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Old page wikitext, before the edit (old_wikitext ) | '{{Taxobox
| name = African golden wolf
| fossil_range = {{Fossil range|1-1.7|0}}<small>[[Early Pleistocene]] – Recent</small>
| status =
| image = Golden jackal small.jpg
| image caption = ''Canis anthus bea'' in [[Ngorongoro Crater]], Tanzania
| regnum = [[Animal]]ia
| phylum = [[Chordate|Chordata]]
| classis = [[Mammal]]ia
| ordo = [[Carnivora]]
| familia = [[Canidae]]
| genus = ''[[Canis]]''
| species ='''''C. anthus'''''
| binomial = ''Canis anthus''<ref name="cuvier"/>
| binomial_authority = [[Frédéric Cuvier|F. Cuvier]], 1820
| subdivision_ranks = [[Subspecies]]
| subdivision =
*''[[Senegalese wolf|C. a. anthus]]''
*''C. a. algirensis''
*''C. a. bea''
*''C. a. riparius''
*''[[Variegated wolf|C. a. soudanicus]]''
}}
The '''African golden wolf''' (''Canis anthus''), also known as the '''African wolf''' or '''thoa''', is a [[canid]] native to [[North Africa|north]] and [[Horn of Africa|northeastern Africa]]. The species is common in [[North Africa|North]] [[West Africa|west]] and north-[[east Africa]], occurring from [[Senegal]] to [[Egypt]] in the east, in a range including [[Morocco]], [[Algeria]], and [[Libya]] in the north to [[Nigeria]], [[Chad]] and [[Tanzania]] in the south.<ref name=iucn>{{IUCN2008|assessors=Jhala, Y.V. & Moehlman, P.D.|year=2008|id=3744|title=Canis aureus|downloaded=22 March 2009}}</ref> It is a desert-adapted canid, and is common in plains and steppe areas, including ones lacking abundant water. It is primarily a predator, targeting [[invertebrate]]s and [[mammal]]s 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.<ref name="e398"/>
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|mtDNA]] and [[Nuclear DNA|nuclear]] [[genome]] demonstrated that it was in fact distinct from the golden jackal, and more closely related to grey wolves and [[coyote]]s. It is nonetheless still close enough to the golden jackal to produce hybrid offspring, as indicated through genetic tests on jackals in Israel<ref name="koepfli"/> and a 19th century captive crossbreeding experiment.<ref name="cuvier3"/> As the [[IUCN]]'s golden jackal page has not been updated since 2008, it has yet to recognise the distinctiveness of the African wolf and give it its own [[conservation status]], with population estimates being completely lacking.<ref>[http://www.theguardian.com/science/grrlscientist/2015/jul/30/golden-jackal-a-new-wolf-species-hiding-in-plain-sight Golden jackal: A new wolf species hiding in plain sight], ''The Guardian'', (30 July, 2015)</ref>
==Physical description==
[[File:Journal.pone.0042740.g006.png|left|thumb|Various ''C. anthus'' [[phenotype]]s, ranging from gracile jackal-like morphs to more robust wolf-like ones.]]
The African wolf is a small canid, with both sexes weighing between 7–15 kg (15.6–33 lbs), and standing 40 cm in height. 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 4–8 teats.<ref name="e398">Estes, Richard (1992). "The behavior guide to African mammals: including hoofed mammals, carnivores, primates". University of California Press. pp. 398–399. ISBN 0-520-08085-8.</ref> 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.<ref name="koepfli"/> The ears are longer in the golden wolf, and the skull has a more elevated forehead.<ref name="mivart41">Mivart, George (1890), ''[https://archive.org/stream/dogsjackalswolve00mivauoft#page/n115/mode/2up Dogs, Jackals, Wolves and Foxes: A Monograph of the Canidæ]'', R.H. Porter, London, pp. 41–43</ref>
{{clear}}
==Taxonomic history==
[[File: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.png|thumb|left|Skull of African wolf (left) and golden jackal (right).]]
[[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 Egypt]]ian city dedicated to [[Anubis]], means "city of the wolf".<ref name="Rueness2011" /><ref name="ferg">{{Cite journal|author=Ferguson, W.W.|year= 1981|title= The systematic of ''Canis aureus lupaster'' (Carnivora : Canidae) and the occurrence of ''Canis lupus'' in North Africa, Egypt and Sinai|journal= Mammalia |volume=4|pages= 459–465}}</ref>
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 [[Arcadia]]n [[Anthus (mythology)|Anthus]] family described by [[Pliny the Elder]] in his ''[[Natural History (Pliny)|Natural History]]'', whose members would draw lots to become [[Werewolf|werewolves]].<ref name="cuvier">{{fr icon}} Cuvier, Frédéric (1824), ''[https://archive.org/stream/HistoirenaturelIIGeof#page/n33/mode/2up Histoire naturelle des mammifères]'', tome 2, A Paris : Chez A. Belin ...</ref> [[Eduard Rüppell]] proposed that the animal was the ancestor of Egyptian [[sighthound]]s, and named it ''Wolf's-hund'' (wolf dog),<ref name="ruppell1826">{{de icon}} Rüppell, Eduard (1826), ''[https://archive.org/stream/atlaszuderreisei00rupp#page/n95/mode/2up Atlas zu der Reise im nördlichen Afrika]'', Frankfurt am Main : Gedruckt und in Commission bei Heinr. Ludw. Brönner, pp. 44–46</ref> while [[Charles Hamilton Smith]] named it "thoa" or "thous dog".<ref name="smith1839">Smith, Charles Hamilton; Jardine, Sir William (1839). ''[https://archive.org/stream/naturalhistoryof139smit#page/192/mode/2up The natural history of dogs : canidae or genus canis of authors ; including also the genera hyaena and proteles]'', Volume I. Edinburgh : W. H. Lizars. pp. 193–194</ref> 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.<ref name="cuvier3">{{fr icon}} Cuvier, Frédéric (1824), ''[https://archive.org/stream/HistoirenaturelIIIGeof#page/n243/mode/2up]'', tome 3, A Paris : Chez A. Belin ...</ref> English biologist [[St. George Jackson Mivart]] emphasised the differences between the African wolf and the golden jackal in his writings:
[[File:Histoire naturelle des mammifères, t. 3 (1824) Canis anthus x aureus.png|thumb|Illustration of golden jackal-African wolf hybrids bred in captivity (1821).]]
