, serve a role in keeping species alive for short
~ ominate the structure of those communities (von Hum- periods until they can be reintroduced into the wild (see
~oldt 1806; Raunkaier 1934; Dansereau 1957; Halle et al. Chapter 15). But captive propagation is of little ultimate
:978). Koppen (1884) even used plant life-form distribu- use if there are no suitable habitats into which to reintro-
__;on to define climates. Holdridge's (1967) widely used life duce the species. Managing species in zoos and botani-
zone system, on the other hand, is based entirely upon cli- cal gardens is expensive, and an animal in a cage or a
~..atic variables. The diversity of schemes reflects the var- plant in a garden lives isolated from its natural physical
:ed goals of biologists who classify large ecological units. 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.)
-10 Tundra

-... -5

Cold ::l
temperate ro .f.j

0... 5


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

~ axes of mean annual precipi-

Warm 20 tation. Within the region
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.)
Global Biodiversity 3.

Ecosystems are often a conservation target because tion of new species and the rate of loss of existing
--.:tical functions that support human populations are species. The rate of species formation depends both on
Ellilintained by this level of biodiversity. As ecosystems events that generate barriers to gene flow (vicariance
-:e degraded by human activities, their ability to provide events), and on the number of species the more
,...__:tical ecosystem services and to support a broad array species, the more ranges there are to be separated, the
: iological functions is compromised, as we discuss in more species available to cross existing barriers, and the
.: e:ail later in this chapter. Biomes across the world are more opportunities for divergence. Rates of loss of
-~g modified at different rates, and by different forces species depend on the kinds and severity of distur-
see Chapter 6). Our focus at the biome level is often to bances, and the strength of factors that permit species to
.. . . ._. erstand the drivers of change in these areas, and how coexist. Many such factors exist but their operation de-
u:r options for mitigating these changes may be used for pends fundamentally on the existence of trade-offs in the
... ~rvation (Millennium Ecosystem Assessment 2005). abilities of species to deal with the factors that influence
their abundance and distributions.
The fossil record, though very incomplete, provides a
Species Richness over Geological Time rough measure of trends in species richness during the
_-:3 number of species on Earth at any moment in time history of life on Earth (Table 2.2). Cellular life in the form
~ ~ e result of the difference between the rate of produc- of bacteria evolved about 3.8 billion years ago (bya); eu-

LE 2@2 Earth's Geological History

40 Chapter 2

karyotic organisms probably evolved about 2 bya. Al- proximately 40,000 species of marine invertebrates have
though many species must have lived in ancient times, been described from the Paleozoic and Mesozoic eras, a
species richness appears to have been low during the first number that increased to about 250,000 in the late Ceno-
2 billion years of Earth's existence. During the late Pre- zoic era. The fossil record of terrestrial animals is much
cambrian, the richer Ediacaran fauna, consisting of poorer, especially among such species-rich groups as in-
strange frond- and disc-shaped, soft-bodied animals and sects, which fossilize poorly. The fossil record of verte-
some forms that appear to be arthropods and echino- brates, particularly mammals, is much better. It indicates
derms, evolved. The first explosion of biodiversity took th at richness, as measured by the number of orders, is
place during the early Cambrian period. Some Cambrian slightly higher today than earlier in the Cenozoic.
species may be members of phyla that left no surviving Terrestrial vascular plants appeared by the early Sil-
descendants. As measured by the number of phyla, ani- urian and their richness increased rapidly during the De-
mallife may have been more diverse during the Cambri- vonian when seed-bearing plants first appeared. Species
an period than at any time since (Gould 1989). richness has continued to increase overall but the num-
The fossil record for marine invertebrates with hard bers of species of ferns and gymnosperms has decreased,
skeletons is good enough to provide a general picture of whereas the number of species of angiosperms has in-
the number of evolutionary lineages present at different creased dramatically (Figure 2.6).
times in the past. The Cambrian explosion was followed,
about 60 million years later, by the extensive radiation of 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
greatly reduced 300 million years later by the great Per-
@ mian mass extinction, which was followed by the Triassic
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

0 700

Jurassic (')

200 600
ro Triassic
::>... Permian
* 500 Angiosperms
'-4ool rJ)
0 Q)
~ 300 . '
0 0.. 400


...-4 '-4ool
~ ~

D evon1an
. ~
~::s 300

N 200 Gymnosperms
Silurian 9.
(') •

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
:5actors: mass extinctions, increasing separation of land ><

:nasses across Earth, and evolution of new life forms and .....
* * '

types of species interactions. Mass extinctions tend to be u

::allowed by increases in rates of speciation because ~

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

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

(A) 22,000


18,000 Amphibians
lr --. , I Birds
UiiU Mammals
• !""'( I I Reptiles
......... 12,000

~;j 10,000

z 8000




(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
broadleaf forests; TCF, tropical and subtropical coniferous forests; TeBF, temperate
• !""'(
Q) 6000 broadleaf and mixed forests; TeCF, temperate coniferous forests; BF, boreal
forests/taiga; TG, tropical and subtropical grasslands, savannas and shrublands;
u TeG, temperate grasslands, savannas, and shrublands; FG, flooded grasslands and
Q) savannas; MG, montane grasslands and shrublands; T, Tundra; MF, Mediterranean
c: forests, woodlands, and scrub; D, deserts and xeric shrublands; M, mangroves.
4000 (Modified from Millennium Ecosystem Assessment 2005.)
z 2000


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

Global Biodiversity 43

Africa •
40 I I I 4 I
I 1

X;- 30
-- Chile
! ;...


