Overview

Brief Summary

Species Abstract

The Painted Hunting Dog is one of the most critically endangered carnivores in Africa, with approximately 5000 individuals extant. The early Holocene species population is estimated at about one half million, and the prehistoric range covered virtually all of the non-desert, non-jungle area of sub-Saharan Africa. In the last several decades, the habitat of L. Pictus has become severely fragmented due to expansion of the human population in Africa, and numbers have dwindled. The species has been historically termed "African Wild Dog", which name is declining in usage, due to pejorative aspects of such a name.

There are five acknowledged subspecies: L.p. pictus (Southern Africa), L.p. lupinus (East Africa), L.p. manguensis (West and Central Africa), L.p. saharicus (Sahara) and L.p. somalicus (Horn of Africa). Based upon the detailed geographic analysis in the following, the last three subspecies can be clearly considered critically endangered due to the extremely low populations in their sub-regions and the ongoing severe threats in those areas.

Until as recently as the 1970s L. pictus had a broad distribution within sub-Saharan Africa. Presently Botswana has one of the most robust populations with an estimated 150 individuals in the northeast part of the Okavango Delta, encompassing the eastern part of the Moremi Game Reserve. The area is approximately 3,000 square kilometres. This region is protected not just by paper legislation, but due to the care and concern of the Botswanan people. In Namibia, approximately 200 wild dogs are estimated to have existed by 2008, with the population dying off by about ten percent per annum. Virtually all of the sightings are in Tsumkwe East and West Caprivi. Livestock grazing and poaching account for the majority of the decline of the species in Namibia.

There is little sexual dimorphism in this species with adults attaining body mass to 36 kg and lengths to 110 cm plus tail extension to an additional 40 cm.(Novak) Pelage exhibits a short fur of mottled yellow, red, black and white patches. This animal is lean and muscular, built for running endurance. Each foot manifests four toes. Characteristically the bushy tail has a white tip. Ears are ovoid and sizable.

The pack chooses a den location which affords a good burrow, but which also ideally has some shade to protect pups from the harsh sun when they first begin to venture from the den. This shade also helps to mitigate against high burrow temperatures in the hot season. A typical burrow landform might consist of loamy soil which was colonised by termites. After construction of the termite mound to pulverise and soften the soil, an aardvark may have used this locale for rooting. A culmination of decades of earthwork by these two pioneer species will then be the formation of a more intricate den by the Painted Hunting Dog pack, building upon the previous subsurface work of others.

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Description

The African wild dog is one of the world's most social and distinctive canids. The short, wiry coat is coloured in blotches of yellow, grey, black and white (2) that give rise to the scientific name of Lycaon pictus or 'painted wolf-like animal' in Greek (3). The hair is short on the limbs and body and longer on the neck. Each dog has a unique colouration pattern, and this is used by researchers to identify individuals (2). The body is thin and muscular, the tail bushy with a white tip and the legs are long (2) (4). Males are slightly larger than females (2). Unlike the other canid species there are only four, rather than five, toes on the front feet (2) (4). These dogs have large rounded ears, which probably help with heat loss as well as keeping track of pack members by picking up long distance vocal signals (2) (4).
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Biology

African wild dogs are highly sociable and exhibit a very unusual social system; within their packs, dogs of the same sex are closely related to each other but not to individuals of the opposite sex, and only the dominant male and female will breed (4) (5) (6). Packs vary in size from 2 to 27 individuals and new packs are formed when subgroups of the same sex (usually siblings) disperse and join up with a subgroup of the opposite sex (5), leading to the unusual configuration of the pack. Only the dominant female will give birth to pups, and births may take place throughout the year, although they are more common between March and June (2). Litter size is the largest of any canid, averaging ten pups (2); these are born within dens where they remain for around three months (6). Initially the mother will stay with her pups and when members of the pack return from hunting they regurgitate food for her. As the pups get older however, all pack members help with feeding and 'baby sitting' of the young dogs (6). Juveniles are fully independent at 16 to 24 months (6) but remain with their pack, females are more likely to disperse, usually leaving in a sub-group with their sisters once they reach two years old (6). Outside of the breeding season, wild dogs are nomadic and wander over large distances in search of prey; home ranges can be as large as 5,000 square kilometres, but are often restricted to areas of less than 200 square kilometres (2). These dogs are carnivorous and hunt their prey by cooperating closely in a group (2). This strategy enables them to hunt prey comprising antelope and ungulates much larger than themselves, to include kudu bulls and wildebeest weighing up to 250 kilograms, as well as ensuring their hunting success is much higher than that of other large carnivorous species (2). Packs set out to hunt in the cool of dawn and dusk, avoiding other predators such as lions. The victim is pulled to the ground and the group descends to feed; pups in the pack are allowed to eat first (6).
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The tri-coloured African wild dog (Lycaon pictus) is probably best known for its organised and well-structured social system (Girman et al., 1997). Pack sizes are based on a trade-off of energetic costs and benefits (Hayward et al., 2006), with larger packs (approx. 20-40 adults) experiencing fitness benefits in terms of hunting success and pup survival (Woodroffe, 2011). Their social structure allows for a dominant breeding pair, with a single breeding female, but rearing of the pups becomes the responsibility of the sub-dominant “helpers” in the pack (Courchamp and Macdonald, 2001).

These mammals are medium sized carnivores (20-28 kg) and mainly prey on hoofed animals (ungulates), such as eland and zebra (although it has been reported that they can bring down juvenile buffalo), hunting at dusk and dawn and consuming more meat per body weight than any other carnivore (Hayward et al., 2006). They typically inhabit open woodland, where competition pressure from other predators such as lions, is slightly reduced (Vucetich and Creel, 2001).

Due to their large range-size requirements, inevitable conflict with humans has been, and continues to be, the primary reason for the decline in numbers of this endangered mammal (IUCN Red List of Threatened Species). Excessive hunting several decades ago drastically reduced the population size of the African wild dogs, now occupying less than 7% of their original geographic range (Woodroffe, 2011).

Conservation bodies such as The African Wildlife Foundation and the African Wild Dog Conservancy focus on monitoring pack sizes and activity, educating local communities and reducing poaching activity in order to stabilise current populations and boost those that are showing a decline. Additionally, the creation of corridors to link scattered populations and increase gene-flow between populations could prove invaluable for the on-going survival of these species (Davies-Mostert et al., 2012).  