{{quote|... 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)<ref name="mivart36">Mivart, George (1890), [https://archive.org/stream/dogsjackalswolve00mivauoft#page/36/mode/2up ''Dogs, Jackals, Wolves and Foxes: A Monograph of the Canidæ''], R.H. Porter, London, pp. 36–37</ref>}}
[[File:Jackalvarieties.jpg|thumb|Comparative illustration of ''[[Canis anthus|C. aureus]]'' (top) and ''C. anthus'' (bottom).]]
Although subsequently [[Synonym (taxonomy)|synonymised]] with the golden jackal, new doubts over its being conspecific with the golden jackal arose in 2011, when several golden "jackal" populations in Egypt and the Horn of Africa classed as ''Canis aureus lupaster'' were found to have [[mtDNA]] sequences more closely resembling those in [[Gray wolf|grey wolves]] than those of golden jackals.<ref name="Rueness2011">{{cite doi|10.1371/journal.pone.0016385}}</ref> These wolf-like mtDNA sequences were confirmed to occur over a range 6,000 km wide, encompassing Algeria, Mali and Senegal. In both these studies, it was proposed to reclassify ''Canis aureus lupaster'' as a subspecies of grey wolf.<ref name="gaubert">{{Cite journal|author=Gaubert P, Bloch C, Benyacoub S, Abdelhamid A, Pagani P, et al|year=2012|title= Reviving the African Wolf ''Canis lupus lupaster'' in North and West Africa: A Mitochondrial Lineage Ranging More than 6,000 km Wide|journal=PLoS ONE|volume=7|issue=8|page=e42740|doi=10.1371/journal.pone.0042740|pmid=22900047|pmc=3416759}}</ref>
In 2015 however, a more thorough comparative study of mitochondrial and [[Nuclear DNA|nuclear]] [[genome]]s on both Eurasian golden jackals and their African counterparts revealed that the latter were in fact a separate species more closely related to grey wolves and coyotes, with a [[genetic divergence]] of around 6.7%,<ref name="koepfli">{{cite journal|last1=Koepfli|first1=K.-P.|last2= Pollinger|first2=J.|last3= Godinho|first3=R.|last4= Robinson|first4=J.|last5= Lea|first5=A.|last6= Hendricks|first6=S.|last7= Schweizer|first7=R. M.|last8= Thalmann|first8=O.|last9= Silva|first9=P.|last10= Fan|first10=Z.|last11= Yurchenko|first11=A. A.|last12= Dobrynin|first12=P.|last13= Makunin|first13=A.|last14= Cahill|first14=J. A.|last15= Shapiro|first15=B.|last16= Álvares|first16=F.|last17= Brito|first17=J. C.|last18= Geffen|first18=E.|last19= Leonard|first19=J. A.|last20= Helgen|first20=K. M.|last21= Johnson|first21=W. E.|last22= O’Brien|first22=S. J.|last23= Van Valkenburgh|first23=B.|last24= Wayne|first24=R. K.|title=Genome-wide Evidence Reveals that African and Eurasian Golden Jackals Are Distinct Species|journal=Current Biology|volume=25|issue= | pages= |url= http://www.cell.com/current-biology/abstract/S0960-9822%2815%2900787-3|date= 2015-08-17|doi= 10.1016/j.cub.2015.06.060|pmid=|pmc=}}</ref> which is greater than that between grey wolves and [[coyote]]s (4%) and that between grey wolves and [[domestic dog]]s (0.2%).<ref>{{cite journal | last1 = Wayne | first1 = Robert K. | year = 1993 | title = Molecular evolution of the dog family | journal = Trends in Genetics | volume = 9 | issue = 6| pages = 218–224 | doi = 10.1016/0168-9525(93)90122-X | pmid=8337763}}</ref> The [[phylogenetic tree]] below is based on nuclear sequences:<ref name="koepfli"/>
{{clade | style = font-size: 90%;line-height:50%
|1={{clade
|1={{clade
|1={{clade
|1=[[Side-striped jackal]]
|2=[[Black-backed jackal]]
}}
|2={{clade
|1={{clade
|1={{clade
|1={{clade
|1={{clade
|1='''African golden wolf''' (formerly African golden jackal), ''Canis anthus''<ref name="koepfli"/>{{rp|at=Fig. 2}}
|2={{clade
|1={{clade
|1=[[Dog]]
|2=[[Grey wolf]]
}}
|2=[[Coyote]]
}}
}}
|2=[[Ethiopian wolf]]
}}
|2='''[[golden jackal|Eurasian golden jackal]]''', ''Canis aureus''<ref name="koepfli"/>{{rp|at=Fig. 2}}
}}
|2=[[Dhole]]
}}
|2={{clade
|1=[[African wild dog]]
}}
}}
}}
}}
}}
It was estimated that the African golden wolf diverged from the wolf-coyote clade 1-1.7 million years ago during the [[Pleistocene]], and that its superficial similarity to the Eurasian golden jackal (particularly in East Africa, where African wolves are similar in size to golden jackals) is a case of [[parallel evolution]]. Considering its phylogenetic position and the canid [[fossil record]], it is likely that the African wolf evolved from larger-sized ancestors which became progressively more jackal-like in size upon populating Africa on account of [[interspecific competition]] with both larger and smaller-sized indigenous [[Carnivora|carnivore]]s. Because of Egypt's [[contiguity]] with [[Israel]], traces of golden wolf DNA were identified in Israeli golden jackals, thus indicating the presence of a hybrid zone.<ref name="koepfli"/>
===Subspecies===
As of 2015, [[MSW3]] still classifies the six African wolf subspecies as part of ''Canis aureus''.<ref name=MSW3>{{MSW3 Wozencraft | id=14000707}}</ref>
Although in the past several attempts have been made to [[Synonym (taxonomy)|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.<ref name="r38">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.</ref>
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.<ref name="r38"/>
