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Morphology

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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.

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Mulheisen, M.; C. Allen and C. Allen 2002. "Lycaon pictus" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Lycaon_pictus.html
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Michael Mulheisen, University of Michigan-Ann Arbor
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Life Expectancy

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Average lifespan
Status: wild:
10.0 years.

Average lifespan
Status: wild:
11.0 years.

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Mulheisen, M.; C. Allen and C. Allen 2002. "Lycaon pictus" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Lycaon_pictus.html
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Michael Mulheisen, University of Michigan-Ann Arbor
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Habitat

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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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Mulheisen, M.; C. Allen and C. Allen 2002. "Lycaon pictus" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Lycaon_pictus.html
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Michael Mulheisen, University of Michigan-Ann Arbor
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Distribution

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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 )

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Mulheisen, M.; C. Allen and C. Allen 2002. "Lycaon pictus" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Lycaon_pictus.html
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Michael Mulheisen, University of Michigan-Ann Arbor
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Trophic Strategy

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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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Mulheisen, M.; C. Allen and C. Allen 2002. "Lycaon pictus" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Lycaon_pictus.html
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Michael Mulheisen, University of Michigan-Ann Arbor
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Benefits

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Lycaon pictus occasionally kills livestock and important game animals.

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Mulheisen, M.; C. Allen and C. Allen 2002. "Lycaon pictus" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Lycaon_pictus.html
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Michael Mulheisen, University of Michigan-Ann Arbor
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Conservation Status

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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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Mulheisen, M.; C. Allen and C. Allen 2002. "Lycaon pictus" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Lycaon_pictus.html
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Michael Mulheisen, University of Michigan-Ann Arbor
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Behavior

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Perception Channels: tactile ; chemical

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Mulheisen, M.; C. Allen and C. Allen 2002. "Lycaon pictus" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Lycaon_pictus.html
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Michael Mulheisen, University of Michigan-Ann Arbor
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Reproduction

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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); sexual ; viviparous

Average birth mass: 318 g.

Average number of offspring: 8.

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

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Mulheisen, M.; C. Allen and C. Allen 2002. "Lycaon pictus" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Lycaon_pictus.html
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Michael Mulheisen, University of Michigan-Ann Arbor
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Crystal Allen, University of Michigan-Ann Arbor
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Biology

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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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Conservation

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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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Description

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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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Habitat

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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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Range

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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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Status

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

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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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Brief Summary

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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).

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MammalMAP: African Wild dogs

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African Wild dogs orLycaon 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 aunique 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.

TheIUCNhas 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 MammalMAPvirtual museumorblog.

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Species Abstract

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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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C.Michael Hogan. 2009. ''Painted Hunting Dog Lycaon pictus. iGoTerra ed. N. Stromberg
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African wild dog

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The African wild dog (Lycaon pictus), also known as the painted dog or Cape hunting dog, is a wild canine native to sub-Saharan Africa. It is the largest wild canine in Africa, and the only extant member of the genus Lycaon, which is distinguished from Canis by dentition highly specialised for a hypercarnivorous diet, and by a lack of dewclaws. It's estimated that there are around 6,600 adults (including 1,400 mature individuals) living in 39 subpopulations that are all threatened by habitat fragmentation, human persecution, and outbreaks of disease. As the largest subpopulation probably comprises fewer than 250 individuals, the African wild dog has been listed as endangered on the IUCN Red List since 1990.[2]

The species is a specialised diurnal hunter of ungulates, which it captures by using its stamina and cooperative hunting to exhaust them. Its natural competitors are lions and spotted hyenas: the former will kill the dogs where possible, whilst hyenas are frequent kleptoparasites.[4]

Like other canids, the African wild dog regurgitates food for its young, but also extends this action to adults, as a central part of the pack's social unit.[5][4][6] The young have the privilege to feed first on carcasses.

The African wild dog has been respected in several hunter-gatherer societies, particularly those of the San people and Prehistoric Egypt.

