The Wild Dog—Africa's Vanishing Carnivore

ORYX VOL 25 NO 3 JULY 1991 The wild dog—Africa's vanishing carnivore

John H. Fanshawe, Lory H. Frame and Joshua R. Ginsberg

This paper presents a synopsis of the current status and distribution of the African wild dog Lycaon pictus, outlines reasons for its decline and discusses recommendations to halt or reverse this decline. A recent review of the status of the species provides evidence that it has disappeared or is in decline throughout its range (sub-Saharan Africa). Relict populations with little or no chance of long-term survival are found in several countries including Algeria and Senegal. Countries believed to contain potentially viable populations are, from north to south, Kenya, Tanzania, Zambia, Zimbabwe, Botswana, and South Africa (only the Kruger National Park).

Background and biology Hwange National Park (Childes, 1988) and South Africa's Kruger (Reich, 1978). Group There are probably fewer adult African wild size is quite variable and, depending on pack dogs Lycaon pictus in protected areas than fecundity and pup survival, as many as 45 black rhinos Diceros bicornis remaining in dogs are occasionally recorded. Litter size Africa (Frame and Fanshawe, in press; averages 10 pups. Denning, which lasts 2 or 3 Cumming et al., 1990). The reduction in the months, is the only time in the year when distribution and abundance of Lycaon pictus, a these nomadic animals stay in one place. Birth monotypic canid restricted to sub-Saharan sex ratio is skewed towards males in both cap- Africa, must receive greater attention, study, tive and wild populations (Frame et al., 1979; and finances if extinction of the species is to be Malcolm and Marten, 1982); in the adult pop- avoided. ulation of the Serengeti, males outnumber It is unlikely that there are more than 5000 females by a ratio of 2:1 (Frame et al, 1979; individuals in all of sub-Saharan Africa. Many Malcolm, 1979). of these individuals are members of widely Lycaon is a truly nomadic animal. Pack scattered packs in unprotected, prey-depleted home range size varies considerably, from 500 areas, and are certainly doomed; the estimate sq km at its smallest (Reich, 1981) up to 1500 is thus reduced to 3000. As 90 per cent of all sq km in both plains (Serengeti: Frame et al., pups are likely to die before reaching maturity 1979) and bush habitats (Hwange: Ginsberg, (Schaller, 1972), the remaining population of unpubl. data). Ranges overlap from 50 to 80 adult African wild dogs may be as low as 2000 per cent. When they are not at the den with (Frame and Fanshawe, in press). young pups, packs rarely sleep in the same Wild dogs are social, communally hunting place on consecutive nights. carnivores, which live in small cohesive packs Immense home ranges, low densities and its typically composed of a dominant breeding extreme mobility engender grave conservation pair, a number of non-breeding adults, and problems for Lycaon: few protected areas are their dependent offspring. Adult males are large enough for populations of 200-300 indi- related to one another, but not to the females, viduals (East, 1981a); fragmentation and which immigrate from other packs. Group size expansion of human populations lead the varies locally, with a mean of 9.8 in the dogs into increased contact with humans, their Serengeti (Frame et al., 1979; Malcolm and domestic animals (Butynski, 1974), and the Marten, 1982), 8.4 in the Zambesi Valley diseases they carry (Dobson and Hudson, (Childes, 1988), and 11 in both Zimbabwe's 1987); an increasing number of roads bisecting 137

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• Countries populations>l B Countries w/ populations 5O-IOO B Countries w/frogmented small populations 0 Extirpated

Figure la. Distribution of African wild dogs 1980 Figure lb. Distribution of African wild dogs 1988 (after Smithers, 1983). (data from Frame and Fanshawe, in press).

