African golden wolf

The African golden wolf (Canis anthus),[2] also known as the Egyptian jackal or grey jackal,[3] , is a canid native to north and northeastern Africa. The species is the descendant of a genetically admixed canid of 72% grey wolf and 28% Ethiopian wolf ancestry.[4] 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, Tunisia and Libya in the north to Nigeria, Chad and Tanzania in the south.[5] It is a desert-adapted canid, and is common in plains and steppe areas, including ones lacking abundant water.[6] In the Atlas Mountains, the species has been sighted in elevations as high as 1,800 metres.[7] 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.[6]

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.[8][9] It is nonetheless still close enough to the golden jackal to produce hybrid offspring, as indicated through genetic tests on jackals in Israel[8] and a 19th-century captive crossbreeding experiment.[10] As the IUCN's golden jackal page has not been updated since 2008, it has yet to recognise the distinctiveness of the African golden wolf; thus its conservation status has not been evaluated and no reliable population estimates are available.[11]

It plays a prominent role in some African cultures; in North African folklore, it is viewed as an untrustworthy animal whose body parts can be used for medicinal or ritualistic purposes,[12][13][14] while it is held in high esteem in Senegal's Serer religion as being the first creature to be created by the god Roog.[15]

African golden wolf
Temporal range: Middle Pleistocene – Recent 0.6-0 Ma
Golden wolf small
Canis anthus bea in Ngorongoro Crater, Tanzania
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Carnivora
Family: Canidae
Genus: Canis
Species:
C. anthus
Binomial name
Canis anthus
Subspecies
Canis anthus range

Local and indigenous names

Linguistic group or area Indigenous name
Afar Wucharia[16]
Amazigh Ouchan asian (for C. a. algirensis)[7]
Ouchan akhatar (for C. a. lupaster)[7]
Amharic ተረ ቀበሮ (Tera kebero)[17]
Arabic ابن آوى (Ibn awa)[18]
ذئب (Deeb)[12]
أبو سليمان (Abu soliman)[12]
Fula Sundu[17]
Hausa Dila[17]
Songhai Nzongo[17]
Swahili (standard)
Swahili (Tanzania)
Bweha wa mbuga[17][19]
Bweha dhahabu[17]
Wolof Tili[17]
English Golden Jackal>

Physical description

Journal.pone.0042740.g006
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,[20] with both sexes weighing 7–15 kg (15–33 lb), and standing 40 cm in height.[6] There is however a high degree of size variation geographically, with Western and Northern African specimens being larger than their East African cousins.[20] 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.[6] 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.[8] The ears are longer in the African wolf, and the skull has a more elevated forehead.[21]

Taxonomic history

Early writings

Beitrag zur Kenntnis der nordafrikanischen Schakale nebst Bemerkungen über deren verhältnis zu den haushunden, insbesondere uordafranischen und altägyptischen Hunderassen (1908) C. l. lupaster & C. a. aureus
Skull of African wolf (left) and golden jackal (right). Note the former's more elevated forehead[21] and narrower muzzle.[8]

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".[22][23]

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),[24] while Charles Hamilton Smith named it "thoa" or "thous dog".[25] 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.[10] English biologist St. George Jackson Mivart emphasised the differences between the African wolf and the golden jackal in his writings:

Histoire naturelle des mammifères, t. 3 (1824) Canis anthus x aureus
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)[26]
Jackalvarieties
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.

In 1965, the Finnish paleontologist Björn Kurtén wrote:

The taxonomy of the Jackals in the Near East is still a matter of dispute. On the basis of skeletal material, however, it can be stated that the Wolf Jackal is specifically distinct from the much smaller Golden Jackal.[27]

In 1981, zoologist Walter Ferguson 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.[23]

21st-century discoveries

Phylogenetic tree of the extant wolf-like canids with timing in millions of years[a]
Caninae 3.5 Ma
3.0
2.5
2.0
0.96
0.60
0.38
0.25
?
0.08
0.031

Dog Tibetan mastiff (transparent background).png

Holarctic grey wolf Dogs, jackals, wolves, and foxes (Plate I).png

Late Pleistocene wolfThe American Museum journal (c1900-(1918)) (Canis dirus) transparent background