{| class="wikitable collapsed"
!Subspecies
!Trinomial authority
!Description
!Range
!Synonyms
|-
|'''Algerian wolf'''<br/>''Canis a. algirensis''
[[File:Canisaureusalgirensis.jpg|150 px]]
|[[Johann Andreas Wagner|Wagner]], 1841
|A dark-coloured subspecies, with a tail marked with three dusky rings. It is similar in size to the [[red fox]].<ref name="l218">Lydekker, Richard (1908). "The Game Animals of Africa". London, R. Ward, limited. p. 218</ref>
|[[Algeria]], [[Morocco]] and [[Tunisia]]
|<small>''barbarus'' (C. E. H. Smith, 1839)</small><br/>
<small>''grayi'' (Hilzheimer, 1906)</small><br/>
<small>''tripolitanus'' (Wagner, 1841)</small>
|-
|'''Senegalese wolf'''<br/>''[[Senegalese Jackal|Canis a. anthus]]''
[[File:Senegalesegoldenjackal.png|150 px]]
|[[Frédéric Cuvier|F. Cuvier]], 1820
|
|[[Senegal]]
|<small>''senegalensis'' (C. E. H. Smith, 1839)</small>
|-
|'''Serengeti wolf'''<br/>''Canis a. bea''
[[File:Golden Jackal, Serengeti.jpg|150 px]]
|Heller, 1914
|
|[[Kenya]], Northern [[Tanzania]]
|
|-
|'''Egyptian wolf'''<br/>''[[Egyptian wolf|Canis a. lupaster]]''
[[File:Lupaster.png|150 px]]
|[[Wilhelm Friedrich Hemprich|Hemprich]] and [[Christian Gottfried Ehrenberg|Ehrenberg]], 1833
| A large subspecies presenting a very grey wolf-like [[phenotype]].<ref name="gaubert"/>
|[[Egypt]], [[Algeria]], [[Mali]], [[Ethiopian Highlands]], and [[Senegal]]
|''C. aureus lupaster''<br>
''C. lupaster''<br>
''C. lupus lupaster''<br>
''C. sacer'' (Hemprich and Ehrenberg, 1833)
|-
|'''Somali wolf'''<br/>''Canis a. riparius''
|[[Wilhelm Friedrich Hemprich|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.<ref name="l460">Lydekker, Richard (1908). "The Game Animals of Africa". London, R. Ward, limited. p. 460.</ref>
|Somalia and coast of [[Ethiopia]] and [[Eritrea]]
|<small>''hagenbecki'' (Noack, 1897)</small><br/>
<small>''mengesi'' (Noack, 1897)</small><br/>
<small>''somalicus'' (Lorenz, 1906)</small>
|-
|'''Variegated wolf'''<br/>''[[Variegated jackal|Canis a. soudanicus]]''
[[File:Paintedthousdog.jpg|150 px]]
|[[Oldfield Thomas|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.<ref name=l460/>
|[[Sudan]] and [[Somalia]]
|<small>''doederleini'' (Hilzheimer, 1906)</small><br/>
<small>''nubianus'' (Cabrera, 1921)</small><br/>
<small>''thooides'' (Hilzheimer, 1906)</small><br/>
<small>''variegatus'' (Cretzschmar, 1826)</small>
|}
==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".<ref name="jhala2004"/> Large groups are rare, and have only been recorded to occur in areas with abundant human waste. Family relationships among golden 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. Golden 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.<ref name="e399">Estes, Richard (1992). "The behavior guide to African mammals: including hoofed mammals, carnivores, primates". University of California Press. pp. 399–400. ISBN 0-520-08085-8.</ref> The pair patrols and [[territorial marking|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.<ref name="e402">Estes, Richard (1992). "The behavior guide to African mammals: including hoofed mammals, carnivores, primates". University of California Press. p. 402. ISBN 0-520-08085-8.</ref>
[[File:Canis anthus threat postures.png|thumb|Threat posture of ''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 mark]]s its territory. [[Animal sexual behavior|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.<ref name="e402–403">Estes, Richard (1992). "The behavior guide to African mammals: including hoofed mammals, carnivores, primates". University of California Press. pp. 402–403. ISBN 0-520-08085-8.</ref>
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.<ref name="r47"/>
[[File:Golden jackal eating agama.jpg|thumb|Serengeti wolf (''C. a. bea'') eating an [[agama (lizard)|agama]]]]
===Communication===
African wolves frequently groom one another, particularly during courtship, during which it can last up to ½ hour. Nibbling of the face and neck is observed during greeting ceremonies. When fighting, the African wolf slams its opponents with its hips, and bites and shakes the shoulder. The species' postures are typically canine, and it has more facial mobility than the black-backed and side-striped jackals, being able to expose its canine teeth like a dog.<ref name="e401–402">Estes, Richard (1992). "The behavior guide to African mammals: including hoofed mammals, carnivores, primates". University of California Press. pp. 401–402. ISBN 0-520-08085-8.</ref>
The vocabulary of the African wolf is similar to that of the domestic dog, with seven sounds having been recorded.<ref name="k22">Kingdon, Jonathan (1988). "East African mammals: an atlas of evolution in Africa, Volume 3, Part 1". University of Chicago Press. pp. 22. ISBN 0-226-43721-3.</ref> The African wolf's vocalisations include howls, barks, growls, whines and cackles.<ref name="e401–402"/> Subspecies can be recognised by differences in their howls.<ref name="k22"/> 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.<ref name="e401–402"/>
===Hunting behaviour===