their habitat threaten the species with greater nine countries. Potentially viable populations mortality from four-wheeled predators (Mech, persist in only six, and these populations are 1989). relatively small and often declining (Figure 1). Even in those countries in which wild dogs exist in relatively large numbers, such as Current status and distribution Zimbabwe, populations are fragmented and concentrated in protected areas (Childes, 1988) Early accounts support a hypothesis that wild and rarely are seen outside of these regions. dogs were once widespread where they have now been eradicated (Frame and Fanshawe, in press), both in protected areas (e.g. Nigeria's Causes of decline Yankari Game Reserve: Sikes, 1964) and entire countries (e.g. Uganda: Frame and Fanshawe, A number of factors are believed to have con- in press). South of the Sahara, it seems likely tributed to the decline of wild dogs: habitat that wild dogs were once found in every habi- loss; conflict with man (Malcolm, 1979; tat except for rain forest and some desert Childes, 1988); disease (Schaller, 1972; P. Kat, in areas. There are records for habitats as unlike- litt.); competition with other predators (Frame, ly as the mountain snows of Kilimanjaro 1986); and loss to vehicle accidents (Fanshawe, (Thesiger, 1970). The species's range must 1989). All these factors are interrelated, each have pushed far into arid lands to the north, being more or less important depending upon because a relict population still exists in the population being considered. Algeria (K. de Smet, pers. comm.) Frame and Fanshawe (in press) give a coun- Habitat loss try-by-country summary of the status and population numbers of the wild dog. African In common with large mammal species wild dogs once occurred in 34 sub-Saharan throughout the continent, wild dogs are facing African countries (Smithers, 1983. The species annual losses of habitat. Because of their large is now extirpated, or nearly so, in 19 of these. home ranges and low population densities, Small or relict populations (n<100) persist in the dogs are more susceptible to fragmenta- 138

tion of continuous habitat than any other dog, yellow-eyed, mud-colored, thick-footed, African large mammals except, perhaps, the head on its forepaws—an eerie embodiment of cheetah Acinonyx jubatus (East, 1981b). the hound of hell' (Alexander, 1986). With the exception of the efforts focused on In contrast, these authors treat lions rather animals such as the African elephant benignly: Loxodonta africana and black rhino, conserva- 'In appearance the lion of the Zambesi val- tion efforts are shifting from saving single ley is a splendid and most majestic creature' species to projects aimed at the integration of (Maugham, 1914). people and wildlife, and the protection of 'I saw him—a big male lion with black- unique habitats. Species like wild dogs and tipped mane and tail, a kingly creature of cheetahs, however, can be viewed as 'flag- habit on his morning stroll, cleaving the tawny ships'. As East (1981b) notes, 'reserves large savanna' (Alexander, 1986). enough to support minimum viable popula- Lions are kingly, majestic: wild dogs savage, tions of cheetah and wild dogs should ensure murderous. Lions cleave the savanna, wild the long-term survival of entire large mammal dogs wallow sullenly. Yet lions kill cattle and communities'. people, they commit infanticide, and scavenge Fragmentation of habitat leads to numerous food from other predators, a behaviour rarely direct and indirect effects. Isolation of sub- exhibited by wild dogs. Although the lan- populations will result in smaller effective guage of popular writing may only reflect population sizes and greater potential for public perception, it also influences and rein- extinction (papers in Soule, 1987); encroach- forces the attitudes of the public. If wild dogs ment and settlement leads to more contact are to survive, it is imperative that the entirely with man and his domestic animals. This has false perception of the animal as a wanton three effects: increased persecution by farmers killer be replaced by a more accurate and, per- in 'defense' of domestic stock and wildlife; haps, more sympathetic view. increased contact with domestic dogs and their diseases; fragmented habitats are usually Government extermination programmes. As traversed by a greater number of better roads recently as the late 1960s, wild dogs existed in than pristine areas, bringing the dogs into many protected areas where their presence greater contact with faster, more numerous, was often met with persistent antagonism. and dangerous traffic (see below). Managers working both inside and outside these areas treated them as vermin (Davison, 1930; Childes, 1988). Persecution Although there are few data documenting Public perception. Arguably, disembowelling the extent of predator control projects, one's dinner is not the prettiest way of killing, Zimbabwe provides a good example of the but it can hardly be called unnatural. Despite pressures Lycaon faced at the height of its per- the work of Goodall and van Lawick in the secution in that country. Before 1975, the num- Serengeti (1970, 1973), wild dogs rarely fare ber of wild dogs is unknown; however, in well in the public eye when compared with 1975, only 500 individuals remained other carnivores such as the lion. This jaun- (Cummings, cited in Childes, 1988). A mini- diced public perception of Lycaon has persist- mum of 3404 were killed from 1956-1975, ed throughout this century: including those shot in Hwange National 'Let us consider for a moment that abomina- Park, with fewer being shot in successive tion—that blot upon the many interesting years, probably as a result of population wild things—the murderous Wild Dog. It will declines (Childes, 1988). be an excellent day for African game and its Fortunately, attitudes among wildlife man- preservation when means can be devised for agers have changed, as witnessed by the writ- its complete extermination' (Maugham, 1914). ings of former Director and Chief Warden of '(I)n the shade of the tailgate lay one sullen Uganda National Parks, Rennie Bere: 139