Etymology and naming

The English language has several names for the African wild dog, including African hunting dog, Cape hunting dog,[7] painted hunting dog,[8] painted dog,[9] painted wolf,[10] and painted lycaon.[11] Though the name "African wild dog" is widely used,[12] "wild dog" is thought by conservation groups to have negative connotations that could be detrimental to its image; one organisation promotes the name "painted wolf",[13][14][15] while the name "painted dog" has been found to be the most likely to counteract negative perceptions.[16]

Taxonomic and evolutionary history

Taxonomy

Phylogenetic tree of the wolf-like canids with timing in millions of years[a] Caninae 3.5 Ma 3.0 2.5 2.0 0.96 0.6 0.38

Domestic dog Tibetan mastiff (white background).jpg

Gray wolf Dogs, jackals, wolves, and foxes (Plate I).jpg

Coyote Dogs, jackals, wolves, and foxes (Plate IX).jpg

African wolf Dogs, jackals, wolves, and foxes (Plate XI).jpg

Golden jackal Dogs, jackals, wolves, and foxes (Plate X).jpg

Ethiopian wolf Dogs, jackals, wolves, and foxes (Plate VI).jpg

Dhole Dogs, jackals, wolves, and foxes (Plate XLI).jpg

African wild dog Dogs, jackals, wolves, and foxes (Plate XLIV).jpg

2.6

Side-striped jackal Dogs, jackals, wolves, and foxes (Plate XIII).jpg

Black-backed jackal Dogs, jackals, wolves, and foxes (Plate XII).jpg

The earliest written reference for the species appears to be from Oppian, who wrote of the thoa, a hybrid between the wolf and leopard, which resembles the former in shape and the latter in colour. Solinus's Collea rerum memorabilium from the third century AD describes a multicoloured wolf-like animal with a mane native to Ethiopia.[11]

The African wild dog was scientifically described in 1820 by Coenraad Jacob Temminck, after examining a specimen from the coast of Mozambique. He named the animal Hyaena picta, erroneously classifying it as a species of hyena. It was later recognised as a canid by Joshua Brookes in 1827, and renamed Lycaon tricolor. The root word of Lycaon is the Greek λυκαίος (lykaios), meaning "wolf-like". The specific epithet pictus (Latin for "painted"), which derived from the original picta, was later returned to it, in conformity with the International Rules on Taxonomic Nomenclature.[17]

Paleontologist George G. Simpson placed the African wild dog, the dhole, and the bush dog together in the subfamily Simocyoninae on the basis of all three species having similarly trenchant carnassials. This grouping was disputed by Juliet Clutton-Brock, who argued that, other than dentition, too many differences exist between the three species to warrant classifying them in a single subfamily.[18]

Evolution

Artistic reconstruction by Mauricio Antón of Xenocyon, a possible ancestral genus

The African wild dog possesses the most specialized adaptations among the canids for coat colour, diet, and for pursuing its prey through its cursorial (running) ability. It has a graceful skeleton, and the loss of the first digit on its forefeet increases its stride and speed. This adaptation allows it to pursue prey across open plains for long distances. The teeth are generally carnassial-shaped, and its premolars are the largest relative to body size of any living carnivoran with the exception of the spotted hyena. On the lower carnassials (first lower molars), the talonid has evolved to become a cutting blade for flesh-slicing, with a reduction or loss of the post-carnassial molars. This adaptation also occurs in two other hypercarnivores – the dhole and the bush dog. The African wild dog exhibits one of the most varied coat colours among the mammals. Individuals differ in patterns and colours, indicating a diversity of the underlying genes. The purpose of these coat patterns may be an adaptation for communication, concealment, or temperature regulation. In 2019, a study indicated that the lycaon lineage diverged from Cuon and Canis 1.7 million years ago through this suite of adaptations, and these occurred at the same time as large ungulates (its prey) diversified.[19]

