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Causes and consequences of species extinctions

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DOI: 10.1515/9781400833023.514

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V.1
Causes and Consequences
of Species Extinctions
Navjot S. Sodhi, Barry W. Brook,
and Corey J. A. Bradshaw

OUTLINE megafauna. This refers to large-bodied (>44 kg) ani­

mals, commonly (but not exclusively) used to refer
1. Introduction
to the large mammal biota of the Pleistocene.
2. Extinction drivers
minimum viable population. This is the number of in­
3. Extinction vulnerability
dividuals in a population required to have a speci­
4. Consequences of extinctions
fied probability of persistence over a given period of
5. Conclusions
time.
The five largest mass die-offs in which 50–95% of species
were eliminated occurred during the Ordovician [490–443
1. INTRODUCTION
million years ago (mya)], Devonian (417–354 mya), Permian
(299–250 mya), Triassic (251–200 mya), and Cretaceous In the Americas, charismatic large-bodied animals
(146–64 mya) periods. Most recently, human actions espe­ (megafauna) such as saber-toothed cats (Smilodon
cially over the past two centuries have precipitated a global spp.), mammoths (Mammuthus spp.), and giant ground
extinction crisis or the ‘‘sixth great extinction wave’’ com­ sloths (Megalonyx jeffersonii) vanished following hu­
parable to the previous five. Increasing human populations man arrival some 11,000–13,000 years ago. Similar
over the last 50,000 years or so have left measurable losses occurred in Australia 45,000 years ago, and in
negative footprints on biodiversity. many oceanic islands within a few hundred years of the
arrival of humans. Classic examples of the loss of is­
land endemics include the dodo (Raphus cucullatus)
GLOSSARY
from Mauritius, moas (e.g., Dinornis maximus) from
Allee effects. These factors cause a reduction in the New Zealand, and elephantbirds (Aepyornis maximus)
growth rate of small populations as they decline from Madagascar. Megafaunal collapse during the late
(e.g., via reduced survival or reproductive success). Pleistocene can largely be traced to a variety of negative
coextinction. Extinction of one species triggers the loss human impacts, such as overharvesting, biological in­
of another species. vasions, and habitat transformation.
extinction debt. This refers to the extinction of species The rate and extent of human-mediated extinctions
or populations long after habitat alteration. are debated, but there is general agreement that ex­
extinction vortex. As populations decline, an insidious tinction rates have soared over the past few hundred
mutual reinforcement occurs among biotic and years, largely as a result of accelerated habitat de­
abiotic processes driving population size downward struction following European colonialism and the sub­
to extinction. sequent global expansion of the human population
extirpation. This refers to extinction of a population during the twentieth century. Humans are implicated
rather than of an entire species. directly or indirectly in the 100- to 10,000-fold in­
invasive species. These are nonindigenous species in­ crease in the ‘‘natural’’ or ‘‘background’’ extinction
troduced to areas outside of their natural range that rate that normally occurs as a consequence of gradual
have become established and have spread. environmental change, newly established competitive
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Species Extinctions 515

interactions (by evolution or invasion), and occasional mortality rates to exceed reproductive replacement
chance calamities such as fire, storms, or disease. The over a sustained period can cause a species to become
current and future extinction rates are estimated using extinct. Such forces may act independently or syner­
a variety of measures such as species–area models and gistically, and it may be difficult to identify a single
changes in the World Conservation Union’s (IUCN) cause of a particular species extinction event. For in­
threat categories over time. Based on the global as­ stance, habitat loss may cause some extinctions directly
sessment of all known species, some 31, 12, and 20% by removing all individuals, but it can also be indirectly
of known amphibian, bird, and mammal species, re­ responsible for an extinction by facilitating the estab­
spectively (by far the best-studied of all animal groups), lishment of an invasive species or disease agent, im­
are currently listed by the IUCN as under threat. proving access to human hunters, or altering biophys­
Just how many species are being lost each year is ical conditions. As a result, any process that causes a
also hotly debated. Various estimates range from a few population to dwindle may ultimately predispose that
thousand to more than 100,000 species being ex­ population to extinction.
tinguished every year, most without ever having been Evidence to date suggests that deforestation is cur­
scientifically described. The large uncertainty comes rently, and is projected to continue to be, the prime
mainly through the application of various species–area direct and indirect cause of reported extirpations. For
relationships that vary substantially among communi­ example, it is predicted that up to 21% of Southeast
ties and habitats. Despite substantial prediction error, Asian forest species will be lost by 2100 because of past
it is nevertheless certain that human actions are causing and ongoing deforestation. Similar projections exist for
the structure and function of natural systems to un­ biotas in other regions.
