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African wolf

The African wolf (Canis lupaster) or golden wolf, formerly known as the African golden jackal, is a canine native to North Africa, West Africa, the Sahel, northern East Africa, and the Horn of Africa. It is listed as least concern on the IUCN Red List.[1] In the Middle Atlas in Morocco, it was sighted in elevations as high as 1,800 m (5,900 ft).[3] It is primarily a predator of invertebrates and mammals as large as gazelle fawns, though larger animals are sometimes taken. Its diet also includes animal carcasses, human refuse, and fruit. The African wolf is a monogamous and territorial species; offspring remain with the family to assist in raising their parents' younger pups.[4]

African wolf
Temporal range: Middle Pleistocene – Recent 0.6-0 Ma
African wolf in Haizer, Algeria
African wolves howling
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Carnivora
Family: Canidae
Genus: Canis
Species:
C. lupaster
Binomial name
Canis lupaster
Subspecies
Synonyms

Canis anthus F. Cuvier, 1820[2]

It was previously classified as an African variant of the golden jackal (Canis aureus), with at that time at least one subspecies (C. a. lupaster) having been classified as a wolf. In 2015, a series of analyses on the species' mitochondrial DNA and nuclear genome demonstrated that it was, in fact, distinct from the golden jackal, and more closely related to the gray wolf and the coyote (Canis latrans).[5][6] It is nonetheless still close enough to the golden jackal to produce hybrid offspring, as indicated through genetic tests on jackals in Israel,[5] and a 19th-century captive crossbreeding experiment.[7] Further studies demonstrated that it is the descendant of a genetically admixed canid of 72% gray wolf (Canis lupus) and 28% Ethiopian wolf (Canis simensis) ancestry.[8]

It plays a prominent role in some African cultures; it was considered sacred in ancient Egypt, particularly in Lycopolis, where it was venerated as a god. In North African folklore, it is viewed as an untrustworthy animal whose body parts can be used for medicinal or ritualistic purposes,[9][10][11] while it is held in high esteem in Senegal's Serer religion as being the first creature to be created by the god Roog.[12]

Local and indigenous names

Description

 
Skull

The African wolf is intermediate in size between the African jackals (L. mesomelas and L. adustus) and the small subspecies of gray wolves,[19] 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.[19] It has a relatively long snout and ears, while the tail is comparatively short, measuring 20 cm in length. Fur color varies individually, seasonally and geographically, though the typical coloration 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-colored. Females bear two to four pairs of teats.[4] Although superficially similar to the golden jackal (particularly in East Africa), the African wolf has a more pointed muzzle and sharper, more robust teeth.[5] The ears are longer in the African wolf, and the skull has a more elevated forehead.[20]

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

Taxonomy

Early writings

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".[21][22]

The African wolf was first recognised as being a separate species from the 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.[2] Eduard Rüppell proposed that the animal was the ancestor of Egyptian sighthounds, and named it Wolfs-hund (wolf dog),[23] while C.H. Smith named it "thoa" or "thous dog".[24]

 
Illustration of golden jackal-African wolf hybrids bred in captivity (1821).

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 pup inherited the golden jackal's shyness, while the other was affectionate toward its human captors.[7] English biologist G.J. Mivart emphasized the differences between the African wolf and the golden jackal in his writings:

... 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 are 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 reason 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)[25]
 
Comparative illustration of C. aureus (top) and C. lupaster (bottom).

The canids present in Egypt in particular were noted to be so much more gray wolf-like than populations elsewhere in Africa that W.F. Hemprich and C.G. Ehrenberg gave them the binomial name Canis lupaster in 1832. Likewise, T.H. Huxley, upon noting the similarities between the skulls of lupaster and Indian wolves, classed the animal as a subspecies of the gray 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.[26]

In 1981, zoologist Walter Ferguson argued in favor of lupaster being a subspecies of the gray 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.[22]

21st-century discoveries

Phylogenetic tree of the wolf-like canids with timing in millions of years[a]
Caninae 3.5 Ma
3.0
2.5
2.0
0.96
0.6
0.38
0.11

African wolf: northwestern Africa 

African wolf: eastern Africa 

Golden jackal  

Ethiopian wolf  

Dhole  

African wild dog  

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 did not correspond to that of the golden jackal or the six other recognized species of the area, but strongly resembled that of the gray 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).[13]

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[19] were found to have mtDNA sequences more closely resembling those found in gray wolves than those of golden jackals.[21] 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 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 the gray wolf.[27]