'Wild dogs hunt in packs, killing wantonly arundinum and waterbuck Kobus ellipsiprymnus far more than they need for food, and by (J. R. Ginsberg, unpubl. data). methods of the utmost cruelty: Lycaon does not kill quickly as the lion but often starts to Sport hunting and totems. It appears that sport devour the antelope ... before life is extinct' hunting of wild dogs has never been popular. (Bere, 1956). Although clients could shoot them as trophies 'Wild dogs are very successful and efficient in Tanzania until 1987 (when their hunting hunters and, in spite of their reputation, are no was banned; Fanshawe, 1989), few did so. more 'cruel' than any other large beasts of Trophy fees were low so professional hunters prey' (Bere, 1975). discouraged their shooting. In addition, skin- Sadly, private landowners whose properties ners disliked handling the carcasses of wild border protected areas have not changed their dogs. view of Lycaon. Many farmers still treat wild dogs as vermin, with reports of private individuals shooting them being disconcertingly Disease common (Childes, 1988; Ginsberg and Lycaon appears particularly sensitive to dis- Macdonald, 1990; Frame and Fanshawe, in ease. Under some circumstances, pathogens press). may be a factor regulating wild dog numbers (Schaller, 1972; Dobson and Hudson, 1986). A Predators of domestic stock and privately owned variety of diseases has been isolated, includ- wildlife. Given the opportunity, there is no ing rabies, distemper, parvo-virus, and doubt that wild dogs will occasionally kill and anthrax, Babesia (van Heerden, 1980) and eat domestic stock. However, little evidence canine ehrlichiosis (van Heerden, 1979). In all has been found for depredation of Masai cat- cases of infection, it appears that wild dogs are tle, sheep and goats by wild dogs in Kenya the victims, not vectors, of disease transmis- (Njungiri, 1990; P. Kat, pers. comm.). With an sion (van Heerden, 1979, 1980). In recent estimated 350 dogs in Zimbabwe, cattle losses years, an increasingly determined effort has are negligible (Childes, 1988). In most species been made by researchers in Kenya, Tanzania, in which studies have been made, losses to Zimbabwe and South Africa to isolate the dis- canid predators are usually exaggerated eases infecting wild dogs. (Ginsberg and Macdonald, 1990). Rabies illustrates how dramatically disease In areas such as north-west Zimbabwe habi- can affect these social carnivores. In 1989, 20 of tat encroachment has been reversed by a shift 22 dogs in a single Masai Mara pack died of from a cattle-based economy to one based on rabies (P. Kat, pers. comm.). An outbreak of wildlife (hunting/ranching/photo safaris). This rabies in domestic dogs living to the north- provides distinct advantages by increasing west of the reserve occurred simultaneously areas of natural habitat, improving prey avail- with this event. Although no rabies could be ability outside protected areas, and providing isolated in a pack that died in the Serengeti in corridors among protected areas. Surprisingly, 1986 (J. Nyange, in litt.), the dogs showed wildlife ranchers are even less tolerant of wild symptoms very similar to those of the Kenyan dogs than cattle ranchers (Childes, 1988). In dogs. Population declines in north-west interviews with ranchers, three complaints Zimbabwe in the early 1980s (Childes, 1988) were repeated: wild dogs take prey that could were correlated with (but not linked causally otherwise be sold for hunting or meat; packs, to) an outbreak of rabies in that area particularly when denning, scare prey into (Kennedy, 1988). thick bush and render hunting difficult or Canine distemper has long been suspected impossible; and even where wild dogs eat rel- to be common in wild dogs (Schaller, 1972), atively inconsequential proportions of the but its isolation from free-ranging animals has total prey available, they appear to preferen- never been successful. In captivity, they tially select rare game, like reedbuck Redunca appear highly susceptible, contracting distem- 140

