Download - Wild and zoo animal interactive management and habitat conservation

Biodiversity and Conservation 4,573-594 (1995)

Wild and zoo animal interactive management and habitat conservation WILLIAM CONWAY Wildlife Conservation Society, Bronx Zoo/Wildlife Conservation Park, Bronx, New York, NY 10460-1099 USA

Received 8 August 1994; revised and accepted 4 September 1994

This review considers interactive management of wild and zoo populations as a strategy to support habitat preservation, help sustain key endangered species, and hasten the evolution of zoos and aquaria as proactive conservation organizations. Interactive management supports key species’ subpopulations in an integrated fashion, using their study in nature as a way to understand wildlife habitats, ecological processes and conservation threats.

In the face of human increase and habitat destruction, the survival of much wildlife will depend upon the utility of fragments of habitat and the survival of relatively small populations whose habitats are reduced or altered and whose numbers are capped. Under such conditions, interactive wild-captive metapopulation strategies may increase the security of key species.

Captive propagation skills and urban locations pre-adapt zoos as headquarters for nature preservation. Thus, a key objective in zoo evolution is to focus upon the species and its habitat as the unit of conservation, not the species alone.

Keywords: interactive management; zoo evolution; habitat preservation; conservation costs.

Introduction

What is wild-zoo animal interactive management? As used here, interactive management (IM) is a strategy of species and habitat preservation that relies upon interpopulational coordination of threatened species’ metapopulations (Hanski and Gilpin, 1991), which includes populations living in nature and in captivity. Although the basic idea of interactive metapopulation management for key species is straight-forward (Foose, 1991), its modification as a strategy for habitat preservation, like any other habitat preservation effort, is complex (May, 1991; Stanley-Price, 1991) for it goes far beyond exchanging animals (Conway, 1989a, 1989b, 1991,1995; Mallinson, 1991). IM’s real strength is found in combining the resources of wildlife managers with those of biologists working in zoos. Thus all the metapopulations of a wild-zoo programme are aimed at preserving both the species and its habitat in nature. IM focuses zoo conservation efforts on the preservation of wildlife communities and habitats as a component in the conservation process rather than upon captive propagation. IM applications are limited, as are those of endangered species propagation, and should not be used as a substitute for habitat preservation or less intensive methods of conservation.

IM identifies ‘key species’ and treats their subpopulations in a more or less integrated fashion, utilizing their study in nature as a way to understand wildlife habitats, ecological processes and conservation threats. Key species may be chosen on the basis of several 0960-3115 0 1995 Chapman & Hall

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criteria in addition to rarity and uniqueness, but in habitat preservation will be most useful when chosen for the significance of their ecological roles and/or ‘flagship’ qualities (see also Stuart, 1991) mindful that where actual population interchanges between nature and captivity occur, many precautions must be taken.

Theoretically, exchanges of animals or germ plasm (wild to captive, captive to wild) between subpopulations of an endangered species in IM would (i) sustain a higher population, more secure than otherwise possible for small circumscribed groups confined in diminished natural habitats; (ii) supplement wild populations where that was found to be the best option for their survival; (iii) enable zoos to maintain greater numbers of endangered species reservoirs by, in some instances, reducing their individual size (but see Willis and Wiese, 1993); and (iv) help to maintain the genetic integrity of such capped populations in both nature and captivity.

In concept, IM would support each of its selected habitats by establishing an ongoing relationship with local communities and authorities, acting as conservation advocate. providing education, inspiration, veterinary and other management expertise, training for local staff, information for local decision-makers and special support for habitat preservation, restoration and expansion of protected areas. It is visualized as a long-term commitment akin to the relationships developed by the Frankfurt Zoological Society in parts of Africa, the Jersey Wildlife Preservation Trusts in Mauritius, Madagascar and elsewhere, Wildlife Conservation Society (formerly New York Zoological Society) in parts of Africa and Latin America (Conway, 1991), and the Minnesota Zoo in Indonesia (Tilson, 1991).

In zoos, captive components of interactive populations are employed as surrogates for wild animals in endangered species research (primarily behavioural, reproductive. veterinary and curatorial) and as inspiration for the development of conservation support and education programmes. Each key species is connected to an in situ wildlife preservation programme. Zoos are already the intellectual home of many conservation efforts but they have the potential to strengthen their roles as a logistic base and source of support for direct habitat preservation.

Translocations between natural wild animal populations have been long used to resolve genetic, demographic, social and behavioural problems, often with regrettable results (Grieg, 1979). Reintroduction from captive-bred populations to areas where they are extinct in nature is as rare as correction of the reasons for disappearance. Long-term interactive management, the subject of this review, has seldom been used but the restoration of the Arabian oryx (Oryx leucoryx) (Stanley-Price, 1989), American bison (Bison bison) (Bridges, 1974) Lord Howe Island wood rail (Trichofimnas sylvestris) (Fullager, 1985) and, perhaps, the bald eagle (Haliaeetus leucocephalus) are comparable. The effort to strengthen the population of the golden lion tamarin (Leontopithecus rosalia) in part of its range in Brazil is a carefully documented interactive management application. although the costs of this pioneering programme may be exceptionally high (Kleiman et al.. 1991).

In this review, it is assumed that IM reintroductions would only be attempted to repopulate an area where the target species is extinct or so threatened that the risk of extinction without IM exceeds the risk of intervention; essentially under the same guidelines as for reintroduction alone (Stanley-Price, 1991: Wilson etal., 1994), or, in some instances. under the guidelines for safe translocation (IUCN, 1990) and where the opportunity for preservation of critical habitat through IM is judged promising.

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Why interactive management?

Why not devote all conservation effort to straightforward habitat preservation and all captive propagation efforts to preserving animals in zoos? The short answers are that IM is yet another way of making support available for habitat preservation and for understanding its needs that would otherwise be unavailable and for sustaining animals in what is left of nature that might otherwise be lost. The longer answer is that nature conservation is on the edge of an appalling chasm of extinction, the threshold of an era where traditional conservation methodologies based upon animals living or being reintroduced in a relatively vast self-sustaining ‘wild’ need to be rethought. We must begin to consider a vastly altered world wherein wildernesses are becoming agricultural developments and where habitat and wildlife population changes are occurring with bewildering speed - and where many wildlife communities will survive only as ever more intensively cared for ‘megazoos’ (Conway, 1989a; Foose, 1989).