ravel. The past five great extinctions shared some im­ Overexploitation is also an important driver of ex­
portant commonalities: (1) they caused a catastrophic tinctions among vertebrates and tends to operate syn­
loss of global biodiversity; (2) they unfolded rapidly (at ergistically with other drivers such as habitat loss. For
least in the context of evolutionary and geological example, roads and trails created to allow logging op­
time); (3) taxonomically, their impact was not random erations to penetrate into virgin forests make previ­
(that is, whole groups of related species were lost while ously remote areas more accessible to human hunters,
other related groups remained largely unaffected); and who can, in turn, cause the decline and eventual ex­
(4) the survivors were often not previously dominant tirpation of forest species. It is estimated that overex­
evolutionary groups. All four of these features are rel­ ploitation is a major threat to at least one-third of
evant to the current biodiversity crisis. This sixth great threatened birds and amphibians, with wildlife cur­
extinction is likely to be most catastrophic in tropical rently extracted from tropical forests at approximately
regions given the high species diversity there (more six times the sustainable rate. In other words, the
than two-thirds of all species) and the large, expanding quantity, and most likely the diversity, of human prey—
human populations that threaten most species there as both fisheries and ‘‘bush’’ (wild) meat—are rapidly
well. diminishing.
The major ‘‘systematic drivers’’ of modern species Megafauna—those species weighing in the tens to
loss are changes in land use (habitat loss degradation hundreds of kilograms—are among the most vulnera­
and fragmentation), overexploitation, invasive species, ble to overexploitation. In general, a species’ genera­
disease, climate change (global warming) connected tion time (interval from birth to reproductive age) is a
to increasing concentration of atmospheric carbon di­ function of body mass (allometry), so larger, longer-
oxide, and increases in nitrogen deposition. Mechan­ lived, and slower-reproducing animal populations are
isms for prehistoric (caused by humans >200 years generally unable to compensate for high rates of har­
ago) extinctions are likely to have been similar: over- vesting. Because slow-breeding large animals, such as
hunting, introduced predators and diseases, and habi­ apes, carnivores (e.g., the lion, Panthera leo), and Af­
tat destruction when early people first arrived in virgin rican elephants (Loxodonta africana), are particularly
landscapes. vulnerable to hunting, the potential for population
recovery in these animals over short time scales is low.
As an example supporting this generality, there is evi­
2. EXTINCTION DRIVERS
dence that 12 large vertebrate species have been ex­
Some events can instantly eliminate all individuals of tirpated from Vietnam, primarily because of excessive
a particular species, such as an asteroid strike, a mas­ hunting, within the past 40 years. The Steller’s sea cow
sive volcanic eruption, or even a rapid loss of large (Hydrodamalis gigas), an aquatic herbivorous mam­
areas of unique and critical habitat because of defor­ mal that inhabited the Asian coast of the Bering Sea,
estation. But ultimately, any phenomena that can cause is the quintessential example of the rapid demise of a
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516 Conservation Biology
species as a result of overexploitation. Discovered in populations through chemical treatments and the elim­
1741, it became extinct by 1768 because of overhunt­ ination of larval habitats.
ing by sailors, seal hunters, and fur traders. This species Perhaps one of the most infamous examples of an
was hunted for food, its skin for making boats, and its invasion catastrophe occurred in the world’s largest
subcutaneous fat for use in oil lamps. freshwater lake—Lake Victoria in tropical East Africa.
The ecosystem and biological community changes Celebrated for its amazing collection of over 600
precipitated by invasive species represent another endemic haplochromine (i.e., formerly of the genus
leading cause of biodiversity loss. Of 170 extinct spe­ Haplochromis) cichlid fishes (Family Cichlidae), the
cies for which causes have been identified reliably, Lake Victoria cichlid community is perhaps one of the
invasive species contributed directly to the demise of most rapid, extensive, and recent vertebrate radiations
91 (54%). In particular, the rates of extinctions oc­ known. There is also a rich community of endemic
curring on islands have been greatly elevated by the noncichlid fish that inhabit the Lake. In addition to the
introduction of novel predators. Several ecological and threats posed to this unique biota by a rapid rise in
life-history attributes of island species, such as their fisheries exploitation, human density, deforestation,
naturally constrained geographic range, small popula­ and agriculture during the past century, without doubt
tion sizes, and particular traits (e.g., lack of flight in the most devastating effect was the introduction of the
birds or lack of thorns in plants) make island biotas predatory Nile perch (Lates niloticus) in the 1950s.
