African wild dog (Lycaon pictus pictus) with springbok.jpg

The African wild dog (''Lycaon pictus''), also called the painted dog or Cape hunting dog, is a wild canine which is a native species 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 is estimated that about 6,600 adults (including 1,400 mature individuals) live 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 antelopes, which it catches by chasing them to exhaustion. Its natural enemies 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 life. The young are allowed 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.

Naming

The English language has several names for the African wild dog, including African hunting dog, Cape hunting dog, painted hunting dog, painted dog, and painted lycaon. One conservation organisation is promoting the name 'painted wolf' as a way of rebranding the species, as wild dog has several negative connotations that could be detrimental to its image. Nevertheless, the name "African wild dog" is still widely used,
However, the name "painted dog" was found to be the most likely to counteract negative perceptions of the species.

Taxonomic and evolutionary history

Taxonomy

The earliest written reference to 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.Smith, C. H. (1839).
Dogs
', W.H. Lizars, Edinburgh, p. 261–69

The African wild dog was first described scientifically 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.Bothma, J. du P. & Walker, C. (1999). ''Larger Carnivores of the African Savannas'', Springer, pp. 130–157,

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.

Evolution

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 possesses 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 except for 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.

The oldest ''L. pictus'' fossil dates back to 200,000 years ago and was found in HaYonim Cave, Israel. 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'', which lived throughout Eurasia and Africa from the Early Pleistocene to the early Middle Pleistocene. Others propose that ''Xenocyon'' should be reclassified as ''Lycaon''. The species ''Canis'' (''Xenocyon'') ''falconeri'' shared the African wild dog's absent first metacarpal ( dewclaw), though its dentition was still relatively unspecialised. 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.

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.

The African wild dog genetically diverged from other canid lineages between and is thought to be isolated from gene transfer with other canid species.

Admixture with the dhole

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. 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.

Subspecies

, five subspecies are recognised by MSW3:

Nevertheless, 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. The original Serengeti and Maasai Mara population of painted dogs is known to have possessed a unique genotypes, but these genotypes may be extinct.

Physical description

The African wild dog is the bulkiest and most solidly built of African canids.Rosevear, D. R. (1974).
The carnivores of West Africa
'. London : Trustees of the British Museum (Natural History). pp. 75–91. . The species stands in shoulder height, measures in head-and-body length and has a tail length of . Body weight of adults range from . On average, dogs from East Africa weigh around while in southern Africa, males reportedly weighed a mean of and females a mean of . By body mass, they are only outsized amongst other extant canids by the gray wolf species complex.Estes, R. (1992). ''The behavior guide to African mammals: including hoofed mammals, carnivores, primates''. University of California Press. pp. 410–419. . Females are generally 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 also 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. 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. The skull is relatively shorter and broader than those of other canids.

The fur of the African wild dog differs significantly from that of other canids, consisting entirely of stiff bristle-hairs with no underfur. It gradually loses its fur as it ages, with older individuals being almost naked. Colour variation is extreme, and may serve in visual identification, as African wild dogs can recognise each other at distances of . 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. 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.

Behaviour

Social and reproductive behaviour

The African wild dog has very strong social bonds, stronger than those of sympatric lions and spotted hyenas; thus, solitary living and hunting are extremely rare in the species.Kingdon, Jonathan (1988). ''East African mammals: an atlas of evolution in Africa, Volume 3, Part 1''. University of Chicago Press. pp. 36–53. . It lives in permanent packs consisting of two to 27 adults and yearling pups. The typical pack size in 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.Nowak, Ronald M. (2005). ''Walker's Carnivores of the World''. Baltimore: Johns Hopkins Press. p. 112. . 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. 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. 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. Males rarely disperse, and when they do, they are invariably rejected by other packs already containing males. 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.

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. During estrus, the female is closely accompanied by a single male, which keeps other members of the same sex at bay. The copulatory tie characteristic of mating in most canids has been reported to be absent or very brief (less than one minute) in African wild dog, possibly an adaptation to the prevalence of larger predators in its environment. 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.