The oldest African wild dog fossil dates back to 200,000 years ago and was found in HaYonim Cave, Israel.[20][1] The evolution of the African wild dog is poorly understood due to the scarcity of fossil finds. Some authors consider the extinct Canis subgenus Xenocyon as ancestral to both the genus Lycaon and the genus Cuon,[21][22][23][24]: p149  which lived throughout Eurasia and Africa from the Early Pleistocene to the early Middle Pleistocene. Others propose that Xenocyon should be reclassified as Lycaon.[1] The species Canis (Xenocyon) falconeri shared the African wild dog's absent first metacarpal (dewclaw), though its dentition was still relatively unspecialised.[1] This connection was rejected by one author because C. (X.) falconeri lacks metacarpal, which is a poor indication of phylogenetic closeness to the African wild dog, and the dentition was too different to imply ancestry.[25]

Another ancestral candidate is the Plio-Pleistocene L. sekowei of South Africa on the basis of distinct accessory cusps on its premolars and anterior accessory cuspids on its lower premolars. These adaptions are found only in Lycaon among living canids, which shows the same adaptations to a hypercarnivorous diet. L. sekowei had not yet lost the first metacarpal absent in L. pictus and was more robust than the modern species, having 10% larger teeth.[25]

The African wild dog genetically diverged from other canid lineages between 1.74 to 1.7 million years ago and is thought to be isolated from gene transfer with other canid species.[26]

Admixture with the dhole

Fossil of Lycaon sekowei, a possible ancestor of the modern African wild dog

In 2018, whole genome sequencing was used to compare the dhole (Cuon alpinus) with the African wild dog. There was strong evidence of ancient genetic admixture between the two species. Today, their ranges are remote from each other; however, during the Pleistocene era the dhole could be found as far west as Europe. The study proposes that the dhole's distribution may have once included the Middle East, from where it may have admixed with the African wild dog in North Africa. However, there is no evidence of the dhole having existed in the Middle East or North Africa.[27]

Subspecies

As of 2005, five subspecies are recognised by MSW3:[28]

Although the species is genetically diverse, these subspecific designations are not universally accepted. East African and Southern African wild dog populations were once thought to be genetically distinct, based on a small number of samples. More recent studies with a larger number of samples showed that extensive intermixing has occurred between East African and Southern African populations in the past. Some unique nuclear and mitochondrial alleles are found in Southern African and northeastern African populations, with a transition zone encompassing Botswana, Zimbabwe and southeastern Tanzania between the two. The West African wild dog population may possess a unique haplotype, thus possibly constituting a truly distinct subspecies.[34] The original Serengeti and Maasai Mara population of painted dogs is known to have possessed a unique genotypes, but these genotypes may be extinct.[35]

Description

Closeup of an African wild dog in Kruger National Park
African wild dog skull (left) compared with that of the grey wolf (right): Note the former's shorter muzzle and fewer molars.

The African wild dog is the bulkiest and most solidly built of African canids.[36] The species stands 60 to 75 cm (24 to 30 in) at the shoulders, measures 71 to 112 cm (28 to 44 in) in head-and-body length and has a tail length of 29 to 41 cm (11 to 16 in). Adults have a weight range of 18 to 36 kg (40 to 79 lb). On average, dogs from East Africa weigh around 20–25 kg (44–55 lb) while in southern Africa, males reportedly weighed a mean of 32.7 kg (72 lb) and females a mean of 24.5 kg (54 lb). By body mass, they are only outsized amongst other extant canids by the gray wolf species complex.[29][37][38] Females are usually 3–7% smaller than males. Compared to members of the genus Canis, the African wild dog is comparatively lean and tall, with outsized ears and lacking dewclaws. The middle two toepads are usually fused. Its dentition differs from that of Canis by the degeneration of the last lower molar, the narrowness of the canines and proportionately large premolars, which are the largest relative to body size of any carnivore other than hyenas.[39] The heel of the lower carnassial M1 is crested with a single, blade-like cusp, which enhances the shearing capacity of the teeth, thus the speed at which prey can be consumed. This feature, termed "trenchant heel", is shared with two other canids: the Asian dhole and the South American bush dog.[7] The skull is relatively shorter and broader than those of other canids.[36]

