Zoo Biology 30 : 1–8 (2011)

COMMENTARY

Buying Time for Wild Animals

With Zoos
William G. Conway
Wildlife Conservation Society, New York

Zoos and aquariums exhibit many rare species, but sustain few for long periods.
Demanding genetic, demographic, and behavioral requirements are a part of the
sustainability challenge, and historical zoo goals and limiting animal management
objectives are another, but they have been overtaken by worldwide wildlife population
contraction and endangerment. New policies are essential for zoo continuance and, if
vanishing species are to be helped by zoo propagation, they must be given priority.
However, zoos have little animal carrying capacity and propagation must be much
more sharply focused. In addition, it is becoming urgent that zoos help to support
parks and reserves and, where possible, manage some especially endangered species
mutually with parks. Zoo Biol 30:1–8, 2011. r 2010 Wiley-Liss, Inc.

Keywords: endangered species propagation; zoo-park collaboration

Inherent in the nature of zoo keeping is a dream of conservation heroics, of

preserving wild species forever. Although zoos may support selected species during
emergencies, it is now clear, in terms of limited carrying capacity as well as genetics,
demography, and behavior, that zoological gardens must also focus on supporting
parks and wildlife environments. Zoos are, in fact, approaching a crossroad. The
global destruction of nature and the difficulty of sustaining wild animal populations
independent of nature increasingly threaten them. What is at stake? Zoos are the
only direct contact the vast majority of humanity has with wildlife’s diversity. Their
potential for advancing public education and conservation by calling attention to
and helping vanishing species and parklands is unique. However, for zoos to become

Grant sponsor: Wildlife Conservation Society.

Correspondence to: William G. Conway, Senior Conservationist, Wildlife Conservation Society,
New York. E-mail: wgcwcs@optonline.net
Received 24 June 2010; Revised 30 August 2010; Accepted 2 September 2010
DOI 10.1002/zoo.20352
Published online 11 October 2010 in Wiley Online Library (wileyonlinelibrary.com).

r 2010 Wiley-Liss, Inc.

2 Conway

major conservation assets, they must address formidable physical and policy changes
as well as species maintenance tasks.
People now influence more than 83% of Earth [Sanderson et al., 2002] and the
Living Planet Index suggests that populations of wild species have diminished by 30%
since 1970. The International Union for Conservation of Nature (IUCN) documents
that 47,677 species are now at risk of extinction, including 12% of the birds, 21% of
the mammals, 32% of the amphibians, and 27% of the reef-building corals [Marton-
Lèfevre, 2010]. Humanity’s numbers and influence continue to expand and no one
knows whether limiting human impacts by democratic means will prove possible.
Climate change is proceeding and rising sea levels are expected to flood low-lying
human population concentrations, agricultural developments, and coastal wildlife
colonies. The stability of many areas protected for wildlife is not promising, as
temperatures rise and reserves become more vulnerable to insect infestations, disease,
and vegetation change as well as to the incursions of agriculturists and developers.
Self-sustaining, nature-regulated populations of large land animals in free-ranging
conditions are becoming uncommon. Saving a species and saving a ‘‘natural’’ habitat
may no longer always be linked in space and time. Protected areas are relatively
independent only where they are so large and ecologically coherent that they require little
care. The ‘‘Greater Yellowstone Ecosystem’’ covering 26 million acres is large enough to
be home for viable populations of such big mammals as bison (Bison bison), elk (Cervus
canadensis), mule deer (Odocoileus hemionus), pronghorn (Antilocapra americana), and
some of their predators, including grizzly bears (Ursus arctos), wolves (Canis lupus), and
cougars (Felis concolor), but very few places are, and Yellowstone is not immune to
climate change—or politics. Even in such big protected areas, it is impractical to sustain
many species of larger mammals [Loarie et al., 2009] without management.
Sustaining big predators that compete with humans and are dependent upon large
prey is especially challenging, but smaller species may also require help to meet genetic,
demographic, and behavioral requirements as well as specialized food needs and threats
from disease, invasive species, and changing climate. As a thought experiment, consider
that one wild tiger (Panthera tigris) in India eats about 50 large (430 kg) prey animals
each year [Karanth et al., 2009]. Multiply by a viable tiger population, perhaps 25
according to Karanth, which few geneticists would consider viable (see note), and then by
as many as the 500 prey necessary to annually produce a 10% edible surplus for each tiger
and, finally, the habitat essential for this herbivore population of 12,500. Thus, a
potentially viable tiger population of 250 might require a larder of 125,000 sizable
herbivores. No wonder some see zoos, with their access to humanity’s kitchen, as lifeboats.