In 2015, a more thorough comparative study of mitochondrial and nuclear genomes on a larger sample of wolf-like African canids from northern, eastern and western Africa showed that they were in fact all distinct from the golden jackal, with a genetic divergence of around 6.7%,[5][28][29] which is greater than that between gray wolves and coyotes (4%) and that between gray wolves and domestic dogs (0.2%).[30] Furthermore, the study showed that these African wolf-like canids (renamed Canis lupaster, or African wolves) were more closely related to gray wolves and coyotes than to golden jackals,[5][31] and that C. l. lupaster merely represents a distinct phenotype of the African wolf rather than an actual gray wolf. The phylogenetic tree below is based on nuclear sequences:[5]

It was estimated that the African wolf diverged from the wolf–coyote clade 1.0–1.7 million years ago, during the Pleistocene, and therefore its superficial similarity to the 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 African wolf DNA were identified in golden jackals in Israel, which adjoins Egypt, thus indicating the presence of a hybrid zone.[5] The study's findings were corroborated that same year by Spanish, Mexican and Moroccan scientists analyzing the mtDNA of wolves in Morocco, who found that the specimens analyzed were distinct from both golden jackals and gray wolves but bore a closer relationship to the latter.[6] Studies on RAD sequences found instances of African wolves hybridizing with both feral dogs and Ethiopian wolves.[32]

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 does not 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.[19] The following year, a major genetic study of Canis species also referred to the African wolf as Canis lupaster.[8]

In 2019, a workshop hosted by the IUCN/SSC Canid Specialist Group recommended that because the specimen identified as Canis anthus Cuvier, 1820 was uncertain, the species should be known as Canis lupaster Hemprich and Ehrenberg, 1832 until Canis anthus can be validated.[33]

Admixture with other Canis species

In 2018, whole genome sequencing was used to compare members of the genus Canis. The study supports the African wolf being distinct from the golden jackal, and with the Ethiopian wolf being genetically basal to both. Two genetically distinct African wolf populations 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 wolf populations was a genetically admixed canid of 72% gray wolf and 28% Ethiopian wolf ancestry. There is evidence of gene flow between African wolves, golden jackals, and gray wolves. One African wolf from the Egyptian Sinai Peninsula showed high admixture with the Middle Eastern gray wolves and dogs, highlighting the role of the land bridge between the African and other continents in canid evolution. African wolves form a sister clade to Middle Eastern gray wolves based on mitochondrial DNA, but to coyotes and gray wolves based on nuclear DNA.[8]

Relationship to the Himalayan wolf

Between 2011 and 2015, two mtDNA studies found that the Himalayan wolf and Indian wolf were closer to the African wolf than they were to the Holarctic gray wolf.[21][5] 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 gray wolf. The Himalayan wolf shares a maternal lineage with the African wolf, and possesses a unique paternal lineage that falls between the gray wolf and the African 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 10 supposed West African subspecies were named or classed almost entirely because of their fur color.[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] However, a study showed that the genetic divergence of all of the African wolves occurred between 50,000 and 10,500 years ago, with most occurring between 30,000 and 16,000 years ago during the Late Glacial Maximum (33,000–16,000 years ago). There were very dry conditions across the Sahara during this period. The study proposes that these wolves were isolated in refugia and therefore isolated for hundreds of generations, leading to genetic divergence.[36]

Behavior

Social and reproductive behaviors

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 in relation to those of the black-backed jackal; although the sexual and territorial behavior 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. l. lupaster (left) and C. l. 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 behavior. 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 meters from the den, even sleeping in the open. Their playing behavior 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 centers 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.[39]

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.[4]

The vocalisations of the African wolf are similar to those of the domestic dog, with seven sounds having been recorded,[17] including 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 and establish territorial status.[4] A comparative analysis of African wolf and some gray wolf subspecies' howls demonstrated that the former's howls bear similarities to those of the Indian wolf, being high-pitched and of relatively short duration.[40]

Hunting behavior

 
A Serengeti wolf (C. l. 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.[39] 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 in mid-air or by pouncing on them while they are on the ground. 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

Distribution and habitat

C. lupaster has a wide range across the upper half of Africa, occurring in Senegal, Burkina Faso, Cameroon, the Central African Republic, Djibouti, Eritrea, Ethiopia, Guinea, Mali, Mauritania, Niger, Somalia, South Sudan, Sudan, Western Sahara, Nigeria, Chad, Morocco, Algeria, Tunisia, Libya, Kenya, Egypt, and Tanzania. 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.[19] 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 and 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.[41]