Indian plains wolf Dogs, jackals, wolves, and foxes (Plate I).png

Himalayan wolf/Tibetan wolf Dogs, jackals, wolves, and foxes (Plate I).png

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

0.11

African golden wolf northwestern AfricaDogs, jackals, wolves, and foxes (Plate XI).jpg

African golden wolf eastern AfricaDogs, 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

Blue shading represents the species Canis lupus

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).[16]

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[20] were found to have mtDNA sequences more closely resembling those found in grey wolves than those of golden jackals.[22] 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.[28]

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%,[8][29][30] which is greater than that between grey wolves and coyotes (4%) and that between grey wolves and domestic dogs (0.2%).[31] 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,[8][32] 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:[8]

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.[8] 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.[9] Studies on RAD sequences found instances of African wolves hybridizing with both feral dogs and Ethiopian wolves.[33]

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, led 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.[20] The following year, a major genetic study of Canis species also referred to the African golden wolf as Canis lupaster.[4]

Admixture with other Canis species

In 2018, whole genome sequencing was used to compare members of the genus Canis. The study supports the African golden wolf being distinct from the golden jackal, and with the Ethiopian wolf being genetically basal to both. There are two genetically distinct African golden wolf populations that exist in northwestern and eastern Africa. This suggests that Ethiopian wolves – or a close and extinct relative – once had a much larger range within Africa to admix with other canids. There is evidence of gene flow between the eastern population and the Ethiopian wolf, which has led to the eastern population being distinct from the northwestern population. The common ancestor of both African golden wolf populations was a genetically admixed canid of 72% grey wolf and 28% Ethiopian wolf ancestry. There is evidence of gene flow between African golden wolves, golden jackals, and grey wolves. One African golden wolf from the Egyptian Sinai Peninsula showed high admixture with the Middle Eastern grey wolves and dogs, highlighting the role of the land bridge between the African and Eurasian continents in canid evolution. African golden wolves form a sister clade to Middle Eastern grey wolves based on mitochondrial DNA, but to coyotes and grey wolves based on nuclear DNA.[4]

Relationship to the Himalayan wolf

Between 2011 and 2015, two mDNA studies found that the Himalayan wolf and Indian grey wolf were closer to the African golden wolf than they were to the Holarctic grey wolf.[22][8] In 2017, a study of mitochondrial DNA, X-chromosome (maternal lineage) markers and Y-chromosome (male lineage) markers found that the Himalayan wolf is genetically basal to the holarctic grey wolf. The Himalayan wolf shares a maternal lineage with the African golden wolf, and possesses a unique paternal lineage that falls between the grey wolf and the African golden wolf.[34]

Subspecies

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.[35]

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.[35]

Subspecies Trinomial authority Description Range Synonyms
Algerian wolf
Canis a. algirensis

Canis anthus algirensis

Wagner, 1841 A dark-coloured subspecies, with a tail marked with three dusky rings. It is similar in size to the red fox.[36] Algeria, Morocco and Tunisia barbarus (C. E. H. Smith, 1839)

grayi (Hilzheimer, 1906)
tripolitanus (Wagner, 1841)

Senegalese wolf
Canis a. anthus

Senegalesegoldenjackal

F. Cuvier, 1820 Similar to lupaster, but smaller and more lightly built, with paler fur and a sharper muzzle.[36] Senegal senegalensis (C. E. H. Smith, 1839)
Serengeti wolf
Canis a. bea

African wolf digging 4

Heller, 1914 Smaller and lighter coloured than the northern forms.[37] Kenya, Northern Tanzania
Egyptian wolf
Canis a. lupaster

Lupaster

Hemprich and Ehrenberg, 1833 A large, stoutly built subspecies with proportionately short ears and presenting a very grey wolf-like phenotype, standing 40.6 cm (16.0 in) in shoulder height and 127 cm (50 in) in body length. The upper parts are yellowish grey tinged with black, while the muzzle, the ears and the outer surfaces of the limbs are reddish yellow. The fur around the mouth is white.[28][36] Egypt, Algeria, Mali, Ethiopian Highlands, and Senegal C. aureus lupaster

C. lupaster
C. lupus lupaster
C. sacer (Hemprich and Ehrenberg, 1833)

Somali wolf
Canis a. riparius
Hemprich and Ehrenberg, 1832 A dwarf subspecies measuring only a dozen inches in shoulder height, it is generally of a greyish-yellow colour, mingled with only a small proportion of black. The muzzle and legs are more decidedly yellow, and the underparts are white.[36] Somalia and coast of Ethiopia and Eritrea hagenbecki (Noack, 1897)

mengesi (Noack, 1897)
somalicus (Lorenz, 1906)

