Editing Wolf (section)

== Taxonomy ==
{{cladogram|title=Canine phylogeny with ages of divergence
|caption=Cladogram and divergence of the grey wolf (including the domestic dog) among its closest extant relatives<ref name=Koepfli-2015/>
|cladogram={{clade| style=font-size:85%;line-height:75%;width:475px;
|sublabel1=''3.50 [[Million years ago|mya]]''<!--E-->
|1={{clade
   |sublabel1=''3.06 mya''<!--F-->
   |1={{clade
      |sublabel1=''2.74 mya''<!--G-->
      |1={{clade
         |sublabel1=''1.92 mya''<!--H-->
         |1={{clade
            |sublabel1=''1.62 mya''<!--I-->
            |1={{clade
               |sublabel1=''1.32 mya''<!--J-->
               |1={{clade
                  |sublabel1=''1.10 mya''<!--K-->
                  |1={{clade
                     |1='''Grey wolf''' [[File:Dogs, jackals, wolves, and foxes (Plate I).png|50 px]]
                     |2=[[Coyote]] [[File:Dogs, jackals, wolves, and foxes (Plate IX).png|50 px]]
                     }}
                  |2=[[African wolf]] [[File:Dogs, jackals, wolves, and foxes (Plate XI).jpg|50 px]]
                  }}
               |2=[[Ethiopian wolf]] [[File:Dogs, jackals, wolves, and foxes (Plate VI).png|50 px]]
               }}
            |2=[[Golden jackal]] [[File:Dogs, jackals, wolves, and foxes (Plate X).png|50 px]]
            }}
         |2=[[Dhole]] [[File:Dogs, jackals, wolves, and foxes (Plate XLI).png|50 px]]
         }}
      |2=[[African wild dog]] [[File:Dogs, jackals, wolves, and foxes (Plate XLIV).png|50 px]]
      }}
   |sublabel2=''2.62 mya''<!--D-->
   |2={{clade
      |1=[[Side-striped jackal]] [[File:Dogs, jackals, wolves, and foxes (Plate XIII).png|50 px]]
      |2=[[Black-backed jackal]] [[File:Dogs, jackals, wolves, and foxes (Plate XII).png|50 px]]
      }}
   }}
}}
}}
In 1758, the Swedish botanist and zoologist [[Carl Linnaeus]] published in his ''[[10th edition of Systema Naturae|Systema Naturae]]'' the [[binomial nomenclature]].<ref name=Linnaeus1758/> ''[[Canis]]'' is the Latin word meaning "[[dog]]",<ref>{{OEtymD|canine}}</ref> and under this [[genus]] he listed the doglike carnivores including domestic dogs, wolves, and [[jackal]]s. He classified the domestic dog as ''Canis familiaris'', and the wolf as ''Canis lupus''.<ref name=Linnaeus1758/> Linnaeus considered the dog to be a separate species from the wolf because of its "cauda recurvata" (upturning tail) which is not found in any other [[canid]].<ref name=Clutton-Brock1995/>

=== Subspecies ===
{{Main|Subspecies of Canis lupus}}
{{Further|Pleistocene wolf}}
In the third edition of ''[[Mammal Species of the World]]'' published in 2005, the [[mammalogist]] [[W. Chris Wozencraft|W. Christopher Wozencraft]] listed under ''C. lupus'' 36 wild subspecies, and proposed two additional subspecies: ''familiaris'' (Linnaeus, 1758) and ''[[Canis lupus dingo|dingo]]'' (Meyer, 1793). Wozencraft included ''hallstromi''—the [[New Guinea singing dog]]—as a [[taxonomic synonym]] for the [[dingo]]. Wozencraft referred to a 1999 [[mitochondrial DNA]] (mtDNA) study as one of the guides in forming his decision, and listed the 38 [[Subspecies of Canis lupus|subspecies of ''C. lupus'']] under the biological [[common name]] of "wolf", the [[nominate subspecies]] being the [[Eurasian wolf]] (''C. l. lupus'') based on the [[type specimen]] that Linnaeus studied in Sweden.<ref name=Wozencraft2005/> Studies using [[Paleogenomics|paleogenomic]] techniques reveal that the modern wolf and the dog are [[sister taxa]], as modern wolves are not closely related to the population of wolves that was first [[Domestication|domesticated]].<ref name=Larson2014/> In 2019, a workshop hosted by the [[IUCN]]/Species Survival Commission's Canid Specialist Group considered the New Guinea singing dog and the dingo to be [[feral dog|feral]] ''Canis familiaris'', and therefore should not be assessed for the [[IUCN Red List]].<ref name=Alvares2019/>

=== 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/>

=== Admixture with other canids ===

{{Main|Canid hybrid}}
[[File:F1 wolf-dog hybrids from Wildlife Park Kadzidlowo, Poland.png|thumb|upright=1.5|alt=Photographs of two wolf–dog hybrids standing outdoors on snowy ground|[[Wolfdog|Wolf–dog hybrids]] in the wild animal park at [[Kadzidłowo]], Poland. Left: product of a male wolf and a female [[spaniel]]; right: from a female wolf and a male [[West Siberian Laika]]]]

In the distant past, there was [[gene flow]] between [[African wolf|African wolves]], [[golden jackal]]s, and grey wolves. The African wolf is a descendant of a genetically admixed canid of 72% wolf and 28% Ethiopian wolf ancestry. One African wolf from the Egyptian [[Sinai Peninsula]] showed admixture with Middle Eastern grey wolves and dogs.<ref name=Gopalakrishnan2018/> There is evidence of gene flow between golden jackals and Middle Eastern wolves, less so with European and Asian wolves, and least with North American wolves. This indicates the golden jackal ancestry found in North American wolves may have occurred before the divergence of the Eurasian and North American wolves.<ref name=Sinding2018/>

The common ancestor of the coyote and the wolf is admixed with a [[ghost population]] of an extinct unidentified canid. This canid was genetically close to the [[dhole]] and evolved after the divergence of the [[African hunting dog]] from the other canid species. The basal position of the [[coyote]] compared to the wolf has been proposed to be due to the coyote retaining more of the mitochondrial genome of this unidentified canid.<ref name=Gopalakrishnan2018/> Similarly, a museum specimen of a wolf from southern China collected in 1963 showed a genome that was 12–14% admixed from this unknown canid.<ref name=Wang2019/> In North America, some coyotes and wolves show varying degrees of past [[Coywolf|genetic admixture]].<ref name=Sinding2018/>

In more recent times, some male [[Italian wolf|Italian wolves]] originated from dog ancestry, which indicates female wolves will breed with male dogs in the wild.<ref name=Iacolina2010/> In the [[Caucasus Mountains]], ten percent of dogs including [[livestock guardian dog]]s, are first generation hybrids.<ref name=Kopaliani2014/> Although mating between golden jackals and wolves has never been observed, evidence of [[jackal-wolf hybrid]]ization was discovered through mitochondrial DNA analysis of jackals living in the Caucasus Mountains<ref name=Kopaliani2014/> and in Bulgaria.<ref name=Moura2013/> In 2021, a genetic study found that the dog's similarity to the extant grey wolf was the result of substantial dog-into-wolf [[gene flow]], with little evidence of the reverse.<ref name="Bergström2020"/>