Skulls of various wolf subspecies from North AmericaPresent and historical range of wild subspecies ofC. lupus. This map uses the more broadly defined North American subspecies of Nowak (1995),[1][2] but see also the map under the section titled North America.
In 1758, the Swedish botanist and zoologistCarl Linnaeus published in hisSystema Naturae thebinomial nomenclature – or the two-word naming – ofspecies.Canis is the Latin word meaning "dog",[3] and under thisgenus he listed the dog-like carnivores including domestic dogs, wolves, and jackals. He classified the domestic dog asCanis familiaris, and on the next page he classified the wolf asCanis lupus.[4] Linnaeus considered the dog to be a separate species from the wolf because of its head, body, andcauda recurvata – its upturning tail – which is not found in any othercanid.[5]
In 1999, a study ofmitochondrial DNA indicated that the domestic dog may have originated from multiple wolf populations, with thedingo andNew Guinea singing dog "breeds" having developed at a time when human populations were more isolated from each other.[6] In the third edition ofMammal Species of the World published in 2005, the mammalogistW. Christopher Wozencraft listed under the wolfCanis lupus some 36 wild subspecies, and proposed two additional subspecies:familiaris Linnaeus, 1758 anddingo Meyer, 1793. Wozencraft includedhallstromi – the New Guinea singing dog – as ataxonomic synonym for the dingo. Wozencraft referred to the mDNA study as one of the guides in forming his decision, and listed the 38 subspecies under the biologicalcommon name of "wolf", with thenominate subspecies being theEurasian wolf (Canis lupus lupus) based on thetype specimen that Linnaeus studied in Sweden.[7] However, the classification of several of thesecanines as either species or subspecies has recently[when?]been challenged.
Sokolov and Rossolimo (1985) recognised nine Old World subspecies of wolf. These wereC. l. lupus,C. l. albus,C. l. pallipes,C. l. cubanenesis,C. l. campestris,C. l. chanco,C. l. desertorum,C. l. hattai, andC. l. hodophilax.[1] In his 1995 statistical analysis of skullmorphometrics, mammalogist Robert Nowak recognized the first four of those subspecies, synonymizedcampestris,chanco anddesertorum withC. l. lupus, but did not examine the two Japanese subspecies. In addition, he recognizedC. l. communis as a subspecies distinct fromC. l. lupus.[1] In 2003, Nowak also recognized the distinctiveness ofC. l. arabs,C. l. hattai,C. l. italicus, andC. l. hodophilax.[9] In 2005,MSW3 includedC. l. filchneri.[7] In 2003, two forms were distinguished in southern China andInner Mongolia as being separate fromC. l. chanco andC. l. filchneri and have yet to be named.[10][11]
A small, "desert-adapted" subspecies that is around 66 cm tall and weighs, on average, about 18 kg.[16] Its fur coat varies from short in the summer to long in the winter, possibly because of solar radiation.[17]
Long sharp face, elevated brows, broad head, large pointed ears, thick woolly pelage and very full brush of medial length. Above, dull earthy-brown; below, with the entire face and limbs yellowish-white.[21]
The fur isfulvous, on the back longer, rigid, with intermixed black and gray hairs; the throat, chest, belly, and inside of the legs pure white; head pale gray-brown; forehead grizzled with short black and gray hairs.[23]
Generally 52–60 cm tall at the shoulders and measures 117 to 124 cm from nose to tail tip. The average weight is 13 to 20 kg.[27] Fur color is mostly sandy- to reddish-brown, but can include tan patterns and can also be occasionally light brown, black or white.[28]
The domestic dog is a divergent subspecies of the gray wolf and was derived from an extinct population of Late Pleistocene wolves.[8][31][32] Throughselective pressure andselective breeding, the domestic dog has developed into hundreds of variedbreeds and shows more behavioral and morphological variation than any other land mammal.[33]