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.[27] 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 meters.[14][3] 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.[13]

Diet

 
Serengeti wolf (C. l. 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.[39] 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. l. anthus and C. l. 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.[27]

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 favored by the latter two species) and being more active during the daytime.[43] 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 antagonistic, 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.[44] Nevertheless, there is at least one record of an African wolf pack adopting a male Ethiopian wolf.[45]

African 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.[46] Wolves will confront a hyena approaching too closely to their dens by taking turns in biting the hyena's hocks until it retreats.[4]

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

In culture

 
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.[47] The wolf was sacred in Lycopolis, whose inhabitants would mummify wolves and store them in chambers, as opposed to other areas of Egypt, where wolves were buried at their place of death. According to Diodorus Siculus in Bibliotheca historica, there were two reasons as to why the wolf was held in such high regard, the first being the animal's affinity to the dog, and the second being a legend that told of how Lycopolis received its name after a pack of wolves repelled an Ethiopian invasion. Plutarch noted in his On the Worship of Isis and Osiris that Lycopolis was the only nome in Egypt where people consumed sheep, as the practice was associated with the wolf, which was revered as a god. The importance of the wolf in Lycopolite culture continued through to the Roman period, where images of the animal were minted on the reverse sides of coins. Herodotus mockingly wrote of a festival commemorating Rhampsinit's descent to the underworld where a priest would be lead by two wolves to the temple of Ceres.[48]

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.[9] Although considered haram in Islamic dietary laws, the wolf is important in Moroccan folk medicine.[10] 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.[11]

The African wolf is not 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.[49]

The African 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.[12][50] 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.[12]

See also

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

References

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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

  • (in Arabic) Le fantôme du moyen Atlas شبح الأطلس المتوسط, AmouddouTV (Feb 13, 2014)
  • (in Arabic) Nouveau!! Nouvelle espèce découverte... اكتشاف نوع جديد من الحيوانات/ حصري, AmouddouTV (Aug 15, 2015)