There are about 97 million more human beings every year, perhaps 2.5 billion pounds of human flesh every twelve months; a proliferation fuelling exponential habitat change. Nearly 90% of this population growth is taking place where the most diverse wildlife dwells, in less developed tropical countries. Deforestation is proceeding at the rate of more than 16.8 million ha each year (UNEP, 1992) and yet some agricultural experts argue that nearly three times as much land as is currently farmed, an additional 2.1 billion ha, could be profitably cultivated (Bongaarts, 1994). We stand to lose hundreds of thousands, perhaps millions, of species of invertebrates (Erwin, 1988; Wilson 1989) and thousands of vertebrates.

Extinctions begin with the loss of numbers. As a species’ biomass declines, reduced populations become ecologically impotent, no longer fulfilling the dynamic habitat functions of seed dispersal, selective cropping, predation and the like. These inadequacies accelerate the momentum of dysfunction, decline and extinction.

Self-sustaining ecosystems, rather than fragments, will be very limited in the future. Although vulnerability and loss of fragmented habitats and wildlife populations characterize the phenomena of extinction (Bierregaard and Lovejoy, 1989; Diamond, 1989; Newmark, 1991; Kattan et al., 1994), destruction elsewhere makes the concentrations of wildlife in tiny parks and reserves important reservoirs of diversity. IM is a way for zoos, working with local communities, to support the survival of such fragments.

The plight of India’s 150 square mile Ranthambhore National Park, an island of diversity in a sea of agriculture, set up 21 years ago as a part of the international rescue effort called Project Tiger, brings the reality home. Two hundred thousand people live within three miles of the park’s core in 60 villages. During the past four years as many as 20 tigers have been killed in the Park ‘for the China tiger bone trade’; only 19 to 25 remain. The deer and other animals which constitute the tiger’s food resources have been decimated. But, the tattered park is the main source of firewood for the villagers and the fodder on which they feed their 150000 head of cattle, goats, buffalo and camels (Ward, 1994).

No matter how well protected, a 150 square mile park and the hunting lands immediately surrounding it are not likely to sustain for long the several hundred tigers necessary to form a viable population, if isolated. Maintaining this fragment’s viability may not be met by managing its food resources (or provisioning), or by translocations. It may simply be hopeless, or it may be a long-term task for IM. Perhaps helping to understand the options,

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expanding the park, helping in the task of aiding nearby villages, supporting local wildlife authorities can save and restore this habitat - help sustain a subpopulation of tigers.

More and more wildlife are destined to survive, if at all, in small, disjunct and fragmentary populations with no room to grow. It is against this background that long-term care strategies such as zoo-wild interactive management and the organizational and community involvement that they imply must be considered.

Employed cautiously, with scientific rigour, and managed in response to both wildlife and human concerns (Conway, 1988; Stanley-Price, 1989: May 1991; Caughley, 1994). interactive management of wild and zoo animal populations could make the difference between extermination and survival for a significant number of treasured wildlife communities and their habitats over the next few decades.

How much can zoos help?

Zoos collaboratively manage metapopulations in small disjunct units, and continue to accrue expertise in this speciality. Although zoo capacity for numbers of species is extremely small, variable and focused upon vertebrates, biodiversity conservation is not just about saving numbers of species. It is about saving functioning systems and communities in which not all species are equal. The judicious management of key species has positive multiplication effects throughout the community. Therefore, special efforts, zoo efforts, to conserve such species, have a role in the future of conservation.

In recent years, zoo managers have bred at least 19% of all the species of mammals and 9% of the birds (Conway, 1986). They have also bred sizeable numbers but small proportions of the amphibians and fish but, of course, only an insignificant proportion of the millions of invertebrates. Those who believe that biodiversity preservation is primarily a numbers exercise correctly state that zoo breedings have contributed little to the preservation of biodiversity - and could add that they never will.

Few taxa, even of vertebrates, have been zoo-bred on a long-sustained basis, certainly less than 250. Until recently, it was not considered a priority to do so - nor were propagation skills sufficiently well developed in zoos until the past decade or so. But reservations about the propagation potential of particular species on an it-hasn’t-been- done basis (Derrickson and Snyder, 1992; Rahbek, 1993) are insufficient.

After all, zoo species capacity is small. Propagation and reintroduction will be relatively uncommon. The fact that zoos would find it difficult efficiently to propagate high numbers of certain specialized forms (Rahbek, 1993; Wilson et al., 1994) does not detract from the significance of their ability to propagate important key species which have no other preservation options.

Potentials are indicated in a recent compilation (calculated from Rahbek, 1993): zoos bred a remarkable 85.7% of the IUCN listed threatened species of mammals that they held in captivity, 87.8% of the birds, and 88.8% of the reptiles. Yet, the total number of threatened taxa in zoos was only 878 of the 2 01.5 listed by IUCN in 1990. Zoos are daunted by the bureaucratic procedures which must be surmounted to acquire rare animals (which can take years in the USA) and captive propagation space is extremely limited.

Nearly ten years ago, I calculated that all of the zoo animal spaces in the world could fit within New York’s Borough of Brooklyn; that there were about 539000 spaces for mammals, birds, reptiles and amphibians, judging from then current collection sizes (Conway, 1986). Recent studies by the IUCN-Captive Breeding Specialist Group (CBSG)


has raised this figure to perhaps 700000 (T. Foose, personal communication), but the Brooklyn-sized (212.7 km2) zoo habitat calculation may still be correct. And, beyond the limitations of space, are those of collaborative animal management.

The necessity for coordinated regional and global strategies is inescapable for long-term propagation of small animal populations (Conway et al., 1984; Conway, 1989~; Maguire and Lacy, 1990; Diebold and Hutchins, 1991; Hutchins et al., 1994; Seal et al., 1994) and immense strides have been made in collaborative zoo collection record-keeping. But the planning, coordination and communication needed to realize propagation goals is a growing financial burden.

Long-term zoo species maintenance efforts for vertebrates, outside of the test tube, may be capped below 500 taxa for the immediate future. The hope that a high proportion of zoo-aquarium collections could so efficiently coordinate their efforts that they might sustain thousands of species for the long term is as unrealistically optimistic as similar expectations for any comparable human effort, or for ecosystem preservation. Nevertheless, zoo capacities for short-term programmes, such as the Arabian oryx’s propagation and reintroduction, are much more promising. They offer uniqe emergency resources.

Help from new technologies?