  • Girman, D. J., Mills, M. G. L., Geffen, E. and Wayne, R. K. 1997. A molecular genetic analysis of social structure, dispersal and interpack relationships of the African wild dog (Lycaon pictus). Behav. Ecol. Sociobiol. 40, 187-198.
  • Courchamp, F. and Macdonald, D. W. 2001. Crucial importance of pack size in the African wild dog Lycaon pictus. Anim. Conserv. 4, 169-174.
  • Vucetich, J. A. and Creel, S. 2001. Ecological interactions, social organisation, and extinction risk in African wild dogs. Conserv. Biol. 13, 1172-1182.
  • Hayward, M. W., O’Brien, J., Hofmeyer, M. and Kerley, G. I. H. 2006. Prey preference of the African wild dog Lycaon pictus (Canidae: Carnivora): Ecological requirements for conservation. J. Mammal. 87, 1122-1131.
  • Woodroffe, R. 2011. Demography of a recovering African wild dog (Lycaon pictus) population. J. Mammal. 92, 305-315.
  • Davies-Mostert, H. T., Kamler, J. F., Mills, M. G. L., Jackson, C. R., Rasmussen, G. S. A., Groom, R. J. and Macdonald, D. W. 2012. Long-distance transboundary dispersal of African wild dogs among protected areas in southern Africa. Afr. J. Ecol. 50, 500-506.
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MammalMAP: African Wild dogs

African Wild dogs or Lycaon pictus (stemming from the Greek word for ‘wolf’ and the latin word for ‘painted’) are the only canid species to lack dewclaws on their forelimbs. Africa’s largest canid – an adult weighs within the range of 18 – 36 kgs.

African Wild dogs form packs that on average consists of 7 – 15 members. Before the recent decline in the population, packs consisting of 100 members have been recorded. Animals within the pack have a unique social concerns and structure – from cooperation during hunts to regurgitating food to feed the young, sick, elderly or wounded.

Typically, wild dogs feed on ungulates that are often twice their body weight. The hunt is lead by the alpha male usually in the morning or early evening. Wild dogs locate prey by sight and pursue it in high speed chases that can reach 55 km/hr until the prey is exhausted.

The IUCN has listed African Wild dogs as an endangered species since 1996. The decline in population size has been attributed to shrinking habitats, conflict with livestock and game farmers, road accidents and infectious diseases.

For more information on MammalMAP, visit the MammalMAP virtual museum or blog.

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Comprehensive Description

African Hunting Dogs have large, rounded ears, a thin body, and long, muscular legs. Their scientific name, Lycaon pictus, literally means "painted or ornate wolf. “ The patterns of their fur, which appear to be painted with patches of brown, red, black, yellow and white, are individually unique.

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Distribution

Range Description

Historical data indicate that African Wild Dogs were formerly distributed throughout sub-Saharan Africa, from desert (Lhotse 1946) to mountain summits (Thesiger 1970), and probably were absent only from lowland rainforest and the driest desert (Schaller 1972). They have disappeared from much of their former range. The species is virtually eradicated from North and West Africa, and greatly reduced in Central Africa and North-east Africa. The largest populations remain in southern Africa (especially northern Botswana, western Zimbabwe, eastern Namibia, and western Zambia) and the southern part of East Africa (especially Tanzania and northern Mozambique).

The current geographic distribution of African Wild Dogs was estimated using data compiled by the IUCN SSC range-wide conservation planning process for Cheetahs and African Wild Dogs, including regional strategies (IUCN SSC 2008, in prep.) and subsequent associated national action plans (www.cheetahandwilddog.org). Current African Wild Dog range was considered to comprise only the “resident range” identified by participants in the IUCN SSC process: this represents land where participants were confident that African Wild Dogs had been confirmed to be resident within the previous 10 years. Land where residence was not confirmed (e.g., possible range, unknown range) was excluded.
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Geographic Range

At one time the distribution of Lycaon pictus was throughout the non-forested and non-desert areas of Africa. Their current distribution is more fragmented. African hunting dogs are now found in Namibia, Botswana, Mozambique, parts of Zimbabwe, Swaziland, and the Transvaal.

Biogeographic Regions: ethiopian (Native )

  • Kingdon, J. 1997. The Kingdon Field Guide to African Mammals. San Diego: Academic Press.
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Historic Range:
Sub-Saharan Africa

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Range

The African wild dog was formerly present throughout sub-Saharan Africa, although it was never locally abundant (2) (4). This species is today restricted to fragmented populations mainly in southern and eastern Africa (2) (4). Potentially viable populations currently exist in Botswana, Kenya, Mozambique, South Africa, Tanzania, Zambia and Zimbabwe (2) (4).
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Physical Description

Morphology

Physical Description

The African hunting dogs' scientific name, Lycaon pictus, reflects the color of their pelage. Lycaon pictus literally means "painted or ornate wolf." The fur appears to be painted with brown, red, black, yellow and white areas. The pattern of colors is different on each animals coat, much like the stripes of zebras. The fur of L. pictus is short, with little or no underfur, and the blackish skin is sometimes visible where fur is sparse. Typically there is dark fur on the head and a white tip on the end of their bushy tail. They have large, rounded ears, a thin body, and long, muscular legs with four toes on each foot. The body length of Lycaon pictus is between 75 and 110 cm, the tail is between 30 and 40 cm long, and they range in weight from 18 to 36 kg. Males and females tend to be approximately the same size.

Range mass: 18 to 36 kg.

Range length: 75 to 110 cm.

Other Physical Features: endothermic ; homoiothermic; bilateral symmetry

Average basal metabolic rate: 33.01 W.

  • Stuart, C., T. Stuart. 1995. Stuart's Field Guide to the Mammals of Southern Africa. Cape Town: Struik.
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Ecology

Habitat

Habitat and Ecology

Habitat and Ecology
African Wild Dogs are generalist predators, occupying a range of habitats including short-grass plains, semi-desert, bushy savannas and upland forest. While early studies in the Serengeti National Park, Tanzania, led to a belief that African Wild Dogs were primarily an open plains species, more recent data indicate that they reach their highest densities in thicker bush (e.g., Selous Game Reserve, Tanzania; Mana Pools National Park, Zimbabwe; and northern Botswana). Several relict populations occupy dense upland forest (e.g., Harenna Forest, Ethiopia; Malcolm and Sillero-Zubiri 2001). African Wild Dogs have been recorded in desert (Lhotse 1946) (although most desert populations are now extirpated), but not in lowland forest. It appears that their current distribution is limited primarily by human activities and the availability of prey, rather than by the loss of a specific habitat type.