per when inoculated with live vaccines, and protected areas. For example: the Dar es showing virtually no antibody response to Salaam-Morogoro highway, which bisects dead-virus vaccines (van Heerden et al., 1980). Mikumi National Park in Tanzania (Frame and In a recent outbreak of anthrax in the Fanshawe, in press); the Bulawayo-Victoria Luangwa Valley in Zambia, several packs of Falls road, which runs parallel to Hwange in wild dogs were found to be infected (K. Zimbabwe, has claimed at least 12 dogs since Saigawa, pers. comm.). In one pack of five 1988 (J. R. Ginsberg, unpubl. data). Accidents adults and eight pups, anthrax killed four of also occur in parks without major highways, the adults and three of the pups. In a separate such as the Serengeti, where three dogs (of a pack, at least two adults contracted the dis- known population of nine individuals) were ease. killed in 1986 (Fanshawe, 1989). As Lycaon, like wolves, frequently use roads for hunting and transit (Reich, 1981), growing road net- Competition works will lead inevitably to more deaths; Loss of kills to larger carnivores such as lions only if areas of high road density abut large and hyaenas possibly poses a threat in some areas of roadless wilderness will wild dogs areas. The absence of wild dogs from survive (Mech, 1989). Ironically, it has been Ngorongoro Crater in Tanzania is often such accidents that have led to a better knowl- thought to result from a high density of spot- edge of the wild dog's range in some areas of ted hyaena Crocuta crocuta (Estes and Kenya (Frame and Fanshawe, in press). Goddard, 1967). Competition with hyaenas has been suggested as a variable responsible for limiting Lycaon numbers in the Serengeti Genetics (Frame, 1986). In particular, in the Serengeti, Living at low densities and in dispersed popu- competition with hyaenas during the late dry lations, wild dogs may be prone to low levels season, a time in which food is already limit- of genetic heterozygosity. This may, in turn, ed, may limit population growth of Lycaon. render them more susceptible to various dis- Recent results suggest that wild dogs can eases and parasites (O'Brien et ah, 1985). defend their kills from hyaenas, but not lions Furthermore, eastern and southern subpopu- (Fanshawe and FitzGibbon, unpubl. data). In lations are genetically distinct (R. Wayne, in this study in the Serengeti, hyaenas were pre- litt.). This divergence has both evolutionary sent at 85 per cent of kills, but only gained and conservation implications and is particu- carcasses when the dogs had finished feeding. larly important to projects contemplating rein- In the Masai Mara, hyaenas are more common troduction. For instance, probably all individ- at the kills of smaller packs. Hyaenas were uals in captivity in the USA are of southern present at 92 per cent of all kills made by a African origin (B. Brewer, North American pack of three adults and five pups; 66 per cent Lycaon Studbook Keeper, pers. comm.). Yet of the kills were parasitized, but only after the those areas in which reintroductions are likely dogs had eaten extensively (T. Fuller and P. to occur are in eastern and western Africa. Kat, pers. comm.). In a small sample of 40 kills Particularly in areas where relict populations from Hwange National Park, hyaena were persist, reintroduction of a distinctly different seen to attend only 10 per cent of kills (J. R. subspecies may be inadvisable. Ginsberg, unpubl. data). Once again, carcasses were only yielded to the scavengers when feeding was essentially over. Current conservation status and research