New genetic technologies enabling more certain identification of taxonomic and even familial relationships and new geographic positioning tools strengthen all aspects of conservation biology, nowhere more so than with the relatively intensive strategies of IM. Theoretically, artificial insemination, embryo transfer and related technologies offer a way not only of increasing species capacity but also of effecting interpopulational moves without some of the social, disease and behavioural disruptions that attend moving individuals. It would be much easier to move germ plasm rather than a new male tiger (Panthera tigris) or black rhino (Diceros bicornis) into the established territories of others. Besides, where frozen sperm, ova and embryos can be long preserved for later use, inbreeding or genetic drift, in nature or in captivity, might be prevented (Dresser, 1988). Thus far, however, the technology has been worked out for less than three dozen wild species of mammals and birds and the chances for broad application appear to be receding, because of limited resources.

What is the numerical potential of interactive zoo-wild management?

Risking the curse of non sequitur, it is stimulating to estimate how interactive metapopulation and habitat preservation, if more generally adopted, might improve the contribution of zoos to biodiversity conservation. Given that there are at least f 100 zoos in the world (of greatly varying capabilities) with a combined attendance exceeding 600 million, this is a significant question.

Consider that there are about 47 500 vertebrate species, 24 500 of these are sharks and bony fish and 23 000 are mammals, birds, reptiles and amphibians. More than 25% of the tetrapods and 10% or more of the fish are threatened with extinction during the next 100 years. to say nothing of the invertebrates (Foose, 1992). Perhaps 12% of the tetrapods have representatives in zoos and aquaria, or 2 700+ species.

If zoos and other captive collections could sustain 300 taxa of key mammals, birds, reptiles and

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amphibians, which have no current opportunity for reintroduction, for 75 years or more. they would provide an alternative to extinction for at least 15% of IUCN listed threatened terrestrial vertebrates. This is a reasonable expectation. given current progress. Few other conservation strategies offer options for species losing their habitats. But. the number of threatened vertebrates is rapidly growing.

If zoos and similar institutions would increase their species care capacity by 2.5% a year. by 2025 the number of taxa they could care for on a sustained basis would be 600+. This may be a realistic expectation but, by 2025,600 will be less than 15% of then threatened species.

However, if zoos made interactive management of key species a high priority by adding the care of significant habitats for 100 key species by 2025, the number of taxa saved might number in the tens of thousands and the ratio of species care and management could be greatly reduced. The in-zoo subpopulation of each IM species might also be reduced making room for other needy species.

Only the integration of habitat preservation support as an essential, ongoing, zoo operation, eventually a prerequisite to the initiation or continuing operation of a zoo or aquarium, is new in this description.

Preserving a species, say an Andean Darwin’s rhea (Pterocnemia darwini garleppi), in captivity can sustain creatures that may otherwise be lost. It can arouse public sympathy and contribute to understanding. Preserving it in tandem with its habitat, and all of the life forms within it -from llareta plants to horned coots, James’ flamingos to southern vicuiias, Tatio frogs to Andean cats, cinclodes, chinchillas, Pentlandt’s tinamous and thousands of species of invertebrates, interactive management could have a conservation multiplier effect beyond calculation.

Interactive management: a proving ground for conservation biology?

Conservation strategies based on interactive species management must call upon every insight of conservation biology, but they also offer the opportunity to test them (May, 1991: Stanley-Price, 1991). In a masterful review, the late Graeme Caughley (1994) described current conservation biology as dwelling within two paradigms: the ‘small population paradigm’ is concerned with populations vulnerable to extinction because their numbers are small, and the potential for population increase thereby limited. It is applicable to most captive animal populations and small wild populations with ‘capped’ numbers: that is, limited opportunity for increase, usually in habitat fragments, reserves or zoos.

Such populations are vulnerable to demographic stochasticity, environmental variation, catastrophes, genetic processes and the like. This paradigm has strong theoretical foundations which have not only guided the development of zoo collection strategies but also conservation biology in general. A salient feature of Caughley’s small-population paradigm, not highlighted by him, is its focus upon single or small groups of species.

In contrast, the ‘declining-population paradigm’ is concerned with identifying external agents responsible for the decline of a species population, and addressing the causes. This paradigm is more empirical than theoretical, is applicable to most wild populations, and is relevant to field conservationists - a ‘First, find out what’s wrong’ approach. Caughley concludes that an analysis based upon one paradigm will have limited applicability to the other, that the declining population paradigm badly needs some theory, but he finds little evidence of the utility or utilization of small population theory in field conservation thus far. In my experience, field conservation more commonly focuses upon habitats and communities than single species.


Inherently small-population based, interactive wild-zoo management straddles Caughley’s paradigms for it also gets its boots muddy in the field and seeks to preserve habitat. The IM conservationist moves back and forth between the controllable captive environment and the complex communities of nature park or reserve, where the targeted subpopulation affects and is affected by other species and factors beyond control or anticipation (Fullager, 1985; Stanley-Price, 1989).

While IM populations will be so small as to fit the small-population paradigm, the immediacy of extrinsic factors will over-ride most of the intrinsic biological ones, placing the empirical declining-population paradigm in the driver’s seat - in a stochastic sense. In the game of survival, a pack of stray dogs, herd of cattle, infectious disease or a family of poachers beats two pair of genetic drifts and an age imbalance on the first hand, but they are not necessarily less susceptible to study, prevention and management.

Thus, in interactive management, success and utility will always be tied to an experimental approach, an investigatory point-of-view. Investigation, research and management experimentation with a project’s ‘key species’ are a way of understanding the system and gaining insights into the community. Such an approach is exemplified in the recovery projects for the Howe Island rail, golden lion tamarin and peregrine falcon (Fafco peregrinus). Meant to be relatively long term with captive population components susceptible to close-up study and testing, zoo based resources and regular on-going management relationships, IM can provide the conservation biologist with unparalleled opportunities to test theory and solve problems. Unfortunately, where IM touches captive propagation, it has the potential of substituting captivity for preservation in nature and the movement of populations, whether from captivity to nature or nature to captivity, carries its own set of dilemmas.

Doubts, problems and dilemmas

The propagation of endangered species in captivity, presumably a part of many IM programmes, could run the danger of replacing habitat preservation, compete for conservation monies that could be used to save the species in nature, focus conservation upon single species rather than wildlife communities, transmit disease to natural populations through reintroductions, or produce populations too altered by long-term captive breeding to be suitable for reintroduction into the wild.

Moving animals from reserves to zoo back-up populations and from captivity to nature is a prescription for a broad spectrum of problems (CBSG, 1991: Stanley-Price, 1991; Woodford and Kock, 1991). Subtracting potential breeders from small populations in nature can reduce their viability, while capture and acclimatization processes are fraught with danger for the animals concerned - and the reintroduction processes of the past have often been hazardous and expensive (Kleiman, 1989; Beck et af., 1994).