:; .u

> (l)
------ ~ Chile

10 c 10

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

lo a iod· rs1 45


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 ·
• 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

0 1000 2000 Miles
I, , , , J,, , ,I,,,,J , , , ,J
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
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- ~
• G
ple, Arctic waters have about 100 species of tunicates, 400 ~
H 100 ----
u 'C\
q., .... ' 0 0 "a
species are known from temperate waters, and more than z •
o/ "'~
600 species inhabit tropical seas. The richness of species,
. 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
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

190 . 80
200 100
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

• •

.~ 100 Puerto •Cuba •• • •


100 • •• •• • ••
Jamaica Sunda Islands


~;J 10
• •
Montserrat 10 •

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 rates of extinction, and thus, also is correlated with high
correlated factor is area, which can influence both speci- species richness. Large areas can facilitate the long-term
ation and extinction rates. survival of populations, which in tum, allows the accu-
One of the first ecological relationships to be estab- mulation of the genetic differences that may prevent hy-
lished empirically was the relationship between area and bridization should the populations come together at
number of species (Arrhenius 1921). This relationship~, some time in the future. This favors greater speciation
which is true for both continental and island biotas rates. Populations of vertebrates in lower latitudes show
(Rosenzweig 1995), is commonly expressed using a greater genetic differentiation, which is consistent with
power function of the form greater capacity for speciation in the Tropics (Martin
2 and McKay 2004). Therefore, area is probably a contrib-
S = cA
utor to high species richness in the Tropics, but so are
where S is the number of species, A is the area, and c and productivity and available energy, as discussed in the
z are constants fitted to the data. On a logarithmic scale, next section.
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-
Species Richness-Energy Relationships
ganisms revealed that values of z range from 0.15 for Available energy affects local species richness because the
continents, to between 0.25 and 0.45 for islands associat- more energy that is available, the more biomass per unit
ed with a continent, to 0.19 for comparisons among con- area that can be supported. More biomass enables more
tinents or biogeographic provinces (Rosenzweig 1995). individuals to coexist in an area, potentially resulting in
The regular relationship between the size of an area and more individuals per species and, hence, lower extinction
the number of species it supports, which is most readily rates. If high productivity is combined with moist tropi-
observed using island data, was a key empirical general- cal conditions, organisms expend relatively little energy
ization in the development of the theory of island bio- maintaining their temperatures and moistures at appro-
geography (MacArthur and Wilson 1967). priate levels, so even more energy is potentially available
The Tropics have a larger climatically similar area for reproduction (Connell and Orias 1964).
than any other ecological zone because the surface area More available energy might also enable species to
of latitudinal bands decreases toward the poles and be- persist on relatively specialized diets (Gaston 2000).
cause temperatures change relatively slowly between There is some evidence that tropical animals may, on av-
the equator and 20°N and 20°5 latitudes. Larger area is erage, have more specialized diets than their temperate
correlated with higher rates of speciation and lower counterparts, but much more research is needed to es-
Global Biodiversity 49

:ablish the degree to which this is true and to identify the available energy to plants, and tree species richness in
~-:-:mensions along which dietary specialization has North America (Currie and Paquin 1987). Similarly, Gen-
eYolved (Beaver 1979; Marquis and Braker 1994). try (1988) demonstrated a strong correlation between
Primary production,-the major source of available en- plant species richness in Neotropical forests and ab-
=::g)" is determined primarily by temperature, moisture, solute annual precipitation, a variable strongly positive-
~1d nutrient availability. Highest terrestrial production ly correlated with productivity.
:S found in regions with high rainfall and year-round However, the relationship between productivity and
...-ar m temperatures. Production drops with elevation species richness is not simple. Many of the world's most
~ecause of lowered temperatures. In many areas of the productive systems, such as estuaries, seagrass beds,
.·.-orld, the amount of precipitation determines total pro- and hot springs, are species-poor. Conversely, plant
iuction. In arid and semiarid regions, annual production species richness is higher in semiarid regions with nutri-
:an be predicted fairly accurately from the amount and ent-deficient soils than in similar areas with richer soils.
~.; tribution of precipitation. The remarkably rich plant communities of the Cape re-
Marine production is also limited by temperature, but gion of South Africa (fynbos) and the extreme southwest
nore often by shortage of nutrients in surface waters corner of Australia (Figure 2.14) are found on highly in-
Longhurst 1998). Organisms and nutrients sink in the fertile soils (Kruger and Taylor 1979; Bond 1983; Rice and
- -ater column. Therefore, processes that keep organisms Westoby 1983). Nitrogen levels in those soils are an order
'-Lose to the surface where photosynthesis is possible, and of magnitude lower than in lowland California and
:hose that bring nutrients and organisms back to the sur- Chile (Specht and Moll1983), where species richness is
:=ace, are major determinants of marine productivity. Pro- lower. Evidence suggests that soil infertility itself is a
. uctivity is highest in areas where vertical turbulence major contributor to the inverse correlation between soil
a p welling) is produced by friction between tidal streams fertility and plant species richness because low fertility
and the seabed and at the confluence of two major ocean favors abilities to exploit slightly different microhabitats
currents; it is lowest in the open ocean (Koblentz-Mishke more efficiently (Tilman 1982, 1985; Cowling et al. 1992;
et al.1970; Bunt 1975; Longhurst 1998). Primary produc- Willis et al. 1996). Fynbos plants are remarkably variable
:i\i ty also is higher near continental margins due to the in leaf morphology (Cody 1986) and growth phenologies
:runoff of nutrients from the land. Finally, primary pro- (Kruger 1981; Cowling 1992). These differences are prob-
uction decreases with depth because light levels de- ably correlated with a corresponding diversity of car-
crease rapidly with depth even in clear water. bon-fixing strategies. In addition, the low palatability of
The hypothesis that available energy contributes to leaves of plants growing on nutrient-poor soils results in
species richness is supported by a correlation between a rapid accumulation of flammable biomass, leading to
___ roductivity and species richness along both latitudinal high frequencies of fires in these summer-dry climates.
and altitudinal gradients. A strong correlation exists be- Soil infertility also may favor high species richness in-
uveen realized annual evapotranspiration, a measure of directly via interactions with seed dispersers. Compared