The African wolf rarely catches hares due to them being faster. [[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.<ref name="e400–401">Estes, Richard (1992). "The behavior guide to African mammals: including hoofed mammals, carnivores, primates". University of California Press. pp. 400–401. ISBN 0-520-08085-8.</ref>
Although it is known to kill animals up to three times its own weight, the golden wolf targets mammalian prey much less frequently than the [[black-backed jackal]] overall.<ref name="e400–401"/> 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 [[hoarding (animal behavior)|caches]] the surplus, which is generally recovered within 24 hours.<ref name="r47"/> 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 [[Old World Vulture|vulture]]s on kills – one can hold dozens of vultures at bay by threatening, snapping and lunging at them.<ref name="e400–401"/>
==Ecology==
===Diet===
[[File:Golden Jackal, navigating Wildebeest, Ngorongoro.jpg|thumb|right|A Serengeti wolf (''C. a. bea'') carefully navigating a herd of [[blue wildebeest]] in the [[Ngorongoro National Park]], [[Tanzania]]]]
In [[West Africa]], the African wolf mostly confines itself to small prey, such as [[hare]]s, [[rat]]s, [[ground squirrel]]s and [[Cane rat|grass cutter]]s. Other prey items include lizards, snakes, and ground-nesting birds, such as [[francolin]]s and [[bustard]]s. It also consumes a large amount of insects, including [[dung beetle]]s, [[larvae]], [[termite]]s and [[grasshopper]]s. It will also kill young [[gazelle]]s, [[duiker]]s and [[warthog]]s.<ref name="r47">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.</ref> In East Africa, it consumes invertebrates and fruit, though 60% of its diet consists of rodents, lizards, snakes, birds, hares and [[Thomson's gazelle]]s.<ref name="jhala2004">Jhala, Y. V. & Moehlman, P. D. 2004. [http://www.canids.org/species/Golden_jackal.pdf Golden jackal ''Canis aureus'']</ref> During the [[wildebeest]] calving season, African wolves will feed almost exclusively on their [[afterbirth]].<ref name="k19">Kingdon, Jonathan (1988). "East African mammals: an atlas of evolution in Africa, Volume 3, Part 1". University of Chicago Press. pp. 19–21. ISBN 0-226-43721-3.</ref> In the [[Serengeti]] and Ngorongoro Crater, less than 20% of its diet comes from scavenging.<ref name="e400–401"/>
===Enemies and competitors===
The African wolf tends to dominate smaller canid species, and has been observed to kill the pups of [[black-backed jackal]]s.<ref name="jhala2004"/> It often eats alongside [[African wild dog]]s, and will stand its ground if the dogs try to harass it.<ref name="e400–401"/> There is at least one record of an African wolf pack adopting a male [[Ethiopian wolf]].<ref name="zm1994">Sillero-Zubiri, C., and D. Gottelli (1994). [http://www.science.smith.edu/msi/pdf/i0076-3519-485-01-0001.pdf Canis simensis]. ''Mammalian Species'' 385: 1–6.</ref>
Wolves will feed alongside [[spotted hyena]]s, 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.<ref name="Kruuk">{{cite book |last=Kruuk |first=Hans |year=1972 |title=The Spotted Hyena: A Study of Predation and Social Behaviour |publisher=University of Chicago Press |location=Chicago |isbn=0-226-45507-6 }}</ref> Wolves will confront a hyena approaching too closely to their dens by taking turns in biting the hyena's hocks until it retreats.<ref name="e400–401"/>
Wolves in the Serengeti are known to carry the [[canine parvovirus]], [[canine herpesvirus]], [[canine coronavirus]] and [[canine adenovirus]].<ref name=jhala2004/>
==In folklore==
In Egyptian folklore, the wolf can cause chickens to faint from fear by simply passing underneath their roosts, and its body parts are associated with various forms of folk magic; placing a wolf's tongue into a house is believed to cause the inhabitants within to argue, and its meat 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]].<ref>Obsorn, Dale. J.; Helmy, Ibrahim (1980). [https://archive.org/stream/contemporaryland05osbo#page/370/mode/2up "The contemporary land mammals of Egypt (including Sinai)"]. Field Museum of Natural History, p. 371</ref>
==See also==
{{commons|Category:Canis anthus|African golden wolf (Canis anthus)}}
*[[Ethiopian wolf]]
{{clear}}
==References==
{{reflist|30em}}
==Further reading==
*[[Hugo van Lawick]] & [[Jane Goodall]] (1971), ''Innocent Killers'', Houghton Mifflin Company Boston
{{Carnivora|Ca.}}
[[Category:Wolves]]
[[Category:Mammals of Africa]]
[[Category:Canis]]' |
New page wikitext, after the edit (new_wikitext ) | '{{Taxobox
| name = African golden wolf
| fossil_range = {{Fossil range|1-1.7|0}}<small>[[Early Pleistocene]] – Recent</small>
| status =
| image = Golden jackal small.jpg
| image caption = ''Canis anthus bea'' in [[Ngorongoro Crater]], Tanzania
| regnum = [[Animal]]ia
| phylum = [[Chordate|Chordata]]
| classis = [[Mammal]]ia
| ordo = [[Carnivora]]
| familia = [[Canidae]]
| genus = ''[[Canis]]''
| species ='''''C. anthus'''''
| binomial = ''Canis anthus''<ref name="cuvier"/>
| binomial_authority = [[Frédéric Cuvier|F. Cuvier]], 1820
| subdivision_ranks = [[Subspecies]]
| subdivision =
*''[[Senegalese wolf|C. a. anthus]]''
*''C. a. algirensis''
*''C. a. bea''
