The IUCN/SSC Canid Specialist Group's
African Wild Dog Status Survey and Action Plan (1997)
Later chapters describe the current distribution and status of Africa's remaining wild dog population, and the threats faced by these populations, before recommending measures for their conservation.
There can be no doubt that African wild dogs (Lycaon pictus) have declined over the last century, and this decline has accelerated in the last 30 years. They were once distributed through much of sub-Saharan Africa, apart from rainforest areas and deserts (Fanshawe et al. 1991; Monod 1928; Schaller 1972). Now, however, they have been extirpated from most of their range - they are extinct in most countries in West and Central Africa, and in the East and the South they are confined to a few areas where human population density remains low (Chapter 3). Today, Africa's wild dog population numbers between 3,000 and 5,500. Most populations outside - and sometimes inside - protected areas may still be declining. Wild dogs are rare compared with other high-profile species in Africa: there are about the same number of wild dogs as there are black rhinos (Diceros bicornis, ~3,000 remaining, Cumming et al. 1990), fewer wild dogs than cheetahs (Acinonyx jubatus, 9-12,000 remaining, Nowell & Jackson 1996), and far fewer wild dogs than African elephants (Loxodonta africana, 100-130,000 remaining, Said et al. 1995).
The ultimate cause of wild dogs' decline has been a combination of persecution and habitat loss. Like other large predators, wild dogs do kill livestock under some circumstances, and have therefore been shot, snared and poisoned in most livestock areas (Chapter3). Worse still, they have been persecuted in the name of animal welfare and conservation. Wild dogs kill their prey by tearing it to pieces or disembowelling it (Kuhme 1965), and this earned them a reputation as cruel and bloodthirsty killers. Game managers' attitudes to them are exemplified by Bere's (1955) observation that they
"...hunt in packs, killing wantonly far more than they need for food, and by methods of the utmost cruelty... When the Uganda national parks were established it was considered necessary, as it had often been elsewhere, to shoot wild dogs in order to give the antelope opportunity to develop their optimum numbers. Fortunately only a few of these creatures have had to be destroyed and their number in the parks does not seem to be particularly large...".
This last remark of Bere's points to a crucial aspect of wild dog ecology: they always live at very low densities, and are rare even where they live in large well-protected habitats with abundant prey (Chapter 4). This makes them unusually susceptible to habitat fragmentation. Growing human populations have caused wild dog habitat to become discontinuous, as large tracts of land have been taken over for livestock grazing and cultivation. As more people have colonized the land, wild dogs have been persecuted and their prey have been depleted. Wild dog populations have, therefore, become increasingly isolated in fragments of habitat with few human inhabitants. Since wild dogs live at such low densities, even the largest of these fragments could support only small populations, which are vulnerable to extinction (Soulé 1987). Worse still, wild dogs were persecuted inside national parks and game reserves, which represented some of the best remaining habitat. This combination of habitat fragmentation, persecution and prey loss explains wild dogs' dramatic decline across most of Africa. As a result of this process, today wild dogs persist only in countries with relatively low human population densities (Chapter 3).
Although wild dog numbers have declined markedly, it is not too late to prevent their extinction. Viable populations remain in several countries in East and southern Africa and, with adequate protection, there is no reason why these populations should not persist. However, to conserve wild dogs we must understand the factors that have led their numbers to fall across Africa in the past, and determine the threats that might cause further decline or extinction in the future. If we can use this knowledge to halt wild dogs' decline, then we can prevent their extinction without the "emergency" measures that have been necessary for some other endangered carnivores (Caughley 1994; Clark 1994; May 1986; Phillips 1995).