Worldwide in association with humans
Increasingly proposed as the speciesCanis familiaris but debated[34][30]
aegyptius Linnaeus, 1758,
alco C. E. H. Smith, 1839,americanus Gmelin, 1792,anglicus Gmelin, 1792,antarcticus Gmelin, 1792,aprinus Gmelin, 1792,aquaticus Linnaeus, 1758,aquatilis Gmelin, 1792,avicularis Gmelin, 1792,borealis C. E. H. Smith, 1839,brevipilis Gmelin, 1792,cursorius Gmelin, 1792,domesticus Linnaeus, 1758,extrarius Gmelin, 1792,ferus C. E. H. Smith, 1839,fricator Gmelin, 1792,fricatrix Linnaeus, 1758,fuillus Gmelin, 1792,gallicus Gmelin, 1792,glaucus C. E. H. Smith, 1839,graius Linnaeus, 1758,grajus Gmelin, 1792,hagenbecki Krumbiegel, 1950,haitensis C. E. H. Smith, 1839,hibernicus Gmelin, 1792,hirsutus Gmelin, 1792,hybridus Gmelin, 1792,islandicus Gmelin, 1792,italicus Gmelin, 1792,laniarius Gmelin, 1792,leoninus Gmelin, 1792,leporarius C. E. H. Smith, 1839,major Gmelin, 1792,mastinus Linnaeus, 1758,melitacus Gmelin, 1792,melitaeus Linnaeus, 1758,minor Gmelin, 1792,molossus Gmelin, 1792,mustelinus Linnaeus, 1758,obesus Gmelin, 1792,orientalis Gmelin, 1792,pacificus C. E. H. Smith, 1839,plancus Gmelin, 1792,pomeranus Gmelin, 1792,sagaces C. E. H. Smith, 1839,sanguinarius C. E. H. Smith, 1839,sagax Linnaeus, 1758,scoticus Gmelin, 1792,sibiricus Gmelin, 1792,suillus C. E. H. Smith, 1839,terraenovae C. E. H. Smith, 1839,terrarius C. E. H. Smith, 1839,turcicus Gmelin, 1792,urcani C. E. H. Smith, 1839,variegatus Gmelin, 1792,venaticus Gmelin, 1792,
The pelt is generally of a grey-fulvous colour, which reddens in summer. The belly and cheeks are more lightly coloured, and dark bands are present on the back and tail tip, and occasionally along the fore limbs.
A small subspecies with pelage shorter than that of northern wolves and with little to no underfur.[39] Fur color ranges from grayish-red to reddish-white with black tips. The dark V-shaped stripe over the shoulders is much more pronounced than in northern wolves. The underparts and legs are more or less white.[40]
North American wolf subspecies distribution according to Goldman (1944) and Hall (1981). Hall split offC. l. griseoalbus from Goldman'sC. l. occidentalis. These subspecies are included inMSW3 2005.
For North America, in 1944 the zoologistEdward Goldman recognized as many as 23 subspecies based onmorphology.[42] In 1959, E. Raymond Hall proposed that there had been 24 subspecies oflupus in North America.[43] In 1970,L. David Mech proposed that there was "probably far too many subspecific designations...in use", as most did not exhibit enough points of differentiation to be classified as separate subspecies.[44] The 24 subspecies were accepted by many authorities in 1981 and these were based on morphological or geographical differences, or a unique history.[45] In 1995, the American mammalogist Robert M. Nowak analyzed data on the skull morphology of wolf specimens from around the world. For North America, he proposed that there were only five subspecies of the wolf. These include a large-toothed Arctic wolf namedC. l. arctos, a large wolf from Alaska and western Canada namedC. l. occidentalis, a small wolf from southeastern Canada namedC. l. lycaon, a small wolf from the southwestern U.S. namedC. l. baileyi and a moderate-sized wolf that was originally found from Texas to Hudson Bay and from Oregon to Newfoundland namedC. l. nubilus.[46][1]
The taxonomic classification ofCanis lupus inMammal Species of the World (3rd edition, 2005) listed 27 subspecies of North American wolf,[7] corresponding to the 24Canis lupus subspecies and the threeCanis rufus subspecies of Hall (1981).[1] The table below shows the extant subspecies, with the extinct ones listed in the following section.