COLLECTION VIABILITY
However, the viability of the collections of the World Association of Zoos
and Aquariums (WAZA) has been carefully studied by Lees and Wilcken [2009], and
their conclusions are not reassuring:

In the World Zoo and Aquarium Conservation Strategy, the world’s leading zoos
commit to focusing their efforts on conserving wildlifey However, to date, this
same group of zoos has largely failed to manage its own populations of wildlife
sustainably despite distinguished calls to action over the past 25 years,
significant scientific input and much organizational effort.

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 Buying Time for Wild Animals 3

Emphasizing genetics and demography, the Lees and Wilcken (L&W) study
makes it clear that zoo collection sustainability failures do not descend from a lack of
understanding concerning the necessarily collective nature of captive preservation
efforts [e.g., Conway, 1982; Soule et al., 1986; and others]. It observes that ‘‘The
science underpinning the management of small populations has been well tested.’’
Nevertheless, L&W’s findings cast serious doubt on the future sustainability of
current zoo collections, to say nothing of their potential contributions to species
preservation.
Although one cannot help but be impressed by the remarkable achievements of
zoos in developing successful care and breeding methodologies for hundreds of
highly specialized birds, mammals, reptiles, amphibians, fishes, and invertebrates,
sustainability requires large numbers and collaborative ongoing species propagation.
No one zoo can for very long maintain a viable population of any sizable terrestrial
vertebrate.
The L&W study produced five collection planning and action recommenda-
tions: (1) Global audit, (2) Global planning, (3) Global species target population
sizes, (4) Global investment in professional species managers, and (5) Global long-
term commitments. Their detailed prescription is a constructive departure point for
the zoo world. Alone, however, it will neither enable zoos to sustain their present
wildlife exhibits nor respond to the global extinction crisis.

BESIDES GENETICS AND DEMOGRAPHY

Although emphasizing genetics and demography, Lees and Wilcken also call
attention to the need to ‘‘make conservation relevant use of zoo space.’’ They suggest
that a part of the strategy might rely upon the addition of new founders [Lacy, 1987]
and upon interactive zoo–wild management. [Conway, 1995; Stanley-Price and Fa,
2007]. However, there is little ‘‘zoo space’’ and most is unsuitable for ongoing
propagation of the species it exhibits. Collection sustainability will require the
creation of ‘‘conservation relevant zoo space’’ and expensive, time-consuming,
revision of zoo exhibition policy. Key is for zoos to make wildlife preservation their
core public service.
Virtually all wild animal populations restricted to reserves will eventually need
some level of curatorial care, such as population management and scientific research
as well as greatly heightened veterinary support and traditional protections. Their
preservation may ultimately entail such zoo tools as accreditation, International
Species Information System, and even Species Survival Plans (SSPs), and zoos and
parks should both be employing such tools as risk and population viability
assessment. Sustaining wildlife in small reserves may also require genetically and
demographically responsive exchanges and translocations. For example, Gusset
et al. [2010] employed focused metapopulation management of African wild dogs
(Lycaon pictus) using fencing, social integration, and translocation for successful
reintroduction. In fact, the best chance of long-term viability for small zoo and park
animal populations may depend upon complementary rescue, medical, reintroduc-
tion, and reservoir functions, despite the health challenges in such processes. Clearly,
zoos should now champion parks in every way that they can, including the provision
of political and financial insulation from those seeking to diminish their protections.