Road kills In a recent review of all Canids (Ginsberg and Throughout their range wild dogs are killed by Macdonald, 1990), the African wild dog ranks vehicles, particularly where major routes cross among the most endangered. The species is 141

classified as Vulnerable by the IUCN World The proceedings of the meeting will elaborate Conservation Union, but reclassification as the conservation options discussed below in Endangered has been recommended greater detail. (Ginsberg and Macdonald, 1990; Frame and Fanshawe, in press). Captive breeding, translocatwn, rehabilitation and Lycaon research has become a growth indus- reintroduction try in recent years. Five field-based projects are under way, two others are planned, and For animals that have complex social systems laboratory research on wild dog genetics is and require learning (hunting, learning exten- also being conducted. Brief details are given sive home range areas, etc.), primary conser- here (from north to south), but they are fully vation efforts should concentrate on the main- discussed in Ginsberg and Macdonald (1990): tenance of adequate habitat for populations (i) Kenya: genetic, ecological and disease mon- that remain in the wild (Imboden, 1989). itoring of a population in the Mara Region (Dr Returning captive-bred dogs to the wild, or Pieter Kat); (ii) Tanzania: continuation of the translocating animals from healthy popula- long-term monitoring in the Serengeti (Dr tions, is often mooted, and has been tried for Markus Borner, Karen Laurenson, Roger and several populations (Frame and Fanshawe, in Jan Burrows and Stephen Lelo); (iii) Tanzania press), and with only some success, notably in (proposed): dog reintroduction to Mkomazi the Umfolozi Reserve, South Africa (Ginsberg Game Reserve (Tony Fitzjohn); (iv) Tanzania and Macdonald, 1990). In fact, pressure is now (funded, to begin 6/91): research into the little increasing to recognize the limitations that known but potentially important population reintroductions can have in the successful in Selous Game Reserve (Scott and Nancy management of mammal populations (Stanley- Creel); (v) Zimbabwe: genetic, disease, ecolog- Price, 1991). ical, and behavioural research, especially in If reintroduction is to be attempted, many terms of conflicts between people and wild issues need to be resolved. No wild popula- dogs (Dr Joshua Ginsberg, Clare Davies); (vi) tion numbers more than 200 or 300 adults Botswana: assessment of wild dogs through- (Frame and Fanshawe, in press); hence, no out, but concentrating in the northern Chobe wild population appears to be large enough to and Ngamiland Districts (Dr John Bulger and enable translocation of more than several Professor Bill Hamilton); (vii) South Africa: packs. Given this constraint, the only obvious long-term monitoring in Kruger National Park source is captive-bred dogs. The greatest need (Dr Gus Mills and Anthony Maddock); (viii) for reintroduction is in western, central, and Namibia: reintroduction of dogs into Etosha National Park (Philip Stander; J. L. Scheepers); eastern Africa, yet most captive dogs are and, (ix) genetic analyses are being carried out southern in origin. Because there are signifi- on blood specimens collected throughout cant genetic differences between southern and Africa (Dr Robert Wayne). eastern populations (R. Wayne, in litt.), these dogs should not be used for reintroduction. Additional areas of concern that must be addressed by any well-planned reintroduction Conservation options include: the risk of introducing disease and parasites; the potential for conflict with neigh- In late 1991 a Population Viability Assessment bouring human populations; the uncertainty Workshop will be held to discuss the problems surrounding the ability of reintroduced packs facing the conservation of the wild dog. The to learn hunting techniques; and the need to meeting will be attended by representatives carefully assess sites for their suitability to from each of the range states that support support a population over a long period. potentially viable wild dog populations, scien- Wild dogs are frequently bred in captivity tists studying wild dogs, and experts con- but there is great variance in the success of cerned with disease and genetics of wild dogs. breeding attempts. The reasons for this large 142