african, wolf, this, article, about, wolf, native, africa, other, wolf, africa, ethiopian, wolf, canis, lupaster, golden, wolf, formerly, known, african, golden, jackal, canine, native, north, africa, west, africa, sahel, northern, east, africa, horn, africa, . This article is about a wolf native to Africa For the other wolf of Africa see Ethiopian wolf The African wolf Canis lupaster or golden wolf formerly known as the African golden jackal is a canine native to North Africa West Africa the Sahel northern East Africa and the Horn of Africa It is listed as least concern on the IUCN Red List 1 In the Middle Atlas in Morocco it was sighted in elevations as high as 1 800 m 5 900 ft 3 It is primarily a predator of invertebrates and mammals as large as gazelle fawns though larger animals are sometimes taken Its diet also includes animal carcasses human refuse and fruit The African wolf is a monogamous and territorial species offspring remain with the family to assist in raising their parents younger pups 4 African wolfTemporal range Middle Pleistocene Recent 0 6 0 MaAfrican wolf in Haizer Algeria source source African wolves howlingConservation statusLeast Concern IUCN 3 1 1 Scientific classificationKingdom AnimaliaPhylum ChordataClass MammaliaOrder CarnivoraFamily CanidaeGenus CanisSpecies C lupasterBinomial nameCanis lupasterHemprich and Ehrenberg 1832 1 SubspeciesC l algirensis C l anthus C l bea C l lupaster C l riparius C l soudanicusSynonymsCanis anthus F Cuvier 1820 2 It was previously classified as an African variant of the golden jackal Canis aureus with at that time at least one subspecies C a lupaster having been classified as a wolf In 2015 a series of analyses on the species mitochondrial DNA and nuclear genome demonstrated that it was in fact distinct from the golden jackal and more closely related to the gray wolf and the coyote Canis latrans 5 6 It is nonetheless still close enough to the golden jackal to produce hybrid offspring as indicated through genetic tests on jackals in Israel 5 and a 19th century captive crossbreeding experiment 7 Further studies demonstrated that it is the descendant of a genetically admixed canid of 72 gray wolf Canis lupus and 28 Ethiopian wolf Canis simensis ancestry 8 It plays a prominent role in some African cultures it was considered sacred in ancient Egypt particularly in Lycopolis where it was venerated as a god In North African folklore it is viewed as an untrustworthy animal whose body parts can be used for medicinal or ritualistic purposes 9 10 11 while it is held in high esteem in Senegal s Serer religion as being the first creature to be created by the god Roog 12 Contents 1 Local and indigenous names 2 Description 3 Taxonomy 3 1 Early writings 3 2 21st century discoveries 3 3 Admixture with other Canis species 3 4 Relationship to the Himalayan wolf 3 5 Subspecies 4 Behavior 4 1 Social and reproductive behaviors 4 2 Communication 4 3 Hunting behavior 5 Ecology 5 1 Distribution and habitat 5 2 Diet 5 3 Enemies and competitors 6 In culture 7 See also 8 Notes 9 References 10 Further reading 11 External linksLocal and indigenous names EditLinguistic group or area Indigenous nameAfar Wucharia 13 Amazigh Ouchan asian 14 Ouchan akhatar 14 Amharic ተረ ቀበሮ Tera kebero 15 Arabic ابن آوى Ibn awa 16 Fula 𞤧𞤵𞤲 𞤣𞤵 Sundu 15 Hausa Kyarkeci 15 Somali EeySonghai Nzongo 15 Swahili standard Swahili Tanzania Bweha wa mbuga 17 Bweha dhahabu 15 Tigrinya ቡኳርያ bukharya 18 Wolof Tili 15 Description Edit Skull The African wolf is intermediate in size between the African jackals L mesomelas and L adustus and the small subspecies of gray wolves 19 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 19 It has a relatively long snout and ears while the tail is comparatively short measuring 20 cm in length Fur color varies individually seasonally and geographically though the typical coloration 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 colored Females bear two to four pairs of teats 4 Although superficially similar to the golden jackal particularly in East Africa the African wolf has a more pointed muzzle and sharper more robust teeth 5 The ears are longer in the African wolf and the skull has a more elevated forehead 20 Various C lupaster phenotypes ranging from gracile jackal like morphs to more robust wolf like ones Taxonomy EditEarly writings Edit 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 21 22 The African wolf was first recognised as being a separate species from the golden jackal by Frederic 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 2 Eduard Ruppell proposed that the animal was the ancestor of Egyptian sighthounds and named it Wolfs hund wolf dog 23 while C H Smith named it thoa or thous dog 24 Illustration of golden jackal African wolf hybrids bred in captivity 1821 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 pup inherited the golden jackal s shyness while the other was affectionate toward its human captors 7 English biologist G J Mivart emphasized the differences between the African wolf and the golden jackal in his writings 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 are 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 reason 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 25 Comparative illustration of C aureus top and C lupaster bottom The canids present in Egypt in particular were