African golden wolf

African golden wolf
Temporal range: Early Pleistocene – Recent 1.3-0 Ma
Canis anthus bea in Ngorongoro Crater, Tanzania
Not evaluated (IUCN 3.1)
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Carnivora
Family: Canidae
Genus: Canis
Species: C. anthus
Binomial name
Canis anthus[1]
F. Cuvier, 1820
Subspecies

The African golden wolf (Canis anthus/Canis lupaster)[2] also known as the golden wolf, African wolf or thoa, is a canid native to north and northeastern Africa. The species is common in north-west and north-east Africa, occurring from Senegal to Egypt in the east, in a range including Morocco, Algeria, and Libya in the north to Nigeria, Chad and Tanzania in the south.[3] It is a desert-adapted canid, and is common in plains and steppe areas, including ones lacking abundant water.[4] In the Atlas Mountains, the species has been sighted in elevations as high as 1,800 metres.[5] It is primarily a predator, targeting invertebrates and mammals as large as gazelle fawns, though larger animals are sometimes taken. Other foodstuffs include animal carcasses, human refuse, and fruit. The African wolf is a monogamous and territorial animal, whose social structure includes yearling offspring remaining with the family to assist in raising their parents' younger pups.[4]

It was previously classified as an African variant of the Eurasian golden jackal, with at least one subspecies (Canis anthus lupaster) having been classified as a grey wolf. In 2015, a series of analyses on the species' mitochondrial DNA and nuclear genome demonstrated that it was in fact distinct from both the golden jackal and the grey wolf, and more closely related to grey wolves and coyotes.[6][7] It is nonetheless still close enough to the golden jackal to produce hybrid offspring, as indicated through genetic tests on jackals in Israel[6] and a 19th-century captive crossbreeding experiment.[8] As the IUCN's golden jackal page has not been updated since 2008, it has yet to recognise the distinctiveness of the African wolf; thus its conservation status has not been evaluated, and no reliable population estimates are available.[9]

It plays a prominent role in some African cultures; in Maghreb and Egyptian folklore, it is viewed as an untrustworthy animal whose body parts can be used for medicinal or ritualistic purposes,[10][11][12] while it is held in high esteem in Senegal's Serer religion as being the first creature to be created by the god Roog.[13]

Local and indigenous names

Physical description

Various C. anthus phenotypes, ranging from gracile jackal-like morphs to more robust wolf-like ones.

The African wolf is intermediate in size between the African jackals (C. mesomelas and C. adustus) and small subspecies of grey wolves,[2] with both sexes weighing 7–15 kg (15–33 lb), and standing 40 cm in height.[4] There is however a high degree of size variation geographically, with Western and Northern African specimens being larger than their East African cousins.[2] It has a relatively long snout and ears, while the tail is comparatively short, measuring 20 cm in length. Fur colour varies individually, seasonally and geographically, though the typical colouration is yellowish to silvery grey, with slightly reddish limbs and black speckling on the tail and shoulders. The throat, abdomen and facial markings are usually white, and the eyes are amber coloured. Females bear two to four pairs of teats.[4] Although superficially similar to the Eurasian golden jackal (particularly in East Africa), the African golden wolf has a more pointed muzzle and sharper, more robust teeth.[6] The ears are longer in the African wolf, and the skull has a more elevated forehead.[18]

Taxonomic history

Early writings

Skull of African wolf (left) and golden jackal (right). Note the former's more elevated forehead[18] and narrower muzzle.[6]

Aristotle wrote of wolves living in Egypt, mentioning that they were smaller than the Greek kind. Georg Ebers wrote of the wolf being among the sacred animals of Egypt, describing it as a "smaller variety" of wolf to those of Europe, and noting how the name Lykopolis, the Ancient Egyptian city dedicated to Anubis, means "city of the wolf".[19][20]

The African golden wolf was first recognised as being a separate species from the Eurasian golden jackal by Frédéric Cuvier in 1820, who described it as being a more elegant animal, with a more melodic voice and a less strong odour. The binomial name he chose for it was derived from the Arcadian Anthus family described by Pliny the Elder in his Natural History, whose members would draw lots to become werewolves.[1] Eduard Rüppell proposed that the animal was the ancestor of Egyptian sighthounds, and named it Wolf's-hund (wolf dog),[21] while Charles Hamilton Smith named it "thoa" or "thous dog".[22] An attempt was also made in 1821 to hybridise the two species in captivity, resulting in the birth of five pups, three of which died before weaning. The two survivors were noted to never play with each other and had completely contrasting temperaments; one inherited the golden jackal's shyness, while the other was affectionate toward its human captors.[8] English biologist St. George Jackson Mivart emphasised the differences between the African wolf and the golden jackal in his writings:

Illustration of golden jackal-African wolf hybrids bred in captivity (1821).
... it is a nice question whether the Common Jackal of North Africa should or should not be regarded as of the same species [as the golden jackal]... Certainly the differences of coloration which exist between these forms is not nearly so great as those which are to be found to occur between the different local varieties of C. lupus. We are nevertheless inclined... to keep the North-African and Indian Jackals distinct... The reasons why we prefer to keep them provisionally distinct is that though the difference between the two forms (African and Indian) is slight as regards coloration, yet it appears to be a very constant one. Out of seventeen skins of the Indian form, we have only found one which is wanting in the main characteristic as to difference of hue. The ears also are relatively shorter than in the North-African form. But there is another character to which we attach greater weight. However much the different races of Wolves differ in size, we have not succeeded in finding any constant distinctive characters in the form of the skull or the proportions of the lobes of any of the teeth. So far as we have been able to observe, such differences do exist between the Indian and North-African Jackals.
— Mivart (1890)[23]
Comparative illustration of C. aureus (top) and C. anthus (bottom).

The canids present in Egypt in particular were noted to be so much more grey wolf-like than populations elsewhere in Africa that Hemprich and Ehrenberg gave them the binomial name Canis lupaster in 1832. Likewise, Thomas Henry Huxley, upon noting the similarities between the skulls of lupaster and Indian grey wolves, classed the animal as a subspecies of grey wolf. However, the animal was subsequently synonymised with the golden jackal by Ernst Schwarz in 1926. This classification remained unquestioned until 1981, when zoologist Walter Ferguson of Tel Aviv University argued in favour of lupaster being a subspecies of grey wolf based on cranial measurements, stating that the classing of the animal as a jackal was based solely on the animal's small size, and predated the discovery of C. l. arabs, which is intermediate in size between C. l. lupus and lupaster.[20]

21st-century discoveries

Further doubts over its being conspecific with the golden jackal of Eurasia arose in December 2002, when a canid was sighted in Eritrea's Danakil Desert whose appearance didn't correspond to that of the golden jackal or the six other recognised species of the area, but strongly resembled that of the grey wolf. The area had previously been largely unexplored because of its harsh climate and embroilment in the Eritrean War of Independence and subsequent Eritrean–Ethiopian War, though local Afar tribesmen knew of the animal, and referred to it as wucharia (wolf).[14]

The animal's wolf-like qualities were confirmed in 2011, when several golden "jackal" populations in Egypt and the Horn of Africa classed as Canis aureus lupaster were found to have mtDNA sequences more closely resembling those found in grey wolves than those of golden jackals.[19] These wolf-like mtDNA sequences were found to occur over a 6,000 km wide area, encompassing Algeria, Mali and Senegal. Furthermore, the sampled African specimens displayed much more nucleotide and haplotype diversity than that present in Indian and Himalayan grey wolves, thus indicating a larger ancestral population, and an effective extant population of around 80,000 females. Both these studies proposed reclassifying Canis aureus lupaster as a subspecies of grey wolf.[24]

In 2015, a more thorough comparative study of mitochondrial and nuclear genomes on a larger sample of wolf-like African canids from north, east and west Africa showed that they were in fact all distinct from the Eurasian golden jackal, with a genetic divergence of around 6.7%,[6] which is greater than that between grey wolves and coyotes (4%) and that between grey wolves and domestic dogs (0.2%).[25] Furthermore, the study showed that these African wolf-like canids (renamed Canis anthus, or African golden wolves) were more closely related to grey wolves and coyotes than to Eurasian golden jackals, and that C. a. lupaster merely represents a distinct phenotype of African wolf rather than an actual grey wolf. The phylogenetic tree below is based on nuclear sequences:[6]