The fur of the African wild dog differs significantly from that of other canids, consisting entirely of stiff bristle-hairs with no underfur.[36] Adults gradually lose their fur as it ages, with older individuals being almost naked.[40] Colour variation is extreme, and may serve in visual identification, as African wild dogs can recognise each other at distances of 50–100 m (160–330 ft).[39] Some geographic variation is seen in coat colour, with northeastern African specimens tending to be predominantly black with small white and yellow patches, while southern African ones are more brightly coloured, sporting a mix of brown, black and white coats.[7] Much of the species' coat patterning occurs on the trunk and legs. Little variation in facial markings occurs, with the muzzle being black, gradually shading into brown on the cheeks and forehead. A black line extends up the forehead, turning blackish-brown on the back of the ears. A few specimens sport a brown teardrop-shaped mark below the eyes. The back of the head and neck are either brown or yellow. A white patch occasionally occurs behind the fore legs, with some specimens having completely white fore legs, chests and throats. The tail is usually white at the tip, black in the middle and brown at the base. Some specimens lack the white tip entirely, or may have black fur below the white tip. These coat patterns can be asymmetrical, with the left side of the body often having different markings from that of the right.[39]

Genetics

The African wild dog has 78 chromosomes, the same number that species of the genus Canis have. However, it is not possible to be bred with other canids.[41][42]

Distribution and habitat

African wild dogs once ranged across much of sub-Saharan Africa, being absent only in the driest deserts and lowland forests. The species has been largely exterminated in North and West Africa, and its population has greatly reduced in Central Africa and northeast Africa. The majority of the species' population now occurs in Southern Africa and southern East Africa; more specifically in countries such as Botswana, Namibia, and Zimbabwe. However, it is hard to track where they are and how many there are because of the loss of habitat.[2] A stable population comprising more than 370 individuals is present in South Africa, particularly the Kruger National Park.[43]

The African wild dog inhabits mostly savannas and arid zones, generally avoiding forested areas.[29] This preference is likely linked to the animal's hunting habits, which require open areas that does not obstruct vision or impede pursuit.[36] Nevertheless, it will travel through scrub, woodland and montane areas in pursuit of prey. Forest-dwelling populations have been identified, including one in the Harenna Forest, a wet montane forest up to 2,400 m (7,900 ft) in altitude in the Bale Mountains of Ethiopia.[44] At least one record exists of a pack being sighted on the summit of Mount Kilimanjaro.[29] In Zimbabwe, the species has been recorded at altitudes of 1,800 m (5,900 ft).[12] In Ethiopia, they have been found at high altitudes; several live wild dog packs have been sighted at altitudes of 1,900 to 2,800 m (6,200 to 9,200 ft), and a dead individual was found in June 1995 at 4,050 m (13,290 ft) on the Sanetti Plateau.[45]

The species is very rare in North Africa, and the remaining populations may be of high conservation value, as they are likely to be genetically distinct from other African wild dog populations. The African wild dog is mostly absent in West Africa, with the only potentially viable population occurring in Senegal's Niokolo-Koba National Park. African wild dogs are occasionally sighted in other parts of Senegal, Guinea and Mali. In Central Africa, the species is extinct in Gabon, the Democratic Republic of Congo and the Republic of Congo. The only viable populations occur in the Central African Republic, Chad and especially Cameroon. The African wild dog is distributed throughout patches in East Africa, having been eradicated in Uganda and much of Kenya.[46]

Behaviour and ecology

Social and reproductive behaviour

Wild dogs with springbok kill in Tswalu Kalahari Reserve
Play fighting after a kill in Tswalu Kalahari Reserve