20 Figure 2.14 Numbers of species of
Eucalyptus and Acacia in southwestern
40 Australia. Notice how many species in
these two genera can be found living
soJ 40
50 55 in proximity. Similar patterns are
found among many other genera of
plants in southwestern Australia.
Acacia Eucalyptus (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 Q)
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.
20 40 60 80 100
1991) and it has unusually high ant species richness (An-
Salinity (0/o)
dersen 1983; Greenslade and Greenslade 1984). Ant-dis-
persed seeds are also common in South African fynbos Figure 2.16 Estimated species richness of aquatic inverte-
(Milewski and Bond 1982). In combination, these factors brates living in waters of different salinity throughout the
world. The two peaks correspond to fresh water (<2°/o salinity)
could result in very high a-species richness and unusu- and seawater (30°/o-40°/o salinity). (Modified from Kinne 1971.)
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). Most highly productive but species-poor systems are
There is a consistent unimodal relationship between distributed as relatively small, fragmented patches
species richness and depth for many marine taxa. Maxi- whose physical environments differ strikingly from those
mum richness of most groups is found at depths be- of the surrounding, more extensive ecosystems. Evolu-
tween 2000 and 4000 meters (Rex 1973, 1983). In contrast, tionary biologists believe that the combination of major
coral species richness peaks at depths between 15 and 30 physical environmental differences and isolation of the
meters because corals, which obtain much of their ener- patches results in fewer species of organisms evolving
gy from photosynthetic algae embedded in their tissues, adaptations to those unusual environments. For the same
are confined to the photic zone (Huston 1994). reason, species adapted to the more common surround-
In addition, in many ecosystems, species richness in- ing environments are less likely to survive well in those
creases with primary production at low production lev- rare habitat types. Both fresh water and seawater are
els, but declines at high levels of primary production widely distributed, but waters of intermediate salinity
(Huston 1994). This phenomenon has been called "the and waters that are more saline than the oceans are rare.
paradox of enrichment" (Rosenzweig 1971) because ad- The species richness of aquatic invertebrates throughout
dition of fertilizer to aquatic and terrestrial plant com- the world parallels this pattern (Figure 2.16).
munities often results in sharp decreases in species rich- Available energy may influence species richness indi-
ness (Huston 1980). rectly via vegetation structure. The structural complexity
of vegetation is strongly correlated with potential evapo-
transpiration and, hence, primary productivity. In most
1 terrestrial environments, plants provide most of the phys-
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-
• ....C 2 plex communities have a greater variety of microclimates,
a greater variety of resources, more ways in which to ex-
0 3 ploit those resources, and more places in which to find
Q) shelter from predators and the physical enviro~ent.
s ~ 4
The richness of species in most taxa is positively corre-
z lated with structural complexity. Structurally simple habi-
100 5 tats, such as the open ocean, grasslands, and cold deserts,
generally support fewer species of organisms than struc-
turally more complex communities, such as forests and
o~ -~------------~----------------~
0 0.1 1 2 coral reefs. Animal groups that exploit the environment in
Area (ha) three dimensions are most sensitive to plant community
structure. A positive correlation between foliage height di-
Figure 2.15 Plant species richness in Borneo and in south- versity and bird species richness exists in many plant com-
western Australia. Curves 1, 2, and 3 represent trees greater
than 10 em in diameter mmixed forests in Borneo. Curves 4 munities on all continents (MacArthur and MacArthur
and 5 are for plots in southwestern Australia. (Modified from 1961; MacArthur 1964). Similarly, the richness of species of
Lamont et al. 1984.) web-building spiders is positively correlated with the het-
Global Biodiversity 51

