Sigma Xi, The Scientific Research Society

Conserving Biodiversity Coldspots: Recent calls to direct conservation funding to the world's
biodiversity hotspots may be bad investment advice
Author(s): Peter Kareiva and Michelle Marvier
Source: American Scientist, Vol. 91, No. 4 (JULY-AUGUST 2003), pp. 344-351
Published by: Sigma Xi, The Scientific Research Society
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Conserving Biodiversity Coldspots

Recent calls todirectconservationfunding to theworld's

biodiversityhots potsmay be bad investmentadvice

Peter Kareiva and Michelle Marvier

[the numbers
fcalforests arechilling.
covering an Every
area the year tropi-
size worth
of dation pausing toexamine
of this conservation thescientific
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co
Poland are destroyed. With them, perhaps ten sider what the consequences of this concep
thousand species arewiped out annually, most may be for the huge expanses of the planet tha
before they can be somuch as cataloged. Some it leaves out in the cold-places we might dub
liken the current calamity to the last episode biodiversity "coldspots."
ofmass extinction-65 million years ago when Does itmake scientific sense to downp
a wayward asteroid killed off the dinosaurs the world's steppes, the Serengeti, the w
along with about two-thirds of the species then Arctic and other relatively species-poor ar
in existence. in favor of biodiversity hotspots? Clea
The rapid loss of tropical forests throughout species richness should be considered wh
theworld and thewidely recognized "biodi- deciding where to invest conservation dolla
versity crisis" have spurred various non- and effort.But there are other relevant facto
governmental conservation organizations and We believe that ecological theory, conside
international agencies to develop strategies for tionof ecosystem services and sociopolitical
protecting natural habitats. But the scale of the alism all argue strongly against placing
crisis is so daunting that conservationists wide- much emphasis on biodiversity hotspots.
Ameicaby xotc ldybrd be forsee ly acceptthatthe e fon fornoher
carneed some lscloed nErhS vroeptnilpolmeome
sort of triage, apr
lamentably, little consideration has been gi
tls.Kaeia s lsafiiaed th lso the ufeedtclmt severe att toalternative frameworks forysetting priorit
whereby limited funds go to the placesdepsrc-enafeyoralzththrereifctm
where
wit te renScoo ofth ton. Nmatnctiosn-6can beenioneast aswoqaniybilgaeauenh
the greatest good done. Experts afflgofeen
have
UnieritofCaifrniana iadwith ateri Unilest offOxfodiars oso prahepaie pce ihes
explored various ways to set priorities, and al- Warming Up To the Idea
BararaandtheEnvronenalconed wth atemia solar prdso he t oe specifiesy thensfr alyn u h
most without exception, rainforests get top Before probing the weak points of hotsp
billing. The reason is simple: These tropical basedstrategies,weshouldexplainprecis
ecosystems harbor more unique species than what data and reasoning have been used to
Peter Kareiva
Clar isUnversty.Michlle
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any other habitat or place. Identifying and pro- fine this approach to conservation. Hotsp
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Nature Conservancy,
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searchinvolvsthestudytfeislnds anatra Med itnansstem sch usually being people's readiness to destroy
as terlalyeovrtepsdcdeher
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maintains plntsand offieldre- rthss
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 41

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Figure 1. Faced with limited resources, many conservationists are concentrating efforts on biodiversity "hotspots," comparatively small ar
eas that each harbor a large number of species that can be found nowhere
fort
he ost
toardosteffetiv
else. The otefcieesi
actonsWIn
authors argue, however, othat such a strategy is problemat
thUoino
deed,~41j
ic, in part because it ignores the requirements of large carnivores, like this polar bear living near the shores of Hudson Bay. These creatures re
quire vast territories,
which may not be particularlyrich in species.
selnot. AsMesadfu olausA