Given wild dogs' current circumstances, this Action Plan has the following aims:
The Action Plan is structured to meet these aims. The remainder of this chapter concerns aspects of wild dogs' natural history that are crucial for understanding the threats they face, and the management options that are possible. Chapter 2 deals with genetic factors important in wild dog conservation, especially their taxonomy and the identification of sub-species. Chapter 3 describes the current status and distribution of wild dog populations across Africa. Chapter 4 outlines the threats faced by wild dogs, and Chapter 5 uses demographic modelling to assess the probability that any of these threats might contribute to the extinction of remaining populations. Chapter 6 draws upon this information to propose measures for the conservation of free-ranging wild dog populations, and Chapter 7 discusses the rôle that captive wild dogs might play in this effort. Chapter 8 describes the additional research that is needed to allow us to refine our strategies for wild dog management. Chapter 9 summarizes the recommendations of Chapters 7 and 8 to propose actions for wild dog conservation in each range state.
The Action Plan also has four appendices. Some of the tactics that we discuss for wild dog conservation involve vaccination against infectious diseases, and immobilization for radio-collaring. Such procedures have been the subject of considerable controversy and we have, therefore, included a full discussion of this issue in Appendix 1. Appendix 2 provides details of some techniques used in current research projects on wild dogs, which may be of use to people directly involved with the management of wild dog populations. Appendix 3 is a list of contributors to this Action Plan, and Appendix 4 gives a detailed bibliography of publications concerning wild dogs.
Many of the problems faced by wild dogs stem from basic features of their natural history. Here we discuss aspects of wild dog biology which are important in understanding the reasons for their decline, and in devising plans for their conservation.
Wild dogs mostly hunt medium-sized antelope; their principal prey in several parts of Africa are summarized in Table1.1. They will chase larger species, such as eland and buffalo, but rarely kill such prey (Creel & Creel 1995; Ginsberg 1992). Wild dogs also take small prey such as hares, lizards and even eggs (Creel & Creel 1995; Ginsberg 1992), but these probably make a fairly small contribution to their diet.
Wild dogs do take livestock in some areas, but this is a fairly rare occurrence. In and around the Masai Mara National Reserve, Kenya, wild dogs ignored livestock (Fanshawe 1989; Fuller & Kat 1990), and in one case in Zimbabwe they ran through a paddock of calves to chase a kudu in the neighbouring paddock (Rasmussen 1996). The only study of wild dog depredation on livestock found that the dogs took far fewer cattle than the farmers believed (Rasmussen 1996). Nevertheless wild dogs can occasionally become a severe problem for livestock, especially smaller stock such as sheep and goats (Chapter4).
|Table 1.1 Principal prey taken by wild dogs in various study sites across Africa. The proportions of prey taken in Serengeti were calculated from data presented in Fanshawe & FitzGibbon (1993); all other data were presented by wild dog atending the IUCN/SSC Canid Specialist Group's Workshop on the Conservation & Recovery of the African wild dog, held in Arusha, Tanzania, in 1992. Data from Hwange provided by J.R.G., data from Kruger provided by M.G.L. Mills, data from Masai Mara provided by P.Kat, data from Moremi provided by J.W.McNutt, data from Namibia provided by L.Scheepers, data from Selous provided by S.Creel, data from Zambia provided by F. Munyenyembe.|
|Study site||Preferred prey|
|Hwange National Park, Zmbabwe||impala (60%)||kudu (30%)||reedbuck (2%)|
|Kruger National Park, South Africa||impala (52%)||kudu (12%)||reedbuck (15%)|
|Masai Mara National Reserve, Kenya||Thompson's gazelle (67%)||impala (17%)||wildebeest (8%)|
|Moremi Game Reserve, Botswana||impala (85%)||kudu||lechwe|
|Selous Game Reserve, Tanzania||impala (69%)||wildebeest (11%)||reedbuck (3%), warthog (3%)|
|Serengeti National Park, Tanzania||Thompson's gazelle (57%)||wildebeest (40%)||Grant's gazelle, zebra|
Wild dogs are intensely social animals, spending almost all of their time in close association with one other. Packs may be as small as a pair, or number as many as 27 adults and yearlings (M.G.L. Mills, pers. comm., R.Burrows 1993, Fuller et al. 1992a) - average pack compositions for various study sites are summarized in Table1.2. Packs are formed when small same-sex sub-groups - usually siblings - leave their natal groups and join up with other sub-groups of the opposite sex. Thus, in newly-formed packs the females are closely related to one another, but not to the males, and the males are closely related to one another, but not to the females (Burrows 1995; Frame et al. 1979; Fuller et al. 1992a). Young born into such packs may remain there, or disperse as yearlings or young adults to form new packs.