erogeneity in heights of the tips of vegetation to which (less specialization, greatervagility,larger ranges), par-
such spiders attach their webs (Greenstone 1984). ticularly in higher latitudes that were most affected by
In contrast, there is no consistent relationship between climate change (Dynesius and Jansson 2000). This com-
v egetation structure and lizard species richness in hot bination of selective factors in response to climate cycles
deserts in North and South America, Africa, and Australia could contribute to the latitudinal gradient of species
(Pianka 1986). The number of lizard species in similar riclmess we see today (Figure 2.17). At a finer spatial
h abitats in African deserts averages twice that in North scale, physical and biological disturbances clearly influ-
.i\merica; Australian deserts are about twice as rich as ence the number of species present in many ecological
African ones. The main differences are due to the presence communities. Small disturbances occur much more fre-
m Australia of non-lizardlike lizards and nocturnal quently than large disturbances. Physical perturbations
species. Only one nocturnal lizard is found at the North include heavy rains, strong winds, landslides, earth-
American sites, whereas there are four nocturnal species quakes, and fires. Biological disturbances, most of which
at the African and eight at the Australian desert sites. are small scale, include activities of predators and para-
Mammal-like species (the monitors) and worm-like sites, tree-falls, activities of competitors, and trampling.
species add to the richness of Australian desert lizard fau- Physical disturbances influence species riclmess by
nas. These differences relate primarily to the long-term destroying habitat structure, selectively killing individ-
evolutionary history of deserts on the three continents, uals of different species, and sterilizing soils. Physical
not to differences in today's vegetation (Pianka 1986). disturbances typically also have important indirect ef-
fects on biodiversity by altering the biological interac-
tions that affect species riclmess. For example, fire, one
Disturbance and Species Richness of the most important terrestrial physical disturbances,

One explanation for the latitudinal patterns of species kills plants and animals, destroys soil organic matter,
richness comes from considerations of the extreme dis- and redistributes nutrients, resulting in a new ecological
turbances caused by climate change over Earth's history community with a dramatically different physical struc-
(Dynesius and Jansson 2000). During the Pleistocene and ture and altered biological interactions.
before, periods of rapid climatic change (influenced by Theoretical and empirical studies both suggest that
orbital oscillations of Earth), which were more intense disturbances do not have consistent effects on species
toward the poles, clearly had enormous influence on richness. The hypothesis that lack of disturbances might
worldwide patterns of species riclmess. Rapid climate favor high species richness was originally advanced by
fluctuations caused increases in extinction rate as species Sanders (1968), and Sanders and Hessler (1969), who
were unable to adapt to new conditions, or migrate to noted that productive estuaries and continental shelves
more hospitable ones, and the more extreme fluctuations in most latitudes have few species of benthic animals,
toward the poles resulted in more extinctions at high lat- but the cold, dark, unproductive floor of the deep sea is
itudes. Importantly, these fluctuations also should have very species-rich. Sanders suggested that the deep sea
changed patterns of speciation, favoring characteristics supported many species because its environments had
that lead to lower speciation rates among the survivors 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
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. Low High
Biological interactions can also influence species rich- Disturbance frequency or intensity
ness. For example, competition could reduce a-richness
if some species exclude others from ecosystems. How- Figure 2.18 Model of the intermediate disturbance hypothe-
sis. Species diversity is lowest at high and low levels (fre-
ever, competition could increase a-richness by favoring quencies or intensities) of disturbance, and highest at an in-
finer habitat segregation among species (Rosenzweig termediate level. (Modified from Connell1978.)
1995). Predation can increase species richness if preda-
tors prey preferentially upon competitive dominants,
thereby preventing competitive exclusion. A well-known and temporal scales (MacArthur 1972; Ricklefs 1987). In
example is the increase in species richness in rocky in- other words, to understand why a particular set of species
tertidal communities of the Pacific Coast of North Amer- exists in a specific place, ecologists must study more than
ica. This increase is a result of selective predation by the local competition, predation, and mutualistic interactions.
sea star Pisaster ochraceus on the competitively dominant Larger-scale processes, such as dispersal, speciation, and
mussel Mytilus californianus (Paine 1974). On the other historical biogeography cannot be ignored.
hand, predation can reduce species richness by prevent- The ecological interactions that appear to influence
ing vulnerable species from living in an area. Experi- current patterns of species richness, in combination with
mental evidence exists for all of these outcomes, but how physical environmental factors, could set absolute limits
often and where these forces exert their influence are un- to species richness. In other words, we can ask if there
known (Orians and Kunin 1991). Analyzers of the litera- are limits to the number of species that can live in a
ture have reached different conclusions, in part because given ecological community. If so, are those limits regu-
they used different criteria for including studies in their larly reached? Is there a critical limit to overlap in re-
samples (Connell1975, 1983; Schoener 1983). source use that prevents more species from being ac-
Some researchers have hypothesized that many organ- commodated unless dramatic changes occur in the
isms are held at sufficiently low population densities such nature of available resources? Does the chemical warfare
that competition rarely occurs (Connell1975). Yet other re- between plants and herbivores set limits to species rich-
searchers have suggested that competition occurs prima- ness or does it offer possibilities for more species? Are
rily during unusually hard times, "competitive crunches," there limits to the size of mimicry systems, and do mim-
when resources are scarce (Wiens 1977). Which of these icry systems allow more species to persist in ecosystems
two scenarios applies depends upon whether low popu- than would be possible without them? Have the rich-
lation densities are caused by harsh physical conditions or ness-generating interactions between plants and their
by scarcity of consumable resources. pollinators and between plants and their seed dispersers
Often species richness is highest at intermediate lev- been exhausted? To answer these questions research
els of disturbance (Figure 2.18). The Intermediate Dis- must be carried out at many different spatial scales. .
turbance Hypothesis states that physical disturbances For many diverse taxa, including gall wasps (Cornell
or predation should augment species richness where 1985), birds (Ricklefs 1987), tiger beetles (Pearson and Ju-
other sources of disturbance are few, but should reduce liano 1993), fishes (Griffiths 1997), and primates (Eeley
species richness when exogenous disturbance is high and Lawes 1999), local species richness appears to in-
(Connell 1978; Abugov 1982). If disturbances are rare, crease along with regional species richness, but at a slow-
species richness may decline owing to competitive ex- er rate. This pattern suggests that rigid limits to local
clusion. If disturbances are common, many species may species richness are not set by competition, predation,
be unable to complete their life cycles during the short and parasitism (Comell1999). Thus, although ecological
intervals between disturbances. interactions are typically very intense in local communi-
ties they apparently influence abundances more strongly
Interactions between local and than numbers of species. Moreover, their influence on the
regional species richness numbers of coexisting species appears to be weaker than
The structure of local ecological communities may be de- the influence of regional processes. A major challenge in
termined by loc~ interactions among species and their en- ecological research is to determine how regional process-
vironments, and by processes operating at larger spatial es act to determine local species richness.
Global ~iodiversity 53