African Wild Dogs mostly hunt medium-sized antelope. Whereas they weigh 20–30 kg, their prey average around 50 kg, and may be as large as 200 kg. In most areas their principal prey are Impala (Aepyceros melampus), Greater Kudu (Tragelaphus strepsiceros), Thomson's Gazelle (Eudorcas thomsonii) and Common Wildebeest (Connochaetes taurinus). They will give chase of larger species, such as Common Eland (Tragelaphus oryx) and African Buffalo (Syncerus caffer), but rarely kill such prey. Small antelope, such as Dik-dik (Madoqua spp.), Steenbok (Raphicerus campestris) and Duiker (tribe Cephalophini) are important in some areas, and warthogs (Phacochoerus spp.) are also taken in some populations. African Wild Dogs also take very small prey such as hares, lizards and even eggs, but these make a very small contribution to their diet.

Generation length
Data on lifetime reproductive success of 19 alpha (breeding) females in western Zimbabwe indicate that 50% of reproductive output was achieved by age 5.5 years (SD 1.35, range 3–8; G.S.A. Rasmussen, unpubl. data). An alternative method, considers the average age of mothers of known litters, without the need for data on lifetime reproductive success. This method gives good agreement with the IUCN recommendations on calculating generation length, indicating a mean female breeding age of 5.7 years from the Zimbabwe dataset. Using this method, data from 18 litters born in Kenya to known-age mothers suggest a mean generation length of 5.0 years (R. Woodroffe, unpubl. data). Both studies suggest a minimum age at first breeding of approximately three years. Based on these data, for convenience we have estimated changes in African Wild Dog populations using a generation time of five years.

Systems
  • Terrestrial
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African hunting dogs are found in grasslands, savannahs and open woodlands. They are widely distributed across the African plains and are not found in jungle areas. Their habitat also includes semi-desert to mountainous areas south of the Sahara Desert in Africa.

Habitat Regions: temperate ; tropical ; terrestrial

Terrestrial Biomes: savanna or grassland ; chaparral ; forest

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The preferred habitats of the African Hunting Dog are savanna and open woodland. Once widespread, hunting dogs now survive in small fragmented populations in Namibia, Botswana, Mozambique,Zimbabwe, Swaziland, and the Transvaal.

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Inhabits a range of habitats, including the plains, semi-desert, bushy savannah, woodlands and upland forest (2).
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Trophic Strategy

Food Habits

African hunting dogs tend to prey on mammals that are about twice their weight. At times they will kill larger animals, and they will also take smaller prey individually. Some of the animals they prey on include small antelope such as impala (Aepyceros melampus) and bush duiker (Sylvicapra grimmia), and old, sick or injured larger animals such as wildebeest (genus Connochaetes) and zebra (genus Equus). On occasion some of the food they get from larger kills may be cached, though very often they never return to the cached food. For the most part Lycaon pictus does not eat plants or insects, except for small amounts of grass. Also African hunting dogs will never scavenge, no matter how fresh the kill is.

Animal Foods: mammals

Foraging Behavior: stores or caches food

Primary Diet: carnivore (Eats terrestrial vertebrates)

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Associations

Known prey organisms

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Predator of impala, duiker, wildebeest and zebra.

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Life History and Behavior

Behavior

Hunting dogs cooperatively hunt small antelopes such as impala and duikers, and old, sick or injured larger animals including wildebeest and zebra. They never scavenge, no matter how fresh the kill.

Highly social animals, hunting dogs form packs of up to 40 members, although average pack size is between 7 and 15. Pack members, led by a dominant female and male pair, cooperate in caring for and feeding the young and wounded or sick adults within their group.

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Life Expectancy

Lifespan/Longevity

Average lifespan

Status: wild:
10.0 years.

Average lifespan

Status: wild:
11.0 years.

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Lifespan, longevity, and ageing

Maximum longevity: 17 years (captivity) Observations: These animals have been estimated to live up to 17 years (Ronald Nowak 1999), though record longevity in captivity is only 15.1 years (Richard Weigl 2005). Further studies are necessary.
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Reproduction

Each African hunting dog pack has a dominant breeding pair. This pair can be identified by their increased tendency to urine mark. They are normally the only pair of pack members to mate and they tend to remain monogamous for life. Their life expectancy is approximately ten years. Generally the dominant pair prevents subordinates from breeding. Breeding suppression between females may often result in aggressive interactions. Occasionally a subordinate female is allowed to mate and rear young.

Mating System: monogamous ; cooperative breeder

Lycaon pictus reaches sexual maturity at approximately 12 to 18 months, though they usually do not mate until much later. The youngest recorded reproduction of a female was at 22 months old. Gestation is approximately ten weeks and pups are usually born between March and July. Litter sizes can vary considerably, from 2 to 20 pups. The smaller litter sizes have been recorded from animals in captivity. Breeding females gives birth to their litters in grass-lined burrows, usually an abandoned aardvark hole. The pups remain in the den with their mother for three to four weeks. Once the pups are brought out of the den they become the responsibility of the whole pack. Pups nurse from other females in the pack as well as from their mother. Weaning can occur as early as 5 weeks. The interval between litters is normally 12 to 14 months.

Breeding interval: Twelve to fourteen months

Breeding season: January to May

Range number of offspring: 2 to 20.

Average number of offspring: 8.

Range gestation period: 60 to 80 days.

Average gestation period: 72.4 days.

Range weaning age: 35 to 90 days.

Range age at sexual or reproductive maturity (female): 12 to 18 months.

Range age at sexual or reproductive maturity (male): 12 to 18 months.

Key Reproductive Features: iteroparous ; seasonal breeding ; gonochoric/gonochoristic/dioecious (sexes separate); viviparous

Average birth mass: 318 g.

Average number of offspring: 8.

Parental Investment: altricial ; post-independence association with parents; extended period of juvenile learning

  • Estes, R. 1991. The Behavior Guide to African Mammals. Berkeley and Los Angeles, California: The University of California Press.
  • Kingdon, J. 1997. The Kingdon Field Guide to African Mammals. San Diego: Academic Press.
  • Nowak, R. 1999. Walker's Mammals of the World Sixth Edition. Baltimore: The Johns Hopkins University Press.
  • Stuart, C., T. Stuart. 1995. Stuart's Field Guide to the Mammals of Southern Africa. Cape Town: Struik.
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Conservation

Conservation Status

IUCN Red List Assessment


Red List Category
EN
Endangered

Red List Criteria
C2a(i)

Version
3.1

Year Assessed
2012

Assessor/s
Woodroffe, R. & Sillero-Zubiri, C.