vulnerable to predation from invading species. For This voracious predator, which can grow to more than
example, the introduction of the brown tree snake 2 m in length, was introduced from lakes Albert and
(Boiga irregularis) shortly after World War II wreaked Turkana (Uganda and Kenya, respectively) to com­
havoc on the biodiversity of the island of Guam in the pensate for depleting commercial fisheries in Lake
South Pacific. In all likelihood, tree snakes were di­ Victoria. Although the Nile perch population remained
rectly responsible for the loss of 12 of 18 native bird relatively low for several decades after its introduc­
species, and they also reduced the populations of other tion, an eventual population explosion in the 1980s
vertebrates such as flying foxes (Pteropus mariannus), caused the devastating direct or indirect extinction of
mainly because of the inability of the island’s native 200– 400 cichlid species endemic to the Lake as well as
species to recognize the novel predator as a threat. the extinction of several noncichlid fish species. Al­
Despite an annual expenditure of US$44.6 million for though many other threats likely contributed to the
the management of this problem, tree snakes on Guam observed extinctions, including direct overexploitation
are still not under control, largely because of their and eutrophication from agriculture and deforestation
ability to penetrate artificial snake barriers such as leading to a change in the algal plankton community,
fences. there are few other contemporary examples of such a
The mosquito Culex quinquefasciatus was inad­ rapid and massive extinction event involving a single
vertently introduced to Hawaii in 1826, and the group of closely related species.
disease-causing parasite (Plasmodium relictum) it car­ Human-mediated climate change represents a po­
ries arrived soon after. Since then, avian malaria (in tentially disastrous sleeping giant in terms of future
conjunction with other threats) has been responsible biodiversity losses. Climate warming can affect species
for the decline and extinction of some 60 species of in five principal ways: (1) alterations of species densi­
endemic forest birds on the Hawaiian Islands. Having ties (including altered community composition and
evolved in the absence of the disease, Hawaiian bird structure); (2) range shifts, either poleward or upward
species were generally unable to cope with the debili­ in elevation; (3) behavioral changes, such as the phe­
tating effects of the novel parasite. However, more nology (seasonal timing of life cycle events) of migra­
than 100 years after the establishment of the disease, tion, breeding, and flowering; (4) changes in mor­
some native thrushes (Myadestes spp.) are now show­ phology, such as body size; and (5) reduction in genetic
ing resistance to the disease. Sadly, many of the re­ diversity that leads to inbreeding depression. A related
maining species, especially forest birds in the family threat for island and coastal biotas is the predicted loss
Drepanididae, are still vulnerable and are now re­ of habitat via inundation by rising sea levels. Although
stricted to altitudes where temperatures are below the large fluctuations in climate have occurred regularly
thermal tolerance limits of the mosquito vector. Global throughout Earth’s history, the implications of an­
warming is predicted to increase the altitudinal distri­ thropogenic global warming for contemporary biodi­
bution of the mosquito, thus spelling doom for disease- versity are particularly pessimistic because of the rate
susceptible birds as mosquito-free habitats disappear. of change and previous heavy modification of land­
The most feasible method of reducing transmission of scapes by humans. Good empirical evidence for some
malaria is to reduce or eliminate vector mosquito of these effects is rare, and speculations abound, but
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Species Extinctions 517

there are already many local or regional examples habitat loss. For example, even if net deforestation
and model-based predictions that support the view rates can be reduced or even halted, the extinction debt
that rapid climate change, acting in concert with other of remnant and secondary forest patches will see the
drivers of species loss and habitat degradation, will be extinction of countless remaining species over this
one of the most pressing conservation issues global interval.
biodiversity faces over the coming centuries.