gauge species richness, which is likely to be ahaergdtatheanrmybeoeac

the er na ifunilNtr r
gue g
component of anyone's figuring of biological
value, and the available data tend to be more pplto rwhnab ri h rcino
complete forplants than for animals. So count-spceinteaathtreurnlyegdd
ing plants is certainly a reasonable approach.asbigtrsk
One just needs to recognize the limitations of Cery h eal fhwhtpt er
. Fcedwit the landesurfaceio the earth."n Their basicy
working with Figre this rather liite reouresmancoseratoft
one-dimensional rep-fiearsujctodagemnbtheee "hpr- cmp
Shat ach arbo a lrge umbe of pecis t scriptie ond iowhr cllec Th atlog tasuhasrtg
resentation of biodiversity.alcnetsilaprsom arglistowecies, lig:Hsos
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destoye great
perils facing species and many differentwaysdethnoinfcs-fetvnsssatp
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to evaluate the Moreover, ikly there is no aselnpot.AMyradfurclagsa
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priori reason for selecting one measure
ermnnt ionanniesiuin of dan-gudtreyasgoiannfetalNtea
his coleae ne frcomlemditrentar srates the hotspo
ger over another. The hotspot approach focus- td,"4 falseiso aclrpat n
es on the percentage of primary vegetation that 3%o l pce nfu etbaegop r
www~ameicanscintist~as
has been destroyed. To qualify as a hotspot, a cnie o2 ososcmrsn 2003ngly-Auurts34
ny14
region must have suffered a loss of at least 70 oftelnsuacofheEr."Terbicp
percent of the original vegetation. Again this srpini ocleta ogals fseisa
definition is reasonable, but ithas its limits-in posbeiasmllnaraspsilendt
particular because the fraction of natural plant gtcnevto is hr epehv l
life that is already gone ismore a statement raydsrydagetda fhbtt l
about land use in the past than a direct indica-thuhheeahos(nsmeterwo
tion of future threat.Thomas D. Sisk ofNorth-haetknutebto)dacowdgte

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Figure 2. Biodiversity hotspots, as defined by Myers et al. 2000 (red), span the globe, but
they fall mostly in the tropics. The influence ofMyers's
hotspot concept is evident in the distribution of the office locations for the three largest nongovernmental conservation organizations (green
dots)?Conservation International, The Nature Conservancy and theWorld Wildlife Fund?shown here with the many office locations in the
United States and Western Europe excluded. (Background colors from satellite imagery, courtesy of ARC Science.)

concept has grown so popular in recent years
within the larger conservation community that
itnow risks eclipsing all other approaches.
Not-So-Hot Consequences
At first glance itmay seem self-evident that
conservation investment should be funneled
into the regions or countries with themost bio
diversity. But the hotspot methodology is logi
cal only if the exclusive goal of conservation is
Figure 3. State ofMontana and the country of Ecuador cover similar areas, but the latter
is home to farmore species. In weighing how to address conservation concerns in these
two places, should all available resources be focused on richer
preserving Ecuador's
natural capital? No, the authors argue; itwould be better to allow a small fraction of
Ecuador's
species to perish if this sacrifice would permit the protection of the same
number of species inMontana, where amount to a larger fraction of the
they would
original flora and fauna. Conservation efforts inMontana would also help to protect the
greater Yellowstone ecosystem (dashed line), the only place in the lower 48 states where
large mammals such as grizzly bears, wolves, bison and elk still roam freely in thewild.
to protect the largest possible number of
species in the smallest possible area. Using
hotspots to set priorities comes intoquestion as
soon as one considers a broader range of
objec
tives, such as maintaining functioning ecosys
tems throughout the world, providing the
greatest variety of distinct plant and animal
lineages for future evolutionary break
throughs, preserving spectacular wild land
scapes that inspire the human spirit or protect
ing nature in a way that provides for the
well-being of people living alongside.
A hypothetical example reveals some of the
unfortunate side effects that arise from empha
sizing hotspots above all else. Consider two ar
eas of roughly equal size, the country of
Ecuador and the state ofMontana. Ecuador is a
renowned biodiversity hotspot, harboring
2,466 vertebrate species and 19,362 vascular
plant species. In contrast,Montana is a biodi
versity coldspot, with only 12 percent of
Ecuador's species richness. Clearly, ifone mea
sures success as protecting the
largest number
of species in the smallest possible area, it
makes sense to ignoreMontana and to concen
trate solely on Ecuador. But assume for themo
ment thatwe desire some level of conservation
effort in both places. Suppose we set a goal of
ensuring protection for 20,000 total species
from these two areas. We could attain that out
come by preserving 18,000 species in Ecuador
and 2,000 species inMontana, or, alternatively,
by safeguarding 19,000 species inEcuador and
1,000 species inMontana. If all thatmatters is
the total number of species protected, these
two strategies are equivalent.
In reality these two choices would have vast
ly different consequences on the ground. Both
would leave Ecuador with thebulk of itsbiodi