Reviewer/s
Hoffmann, M. & Hilton-Taylor, C.

Contributor/s
Rasmussen, G.

Justification
African Wild Dogs have disappeared from much of their former range. Their population is currently estimated at approximately 6,600 adults in 39 subpopulations, of which only 1,400 are mature individuals. Population size is continuing to decline as a result of ongoing habitat fragmentation, conflict with human activities, and infectious disease. Given uncertainty surrounding population estimates, and the species’ tendency to population fluctuations, the largest subpopulations might well number <250 mature individuals, thereby warranting listing as Endangered under criterion C2a(i).

History
  • 2008
    Endangered
  • 2004
    Endangered
  • 1996
    Endangered
  • 1994
    Endangered
    (Groombridge 1994)
  • 1990
    Endangered
    (IUCN 1990)
  • 1988
    Vulnerable
    (IUCN Conservation Monitoring Centre 1988)
  • 1986
    Vulnerable
    (IUCN Conservation Monitoring Centre 1986)
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Current Listing Status Summary

Status: Endangered
Date Listed: 01/23/1984
Lead Region: Foreign (Region 10) 
Where Listed: Entire


Population detail:

Population location: Entire
Listing status: E

For most current information and documents related to the conservation status and management of Lycaon pictus , see its USFWS Species Profile

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Lycaon pictus is listed as endangered by the IUCN and the United States Endangered Species Act. Habitat loss and diseases that are spread by domestic animals jeopardize the remaining African hunting dog populations.

US Federal List: endangered

CITES: no special status

IUCN Red List of Threatened Species: endangered

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IUCN Red List Status: ENDANGERED

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Status

Classified as Endangered (EN) on the IUCN Red List (1).
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Population

Population
Background
African Wild Dogs are rarely seen, even where they are relatively common, and it appears that populations have always existed at very low densities. Table 2 in the additional supporting information (see link below) presents estimates of the sizes of African Wild Dog subpopulations occupying all discrete areas of resident range identified by participants in the range-wide conservation planning process (IUCN SSC 2008, 2009, in prep.). These comprise a total of 39 distinct subpopulations estimated to range in size from two to 276 mature individuals.

Proportion of mature individuals
Estimating the number of “mature individuals” is challenging, because African Wild Dogs are obligate cooperative breeders: within a pack, the alpha male and female are the parents of the majority of surviving pups (Girman et al. 1997). The IUCN Red List User Guidelines (IUCN 2010) define mature individuals as “individuals known, estimated or inferred to be capable of reproduction”, but do not specify the time period within which reproduction is considered possible. The User Guidelines go on to state “in many taxa there is a pool of non-reproductive (e.g., suppressed) individuals that will quickly become reproductive if a mature individual dies. These individuals can be considered to be capable of reproduction”.

In African Wild Dogs, a high proportion of individuals are indeed reproductively suppressed, but these animals do not always become reproductive “quickly” if an alpha individual dies. In a mature pack, most pack members are offspring of the alpha pair; for these animals, death of an alpha would usually not open up a breeding opportunity because no unrelated mates would be available within the pack. In our experience, death of an alpha often leads to disintegration of the pack, with no breeding until new packs are formed. Given these complexities, and in keeping with the spirit of capturing a “snapshot” of current conditions, we have chosen to define mature individuals as those considered capable of reproduction within the current breeding season. The number of mature individuals thus comprises the number of alpha males and females, and the number of sub-dominant (i.e. non-alpha) animals which breed successfully.

Table 1 in the additional supporting information (see link below) provides demographic data to allow the estimation of numbers of mature individuals (Nm) from the census population of adults and yearlings (Nc). Using these data, the number of alpha males (NaM) and females (NaF) would be estimated thus:

NaM = Nc x 0.55 x 0.176
NaF = Nc x 0.45 x 0.215

Reassuringly, these equations return approximately equal estimates of the numbers of alpha males and alpha females.

No published estimate is available of the proportion of adults and yearlings which breed successfully as sub-dominants. However, Girman et al. (1997) report the proportions of surviving pups with sub-dominant mothers and/or fathers. Using a simple assumption that litter size was unrelated to social status, we therefore estimate the number of sub-dominant breeders (Nsub) as:

Nsub = (NaM x 0.10) + (NaF x 0.08)

The number of mature individuals is then calculated as

Nm = NaM + NaF + Nsub

We made exploratory calculations to investigate the effect of including the probability of currently-suppressed individuals surviving and breeding in subsequent years. These calculations became highly complex, and risked double-counting animals which might breed as sub-dominants in the current year and then attain alpha (dominant) status in later years. It is also important to note that, in the previous listing, the number of mature individuals was estimated simply as the number of alpha animals (i.e., NaM + NaF). These alternative ways of calculating numbers of mature individuals contribute to considerable uncertainty about African Wild Dog numbers.

Current population size
Table 2 in the additional supporting information (see link below) presents estimates of the sizes of African Wild Dog subpopulations occupying all discrete areas of resident range identified by participants in the range-wide conservation planning process (IUCN SSC 2008, 2009, in prep.). These comprise a total of 39 distinct sub-populations estimated to range in size from two to 276 mature individuals. Few wild dog subpopulations have been systematically monitored, and these estimates are subject to considerable imprecision. This imprecision, combined with uncertainty about the calculation of mature individuals, and the species’ propensity for substantial population fluctuations, means that these population estimates should be viewed with great caution. Lack of precision has important consequences for Red List assessment; the two largest sub-populations (estimated at 268 and 276 mature individuals) are only slightly larger than one of the threshold sub-population sizes used by the Red List criteria (250 mature individuals: EN C2(i)). Had only alpha animals been considered mature individuals (as in the previous assessment), the sizes of these two largest sub-populations fall to 246 and 253 respectively. This uncertainty needs to be taken into account in comparing estimates of African Wild Dogs’ population sizes with IUCN Red List criteria.

Change in population size
Data on African Wild Dogs’ past global population size are taken from Ginsberg and Woodroffe (1997). It is important to note, once again, that few of these population estimates are based on systematic monitoring, and all should be viewed with caution. Assessing changes in population size is complicated by the fact that a less complete dataset was available in 1997 than in 2012. As a result of these improved data, the global estimate of African Wild Dog population size is in fact higher for 2012 than for 1997. However, this difference reflects the greater area surveyed in 2012.