The domestic dog is a divergent subspecies of the gray wolf and was derived from an extinct population of Late Pleistocene wolves.[8][31][32] Throughselective pressure andselective breeding, the domestic dog has developed into hundreds of variedbreeds and shows more behavioral and morphological variation than any other land mammal.[33]
aegyptius Linnaeus, 1758,
alco C. E. H. Smith, 1839,americanus Gmelin, 1792,anglicus Gmelin, 1792,antarcticus Gmelin, 1792,aprinus Gmelin, 1792,aquaticus Linnaeus, 1758,aquatilis Gmelin, 1792,avicularis Gmelin, 1792,borealis C. E. H. Smith, 1839,brevipilis Gmelin, 1792,cursorius Gmelin, 1792,domesticus Linnaeus, 1758,extrarius Gmelin, 1792,ferus C. E. H. Smith, 1839,fricator Gmelin, 1792,fricatrix Linnaeus, 1758,fuillus Gmelin, 1792,gallicus Gmelin, 1792,glaucus C. E. H. Smith, 1839,graius Linnaeus, 1758,grajus Gmelin, 1792,hagenbecki Krumbiegel, 1950,haitensis C. E. H. Smith, 1839,hibernicus Gmelin, 1792,hirsutus Gmelin, 1792,hybridus Gmelin, 1792,islandicus Gmelin, 1792,italicus Gmelin, 1792,laniarius Gmelin, 1792,leoninus Gmelin, 1792,leporarius C. E. H. Smith, 1839,major Gmelin, 1792,mastinus Linnaeus, 1758,melitacus Gmelin, 1792,melitaeus Linnaeus, 1758,minor Gmelin, 1792,molossus Gmelin, 1792,mustelinus Linnaeus, 1758,obesus Gmelin, 1792,orientalis Gmelin, 1792,pacificus C. E. H. Smith, 1839,plancus Gmelin, 1792,pomeranus Gmelin, 1792,sagaces C. E. H. Smith, 1839,sanguinarius C. E. H. Smith, 1839,sagax Linnaeus, 1758,scoticus Gmelin, 1792,sibiricus Gmelin, 1792,suillus C. E. H. Smith, 1839,terraenovae C. E. H. Smith, 1839,terrarius C. E. H. Smith, 1839,turcicus Gmelin, 1792,urcani C. E. H. Smith, 1839,variegatus Gmelin, 1792,venaticus Gmelin, 1792,
Two forms are known – a small, reddish-brown colored form called the Algonquin wolf; and a slightly larger, more grayish-brown form called theGreat Lakes wolf, which is an admixture of the Algonquin wolf and other gray wolves.[70]
The Algonquin form occupies centralOntario and southwesternQuebec, particularly in and nearby protected areas, such asAlgonquin Provincial Park inOntario, and possibly extreme northeastern U.S. and western New Brunswick. The Great Lakes form occupies northern Ontario,Wisconsin andMinnesota, the Upper Peninsula ofMichigan and southernManitoba. Overlaps of the two forms occur, with intermixing in the southern portions of northern Ontario.