The importance of biodiversity

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

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

A ......
.... --- ...

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

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

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


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

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

0 20
--- 40 60 80 100
l( ater flow, carbon sequestration, decomposition, and Richness (0/o)
p rimary and secondary production (Table 2.4). Claire
Figure 2.19 Ecosystem function could increase quickly with
Kremen discusses the importance of one of these servic- species richness (A), linearly (B), or as an accelerating func-
es, pollination, in Box 2.2. tion (C). Increases in ecosystem function with species richness
If clean air and water, fertile soils, pollination of crops, could occur due to increases in ecological functionality either
and other "services" depend upoh the maintenance of a because of changes in the number, abundance, or functional
b road diversity of native species, this becomes a power- traits held by the species in the community. (Modified from
Kremen 2005.)
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- interact to influence the magnitude and stability of
rich plots than in species-poor plots. Many observation- ecosystem processes (Figure 2.19). For example, are rela-
al, experimental and theoretical studies have found that tively few or many species required for high values of
species-rich communities have higher values of some some ecosystem process (curve A versus C in Figure
metric of ecosystem functioning than species-poor com- 2.19)? Or is the relative abundance of key species, or the
munities. If generally true, then preserving as many diversity of functional groups, most strongly related to
species as possible should be a primary goal. ecosystem processes?
Yet, ecologists still know little about how attributes of Considerable debate has sprung up regarding the inter-
communities including species richness, relative abun- pretation of experimental investigations of relationships
·d ances, and functional traits of individual species (such between species richness and ecosystem processes. The
as ability to fix nitrogen or seed dispersal effectiveness)- importance of rare species to ecosystem functioning is
poorly known, and many studies show that a particular
TABLE 2.4 Examples of Ecosystem Services 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-
Source: Costanza et al. 1997 and Millenium Ecosystem Assessment munities are complex enough to show that functional di-
2005. versity of species (i.e., differences in the roles they play in
5 Chapt r 2

Global Biodiversity 55

the ecosystem), and not species richness, influences the

The Future of Biodiversity Studies
rate of decomposition (Heemsbergen et al. 2004). The incomplete state of our knowledge of the identities,
Increasingly researchers are using large-scale removal taxonomic relationships, and distributions of the vast ma-
experiments and comparative studies of communities jority of the world's organisms means that the primary
1vith varying species membership and abundances to work of cataloging biodiversity is yet to be done. Today
gauge the role of individual species as well as diversity relatively few scientists are being trained as taxonomists.
on ecosystem functioning (e.g., Kremen et al. 2002a,b; Therefore, increasing the cadre of competent taxonomists,
Diaz et al. 2003). A number of biologists believe that eco- particularly in tropical nations, is an important goal
logical differences among species should create the con- (Mikkelsen and Cracraft 2001; Gotelli 2004). How this in-
ditions for an increase in ecosystem process with diver- ventory should be carried out is the subject of much de-
sity, while others believe that only the attributes of a few bate. Some biologists have committed themselves to an
d ominant species will control that relationship (Figure intense global surve~ aimed at the discovery and classifi-
2.20). A growing consensus focuses on the importance of cation of all species (e.g., the Species 2000 Project). Others,
differences in the ecological roles of species, and the joint pointing to the shortage of people, funds, and time, be-
contributions these differences make (Diaz and Cabido lieve that the only realistic hope lies in the rapid recogni-
2001; Lareau et al. 2001). However, measuring the signif- tion and preservation of those threatened habitats that
icance of a species for a particular ecosystem process is contain the largest number of endemic species (Conserva-
m uch harder than counting numbers of species or esti- tion International strategy), or to conserve representatives
m ating population densities. Full understanding of how of the habitats of every ecoregion (World Wildlife Fund
biodiversity affects the ability of an ecosystem to provide and The Nature Conservancy strategies). They give the in-
goods and services over time awaits much further work. ventory task a lower immediate priority.

Species pool 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.)