To try to overcome this problem, Table 3 in the additional supporting information (see link below) compares estimates of subpopulation size in 1997 (taken from Ginsberg and Woodroffe 1997) with estimates of African Wild Dog subpopulations from the same areas in 2012. Subpopulations known in 2012 but not known in 1997 are excluded. However, sub-populations known to be absent in 1997 (e.g. the managed metapopulation in South Africa, and the sub-population in Laikipia District, Kenya) are included in Table 3 . Hence, Table 3 represents our best assessment of a like-with-like comparison of African Wild Dog numbers in 1997 and 2012

Causes of decline
The causes of African Wild Dogs’ decline are reasonably well understood and include extreme sensitivity to habitat fragmentation as a consequence of wide-ranging behaviour, conflict with livestock and game farmers, accidental killing by people in snares and road accidents, and infectious disease. All of these causes are associated with human encroachment on African Wild Dog habitat and, as such, have not ceased and are unlikely to be reversible across the majority of the species’ historical range.

Fluctuations in population size
Populations of African Wild Dogs are prone to marked fluctuations at a variety of temporal and geographical scales which are likely to both increase extinction risks and undermine the precision of population estimates. At the local scale, a combination of high mortality, high fecundity, and dispersal by both sexes means that pack size fluctuates substantially over short periods (Figure 1 in the attached PDF), although fluctuation in numbers of mature individuals would be less dramatic. Because African Wild Dogs are seasonal breeders across most of their remaining geographic range, fluctuations may be synchronised across packs (see Figure 1 in the additional supporting information).

The same demographic characteristics – high mortality, high fecundity, and long-distance dispersal – likewise lead to fluctuations at the population scale. This pattern is further exaggerated by the species’ susceptibility to infectious disease which can cause rapid die-offs. Local extinctions are not uncommon, and are often both rapid and unanticipated. Figure 2 in the additional supporting information presents data from three relatively well-documented cases of local extinction involving small wild dog subpopulations affected by rabies.

Similar die-offs have been documented in larger African Wild Dog populations. For example, five of 12 study packs in Botswana (Alexander et al. 2010) and three of eight study packs in Kenya (Woodroffe 2011) have been reported as having died within short time periods during disease outbreaks. Although these relatively large study populations recovered, the majority of African Wild Dog sub-populations are estimated to comprise ≤20 mature individuals and could be severely compromised by outbreaks of this size.

For comparison, under good conditions African Wild Dog populations are also able to grow relatively quickly. African Wild dogs’ capacity for very long-range dispersal means that sub-populations sometimes reappear unexpectedly and grow rapidly; examples include natural recoveries in Samburu and Laikipia Districts, Kenya (Woodroffe 2011), Savé Valley Conservancy in Zimbabwe (Pole 2000), and the Serengeti ecosystem of Tanzania (Marsden et al. 2011). As an example of the speed of recovery, the subpopulation in Laikipia District, Kenya, grew from 0 in 1999 to 17 adults and yearlings in two packs by 2000, and by 2006 had increased 10-fold to 170 adults and yearlings (Woodroffe 2011).

On the basis of this evidence, we conclude that African Wild Dogs show substantial population fluctuations, but may not experience extreme fluctuations in sub-population size as outlined in the Red List guidelines. Nevertheless, the substantial fluctuations which do occur contribute to further uncertainty about sub-population sizes.


Follow the link below for additional supporting information.

Population Trend
Decreasing
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African Hunting Dogs require large home ranges to support viable populations. In recent years, human encroachment has reduced and fragmented their habitat; domesticated dogs have introduced diseases such canine distemper and rabies with devastating consequences for hunting dog populations.

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Threats

Major Threats
The principal threat to African Wild Dogs is habitat fragmentation, which increases their contact with people and domestic animals, resulting in human-wildlife conflict and transmission of infectious disease. The important role played by human-induced mortality has two long-term implications. First, it makes it likely that, outside protected areas, African Wild Dogs may be unable to coexist with increasing human populations unless land use plans and other conservation actions are implemented. Second, African Wild Dog ranging behaviour leads to a very substantial "edge effect", even in large reserves. Simple geometry dictates that a reserve of 5,000 km² contains no point more than 40 km from its borders – a distance well within the range of distances travelled by a pack of African Wild Dogs in their usual ranging behaviour. Thus, from an African Wild Dog's perspective, a reserve of this size (fairly large by most standards) would be all edge. As human populations rise around reserve borders, the risks to African Wild Dogs venturing outside are also likely to increase. Under these conditions, only the very largest unfenced reserves will be able to provide any level of protection for African Wild Dogs. In South Africa, “predator proof” fencing around small reserves has proved reasonably effective at keeping dogs confined to the reserve, but such fencing is not 100% effective (Davies-Mostert et al. 2009) and is unlikely to be long-term beneficial for wildlife communities.

Even in large, well-protected reserves, or in stable populations remaining largely independent of protected areas (as in northern Botswana), African Wild Dogs live at low population densities. Predation by Lions, and perhaps competition with Spotted Hyaenas, contribute to keeping African Wild Dog numbers below the level that their prey base could support. Such low population density brings its own problems. The largest areas contain only relatively small wild dog populations; for example, the Selous Game Reserve, with an area of 43,000 km² (about the size of Switzerland), is estimated to contain about 800 African Wild Dogs. Most reserves, and probably most African Wild Dog populations, are smaller. For example, the population in Niokolo-Koba National Park and buffer zones (about 25,000 km²) is likely to be not more than 50–100 dogs. Such small populations are vulnerable to extinction. "Catastrophic" events such as outbreaks of epidemic disease may drive them to extinction when larger populations have a greater probability of recovery – such an event seems to have led to the local extinction of the small African Wild Dog population in the Serengeti ecosystem on the Kenya-Tanzania border. Problems of small population size will be exacerbated if, as seems likely, small populations occur in small reserves or habitat patches. As discussed above, animals inhabiting such areas suffer a strong "edge effect". Thus, small populations might be expected to suffer disproportionately high mortality as a result of their contact with humans and human activity.
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African wild dogs require large home ranges to support viable populations and recent habitat fragmentation has caused a population decline (4). Wild dogs traditionally have a reputation for attacking livestock, and despite this rarely occurring in practice, they are therefore often persecuted wherever they come into contact with humans (2) (4). In addition, road accidents and incidental snaring have recently become an important cause of mortality (2). Wild dogs are susceptible to disease; particularly those carried by domestic dogs such as canine distemper and rabies. A final threat that keeps wild dog populations low is competition and predation with the other large carnivores of the African savanna, such as lions and spotted hyaenas (6).
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Management

Conservation Actions

Conservation Actions
Conservation strategies have been developed for the species in all regions of Africa Sillero-Zubiri et al. 2004, Woodroffe et al. 1997, (IUCN SSC 2008, 2009, in prep.), and many range states have used these strategies as templates for their own national action plans (Department of Wildlife and National Parks 2008, Wildlife Service 2010). Although each regional strategy was developed independently through a separate participatory process, the three strategies have a similar structure, comprising objectives aimed at improving coexistence between people and African Wild Dogs, encouraging land use planning to maintain and expand wild dog populations, building capacity for wild dog conservation within range states, outreach to improve public perceptions of wild dogs at all levels of society, ensuring a policy framework compatible with wild dog conservation. These strategies are accessible at www.cheetahandwilddog.org.