Although some risks are species-specific, and can be used in determining the suitability of particular interactive applications, IM differs from conventional capture and reintroduction or translocation programmes in that it anticipates ongoing interpopulational exchanges. This not only means more moves, more monitoring, and more management but, on the positive side, more research, more zoo-based medical and curatorial resources and more opportunity for public education, response and involvement. The challenges of related kinds of management have been extensively reviewed by many including Seal et al. (1994), Kleiman (1989). Stanley Price (1991), Stuart

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(1991) Woodford and Kock (1991) Wilson et al. (1994) and Caughley (1994); the same concerns crop up again and again. Even though the goal of IM is to use zoo support for habitat and species preservation, these concerns deserve re-statement and re-examination.

Will interactive management reduce the impetus for habitat preservation?

Will interactive management programmes feed a public perception that the species is the unit of conservation, not the habitat? Will the political suggestion that a species might be captive bred and reintroduced in the future, as an alternate to preserving it in nature, cloud the real issue (e.g. Zyla, 1994)? The danger is real with simplistic ‘breed and reintroduce’ programmes.

Consider the disproportionate attention and dollars given to construction of expensive propagation facilities for giant pandas (Ailuropoda melanoleuca) in China, rather than to the essentials of habitat protection and poaching control (Schaller, 1993). However, it is debatable whether US government support for the propagation of black-footed ferrets (Mustela nigripes), California condors (Gymnogyps californianus), whooping cranes (Grus americana), peregrine falcons and bald eagles (Haliaeetus leucocephalus) competed with other better uses. Compensatory habitat preservation efforts would almost certainly not have occurred in time for these species (or were not possible, e.g. alleviating DDE effects on birds of prey) and it may be that interest in these species generated funds that would not otherwise have been available had none of these programmes taken place. IM programmes would have sought to integrate habitat preservation within them.

Cost of conservation

Cost-of-conservation accounting is confused when conservationists ignore the fact that monetary appropriations are not freely interchangeable but determined by their sources and purposes. Rahbek (1993) referring to the cost of US peregrine falcon propagation and restoration work (Cade, 1988) observed, ‘Consider how much just a fraction of these resources would mean if used elsewhere for the preservation of biodiversity’. He goes on to suggest that protection of Ecuador’s Podocarpus National Park would have been a much better investment. But the US peregrine falcon appropriations could not have been raised for Podocarpus.

Fortunately, propagation programmes, such as that for the peregrine falcon, can become powerful levers for increasing public awareness and the preservation of habitat (Durrell and Mallinson, 1987; Mallinson, 1994). The American bison’s re-establishment from zoo, farm and translocated stocks in the US West between 1907 and 1917 (Bridges, 1974) and the recent effort to establish new populations of the golden lion tamarin in the Poco das Antas Reserve in Brazil (Kleiman, 1989) not only led to the preservation of critical habitat and many other species but also to the establishment of new levels of community involvement in wildlife conservation.

Even where habitat has been so altered as to make its preservation in an original state impossible, reintroduction from captivity can become a fulcrum for restoration programmes and sympathetic community attitudes. Further examples are the exemplary work of the Wildlife Preservation Trust for Mauritius pink pigeons (Columba mayeri) (Jones et al., 1992) and kestrels (Falco punctatus) (Jones et al., 1991).

The lesson is not that the public’s seemingly inevitable single species-focus is bad for conservation but that properly used, it can be a useful key to the preservation of systems.


Spreading disease

The potential for infecting critical captive or wild parts of a metapopulation with infectious disease during exchanges of individuals between subgroups is a major obstacle to almost any kind of interactive management with wild or domestic animals (Woodford and Kock, 1991). In interactive management, used only after carefully considering other conservation options, the problem should be of less significance.

Vulnerability can be unexpected, as in an outbreak of canine distemper, perhaps obtained from domestic dogs, which has recently killed at least 60 of the 3000 lions (Panthera led) in Tanzania’s Serengeti National Park (Morell, 1994). This is the same disease that nearly extirpated the black-footed ferret when accidentally introduced into the captive population after it had wiped out the wild population (Thorne and Williams, 1988).

Hess (1994) observes that any conservation activity that increases the movement of individuals among populations, even the development of ‘conservation corridors’, may spread disease. Mainly for fear of disease transmission, Wilson et al. (1994) referring especially to the Puerto Rican parrot (Amazona vittata), argue against a proposal by Lacy (1987) that a managed population be subdivided and dispersed into several breeding subgroups to avoid catastrophic loss, exchanging a few individuals between populations each generation to reduce genetic drift-thereby creating an interactive metapopulation. Lacy’s and Wilson’s views present aspects of the conflict between Caughley’s ‘small population paradigm’ and ‘declining population paradigm’.

Wilson et al. (1994) offer four more or less theoretical propositions that, if fully accepted, would greatly diminish consideration of reintroduction and interactive management applications from all zoo and most governmental animal propagation programmes: (i) endangered species which have gone through genetic bottlenecks may have heightened susceptibility to disease; (ii) zoos and other multi-species facilities are high risk environments favouring the transmission of pathogens; (iii) breeding of K-selected species (such as the Puerto Rican parrot) in which mate selection is idiosyncratic is more difficult in small subdivided groups (d la Lacy) and (iv) transfers between small populations to overcome this problem and enhance mating increases the possibility of transmitting disease.

The evidence that some endangered species which have been through a genetic bottleneck (e.g. cheetahs mentioned in Wilson) are exceptionally vulnerable to disease is equivocal (Lande, 1988; Caughley, 1994). Endangered status and diminished numbers are not necessarily equivalent to a genetic bottleneck, as recent studies of, for example, the diminished black rhinos (Diceros bicornis) have shown (G. Amato, personal communication). Nevertheless, such animals should be treated as though they are exceptionally vulnerable.

The generalization that multi-species as compared with single-species facilities are high risk environments makes intuitive sense, but is far too broad without further description. Not only are they likely to have personnel with far greater expertise and resources but few single-species facilities outside of laboratories have enjoyed the tradition of high quality medical care and pathology reporting routine in many major zoos. The very short time (less than ten years) over which closely managed endangered species breeding programmes have been in place in zoos (see below) makes the use of older data dubious. Zoo disease reservoir assumptions fail to cite comparative data from wild, domestic animal and human populations. Actually, translocation from extant wild populations may pose

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the larger disease danger (Grieg, 1979) not to mention the burgeoning domestics. Sustaining some IM species in multi-species institutions may provide them the best care - a species-specific matter requiring careful evaluation,

Mate selection in K-selected species is species-specific and not fundamentally impractical in small groups. The majority of the 70+ SSP species, from gorillas (Gorilla gorilla) to Andean condors (Vuftur gryphus), are K-selected, and bred in small groups. Breeding populations may be small in intensive management.