Side-striped jackal



Black-backed jackal












Dog



Gray wolf




Coyote




African golden wolf




Golden jackal




Ethiopian wolf




Dhole





African wild dog





It was estimated that the African golden wolf diverged from the wolf–coyote clade 1.0–1.7 million years ago, during the Pleistocene, and therefore its superficial similarity to the Eurasian golden jackal (particularly in East Africa, where African wolves are similar in size to golden jackals) would be a case of parallel evolution. Considering its phylogenetic position and the canid fossil record, it is likely that the African wolf evolved from larger ancestors that became progressively more jackal-like in size upon populating Africa on account of interspecific competition with both larger and smaller indigenous carnivores. Traces of golden wolf DNA were identified in golden jackals in Israel, which adjoins Egypt, thus indicating the presence of a hybrid zone.[6] The study's findings were corroborated that same year by Spanish, Mexican and Moroccan scientists analysing the mtDNA of wolves in Morocco, who found that the specimens analysed were distinct from both Eurasian golden jackals and grey wolves but bore a closer relationship to the latter.[7] Studies on RAD sequences found instances of African wolves hybridizing with both feral dogs and Ethiopian wolves.[26]

In 2017, it was proposed by scientists at the Oslo and Helsinki Universities that the binomial name C. anthus was a nomen dubium, on account of the fact that Cuvier's 1820 description of the holotype, a female collected from Senegal, seems to be describing the side-striped jackal rather than the actual African wolf, and doesn't match the appearance of a male specimen described by Cuvier in his later writings. This ambiguity, coupled with the disappearance of the holotype's remains, lead to the scientists proposing giving priority to Hemprich and Ehrenberg's name C. lupaster, due to the type specimen having a more detailed and consistent description, and its remains being still examinable at the Museum für Naturkunde.[2]

Relationship to the Himalayan wolf

In 2017 a study of mitochondrial DNA and other DNA sequences found that the Himalayan wolf formed a basal distinct monophyletic clade relative to the holarctic gray wolf. Its degree of divergence to the holarctic gray wolf is similar to the degree of divergence of the African golden wolf from the holarctic gray wolf. The Himalayan wolf shares an X-chromosome haplotype with the African golden wolf. It possesses a unique Y-chromosome haplotype that falls phylogenetically between the holarctic gray wolf and the African golden wolf.[27]

Subspecies

As of 2015, MSW3 still classifies the six African wolf subspecies as part of Canis aureus.[28]

Although in the past several attempts have been made to synonymise many of the proposed names, the taxonomic position of West African wolves, in particular, is too confused to come to any precise conclusion, as the collected study materials are few. Prior to 1840, six of the ten supposed West African subspecies were named or classed almost entirely because of their fur colour.[29]

The species' display of high individual variation, coupled with the scarcity of samples and the lack of physical barriers on the continent preventing gene flow, brings into question the validity of some of the West African forms.[29]

Behaviour

Social and reproductive behaviours

The African wolf's social organisation is extremely flexible, varying according to the availability and distribution of food. The basic social unit is a breeding pair, followed by its current offspring, or offspring from previous litters staying as "helpers".[15] Large groups are rare, and have only been recorded to occur in areas with abundant human waste. Family relationships among African wolves are comparatively peaceful compared to those of the black-backed jackal; although the sexual and territorial behaviour of grown pups is suppressed by the breeding pair, they are not actively driven off once they attain adulthood. African wolves also lie together and groom each other much more frequently than black-backed jackals. In the Serengeti, pairs defend permanent territories encompassing 2–4 km², and will vacate their territories only to drink or when lured by a large carcass.[4] The pair patrols and marks its territory in tandem. Both partners and helpers will react aggressively towards intruders, though the greatest aggression is reserved for intruders of the same sex; pair members do not assist each other in repelling intruders of the opposite sex.[4]

Threat postures in C. a. lupaster (left) and C. a. anthus (right)