Gaps in knowledge
Several pieces of information are needed to enable more effective conservation of African wild dogs. These include: 1) development of cost-effective methods for surveying wild dogs across large geographical scales; 2) surveys of wild dog distribution and status, particularly in Algeria, Angola, Central African Republic, Chad, Somalia, South Sudan, and Sudan; 3) development of locally-appropriate and effective means to reduce conflict between wild dogs and farmers; 4) establishing which techniques will be most effective and sustainable for protecting wild dogs from disease; and 5) determining the landscape features which facilitate (or prevent), wild dog movement over long distances and hence promote (or block) landscape connectivity.
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Conservation

The current population of African wild dogs is estimated to be less than 5,500 individuals (2). The majority of dogs are found in medium sized populations, which are extremely vulnerable to sudden environmental change. Due to their expansive home ranges, large areas of contiguous habitat are the key to the survival of this species and it is estimated that protected areas need to be greater than 10,000 square kilometres in order to prevent detrimental contact with local human settlements (2) (4). Consequently even most of the larger reserves cannot fully contain African wild dog packs, leading to clashes with humans (2). Preventing persecution through education is also a priority of the conservation action plan (2) (4), in an effort to preserve this most intriguing and unique canid.
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Relevance to Humans and Ecosystems

Benefits

Economic Importance for Humans: Negative

Lycaon pictus occasionally kills livestock and important game animals.

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Wikipedia

African Wild Dog

Lycaon pictus!<-- This template has to be "warmed up" before it can be used, for some reason -->

Animalia

Lycaon pictus is a large canid found only in Africa, especially in savannas and other lightly wooded areas. It is commonly called the African Hunting Dog, African Wild Dog, Painted Wolf, Cape Hunting Dog, Painted Dog, Painted Hunting Dog, Spotted Dog or Ornate Wolf.

Contents

Anatomy and reproduction

Skull of an African wild dog
At Denver Zoo, Colorado, USA

The scientific name "Lycaon pictus" is derived from the Greek for "wolf" and the Latin for "painted". It is the only canid species to lack dewclaws on the forelimbs.

Adults typically weigh 18–34 kilograms (40–75 lb).[2] A tall, lean animal, it stands about 75 centimetres (30 in) at the shoulder, with a head and body length averaging about 100 centimetres (39 in) long with a tail of 30 to 45 centimetres (12 to 18 in). Animals in southern Africa are generally larger than those in eastern or western Africa.

There is little sexual dimorphism, though judging by skeletal dimensions, males are usually 3-7% larger. It has a dental formula of

Dentition
3.1.4.2
3.1.4.3

for a total of 42 teeth. The premolars are relatively large compared with those of other canids, allowing it to consume a large quantity of bone, much like hyenas.[3] The heel of the lower carnassial M1 is crested with a single cusp, which enhances the shearing capacity of teeth and thus the speed at which prey can be consumed. This feature is called trenchant heel and is shared with two other canids: the Asian Dhole and the South American Bush Dog.

A study established that the African Wild Dog had a Bite Force Quotient of 142, the highest of any extant mammal of the order Carnivora.[4] The BFQ is essentially the strength of bite as measured against the animal's mass.

The African Wild Dog reproduces at any time of year, although mating peaks between March and June during the second half of the rainy season. Litters can contain 2-19 pups, though 10 is the most usual number.[5] The time between births is usually 12–14 months, though it can also be as short as 6 months if all of the previous young die. The typical gestation period is approximately 70 days.[6][not in citation given] Pups are usually born in an abandoned den dug by other animals such as those of the Aardvark. Weaning takes place at about 10 weeks. After 3 months, the den is abandoned and the pups begin to run with the pack. At the age of 8–11 months they can kill small prey, but they are not proficient until about 12–14 months, at which time they can fend for themselves. Pups reach sexual maturity at the age of 12–18 months.

Females will disperse from their birth pack at 14–30 months of age and join other packs that lack sexually mature females. Males typically do not leave the pack they were born to. This is the opposite situation to that in most other social mammals, where a group of related females forms the core of the pack or similar group. In the African Wild Dog, the females compete for access to males that will help to rear their offspring. In a typical pack, males outnumber females by a factor of two to one, and only the dominant female is usually able to rear pups. This unusual situation may have evolved to ensure that packs do not over-extend themselves by attempting to rear too many litters at the same time.[7] The species is also unusual in that other members of the pack including males may be left to guard the pups whilst the mother joins the hunting group; the requirement to leave adults behind to guard the pups may decrease hunting efficiency in smaller packs.[8]

A captive breeding and translocation program at Mkomazi Game Reserve, the first of its kind in East Africa, was founded in 1995 to provide dogs for a multinational effort to stabilize their numbers and to reintroduce the species to its traditional homeland. The dogs are allocated to four breeding compounds to maximize genetic diversity. An extensive veterinary program has been set up to improve their immunity to disease.