*''C. a. riparius''
*''[[Variegated wolf|C. a. soudanicus]]''
}}
The '''African golden wolf''' (''Canis anthus''), also known as the '''African wolf''' or '''thoa''', is a [[canid]] native to [[North Africa|north]] and [[Horn of Africa|northeastern Africa]]. The species is common in [[North Africa|North]] [[West Africa|west]] and north-[[east Africa]], occurring from [[Senegal]] to [[Egypt]] in the east, in a range including [[Morocco]], [[Algeria]], and [[Libya]] in the north to [[Nigeria]], [[Chad]] and [[Tanzania]] in the south.<ref name=iucn>{{IUCN2008|assessors=Jhala, Y.V. & Moehlman, P.D.|year=2008|id=3744|title=Canis aureus|downloaded=22 March 2009}}</ref> It is a desert-adapted canid, and is common in plains and steppe areas, including ones lacking abundant water. It is primarily a predator, targeting [[invertebrate]]s and [[mammal]]s 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.<ref name="e398"/>
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|mtDNA]] and [[Nuclear DNA|nuclear]] [[genome]] demonstrated that it was in fact distinct from the golden jackal, and more closely related to grey wolves and [[coyote]]s. It is nonetheless still close enough to the golden jackal to produce hybrid offspring, as indicated through genetic tests on jackals in Israel<ref name="koepfli"/> and a 19th century captive crossbreeding experiment.<ref name="cuvier3"/> As the [[IUCN]]'s golden jackal page has not been updated since 2008, it has yet to recognise the distinctiveness of the African wolf and give it its own [[conservation status]], with population estimates being completely lacking.<ref>[http://www.theguardian.com/science/grrlscientist/2015/jul/30/golden-jackal-a-new-wolf-species-hiding-in-plain-sight Golden jackal: A new wolf species hiding in plain sight], ''The Guardian'', (30 July, 2015)</ref>
==Physical description==
[[File:Journal.pone.0042740.g006.png|left|thumb|Various ''C. anthus'' [[phenotype]]s, ranging from gracile jackal-like morphs to more robust wolf-like ones.]]
The African wolf is a small canid, with both sexes weighing between 7–15 kg (15.6–33 lbs), and standing 40 cm in height. 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 4–8 teats.<ref name="e398">Estes, Richard (1992). "The behavior guide to African mammals: including hoofed mammals, carnivores, primates". University of California Press. pp. 398–399. ISBN 0-520-08085-8.</ref> 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.<ref name="koepfli"/> The ears are longer in the golden wolf, and the skull has a more elevated forehead.<ref name="mivart41">Mivart, George (1890), ''[https://archive.org/stream/dogsjackalswolve00mivauoft#page/n115/mode/2up Dogs, Jackals, Wolves and Foxes: A Monograph of the Canidæ]'', R.H. Porter, London, pp. 41–43</ref>
{{clear}}
==Taxonomic history==
[[File: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.png|thumb|left|Skull of African wolf (left) and golden jackal (right).]]
[[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 Egypt]]ian city dedicated to [[Anubis]], means "city of the wolf".<ref name="Rueness2011" /><ref name="ferg">{{Cite journal|author=Ferguson, W.W.|year= 1981|title= The systematic of ''Canis aureus lupaster'' (Carnivora : Canidae) and the occurrence of ''Canis lupus'' in North Africa, Egypt and Sinai|journal= Mammalia |volume=4|pages= 459–465}}</ref>
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 [[Arcadia]]n [[Anthus (mythology)|Anthus]] family described by [[Pliny the Elder]] in his ''[[Natural History (Pliny)|Natural History]]'', whose members would draw lots to become [[Werewolf|werewolves]].<ref name="cuvier">{{fr icon}} Cuvier, Frédéric (1824), ''[https://archive.org/stream/HistoirenaturelIIGeof#page/n33/mode/2up Histoire naturelle des mammifères]'', tome 2, A Paris : Chez A. Belin ...</ref> [[Eduard Rüppell]] proposed that the animal was the ancestor of Egyptian [[sighthound]]s, and named it ''Wolf's-hund'' (wolf dog),<ref name="ruppell1826">{{de icon}} Rüppell, Eduard (1826), ''[https://archive.org/stream/atlaszuderreisei00rupp#page/n95/mode/2up Atlas zu der Reise im nördlichen Afrika]'', Frankfurt am Main : Gedruckt und in Commission bei Heinr. Ludw. Brönner, pp. 44–46</ref> while [[Charles Hamilton Smith]] named it "thoa" or "thous dog".<ref name="smith1839">Smith, Charles Hamilton; Jardine, Sir William (1839). ''[https://archive.org/stream/naturalhistoryof139smit#page/192/mode/2up The natural history of dogs : canidae or genus canis of authors ; including also the genera hyaena and proteles]'', Volume I. Edinburgh : W. H. Lizars. pp. 193–194</ref> 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.<ref name="cuvier3">{{fr icon}} Cuvier, Frédéric (1824), ''[https://archive.org/stream/HistoirenaturelIIIGeof#page/n243/mode/2up]'', tome 3, A Paris : Chez A. Belin ...</ref> English biologist [[St. George Jackson Mivart]] emphasised the differences between the African wolf and the golden jackal in his writings:
[[File:Histoire naturelle des mammifères, t. 3 (1824) Canis anthus x aureus.png|thumb|Illustration of golden jackal-African wolf hybrids bred in captivity (1821).]]