The African wolf's courtship rituals are remarkably long, during which the breeding pair remains almost constantly together. Prior to mating, the pair patrols and scent marks its territory. Copulation is preceded by the female holding her tail out and angled in such a way that her genitalia are exposed. The two approach each other, whimpering, lifting their tails and bristling their fur, displaying varying intensities of offensive and defensive behaviour. The female sniffs and licks the male's genitals, whilst the male nuzzles the female's fur. They may circle each other and fight briefly. The copulatory tie lasts roughly four minutes. Towards the end of estrus, the pair drifts apart, with the female often approaching the male in a comparatively more submissive manner. In anticipation of the role he will take in raising pups, the male regurgitates or surrenders any food he has to the female. In the Serengeti, pups are born in December–January, and begin eating solid food after a month. Weaning starts at the age of two months, and ends at four months. At this stage, the pups are semi-independent, venturing up to 50 metres from the den, even sleeping in the open. Their playing behaviour becomes increasingly more aggressive, with the pups competing for rank, which is established after six months. The female feeds the pups more frequently than the male or helpers do, though the presence of the latter allows the breeding pair to leave the den and hunt without leaving the litter unprotected.[4]

The African wolf's life centres around a home burrow, which usually consists of an abandoned and modified aardvark or warthog earth. The interior structure of this burrow is poorly understood, though it is thought to consist of a single central chamber with 2–3 escape routes. The home burrow can be located in both secluded areas or surprisingly near the dens of other predators.[32]

Communication

African wolves frequently groom one another, particularly during courtship, during which it can last up to ½ hour. Nibbling of the face and neck is observed during greeting ceremonies. When fighting, the African wolf slams its opponents with its hips, and bites and shakes the shoulder. The species' postures are typically canine, and it has more facial mobility than the black-backed and side-striped jackals, being able to expose its canine teeth like a dog.[4]

The vocabulary of the African wolf is similar to that of the domestic dog, with seven sounds having been recorded.[17] The African wolf's vocalisations include howls, barks, growls, whines and cackles.[4] Subspecies can be recognised by differences in their howls.[17] One of the most commonly heard sounds is a high, keening wail, of which there are three varieties; a long single toned continuous howl, a wail that rises and falls, and a series of short, staccato howls. These howls are used to repel intruders and attract family members. Howling in chorus is thought to reinforce family bonds, as well as establish territorial status.[4] A comparative analysis of African wolf and some grey wolf subspecies' howls demonstrated that the former's howls bear similarities to those of the Indian grey wolf, being high pitched and of relatively short duration.[33]

Hunting behaviour

A Serengeti wolf (C. a. bea) navigating through a herd of blue wildebeest in the Ngorongoro National Park, Tanzania

The African wolf rarely catches hares, due to their speed. Gazelle mothers (often working in groups of two or three) are formidable when defending their young against single wolves, which are much more successful in hunting gazelle fawns when working in pairs. A pair of wolves will methodically search for concealed gazelle fawns within herds, tall grass, bushes and other likely hiding places.[4]

Although it is known to kill animals up to three times its own weight, the African wolf targets mammalian prey much less frequently than the black-backed jackal overall.[4] On capturing large prey, the African wolf makes no attempt to kill it; instead it rips open the belly and eats the entrails. Small prey is typically killed by shaking, though snakes may be eaten alive from the tail end. The African wolf often carries away more food than it can consume, and caches the surplus, which is generally recovered within 24 hours.[32] When foraging for insects, the African wolf turns over dung piles to find dung beetles. During the dry seasons, it excavates dung balls to reach the larvae inside. Grasshoppers and flying termites are caught either by pouncing on them while they are on the ground or are caught in mid-air. It is fiercely intolerant of other scavengers, having been known to dominate vultures on kills – one can hold dozens of vultures at bay by threatening, snapping and lunging at them.[4]

Ecology

Habitat

The African wolf inhabits a number of different habitats; in Algeria it lives in Mediterranean, coastal and hilly areas (including hedged farmlands, scrublands, pinewoods and oak forests), while populations in Senegal inhabit tropical, semi-arid climate zones including Sahelian savannahs. Wolf populations in Mali have been documented in arid Sahelian massifs.[24] In Egypt, the African wolf inhabits agricultural areas, wastelands, desert margins, rocky areas, and cliffs. At Lake Nasser, it lives close to the lakeshore.[16] In 2012, African wolves were photographed in Morocco's Azilal Province at an elevation of 1,800 metres.[5][34] It apparently does well in areas where human density is high and natural prey populations low, as is the case in the Enderta district in northern Ethiopia.[35]