|Table 1.2 Pack compositions of wild dogs in various study sites across Africa. Data for Hwange, Kruger, Masai Mara, Moremi and Serengeti are from Fuller et al. (1992a), and data from Selous are from Creel & Creel (1995).|
|Hwange National Park, Zimbabwe||5 packs||7.8||3.2||5.4|
|Kruger National Park, South Africa||8 packs||4.8||2.1||5.8|
|Masai Mara National Reserve, Kenya||6 packs||4.2||4.0||8.8|
|Moremi Game Reserve, Botswana||8 packs||4.3||2.5||8.3|
|Selous Game Reserve, Tanzania||6 packs||7.7||4.3||6.3|
|Serengeti National Park, Tanzania||7 packs||6.6||6.0||11.2|
Members of wild dog packs hunt cooperatively. By hunting together, they can capture prey much larger than themselves which would not otherwise be accessible to them. Wild dogs weigh 20-25kg, but their prey average around 50kg, and may be as large as 200kg (Creel & Creel 1995; Malcolm & van Lawick 1975).
Wild dog hunts are almost always preceded by a ?social rally' which is believed to coordinate the pack in preparation for hunting (Estes & Goddard 1967; Kuhme 1965). Once prey sight the dogs, they may flee, or stand and defend themselves alone or as a herd (Creel & Creel 1995; Kuhme 1965). During chases, wild dogs may run at speeds of up to 60km/h, and are specially adapted to deal with the heat stress that this involves (Taylor et al. 1971). During such chases, wild dogs are spaced around the running prey so that a member of the pack can intercept the quarry as it turns. After this dog has made the first grab, other pack members cooperate to drag the quarry to a halt (Creel & Creel 1995; Estes & Goddard 1967; Kuhme 1965).
Once the quarry has been brought to bay, one or a few dogs may distract it from the front, while others attack from behind and begin to disembowel it (Kuhme 1965). Alternatively, one pack member may restrain the head of the prey by biting its nose, and holding on while the others make the kill (Creel & Creel 1995; Malcolm & van Lawick 1975). When hunting ungulate calves, some members of a wild dog pack may distract the mother while the remainder attack her calf.
As a result of such cooperative hunting, each pack member has a higher foraging success (measured as kg killed per km chased) than it would if it hunted alone (Creel & Creel 1995). Larger packs are also better able to defend their kills against scavenging hyaenas (Fanshawe & FitzGibbon 1993).
In most wild dog packs, a single dominant female is the mother of all the pups, although two or even three females may breed on some occasions (Fuller et al. 1992a). However, all pack members are involved in caring for the pups (Frame et al. 1979; Malcolm & Marten 1982; van Heerden & Kuhn 1985). Such additional care is vital if pups are to survive: packs rarely manage to raise any pups if they contain fewer than four members (S.R. Creel pers. comm.).
The pups are born in a den, where they remain for the first three months of life. The mother is confined to the den during early lactation, and relies on other pack members to feed her at this time. Wild dogs deliver food to the mother by regurgitation; later on, they regurgitate to the pups as well (Malcolm & Marten 1982). Some pack members also ?babysit' the pups, and chase predators away from the den (Malcolm & Marten 1982).
Perhaps because so many helpers are available to assist with pup care (Creel & Creel 1991), wild dogs' litters are enormous: litters number 10-11 pups on average and occasionally contain as many as 21 pups (Fuller et al. 1992a). Pup mortality may be high, however. There is some evidence to suggest that more pups survive in packs where there are more helpers to assist with their care, but this is certainly not always the case (S.R. Creel pers. comm., Burrows 1995; Fuller et al. 1992a; Malcolm & Marten 1982).