Social structure

In packs, there are separate male and female hierarchies that will split up if either of the alphas die. In the female group, the oldest will have alpha status over the others, so a mother will retain her alpha status over her daughters. For the males, in contrast the youngest male or the father of the other males will be dominant. When two such loner separate-gender groups meet, if unrelated they can form a pack together. Dominance is established without blood-shed, as most dogs within a group tend to be related to one another in some way, and when this is not the case they have a submission-based hierarchy, instead of a dominance based one. Submission and nonaggression is emphasized heavily, even over food they will beg energetically instead of fight. This is likely because of their manner of raising huge litters of dependent pups, so if one individual is injured the entire pack would not be able to provide as much.[9][dead link]

Unrelated African Wild Dogs sometimes join up in packs, but this is usually temporary. Occasionally, instead unrelated cape dogs will attempt hostile takeovers of packs.[10]

Hunting and diet

Dogs with a wildebeest carcass

The African Wild Dog hunts in packs. Like most members of the dog family, it is a cursorial hunter, meaning that it pursues its prey in a long, open chase. Nearly 80% of all wild dog hunts end in a kill; for comparison, the success rate of lions, often viewed as ultimate predators, is only 30%. Schaller found that 9 of 10 wild dog hunts in the Serengeti ended in kills.[11] Members of a pack vocalize to help coordinate their movements. Its voice is characterized by an unusual chirping or squeaking sound, similar to a bird. Wild dogs frequently kill larger prey via disemboweling, a technique that is rapid but has caused this species to have a negative, ferocious reputation. Even some early wildlife conservationists, such as Carl Akeley[12] delighted in killing entire wild dog packs.[13]

After a successful hunt hunters regurgitate meat for those that remained at the den during the hunt, such as the dominant female and the pups. They will also feed other pack members, such as the sick, injured, the very old that cannot keep up, or those who stayed back to watch the pups.

The African Wild Dog's main prey varies among populations but always centers around medium-to-large sized ungulates, such as the impala, Thomson's Gazelle, Springbok, kudu, reedbuck, and wildebeest calves. The most frequent single prey species depends upon season and local availability. For example, in the Serengeti in the 1970s wildebeest (mostly calves) were the most frequently taken species (57%) from January to June, but Thompsons gazelle were the most frequently taken (79%) during the rest of the year.[14] In the Selous Game Reserve, the most frequent prey is impala.[15] While the vast majority of its diet is made up of mammal prey, it sometimes hunts large birds, especially Ostriches.[7] Other predators, mainly lions, sometimes steal the prey that Wild Dogs catch.[16]

Some packs will also include large animals in their prey, such as zebras and warthogs. The frequency and success rates of hunting zebra and warthogs varies widely among specific packs (whereas the rates for wildebeest and smaller ungulates do not). Hunting larger prey requires a closely coordinated attack, beginning with a rapid charge to stampede the herd. One African Wild Dog then grabs the victim's tail, while another attacks the upper lip or nose, and the remainder disembowel the animal while it is immobilised. Male wild dogs usually perform the task of grabbing warthogs by the nose.[17] This behaviour is also used on other large dangerous prey, such as the African Buffalo, giraffe calves, and large antelope—even the one-ton Giant Eland.

Remarkably, this large-animal hunting tactic appears to be a learned behavior, passed on from generation to generation within specific hunting packs, rather than an instinctive behaviour found commonly within the species. Some studies have also shown that other information, such as the location of watering holes, may be passed on in a similar fashion.

Distribution and threats

The home range of packs varies enormously, depending on the size of the pack and the nature of the terrain. In the Serengeti, the average dog density (prior to the local extinction of the species) was 1 dog per 208 square kilometers (80 square miles), whereas in the Selous Game Reserve the average density was 1 dog every 25 square kilometers (9.6 square miles).[18] Their preferred habitat is deciduous forests because of large prey herd size, lack of competition from other carnivores, and better sites for denning.[19] In the Serengeti, the average range has been estimated at 1,500 square kilometres (580 square miles), although individual ranges overlap extensively.[7]

An African Wild Dog in a captive breeding program at Monarto Zoo, South Australia.

There were once approximately 500,000 African Wild Dogs in 39 countries, and packs of 100 or more were not uncommon. Now there are only about 3,000-5,500 in fewer than 25 countries,[1] or perhaps only 14 countries.[20] They are primarily found in eastern and southern Africa, mostly in the two remaining large populations associated with the Selous Game Reserve in Tanzania and the population centered in northern Botswana and eastern Namibia. Smaller but apparently secure populations of several hundred individuals are found in Zimbabwe, South Africa (Kruger National Park), and in the Ruaha/Rungwa/Kisigo complex of Tanzania. Isolated populations persist in Zambia, Kenya, and Mozambique.

The African Wild Dog is endangered by human overpopulation,[21] habitat loss and hunting. It uses very large territories (and so can persist only in large wildlife protected areas), and it is strongly affected by competition with larger carnivores that rely on the same prey base, particularly the lion and the Spotted Hyena. Lions often will kill as many wild dogs as they can but do not eat them. Hyenas usually follow them to steal their kills. One on one the hyena is much more powerful than the Wild Dog but a large group of Wild Dogs can successfully chase off a small number of hyenas because of their teamwork. It is also killed by livestock herders and game hunters, though it is typically no more (perhaps less) persecuted than other carnivores that pose more threat to livestock. Most of Africa's national parks are too small for a pack of wild dogs, so the packs expand to the unprotected areas, which tend to be ranch or farm land. Ranchers and farmers protect their domestic animals by killing the wild dogs. Like other carnivores, the African Wild Dog is sometimes affected by outbreaks of viral diseases such as rabies, distemper, and parvovirus. Although these diseases are not more pathogenic or virulent for wild dogs, the small size of most wild dog populations makes them vulnerable to local extinction due to diseases or other problems.[1]

The Painted Dog Conservation (PDC) effort, based in Hwange National Park, western Zimbabwe, works with local communities to create new strategies for conserving the wild dog and its habitat.

An African Wild Dog in repose

Name controversy

A controversy began in the late 1990s when conservationists working to protect Lycaon pictus said that their most common name, "African Wild Dog", was a source of confusion and prejudice. Conservationist Greg Rasmussen wrote in 1998:

"The name 'wild dog' developed during an era of persecution of all predators when the name applied to feral dogs, hyenas, jackals and the cape hunting dogs (Pringle, 1980). 'Painted' aside from being a direct translation of the specific epithet, accurately describes the unique varicoloured markings of each individual. Apart from being misleading, continued use of the name 'wild dog' does little more than further fuel negative attitude and prejudice which is detrimental to conservation efforts."

Rasmussen is one of the founders of the Painted Dog Conservation, which was originally founded in 1992 as Painted Dog Research Project. He advocates using the name "Painted Dog".