{{quote|... 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)<ref name="mivart36">Mivart, George (1890), [https://archive.org/stream/dogsjackalswolve00mivauoft#page/36/mode/2up ''Dogs, Jackals, Wolves and Foxes: A Monograph of the Canidæ''], R.H. Porter, London, pp. 36–37</ref>}}
[[File:Jackalvarieties.jpg|thumb|Comparative illustration of ''[[Canis anthus|C. aureus]]'' (top) and ''C. anthus'' (bottom).]]
Although subsequently [[Synonym (taxonomy)|synonymised]] with the golden jackal, new doubts over its being conspecific with the golden jackal arose in 2011, when several golden "jackal" populations in Egypt and the Horn of Africa classed as ''Canis aureus lupaster'' were found to have [[mtDNA]] sequences more closely resembling those in [[Gray wolf|grey wolves]] than those of golden jackals.<ref name="Rueness2011">{{cite doi|10.1371/journal.pone.0016385}}</ref> These wolf-like mtDNA sequences were confirmed to occur over a range 6,000 km wide, encompassing Algeria, Mali and Senegal. In both these studies, it was proposed to reclassify ''Canis aureus lupaster'' as a subspecies of grey wolf.<ref name="gaubert">{{Cite journal|author=Gaubert P, Bloch C, Benyacoub S, Abdelhamid A, Pagani P, et al|year=2012|title= Reviving the African Wolf ''Canis lupus lupaster'' in North and West Africa: A Mitochondrial Lineage Ranging More than 6,000 km Wide|journal=PLoS ONE|volume=7|issue=8|page=e42740|doi=10.1371/journal.pone.0042740|pmid=22900047|pmc=3416759}}</ref>
In 2015 however, a more thorough comparative study of mitochondrial and [[Nuclear DNA|nuclear]] [[genome]]s on both Eurasian golden jackals and their African counterparts revealed that the latter were in fact a separate species more closely related to grey wolves and coyotes, with a [[genetic divergence]] of around 6.7%,<ref name="koepfli">{{cite journal|last1=Koepfli|first1=K.-P.|last2= Pollinger|first2=J.|last3= Godinho|first3=R.|last4= Robinson|first4=J.|last5= Lea|first5=A.|last6= Hendricks|first6=S.|last7= Schweizer|first7=R. M.|last8= Thalmann|first8=O.|last9= Silva|first9=P.|last10= Fan|first10=Z.|last11= Yurchenko|first11=A. A.|last12= Dobrynin|first12=P.|last13= Makunin|first13=A.|last14= Cahill|first14=J. A.|last15= Shapiro|first15=B.|last16= Álvares|first16=F.|last17= Brito|first17=J. C.|last18= Geffen|first18=E.|last19= Leonard|first19=J. A.|last20= Helgen|first20=K. M.|last21= Johnson|first21=W. E.|last22= O’Brien|first22=S. J.|last23= Van Valkenburgh|first23=B.|last24= Wayne|first24=R. K.|title=Genome-wide Evidence Reveals that African and Eurasian Golden Jackals Are Distinct Species|journal=Current Biology|volume=25|issue= | pages= |url= http://www.cell.com/current-biology/abstract/S0960-9822%2815%2900787-3|date= 2015-08-17|doi= 10.1016/j.cub.2015.06.060|pmid=|pmc=}}</ref> which is greater than that between grey wolves and [[coyote]]s (4%) and that between grey wolves and [[domestic dog]]s (0.2%).<ref>{{cite journal | last1 = Wayne | first1 = Robert K. | year = 1993 | title = Molecular evolution of the dog family | journal = Trends in Genetics | volume = 9 | issue = 6| pages = 218–224 | doi = 10.1016/0168-9525(93)90122-X | pmid=8337763}}</ref> The [[phylogenetic tree]] below is based on nuclear sequences:<ref name="koepfli"/>
{{clade | style = font-size: 90%;line-height:50%
|1={{clade
|1={{clade
|1={{clade
|1=[[Side-striped jackal]]
|2=[[Black-backed jackal]]
}}
|2={{clade
|1={{clade
|1={{clade
|1={{clade
|1={{clade
|1='''African golden wolf''' (formerly African golden jackal), ''Canis anthus''<ref name="koepfli"/>{{rp|at=Fig. 2}}
|2={{clade
|1={{clade
|1=[[Dog]]
|2=[[Grey wolf]]
}}
|2=[[Coyote]]
}}
}}
|2=[[Ethiopian wolf]]
}}
|2='''[[golden jackal|Eurasian golden jackal]]''', ''Canis aureus''<ref name="koepfli"/>{{rp|at=Fig. 2}}
}}
|2=[[Dhole]]
}}
|2={{clade
|1=[[African wild dog]]
}}
}}
}}
}}
}}
It was estimated that the African golden wolf diverged from the wolf-coyote clade 1-1.7 million years ago during the [[Pleistocene]], and that its superficial similarity to the Eurasian golden jackal (particularly in East Africa, where African wolves are similar in size to golden jackals) is a case of [[parallel evolution]]. Considering its phylogenetic position and the canid [[fossil record]], it is likely that the African wolf evolved from larger-sized ancestors which became progressively more jackal-like in size upon populating Africa on account of [[interspecific competition]] with both larger and smaller-sized indigenous [[Carnivora|carnivore]]s. Because of Egypt's [[contiguity]] with [[Israel]], traces of golden wolf DNA were identified in Israeli golden jackals, thus indicating the presence of a hybrid zone.<ref name="koepfli"/>
===Subspecies===
As of 2015, [[MSW3]] still classifies the six African wolf subspecies as part of ''Canis aureus''.<ref name=MSW3>{{MSW3 Wozencraft | id=14000707}}</ref>
Although in the past several attempts have been made to [[Synonym (taxonomy)|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.<ref name="r38">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.</ref>