Low richness
Low functional diversity

Dominance of species Complimentarity among Complimentarity among species

\vith particular traits functional groups of species with different functional roles

Lower Functioning ECOSYSTEM PROCESSES Higher Functioning

56 Chapter 2

E. 0. Wilson (1992) strongly advocated a strategy that While a reasonably complete inventory of the biota of a
combines global surveys designed to achieve a complete region, or significant components of it, is being achieved, a
biodiversity inventory in 50 years with special attention monitoring program also needs to be established. Such a
to areas of unusually high species richness ...a nd en- program should be designed to detect trends in biodiver-
demism. His strategy has three components. The first is sity and to identify impending problems to which atten-
a Rapid Assessment Program (RAP) that would investi- tion should be directed (National Resource Council2000).
gate, within a few years, poorly known ecosystems. RAP Without a monitoring program society cannot know if its
teams would be formed, consisting of experts on groups efforts to preserve biodiversity are succeeding or why
such as flowering plants, reptiles, mammals, birds, fish- and where they are failing.
es, and butterflies that are well enough known to be in- Human societies will need to greatly increase their in-
ventoried quickly and accurately. These groups would vestments in biodiversity studies during future decades
then serve as proxies for the entire biota (see Essay 2.3 by if we are to understand the patterns of biodiversity more
Lily Rodriguez; for a discussion of applications to prior- fully. The scope of the untapped wealth residing in bio-
ity setting in biodiversity conservation see Chapter 14). diversity, as well as the importance of inventorying bio-
The next stage would be to establish research stations in diversity and understanding ecological relationships
areas believed to be major hotspots of diversity. Invento- among species, is unappreciated. It is doubtful that hu-
ries and ecological studies would then be carried out mans can devise a sustainable future without a more
there and in surrounding regions. The third stage, with complete knowledge of biodiversity.
a time frame of 50 years, would combine the inventories The relationship between area and species richness
from RAP and the intensive studies at a small number of has major practical implications for the location, design,
research stations with monographic studies of many and management of parks and reserves that are estab-
groups of organisms to provide a more complete picture lished and maintained to preserve biodiversity. There is
of global biodiversity and its distribution. 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- pating in inventories over the last During those overflights, team mem-
ment Program (RAP, of Conservation decade. bers select points for easy access to as
International, CI) or as Rapid Biological Multidisciplinary teams are formed many different types of habitats as pos-
Inventories (RBI, by The Field Museum for each expedition, usually with a core sible, verify any unique characteristics
of Chicago), rapid inventories are a rel- group of well-recognized scientists (tax- seen in the images, and assess impacts
atively new conservation tool designed onomists and ecologists), combined of human activities in the area to be
to assess the biological importance of with resident field biologists, who are surveyed. Final decisions on sites are
priority sites for conservation. Using both specialists and specialists-in-train- made based on accessibility and trans-
short field trips, usually of no more ing. Each specialist selects the methods portation. Although relatively expen-
than four weeks, these inventories eval- and groups that will be the focus of its sive, helicopters are the best way to
uate the biodiversity values of sites of work, based on abundance, rarity or reach remote areas, and to relocate the
global conservation interest, providing uniqueness, and capacity to document team between main campsites. To maxi-
scientifically based recommendations the group in a short period of time. mize time for the inventory, camps and
for prompt establishment of protected None of these inventories are meant to trails should be prepared prior to
areas and conservation action plans. yield complete information on the site. arrival of the team. While preparing the
Identifying conservation targets and Planning the trip and identifying trails (usually loop trails), satellite
their threats, and generating recom- 2-4 campsites are crucial steps to campsites can be prepared and used by
mendations for immediate conservation ensure exciting results. Satellite the team according to distances, inten-
action are the main goals of these imagery is used first to assess the gen- sity of work in different habitats, taxa,
inventories. eral area and locate points of main and technique requirements.
RAP or RBI results rely highly on interest. The preliminary selection of Different approaches are needed for
the expertise of the team undertaking sites to inventory is made based on this each of the principle taxonomic groups
the inventory, as there is no single first step, combined with a review of addressed in rapid inventories. Plants
methodology that is or can be used in any previous research in the area or provide the best characterization of
all places with all taxa. Here, I provide nearby locations, as well as local habitat types and communities associ-
some insights and recommendations knowledge. Overflights to the area are ated with the area and are therefore a
based on my experience after partici- then made by a few team members. must in any rapid inventory. Transects,
Global Biodiversity 57