One glimpse of a possible future crisis comes
3. EXTINCTION VULNERABILITY
from the highland forests of Monterverde (Costa Rica),
where 40% (20 of 50) of frog and toad species dis­ Certain life-history, behavioral, morphological, and
appeared following synchronous population crashes in physiological characteristics appear to make some spe­
1987, with most crashes linked to a rapid progressive cies more susceptible than others to the extinction
warming and drying of the local climate. The locally drivers described above. In general, large-sized species
endemic golden toad (Bufo periglenes) was one of the with a restricted distribution that demonstrate habitat
high-profile casualties in this area. It has been sug­ specialization tend to be at greater risk of extinction
gested that climate warming resulted in a retreat of the from human agency than others within their respective
clouds and a drying of the mountain habitats, making taxa (e.g., Javan rhinoceros, Rhinoceros sondaicus),
amphibians more susceptible to fungal and parasite especially to processes such as rapid habitat loss.
outbreaks. Indeed, the pathogenic chytrid fungus Ba­ Because of their high habitat specificity and/or low
trachochytrium dendrobatidis, which grows on am­ population densities, rare species may be more prone to
phibian skin and increases mortality rates, has been extinction than common species. The size of a species’
implicated in the loss of harlequin frogs (Atelopus spp.) range is also a major determinant of its extinction
in Central and South America and reductions in proneness. Small ranges may make species more vul­
other amphibian populations elsewhere. It is hypoth­ nerable to stochastic perturbations, even if local abun­
esized that warm and dry conditions may stress am­ dance is high; for example, proportionally more
phibians and make them more vulnerable to the fungal passerines (perching birds) with relatively small geo­
infection. graphic ranges in the Americas are at risk of extinction
Irrespective of the reason for a population’s decline than their more widely distributed counterparts. Such
from a large to small population size, unusual (and trends are worrisome because those species with
often random and detrimental) events assume promi­ shrinking ranges as a result of adverse human activities
nence at low abundances. For instance, although become particularly vulnerable to other drivers such as
competition among individuals is reduced at low den­ climate change. Habitat loss also reduces the patch
sities and can induce a population rebound, a coun­ sizes necessary for species requiring large home ranges,
tervailing phenomenon known as the ‘‘Allee effect’’ can making them vulnerable to extinction from a loss of
act to draw populations toward extinction by (for in­ subpopulation connectedness, reduced dispersal ca­
stance) disrupting behavioral patterns that depend on pacity, and the ensuing lower population viability.
numbers (e.g., herd defense against predators) or by Larger-bodied vertebrates are considered to be more
genetic threats such as inbreeding depression. Small extinction-prone than smaller-bodied ones when the
populations, dominated by chance events and Allee threatening process unfolds rapidly or intensely. In­
effects, are often considered to have dipped below their deed, threatened mammals are an order of magnitude
‘‘minimum viable population’’ size. Thus, once a major heavier than nonthreatened ones. A common expla­
population decline has occurred (from habitat loss, nation for this trend is that body size is inversely cor­
overexploitation, or in response to many other possible related with population size, making large-bodied an­
stressors), an ‘‘extinction vortex’’ of positive feed­ imals less abundant and more vulnerable to chronic
back loops can doom species to extinction, even if the environmental perturbations (while being buffered
original threats have been alleviated. Further, many against short-term environmental fluctuations). The
species may take decades to perish following habitat extinction proneness of large-bodied animals to human
degradation. Although some species may withstand activities is further enhanced because of other corre­
the initial shock of land clearing, factors such as the lated traits, such as their requirement of large area,
lack of food resources, breeding sites, and dispersers greater food intake, high habitat specificity, and lower
may make populations unviable, and they eventually reproductive rate.
succumb to extinction. This phenomenon evokes the Large species can also be more vulnerable to human
concept of ‘‘living-dead’’ species, or those ‘‘committed persecution such as hunting, whereas smaller species
to extinction.’’ The eventual loss of such species is are generally more vulnerable to habitat loss. It is im­
referred to as the ‘‘extinction debt’’ caused by past portant, however, to be cautious when constructing
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518 Conservation Biology
generalized rules regarding the role of body size in the penetrating smaller forest fragments. Absence of some
extinction process. Because they have a slower repro­ insectivorous bird species from small fragments may
ductive rate, larger parrots are more vulnerable to not be related to food scarcity; rather, it may result
overexploitation than smaller finches, despite fewer from their poorer dispersal abilities. The ability to
numbers of the former being captured for the pet trade. disperse in birds and insects depends on morphological
However, some smaller species (e.g., white-eyes, Zos­ characteristics such as wing loading, and physiological
terops spp.) with small population sizes are also vul­ restrictions such as intolerance to sunlight when mov­
nerable to extinction because of heavy harvest rates for ing within the nonforested matrix landscape separat­
the pet trade, suggesting that only when the threatening ing fragments. As a result, poor dispersal ability may
processes are approximately equivalent will the larger make certain species vulnerable to extinction because
of two species being compared demonstrate a higher they cannot readily supplement sink habitats (habi­
risk of extinction. In addition to body size, other tats in which populations cannot replace themselves),
morphological characteristics affect extinction prone­ supporting otherwise unviable subpopulations, or
ness. For instance, large investment in secondary sexual colonize new areas. Because of poor dispersal abil­
characteristics may render highly dimorphic species ity, patchy distributions, and generally low popula­
less adaptable in a changing environment or more at­ tion densities, the genetic diversity of species in
tractive to specimen or pet-trade collectors. fragmented landscapes may be difficult to maintain,
When an environment is altered abruptly or sys­ with the resulting inbreeding depression further re­
tematically at a rate above normal background change, ducing population size toward extinction. However,
or beyond the capacity of adaptation via natural se­ clear and quantitative demonstrations of the role of
lection, specialist species with narrow ecological niches life-history traits in the extinction process of biotas are
often bear the brunt of progressively unfavorable still rare.