noted to be so much more gray wolf like than populations elsewhere in Africa that W F Hemprich and C G Ehrenberg gave them the binomial name Canis lupaster in 1832 Likewise T H Huxley upon noting the similarities between the skulls of lupaster and Indian wolves classed the animal as a subspecies of the gray wolf However the animal was subsequently synonymised with the golden jackal by Ernst Schwarz in 1926 In 1965 the Finnish paleontologist Bjorn Kurten 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 26 In 1981 zoologist Walter Ferguson argued in favor of lupaster being a subspecies of the gray 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 22 21st century discoveries Edit Phylogenetic tree of the wolf like canids with timing in millions of years a Caninae 3 5 Ma 3 0 2 5 2 0 0 96 0 6 0 38 Domestic dog Gray wolf Coyote 0 11 African wolf northwestern Africa African wolf eastern Africa Golden jackal Ethiopian wolf Dhole African wild dog 2 6 Side striped jackal Black backed jackal 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 did not correspond to that of the golden jackal or the six other recognized species of the area but strongly resembled that of the gray 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 13 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 19 were found to have mtDNA sequences more closely resembling those found in gray wolves than those of golden jackals 21 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 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 the gray wolf 27 In 2015 a more thorough comparative study of mitochondrial and nuclear genomes on a larger sample of wolf like African canids from northern eastern and western Africa showed that they were in fact all distinct from the golden jackal with a genetic divergence of around 6 7 5 28 29 which is greater than that between gray wolves and coyotes 4 and that between gray wolves and domestic dogs 0 2 30 Furthermore the study showed that these African wolf like canids renamed Canis lupaster or African wolves were more closely related to gray wolves and coyotes than to golden jackals 5 31 and that C l lupaster merely represents a distinct phenotype of the African wolf rather than an actual gray wolf The phylogenetic tree below is based on nuclear sequences 5 It was estimated that the African wolf diverged from the wolf coyote clade 1 0 1 7 million years ago during the Pleistocene and therefore its superficial similarity to the 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 African wolf DNA were identified in golden jackals in Israel which adjoins Egypt thus indicating the presence of a hybrid zone 5 The study s findings were corroborated that same year by Spanish Mexican and Moroccan scientists analyzing the mtDNA of wolves in Morocco who found that the specimens analyzed were distinct from both golden jackals and gray wolves but bore a closer relationship to the latter 6 Studies on RAD sequences found instances of African wolves hybridizing with both feral dogs and Ethiopian wolves 32 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 does not 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 fur Naturkunde 19 The following year a major genetic study of Canis species also referred to the African wolf as Canis lupaster 8 In 2019 a workshop hosted by the IUCN SSC Canid Specialist Group recommended that because the specimen identified as Canis anthus Cuvier 1820 was uncertain the species should be known as Canis lupaster Hemprich and Ehrenberg 1832 until Canis anthus can be validated 33 Admixture with other Canis species Edit In 2018 whole genome sequencing was used to compare members of the genus Canis The study supports the African wolf being distinct from the golden jackal and with the Ethiopian wolf being genetically basal to both Two genetically distinct African wolf populations 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 wolf populations was a genetically admixed canid of 72 gray wolf and 28 Ethiopian wolf ancestry There is evidence of gene flow between African wolves golden jackals and gray wolves One African wolf from the Egyptian Sinai Peninsula showed high admixture with the Middle Eastern gray wolves and dogs highlighting the role of the land bridge between the African and other continents in canid evolution African wolves form a sister clade to Middle Eastern gray wolves based on mitochondrial DNA but to coyotes and gray wolves based on nuclear DNA 8 Relationship to the Himalayan wolf Edit Between 2011 and 2015 two mtDNA studies found that the Himalayan wolf and Indian wolf were closer to the African wolf than they were to the Holarctic gray wolf 21 5 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 gray wolf The Himalayan wolf shares a maternal lineage with the African wolf and possesses a unique paternal lineage that falls between the gray wolf and the African wolf 34 Subspecies Edit 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 10 supposed West African subspecies were named or classed almost entirely because of their fur color 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 However a study showed that the genetic divergence of all of the African wolves occurred between 50 000 and 10 500 years ago with most occurring between 30 000 and 16 000 years ago during the Late Glacial Maximum 33 000 16 000 years ago There were