Variegated wolf
Canis a. soudanicus

Atti della Societitaliana di scienze naturali e del Museo civico di storia naturale di Milano (1902) (20160257140)

Thomas, 1903 A small subspecies standing 38 cm (15 in) at the shoulder, and measuring 102 cm (40 in) in length. The fur is generally pale stone-buff, with blotches of black.[36] Sudan and Somalia doederleini (Hilzheimer, 1906)

nubianus (Cabrera, 1921)
thooides (Hilzheimer, 1906)
variegatus (Cretzschmar, 1826)

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".[17] 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.[6] 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.[6]

Canis anthus threat postures
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.[6]

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.[38]

Communication

African wolves frequently groom one another, particularly during courtship, during which it can last up to 30 minutes. 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.[6]

The vocabulary of the African wolf is similar to that of the domestic dog, with seven sounds having been recorded.[19] The African wolf's vocalisations include howls, barks, growls, whines and cackles.[6] Subspecies can be recognised by differences in their howls.[19] 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.[6] 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.[39]

Hunting behaviour

Golden Wolf, navigating Wildebeest, Ngorongoro
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.[6]

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.[6] 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.[38] 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.[6]

Ecology

Distribution and habitat

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.[20] 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]

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.[28] In Egypt, the African wolf inhabits agricultural areas, wastelands, desert margins, rocky areas, and cliffs. At Lake Nasser, it lives close to the lakeshore.[18] In 2012, African wolves were photographed in Morocco's Azilal Province at an elevation of 1,800 metres.[7][41] 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.[42] This wolf has been reported in the very dry Danakil Depression desert on the coast of Eritrea, in eastern Africa.[43]

Diet

Golden wolf eating agama
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.[38] In East Africa, it consumes invertebrates and fruit, though 60% of its diet consists of rodents, lizards, snakes, birds, hares and Thomson's gazelles.[17] During the wildebeest calving season, African wolves will feed almost exclusively on their afterbirth.[19] In the Serengeti and Ngorongoro Crater, less than 20% of its diet comes from scavenging.[6] 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.[28]

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.[44] Nevertheless, the African wolf has been known to kill the pups of black-backed jackals,[17] but has in turn been observed to be dominated by adults during disputes over carcasses.[19] It often eats alongside African wild dogs, and will stand its ground if the dogs try to harass it.[6] 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.[45] Nevertheless, there is at least one record of an African wolf pack adopting a male Ethiopian wolf.[46]

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.[47] Wolves will confront a hyena approaching too closely to their dens by taking turns in biting the hyena's hocks until it retreats.[6]

Wolves in the Serengeti are known to carry the canine parvovirus, canine herpesvirus, canine coronavirus and canine adenovirus.[17]

In folklore

Amulet of a Jackal LACMA 45.23.31
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.[48] According to Diodorus Siculus, the Egyptian city of Lykopolis was named in honour of a pack of wolves that repelled an Ethiopian invasion.[49] 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.[12]

Although considered haram in Islamic dietary laws, the wolf is important in Moroccan folk medicine.[13] 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.[14]

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.[50]

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.[15][51] 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.[15]

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

See also

References

  1. ^ a b (in French) Cuvier, Frédéric (1824), Histoire naturelle des mammifères, tome 2, A Paris : Chez A. Belin ...
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Further reading

  • Cheryl Lyn Dybas, "A Wolf in Jackal's Clothing", Africa Geographic (July 2012)
  • Cheryl Lyn Dybas, "In the Long Shadow of the Pyramids and Beyond: Glimpse of an African... Wolf?", International Wolf Center (Spring 2015)
  • Hugo van Lawick & Jane Goodall (1971), Innocent Killers, Houghton Mifflin Company Boston