variation in breeding success among zoos are lands, such programmes could, potentially, poorly understood (Ginsberg and Macdonald, reduce the rate and frequency of infection. As 1990) but are being investigated (J. R. packs appear to succumb rapidly once a single Ginsberg, unpubl. data). Although captive dog is infected, removal might be considered. populations may be self-sustaining numerical- Achieving this before infection spreads is ly (Frankel and Soule, 1981), only a few zoos unlikely. Morever, if a beneficial genetic resis- breed wild dogs with any success, so fewer tance to disease exists in some individuals, and fewer founder animals will be represent- their removal would have negative long-term ed in captive groups. A regional studbook for effects. zoos in North America is near completion (B. Direct inoculation of Lycaon appears to be Brewer, Brookfield Zoo, pers. comm.). This the best option, but data from zoos suggest studbook should clarify the effective popula- that finding vaccines that are efficacious and tion size of captive populations in North safe may be difficult. This is especially true of America. distemper (van Heerden et ah, 1980). Testing of new vaccines, such as recombinant rabies vaccine, must be carried out under controlled Disease control conditions before widespread inoculations are With such an intensely social nature, wild attempted. If promising vaccines are found, dogs are very susceptible to the rapid passage tests should be made in field projects planned of disease within a group. Three management to allow for proper assessment. options are open to combat disease: control pathogens in sympatric domestic dog popula- Legislation tions; remove sick individuals before they infect other pack members; or inoculate the In recent years, wild dogs have been afforded wild dogs directly. at least limited protection in all the countries Throughout much of Africa, the costs and where they occur in relatively large numbers logistics of controlling disease in domestic (n =200-500). Hunting has been banned in dogs would be prohibitive. If, however, efforts Tanzania and Botswana, and near-complete are focused on areas bordering protected protection has been enacted in Zimbabwe. At

African wild dogs during a pre-hunt greeting (/. R. Ginsberg). 143

national level, a legal framework for the pro- Recommendations tection of endangered species is a critical tool towards their conservation. The following recommendations amplify Internationally, placing the wild dog on the those made in Ginsberg and Macdonald list of endangered mammals has been pro- (1990). posed (Ginsberg and Macdonald, 1990; Frame 1. The status of the African wild dog should and Fanshawe, in press). Wild dogs are clearly be changed to Endangered by the IUCN, endangered, and such a reclassification will World Conservation Union. Reclassification aid in soliciting support for conservation will have little effect in stopping illegal hunt- efforts. A second proposal, to list the dogs on ing or vermin control, or in arresting disease. CITES (Convention on Trade in Endangered It will, however, lend support to improving Species) under Appendix II has far less value. legislation and enforcement in range states. This Convention is confined to regulating Furthermore, it will assist efforts to enact pro- trade in endangered species and there is no grammes (education and disease prevention) evidence for even a small international trade that may lead to an improved potential for the in wild dog products. survival of the species. 2. Efforts should be made to ensure adequate support for populations in the target Education countries of Kenya, Tanzania, Zambia, Perhaps the act that would have the greatest Zimbabwe, Botswana and in South Africa's immediate impact on the conservation of wild Kruger National Park. The role of wild dogs in dogs would be to improve their image, both in tourism, and their value as a tourist attraction Africa and internationally. As most popula- in these countries, must be exploited. Funding tions of wild dogs move in and out of protect- for protection and research needs to be pro- ed areas, local people's attitudes and percep- vided, especially if the effects of disease are to tions must be encouraged towards a benign be successfully countered. Large parks are view. Various methods, including talks in vital for Lycaon and parks that are large churches and schools, popular articles pub- enough to protect the species will also serve to lished locally (e.g. Njunjiri, 1990), and meet- conserve the entire faunal and floral commu- ings with local farmers and ranchers, must be nity there. pursued. The tourist value of the dogs should 3. Surveys of the species should be made also be promoted. Visitors to a den in the urgently in areas that are very poorly known, Serengeti spent an average of 38 minutes with but where wild dogs are believed to exist at the pack (J. H. Fanshawe, unpubl. data), com- reasonable densities, e.g. the Selous in south- pared with an average of 10 minutes with ern Tanzania; Luangwa and Kafue National lions (Schaller, 1972). Parks, Zambia. 4. Relict populations must be assessed for long-term survival. No healthy West African Genetics population (>100) is known and surveys need Given that there appear to be large genetic dif- to be undertaken in large reserves where wild ferences among wild dogs occurring in differ- dogs still eke out a precarious existence, e.g. ent parts of Africa, greater research is needed Niokolo-Koba National Park in Senegal on the population genetics of this species. In (Figure lb); the Bouda-Ndiida Reserve particular, samples from populations in west- National Park in Cameroon; Bamingui- ern Africa are needed. In addition, an under- Bangoran National Park in the Central African standing of the intrapopulation genetics of rel- Republic; and Ouadi Rime-Ouadi Achim atively healthy populations of wild dogs Forest Reserve in Tchad. would be of great assistance to those contem- 5. As the great majority of Lycaon in captivi- plating reintroduction in areas where wild ty are of southern origin, consideration should dogs have been extirpated. be given to including dogs from western and 144