The African wild dog have strong social bonds, stronger than those of sympatric lions and spotted hyenas; thus, solitary living and hunting are extremely rare in the species.[47] It lives in permanent packs consisting of two to 27 adults and yearling pups. The typical pack size in the Kruger National Park and the Maasai Mara is four or five adults, while packs in Moremi and Selous contain eight or nine. However, larger packs have been observed and temporary aggregations of hundreds of individuals may have gathered in response to the seasonal migration of vast springbok herds in Southern Africa.[48] Males and females have separate dominance hierarchies, with the latter usually being led by the oldest female. Males may be led by the oldest male, but these can be supplanted by younger specimens; thus, some packs may contain elderly male former pack leaders. The dominant pair typically monopolises breeding.[39] The species differs from most other social species in that males remain in the natal pack, while females disperse (a pattern also found in primates such as gorillas, chimpanzees, and red colobuses). Furthermore, males in any given pack tend to outnumber females 3:1.[29] Dispersing females join other packs and evict some of the resident females related to the other pack members, thus preventing inbreeding and allowing the evicted individuals to find new packs of their own and breed.[39] Males rarely disperse, and when they do, they are invariably rejected by other packs already containing males.[29] Although arguably the most social canid, the species lacks the elaborate facial expressions and body language found in the gray wolf, likely because of the African wild dog's less hierarchical social structure. Furthermore, while elaborate facial expressions are important for wolves in re-establishing bonds after long periods of separation from their family groups, they are not as necessary to African wild dogs, which remain together for much longer periods.[18]

African wild dog populations in East Africa appear to have no fixed breeding season, whereas those in Southern Africa usually breed during the April–July period.[47] During estrus, the female is closely accompanied by a single male, which keeps other members of the same sex at bay.[29] The copulatory tie characteristic of mating in most canids has been reported to be absent[49] or very brief (less than one minute)[50] in African wild dog, possibly an adaptation to the prevalence of larger predators in its environment.[51] The gestation period lasts 69–73 days, with the interval between each pregnancy being 12–14 months typically. The African wild dog produces more pups than any other canid, with litters containing around six to 16 pups, with an average of 10, thus indicating that a single female can produce enough young to form a new pack every year. Because the amount of food necessary to feed more than two litters would be impossible to acquire by the average pack, breeding is strictly limited to the dominant female, which may kill the pups of subordinates. After giving birth, the mother stays close to the pups in the den, while the rest of the pack hunts. She typically drives away pack members approaching the pups until the latter are old enough to eat solid food at three to four weeks of age. The pups leave the den around the age of three weeks and are suckled outside. The pups are weaned at the age of five weeks, when they are fed regurgitated meat by the other pack members. By seven weeks, the pups begin to take on an adult appearance, with noticeable lengthening in the legs, muzzle, and ears. Once the pups reach the age of eight to 10 weeks, the pack abandons the den and the young follow the adults during hunts. The youngest pack members are permitted to eat first on kills, a privilege which ends once they become yearlings.[29] African wild dogs have an average lifespan of about 10 to 11 years in the wild.[52]

When separated from the pack, an African wild dog becomes depressed and can die as a result of broken heart syndrome.[53][54]

Male/female ratio

Packs of African wild dogs have a high ratio of males to females. This is a consequence of the males mostly staying with the pack whilst female offspring disperse and is supported by a changing sex-ratio in consecutive litters. Those born to maiden females contain a higher proportion of males, second litters are half and half and subsequent litters biased towards females with this trend increasing as females get older. As a result, the earlier litters provide stable hunters whilst the higher ratio of dispersals amongst the females stops a pack from getting too big.[55][5]

Sneeze communication and "voting"

African wild dog pack in Kruger National Park

Populations in the Okavango Delta have been observed "rallying" before setting out to hunt. Not every rally results in a departure, but departure becomes more likely when more individual dogs "sneeze". These sneezes are characterized by a short, sharp exhale through the nostrils.[56] When members of dominant mating pairs sneeze first, the group is much more likely to depart. If a dominant dog initiates, around three sneezes guarantee departure. When less dominant dogs sneeze first, if enough others also sneeze (about 10), then the group will go hunting. Researchers assert that wild dogs in Botswana, "use a specific vocalization (the sneeze) along with a variable quorum response mechanism in the decision-making process [to go hunting at a particular moment]".[57]