very dry conditions across the Sahara during this period The study proposes that these wolves were isolated in refugia and therefore isolated for hundreds of generations leading to genetic divergence 36 Subspecies Trinomial authority Description Range SynonymsAlgerian wolfC l algirensis Wagner 1841 A dark colored subspecies with a tail marked with three dusky rings It is similar in size to the red fox 37 Algeria Morocco and Tunisia barbarus C E H Smith 1839 grayi Hilzheimer 1906 tripolitanus Wagner 1841 Senegalese wolfC l anthus F Cuvier 1820 Similar to lupaster but smaller and more lightly built with paler fur and a sharper muzzle 37 Senegal senegalensis C E H Smith 1839 Serengeti wolfC l bea Heller 1914 Smaller and lighter colored than the northern forms 38 Kenya northern TanzaniaEgyptian wolfC l lupaster Hemprich and Ehrenberg 1833 A large stoutly built subspecies with proportionately short ears and presenting a very gray 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 gray 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 27 37 Egypt Algeria Mali Ethiopian Highlands and Senegal C aureus lupasterC lupus lupasterC lupasterC sacer Hemprich and Ehrenberg 1833 Somali wolfC l riparius Hemprich and Ehrenberg 1832 A dwarf subspecies measuring only 12 inches in shoulder height it is generally of a grayish yellow color mingled with only a small proportion of black The muzzle and legs are more decidedly yellow and the underparts are white 37 Somalia and the coast of Ethiopia and Eritrea hagenbecki Noack 1897 mengesi Noack 1897 somalicus Lorenz 1906 Variegated wolfor Nubian wolfC l soudanicus 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 37 Sudan and Somalia doederleini Hilzheimer 1906 nubianus Cabrera 1921 thooides Hilzheimer 1906 variegatus Cretzschmar 1826 Behavior EditSocial and reproductive behaviors Edit 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 in relation to those of the black backed jackal although the sexual and territorial behavior 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 l lupaster left and C l 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 behavior 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 meters from the den even sleeping in the open Their playing behavior 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 centers 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 39 Communication Edit 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 4 The vocalisations of the African wolf are similar to those of the domestic dog with seven sounds having been recorded 17 including 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 and establish territorial status 4 A comparative analysis of African wolf and some gray wolf subspecies howls demonstrated that the former s howls bear similarities to those of the Indian wolf being high pitched and of relatively short duration 40 Hunting behavior Edit A Serengeti wolf C l 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 39 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 in mid air or by pouncing on them while they are on the ground 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 EditDistribution and habitat Edit C lupaster has a wide range across the upper half of Africa occurring in Senegal Burkina Faso Cameroon the Central African Republic Djibouti Eritrea Ethiopia Guinea Mali Mauritania Niger Somalia South Sudan Sudan Western Sahara Nigeria Chad Morocco Algeria Tunisia Libya Kenya Egypt and Tanzania 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 19 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 and 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 41 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 27 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 meters 14 3 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 13 Diet Edit Serengeti wolf C l 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 39 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 l anthus and C l 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 27 Enemies and competitors Edit 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 favored by the latter two species and being more active during the daytime 43 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 antagonistic 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 44 Nevertheless there is at least one record of an African wolf pack adopting a male Ethiopian wolf 45 African 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 46 Wolves will confront a hyena approaching too closely to their dens by taking turns in biting the hyena s hocks until it retreats 4 African wolves in the Serengeti are known to carry the canine parvovirus canine herpesvirus canine coronavirus and canine adenovirus 15 In culture EditMain article Wolves in folklore religion and mythology 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 47 The wolf was sacred in Lycopolis whose inhabitants would mummify wolves and store them in chambers as opposed to other areas of Egypt where wolves were buried at their place of death According to Diodorus Siculus in Bibliotheca historica there were two reasons as to why the wolf was held in such high regard the first being the animal s affinity to the dog and the second being a legend that told of how Lycopolis received its name after a pack of wolves repelled an Ethiopian