q:uadrat plots, collections, and photo- the inventory with rare records or new regional faunas. Because aquatic and
graphs are some of the techniques that species and the presence of suites of terrestrial turtles as well as crocodilians
can be used to characterize the flora, species can help relate basins and are frequently hunted they are always
providing quantitative information, aquatic communities (Figure A). included and evaluated as target
::eld observations and general collec- Insects, because of their immense species for conservation actions. Tissue
::ons of the diversity of plants. Focal species richness, can rarely be used as a collection of amphibians (especially in
groups, such as ferns or understory group for rapid inventories. The best mountain ecosystems, where popula-
~egetation, are important as they pro- choice is to take experts on specific tax- tions declines due to disease appear to
-tide better resolution of habitat differ- onomic groups that are best known occur more often) is now recom-
ences than do trees. Groups of special and easily seen in few days. Butterflies, mended to be able to evaluate possible
-.--alue such as orchids may also add to beetles, dung beetles, and spiders are causes of decline in amphibian popula-
~.,_e richness of the results. some candidates for rapid inventories, tions.
Birds are one. of the best-documented but, as always, this will depend on the Geology, soil diversity, freshwater
groups around the world. Because of availability of experts and the scope of characteristics, and substratum are sur-
:heir high species numbers, and the fact knowledge of the group. veyed to understand and document the
that the range, distribution, and biology Mammals (especially large mam- uniqueness of the area and to indicate
is reasonably well known for most mals) experience heavy human pres- species distributions and plant commu-
species or groups of species, birds are sures and are important to document, nity types. An additional but extremely
used as habitat indicators. Avian data providing a primary indica tor of the important aspect is to include human
can be used to provide quick compar- conservation quality of the area. Tracks dimensions in the rapid inventory
isons among sites, and to guide consid- and visual encounters are used most through rapid social asset inventories.
erations such as connectivity, size of the frequently to document mammalian This consists of visiting the communi-
nrotected area, and conservation status
species. However, small mammals such ties surrounding the site surveyed, and
and perturbation of the site. Most com- as rodents and bats are speciose groups is achieved through a diversity of tech-
monly used methods involve auditory that if specifically surveyed can pro- niques (interviews of local leaders,
surveys along trails, including song vide a more detailed evaluation of the authorities, key people of the commu-
recording and point counts. Assessing site. A trade-off between time and nity, and focal groups; meetings with
the status of local populations of hunted value is usually applied here, with bats the whole community; and visits to
species is an important step for manage- (which are easily censused with mist their crop fields), the team explores the
ment recommendations. nets) being one of the best choices. history, demography, economy, social
Fishes are the main community Because of their sensitivity to distur- organization and structure, and the use
associated with aquatic habitats. Often bance, frogs are usually included in of natural resources. While identifying
easy to collect, they can be captured rapid inventories. Easily located by patterns of social organization and
1-vith small nets that are for scientific their calls, auditory transects and opportunities for capacity building,
use only. A high percentage of the fish visual encounters are the most com- these· inventories contribute to hone the
community can thus be surveyed. mon techniques employed to survey recommendations, and to engage local
Interviewing local people with photo- for frogs. Snakes, because of their low participation for future conservation
graphs in hand provides rich informa- population densities and secretive action.
tion on consumed species and seasonal habits, are difficult to include; however, Rapid inventories should not be
variations. Usually fishes contribute to they can add value as indicators of 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
Figure A Members of a RAP team examining fish specimens collected to them but that can be of high scien-
that day. (Photograph courtesy of L. Rodriguez.) 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
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
Presenting results to local, regional,
and national key players is another
important issue. Immediately after gatoni
completing fieldwork and before leav- Tambopata
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 over the long term (Figure 2.21). Each megareserve in-
those species with the largest aerial requirements (dis- cludes natural areas and areas managed for economical-
cussed in Chapters 7 and 14). Many of the national parks ly valuable products. Some conservation areas remain
of the United States have already lost their largest mam- the homes of indigenous people who continue to use the
mal species, and the trend continues (Newmark 1987). environment in their traditional ways. The largest of the
Therefore, additional studies of the species-area rela- Costa Rican reserves, La Amistad Biosphere Reserve, is
tionship and its causes are essential to inform manage- a mosaic of more than 500,000 ha and includes three na-
ment of protected areas. Meanwhile, existing knowledge tional or international parks, a large biological reserve,
of species-area relationships is being used to guide es- five Indian reservations, and two large forest reserves.
tablishment of protected areas. A notable example is the On a theoretical level, biodiversity studies continue to
megareserve system of Costa Rica, which is designed to be needed to resolve the many uncertainties surround-
preserve about 80°/o of the biodiversity of the country ing the historical and present-day ecological processes
Global Biodiversity 59

Figure 2.21 The extent of Costa

Rica's conservation areas as of 1996.
Nicaragua Shaded areas indicate protected areas,
which are organized into 10 distinct
conservation areas.
Caribbean Sea

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


that determine today' s patterns of biodiversity. Modern versity on Earth that so fascinate us today. This knowl-
phylogenetic techniques that enable systematists to de- edge is also being used to attempt to reduce the rates of
velop soundly based phylogenies are being combined species extinction and to restore landscapes so that they
w ith biogeographical studies to provide a more com- can continue to support the array of species originally
plete picture of the history of the distribution of life on found in them and the evolutionary processes that gener-
Earth. Processes operating over ecological time frames ate new species. For example, to promote survival of ex-
are increasingly being studied using manipulative ex- isting species and the evolutionary processes that gener-
periments in which some or all species are removed ate new species, devoting large areas to biodiversity
from restricted areas, or species are introduced. Many preservation will be critical.
.unatural experiments," are being studied to gain ecolog- Recently, the Millennium Ecosystem Assessment
ical insights from them volcanic eruptions that elimi- (MA), a large coalition of international development and
nate the biotas of islands, the massive inadvertent move- conservation organizations, governments, and scientists
ment of species around the world by human travel; has come together to assess the status of Earth's ecosys-
deliberate introductions for agricultural, aesthetic, or tems, the goods and services they provide, and the like-
pest-control purposes; and habitat fragmentation by ly effects of potential pathways of human economic de-
conversion of natural .landscapes to ones dominated by velopment on the future provisioning of these services
highly modified communities that are managed to chan- and human well-being (Figure 2.22). The MA focuses
nel most of their productivity to human uses. Such sce- both globally, and on subglobal regions of particular
narios provide opportunities to examine the results of concern due to the difficulty of human existence or po-
manipulations over longer time frames than is possible tential for serious declines in human welfare in these re-
with investigator-initiated experiments. gions. Published in 2005, this is the first comprehensive
Studies of the influences of human activities on species assessment of the condition and status of global ecosys-
distributions and species richness are adding rapidly to tem services. The information summarized in the MA
our understanding of the roles of the varied processes that will be used to guide development policy both regional-
interact to cause the patterns in the distribution of biodi- ly and globally. In addition, the MA will help focus re-
Global Biodiversity 61