External links

Anubis

Anubis (; Ancient Greek: Ἄνουβις, Egyptian: jnpw, Coptic: ⲁⲛⲟⲩⲡ Anoup) is the Greek name of a god associated with mummification and the afterlife in ancient Egyptian religion, usually depicted as a canine or a man with a canine head. Archeologists have identified Anubis's sacred animal as an Egyptian canid, the African golden wolf.Like many ancient Egyptian deities, Anubis assumed different roles in various contexts. Depicted as a protector of graves as early as the First Dynasty (c. 3100 – c. 2890 BC), Anubis was also an embalmer. By the Middle Kingdom (c. 2055 – 1650 BC) he was replaced by Osiris in his role as lord of the underworld. One of his prominent roles was as a god who ushered souls into the afterlife. He attended the weighing scale during the "Weighing of the Heart," in which it was determined whether a soul would be allowed to enter the realm of the dead. Despite being one of the most ancient and "one of the most frequently depicted and mentioned gods" in the Egyptian pantheon, Anubis played almost no role in Egyptian myths.Anubis was depicted in black, a color that symbolized regeneration, life, the soil of the Nile River, and the discoloration of the corpse after embalming. Anubis is associated with Wepwawet (also called Upuaut), another Egyptian god portrayed with a dog's head or in canine form, but with grey or white fur. Historians assume that the two figures were eventually combined. Anubis' female counterpart is Anput. His daughter is the serpent goddess Kebechet.

Arabian wolf

The Arabian wolf (Canis lupus arabs) is a subspecies of gray wolf which lives on the Arabian peninsula. It is the smallest sized wolf known. It is a desert-adapted wolf that normally lives in small groups and is omnivorous, eating carrion and garbage as well as small to medium-sized prey.

Black-backed jackal

The black-backed jackal (Canis mesomelas) is a canid native to two areas of Africa, separated by roughly 900 km.

One region includes the southernmost tip of the continent, including South Africa, Namibia, Botswana, and Zimbabwe. The other area is along the eastern coastline, including Kenya, Somalia, Djibouti, Eritrea, and Ethiopia. It is listed by the IUCN as least concern, due to its widespread range and adaptability, although it is still persecuted as a livestock predator and rabies vector.Compared to other members of the genus Canis, the black-backed jackal is a very ancient species, and has changed little since the Pleistocene, being the most basal wolf-like canine, alongside the closely related side-striped jackal. It is a fox-like animal with a reddish coat and a black saddle that extends from the shoulders to the base of the tail. It is a monogamous animal, whose young may remain with the family to help raise new generations of pups. The black-backed jackal is not a fussy eater, and feeds on small to medium-sized animals, as well as plant matter and human refuse.

Canidae

The biological family Canidae

(from Latin, canis, “dog”) is a lineage of carnivorans that includes domestic dogs, wolves, coyotes, foxes, jackals, dingoes, and many other extant and extinct dog-like mammals. A member of this family is called a canid (, ).The cat-like feliforms and dog-like caniforms emerged within the Carnivoramorpha 43 million years before present. The caniforms included the fox-like genus Leptocyon whose various species existed from 34 million years ago (Mya) before branching 11.9 Mya into Vulpini (foxes) and Canini (canines).Canids are found on all continents except Antarctica, having arrived independently or accompanied human beings over extended periods of time. Canids vary in size from the 2-m-long (6 ft 7 in) gray wolf to the 24-cm-long (9.4 in) fennec fox. The body forms of canids are similar, typically having long muzzles, upright ears, teeth adapted for cracking bones and slicing flesh, long legs, and bushy tails. They are mostly social animals, living together in family units or small groups and behaving co-operatively. Typically, only the dominant pair in a group breeds, and a litter of young is reared annually in an underground den. Canids communicate by scent signals and vocalizations. They are very intelligent. One canid, the domestic dog, long ago entered into a partnership with humans and today remains one of the most widely kept domestic animals.

Canis

Canis is a genus of the Canidae containing multiple extant species, such as wolves, coyotes, jackals, dingoes, and dogs. Species of this genus are distinguished by their moderate to large size, their massive, well-developed skulls and dentition, long legs, and comparatively short ears and tails.

East African wild dog

The East African wild dog (Lycaon pictus lupinus) is a subspecies of African wild dog native to East Africa. It is distinguished from the nominate Cape subspecies by its smaller size and much blacker coat.Its range is patchy, having been eradicated in Uganda and much of Kenya. A small population occupies an area encompassing South Sudan, northern Kenya and probably northern Uganda. It is almost certainly extinct in Rwanda and Burundi. Nevertheless, it remains somewhat numerous in southern Tanzania, particularly in the Selous Game Reserve and Mikumi National Park, both of which are occupied by what could be Africa's largest wild dog population.Artistic depictions of African wild dogs are prominent on cosmetic palettes and other objects from Egypt's predynastic period, likely symbolising order over chaos, as well as the transition between the wild (represented by the African golden wolf) and the domestic (represented by the dog). Predynastic hunters may have also 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.