Still, any regrouping of animals risks a transfer of disease, as do their natural movements in nature. Wild species never live in a disease vacuum. For example, 1993-4 studies conducted by Wildlife Conservation Society field veterinarian William B. Karesh and his colleagues at four seabird colonies in Peru and Argentina focusing on penguins, (Spheniscus humboldti) and (S. mageffanicus), found Chlamydia (psittacosis) in 38 of 60 penguins, Salmonella in two, and paramyxoviruses (Newcastle’s disease) in more than 25% of all tested. Karesh et al. (1994) collected blood and faecal samples from 77 free-ranging duikers of five species in Zaire’s Ituri Forest finding positive antibody titres for leptospirosis, bluetongue, infectious bovine rhinotracheitis, and epizootic haemorrhagic disease in almost all. The implications for translocations as well as for zoo-wild interactive management are essentially the same as those dealt with in the regular acquisition of animals from other collections or from nature. The IUCN Captive Breeding Specialist Group has initiated analysis of this little-studied piece of the conservation puzzle (CBSG. 1991).

Difficult guidelines, prohibitive regulations

The difficulty of generalized prescriptions for disease concerns is further evident in five thoughtful recommendations by Derrickson and Snyder (1992) developed for captive parrot breeding but based upon the four imperfect propositions above:

(i) Facilities should, whenever possible, be located in the species’ natural range. (ii) The species should be housed in two or three geographically isolated, single-species

facilities. (iii) The facilities should be staffed by individuals who are not simultaneously caring for

other captive birds. (iv) As much as possible, facilities should be in areas free from arthropod vectors and

feral populations of exotic birds. (v) Established husbandry protocols should emphasize disease prevention.

Unfortunately, most species at risk are in less developed countries which may not be able to provide the described facilities, expert personnel and desired protocols within a species’ natural range (or are unwilling to do so). Poor caging and care within a natural range is no substitute for proper housing elsewhere. Although disease is not always a respecter of taxonomy, the provisions set forth above which call for local venues, mostly with psittacines in mind, are probably not appropriate for most reptiles, amphibians, and fish which often require high technology climate-controlled captive habitats. An attempt to visualize the facilities and expertise required for a broad spectrum of species will produce many other exceptions - not the least of which is local political instability.

Use of local conservation funds for such efforts could compete with habitat preservation whereas propagation abroad probably would not. Charitable entities will rarely be able to sustain expert personnel in most such areas over the long term except, perhaps. in IM


programmes, and such efforts should be a part of interactive management. Where that can be done, such projects can become highly important assets in arousing local interest in conservation (Durrell and Mallinson, 1987). However, the main pre-requisite for such propagation facilities is that they be the best and safest possible whatever their location.

In this connection, the use of the term in situ by Wilson et al. (1994) and by Durrell and Mallinson (1987) to describe captive breeding facilities located in a species’ home range is misleading. A cage does not maintain its denizens in place, their natural habitat, i.e. ‘in situ’, whether or not within their range. Disease exposures are unlikely to be the same as those in nature and even photoperiod and micro-climate .may be compromised.

Although conservation biologists tend to think of disease risks in terms of wild species, the world’s more than ten billion domestic animals constitute a vast and growing source of potential infection and disease evolution, as well as competition to wildlife and destruction of their habitats. Their significance may outweigh all other novel sources of infectious disease for threatened species (Woodford and Kock, 1991).

Today, veterinary rules controlling the movement of animals and designed to prevent the spread of disease to or in domestic livestock have become highly restrictive. Acquisition of wild animals, reintroduction and interactive metapopulation management may be precluded by the need to transfer animals across political-veterinary boundaries, unless special dispensations can be obtained for endangered species.

Are captive populations genetically unrepresentative of their founders and therefore unsuitable for reintroduction?

Several species now found only in zoos are descended from extraordinarily small founder groups, for example the Pere David deer (Elaphurus davidianus) from three founders over 75 years ago (M. Jones, personal communication). Historically, when wild animals were long-sustained in captivity, some percentage of original heterozygosity was liable to be lost through deliberate or inadvertent selection, especially with species that entered the pet or private fancier trade, such as psittacines and pheasants. Russell Lande has modelled such situations as well as many others affecting small populations (Lande, 1992). Zoos once paid little attention to genetics and sometimes little to subspecific identification. Consequently, critics have disparaged their ability to manage the genetics of species propagation for conservation. In fact, the terms ‘genetics ‘, ‘selection’, and ‘inbreeding’ do not even appear in the index of the basic zoo mammal management reference of 1964 (Crandall, 1964).

Today, the question is not solely whether captive populations are genetically unrepresentative of their founding populations, for any small subset of a large population, wild as well as captive, has lost rare alleles. The worry is whether a small captive population is suffering a crippling loss of genetic attributes important to a life at liberty in a particular habitat. Thus, application of the term ‘domestication’ to such unidentified losses of heterozygosity over a few years of captivity of long-generation-time wild animals in captivity is more editorial than informational (Derrickson and Snyder, 1992). Genes, like species, are not created equal and the significance of genetic changes over short periods in zoos, relative to the ability of a species to survive in a ‘nature’ which itself has changed, must be treated case by case.

A specific concern is that local animal populations may contain coadapted gene complexes essential to their survival that could be lost in managed populations or attenuated by population supplementations (Templeton, 1986). Such a worry is inconsistent with the expectation that those future habitats where IM is practiced will be

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greatly changed as well as reduced from their original condition, or that reintroduced populations will change further upon reintroduction. After all, IM is an intensive approach to sustaining species which have precariously small populations living mostly in areas which have been damaged by humans. We anticipate that these animals must live in an altered state where historically coadapted gene complexes may be of reduced relevance and where hostile effects are dealt with by management.