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4 Conway

Some zoos are now connecting their guests with the sharp end of
conservation—finance. Upon opening its ‘‘Congo Gorilla Forest’’ in 1999, the
Bronx Zoo not only imposed a special admission fee to support wildlife conservation
in tropical African forests, but also provided visitors with the opportunity to choose
among ways their fees could be spent with touchscreen voting machines. By 2009,
US$10.6 million had been raised and expended on African forest wildlife
conservation from this source alone. Zoo Boise (Boise, Idaho) has tailored such
an approach even more broadly and is seeking to use a large part of its admission
fees for conservation by defining its zoo as: ‘‘a garden or park where wild animals are
kept for exhibition for the primary purpose of generating funds for the conservation of
animals in the wild’’ [S. Burns and J. Beinemann, personal communication].
In 1978, an imaginative and global network of conservation professionals, the
Conservation Breeding Specialist Group (CBSG) was organized as a part of the
Species Survival Commission (SSC) of the IUCN. Its forte is increasing conservation
effectiveness through the use of science-based, collaborative processes that bring
together people with diverse perspectives and knowledge. It has become a powerful
link between governments, conservation organizations, and others in the conserva-
tion community and, in particular, provides a science-based innovator, evaluator,
and convener unique in the wildlife conservation field and with a special interest in
the potential of zoos to help sustain species.
At best, captive propagation and veterinary help can aid but a tiny proportion
of threatened species, most importantly creatures that nothing else can help. To do
so, however, it must not only meet genetic and demographic concerns, but also
address carrying capacity, propagation management, design and exhibition policies
and, especially, restoration of wildlife to nature or to the novel systems that may
replace it.

CARRYING CAPACITY AND PROPAGATION

In 1982, there were approximately 539,000 mammal, bird, reptile, and amphibia
animal spaces in the 635 collections listed in the International Zoo Yearbook [Conway,
1986]. Even if this space has doubled since 1982, all the zoo animal spaces in the world
could fit within New York’s 212.7 km2 Borough of Brooklyn.
Despite the hard work of zoo scientists associated with such efforts as the
Association of Zoos and Aquariums (AZA), SSPs, the European Association of
Zoos and Aquariums, and the European Endangered Species Programs (EEP), and
their remarkable payment-free transfer of expensive animals, mutual zoo species
selection, and management of vanishing species remains a low zoo priority. In a
penetrating overview of the effectiveness of ex-situ zoo breeding efforts, Baker [2007]
applied criteria established to assess the likelihood of long-term population viability
to, for example, an analysis of 10 species recommended for SSP management in the
AZA Old World Monkey Regional Collection Plan. She concluded that none ‘‘has
much chance of surviving for an extended period of time.’’
Checking current figures, I found that only a tiny fraction of SSP
Cercopithecines is kept in appropriately composed ‘‘breeding’’ populations. In
nature, most live in groups. In zoos, they are commonly exhibited in pairs where they
rarely breed. For example, in 2007, the 40 Diana monkeys (Cercopithecus diana)
separated among 16 AZA zoos produced no births. Even mandrills (Mandrillus

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 Buying Time for Wild Animals 5

sphinx), which naturally live in large multimale troops, are mostly kept in
unproductive pairs (108 mandrills in 35 zoos, only 2 born in 2009). The situation
is much the same with mangabeys (Cercocebus) and langurs (Presbytis, Trachy-
pithecus). Some zoo space is also fragmented between populations of closely related
subspecies, such as tigers. The 350 tigers in AZA zoos are divided among five
subspecies.
Where mate choice is extremely limited, best propagation practices cannot be
applied. The depressive effects of undersized groupings plus lack of SSP
collaboration also affects many zoo populations of nonsocial carnivores, birds,
and reptiles. It is not that SSP has been tried and found wanting, but that it has been
found difficult and left untried. Unpublished AZA studies by Bob Wiese (San Diego
Zoo) and Steve Thompson (Lincoln Park Zoo) suggest that the majority of SSP
populations ‘‘neither approach nor show appropriate progress toward viability.’’
There are only 115 species in the AZA SSP, mostly considered unsustainable.
Despite the difficulty of administering SSP and EEP programs, for the present, zoos
have no convincing substitutes for sustaining the viability of small isolated zoo
populations.
It is not surprising that rhinos require more space than frogs and tigers more than
turtles, but it highlights how zoo species selection and carrying capacity are
indissolubly linked. Captive care, such as ecology in nature, is scale dependent.
Nevertheless, agreeing on species selection and providing carrying capacity for
vanishing species and collection sustainability is urgent. Unfortunately, straightforward
taxonomic and rarity criteria are rarely applicable. Time-limited populations and range
state dependent negotiation may have to suffice. In any case, without new exhibition
policies, zoos will be unable to sustain rare wildlife or their present collections.