Diet

Serengeti wolf (C. a. bea) eating an agama

In West Africa, the African wolf mostly confines itself to small prey, such as hares, rats, ground squirrels and cane rats. Other prey items include lizards, snakes, and ground-nesting birds, such as francolins and bustards. It also consumes a large amount of insects, including dung beetles, larvae, termites and grasshoppers. It will also kill young gazelles, duikers and warthogs.[32] In East Africa, it consumes invertebrates and fruit, though 60% of its diet consists of rodents, lizards, snakes, birds, hares and Thomson's gazelles.[15] During the wildebeest calving season, African wolves will feed almost exclusively on their afterbirth.[17] In the Serengeti and Ngorongoro Crater, less than 20% of its diet comes from scavenging.[4] In Senegal, where both C. a. anthus and C. a. lupaster coexist, some degree of niche segregation is apparent in their choice of prey; the former is reputed to feed primarily on lambs, whereas the latter attacks larger prey, such as sheep, goats and cattle.[24]

Enemies and competitors

The African wolf generally manages to avoid competing with black-backed and side-striped jackals by occupying a different habitat (grassland, as opposed to the closed and open woodlands favoured by the latter two species) and being more active during the daytime.[36] Nevertheless, the African wolf has been known to kill the pups of black-backed jackals,[15] but has in turn been observed to be dominated by adults during disputes over carcasses.[17] It often eats alongside African wild dogs, and will stand its ground if the dogs try to harass it.[4] Encounters with Ethiopian wolves are usually agonistic, with Ethiopian wolves dominating African wolves if the latter enter their territories, and vice versa. Although African wolves are inefficient rodent hunters and thus not in direct competition with Ethiopian wolves, it is likely that heavy human persecution prevents the former from attaining numbers large enough to completely displace the latter.[37] Nevertheless, there is at least one record of an African wolf pack adopting a male Ethiopian wolf.[38]

Wolves will feed alongside spotted hyenas, though they will be chased if they approach too closely. Spotted hyenas will sometimes follow wolves during the gazelle fawning season, as wolves are effective at tracking and catching young animals. Hyenas do not take to eating wolf flesh readily; four hyenas were reported to take half an hour in eating one. Overall, the two animals typically ignore each other when no food or young is at stake.[39] Wolves will confront a hyena approaching too closely to their dens by taking turns in biting the hyena's hocks until it retreats.[4]

Wolves in the Serengeti are known to carry the canine parvovirus, canine herpesvirus, canine coronavirus and canine adenovirus.[15]

Range and status

Fossil finds dating back to the Pleistocene indicate that the species' range was not always restricted to Africa, with remains having been found in the Levant and Saudi Arabia.[2]

In Tanzania, the African wolf is limited to a small area of the north between the western slopes of Mount Kilimanjaro and the centre of the Serengeti. In the latter area, it occurs mostly in the short-grass plains, the floor of the Ngorongoro Crater, and the plains between the Olmoti and Empakai Craters, being relatively rare in Serengeti National Park, Loliondo and the Maswa game reserve. The species also inhabits the Lake Natron area and West Kilimanjaro. It is sometimes found in the northern part of Arusha National Park, and as far south as Manyara. In areas where it is common, such as the short-grass plains of Serengeti National Park and the Ngorongoro Crater, population densities can range between 0.5-1.5 specimens per km2. A population decrease of 60% has been recorded in the southern plains of Serengeti National Park since the early 1970s, though the reasons are unknown.[40]

In folklore

Wolf-shaped bronze amulet from Egypt's Ptolemaic Period (711 - 30 BCE).

The wolf was the template of numerous Ancient Egyptian deities, including Anubis, Wepwawet and Duamutef.[41] According to Diodorus Siculus, the Egyptian city of Lykopolis was named in honour of a pack of wolves that repelled an Ethiopian invasion.[42] Arab Egyptian folklore holds that the wolf can cause chickens to faint from fear by simply passing underneath their roosts, and associates its body parts with various forms of folk magic: placing a wolf's tongue in a house is believed to cause the inhabitants to argue, and its meat is thought to be useful in treating insanity and epilepsy. Its heart is believed to protect the bearer from wild animal attacks, while its eye can protect against the evil eye.[10]

Although considered haram in Islamic dietary laws, the wolf is important in Moroccan folk medicine.[11] Edvard Westermarck wrote of several remedies derived from the wolf in Morocco, including the use of its fat as a lotion, the consumption of its meat to treat respiratory ailments, and the burning of its intestines in fumigation rituals meant to increase the fertility of married couples. The wolf's gall bladder was said to have various uses, including curing sexual impotence and serving as a charm for women wishing to divorce their husbands. Westermarck noted, however, that the wolf was also associated with more nefarious qualities: it was said that a child who eats wolf flesh before reaching puberty will be forever cursed with misfortune and that scribes and saintly persons refrain from consuming it even in areas where it is socially acceptable, as doing so would render their charms useless.[12]