Grieg (1979) has documented several instances where translocations ran foul of seeming genetic adaptations to photoperiod and/or climate which affected breeding season and hardiness. Such effects may be a foretaste of problems global warming will pose for conservation. However, the recommendation that we sustain the full panoply of subspecies and ecotypes (if we knew how to recognize them) (Grieg, 1979) is less realistic than Templeton’s suggestion (1986) that our goal should be ‘preserving evolutionary lineages’. not a current ‘constellation of present day traits that defines a rigid category we call species’. The only realistic ways of sustaining many wildlife communities under the dramatic changes expected will be more intensive levels of care. The observation that the subdivided populations of a metapopulation retain variation across the subpopulations better than a panmictic population (Lacy, 1987; Lande, 1992) may be consistent with a zoo-wild interactive model.

Collaborative interzoo management of certain animals as metapopuIations, sensitive to the effects of genetic drift, inbreeding, demography and selection, began to develop in the 1970s (Conway, 1974). a process hastened by the advisory efforts of US Seal, the work of T. Foose for AAZPA and by IS. Ralls and her collaborators (e.g. Ralls and Ballou, 1983). The American Association of Zoological Parks and Aquariums (AZA) published its Species Survival Plan (SSP) just ten years ago (Conway et al., 1984) the first broad scale attempt at collaborative genetic and demographic management of wild animals in the zoo world. Thus, to generalize from the genetic mismanagement of early years about present zoo capabilities, is to misunderstand the data’s history.

Captive breeding of small populations cannot be expected to increase a species’ genetic fitness for reintroduction, but worry that it leads to loss of needed heterozygosity and of fitness for living in nature more rapidly than would occur with a similar-sized population in a fragment of nature, ignores the fact that closely managed captive populations can equalize family lines and sex ratio with a faithfulness impossible in nature. The result is preservation of greater genetic variability at much lower population numbers than wild populations of the same species (Frankel and Soul& 1981). Well-managed interactive populations, because of increased size and the selective pressures upon the majority subpopulations in nature, might be able to do better than either alone (see Lande, 1988, 1993). Nevertheless, IM might also slow if not halt adaptive evolution; a problem on a time scale of secondary importance to endangered species. especially K-selected species. considering the rapid rate of current habitat destruction.

It must be remembered that IM seeks to deal with very small capped populations in fragmentary habitats under some degree of management. Indeed, as we look for chinks in conservation methodologies, we should be mindful that even the selective effects of zoo propagation are likely to happen on a time scale of little immediacy to seriously endangered long-generation time, K-selected species, and that we have little evidence about whether specific captive populations are either genetically unrepresentative of their founders or genetically unsuitable for living at liberty.

Ecology and ethology provide far higher hurdles than genetics for the re-establishment


of captive-bred animals in an altered fragment of former habitat. However, if IM is employed as intended, a strategy used when preservation of habitat or key species in nature without such efforts is not possible or, possibly, to address compelling research, conservation support or education needs, some of these worries become moot.

Utility of interactive wild-captive species management

If interpopulational management of the disjunct fragments of a population is unavoidable when its numbers are precariously small, what is ‘precariously small’? SoulC (1987) states the obvious in noting that there can be no magic number. Each is species, site and time specific - and we rarely have accurate estimates of wild animal numbers. If a population’s numbers are judged perilously low, interactive zoo-wild metapopulation management (as a subset of translocation and reintroduction technology and their guidelines; see Griffith et al., 1989; Beck et al., 1994) would most often be brought into play when:

(i) Risks of inaction exceed risks of intervention to deal with genetic, demographic or other limiting factors.

(ii) Translocation between wild populations is unavailable or inadvisable. (iii) Available habitat is limited, might be made more secure or useful by IM and the

population expansion of the target species is expected to be capped below minimum viable numbers.

(iv) The species is qualified for IM by reason of its suitability for transfer and intensive management, captive propagation, and availability of appropriate captive and wild habitat.

(v) Political winds are favourable and a qualified body of conservationists is financially prepared to make a long-term commitment to species and habitat preservation.

Interactive wild-zoo metapopulation strategies seek to emphasize habitat preservation by: (i) Increased metapopulation size and security for small capped populations in the absence of less intensive options and preservation of part of a species’ population away from extinction’s ‘Evil Quartet’: overkill, habitat destruction and fragmentation, impact of introduced species, and chains of extinction (Diamond, 1989). (ii) Preservation of habitat and supplementation of key species in otherwise undersized habitat patches for subpopulations too small to be viable except as a part of an interactive wild-captive metapopulation. (iii) The creation of public awareness and support for endangered species and habitats, establishment of programmes in habitat preservation and long-term institutional commitments to specific habitat preservation efforts, technology transfer, training and (in some instances) conservation support relationships with developing countries or communities. (iv) Enhanced basic and applied conservation research.

Is it more expensive to preserve animals in captivity?

Common sense suggests that the more intensive a species preservation strategy, the greater its management costs, so too with IM. Thus zoo animal care costs as calculated by Conway (1986) are often used to illustrate that ex situ care is far more expensive than preservation of a species in nature. This is usually a comparison of apples and oranges where zoos and exotic wildlife are concerned.

Preservation’s costs are likely to be seen differently by politicians and conservationists.

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The latter have based their preservationist arguments on the dollars it costs to care for endangered animals in zoos (e.g. Leader-Williams, 1990; Derrickson and Snyder, 1992) without much regard as to where those dollars come from or how comparative costs are measured. They have tacitly assumed that the community or government in the home range of species X concurs with them in placing a high value on its preservation. But the former may judge the value of local species differently and consider that the costs. monetary and political, of restraining human destruction of wildlife and use of its habitats are impossibly greater.

In contrast, from a zoo’s point of view, ex situ care and propagation must proceed to assure the continuance of exhibits, and are funded for the most part from monies that would otherwise be used for other kinds of education or recreation - with the important exception of local species propagation. Thus animal maintenance in foreign zoos usually has very little conservation cost, competing for no conservation pocketbook. Whatever its value, the long-term propagation of an endangered species from a less-developed country in a foreign zoo moves the expense of its upkeep into the framework of a distant economy. It utilizes resources, from municipal tax dollars to gate admissions, that would not otherwise be applied to the preservation of an exotic species or its far-off habitat. However, if this propagation effort, local or foreign, provides an excuse for authorities to avoid protecting habitat, or the species in nature, it is unbearably expensive.

Unfortunately. to the community living on the edge of a tiger’s range, hunting a green turtle population, or competing for freedom of resource-use and life-style with elephants or warblers, the investments of restraint and care required for preservation may exceed any perceived return of local consequence. Nevertheless, those who argue that it is too costly to preserve wild animals by captive propagation are right - because artificial propagation of nature is beside the point. But it is not cheaper to preserve animals in nature, be they pink pigeons, Siberian tigers, American bison Arabian oryx or golden lion tamarins, if it is not really going to be done.