invasion Plutarch noted in his On the Worship of Isis and Osiris that Lycopolis was the only nome in Egypt where people consumed sheep as the practice was associated with the wolf which was revered as a god The importance of the wolf in Lycopolite culture continued through to the Roman period where images of the animal were minted on the reverse sides of coins Herodotus mockingly wrote of a festival commemorating Rhampsinit s descent to the underworld where a priest would be lead by two wolves to the temple of Ceres 48 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 9 Although considered haram in Islamic dietary laws the wolf is important in Moroccan folk medicine 10 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 11 The African wolf is not common in Neolithic rock art though it does occasionally appear a definite portrayal is shown on the Kef Messiouer cave in Algeria s Tebessa 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 49 The African 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 12 50 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 12 See also EditEthiopian wolfNotes Edit For a full set of supporting references refer to the note a in the phylotree at Evolution of the wolf Wolf like canidsReferences Edit a b c Hoffmann M amp Atickem A 2019 Canis lupaster IUCN Red List of Threatened Species 2019 e T118264888A118265889 doi 10 2305 IUCN UK 2019 1 RLTS T118264888A118265889 en Retrieved 19 November 2021 a b Cuvier F 1824 Le Chacal du Senegal The Senegalese jackal Histoire naturelle des mammiferes in French Vol Tome 2 Paris FR A Belin pp 34 36 via Internet Archive archive org a b in Spanish Moliner V U Ramirez C Gallardo M amp Idrissi H R 2012 Detectan el lobo en Marruecos gracias al uso del foto trampeo Quercus 319 14 15 ISSN 0212 0054 a b c d e f g h i j k l m n o Estes R 1992 The behavior guide to African mammals including hoofed mammals carnivores primates University of California Press pp 398 404 ISBN 0 520 08085 8 a b c d e f g h 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 Alvares 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 Genome wide evidence reveals that African and Eurasian golden jackals are distinct species Current Biology 25 16 2158 2165 doi 10 1016 j cub 2015 06 060 PMID 26234211 a b in Spanish Urios Vicente Donat Torres Maria P Ramirez Carlos Monroy Vilchis Octavio Hamid Rgribi Idrissi 2015 El analisis del genoma mitocondrial del canido estudiado en Marruecos manifiesta que no es ni lobo Canis lupus ni chacal euroasiatico Canis aureus figshare doi 10 6084 m9 figshare 1524971 a b Cuvier F 1824 Histoire naturelle des mammiferes Natural History of the Mammals in French Vol tome 3 Paris FR Chez A Belin via Internet Archive archive org a b c Gopalakrishnan Shyam Sinding Mikkel Holger S Ramos Madrigal Jazmin Niemann Jonas Samaniego Castruita Jose A Vieira Filipe G Caroe Christian de Montero Marc Manuel Kuderna Lukas Serres Aitor Gonzalez Basallote Victor Manuel Liu Yan Hu Wang Guo Dong Marques Bonet Tomas Mirarab Siavash Fernandes Carlos Gaubert Philippe Koepfli Klaus Peter Budd Jane Rueness Eli Knispel Heide Jorgensen Mads Peter Petersen Bent Sicheritz Ponten Thomas Bachmann Lutz Wiig Oystein Hansen Anders J Gilbert M Thomas P 2018 Interspecific gene flow shaped the evolution of the genus Canis Current Biology 28 21 3441 3449 e5 doi 10 1016 j cub 2018 08 041 PMC 6224481 PMID 30344120 a b Osborn D amp Helmy I 1980 Canis aureus lupaster Hemprich and Ehrenberg 1833 The contemporary land mammals of Egypt including Sinai Chicago Field Museum of Natural History pp 360 371 a b Bidwell M Bidwell R 2005 The Traveller s Companion Second ed Tauris Parke Paperbacks ISBN 1845111079 a b Westermarck E 2013 Ritual and Belief in Morocco Vol II Routledge Revivals ISBN 978 1317912613 a b c Gravrand H 1990 La Civilisation Sereer Pangool Les Nouvelles Editions Africaines du Senegal in French Vol 2 pp 201 203 ISBN 2 7236 1055 1 a b c Tiwari Jugal Sillero Zubiri Claudio 2004 Unidentified canid in the Danakil desert of Eritrea Horn of Africa PDF Canid News 7 5 1 3 Archived from the original PDF on 2007 10 12 Retrieved 2015 08 13 a b c in French Rachid Tarik Le loup apparait dans la region d Azilal Le Matin April 29 2013 a b c d e f g h i j Jhala Y V amp Moehlman P D 2004 Golden jackal Canis aureus Archived 2007 10 12 at the Wayback Machine In Sillero Zubiri C Hoffman M amp 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 a b Hoath R 2009 A Field Guide to the Mammals of Egypt Cairo American University in Cairo Press pp 70 73 ISBN 978 9774162541 a b c d e Kingdon J 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 Aerts R 2019 Forest and woodland vegetation in the highlands of Dogu a Tembien In Nyssen J Jacob M Frankl A eds Geo trekking in Ethiopia s Tropical Mountains The Dogu a Tembien District GeoGuide Cham SpringerNature pp 233 250 doi 10 1007 978 3 030 04955 3 15 ISBN 978 3 030 04954 6 S2CID 199103486 a b c d e 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 Mivart G 1890 The Nort African Jackal Dogs Jackals Wolves and Foxes A Monograph of the Canidae London R H Porter pp 41 43 a b c Rueness Eli Knispel Asmyhr Maria Gulbrandsen Sillero Zubiri Claudio MacDonald David W Bekele Afework Atickem Anagaw Stenseth Nils C 2011 The cryptic African wolf Canis aureus lupaster is not a golden jackal and is not endemic to Egypt PLOS ONE 6 1 