346 American Scientist, Volume 91

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versity intact (82 percent or 87 percent, if,forar leagues recently valued at about $2,000 per
gument's sake, one considers just vertebrates hectare per year. But hotspot analyses overlook
and vascular plants) and, presumably, with rea the value ofwetlands?and if s easy tounder
no en
sonably well-functioning ecosystems. But shift stand why: A typical Spartinamarsh has
from the first to the second strategy cuts the demic and no more than 20 or 30 plant
ing plants
fraction of species protected inMontana se species total. Still, tidalmarshes offer consider
verely (from 74 percent to 37 percent).You able ecosystem services, such as flood regula
wouldn't have to be a scientist to notice thedif tion,waste treatment and fisheries production,
ference between saving three-quarters versus with an estimated annual value of nearly
two-fifthsof the species in the state. But even $10,000 per hectare per year. Clearly, marshes
the best-designed scientific monitoring pro are precious resources, not just forpeople but
grams would be hard pressed to register the also for the enormous variety of wildlife that
difference between having 87 percent or 82 per depends on clean water. Yet by any sort of
come
cent of the species under protection in Ecuador. hotspot reckoning, thesemarshes would
This example iUuminates a major flaw with out at the very bottom of the pile.
approaches to conservation that are solely This difficultywould not arise if the stated
based on hotspots. Ifwe measure success sim conservation goal were to preserve the func
up total species protected, we tioning of the planet's ecosystems to themaxi
ply by tallying
risk the folly of allowing major ecosystems to mum extent possible. This objective would not
a desire to
degrade beyond repair simply because theydo necessarily be incompatible with
not provide lengthy species lists. For instance, save species: Many empirical studies have
the Yellowstone ecosystem, which includes shown that ecosystem services, such as pro
parts of Montana, Idaho and Wyoming, is duction of plant biomass, retention of nutri
species poor, relatively speaking. Yet this region ents, resistance to drought, pollination of crops
harbors the last assemblage of largemammals and decomposition of organic matter, decline
and carnivores in the lower 48 states (grizzly with major losses inbiodiversity. But one strik
bears, wolves, bison, elk and so forth).And be ing feature of the relation between ecosystem
cause the Yellowstone ecosystem contains the services and biodiversity is that it is not linear.
world's first national park, it is obviously an Rather, the benefits of biodiversity are quickly
important locale for continued conservation, realized with an initial accumulation of species
despite the relative paucity of species that exist and thereafter remain constant, so thatprotect
there. So by itself, the number of species saved ingmore species does not forever translate into
is an inadequate barometer of success. Other more or better ecosystem services.
dimensions of the problem need to be consid
ered as well, including what environmentalists
often describe as "ecosystem services."