Evolution of zoos as proactive habitat conservation centres

As the structure of IM suggests, the evolution of zoos as conservation centres may have the potential to draw upon new sources of support unavailable to other conservation efforts. Their locations, primarily in urban population centres, position zoos to affect public opinion and even to deal with central government decision-makers. Many can provide experts broadly experienced in the techniques of close-up animal care and management who can assist in monitoring and managing small wild animal populations.

Direct habitat preservation, whether or not associated with interactive animal management, is becoming an ever more powerful element in the thinking of zoo biologists. AZA responded to the proposal that it create an In Situ Conservation Committee (now. Field Conservation Committee) pursuant to a paper proposing it (Conway, 1995) within three months. Conservation partnerships between zoos and national parks and reserves are a growing phenomenon (Durrell and Mallinson, 1987; Mallinson, 1991; Tilson, 1991). emphasizing research and sustaining relationships (Conway, 1989a. 1991,1995). The 1991 AAZPA (AZA) Annual Report on Conservation and Science lists 390 conservation projects being supported in 63 countries by accredited North American zoos and aquariums (Hutchins et al., 1991). The 1992-93 issue reports a near doubling of projects. For example, the Wildlife Conservation Society (founded in 1895 as the New York


Zoological Society) currently conducts more than 240 conservation and conservation science projects in 45 less developed countries. With IM as a stimulus, zoos and aquaria may more rapidly justify a focus on basic habitat preservation to parochial trustees and municipal authorities.

But, endangered species have little market value

Although interactive management must be seen in a world of ever smaller more fragmented habitats, ever more edge effect, ever more alien species, it must also be considered in the light of the largely insignificant market value of threatened species - except for giant pandas. It is interesting, in this connection, that most recent IM programmes have accorded animal ownership to the country of origin, not to the foreign zoo stewards of specimens and financiers of conservation action.

AZA has proposed an interactive giant panda conservation plan to Chinese authorities and preliminary agreements were signed in 1994. The plan seeks to help preserve and expand panda natural habitat rather than to reinforce panda populations in habitat fragments with zoo-bred animals. Panda husbandry is far from being able to offer an interactive metapopulation approach. The programme will be paid for, it is hoped, by zoo revenues from panda exhibits to be provided from China’s existing captive pandas. But this controversial and experimental plan, which will include efforts to improve panda captive propagation, is not a generally useful model. No other species has created such sustained public interest - nor will pandas sustain it if they become more generally exhibited.

If the priority of habitat preservation is clear to zoo-based conservationists, the way to finance it from zoos is not. Most zoos and aquaria were created as local cultural and educational facilities, meant to serve the immediate community which supports them. Few make a profit. How then shall they justify investing in the preservation of distant habitats? What current activities can be set aside to make support available for IM? What new sources of support would IM bring? The fact that an increasing number of institutions have such programmes suggests that there are specialized resources. Yet, outside of the panda effort noted above, most zoos have not yet obtained substantial conservation support from their attendees.

North American attendance at AZA’s 162 accredited zoos and aquaria is nearly 100 million. A $1.00 conservation surcharge would provide a reliable source of almost overhead-free conservation support, but *much time will have to be devoted to the education of local authorities to win approval for such a policy. New York’s Wildlife Conservation Society spent nearly $12 million on field conservation programmes in 1993 but, with basic support at its zoo base in the Bronx, the programme was largely free of overhead. Yet this is the largest programme of its kind and constantly struggling to fund itself. The obvious way, with admissions to view endangered species, has not been successfully developed, yet.

Neither acquisition nor propagation of endangered species pay for themselves. Despite the suspicions of some journalists (Ormrod, 1994) there is little market for endangered species - alive.

Wildlife farming and ranching have parallels with interactive management in their attempts at sustainable use of species with declining populations in nature - but their record has not been encouraging. Four Latin American examples are discussed in Robinson and Redford (1991). Ecotourism is advancing in sophistication and usefulness, and many zoos are enlarging their travel programmes. But such programmes rarely make

sxx Conway

significant profits. It should also be remembered that the vast majority of wildlife exists outside of parks and reserves at the mercy of the towns and communities which share the land with it. Community-based conservation and interactive management plans which bring benefits to local communities are largely unexplored and they may have a special role in wildlife conservation’s future.

Discussion

Interactive zoo-wild metapopulation management, as described here, is a method of sustaining key species in habitat patches too small or damaged to fulfil the needs of a population of viable numbers of key species and which seeks to focus urban zoos on preservation of the habitats of the creatures they exhibit. It is set forth in the expectation that habitat destruction will continue for most of the next century at a rate comparable to that of the past decade, or worse.

IM is a demanding, special case conservation approach. Its support, monitoring, required ecogardening, species weeding and planting, will often be too multi-layered to understand and manage, and too hard to pay for, or too politically challenging, for general use. As with any conservation endeavour. chances of human error and loss increase with complexity.

Effects upon other rare species

Multiplier effects must be anticipated in reintroduction and interpopulational exchanges for many of the species managed in IM may be medium to large size vertebrates. Their ecological impacts upon the environments to which they have been restored may be dramatic, especially where dependent upon a reduced food base. For example, India’s endangered wolves (Cunis 1upi.s pallipes) are now killing its endangered blackbuck antelope (Antelope cervicapra) at the rate of 35-39 blackbuck per annum per wolf (Jhala, 1993). Reserve managers should remember Terborgh’s observations about ‘The big things that run the world . . .’ (1988).

Reduction of diversity in species parks and megazoos

Such intensive conservation action runs the danger of creating Godelian management tangles beyond comprehension. Because of the complexity of multi-species management, we will probably simplify the habitat patches we manage in proportion to the level of our management activity, thereby reducing their biodiversity.

Zoo experience with multi-species exhibits and observation of isolated or fenced parks, such as Kenya’s Nakuru National Park, Abedares N.P., tiny Saiwa Park or Lwambe Valley N.P., Rwanda’s Virunga N.P., Venezuela’s Henri Pittier or San Esteban N.P., suggests that the larger a species’ biomass in relation to its available habitat fragment, the more intensive and zoo-like must be its management in the effort to preserve the reserve’s diversity. Thus patches managed for the maximum number of a particular species. ‘species parks’, could become less diverse, more homogenous and more farm-like.

That said, interactive species-area management models will vary with the biology of the species of primary concern and the ecological and political framework within which they live. But the larger the proportion of a species population that the management programme is able to keep in nature. the more responsive (declining population paradigm), less expensive and less intensive (small population paradigm) we can expect


management to be, if the fragment of ecosystem can function at all (Western and Gichohi, 1993).

