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Canis
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==Taxonomy== The [[genus]] ''Canis'' ([[Carl Linnaeus]], 1758) was published in the [[10th edition of Systema Naturae]]<ref name=linnaeus1758/> and included the dog-like carnivores: the domestic dog, wolves, coyotes and jackals. All species within ''Canis'' are [[Phylogenetics|phylogenetically]] closely related with 78 [[chromosome]]s and can potentially [[hybrid (biology)|interbreed]].<ref name=wayne1999/> In 1926, the [[International Commission on Zoological Nomenclature]] (ICZN) in Opinion 91 included Genus ''Canis'' on its ''Official Lists and Indexes of Names in Zoology''.<ref name=iczn1926/> In 1955, the ICZN's Direction 22 added ''[[Dog|Canis familiaris]]'' as the [[type species]] for genus ''Canis'' to the official list.<ref name=iczn1955/> {{Blockquote|''Canis'' is primitive relative to ''Cuon'', ''Lycaon'', and ''Xenocyon'' in its relatively larger canines and lack of such dental adaptations for hypercarnivory as m1–m2 metaconid and entoconid small or absent; M1–M2 hypocone small; M1–M2 lingual cingulum weak; M2 and m2 small, may be single-rooted; m3 small or absent; and wide palate.|[[Richard H. Tedford]]<ref name=Tedford2009/>}} The cladogram below is based on the [[DNA]] phylogeny of Lindblad-Toh ''et al''. (2005),<ref name=LindbladToh2005/> modified to incorporate recent findings on ''Canis'' species,<ref name=Koepfli2015/><ref>{{Cite journal |last1=Wilson |first1=Paul J. |last2=Grewal |first2=Sonya |last3=Lawford |first3=Ian D. |last4=Heal |first4=Jennifer NM |last5=Granacki |first5=Angela G. |last6=Pennock |first6=David |last7=Theberge |first7=John B. |last8=Theberge |first8=Mary T. |last9=Voigt |first9=Dennis R. |last10=Waddell |first10=Will |last11=Chambers |first11=Robert E. |date=2011-02-15 |title=DNA profiles of the eastern Canadian wolf and the red wolf provide evidence for a common evolutionary history independent of the gray wolf |url=https://cdnsciencepub.com/doi/abs/10.1139/z00-158 |journal=Canadian Journal of Zoology |volume=78 |issue=12 |pages=2156–2166 |language=en |doi=10.1139/z00-158|url-access=subscription }}</ref> {{Clade |style=font-size:85%; line-height:85% |label1=''Canis'' |1={{Clade |1={{Clade |1={{Clade |1={{Clade |1={{Clade |1={{Clade |1=''[[Canis latrans]]'' (coyote) [[File:Dogs, jackals, wolves, and foxes (Plate IX).png|50 px]] |2={{Clade |1=''[[Canis rufus]]'' (red wolf) [[File:Dogs, jackals, wolves, and foxes (Plate V) C. l. rufus mod.png|50 px]] |2=''[[Canis lycaon]]'' (eastern wolf) [[File:Dogs, jackals, wolves, and foxes (Plate V).png|50 px]] }} }} |2={{Clade |1=''[[Canis lupus]]'' (gray wolf) [[File:Dogs, jackals, wolves, and foxes (Plate I).png|50 px]] |2=''[[Canis familiaris]]'' (domestic dog) <span style="{{MirrorH}}">[[File:202104 Dog.svg|50 px]]</span> }} }} |2=''[[Canis lupaster]]'' ([[African golden wolf]]) [[File:Dogs,_jackals,_wolves,_and_foxes_(Plate_XI).png|50px]] }} |2=''[[Canis simensis]]'' ([[Ethiopian wolf]]) [[File:Dogs, jackals, wolves, and foxes (Plate VI).png|50 px]] }} |2=''[[Canis aureus]]'' ([[golden jackal]]) [[File:Dogs, jackals, wolves, and foxes (Plate X).png|50 px]] }} }} }} In 2019, a workshop hosted by the [[IUCN]]/SSC Canid Specialist Group recommends that because DNA evidence shows the [[side-striped jackal]] (''Canis adustus'') and [[black-backed jackal]] (''Canis mesomelas'') to form a monophyletic lineage that sits outside of the ''Canis''/''Cuon''/''Lycaon'' clade, that they should be placed in a distinct genus, ''Lupulella'' Hilzheimer, 1906 with the names ''Lupulella adusta'' and ''Lupulella mesomelas''.<ref name=Alvares2019/> ===Evolution=== :''See further: [[Canidae#Evolution|Evolution of the canids]]'' The fossil record shows that [[Feliformia|feliform]]s and [[Caniformia|caniform]]s emerged within the clade [[Carnivoramorpha]] 43 million [[Years before present|YBP]].