Earth's Life-Support System

Society depends on the products and services
ecosystems provide to a fargreater extent than
most of us realize. When asked what nature
gives them, most people could name a few
foods, drugs and building materials derived
fromwild species. But theywould probably
take for granted many other products of na
ture, including clean air, abundant freshwater,
fertile soil and a benign climate. In short, they
would fail to recognize the critical role that di
verse communities of species play in fostering
a healthy and predictable environment.
Would not hotspot-based conservation
strategies automatically take such effects into
account? Hardly. Investing conservation efforts
only in hotspots could lead us to ignore and
potentially lose some of our most valuable
ecosystems simply because they harbor few
plant species. Consider, for example, the fact
that hotspot analyses so often point to tropical
forestsas areas of highest priority.These forests
Figure 4. Wetlands, such as this one near Mobile, Al
do indeed provide important ecosystem ser services. Yet these
abama, provide valuable ecosystem
vices, such as climate regulation and nutrient places may host only a small number of plant species. A
an economist
cycling, which Robert Costanza, typical Spartina marsh, for example, harbors fewer than
at the University of Maryland, and his col 30. (Courtesy of the U.S. Army Corps of Engineers.)

www.americanscientist.org
2003 July-August 347

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 It is not clear how to predict where this satu
ration effectwill manifest itselfwhen one con
siders a variety of possible benefits. But ifwe ac
cept that benefit curves do level off,we must
accept that thereare dirninishing returns forpro
tecting ever more species in any particular
ecosystem. Given this pattern, a logical goal
might be to ensure thatno major ecosystem suf
fersgreater than a 10 percent loss, or a 20 percent
loss, or a 50 percent loss of diversity.
Although scientists lack the knowledge to
assign values to the services provided by the
ecosystems in all countries or regions, what
matters is obviously not how many species
there are as much as what percentage of the
native diversity remains sheltered from de
struction. The hotspot approach would result
in high levels of protection for a few species
rich areas to the neglect ofmany others. Thus,
setting conservation priorities using only
an un
hotspots as a guide could well bring on
fortunate side effect:more degradation of glob
al ecosystems thanwould take place ifa more
broadly based strategywere used.
Thicker Branches on the Tree of Life
Most conservationists emphasize ecological or
ecosystem value when discussing the need
preserve biodiversity. But they often neglect an
aspect of biodiversity that is just as important: points to a greater need for conservation in
itsworth as a resource for future evolutionary
innovation. In this regard, one has to consider
more than just species. For example, a greater
amount of evolutionary history and biological
distinctiveness is lostwhen the last species of
an entire genus or family becomes extinct than wildlife should receive urgent attention. After
when a species with many close relatives dis
io 15
plantspecies richnessbeforedrought
Figure 5. Ecosystem functioning generally improves as the number of species rises, but
the relation is not linear. Typically, the benefits of increasing the species count
and then level out. Here, one sees that the resistance of an ecosystem to drought,
quickly
measured as the ratio of biomass after the drought to the biomass before, depends
number?but to a from Tilman and 1994.)
species only point. (Adapted Downing
348 American Scientist, Volume 91
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appears. Some evolutionary biologists have
challenged the current focus on biodiversity
hotspots as myopic, because it neglects the
unique value of distinct evolutionary lineages,
which represent very different life histories
and forms.Rather than simply directing efforts
toward areas with rich sets of endemic species,
conservation organizations might better con
centrate on saving higher taxonomic groups
under threat.
One could, for example, try to rescue genera
that are in danger of being entirely lost. To test
this idea, we looked at the distribution of
mammal and bird genera with more than one
species forwhich theWorld Conservation
Union (IUCN) has listed all their constituent
species as critically endangered, endangered,
vulnerable or dependent on conservation. For
each of themembers of these highly threatened
genera, we then searched IUCN's latest
"redlist" database of endangered species and
recorded all countries inwhich these rare crea
tures still occur naturally or have been reintro
duced. We then ranked the countries according
to thenumber of highly threatened genera that
can be found there.
This procedure provides a list that differs
on therichness of
to greatly fromone based solely
endemic plants. In particular, this exercise
cer
tainAfrican nations than does a hotspot-based
approach. For example, Kenya, with only 265
endemic plant species, has never been identi
fied as a biodiversity hotspot. Yet our ranking
suggests that the conservation of Kenyan
all, this nation is home to species belonging to
each of the following eight highly threatened
mammalian genera: Alcelaphus (hartebeest),
Connochaetes (gnu),Hippotragus (a type of ante
lope), Oryx (another type of antelope), Otomops
(a type of bat), Redunca (reedbuck), Rhynchocyon
(elephant shrew) and Surdisorex (mole shrew).
Another shortcoming of thehotspot method
one looks care
ology becomes apparent when
fully at how it is often applied. Although Myers
and his colleagues initially proposed hotspots
as a means of setting conservation priorities at a
very large scale and in a coarse manner, the no
tion of getting themost species per unit area of
land protected ("efficiency") has been translat
ed tomuch smaller spatial scales with poten
tially unfortunate consequences. For example,
in thepast few years two influential analyses of
biodiversity within theU.S. have been used to
show how conservationists might efficiently
protect species by focusing on just a few small
clusters of critical counties. This strategy
would, however, fail to protect adequately
accrue those species that require large tracts of rela
tively undisturbed habitat.
on An added worry surfaces when you take a
on the
long-term, evolutionary perspective
Figure 6. Conservation priorities can be set using any number of different strategies. Here the authors examined the distribution of bird and
mammal genera that are in danger of being entirely wiped out and scored countries according to how many of these genera are still found there.
The resulting map shows some similarities with the hotspots ofMyers (dark shading) but also many differences.