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.<ref name="r38"/>
{| class="wikitable collapsed"
!Subspecies
!Trinomial authority
!Description
!Range
!Synonyms
|-
|'''Algerian wolf'''<br/>''Canis a. algirensis''
[[File:Canisaureusalgirensis.jpg|150 px]]
|[[Johann Andreas Wagner|Wagner]], 1841
|A dark-coloured subspecies, with a tail marked with three dusky rings. It is similar in size to the [[red fox]].<ref name="l218">Lydekker, Richard (1908). "The Game Animals of Africa". London, R. Ward, limited. p. 218</ref>
|[[Algeria]], [[Morocco]] and [[Tunisia]]
|<small>''barbarus'' (C. E. H. Smith, 1839)</small><br/>
<small>''grayi'' (Hilzheimer, 1906)</small><br/>
<small>''tripolitanus'' (Wagner, 1841)</small>
|-
|'''Senegalese wolf'''<br/>''[[Senegalese Jackal|Canis a. anthus]]''
[[File:Senegalesegoldenjackal.png|150 px]]
|[[Frédéric Cuvier|F. Cuvier]], 1820
|
|[[Senegal]]
|<small>''senegalensis'' (C. E. H. Smith, 1839)</small>
|-
|'''Serengeti wolf'''<br/>''Canis a. bea''
[[File:Golden Jackal, Serengeti.jpg|150 px]]
|Heller, 1914
|
|[[Kenya]], Northern [[Tanzania]]
|
|'''Somali wolf'''<br/>''Canis a. riparius''
|[[Wilhelm Friedrich Hemprich|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.<ref name="l460">Lydekker, Richard (1908). "The Game Animals of Africa". London, R. Ward, limited. p. 460.</ref>
|Somalia and coast of [[Ethiopia]] and [[Eritrea]]
|<small>''hagenbecki'' (Noack, 1897)</small><br/>
<small>''mengesi'' (Noack, 1897)</small><br/>
<small>''somalicus'' (Lorenz, 1906)</small>
|-
|'''Variegated wolf'''<br/>''[[Variegated jackal|Canis a. soudanicus]]''
[[File:Paintedthousdog.jpg|150 px]]
|[[Oldfield Thomas|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.<ref name=l460/>
|[[Sudan]] and [[Somalia]]
|<small>''doederleini'' (Hilzheimer, 1906)</small><br/>
<small>''nubianus'' (Cabrera, 1921)</small><br/>
<small>''thooides'' (Hilzheimer, 1906)</small><br/>
<small>''variegatus'' (Cretzschmar, 1826)</small>
|}
==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".<ref name="jhala2004"/> Large groups are rare, and have only been recorded to occur in areas with abundant human waste. Family relationships among golden 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. Golden 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.<ref name="e399">Estes, Richard (1992). "The behavior guide to African mammals: including hoofed mammals, carnivores, primates". University of California Press. pp. 399–400. ISBN 0-520-08085-8.</ref> The pair patrols and [[territorial marking|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.<ref name="e402">Estes, Richard (1992). "The behavior guide to African mammals: including hoofed mammals, carnivores, primates". University of California Press. p. 402. ISBN 0-520-08085-8.</ref>
[[File:Canis anthus threat postures.png|thumb|Threat posture of ''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 mark]]s its territory. [[Animal sexual behavior|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.<ref name="e402–403">Estes, Richard (1992). "The behavior guide to African mammals: including hoofed mammals, carnivores, primates". University of California Press. pp. 402–403. ISBN 0-520-08085-8.</ref>
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.<ref name="r47"/>
[[File:Golden jackal eating agama.jpg|thumb|Serengeti wolf (''C. a. bea'') eating an [[agama (lizard)|agama]]]]
===Communication===
African wolves frequently groom one another, particularly during courtship, during which it can last up to ½ hour. Nibbling of the face and neck is observed during greeting ceremonies. When fighting, the African wolf slams its opponents with its hips, and bites and shakes the shoulder. The species' postures are typically canine, and it has more facial mobility than the black-backed and side-striped jackals, being able to expose its canine teeth like a dog.<ref name="e401–402">Estes, Richard (1992). "The behavior guide to African mammals: including hoofed mammals, carnivores, primates". University of California Press. pp. 401–402. ISBN 0-520-08085-8.</ref>
The vocabulary of the African wolf is similar to that of the domestic dog, with seven sounds having been recorded.<ref name="k22">Kingdon, Jonathan (1988). "East African mammals: an atlas of evolution in Africa, Volume 3, Part 1". University of Chicago Press. pp. 22. ISBN 0-226-43721-3.</ref> The African wolf's vocalisations include howls, barks, growls, whines and cackles.<ref name="e401–402"/> Subspecies can be recognised by differences in their howls.<ref name="k22"/> 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.<ref name="e401–402"/>
===Hunting behaviour===
The African wolf rarely catches hares due to them being faster. [[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.<ref name="e400–401">Estes, Richard (1992). "The behavior guide to African mammals: including hoofed mammals, carnivores, primates". University of California Press. pp. 400–401. ISBN 0-520-08085-8.</ref>