Inbreeding avoidance

Because the African wild dog largely exists in fragmented, small populations, its existence is endangered. Inbreeding avoidance by mate selection is a characteristic of the species and has important potential consequences for population persistence.[58] Inbreeding is rare within natal packs. Inbreeding may have been selected against evolutionarily because it leads to the expression of recessive deleterious alleles.[59] Computer simulations indicate that all populations continuing to avoid incestuous mating will become extinct within 100 years due to the unavailability of unrelated mates.[58] Thus, the impact of reduced numbers of suitable unrelated mates will likely have a severe demographic impact on the future viability of small wild dog populations.[58]

Hunting and diet

African wild dog pack consuming a blue wildebeest in Madikwe Game Reserve, South Africa

The African wild dog is a specialised pack hunter of common medium-sized antelopes.[60] It and the cheetah are the only primarily diurnal African large predators.[29] The African wild dog hunts by approaching prey silently, then chasing it in a pursuit clocking at up to 66 km/h (41 mph) for 10–60 minutes.[48] The average chase covers some 2 km (1.2 mi), during which the prey animal, if large, is repeatedly bitten on the legs, belly, and rump until it stops running, while smaller prey is simply pulled down and torn apart.[5]

African wild dogs adjust their hunting strategy to the particular prey species. They will rush at wildebeest to panic the herd and isolate a vulnerable individual, but pursue territorial antelope species (which defend themselves by running in wide circles) by cutting across the arc to foil their escape. Medium-sized prey is often killed in 2–5 minutes, whereas larger prey such as wildebeest may take half an hour to pull down. Male wild dogs usually perform the task of grabbing dangerous prey, such as warthogs, by the nose.[61] A species-wide study showed that by preference, where available, five prey species were the most regularly selected, namely the greater kudu, Thomson's gazelle, impala, Cape bushbuck and blue wildebeest.[60][62] More specifically, in East Africa, its most common prey is the Thomson's gazelle, while in Central and Southern Africa, it targets impala, reedbuck, kob, lechwe and springbok.[29] and smaller prey such as common duiker, dik-dik, hares, spring hares, insects and cane rats.[47][63] Staple prey sizes are usually between 15 and 200 kg (33 and 441 lb), though some local studies put upper prey sizes as variously 90 to 135 kg (198 to 298 lb). In the case of larger species such as kudu and wildebeest, calves are largely but not exclusively targeted.[60][64][65] However, certain packs in the Serengeti specialized in hunting adult plains zebras weighing up to 240 kg (530 lb) quite frequently.[66] Another study claimed that some prey taken by wild dogs could weigh up to 289 kg (637 lb).[67] One pack was recorded to occasionally prey on bat-eared foxes, rolling on the carcasses before eating them. African wild dogs rarely scavenge, but have on occasion been observed to appropriate carcasses from spotted hyenas, leopards, cheetahs, lions, and animals caught in snares.[12]

Hunting success varies with prey type, vegetation cover and pack size, but African wild dogs tend to be very successful: often more than 60% of their chases end in a kill, sometimes up to 90%.[68] An analysis of 1,119 chases by a pack of six Okavango wild dogs showed that most were short distance uncoordinated chases, and the individual kill rate was only 15.5 percent. Because kills are shared, each dog enjoyed an efficient benefit–cost ratio.[69][70]

Small prey such as rodents, hares and birds are hunted singly, with dangerous prey such as cane rats and Old World porcupines being killed with a quick and well-placed bite to avoid injury. Small prey is eaten entirely, while large animals are stripped of their meat and organs, leaving the skin, head, and skeleton intact.[47][71] The African wild dog is a fast eater, with a pack being able to consume a Thomson's gazelle in 15 minutes. In the wild, the species' consumption is 1.2–5.9 kg (2.6–13.0 lb) per African wild dog a day, with one pack of 17–43 individuals in East Africa having been recorded to kill three animals per day on average.[12]

Unlike most social predators, African wild dogs will regurgitate food for other adults as well as young family members.[47] Pups old enough to eat solid food are given first priority at kills, eating even before the dominant pair; subordinate adult dogs help feed and protect the pups.[72]