Species priorities

Species that create conditions which help to sustain greater or specialized biodiversity in some habitats (‘keystone species’), deserve particular consideration for IM, but there are many other special cases. ‘Flagship species’ upon which public sympathy may fasten, and large animals which by their nature have always been comparatively rare, are sure to become priorities in IM. Simon Stuart and several IUCN SSC Specialist Groups have made a start at such lists for reintroductions (Stuart, 1991) while CBSG’s Ulysses Seal has made a global effort to focus both zoos and other conservation groups on this problem. However, the occasional argument that a species should not be bred unless there is a chance for its reintroduction requires a prescience of conservationists not evident elsewhere in the conduct of human affairs. Much of the task is to preserve options and there is, as yet, no consensus.

‘Scientifically managed captive populations that can interact genetically and demographically with wild populations’ are a stated objective of the Conservation Assessment and Management Plan (CAMP) process of the IUCN-Species Survival Commission’s Captive Breeding Specialist Group (Ellis-Joseph and Seal, 1992; Foose and Seal, 1992). The CBSG workshop process, initiated in 1991, has focused attention on species priorities. Bringing together a broad spectrum of local authorities, field and zoo biologists and based in part upon Population Viability Analysis and/or Population and Habitat Viability Analysis approaches (Gilpin and Soul& 1986) and using the impressive VORTEX computer program (Lacy and Kreeger, 1992), CBSG Workshops have often concluded that a species or its habitat is not likely to long survive current habitat destruction and hunting pressure and recommended the development of back-up captive populations at some level.

The process has been remarkably successful in unearthing previously unpublished information, inspiring concern on the part of government authorities and creating new relationships between conservation experts and decision-makers. But it has also unearthed critics who have questioned data emanating from some of its workshops and, especially, poorly delimited recommendations for the captive propagation of too many declining species. The recommendations have too often been unrealistic with regard to propagation potential and, more important, not responsive to the plight of the species, and should not be confused with IM.

However, the CAMP process is an exceptionally open one, deliberately evolutionary. Its proven heuristic value and constant refinement and expansion by CBSG have made it one of the most imaginative and productive organizing forces for species conservation today. Each IM programme, as defined here, would profit from the scrutiny of the CBSG workshop approach, as well as conventional feasibility studies.

Collaboration and coordination with wildlife authorities

The inherent complexity of collaborative projects, of communication, between different organizations or people of different cultures can deter interactive conservation of any kind. When US scientists Joel Berger and Carol Cunningham, studying black rhinos in Namibia, suggested that the government’s policy of dehorning might be counter- productive as a conservation strategy (Berger and Cunningham, 1994) permission to

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further their studies was promptly withdrawn. However, George Schaller continues to be invited by Chinese authorities to participate in their conservation and science programmes despite publishing a book critical of China’s giant panda conservation efforts (Schaller. 1993).

That so many international conservation programmes are in place, and their numbers multiplying each year, suggests that the limits of tolerance and goodwill are surprisingly broad and not a barricade to most interactive management efforts. AZA is developing preliminary guidelines for such programmes but the most significant element in zoo-wild interactive conservation strategies is the zoo-wildlife authority connection.

The Wildlife Conservation Society, now conducting field conservation in 46 nations, has found that the most fruitful relationships are the product of long-term commitment and local involvement. The golden lion tamarin, black-footed ferret, and Arabian oryx projects are IM examples and their approach may be likened to that of field conservation. The majority of WCS field projects begin as scientific studies of important ecosystems, wildlife communities or key species. Knowledge of local conditions, authorities and community attitudes are built-up over time. Expertise is accumulated and made available to local decision-makers, Local people are added to the project, training is provided and, sometimes, advanced education supported. The development of local conservation NGOs is aided and also that of the wildlife authorities themselves. Today, more than 40% of all WCS overseas conservation projects are run by local scientists on WCS’s staff; two hold endowed chairs with the Society.

Conclusion

Interactive zoo-wild management, as discussed here, is aimed at the preservation of wildlife habitats and communities, not intensive propagation of species one by one. Its applications, like the fragments of nature it seeks to sustain, may be few and far between but their contribution to the conservation of biodiversity would be immensely larger than comparable efforts in captive propagation. IM seeks to take advantage of intrinsic zoo concerns, making habitat preservation the highest zoo conservation priority.

In the darkness of excessive human increase and habitat destruction, the survival of much wildlife will depend upon the utility of managed ecosystem fragments as sanctuaries of biodiversity. To sustain significant representations of the diversity of such areas, composed of relatively small populations whose habitats are reduced or altered and whose numbers are capped, the care of both habitats and wildlife will have to be sophisticated, sometimes even zoo-like. Key species, large or highly specialized animals and plants, rare per unit area, will require particular effort.

Interactive wild-zoo endangered species management is a conservation strategy for a changed environment. It is a demanding option for sustaining precariously small wildlife populations living mostly in areas which have been damaged by human action. It anticipates that these animals and plants must survive in an altered state where hostile effects and deficits are dealt with by management, as minimal a management as possible, for managers are human. But it also anticipates that there will be more and more such ‘altered states’. Managing ‘key species’ subpopulations in an integrated fashion, it uses their study in nature as a way to understand wildlife habitats, ecological processes and conservation threats. And it brings zoo biologists and zoo support to the preservation of habitat.


Preoccupation with wildlife, captive propagation skills and urban locations pre-adapt zoos to become more effective and involved headquarters for nature preservation. Interactive management projects could hasten their evolution into proactive conservation organizations. Ultimately, any endangered species in a zoo should be part of an interactive management programme, if not genetic then of other forms of support.

Zoos deal with wildlife in humanity’s closest, most constant and most interdependent relationship with wild animals. They have compelling and profound reasons to provide not only local educational and recreational services but also expertise, advocacy, support, training and conservation science to the care of the habitats from which their exhibits derive. It is preservation of the source.

Acknowledgements

I especially thank John Robinson, Fred Koontz, Richard Lattis, Bob Cook, George Amato, and Dan Wharton who read an early draft of this paper and made numerous suggestions which greatly improved it. I thank Ben Beck, Don Bruning, Claudio Campagna, Jim Doherty, Helen Gichohi, Guillermo Harris, Michael Hutchins, Ullas Karanth, and David Western for helpful discussions and information, and Jeremy Mallinson who suggested this paper and contributed useful information.

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