eastern Africa in captive stock. The size of 3-20. these captive populations, or in fact the ability East, R. 1981b. Species-area curves and populations to collect individuals for captive breeding, will of large mammals in African savanna reserves. depend on population sizes in the wild. Biol. Conserv. 21,111-126. Estes, R.D. and Goddard, J. 1967. Prey selection and Groups of differing genetic origin should be hunting behavior of the African Wild Dog. /. bred separately, and the Studbook in the USA Wildl. Manage. 31,52-70. should be expanded to include collections Fanshawe, J.H. 1989. Serengeti's Painted Wolves. world-wide. Natural History, 56-67. Frame, G.W. 1986. Carnivore competition and resource use in the Serengeti ecosystem of Acknowledgments Tanzania. PhD Thesis, Utah State University, The authors would like to acknowledge the helpful Logan, Utah. criticisms of Clare FitzGibbon, Ulie Seal, Markus Frame, L.H. and Fanshawe, J.H. In press. African Borner, David Macdonald, Leo Kiinkel, Mark Wild Dog Lycaon pictus: a Survey of Status and Stanley Price, Scott Creel, Todd Fuller, Pieter Kat, Distribution 1985-1988. IUCN/SSC, Canid Jonathan Scott and Chris Thouless. Specialist Group Report, IUCN, Gland, Switzerland. Frame, L.H. and Frame, G.W. 1976. Female African Request Wild Dogs emigrate. Nature, 263,227-229. Frame, L.H., Malcolm, J.R., Frame, G.W. and van Although we are beginning to appreciate the extent La wick, L.H. 1979. The Social Organisation of of Lyceum's decline, the data available for many African Wild Dogs (Lycaon pictus) on the Serengeti countries, in West and Central Africa especially, are Plains, Tanzania, 1967-78. Z. Tierpsychol. 50, few and far between. Any readers who feel they can 225-249. contribute information are welcome to contact Frankel, O.H. and Soule, M.E. 1981. Conservation and Joshua Ginsberg, Deputy Chairman IUCN Canid Evolution. Cambridge University Press, Specialist Group, Wildlife Conservation Research Cambridge. Unit, Department of Zoology, University of Oxford, South Parks Road, Oxford OX1 3PS, UK. Ginsberg, J.R. and Macdonald D.M. 1990. Foxes, Wolves and Jackals: An Action Plan for the Conservation ofCanids. IUCN, Gland, Switzerland. References Goodall, J.M. and van Lawick, H. 1970. Innocent Killers. Collins, London. Alexander, S. 1986. The Serengeti: The Glory of Life. van Heerden, J. 1979. The transmission of canine Nat. Geo. 169,585-601. ehrlichiosis to the Wild Dog Lycaon pictus Bere, R.M. 1956. The African Wild Dog. Oryx, 3, (Temminck) and Black-backed Jackal Canis 180-182. mesomelas Schreber. /. S. Afr. Vet. Ass. 50, 245-248. Bere, R.M. 1975. Mammals of East and Central Africa. van Heerden, J. 1980. The transmission of Babesia Longman, London. canis to the Wild Dog Lycaon pictus (Temminck) Butynski, T.M. 1974. An encounter between African and black-backed jackal Canis mesomelas Schreber. Wild Dog and domestic dog. E. Afr. Wildl. J. 12, /. S. Afr. Vet. Ass. 51,119-120. 243. van Heerden, J. 1981. The role of integumental Childes, S.L. 1985. The past history, present status glands in the social and mating behaviour of the and distribution of the hunting dog Lycaon pictus hunting dog Lycaon pictus (Temminck, 1820). in Zimbabwe. Biol. Conserv. 44, 301-316. Onderstepoort}. Vet. Res. 48,19-21. Cumming, M., du Toit, D.H. and Stuart, S.N. 1990. van Heerden, J., Swart, W.H. and Meltzer, D.G.A. African Elephants and Rhinos: Status Survey and 1980. Serum antibody levels before and after Conservation Action Plan. IUCN, Gland, administration of live canine distemper vaccine to Switzerland. the Wild Dog Lycaon pictus. J. S. Afr. Vet. Ass. 51, Davison, T. 1930. Report on Wankie Game Reserve, 283-284. October 1928-March 1930. Wankie National Park Imboden, C. 1989. Parrots, trade and captive breed- Notes, 21,1978. ing. World Birdwatch, 11,2. Dobson, A.P. and Hudson, P.J. 1986. Parasites, dis- Kennedy, D.J. 1988. An outbreak of rabies in North- ease and the structure of ecological communities. Western Zimbabwe 1980-1983. Vet. Record, 122, TREE, 1,111-15. 129-133. East, R. 1981a. Area requirements and conservation Kruuk, H. and Turner, M. 1967. Comparative notes status of large African mammals. Nyala, 7 (1), on predation by lion, leopard, cheetah and Wild

