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===Pleistocene epoch=== During the [[Pleistocene]], the North American wolf line appeared, with ''[[Canis edwardii]]'', clearly identifiable as a wolf, and ''[[Canis rufus]]'' appeared, possibly a direct descendant of ''C. edwardii''. Around 0.8 Mya, ''[[Canis ambrusteri]]'' emerged in North America. A large wolf, it was found all over North and Central America and was eventually supplanted by the dire wolf, which then spread into South America during the Late Pleistocene.<ref name=Larson>{{cite web |author=Larson, Robert |title=Wolves, coyotes and dogs (Genus ''Canis'') |series=The Midwestern United States 16,000 years ago |publisher=Illinois State Museum |url=http://exhibits.museum.state.il.us/exhibits/larson/canis.html |access-date=7 June 2014}}</ref> By 0.3 Mya, a number of subspecies of the gray wolf (''C. lupus'') had developed and had spread throughout Europe and northern Asia.<ref>{{cite journal |author=Nowak, R. |year=1992 |title=Wolves: The great travelers of evolution |journal=International Wolf |volume=2 |issue=4 |pages=3–7}}</ref> The gray wolf colonized North America during the late [[Rancholabrean]] era across the Bering land bridge, with at least three separate invasions, with each one consisting of one or more different Eurasian gray wolf clades.<ref name=Chambers>{{cite journal |author1=Chambers, S.M. |author2=Fain, S.R. |author3=Fazio, B. |author4=Amaral, M. |year=2012 |title=An account of the taxonomy of North American wolves from morphological and genetic analyses |journal=North American Fauna |volume=77 |pages=1–67 |doi=10.3996/nafa.77.0001 |doi-access=free }}</ref> MtDNA studies have shown that there are at least four extant ''C. lupus'' lineages.<ref name="gaubert">{{cite journal|author1=Gaubert, P. |author2=Bloch, C. |author3=Benyacoub, S. |author4=Abdelhamid, A. |author5=Pagani, P. |display-authors=etal |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 |bibcode=2012PLoSO...742740G|doi-access=free }}</ref> The dire wolf shared its habitat with the gray wolf, but became extinct in a large-scale extinction event that occurred around 11,500 years ago. It may have been more of a scavenger than a hunter; its molars appear to be adapted for crushing bones and it may have gone extinct as a result of the extinction of the large herbivorous animals on whose carcasses it relied.<ref name=Larson/> In 2015, a study of mitochondrial genome sequences and whole-genome nuclear sequences of African and Eurasian canids indicated that extant wolf-like canids have colonized Africa from Eurasia at least five times throughout the Pliocene and Pleistocene, which is consistent with fossil evidence suggesting that much of African canid fauna diversity resulted from the immigration of Eurasian ancestors, likely coincident with Plio-Pleistocene climatic oscillations between arid and humid conditions. When comparing the African and Eurasian golden jackals, the study concluded that the African specimens represented a distinct monophyletic lineage that should be recognized as a separate species, ''Canis anthus'' ([[African golden wolf]]). According to a phylogeny derived from nuclear sequences, the Eurasian golden jackal (''Canis aureus'') diverged from the wolf/coyote lineage 1.9 [[MYA (unit)|Mya]], but the African golden wolf separated 1.3 Mya. Mitochondrial genome sequences indicated the Ethiopian wolf diverged from the wolf/coyote lineage slightly prior to that.<ref name=Koepfli-2015>{{cite journal |last1=Koepfli |first1=Klaus-Peter |last2=Pollinger |first2=John |last3=Godinho |first3=Raquel |last4=Robinson |first4=Jacqueline |last5=Lea |first5=Amanda |last6=Hendricks |first6=Sarah |last7=Schweizer |first7=Rena M. |last8=Thalmann |first8=Olaf |last9=Silva |first9=Pedro |last10=Fan |first10=Zhenxin |last11=Yurchenko |first11=Andrey A. |last12=Dobrynin |first12=Pavel |last13=Makunin |first13=Alexey |last14=Cahill |first14=James A. |last15=Shapiro |first15=Beth |last16=Álvares |first16=Francisco |last17=Brito |first17=José C. |last18=Geffen |first18=Eli |last19=Leonard |first19=Jennifer A. |last20=Helgen |first20=Kristofer M. |last21=Johnson |first21=Warren E. |last22=o'Brien |first22=Stephen J. |last23=van Valkenburgh |first23=Blaire |last24=Wayne |first24=Robert K. |display-authors=6 |year=2015 |title=Genome-wide evidence reveals that African and Eurasian golden jackals are distinct species |journal=Current Biology |volume=25 |issue=16 |pages=2158–2165 |pmid=26234211 |doi=10.1016/j.cub.2015.06.060 |doi-access=free|bibcode=2015CBio...25.2158K }}</ref>{{rp|S1}}
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