common species typically dominate ecosystem 6. Climatic oscillations producing swift, radical changes
processes. in climate increased extinction rates and likely de-
_ 1.- \bout 1.75 million living and 300,000 fossil species creased speciation rates toward the poles, thus acting
have been described, but estimates of the total num- as a strong driver of latitudinal patterns in species
ber of living species range from 10 mi11ion to as high richness.
as 50 million or more. Although species richness is 7. Both physical (heavy rains, strong winds, landslides,
higher on land than in the oceans, 34 of the 35 extant earthquakes, and fires) and biological disturbances
animal phyla have marine representatives, and 16 (tree-falls and activities of predators and competi-
are exclusively marine. tors) influence species richness but their effects are
.i. Cellular life in the form of bacteria evolved about 3.8 highly varied. Most disturbances are small and di-
billion years ago; eukaryotic organisms evolved rectly affect only small areas, but they may influence
about 2 billion years ago. The first major explosion more distant regions via indirect effects. Strong com-
of biodiversity took place in the early Cambrian pe- petition and predation may eliminate species from
riod, and except during times of mass extinctions, particular areas but intermediate levels of predation,
the number of species has increased since then. by preventing competitive exclusion, may increase
ore species are probably alive today than at any local species richness.
other time in the history of life, even though some 8. Despite the demonstrated importance of local bio-
taxa had more species in the past than they do today. logical interactions, local species richness in many
The number of species present at any moment is the taxa appears to increase without apparent limit along
result of the difference in the rate of formation of with increasing regional species richness. This sur-
new species and the rate of extinction of existing prising result suggests either that local interactions
species. High rates of speciation have been favored exert strong effects on abundances of species but not
by mass extinctions, the breakup of the continents, on their numbers, or that the influence of regional
and the evolution of more diverse body plans that processes on numbers of species in local communi-
enabled animals to burrow, swim, and fly. ties is strong enough to override local influences.
~· Several broad patterns characterize the current dis- 9. The state of knowledge of Earth's biodiversity is so
tribution of species. Areas that have experienced poor that the primary work of cataloging biodiver-
long geographical isolation and that have great sity is yet to be done. Resources currently devoted to
topographic relief often support many endemic this task are inadequate, especially given the rate at
species. Tropical regions often have both high which species are becoming extinct. Better informa-
species richness and endemism, and islands have tion on biodiversity, its distribution, at1d its causes is
high endemism of both marine and terrestrial needed for wise management of Earth's biotic re-

species. sources. Further, research that allows us to under-
- . Among most taxa, more species live in tropical re- stand the contributions of biodiversity to human
gions than at higher latitudes. The large land area of welfare, and the effects of development on ecosys-
the Tropics contributes to this pattern. Species rich- tem services is needed to help us preserve the rich-
ness is also positively correlated with available ener- ness of biodiversity and human existence.
gy, which enables more individuals to be supported
per unit area, and with structural complexity, which . -

Pleases refer te the;~e}fsite~www.s:Un.au~r.c(i)Pl/~oq;p:l. s~¥

is positively correlated with available energy. On fqr Stlggesjed ~ea~~' ~eb l~s,_,,adqitiol).al ques¥ons,
land, structure is provided primarily by vascular aha supplementary resouices.~w -,~'{' !'7 ~- -~i ' ,,ifl't Yf~ '
plants, whereas in many marine communities, ani- *

. . . ·,1S,;~ -~-1>:' . _,Jf~ . ;·~~t- -~_;;i; '<·..... .;o

mals such as corals generate most structure. There is

a positive correlation between productivity and Questions for Discussion
species richness at low levels of productivity, but
species richness typically declines at higher produc- 1. The history of life has been punctuated by five
tivities. Some highly productive systems, such as salt episodes during which extinction rates were very
marshes, seagrass beds, and hot springs are species- high. If extinction is a normal process, and if life has
poor. Plant species richness is extremely high in rediversified after each mass extinction, why should
some unproductive semiarid regions with poor soils. we be worried about the prospects of high extinction
Island communities are poorer in species than com- rates during this century? How does the current ex-
parable mainland communities at all latitudes. tinction spasm differ from previous ones?
62 Chapter 2

2. Given that millions of species are yet to be described biologists. For what purposes might conservation
and named, how should the limited human and fi- biologists wish to use weighted indices instead of
nancial resources available for taxonomic research simple lists of species?
be allocated? Should attention be concentrated on 4. Many conservation efforts are directed at particular
poorly known taxa? Should efforts be directed to-
local areas harboring rare species or having high
ward areas threatened with habitat destruction so
species richness. Why is concentrating only on local
that species can be collected before they are elimi- problems insufficient as an effective conservation
nated? Should major efforts be directed to obtain
complete "all taxa" surveys of selected areas? How
and by whom should these decisions be made? 5. For which animal taxa would you expect species
richness to be most positively correlated with plant
3. Indices of species diversity that are weighted by
community structure? Mammals? Amphibians? In-
abundance, biomass, or productivity are used fre-
sects? Why?
quently by ecologists, but seldom by conservation