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Dog in the Serengeti area, East Africa. Mammalia, Sikes, S.K. 1964. A Game Survey of the Yankari 31,1-27. Reserve of Northern Nigeria. Nigerian Field, 29, van Lawick, H. 1973. Solo: The Story of an African 54-82,127-141. Wild Dog. Collins, London. Smithers, R.H.N. 1983. Mammals of the Southern Malcolm, J.R. 1979. Social organisation and commu- African Sub-region. University of Pretoria, nal rearing in African Wild Dogs. PhD Thesis, Republic of South Africa. Harvard University, Cambridge. Soul£, M.E. 1987. Viable Populations for Conservation. Malcolm, J.R. and Marten, K. 1982. Natural selection Cambridge University Press, Cambridge. and the communal rearing of pups in African Schaller, G.B. 1972. The Serengeti Lion. Chicago Wild Dogs (Lycaon pictus). Behav. Ecol. Sociobiol. 10, University Press, Chicago. 1-13. Stanley-Price, M.R. 1991. In press. A review of mam- Malcolm, J.R. and van Lawick, H. 1975. Notes on mal reintroductions, and the role of the Wild Dogs hunting zebras. Mammalia, 39, 231-240. Reintroductions Specialist Group of IUCN/SSC. Maugham, R.C.F. 1914. Wild Game in Zambesia. John Symp. Zool. Soc. {Land). 62. Murray, London. Thesiger, W. 1970. Wild dog at 5894 m (19,340 ft). E. Mech, L.D. 1989. Wolf population survival in areas Aft. Wildl. ]. 8,202. of high road density. Am. Midi. Nat. 121, 387-389. Njungiri, P. 1990. The dogs in danger. The Standard, Nairobi, 4/4/1990, p. 14. John H. Fanshawe, Wildlife Conservation Research O'Brien, S.J., Roelke, M.E., Newman, A., Winkler, Unit, Department of Zoology, University of Oxford, C.A., Meltzer, K.D., Colly, L., Evermann, J.F., South Parks Road, Oxford OX1 3PS, UK. Current Bush, M. and Wildt, D.E. 1985. Genetic basis for address: Box 95, Watamu, Kenya. species vulnerability in the cheetah. Science, 227, Lory H. Frame, s/c ADEFA, B.P. 5570, 1101-1103. Ouagadougou, Burkina Faso. Reich, A. 1978. The behavior and ecology of the Joshua R. Ginsberg, Wildlife Conservation Research African Wild Dog (Lycaon pictus) in the Kruger Unit, Department of Zoology, University of Oxford, National Park. PhD Thesis, Yale University, New South Parks Road, Oxford OX1 3PS, UK. Reprint Haven, Connecticut. requests to Joshua R. Ginsberg.

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