Enemies and competitors

Pack confronting a spotted hyena in Sabi Sand Game Reserve

Lions dominate African wild dogs and are a major source of mortality for both adults and pups.[73] Population densities are usually low in areas where lions are more abundant.[74] One pack reintroduced into Etosha National Park were wiped out by lions. A population crash in lions in the Ngorongoro Conservation Area during the 1960s resulted in an increase in African wild dog sightings, only for their numbers to decline once the lions recovered.[73] As with other large predators killed by lion prides, the dogs are usually killed and left uneaten by the lions, indicating the competitive rather than predatory nature of the lions' dominance.[75][76] However, a few cases have been reported of old and wounded lions falling prey to African wild dogs.[77][78] On occasion, packs of wild dogs have been observed defending pack members attacked by single lions, sometimes successfully. One pack in the Okavango in March 2016 was photographed by safari guides waging "an incredible fight" against a lioness that attacked a subadult dog at an impala kill, which forced the lioness to retreat, although the subadult dog died. A pack of four wild dogs was observed furiously defending an old adult male dog from a male lion that attacked it at a kill; the dog survived and rejoined the pack.[79]

African wild dogs commonly lose their kills to larger predators.[80] Spotted hyenas are important kleptoparasites[73] and follow packs of African wild dogs to appropriate their kills. They typically inspect areas where wild dogs have rested and eat any food remains they find. When approaching wild dogs at a kill, solitary hyenas approach cautiously and attempt to take off with a piece of meat unnoticed, though they may be mobbed in the attempt. When operating in groups, spotted hyenas are more successful in pirating African wild dog kills, though the latter's greater tendency to assist each other puts them at an advantage against spotted hyenas, which rarely work cooperatively. Cases of African wild dogs scavenging from spotted hyenas are rare. Although African wild dog packs can easily repel solitary hyenas, on the whole, the relationship between the two species is a one-sided benefit for the hyenas,[81] with African wild dog densities being negatively correlated with high hyena populations.[82] Beyond piracy, cases of interspecific killing of African wild dogs by spotted hyenas are documented.[83] African wild dogs are apex predators, only fatally losing contests to larger social carnivores.[84] When briefly unprotected, wild dog pups may occasionally be vulnerable to large eagles, such as the martial eagle, when they venture out of their dens.[85]

Threats and conservation

The African wild dog is primarily threatened by habitat fragmentation, which results to human–wildlife conflict, transmission of infectious diseases and high mortality rates.[2] Surveys in the Central African Republic's Chinko area revealed that the African wild dog population decreased from 160 individuals in 2012 to 26 individuals in 2017. At the same time, transhumant pastoralists from the border area with Sudan moved in the area with their livestock.[84]

The African Wild Dog Conservancy, a non-profit, 501(c)(3), non-governmental organization, began working in 2003 to conserve the African wild dog in northeastern and coastal Kenya, a convergence zone of two biodiversity hotspots. This area largely consists of community lands inhabited by pastoralists. With the help of local people, a pilot study was launched confirming the presence of a population of wild dogs largely unknown to conservationists.[86] Over the next 16 years, local ecological knowledge revealed this area to be a significant refuge for wild dogs and an important wildlife corridor connecting Kenya’s Tsavo National Parks with the Horn of Africa in an increasingly human-dominated landscape. This project has been identified as a wild dog conservation priority by the IUCN/SSC Canid Specialist Group.[87][88]

In culture

Ancient Egypt

Cosmetic palette from the Naqada III period depicting African wild dogs, Louvre.

Depictions of African wild dogs are prominent on cosmetic palettes and other objects from Egypt's predynastic period, likely symbolising order over chaos and the transition between the wild and the domestic dog. Predynastic hunters may have identified with the African wild dog, as the Hunters Palette shows them wearing the animals' tails on their belts. By the dynastic period, African wild dog illustrations became much less represented, and the animal's symbolic role was largely taken over by the wolf.[89][90]

Ethiopia

According to Enno Littmann, the people of Ethiopia's Tigray Region believed that injuring a wild dog with a spear would result in the animal dipping its tail in its wounds and flicking the blood at its assailant, causing instant death. For this reason, Tigrean shepherds used to repel wild dog attacks with pebbles rather than with edged weapons.[91]