canadensis de Blainville, 1843,ungavensis Comeau, 1940[71]
The Algonquin form is currently (2025) recognized as the speciesCanis lycaon[72] by the American Society of Mammalogists, but its taxonomy is still debated.[73]
TheC. l. occidentalis of Nowak (1995) synonymizesalces,columbianus,griseoalbus,mackenzii,pambasileus andtundrarum, which is the currently (2025) recognized classification.[1]
The second largest subspecies of wolf, second in skull and tooth proportions only tooccidentalis (see chart above), with fur that is black, white or a mixture of both in color.[82]
variabilis Wied-Neuwied, 1841.[87]Previously thought extinct in 1926, the Great Plains wolf's descendants were found in the northeastern region of the United States and have become federally protected since 1974.[88]
As of 2025 the classification of the more broadly definedC. l. nubilus of Nowak (1995) synonymizesbeothucus,fuscus,hudsonicus,irremotus,labridorius,manningi,mogollonensis,monstrabilis andyoungi, in which case the subspecies is extant in Canada (see infobox map).[1]
Has a brownish or cinnamon pelt, with gray and black shading on the back and tail. Generally intermediate in size between other North American wolf subspecies and thecoyote. Like other wolves, it has almond-shaped eyes, a broad muzzle and a wide nose pad though, like the coyote, its ears are proportionately larger. It has a deeper profile, a longer and broader head than the coyote, and has a less prominent ruff than other wolves.[89]
Currently considered a distinct species,Canis rufus, but this proposal is still debated.[2] As a species, the red wolf would have the following subspecies:
Thelargest subspecies of all known extinct and extant wolves fromWestern Europe. The wolf's long bones are 10% longer than those of extant European wolves, 12% larger than those ofC. l. santenaisiensis and 20% longer than those ofC. l. lunellensis.[94] The teeth are robust, the posterior denticules on the lower premolars p2, p3, p4 and upper P2 and P3 are highly developed, and the diameter of the lower carnassial (m1) were larger than any known European wolf.[94]
A jet-black subspecies that is described as having been extremely similar to thered wolf in both size and weight.[108] This subspecies became extinct in 1908.[109]
A slender, short-legged subspecies with light, tawny-colored fur. The dark bands present on the forelimbs of the mainland Italian wolf were absent or poorly defined in the Sicilian wolf.
In 2019, a workshop hosted by theIUCN/SSC Canid Specialist Group considered the New Guinea singing dog and the dingo to be feral dogs (Canis familiaris).[127] In 2020, a literature review of canid domestication stated that modern dogs were not descended from the sameCanis lineage as modern wolves, and proposed that dogs may be descended from a Pleistocene wolf closer in size to a village dog.[128] In 2021, theAmerican Society of Mammalogists also considered dingos a feral dog (Canis familiaris) population.[30]
Giuseppe Altobello's 1925 comparative illustration of the skulls and dentition ofC. l. lupus (a) andC. l. italicus (b). The distinct status of the latter is currently unrecognized by MSW3.
TheItalian wolf (or Apennine wolf) was first recognised as a distinct subspecies (Canis lupus italicus) in 1921 by zoologist Giuseppe Altobello.[129] Altobello's classification was later rejected by several authors, includingReginald Innes Pocock, whosynonymisedC. l. italicus withC. l. lupus.[130] In 2002, the noted paleontologist R.M. Nowak reaffirmed the morphological distinctiveness of the Italian wolf and recommended the recognition ofCanis lupus italicus.[130] A number of DNA studies have found the Italian wolf to be genetically distinct.[131][132] In 2004, the genetic distinction of the Italian wolf subspecies was supported by analysis which consistently assigned all the wolfgenotypes of a sample inItaly to a single group. This population also showed a uniquemitochondrial DNA control-regionhaplotype, the absence of privatealleles and lowerheterozygosity at microsatelliteloci, as compared to other wolf populations.[133] In 2010, a genetic analysis indicated that a single wolf haplotype (w22) unique to the Apennine Peninsula and one of the two haplotypes (w24, w25), unique to the Iberian Peninsula, belonged to the same haplogroup as the prehistoric wolves of Europe. Another haplotype (w10) was found to be common to the Iberian peninsula and the Balkans. These three populations with geographic isolation exhibited a near lack of gene flow and spatially correspond to three glacial refugia.[134]
The taxonomic referenceMammal Species of the World (3rd edition, 2005) does not recognizeCanis lupus italicus; however,NCBI/Genbank publishes research papers under that name.[135]