Egyptian wolf

The Egyptian wolf (Canis anthus lupaster) is a subspecies of African golden wolf native to northern, eastern and western Africa.

El Feidja National Park

El Feidja National Park is located in Northwest Tunisia and has an area of 2,765 hectares (6,830 acres). It is home to many animals, notably, Barbary stag, African golden wolf, and Barbary boar.

Ethiopian wolf

The Ethiopian wolf (Canis simensis) is a canid native to the Ethiopian Highlands. It is similar to the coyote in size and build, and is distinguished by its long and narrow skull, and its red and white fur. Unlike most large canids, which are widespread, generalist feeders, the Ethiopian wolf is a highly specialised feeder of Afroalpine rodents with very specific habitat requirements. It is one of the world's rarest canids, and Africa's most endangered carnivore.The species' current range is limited to seven isolated mountain ranges at altitudes of 3,000–4,500 m, with the overall adult population estimated at 360–440 individuals in 2011, more than half of them in the Bale Mountains.The Ethiopian wolf is listed as endangered by the IUCN, on account of its small numbers and fragmented range. Threats include increasing pressure from expanding human populations, resulting in habitat degradation through overgrazing, and disease transference and interbreeding from free-ranging dogs. Its conservation is headed by Oxford University's Ethiopian Wolf Conservation Programme, which seeks to protect wolves through vaccination and community outreach programs.

Golden jackal

The golden jackal (Canis aureus) is a wolf-like canid that is native to Southeast Europe, Southwest Asia, South Asia, and regions of Southeast Asia. Compared with the Arabian wolf, which is the smallest of the gray wolves (Canis lupus), the jackal is smaller and possesses shorter legs, a shorter tail, a more elongated torso, a less-prominent forehead, and a narrower and more pointed muzzle. The golden jackal's coat can vary in color from a pale creamy yellow in summer to a dark tawny beige in winter. It is listed as Least Concern on the IUCN Red List due to its widespread distribution and high density in areas with plenty of available food and optimum shelter.

The ancestor of the golden jackal is believed to be the extinct Arno river dog that lived in Mediterranean Europe 1.9 million years ago. It is described as having been a small, jackal-like canine. Genetic studies indicate that the golden jackal expanded from India around 20,000 years ago towards the end of the last ice age. The oldest golden jackal fossil, found at the Ksar Akil rock shelter near Beirut, Lebanon, is 20,000 years old. The oldest golden jackal fossils in Europe were found in Greece and are 7,000 years old. There are seven subspecies of the golden jackal. The golden jackal is more closely related to the gray wolf, coyote, African golden wolf, and Ethiopian wolf than it is to the African black-backed jackal or side-striped jackal. It is capable of producing fertile hybrids with both the gray wolf and the African golden wolf. Jackal–dog hybrids called Sulimov dogs are in service at the Sheremetyevo Airport near Moscow where they are deployed by the Russian airline Aeroflot for scent-detection.

Golden jackals are abundant in valleys and beside rivers and their tributaries, canals, lakes, and seashores. They are rare in foothills and low mountains. The golden jackal is a social species, the basic social unit of which consists of a breeding pair and any young offspring. It is very adaptable, with the ability to exploit food ranging from fruit and insects to small ungulates. They will attack domestic fowl and domestic mammals up to the size of domestic water buffalo calves. The jackal's competitors are the red fox, wolf, jungle cat, forest wildcat, and, in the Caucasus, the raccoon, and, in Central Asia, the steppe wildcat. The jackal is expanding beyond its native grounds in Southeast Europe into Central Europe, occupying areas where there are few or no wolves.

Himalayan wolf

The Himalayan wolf (Canis lupus) is a canine of debated taxonomy. It is distinguished by its genetic markers, with mitochondrial DNA indicating that it is genetically basal to the Holarctic grey wolf, genetically the same wolf as the Tibetan wolf, and has an association with the African golden wolf (Canis anthus). There are no striking morphological differences between the wolves from the Indian Himalayas and those from Tibet. The wolf is found in northern India in the Ladakh region of eastern Kashmir, and the Lahaul and Spiti region in the northeastern part of Uttarakhand and Himachal Pradesh . It is also found in the Himalayan regions of Nepal.