San people

The African wild dog also plays a prominent role in the mythology of Southern Africa's San people. In one story, the wild dog is indirectly linked to the origin of death, as the hare is cursed by the moon to be forever hunted by African wild dogs after the hare rebuffs the moon's promise to allow all living things to be reborn after death.[92] Another story has the god Cagn taking revenge on the other gods by sending a group of men transformed into African wild dogs to attack them, though who won the battle is never revealed.[93] The San of Botswana see the African wild dog as the ultimate hunter and traditionally believe that shamans and medicine men can transform themselves into wild dogs. Some San hunters will smear African wild dog bodily fluids on their feet before a hunt, believing that doing so will give them the animal's boldness and agility. Nevertheless, the species does not figure prominently in San rock art, with the only notable example being a frieze in Mount Erongo showing a pack hunting two antelopes.[93]

Ndebele

The Ndebele have a story explaining why the African wild dog hunts in packs: in the beginning, when the first wild dog's wife was sick, the other animals were concerned. An impala went to hare, who was a medicine man. Hare gave Impala a calabash of medicine, warning him not to turn back on the way to Wild Dog's den. Impala was startled by the scent of a leopard and turned back, spilling the medicine. A zebra then went to Hare, who gave him the same medicine along with the same advice. On the way, Zebra turned back when he saw a black mamba, thus breaking the gourd. A moment later, a terrible howling is heard: Wild Dog's wife had died. Wild Dog went outside and saw Zebra standing over the broken gourd of medicine, so Wild Dog and his family chased Zebra and tore him to shreds. To this day, African wild dogs hunt zebras and impalas as revenge for their failure to deliver the medicine which could have saved Wild Dog's wife.[94]

In media

Documentary

  • A Wild Dog's Tale (2013), a single painted dog (named Solo by researchers) befriends hyenas and jackals in Okavango, hunting together. Solo feeds and cares for jackal pups.[95][96]
  • The Pale Pack, Savage Kingdom, Season 1 (2016), was the story of Botswana African wild dog pack leaders Teemana and Molao written and directed by Brad Bestelink, and narrated by Charles Dance premiered on National Geographic.[97][98]
  • Dynasties (2018 TV series), episode 4, Produced by Nick Lyon: Tait is the elderly matriarch of a pack of painted wolves in Zimbabwe's Mana Pools National Park. Her pack is driven out of their territory by Tait's daughter, Blacktip, the matriarch of a rival pack in need of more space for their large family of 32. Their combined territory also shrunk over Tait's lifetime due to the expansion of human, hyena and lion territories. Tait leads her family into the territory of a lion pride in the midst of a drought, with Blacktip's pack in an eight month long pursuit. When Tait died, the pack was observed performing a rare "singing", the purpose of which is unclear.[99]

See also

Explanatory notes

  1. ^ For a full set of supporting references refer to the note (a) in the phylotree at Evolution of the wolf#Wolf-like canids

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African wild dog: Brief Summary

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The African wild dog (Lycaon pictus), also known as the painted dog or Cape hunting dog, is a wild canine native to sub-Saharan Africa. It is the largest wild canine in Africa, and the only extant member of the genus Lycaon, which is distinguished from Canis by dentition highly specialised for a hypercarnivorous diet, and by a lack of dewclaws. It's estimated that there are around 6,600 adults (including 1,400 mature individuals) living in 39 subpopulations that are all threatened by habitat fragmentation, human persecution, and outbreaks of disease. As the largest subpopulation probably comprises fewer than 250 individuals, the African wild dog has been listed as endangered on the IUCN Red List since 1990.

The species is a specialised diurnal hunter of ungulates, which it captures by using its stamina and cooperative hunting to exhaust them. Its natural competitors are lions and spotted hyenas: the former will kill the dogs where possible, whilst hyenas are frequent kleptoparasites.

Like other canids, the African wild dog regurgitates food for its young, but also extends this action to adults, as a central part of the pack's social unit. The young have the privilege to feed first on carcasses.

The African wild dog has been respected in several hunter-gatherer societies, particularly those of the San people and Prehistoric Egypt.

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