problem. By focusing on conserving themost variables can be cast into the ideal metric for
species in the smallest possible area (for the sake scoring urgency. This leaves scientifically
of cost-effectiveness), conservationists may in minded conservationists with the unanswered
advertently be altering the course of evolution. question: What then should we do?
How? Protecting hotspots at small scales favors One thing is tomake sure that the problem is
species that can live in relatively restricted areas. framed properly. In recent times,biologists have
Thus we might expect rodents to enjoy high achieved a major conceptual advance by realiz
speciation rates relative to extinction rates. At ing that conservation programs must span po
the other extreme, species that require vast ter litical boundaries and that for planning
ritories, and thus cannot be contained in cost-ef purposes biologically defined areas such as
fectivehotspots, will sufferdisproportionate ex "eco-regions" represent natural units. Although
tinction relative to speciation. Hence primates thismindset is useful forquantifying biological
and large carnivores would be expected towane value, it is not terriblyhelpful formaking deci
relative to their smaller mammalian counter sions about the feasibilityor cost-effectivenessof
parts. Recent analyses of vertebrate extinction a particular conservation project. When an or
and speciation rates suggest that this is exactly ganization decides towork in a region, itwill
what is happening. Thus, as Donald A. Levin necessarily have to deal with the regulatory
(of theUniversity of Texas atAustin) and Phillip agencies, legal institutions and people of indi
S. Levin (of theNational Marine Fisheries Ser vidual nations. Thus, although our conservation
vice) argued in these pages not too long ago goals may be biological, as long as feasibility
(Macroscope, January-February 2002), the Earth and cost-effectiveness are important, countries
may well end up with a paucity of primates and will remain essential units for consideration.
rhinoceroses, and a surplus of rodents. One might thus be tempted to try to identify
One hopes that eventuality will not actually a comprehensive list of countries having great
come to pass. Indeed, one hopes that society biological value and where an immediate
will ultimately prove able to protect habitats threat looms?but also where action is cost-ef
and ecosystems throughout theworld so that fective and feasible. Unfortunately, conserva
evolution will be allowed to take a natural tionists typically lack the information needed
course. But in the short term it's clear that to assess all of the relevant variables. For in
some tough decisions need to be made in allo stance, ecosystem services have not been wide
cating
scarce resources. Thus far, most conser
ly calculated, current rates of habitat loss are
vationists have focused on biological value as only erratically available, and the uniqueness
themajor criterion for setting priorities. But or irreplaceability of particular environments
others have explored different sides of the are complicated to assess. One solution is to
problem, including the degree of immediate investigate many different approaches to see
threat to biodiversity, general feasibility and whether certain areas are consistently identi
cost-effectiveness. Although there are many fied as being of high priority. In other words,
ways to go about doing this, it's hard to argue are some conclusions robust, no matter which
from scientific principles that any simple set of criteria are used?

www.americanscientist.org 2003 July-August 349

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Clockwise fromleft:
Nigel J.Dennis, TomMcHueh,
TimDavis, MerlinD. T?Hie/PhotoResearchers,Inc.