Taxonomy

Lycaon pictus is the only extant species in the genus Lycaon. An extinct species from the African Pleistocene, Lycaon sekowei, is a possible ancestor of the modern species.[22] Another possible ancestor of these (and several other canids) from the Pleistocene of Eurasia is Xenocyon lycaonoides; this is sometimes also placed in Lycaon.[23]

There are five recognized subspecies of Lycaon pictus:[24]

  • Lycaon pictus pictus
  • Lycaon pictus lupinus
  • Lycaon pictus manguensis
  • Lycaon pictus sharicus
  • Lycaon pictus somalicus

Wild Dog research

African Wild Dogs in Perth Zoo

Founded as the Botswana Wild Dog Research Project in 1989, the Botswana Predator Conservation Trust (BPCT) has expanded to cover all the large carnivore species in Botswana. It is one of the longest running large predator research projects in Africa and one of only a handful of its caliber worldwide. BPCT research on wild dogs has made it abundantly clear that the health and welfare of the entire predator population is a key indication of overall health of the ecosystem. The Government of Botswana, also acknowledging that appropriate and necessary resource management cannot be undertaken in the absence of accurate information about its natural resources, has entrusted BPCT with the task of leading northern Botswana's conservation and research initiatives on all large carnivores and their associated habitats. The Okavango Delta, where most of BPCT's research takes place, is a freshwater wetland of global importance. It is the largest Ramsar (International Convention on Wetlands) site on earth and was granted IUCN world heritage status by the World Conservation Union (IUCN).

Fossil of Lycaon sekowei, a possible ancestor of modern wild dogs

Under the leadership of Dr JW "Tico" McNutt, a number of international graduate students, Botswana national students, and local staff make up a strong field team that works on African Wild Dogs (Lycaon pictus), leopards (Panthera pardus), cheetahs (Acinonyx jubatus), Spotted Hyenas (Crocuta crocuta), and lions (Panthera leo).

Interspecies adoption

In 2009 at the Pittsburgh Zoo, a female mixed breed domestic dog was brought in to nurse nine African wild dog pups, after the pups' mother had died. The nursing was going successfully, and the pups had gained weight. This is the first time that a domestic dog has ever been documented nursing African wild dog pups.[25]

References

  1. ^ a b c McNutt et al. (2008). Lycaon pictus. In: IUCN 2008. IUCN Red List of Threatened Species. Downloaded on 06 May 2008. Database entry includes justification for why this species is endangered
  2. ^ Sillero-Zubiri, C., Hoffmann, M. and Macdonald, D.W. (2004) Canids: Foxes, Wolves, Jackals and Dogs: Status Survey and Conservation Action Plan. IUCN/SSC Canid Specialist Group, IUCN, Gland, Switzerland and Cambridge, UK, cited in Arkive: African Wild Dog
  3. ^ "African wild dog (Lycaon pictus)". Lioncrusher's Domain. http://www.lioncrusher.com/animal.asp?animal=2. Retrieved 2007-06-08. 
  4. ^ Wroe, Stephen & McHenry, Colin (2004-10-16). "Bite club: comparative bite force in big biting mammals and the prediction of predatory behaviour in fossil taxa". Proc. R. Soc. B 272 (1563): 619. doi:10.1098/rspb.2004.2986. PMID 15817436. PMC 1564077. http://intern.forskning.no/dokumenter/wroe.pdf. 
  5. ^ Animal Info - African Wild Dog
  6. ^ "BBC Wildfacts". http://www.bbc.co.uk/nature/wildfacts/factfiles/157.shtml. Retrieved 2008-05-31. 
  7. ^ a b c Malcolm, James (1984). Macdonald, D.. ed. The Encyclopedia of Mammals. New York: Facts on File. pp. 31. ISBN 0-87196-871-1. 
  8. ^ Franck Courchamp, Gregory S. A. Rasmussen, and David W. Macdonald (2002). "Small pack size imposes a trade-off between hunting and pup-guarding in the painted hunting dog Lycaon pictus". International Society for Behavioral Ecology. http://www.ese.u-psud.fr/epc/conservation/PDFs/Babysitting.pdf. 
  9. ^ Rebecca Postanowicz (2008). "African Wild Dogs". http://www.lioncrusher.com/animal.asp?animal=2. 
  10. ^ Roger Burrows (2002). "Wild Dog Pack Formation". http://www.africanconservation.org/wilddogs/burrowslycaonpack.html. 
  11. ^ Schaller, George B. Golden Shadows, Flying Hoovers. 1973. p 277.
  12. ^ Akeley, Lions, Gorillas, and Their Neighbors.
  13. ^ Schaller, George B. Golden Shadows, Flying Hooves. 1973. p215
  14. ^ Schaller, George B. Golden Shadows, Flying Hooves. 1973. p225.
  15. ^ "Hope Rises for Africa's Wild Dog." Morell, Virginia. International Wildlife. 26(3): p35.
  16. ^ An example of this can be seen in this video: http://www.youtube.com/watch?v=3A-ZdJjG5Vg
  17. ^ "Hope Rises for Africa's Wild Dog." Morell, Virginia. International Wildlife. 26(3):p35.
  18. ^ "Hope Rises for Africa's Wild Dog." Morell, Virginia. International Wildlife. 26(3):p32.
  19. ^ Creel, Scott & Creel, Nancy (2002). Krebs, J. and Clutton-Brock, T.. ed. The African Wild Dog: Behavior, Ecology, and Conservation. Princeton: Princeton University Press. p. 59. ISBN 0-69101-654-2. 
  20. ^ Borrell, Brendan (2009-08-19). "Endangered in South Africa: Those Doggone Conservationists". Slate. http://www.slate.com/id/2225607/entry/2225663/. 
  21. ^ C. Michael Hogan. 2009
  22. ^ Adam Hartstone-Rose, Lars Werdelin, Darryl J. De Ruiter, Lee R. Berger and Steven E. Churchill, "The Plio-pleistocene Ancestor of Wild Dogs, Lycaon sekowei N. Sp, Journal of Paleontology; March 2010; 84(2), pp. 299-308; DOI: 10.1666/09-124.1 [1]
  23. ^ Martínez-Navarro, B., and L. Rook (2003). "Gradual evolution in the African hunting dog lineage: systematic implications". Comptes Rendus Palevol 2: 695–702. doi:10.1016/j.crpv.2003.06.002. 
  24. ^ Wozencraft, W. Christopher (16 November 2005). "Order Carnivora (pp. 532-628)". In Wilson, Don E., and Reeder, DeeAnn M., eds. Mammal Species of the World: A Taxonomic and Geographic Reference (3rd ed.). Baltimore: Johns Hopkins University Press, 2 vols. (2142 pp.). ISBN 978-0-8018-8221-0. OCLC 62265494. http://www.bucknell.edu/msw3. 
  25. ^ Dog from North Side shelter becomes surrogate for African painted pups, Pittsburgh Post-Gazette, November 4, 2009

Bibliography

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