Jackal

Jackals are medium-sized omnivorous mammals of the genus Canis, which also includes wolves, coyotes and the domestic dog. While the word "jackal" has historically been used for many small canids, in modern use it most commonly refers to three species: the closely related black-backed jackal and side-striped jackal of sub-Saharan Africa, and the golden jackal of south-central Eurasia, which is more closely related to other members of the genus Canis.

Jackals and coyotes (sometimes called the "American jackal") are opportunistic omnivores, predators of small to medium-sized animals and proficient scavengers. Their long legs and curved canine teeth are adapted for hunting small mammals, birds, and reptiles, and their large feet and fused leg bones give them a physique well-suited for long-distance running, capable of maintaining speeds of 16 km/h (9.9 mph) for extended periods of time. Jackals are crepuscular, most active at dawn and dusk.

Their most common social unit is a monogamous pair, which defends its territory from other pairs by vigorously chasing intruding rivals and marking landmarks around the territory with their urine and feces. The territory may be large enough to hold some young adults, which stay with their parents until they establish their own territories. Jackals may occasionally assemble in small packs, for example, to scavenge a carcass, but they normally hunt either alone or in pairs.

List of fauna of Sudan and South Sudan

Fauna of Sudan and South Sudan include:

Aardvark

Aardwolf

African buffalo

African bush elephant

African civet

African golden wolf

African leopard

Ball Python

Banded mongoose

Lion

Barbary sheep

Black-backed jackal

Blue duiker

Bohor reedbuck

Bongo

Bushbuck

Cape hyrax

Common duiker

Common genet

Congo lion

Dama gazelle

Dorcas gazelle

Dugong

Gemsbok

Giant eland

Giant forest hog

Grant's gazelle

Grant's zebra

Greater kudu

Grevy's zebra

Hartebeest

Hippopotamus

Klipspringer

Kob

Maneless zebra

Marsh mongoose

Nile lechwe

North African ostrich

Northern white rhinoceros

Nubian giraffe

Nubian wild ass

Okapi

Oribi

Pale fox

Plains zebra

Red fox

Red river hog

Roan antelope

Rothschild's giraffe

Rueppell's fox

Side-striped jackal

Sitatunga

Somali wild ass

Somali wild dog

Spotted hyena

Striped hyena

Sudan cheetah

Temminck's pangolin

Thomson's gazelle

Warthog

Waterbuck

Yellow-backed duiker

Lutrogale

Lutrogale is a genus of otters, with only one extant species—the smooth-coated otter.

Senegalese wolf

The Senegalese wolf (Canis anthus anthus), also known as the grey jackal, slender jackal or Anthus, is the nominate subspecies of the African golden wolf native to Senegal.

Side-striped jackal

The side-striped jackal (Canis adustus) is a species of jackal, native to east and southern Africa. Unlike its cousin, the smaller black-backed jackal, which dwells in open plains, the side-striped jackal primarily dwells in woodland and scrub areas.

Tibetan wolf

The Tibetan wolf (Canis lupus filchneri) is a subspecies of the gray wolf that is native to China in the regions of Gansu, Qinghai, and the Tibetan Autonomous Region. It is distinguished by its genetic markers, with whole genome sequencing indicating that it is the most genetically divergent wolf population, and mitochondrial DNA sequencing indicating that it is genetically the same wolf as the Himalayan wolf, is genetically basal to the Holarctic grey wolf, and has an association with the African golden wolf (Canis anthus).

Variegated wolf

The variegated wolf (Canis anthus soudanicus), also known as the Nubian wolf, is a subspecies of African golden wolf native to Sudan and Somalia. It is smaller and more lightly built than the Egyptian wolf, standing 38 cm (15 in) at the shoulder, and 102 cm (40 in) in length. Compared with more the wolf-like Egyptian wolf, the variegated wolf is built more like a greyhound. The ears are somewhat larger than the Egyptian wolf's and the body colour is generally pale stone-buff, with blotches of black. Its main habitat consists of rocky regions, where it feeds on small mammals and birds. The subspecies has been encountered in elevations of up to 5000 feet in the highlands of Abyssinia.

Extant Carnivora species

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