conditions such habitat loss and degradation. For in­
stance, highly specialized forest-dependent taxa are
4. CONSEQUENCES OF EXTINCTIONS
acutely vulnerable to extinction following deforesta­
tion and forest fragmentation. Possible mechanisms The extinction of certain species such as large preda­
include reductions in breeding and feeding sites, in­ tors and pollinators may have more devastating eco­
creased predation, elevated soil erosion and nutrient logical consequences than the extinction of others.
loss, dispersal limitation, enhanced edge effects, and Ironically, avian vulnerability to predation is often
other stressors. Conversely, non-forest-dependent spe­ exacerbated when certain large predatory species be­
cies or those that prefer open habitats are often better come rarer in tropical communities. For example, al­
able to persist in disturbed landscapes and may even be though large cats such as jaguars (Panthera onca) do
favored by having fewer competitors or expanded not prey on small birds directly, they exert a limiting
ranges following deforestation. It is important to be force on smaller predators such as medium-sized and
aware that in relatively stable systems, evolution en­ small mammals (mesopredators), which become more
genders the speciation of taxa that occupy all available abundant with the former species’ decline. The cor­
niches so both specialist and generalist species can co­ ollary is that abundant mesopredators inflict an above-
exist. As a result, the rapid pace of habitat and climate average predation rate on the eggs and nestlings of
change renders specialization a modern ‘‘curse’’ in small birds. Although this ‘‘mesopredator-release’’
evolutionary terms. hypothesis has been applied largely to mammals (e.g.,
Foraging specialization is one mechanism that can Australian dingoes, Canis lupus, suppressing foxes and
compromise a species’ ability to persist in altered cats; coyotes in California controlling cat abundance),
habitats. Many studies have shown that frugivorous the loss of large predatory birds such as the harpy eagle
and insectivorous birds are more extinction-prone than (Harpia harpyja) may have similar ecosystem effects.
other avian feeding guilds, with the lack of year-round Similar mesopredator release has been demonstrated
access to fruiting plants in fragmented forests being the for the first time in the marine environment, where the
culprit for the former. A number of hypotheses have overexploitation of large pelagic sharks resulted in an
been proposed to explain the disappearance of insec­ increase in rays and skates that eventually suppressed
tivorous birds from deforested or fragmented areas. commercially important scallop populations. Likewise,
First, deforestation may impoverish the insect fauna does the disappearance of a competitor result in the
and reduce selected insectivore microhabitats (e.g., niche expansion and higher densities of subordinate
dead leaves). Second, insectivores may be poor dis­ species? This phenomenon has been observed between
persers and have near-ground nesting habits, the latter unrelated taxa—the extinction of insectivorous birds
trait making them more vulnerable to nest predators from scrub forests of West Indian islands correlated
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Species Extinctions 519

with the subsequent higher biomass of competing forest bees were worth US$60,000 to a 1100-ha farm.
Anolis lizards. A forest patch as small as 20 ha located near farms
Conservation biologists have traditionally focused can increase coffee yield and thus bring large eco­
on the study of the independent declines, extirpations, nomic benefits to the farmers. Such findings illustrate
or extinctions of individual species while paying rela­ the imperative of preserving native forests near agro­
tively less attention to the possible cascading effects of forestry systems to facilitate the travel by forest-
species coextinctions (e.g., hosts and their parasites). dependent pollinating insects.