As well as a dominant, breeding female, each pack also has a dominant male (Frame et al. 1979; Malcolm & Marten 1982). Both mating behaviour and genetic analysis indicate that the dominant male fathers most (but not all) of the pups (D.Girman pers. comm., Malcolm & Marten 1982). However, dominant males are usually no more assiduous in caring for the pups than are other males in the pack (Malcolm & Marten 1982).
Since wild dog females cannot breed without assistance, in most cases the pack, rather than the individual, should be considered the basic unit within the population.
Wild dogs have enormous home ranges (Table1.3), much larger than would be expected on the basis of their body size (Gittleman & Harvey 1982). Packs are confined to relatively small areas when they are feeding young pups at a den, but outside the denning period they are truly nomadic. For example, in Serengeti home ranges were 50-260km² during denning, but 1,500-2,000km² at other times (Burrows 1995), and a pack in Kruger ranged over 80km² when denning, but 885km² after denning (Gorman et al. 1992).
|Table 1.3 Home ranges of wild dogs in various study sites across Africa. Data for Aitong are from Fuller & Kat (1990), data from Hwange, Kruger, Moremi and Serengeti are from Fuller et al. (1992a), and data from Selous are from Creel & Creel (1995).|
|Study site||Sample||Home range size|
|Aitong, near Masai Mara, Kenya||1 pack||659 km²|
|Hwange National Park, Zimbabwe||4 packs||423 km² (range 260-633 km²|
|Kruger National Park, South Africa||20 packs||553 km² (ran ge 150-1,110 km²)|
|Moremi Game Reserve, Botswana||9 packs||617 km² (range 375-1,050 km²)|
|Selous Game Reserve, Tanzania||6 packs||438 km² (range 620-2,460 km²)|
|Serengeti National Park, Tanzania||5 packs||1,318 km² (range 620-2,460 km²)|
The home ranges of different wild dog packs may overlap considerably, but they rarely enter one anothers' core areas and so their ranges are, to some extent, exclusive (Fuller et al. 1992a). As a result, wild dogs' large home ranges translate into very low population densities (Table1.4). The reasons why wild dogs live at such low densities are not clear, but several studies indicate that their numbers are rarely limited by the availability of ungulate prey (Creel & Creel 1996; Fuller et al. 1992a; Mills & Biggs 1993). This issue is discussed in detail in Chapter 4.
|Table 1.4 Population densities of wild dogs in various study sites across Africa. Data for Aitong are from Fuller & Kat (1990), data for Hluhluwe, Hwange, Selous and Serengeti are from Creel & Creel (1996), and data for Kruger are from Mills & Biggs (1993).|
|Study site||Population density (adults/100 km²)|
|Aitong, near Masai Mara, Kenya||2..6-4.6|
|Hluhluwe-Umfolozi Park, South Africa||3.3|
|Hwange National Park, Zimbabwe||1.5|
|Kruger National Park, South Africa||2.0|
|Selous Game Reserve, Tanzania||4|
|Serengeti National Park, Tanzania|
Even wild dog packs which inhabit protected areas may travel extensively outside the reserve borders - where they encounter human activity and threats such as roads, snares and livestock farmers likely to persecute them (Chapter 4). Wild dogs dispersing away from their natal packs range even more widely - they have been followed for hundreds of kilometres (Fuller et al. 1992b) and single wild dogs, or single-sex groups, are occasionally reported from countries such as the Democratic Republic of Congo and Uganda, where there has been no resident wild dog population for some years (Chapter3).
In this chapter, we have outlined the background to the problems faced by wild dogs today, and given brief details of their natural history. An important conclusion is that the pack, rather than the individual, should be considered the basic unit of wild dog populations.
The next chapter will discuss wild dog taxonomy and other aspects of wild dog genetics important in their conservation.
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© 1997 International Union for the Conservation of Nature and Natural Resources.