Although it is known to kill animals up to three times its own weight, the golden wolf targets mammalian prey much less frequently than the [[black-backed jackal]] overall.<ref name="e400–401"/> 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 [[hoarding (animal behavior)|caches]] the surplus, which is generally recovered within 24 hours.<ref name="r47"/> 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 [[Old World Vulture|vulture]]s on kills – one can hold dozens of vultures at bay by threatening, snapping and lunging at them.<ref name="e400–401"/>
==Ecology==
===Diet===
[[File:Golden Jackal, navigating Wildebeest, Ngorongoro.jpg|thumb|right|A Serengeti wolf (''C. a. bea'') carefully navigating a herd of [[blue wildebeest]] in the [[Ngorongoro National Park]], [[Tanzania]]]]
In [[West Africa]], the African wolf mostly confines itself to small prey, such as [[hare]]s, [[rat]]s, [[ground squirrel]]s and [[Cane rat|grass cutter]]s. Other prey items include lizards, snakes, and ground-nesting birds, such as [[francolin]]s and [[bustard]]s. It also consumes a large amount of insects, including [[dung beetle]]s, [[larvae]], [[termite]]s and [[grasshopper]]s. It will also kill young [[gazelle]]s, [[duiker]]s and [[warthog]]s.<ref name="r47">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.</ref> In East Africa, it consumes invertebrates and fruit, though 60% of its diet consists of rodents, lizards, snakes, birds, hares and [[Thomson's gazelle]]s.<ref name="jhala2004">Jhala, Y. V. & Moehlman, P. D. 2004. [http://www.canids.org/species/Golden_jackal.pdf Golden jackal ''Canis aureus'']</ref> During the [[wildebeest]] calving season, African wolves will feed almost exclusively on their [[afterbirth]].<ref name="k19">Kingdon, Jonathan (1988). "East African mammals: an atlas of evolution in Africa, Volume 3, Part 1". University of Chicago Press. pp. 19–21. ISBN 0-226-43721-3.</ref> In the [[Serengeti]] and Ngorongoro Crater, less than 20% of its diet comes from scavenging.<ref name="e400–401"/>
===Enemies and competitors===
The African wolf tends to dominate smaller canid species, and has been observed to kill the pups of [[black-backed jackal]]s.<ref name="jhala2004"/> It often eats alongside [[African wild dog]]s, and will stand its ground if the dogs try to harass it.<ref name="e400–401"/> There is at least one record of an African wolf pack adopting a male [[Ethiopian wolf]].<ref name="zm1994">Sillero-Zubiri, C., and D. Gottelli (1994). [http://www.science.smith.edu/msi/pdf/i0076-3519-485-01-0001.pdf Canis simensis]. ''Mammalian Species'' 385: 1–6.</ref>
Wolves will feed alongside [[spotted hyena]]s, 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.<ref name="Kruuk">{{cite book |last=Kruuk |first=Hans |year=1972 |title=The Spotted Hyena: A Study of Predation and Social Behaviour |publisher=University of Chicago Press |location=Chicago |isbn=0-226-45507-6 }}</ref> Wolves will confront a hyena approaching too closely to their dens by taking turns in biting the hyena's hocks until it retreats.<ref name="e400–401"/>
Wolves in the Serengeti are known to carry the [[canine parvovirus]], [[canine herpesvirus]], [[canine coronavirus]] and [[canine adenovirus]].<ref name=jhala2004/>
==In folklore==
In Egyptian folklore, the wolf can cause chickens to faint from fear by simply passing underneath their roosts, and its body parts are associated with various forms of folk magic; placing a wolf's tongue into a house is believed to cause the inhabitants within to argue, and its meat 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]].<ref>Obsorn, Dale. J.; Helmy, Ibrahim (1980). [https://archive.org/stream/contemporaryland05osbo#page/370/mode/2up "The contemporary land mammals of Egypt (including Sinai)"]. Field Museum of Natural History, p. 371</ref>
==See also==
{{commons|Category:Canis anthus|African golden wolf (Canis anthus)}}
*[[Ethiopian wolf]]
{{clear}}
==References==
{{reflist|30em}}
==Further reading==
*[[Hugo van Lawick]] & [[Jane Goodall]] (1971), ''Innocent Killers'', Houghton Mifflin Company Boston
{{Carnivora|Ca.}}
[[Category:Wolves]]
[[Category:Mammals of Africa]]
[[Category:Canis]]' |
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-|'''Egyptian wolf'''<br/>''[[Egyptian wolf|Canis a. lupaster]]''
-[[File:Lupaster.png|150 px]]
-|[[Wilhelm Friedrich Hemprich|Hemprich]] and [[Christian Gottfried Ehrenberg|Ehrenberg]], 1833
-| A large subspecies presenting a very grey wolf-like [[phenotype]].<ref name="gaubert"/>
-|[[Egypt]], [[Algeria]], [[Mali]], [[Ethiopian Highlands]], and [[Senegal]]
-|''C. aureus lupaster''<br>
-''C. lupaster''<br>
-''C. lupus lupaster''<br>
-''C. sacer'' (Hemprich and Ehrenberg, 1833)
-|-
|'''Somali wolf'''<br/>''Canis a. riparius''
|[[Wilhelm Friedrich Hemprich|Hemprich and Ehrenberg]], 1832
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