TheIberian wolf was first recognised as a distinct subspecies (Canis lupus signatus) in 1907 by zoologistÁngel Cabrera. The wolves of the Iberian peninsula havemorphologically distinct features from otherEurasian wolves and each are considered by their researchers to represent their own subspecies.[136][137]
The Himalayan wolf is distinguished by itsmitochondrial DNA, which is basal to all other wolves. The taxonomic name of this wolf is disputed, with the speciesCanis himalayensis being proposed based on two limited DNA studies.[139][140][141] In 2017, a study ofmitochondrial DNA,X-chromosome (maternal lineage) markers andY-chromosome (male lineage) markers found that the Himalayan wolf was geneticallybasal to the Holarctic grey wolf and has an association with theAfrican golden wolf.[142]
In 2019, a workshop hosted by theIUCN/SSC Canid Specialist Group noted that the Himalayan wolf's distribution included the Himalayan range and the Tibetan Plateau. The group recommends that this wolf lineage be known as the "Himalayan wolf" and classified asCanis lupus chanco until a genetic analysis of the holotypes is available.[127] In 2020, further research on the Himalayan wolf found that it warranted species-level recognition under the Unified Species Concept, the Differential Fitness Species Concept, and the Biological Species Concept. It was identified as an Evolutionary Significant Unit that warranted assignment onto theIUCN Red List for its protection.[143]
TheIndian plains wolf is a proposed clade within the Indian wolf (Canis lupus pallipes) that is distinguished by its mitochondrial DNA, which is basal to all other wolves except for the Himalayan wolf. The taxonomic status of this wolf clade is disputed, with the separate speciesCanis indica being proposed based on two limited DNA studies.[139][140] The proposal has not been endorsed because it relied on a limited number of museum and zoo samples that may not have been representative of the wild population, and a call for further fieldwork has been made.[141]
The taxonomic referenceMammal Species of the World (3rd edition, 2005) does not recognizeCanis indica; however,NCBI/Genbank lists it as a new subspecies,Canis lupus indica.[144]
In 2017, a comprehensive study found that the gray wolf was present across all of mainland China, both in the past and today. It exists in southern China, which refutes claims made by some researchers in the Western world that the wolf had never existed in southern China.[145][146] This wolf has not been taxonomically classified.[10][11]
In 2019, a genomic study on the wolves of China included museum specimens of wolves from southern China that were collected between 1963 and 1988. The wolves in the study formed three clades: northern Asian wolves that included those from northern China and eastern Russia,Himalayan wolves from the Tibetan Plateau, and a unique population from southern China. One specimen fromZhejiang Province in eastern China shared gene flow with the wolves from southern China; however, its genome was 12–14 percent admixed with a canid that may be the dhole or an unknown canid that predates the genetic divergence of thedhole. The wolf population from southern China is believed to still exist in that region.[147]
A study of the three coastal wolves indicates a close phylogenetic relationship across regions that are geographically and ecologically contiguous, and the study proposed thatCanis lupus ligoni (theAlexander Archipelago wolf),Canis lupus columbianus (theBritish Columbian wolf), andCanis lupus crassodon (theVancouver Coastal Sea wolf) should be recognized as a single subspecies ofCanis lupus, synonymized asCanis lupus crassodon.[148] They share the same habitat and prey species, and form one study's six identified North Americanecotypes – a genetically and ecologically distinct population separated from other populations by their different types of habitat.[149][150]
The eastern wolf has two proposals over its origin. One is that the eastern wolf is a distinct species (C. lycaon) that evolved in North America, as opposed to the gray wolf that evolved in the Old World, and is related to the red wolf. The other is that it is derived from admixture between gray wolves, which inhabited the Great Lakes area and coyotes, forming a hybrid that was classified as a distinct species by mistake.[151]
The red wolf is an enigmatic taxon, of which there are two proposals over its origin. One is that the red wolf is a distinct species (C. rufus) that has undergone human-influenced admixture with coyotes. The other is that it was never a distinct species but was derived from past admixture between coyotes and gray wolves, due to the gray wolf population being eliminated by humans.[151]
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