The African wolf isn't common in Neolithic rock art, though it does occasionally appear; a definite portrayal is shown on the Kef Messiouer cave in Algeria's Tébessa Province, where it is shown feeding on a wild boar carcass alongside a lion pride. It plays a role in Berber mythology, particularly that of the Ait Seghrouchen of Morocco, where it plays a similar role in folktales as the red fox does in Medieval European fables, though it is often the victim of the more cunning hedgehog.[43]

The wolf plays a prominent role in the Serer religion's creation myth, where it is viewed as the first living creature created by Roog, the Supreme God and Creator.[13][44] In one aspect, it can be viewed as an Earth-diver sent to Earth by Roog, in another, as a fallen prophet for disobeying the laws of the divine. The wolf was the first intelligent creature on earth, and it is believed that it will remain on earth after human beings have returned to the divine. The Serers believe that, not only does it know in advance who will die, but it traces the tracks in advance of those who will go to funerals. The movements of the wolf are carefully observed, because the animal is viewed as a seer who came from the transcendence and maintains links with it. Although believed to be rejected in the bush by other animals and deprived of its original intelligence, it is still respected because it dared to resist the supreme being who still keeps it alive.[13]

See also

References

  1. 1 2 (in French) Cuvier, Frédéric (1824), Histoire naturelle des mammifères, tome 2, A Paris : Chez A. Belin ...
  2. 1 2 3 4 5 Viranta, Suvi; Atickem, Anagaw; Werdelin, Lars; Stenseth, Nils Chr. (2017). "Rediscovering a forgotten canid species". BMC Zoology. 2. doi:10.1186/s40850-017-0015-0.
  3. ↑ Jhala, Y.V. & Moehlman, P.D. (2008). "Canis aureus". IUCN Red List of Threatened Species. Version 2008. International Union for Conservation of Nature. Retrieved 22 March 2009.
  4. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 Estes, Richard (1992). "The behavior guide to African mammals: including hoofed mammals, carnivores, primates". University of California Press. pp. 398–404. ISBN 0-520-08085-8.
  5. 1 2 3 4 (in French) Rachid Tarik, "Le loup apparaît dans la région d'Azilal", Le Matin (April 29, 2013)
  6. 1 2 3 4 5 6 7 Koepfli, K.-P.; Pollinger, J.; Godinho, R.; Robinson, J.; Lea, A.; Hendricks, S.; Schweizer, R. M.; Thalmann, O.; Silva, P.; Fan, Z.; Yurchenko, A. A.; Dobrynin, P.; Makunin, A.; Cahill, J. A.; Shapiro, B.; Álvares, F.; Brito, J. C.; Geffen, E.; Leonard, J. A.; Helgen, K. M.; Johnson, W. E.; O'Brien, S. J.; Van Valkenburgh, B.; Wayne, R. K. (2015-08-17). "Genome-wide Evidence Reveals that African and Eurasian Golden Jackals Are Distinct Species". Current Biology. 25: 2158–65. PMID 26234211. doi:10.1016/j.cub.2015.06.060.
  7. 1 2 (in Spanish) Urios, Vicente; Donat-Torres, Maria P.; Ramírez, Carlos; Monroy-Vilchis, Octavio; Hamid Rgribi-Idrissi (2015): El análisis del genoma mitocondrial del cánido estudiado en Marruecos manifiesta que no es ni lobo (Canis lupus) ni chacal euroasiático (Canis aureus). figshare. doi:10.6084/m9.figshare.1524971
  8. 1 2 (in French) Cuvier, Frédéric (1824), Histoire naturelle des mammifères, tome 3, A Paris : Chez A. Belin ...
  9. ↑ Golden jackal: A new wolf species hiding in plain sight, The Guardian, (30 July 2015)