<ref name=flynn2005/> The caniforms included the fox-like genus ''[[Leptocyon]]'', whose various species existed from 24 million YBP before branching 11.9 million YBP into ''[[Vulpes]]'' (foxes) and Canini (canines). The jackal-sized ''[[Eucyon]]'' existed in North America from 10 million YBP and by the [[Early Pliocene]] about 6-5 million YBP the coyote-like ''Eucyon davisi''<ref name=fossilworks5/> invaded Eurasia. The canids that had emigrated from North America to Eurasia – ''[[Eucyon]]'', ''[[Vulpes]]'', and ''[[Nyctereutes]]'' – were small to medium-sized predators during the Late Miocene and Early Pliocene but they were not the top predators. [[File:C. dirus, C. lupus, C. lycaon, C. rufus, C. latrans, C. anthus C. aureus & C. mesomelas skulls.jpg|thumb|Skulls of dire wolf (''Aenocyon dirus''), gray wolf (''C. lupus''), eastern wolf (''C. lycaon''), red wolf (''C. rufus''), coyote (''C. latrans''), African golden wolf (''C. lupaster''), golden jackal (''C. aureus'') and black-backed jackal (''Lupulella mesomelas'')]] For ''Canis'' populations in the New World, ''Eucyon'' in North America gave rise to early North American ''Canis'' which first appeared in the [[Miocene]] (6 million YBP) in south-western United States and Mexico. By 5 million YBP the larger ''[[Canis lepophagus]]'', ancestor of wolves and coyotes, appeared in the same region.<ref name="wang2008" />{{rp|p58}} Around 5 million years ago, some of the Old World ''Eucyon'' evolved into the first members of ''Canis'',<ref name="Perri-2021">{{Cite journal|last1=Perri|first1=Angela R.|last2=Mitchell|first2=Kieren J.|last3=Mouton|first3=Alice|last4=Álvarez-Carretero|first4=Sandra|last5=Hulme-Beaman|first5=Ardern|last6=Haile|first6=James|last7=Jamieson|first7=Alexandra|last8=Meachen|first8=Julie|last9=Lin|first9=Audrey T.|last10=Schubert|first10=Blaine W.|last11=Ameen|first11=Carly|date=2021-01-13|title=Dire wolves were the last of an ancient New World canid lineage|url=https://www.nature.com/articles/s41586-020-03082-x|journal=Nature|volume=591|issue=7848|language=en|pages=87–91|doi=10.1038/s41586-020-03082-x|pmid=33442059|bibcode=2021Natur.591...87P|s2cid=231604957 |issn=1476-4687}}</ref> and the position of the canids would change to become a dominant predator across the [[Palearctic realm|Palearctic]]. The wolf-sized ''[[Evolution of the wolf#Canis chihliensis|C. chihliensis]]'' appeared in northern China in the Mid-Pliocene around 4-3 million YBP. This was followed by an explosion of ''Canis'' evolution across Eurasia in the Early Pleistocene around 1.8 million YBP in what is commonly referred to as the ''wolf event''. It is associated with the formation of the [[mammoth steppe]] and continental glaciation. ''Canis'' spread to Europe in the forms of ''[[Canis arnensis|C. arnensis]]'', ''[[Canis etruscus|C. etruscus]]'', and ''[[Canis falconeri|C. falconeri]]''.<ref name=wang2008/>{{rp|p148}} However, a 2021 genetic study of the [[dire wolf]] (''Aenocyon dirus''), previously considered a member of ''Canis'', found that it represented the last member of an ancient lineage of canines originally indigenous to the New World that had diverged prior to the appearance of ''Canis'', and that its lineage had been distinct since the Miocene with no evidence of introgression with ''Canis''. The study hypothesized that the [[Neogene]] canids in the New World, ''[[Armbruster's wolf|Canis armbrusteri]]'' and ''[[Canis edwardii]]'', were possibly members of the distinct dire wolf lineage that had [[Convergent evolution|convergently evolved]] a very similar appearance to members of ''Canis''. True members of ''Canis'', namely the [[Gray Wolf|gray wolf]] and [[coyote]], likely only arrived in the New World during the [[Late Pleistocene]], where their dietary flexibility and/or ability to hybridize with other canids allowed them to survive the [[Quaternary extinction event]], unlike the dire wolf.<ref name="Perri-2021" /> ''[[Xenocyon]]'' (strange wolf) is an extinct [[subgenus]] of ''Canis''.<ref name=rook1994/> The diversity of the ''Canis'' group decreased by the end of the [[Early Pleistocene]] to the [[Middle Pleistocene]] and was limited in Eurasia to the small wolves of the ''[[Canis mosbachensis]]–Canis variabilis'' group and the large hypercarnivorous [[Xenocyon#Canis (Xenocyon) lycaonoides|''Canis (Xenocyon) lycaonoides'']].<ref name=sotnikova2010/> The hypercarnivore ''Xenocyon'' gave rise to the modern [[dhole]] and the [[African wild dog]].<ref name=wang2008/>{{rp|p149}}
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