Figure 7. Although it does not appear on any hotspots list, Kenya deserves urgent attention because it harbors members of eight highly threat
ened mammalian genera, including the four shown here: Hippotragus (a type of antelope, left), Rhynchocyon (elephant shrew, middle top), Oto
mops (a type of bat, middle bottom) and Oryx (another type of antelope, right).

In this spirit,we have invested quite a bit of
time recently exploring all sorts ofmethods for
setting conservation priorities. And what we've
noticed is that some countries seem always to
come out on top. A few?such as India, Viet
nam, South Africa, thePhilippines and Brazil?
overlap with the top 25 hotspots thatMyers
and his colleagues identified. Interestingly,
these five countries have relatively small per
centages of their land under conservation pro
tection,which suggests that theymay be rela
tively sure bets, given that so much room for
progress exists there.Yet other nations appear
ing on a great many of our priority lists have
never been identified as biodiversity hotspots.

Dispensing with Lists

Biodiversity hotspots represent an initial and
pioneering effort at establishing conservation
priorities. But by relying toomuch on counts of
plant species, this approach loses sight of
whole ecosystems, habitats and the needs of
people. To work themetaphor a little harder,
you might say thatbiodiversity hotspots leave
toomany places and people out in the cold.
Because conservation threats are now
many
global in their origin and scope (for example,
climate change and invasive species), place
based priorities risk disenfranchising too
number of endangered
many people from the challenge at hand. In
plantspecies
119 to77 deed, on reflection,we worry that the initially
8 to 18 appealing notion of getting themost species or
5 to7
greatest biological value per unit area is, in
133 or 4
2 fact, a thoroughly misleading strategy. How
1 much of a victory would it actually be ifpeo
l__ 10or no data ple did manage to conserve only the 1.4 per
cent of the Earth's land surface that contains
8. Using the threat to local species to identify certain areas as biodiversity
Figure almost half theworld's vascular plants? The
hotspots, a strategy
originally intended to establish conservation priorities on a broad,
scale, has also been to evaluate small areas such as U.S. coun reality is that people must make conservation
regional employed
ties?with great potential formisapplication, according to the authors. Although one progress everywhere. Doing that requires not
could concentrate conservation efforts using maps such as this one, the
a ranking of theoretically deserving places but
economically
counties" are commonly
identified too small and too disjoint to provide ad a prioritization that takes into account the ef
"hotspot
equate range for highly valued carnivores such as panthers, mountain lions, grizzly fectiveness of past conservation efforts.A per
bears and wolves. (Adapted from Dobson et al. 1997). formance-based system would not only hold

350 American Scientist, Volume 91

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conservation
organizations
more accountable, mantra runs the risk of leaving theworld with
itwould also provide incentives to countries a sizable collection of species in a few areas but
attempting
to
implement
conservation mea with an environment that is otherwise largely
sures, so that those nations demonstrating
suc
degraded. Rather than trying to identifydense
cesses on the ground would be more likely to concentrations of species
on a
map,
we and
receive funding in the future.Anyone who has other conservationists should be more flexible
worked fora while in conservation knows that and should be prepared to reward effective ac
certain people and certain leaders can over tions on the ground as they happen. Ifwe do
come enormous obstacles and do wonders in so,we will surely discover plenty of coldspots
themost unlikely places. Yet none of the es deserving of our attention.
tablished priority-setting schemes recognizes
such human factors. Bibliography
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Unwavering support for the protection of For relevant Web links, consult this issue of American
hotspots oversimplifies the difficult decisions Scientist Online:
thatmust be made in deciding which projects
to fund and where to investmoney. Although http: / /www.americanscientist.org/

biodiversity hotspots are indeed an academi template / IssueTOC / issue /394

cally appealing idea, blind adherence to this

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