  10. 1 2 3 4 Osborn, Dale. J.; Helmy, Ibrahim (1980). "The contemporary land mammals of Egypt (including Sinai)". Field Museum of Natural History, pp. 360-371
  11. 1 2 Bidwell, M. & Bidwell, R. (2005), The Traveller's Companion, Second Edition, Tauris Parke Paperbacks, p. 228, ISBN 1845111079
  12. 1 2 Westermarck, E. (2013), Ritual and Belief in Morocco: Vol. II, Routledge Revivals, ISBN 1317912616
  13. 1 2 3 (in French) Gravrand, Henry (1990), "La Civilisation Sereer - Pangool", vol. 2. Les Nouvelles Editions Africaines du Senegal, pp. 201−203, ISBN 2-7236-1055-1
  14. 1 2 Tiwari, J. K.; Sillero-Zubiri, C. (2004). "Unidentified Canid in Horn of Africa" (PDF). Canid News. 7: 5.
  15. 1 2 3 4 5 6 7 8 9 10 11 Jhala, Y. V. & Moehlman, P. D. 2004. Golden jackal Canis aureus Archived 2007-10-12 at the Wayback Machine.. In Sillero-Zubiri, C., Hoffman, M. & MacDonald, D. W., ed., Canids: Foxes, Wolves, Jackals and Dogs - 2004 Status Survey and Conservation Action Plan, 156-161. IUCN/SSC Canid Specialist Group, ISBN 2-8317-0786-2
  16. 1 2 Hoath, Richard (2009), A Field Guide to the Mammals of Egypt, American Univ in Cairo Press, pp. 70-73, ISBN 9774162544
  17. 1 2 3 4 5 Kingdon, Jonathan (1988). "East African mammals: an atlas of evolution in Africa, Volume 3, Part 1". University of Chicago Press. p. 18-22. ISBN 0-226-43721-3.
  18. 1 2 Mivart, George (1890), Dogs, Jackals, Wolves and Foxes: A Monograph of the Canidæ, R.H. Porter, London, pp. 41–43
  19. 1 2 Rueness, E. K.; Asmyhr, M. G.; Sillero-Zubiri, C.; MacDonald, D. W.; Bekele, A.; Atickem, A.; Stenseth, N. C. (2011). Gilbert, Thomas M, ed. "The Cryptic African Wolf: Canis aureus lupaster is Not a Golden Jackal and is Not Endemic to Egypt". PLoS ONE. 6 (1): e16385. PMC 3027653 Freely accessible. PMID 21298107. doi:10.1371/journal.pone.0016385.
  20. 1 2 Ferguson, W.W. (1981). "The systematic of Canis aureus lupaster (Carnivora : Canidae) and the occurrence of Canis lupus in North Africa, Egypt and Sinai". Mammalia. 4: 459–465. doi:10.1515/mamm.1981.45.4.459.
  21. ↑ (in German) Rüppell, Eduard (1826), Atlas zu der Reise im nördlichen Afrika, Frankfurt am Main : Gedruckt und in Commission bei Heinr. Ludw. Brönner, pp. 44–46
  22. ↑ Smith, Charles Hamilton; Jardine, Sir William (1839). The natural history of dogs : canidae or genus canis of authors; including also the genera hyaena and proteles, Volume I. Edinburgh : W. H. Lizars. pp. 193–194
  23. ↑ Mivart, George (1890), Dogs, Jackals, Wolves and Foxes: A Monograph of the Canidæ, R.H. Porter, London, pp. 36–37
  24. 1 2 3 4 Gaubert P, Bloch C, Benyacoub S, Abdelhamid A, Pagani P, et al. (2012). "Reviving the African Wolf Canis lupus lupaster in North and West Africa: A Mitochondrial Lineage Ranging More than 6,000 km Wide". PLoS ONE. 7 (8): e42740. PMC 3416759 Freely accessible. PMID 22900047. doi:10.1371/journal.pone.0042740.
  25. ↑ Wayne, Robert K. (1993). "Molecular evolution of the dog family". Trends in Genetics. 9 (6): 218–224. PMID 8337763. doi:10.1016/0168-9525(93)90122-X.
  26. ↑ Bahlk, S. H. (2015). Can hybridization be detected between African wolf and sympatric canids? . Master of Science Thesis. Center for Ecological and Evolutionary Synthesis Department of Bioscience Faculty of Mathematics and Natural Science, University of Oslo, Norway