e16385 Bibcode 2011PLoSO 616385R doi 10 1371 journal pone 0016385 PMC 3027653 PMID 21298107 a b 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 45 4 459 465 doi 10 1515 mamm 1981 45 4 459 S2CID 86455216 Ruppell E 1826 Canis anthus Harrier dog Atlas zu der Reise im nordlichen Afrika Atlas of a Voyage through Northern Africa in German Frankfurt am Main Senckenbergische Naturforschende Gesellschaft pp 44 46 via Internet Archive archive org Smith C H Jardine W 1839 The Natural History of Dogs Vol I Edinburgh UK W H Lizars pp 193 194 via Internet Archive archive org Canidae or genus Canis of authors including also the genera hyaena and proteles Mivart G J 1890 Dogs Jackals Wolves and Foxes A monograph of the Canidae London UK R H Porter pp 36 37 via Internet Archive archive org Kurten B 1965 The Carnivora of the Palestine Caves Acta Zoologica Fennica Societas pro fauna et flora Fennica 107 41 hdl 10138 37761 a b c d 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 Bibcode 2012PLoSO 742740G doi 10 1371 journal pone 0042740 PMC 3416759 PMID 22900047 Lewis M E 2017 Chapter 2 Carnivore guilds and hominim dispersals In Boivin Nicole Crassard Remy Petraglia Michael eds Human Dispersal and Species Movement Cambridge University Press p 52 ISBN 9781107164147 Canis anthus F Cuvier 1820 Integrated Taxonomic Information System itis gov U S Geological Survey Retrieved 8 November 2017 Wayne Robert K 1993 Molecular evolution of the dog family Trends in Genetics 9 6 218 224 doi 10 1016 0168 9525 93 90122 X PMID 8337763 Zachos Frank E 2016 Chapter 6 Species delimitations Species Concepts in Biology Historical development theoretical foundations and practical relevance Springer p 158 ISBN 9783319449647 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 Alvares Francisco Bogdanowicz Wieslaw Campbell Liz A D Godinho Rachel Hatlauf Jennifer Jhala Yadvendradev V Kitchener Andrew C Koepfli Klaus Peter Krofel Miha Moehlman Patricia D Senn Helen Sillero Zubiri Claudio Viranta Suvi Werhahn Geraldine 28 30 May 2019 Workshop conclusions and recommendations PDF Old World Canis spp with taxonomic ambiguity CIBIO Vairao Portugal IUCN SSC Canid Specialist Group Retrieved 6 March 2020 via canids org CBC Werhahn G Senn H Kaden J Joshi J Bhattarai S Kusi N Sillero Zubiri C MacDonald D 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 Bibcode 2017RSOS 470186W doi 10 1098 rsos 170186 PMC 5493914 PMID 28680672 a b 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 Sarabia Carlos Vonholdt Bridgett Larrasoana Juan C Urios Vicente Leonard Jennifer A 2021 Pleistocene climate fluctuations drove demographic history of African wolves Ca nis lupaster Molecular Ecology 30 23 6101 6120 doi 10 1111 mec 15784 hdl 10045 114701 PMID 33372365 S2CID 229715430 a b c d e Lydekker Richard 1908 The Game Animals of Africa London R Ward limited pp 459 461 Heller E 1914 Four new subspecies of large mammals from equatorial Africa Smithsonian Miscellaneous Collections 61 No 22 Washington DC Smithsonian Institution a b c 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 Hennelly L Habib B Root Gutteridge H Palacios V Passilongo D 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 doi 10 1093 cz zox001 PMC 5804178 PMID 29491993 Foley C et al 2014 A Field Guide to the Larger Mammals of Tanzania Princeton University Press pp 102 103 ISBN 1400852803 Densities of spotted hyaena Crocuta crocuta and African wolf Canis lupaster increase with increasing anthropogenic influence Mammalian Biology 85 60 69 2017 02 17 doi 10 1016 j mambio 2017 02 004 hdl 10067 1418230151162165141 S2CID 90177487 Retrieved 2021 02 23 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 4 313 323 doi 10 1111 j 1365 2028 1989 tb01025 x Gutema T M Foraging ecology and trophic niche overlap between sympatric African wolf and Ethiopian wolf in the Ethiopian Highlands Archived 2016 10 06 at the Wayback Machine The Rufford Foundation November 2015 Sillero Zubiri C Gottelli D 1994 Canis simensis PDF Mammalian Species 485 1 6 doi 10 2307 3504136 JSTOR 3504136 S2CID 253988540 Archived from the original PDF on 2015 09 24 Retrieved 2015 08 01 Kruuk H 1972 The Spotted Hyena A Study of Predation and Social Behaviour Chicago University of Chicago Press ISBN 978 0 226 45507 5 Remler P 2010 Egyptian Mythology A to Z Infobase Publishing p 99 ISBN 978 1438131801 Wilkinson J G 1847 The Manners and Customs of the Ancient Egyptians Vol 5 John Murray Trecolle G Camps G 1993 Chacal Encyclopedie Berbere Aix en Provence Capsa Cheval 12 12 1857 1859 doi 10 4000 encyclopedieberbere 2099 Thiaw Issa laye 2009 Mythe de la creation du monde selon les sages sereer Plaidoyer pour le dialogue interreligieux Enracinement et Ouverture in French Dakar Konrad Adenauer Stiftung pp 45 50 59 61 Further reading EditCheryl 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 amp Jane Goodall 1971 Innocent Killers Houghton Mifflin Company BostonExternal links Edit Wikispecies has information related to African golden wolf Canis lupaster in Arabic Le fantome du moyen Atlas شبح الأطلس المتوسط AmouddouTV Feb 13 2014 in Arabic Nouveau Nouvelle espece decouverte اكتشاف نوع جديد من الحيوانات حصري AmouddouTV Aug 15 2015 Retrieved from https en wikipedia org w index php title African wolf amp oldid 1131380488, wikipedia, wiki, book, books, library,

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