DESIGN AND EXHIBITION POLICY

Given the inability of any one zoo to sustain enough of a larger species to
achieve viable populations, exhibit planning requires the ‘‘specification’’ of future
species sources as much as of design and building materials. Exhibits should be
thought of as species preservation units. In any case, 21st century animal exhibit plans
lacking sustainability programming are building a road to nowhere. Nowadays, a
zoo species may have to be considered in the context that it may be among the last of
its kind, part of an international propagation effort, and a long-term commitment.
The future of zoos could become the development of specialized species plans
and of zoo operating models exhibiting fewer kinds of animals more compellingly
and jointly with parks. Relatively small but dedicated collaborations as well as broad
international programs may turn out to be effective models for some parts of the
global effort. Consider birds.
Sustained breeding of diverse bird species in zoos is rare, whereas specialized
programs have been famously successful. The International Crane Foundation, The
Peregrine Fund, and the Wildfowl & Wetlands Trust, Slimbridge, are examples,
along with specialized programs, in a number of zoos. In contrast, traditional zoo
bird collections exhibiting scores of species, often in the same aviary, are usually
incompatible with sustainable propagation—or best care. Successful specialized
propagation programs for bats, beetles, snakes, and fish underscore the point:
Specialization is key to every successful threatened species propagation program.

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6 Conway

Fortunately, useful propagation of endangered species need not always be long

term if combined with reintroduction. In 1988, the last wild California condors
(Gymnogyps californianus) were brought into captivity. There were only 22. There
are now 325, 168 flying free in Arizona, California, and Baja Mexico. There were but
16 whooping cranes (Grus americana) left when their captive propagation began;
presently there are approximately 400, 200 in captivity and 200 in nature. Mauritius
parakeets (Psittacula eques) were increased from less than 12 to 343 by 2007, and
black-footed ferrets (Mustela nigripes) from 18 (7 breeders) to about 750 at present.
Arabian oryx (Oryx leucoryx), American bison (B. bison), Wyoming toads (Bufo
baxteri), and many other declining species have been increased in captivity,
reintroduced, and then managed more or less free-ranging with considerable
success—despite the diminution of original habitat but not without continuing costs.
If zoos are to support species in trouble, whether from lost habitat and overhunting,
such as Sumatran rhinos (Dicerorhinus sumatrensis) and saolas (Pseudoryx
nghetinhensis), or disease, such as Tasmanian devils (Sarcophilus harrisii) and many
frogs, they must reduce the number of species they maintain that do not need help
and specialize in species that do.
The Amphibian Ark is a specialized multiinstitution program addressing the
worldwide collapse of amphibian populations, by providing support for species that
cannot currently be adequately protected in the wild. Its principal partners are
WAZA, the CBSG, and the IUCN/SSC Amphibian Specialist Group. A research
and breeding program is underway in both zoo and nonzoo institutions and half-
dozen countries. Five AZA facilities, the San Diego Zoo, Fossil Rim, The Wilds,
White Oak, and National Zoo’s Conservation and Research Center, have created a
consortium of spacious ‘‘Conservation Centers for Species Survival’’ (‘‘C2S2’’).
Beyond an emphasis upon ungulates, it is working with the reintroduction of
California condors, black-footed ferrets, desert tortoises (Gopherus agassizi), kit
foxes (Vulpes velox), and Attwater’s prairie chickens (Tympanuchus cupido attwateri).
Many specialized efforts are underway including some for bats, wolves (C. lupus),
and various snakes, but many more are needed. Yet, species selection for zoo
propagation is in its infancy and the ground keeps shifting under the conserva-
tionist’s feet. New evidence [Huey et al., 2010] suggests that lizards may be as
threatened as frogs.