  27. ↑ Werhahn, Geraldine; Senn, Helen; Kaden, Jennifer; Joshi, Jyoti; Bhattarai, Susmita; Kusi, Naresh; Sillero-Zubiri, Claudio; MacDonald, David W. (2017). "Phylogenetic evidence for the ancient Himalayan wolf: Towards a clarification of its taxonomic status based on genetic sampling from western Nepal". Royal Society Open Science. 4 (6): 170186. doi:10.1098/rsos.170186.
  28. ↑ Wozencraft, W.C. (2005). "Order Carnivora". In Wilson, D.E.; Reeder, D.M. Mammal Species of the World: A Taxonomic and Geographic Reference (3rd ed.). Johns Hopkins University Press. pp. 532–628. ISBN 978-0-8018-8221-0. OCLC 62265494.
  29. 1 2 Rosevear, Donovan Reginald (1974). "The carnivores of West Africa". London : Trustees of the British Museum (Natural History). pp. 38–44. ISBN 1-175-10030-7.
  30. 1 2 3 4 5 Lydekker, Richard (1908). "The Game Animals of Africa". London, R. Ward, limited. pp. 459-461
  31. ↑ Heller, E. 1914. Four new subspecies oflarge mammals from equatorial Africa. Smithsonian Miscellaneous Collections, 61, No. 22. Washington, DC: Smithsonian Institution
  32. 1 2 3 Rosevear, Donovan Reginald (1974). "The carnivores of West Africa". London : Trustees of the British Museum (Natural History). pp. 46–47. ISBN 1-175-10030-7.
  33. ↑ Lauren Hennelly, Bilal Habib, Holly Root-Gutteridge, Vicente Palacios and Daniela Passilongo. (2017) "Howl variation across Himalayan, North African, Indian, and Holarctic wolf clades: tracing divergence in the world’s oldest wolf lineages using acoustics". Current Zoology 63:3, 341-348.
  34. ↑ (in Spanish) Moliner, V. U., Ramírez, C., Gallardo, M. & Idrissi, H. R. (2012), "Detectan el lobo en Marruecos gracias al uso del foto-trampeo", Quercus, 319:14-15, ISSN 0212-0054
  35. ↑ Yirga, G. et al. "Densities of spotted hyaena (Crocuta crocuta) and African golden wolf (Canis anthus) increase with increasing anthropogenic influence", Elsevier, Volume 413, Part B, 22 August 2016, Pages 7–14
  36. ↑ Fuller, T.K.; Biknevicius, A.R.; Kat, P.W.; Van Valkenburgh, B.; Wayne, R.K. (1989). "The ecology of three sympatric jackal species in the rift valley of kenya". African Journal of Ecology. 27: 313–323. doi:10.1111/j.1365-2028.1989.tb01025.x.
  37. ↑ Gutema, T. M., Foraging ecology and trophic niche overlap between sympatric African wolf and Ethiopian wolf in the Ethiopian Highlands, The Rufford Foundation (November 2015)
  38. ↑ Sillero-Zubiri, C.; Gottelli, D. (1994). "Canis simensis" (PDF). Mammalian Species. 385: 1–6.
  39. ↑ Kruuk, Hans (1972). The Spotted Hyena: A Study of Predation and Social Behaviour. Chicago: University of Chicago Press. ISBN 0-226-45507-6.
  40. ↑ Foley, C. et al. (2014), A Field Guide to the Larger Mammals of Tanzania, Princeton University Press, pp. 102-103, ISBN 1400852803
  41. ↑ Remler, P. (2010), Egyptian Mythology, A to Z, Infobase Publishing, p. 99, ISBN 1438131801
  42. ↑ Burton, Anne (1973), Diodorus Siculus, Book 1: A Commentary, BRILL, p.259, ISBN 9004035141
  43. ↑ G. Trécolle & G. Camps, « Chacal », in 12 | Capsa – Cheval, Aix-en-Provence, Edisud (« Volumes », no 12) , 1993 [online], published online 1 March 2012, consulted 19 November 2015. URL : http:// encyclopedieberbere.revues.org/2099
  44. ↑ (in French) Thiaw, Issa laye, "Mythe de la création du monde selon les sages sereer", pp. 45−50, 59−61 [in] "Enracinement et Ouverture" – "Plaidoyer pour le dialogue interreligieux", Konrad Adenauer Stiftung (23–24 June 2009), Dakar

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