However, it is likely that many coextinctions between
interdependent taxa have occurred, but most have gone
unnoticed in these relatively understudied systems. For 5. CONCLUSIONS
example, an extinct feather louse (Columbicola ex­ Although extinctions are a normal part of evolution,
tinctus) was discovered in 1937, 23 years after likely human modifications to the planet in the last few
coextinction with its host passenger pigeon (Ectopistes centuries, and perhaps even millennia, have greatly
migratorius). Ecological processes disrupted by ex­ accelerated the rate at which extinctions occur. Habitat
tinction or species decline may also lead to cascading loss remains the main driver of extinctions, but it may
and catastrophic coextinctions. Frugivorous animals act synergistically with other drivers such as over­
and fruiting plants on which they depend have a key harvesting and pollution, and, in the future, climate
interaction linking plant reproduction and dispersal change. Large-bodied species, rare species, and habitat
with animal nutrition. Thus, the two interdependent specialists are particularly prone to extinction as a re­
taxa are placed in jeopardy by habitat degradation. sult of rapid human modifications of the planet. Ex­
Many trees produce large, lipid-rich fruits adapted for tinctions can disrupt vital ecological processes such as
animal dispersal, so the demise of avian frugivores may pollination and seed dispersal, leading to cascading
have serious consequences for forest regeneration, even losses, ecosystem collapse, and a higher extinction rate
if the initial drivers of habitat loss and degradation are overall.
annulled.
Essential ecosystem functions provided by forest
invertebrates are also highly susceptible when species FURTHER READING
are lost after habitat loss and degradation. Acting as
Brook, Barry W., Navjot S. Sodhi, and Peter K. L. Ng. 2003.
keystone species in Southeast Asian rainforests, figs Catastrophic extinctions follow deforestation in Singa­
rely on tiny (1–2 mm) species-specific wasps for their pore. Nature 424: 420–423. This is one of few papers
pollination. Some fig wasps may have limited dispersal reporting broad-scale extinctions driven by tropical de­
ability, suggesting that forest disturbance can reduce forestation.
wasp densities and, by proxy, the figs that they polli­ Clavero, Miguel, and Emili Garcia-Berthou. 2005. Invasive
nate. Similarly, dung beetles are essential components species are a leading cause of animal extinctions. Trends in
of ecosystem function because they contribute heavily Ecology and Evolution 20: 110. The article highlights that
to nutrient-recycling processes, seed dispersal, and the invasive species represent one of the primary threats to
reduction of disease risk associated with dung accu­ biodiversity.
Dirzo, Rudolfo, and Peter J. Raven. 2003. Global state of
mulation. In Venezuela, heavier dung beetles were
biodiversity and loss. Annual Review of Environment and
more extinction-prone than lighter species on artifi­ Resources 28: 137–167. The article constitutes a major
cially created forested islands, which predicts particu­ review of the state of the modern global biodiversity and
larly dire ecosystem functional loss given the former its associated losses.
group’s greater capacity to dispose of dung. Fagan, William F., and E. E. Holmes. 2006. Quantifying the
Almost all flowering plants in tropical rainforests extinction vortex. Ecology Letters 9: 51–60. This is the
are pollinated by animals, and an estimated one-third only study yet to quantify the final phases of extinction in
of the human diet in tropical countries is derived from vertebrates for which date of extinction was known.
insect-pollinated plants. Therefore, a decline of forest- IUCN Red List of threatened species. Download from http:
dwelling pollinators impedes plant reproduction not //www.iucnredlist.org. This presents an up-to-date clas­
sification of and reasons for a listed species’ conservation
only in forests but also in neighboring agricultural
status.
areas visited by these species. Lowland coffee (Coffea Koh, Lian P., Robert R. Dunn, Navjot S. Sodhi, Robert
canephora) is an important tropical cash crop, and it K. Colwell, Heather C. Procter, and Vince S. Smith.
depends on bees for cross-pollination. A study in Costa 2004. Species co-extinctions and the biodiversity crisis.
Rica found that forest bees increased coffee yield by Science 305: 1632–1634. This models how loss of a spe­
20% in fields within 1 km of the forest edge. Between cies could indirectly result in the extinction of dependent
2000 and 2003, the pollination services provided by species.
 Copyrighted Material
520 Conservation Biology
Pimm, Stuart L., and Peter Raven. 2000. Extinction by to coffee production. Proceedings of the National Acad­
numbers. Nature 403: 843–845. The article summarizes emy of Sciences U.S.A. 34: 12579–12582. This work
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Ricketts, Taylor H., Gretchen C. Daily, Paul R. Ehrlich, and assessing the loss of ecosystem functions caused by avian
C. D. Michener. 2004. Economic value of tropical forest declines.

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