Ratio male/female

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.

Sneeze communication and "voting"

African wild dog populations in the Okavango Delta have been observed "rallying" before they set 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. 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 o go hunting at a particular moment.

Inbreeding avoidance

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

Hunting and feeding behaviour

The African wild dog is a specialised pack hunter of common medium-sized antelopes.. It and the cheetah are the only primarily diurnal African large predators. The African wild dog hunts by approaching prey silently, then chasing it in a pursuit clocking at up to for 10–60 minutes. The average chase covers some , 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.

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.

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%. Despite their smaller size, they are much more consistently successful than lion (27–30%) and hyena (25–30%), but African wild dogs commonly lose their kills to these two large predators. 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.

Small prey such as rodents, hares and birds are hunted singly, with dangerous prey such as cane rats and 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. 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 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.

Unlike most social predators, African wild dogs will regurgitate food for other adults as well as young family members. 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.

Ecology

Habitat

The African wild dog is mostly found in savanna and arid zones, generally avoiding forested areas. This preference is likely linked to the animal's hunting habits, which require open areas that do not obstruct vision or impede pursuit. Nevertheless, it will travel through scrub, woodland and montane areas in pursuit of prey. Forest-dwelling populations of African wild dogs have been identified, including one in the Harenna Forest, a wet montane forest up to in altitude in the Bale Mountains of Ethiopia. At least one record exists of a pack being sighted on the summit of Mount Kilimanjaro. In Zimbabwe, the species has been recorded at altitudes of . In Ethiopia, this species has been found at great altitudes; several live wild dog packs have been sighted at altitudes of from 1,900 to 2,800 m, and a dead individual was found in June 1995 at on the Sanetti Plateau.

Diet

A species-wide study showed that by preference, where available, five species were the most regularly selected prey, namely the greater kudu, Thomson's gazelle, impala, Cape bushbuck and blue wildebeest. More specifically, in East Africa, its most common prey is Thomson's gazelle, while in Central and Southern Africa, it targets impala, reedbuck, kob, lechwe and springbok. and smaller prey such as dik-dik, hares, spring hares, insects and cane rats. Staple prey sizes are usually between , though some local studies put upper prey sizes as variously . In the case of larger species such as kudu and wildebeest, calves are largely but not exclusively targeted. However, certain packs in the Serengeti specialized in hunting adult plains zebras weighing up to quite frequently. Another study claimed that some prey taken by wild dogs could weigh up to . 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.

Enemies and competitors

Lions dominate African wild dogs and are a major source of mortality for both adults and pups. Population densities of African wild dogs are low in areas where lions are more abundant. One pack reintroduced into Etosha National Park was destroyed by lions. A population crash in lions in the Ngorongoro Crater during the 1960s resulted in an increase in African wild dog sightings, only for their numbers to decline once the lions recovered. 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 larger species' dominance. However, a few cases have been reported of old and wounded lions falling prey to African wild dogs.Schaller, G. B. (1972). ''The Serengeti lion: A study of predator-prey relations''. Chicago: University of Chicago Press. p. 188. . 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.

Spotted hyenas are important kleptoparasites and follow packs of African wild dogs to appropriate their kills. They typically inspect areas where African wild dogs have rested and eat any food remains they find. When approaching African 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, with African wild dog densities being negatively correlated with high hyena populations. Beyond piracy, cases of interspecific killing of African wild dogs by spotted hyenas are documented. African wild dogs are apex predators, normally only fatally losing contests to larger social carnivores, although anecdotally Nile crocodiles may opportunistically and rarely prey upon a wild dog. 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.

Distribution

African wild dogs once ranged across much of sub-Saharan Africa, being absent only in the driest desert regions and lowland forests. The species has been largely exterminated in North and West Africa, and has been greatly reduced in number in Central Africa and northeast Africa. The majority

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