Zoo Biology 30 : 18 (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 30:18, 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 difculty 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 wildlifes 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.

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major conservation assets, they must address formidable physical and policy changes as well as species maintenance tasks. People now inuence 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` Lefevre, 2010]. Humanitys numbers and inuence 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 ood 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 changeor 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, nally, 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 humanitys 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 worlds 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, signicant scientic input and much organizational effort.
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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&Ws ndings 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, shes, 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 ve collection planning and action recommendations: (1) Global audit, (2) Global planning, (3) Global species target population sizes, (4) Global investment in professional species managers, and (5) Global longterm 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 zoowild 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 scientic 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, reintroduction, 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 nancial insulation from those seeking to diminish their protections.
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Some zoos are now connecting their guests with the sharp end of conservationnance. 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 dening 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 conservation community and, in particular, provides a science-based innovator, evaluator, and convener unique in the wildlife conservation eld 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 t within New Yorks 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 gures, 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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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, Trachypithecus). Some zoo space is also fragmented between populations of closely related subspecies, such as tigers. The 350 tigers in AZA zoos are divided among ve 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 difcult 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 difculty 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 sufce. 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 specication 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 propagationor best care. Successful specialized propagation programs for bats, beetles, snakes, and sh underscore the point: Specialization is key to every successful threatened species propagation program.
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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 ying 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 successdespite 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 halfdozen countries. Five AZA facilities, the San Diego Zoo, Fossil Rim, The Wilds, White Oak, and National Zoos 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 Attwaters 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 conservationists 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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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 signicantly 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 fulll 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 specic taxa; (2) manage species collaboratively in units large enough to fulll the behavioral, genetic, and demographic requirements of propagation in the light of such monitoring technologies as population viability and risk assessments. Twenty-ve tigers or Wyoming toads will not sufce; (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 zoos ultimate education and conservation goal. BUYING TIME The retention of biodiversity is among the most difcult 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 Earths
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wildlife is starvation, slaughter, and extinction. No credible forecasts suggest that a signicant 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. REFERENCES
Baker A. 2007. Animal ambassadors: an analysis of the effectiveness and conservation impact of ex situ breeding efforts. In: Zimmermann A, Hatchwell M, Dickie L, West C, editors. Zoos in the 21st century: catalysts for conservation? New York: Cambridge University Press. p 139154. Cade T. 2000. Progress in translocation of diurnal raptors. In: Chancellor RD, Meyburg B-U, editors. Raptors at risk. Blaine, WA: WWGBP/ Hancock House. p 343372. Conway W. 1982. The species survival plan: tailoring long-term propagation species by species. AAZPA 1982 Annu Conf Proc 611. Conway W. 1986. The practical difculties and nancial implications of endangered species breeding programs. Int Zoo Yrbk 24/25:210219. Conway W. 1995. Wild and zoo animal interactive management and habitat conservation. Biodivers Conserv 4:573594. Gusset M, Stewart G, Bowler D, Pullin A. 2010. Wild dog reintroductions in South Africa: a systematic review and cross-validation of an endangered species recovery programme. J Nat Conserv 18:230234. Huey R, Losos J, Moritz C. 2010. Are lizards toast? Science 328:832833. Karanth KU, Goodrich J, Vaidyanathan S, Reddy G. 2009. Landscape-scale, ecology-based management of wild tiger populations. Unpublished report for the Kathmandu Global Tiger Workshop, 23 pages. Note: This is a detailed discussion of the problems of preserving tigers. The authors suggest that 25 is a workable number of tigers in a viable population. The number of effective breeders in such a population (Ne) seems unlikely to be more than 12 or 15. Few geneticists or demographers would consider such numbers viable unless connected in a larger metapopulation. Lacy R. 1987. Loss of genetic diversity from managed populations: interacting effects of drift, mutation, immigration selection and population subdivision. Conserv Biol 1:143158. Less intensively managed populations in parks and reserves will require much larger populations to achieve similar Ne, although Lacy calculates that a population with an Ne of 120 could be managed indenitely through the addition of ve new founders in each generation, presuming the management of multiple populations. Lees C, Wilcken J. 2009. Sustaining the Ark: the challenges faced by zoos in maintaining viable populations. Int Zoo Yrbk 43:618. Loarie S, Duffy P, Hamilton H, Asner G, Field C, Ackerly D. 2009. The velocity of climate change. Nature 462:10521055. ` Marton-Lefevre J. 2010. Biodiversity is our life. Science 327:1179. Sanderson E, Jaiteh M, Levy M, Redford K, Wannebo A, Woolmer G. 2002. The human footprint and the last of the wild. BioScience 52:891904. Soorae PS, editor. 2008. Global re-introduction perspectives: reintroduction case studies from around the globe. Abu Dhabi, UAE: IUCN/SSC Re-introduction Specialist Group. 284p. Soule M, Gilpin M, Conway W, Foose T. 1986. The millennium ark: how long a voyage, how many staterooms, how many passengers? Zoo Biol 5:101113. Stanley-Price M, Fa J. 2007. Reintroduction from zoos: a conservation guiding light or a shooting star? In: Zimmermann A, Hatchwell M, Dickie L, West C, editors. Zoos in the 21st century: catalysts for conservation? New York: Cambridge University Press. p 155177. Swaisgood R. 2010. The conservation-welfare nexus in reintroduction programs. Anim Welf 19:125137.

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