RESTORATION TO NATURE
Despite successful reintroductions of bison, whooping cranes, condors,
peregrines (Falco peregrinus), and black-footed ferrets, most early restoration
attempts were unsuccessful, but reintroduction science is evolving [see: Soorae, 2008,
but also Swaisgood, 2010]. Reintroduction attempts grew from perhaps 100 species
in the early 1990s to several hundreds by 2005, 74% of them mammals and birds and
with increasing success. For example, between 1986 and 2000, 21 of 28 raptor
reintroductions resulted in the establishment of breeding populations and most used
captive-bred young [Cade, 2000]. A remarkable 80% of bird reintroductions in New
Zealand have proven successful.
In 2008, a ‘‘First International Wildlife Reintroduction Conference’’ was
organized in Chicago by Joanne Earnhardt of the Lincoln Park Zoo, conservation
biologist Devra Kleiman, and Frederic Launay, Chair of the IUCN/SSC Reintroduction

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 Buying Time for Wild Animals 7

Specialist Group. It took place at the Lincoln Park Zoo, and was an update of what
was known and successful and what was not and was not known.
Earnhardt emphasized that most reintroductions have been very poorly
documented, which stimulated her creation of an ‘‘Avian Reintroduction &
Translocation Database’’ on the Web. By January 2010, it included 1,762 release
events of 173 species, and restoration, reintroduction, interaction, and translocation
seem likely to become a focus of both future science and conservation action.
A species sustained ex-situ for several generations may come to restoration
significantly changed, behaviorally if not genetically. Its former habitat, plants,
climate, and remaining animals may be even more changed, and the new species
associations such changes bring will make it all the harder to reintroduce species
removed from the ecological reassortment taking place over lengthy periods of time.
This argues for a continuous interchange of captive and wild populations to sustain
the viability of ex-situ populations for reintroduction, even though readjustments
may require intensive management [Swaisgood, 2010]. Thus far, however,
reintroduction is proving an inspiring way to arouse support for wildlife and
habitat preservation. Who would have thought that condors, black-footed ferrets,
whooping cranes, and wolves would have a further chance in America, or echo
parakeets in Mauritius and kakapos (Strigops habroptilus) and saddlebacks
(Creadion carunculatus) in New Zealand? Who could doubt that if zoos still had
Carolina parakeets (Conuropsis carolinensis), thylacines (Thylacinus cynocephalus),
pink-headed ducks (Rhodonessa caryophyllacea), sea minks (Mustela macrodon), or
dodos (Raphus cucullatus), each would become an incentive for wildlife restoration
and a powerful stimulus for environmental education and the preservation of
protected areas?
But, for zoos to fulfill their conservation potentials, they must address not only
the ongoing task of better meeting the biological requirements of wild animals, but also
that of revising zoos as reservoirs of rare wildlife and of parkland support. Despite the
all too apparent obstacles, the survival of zoo and park animal populations is likely to
be more successful where their management can be mutually supportive.
In summary, it is suggested that zoos seek to: (1) specialize in agreed SSP
species and other cooperative breeding programs, reducing collection species
numbers, and tailoring facilities and programs so as to provide the best conditions,
expertise, and population resilience for specific taxa; (2) manage species collabora-
tively in units large enough to fulfill the behavioral, genetic, and demographic
requirements of propagation in the light of such monitoring technologies as
population viability and risk assessments. Twenty-five tigers or Wyoming toads will
not suffice; (3) share rare species management and medical care with reserves and
support planned reintroductions, translocations, and related efforts; and (4)
champion protected area needs and involve zoo visitors in their support as well as
that of the zoo. The survival of protected wildlife is becoming the zoo’s ultimate
education and conservation goal.

BUYING TIME
The retention of biodiversity is among the most difficult challenges facing
contemporary cultures. Society has not yet come to terms with the fact that the
alternative to protection, management, and care for a growing percentage of Earth’s

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8 Conway

wildlife is starvation, slaughter, and extinction. No credible forecasts suggest that a

significant representation of wild lands and wildlife will remain when and if human
population growth and habitat destruction cease. During the next 100 years, the only
hope for many wonderful and charismatic species will be zoo care and park
stewardship. The potential zoo role in the intensifying nature of wildlife conservation
is unique, and it is up to zoos as well as parks to save as many species as they can for
as long as they can, to buy time for wildlife.

ACKNOWLEDGMENTS
Elizabeth Bennett, Onnie Byers, Robert Lacy, Jim Doherty, John Robinson,
and David Western read an earlier version of this article and provided many helpful
suggestions and insights. Four anonymous Zoo Biology reviewers made important
and constructive criticisms. I gratefully acknowledge the long-term support of the
Wildlife Conservation Society that has made so much of my work possible.

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