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=== Evolution === {{Main|Evolution of the wolf}} {{Further|Domestication of the dog}} [[File:Canis mosbachensis Wikipedia Juandertal.jpg|thumb|right|Life restoration of ''[[Canis mosbachensis]]'', the wolf's immediate ancestor]] The [[phylogenetic]] descent of the extant wolf ''C. lupus'' from the earlier ''[[Canis mosbachensis|C. mosbachensis]]'' (which in turn descended from ''[[Canis etruscus|C. etruscus]]'') is widely accepted.{{sfn|Mech|Boitani|2003|pp=239–245}} Among the oldest fossils of the modern grey wolf is from Ponte Galeria in Italy, dating to 406,500 ± 2,400 years ago.<ref name=":2">{{Cite journal |last1=Iurino |first1=Dawid A. |last2=Mecozzi |first2=Beniamino |last3=Iannucci |first3=Alessio |last4=Moscarella |first4=Alfio |last5=Strani |first5=Flavia |last6=Bona |first6=Fabio |last7=Gaeta |first7=Mario |last8=Sardella |first8=Raffaele |date=2022-02-25 |title=A Middle Pleistocene wolf from central Italy provides insights on the first occurrence of Canis lupus in Europe |journal=Scientific Reports |language=en |volume=12 |issue=1 |page=2882 |doi=10.1038/s41598-022-06812-5 |issn=2045-2322 |pmc=8881584 |pmid=35217686|bibcode=2022NatSR..12.2882I }}</ref> Remains from Cripple Creek Sump in Alaska may be considerably older, around 1 million years old,<ref name=Tedford2009/> though differentiating between the remains of modern wolves and ''C. mosbachensis'' is difficult and ambiguous, with some authors choosing to include C. ''mosbachensis'' (which first appeared around 1.4 million years ago) as an early subspecies of ''C. lupus.''<ref name=":2" /> Considerable morphological diversity existed among wolves by the [[Late Pleistocene]]. Many Late Pleistocene wolf populations had more robust skulls and teeth than modern wolves, often with a shortened [[Rostrum (anatomy)#Vertebrates|snout]], a pronounced development of the [[temporalis]] muscle, and robust [[premolar]]s. It is proposed that these features were specialized adaptations for the processing of carcass and bone associated with the hunting and scavenging of [[Pleistocene megafauna]]. Compared with modern wolves, some Pleistocene wolves showed an increase in tooth breakage similar to that seen in the extinct [[dire wolf]]. This suggests they either often processed carcasses, or that they competed with other carnivores and needed to consume their prey quickly. The frequency and location of tooth fractures in these wolves indicates they were habitual bone crackers like the modern [[spotted hyena]].<ref name=Thalmann2018/> [[Genome|Genomic]] studies suggest modern wolves and dogs descend from a common ancestral wolf population.<ref name=Freedman2014/><ref name=Skoglund2015/><ref name=Fan2016/> A 2021 study found that the [[Himalayan wolf]] and the [[Indian wolf|Indian plains wolf]] are part of a [[Lineage (evolution)|lineage]] that is [[Basal (phylogenetics)|basal]] to other wolves and [[Cladogenesis|split]] from them 200,000 years ago.<ref name=Hennelly2021/> Other wolves appear to share most of their common ancestry much more recently, within the last 23,000 years (around the peak and the end of the [[Last Glacial Maximum]]), originating from [[Siberia]]<ref name=":0">{{Cite journal |last1=Bergström |first1=Anders |last2=Stanton |first2=David W. G. |last3=Taron |first3=Ulrike H. |last4=Frantz |first4=Laurent |last5=Sinding |first5=Mikkel-Holger S. |last6=Ersmark |first6=Erik |last7=Pfrengle |first7=Saskia |last8=Cassatt-Johnstone |first8=Molly |last9=Lebrasseur |first9=Ophélie |last10=Girdland-Flink |first10=Linus |last11=Fernandes |first11=Daniel M. |last12=Ollivier |first12=Morgane |last13=Speidel |first13=Leo |last14=Gopalakrishnan |first14=Shyam |last15=Westbury |first15=Michael V. |date=2022-07-14 |title=Grey wolf genomic history reveals a dual ancestry of dogs |journal=Nature |language=en |volume=607 |issue=7918 |pages=313–320 |doi=10.1038/s41586-022-04824-9 |issn=0028-0836 |pmc=9279150 |pmid=35768506|bibcode=2022Natur.607..313B }}</ref> or [[Beringia]].<ref name=":1">{{Cite journal |last1=Loog |first1=Liisa |last2=Thalmann |first2=Olaf |last3=Sinding |first3=Mikkel-Holger S. |last4=Schuenemann |first4=Verena J. |last5=Perri |first5=Angela |last6=Germonpré |first6=Mietje |last7=Bocherens |first7=Herve |last8=Witt |first8=Kelsey E. |last9=Samaniego Castruita |first9=Jose A. |last10=Velasco |first10=Marcela S. |last11=Lundstrøm |first11=Inge K. C. |last12=Wales |first12=Nathan |last13=Sonet |first13=Gontran |last14=Frantz |first14=Laurent |last15=Schroeder |first15=Hannes |date=May 2020 |title=Ancient DNA suggests modern wolves trace their origin to a Late Pleistocene expansion from Beringia |journal=Molecular Ecology |language=en |volume=29 |issue=9 |pages=1596–1610 |doi=10.1111/mec.15329 |issn=0962-1083 |pmc=7317801 |pmid=31840921|bibcode=2020MolEc..29.1596L }}</ref> While some sources have suggested that this was a consequence of a [[population bottleneck]],<ref name=":1" /> other studies have suggested that this a result of [[gene flow]] homogenising ancestry.<ref name=":0" /> A 2016 genomic study suggests that Old World and New World wolves split around 12,500 years ago followed by the [[Genetic divergence|divergence]] of the lineage that led to dogs from other Old World wolves around 11,100–12,300 years ago.<ref name=Fan2016/> An extinct [[Megafaunal wolf|Late Pleistocene wolf]] may have been the ancestor of the dog,<ref name=Freedman2017/><ref name=Thalmann2018/> with the dog's similarity to the extant wolf being the result of [[genetic admixture]] between the two.<ref name=Thalmann2018/> The dingo, [[Basenji]], [[Tibetan Mastiff]] and Chinese indigenous breeds are basal members of the domestic dog clade. The divergence time for wolves in Europe, the Middle East, and Asia is estimated to be fairly recent at around 1,600 years ago. Among New World wolves, the [[Mexican wolf]] diverged around 5,400 years ago.<ref name=Fan2016/>
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