Subspecies of Canis lupus
|Canis lupus species
Temporal range: 0.7–0 Ma Middle Pleistocene – Recent
|Skulls of various gray wolf subspecies from North America|
Numerous and disputed
|Historical range of wild subspecies of C. lupus|
Canis lupus has 37 subspecies currently described, including the dingo, Canis lupus dingo, and the domestic dog, Canis lupus familiaris, and many subspecies of wolf throughout the Northern Hemisphere. The nominate subspecies is Canis lupus lupus.
Canis lupus is assessed as least concern by the IUCN, as its relatively widespread range and stable population trend mean that the species, at global level, does not meet, or nearly meet, any of the criteria for the threatened categories. However, some local populations are classified as endangered, and some subspecies are endangered or extinct.
Biological taxonomy is not fixed, and placement of taxa is reviewed as a result of new research. The current categorization of subspecies of Canis lupus is shown below. Also included are synonyms, which are now discarded duplicate or incorrect namings, or in the case of the domestic dog synonyms, old taxa referring to subspecies of domestic dog which, when the dog was declared a subspecies itself, had nowhere else to go. Common names are given but may vary, as they have no set meaning.
- 1 Taxonomy
- 2 History of classification
- 3 List of subspecies
- 4 Disputed subspecies and species
- 5 See also
- 6 References
- 7 External links
A subspecies is the taxonomic rank below species. When geographically separate populations of a species exhibit recognizable phenotypic differences, biologists may identify these as separate subspecies; a subspecies is a recognized local variant of a species. The thirty-seven subspecies of Canis lupus are listed in Mammal Species of the World third edition that was published in 2005. The nominate subspecies is the Eurasian wolf (Canis lupus lupus), also known as the common wolf. The subspecies includes the domestic dog, dingo, eastern wolf and red wolf. However, the classification of several as either species or subspecies has recently been challenged.
For Eurasia, in 1995 mammalogist Robert Nowak recognized five subspecies from Eurasia based on skull morphology; C. l. lupus, C. l. albus, C. l. pallipes, C. l. cubanensis and C. l. communis. In 2003, Nowak also recognized the distinctiveness of C. l. arabs, C. l. hattai, C. l. hodophilax and C. l. lupaster.
For North America, in 1944 the zoologist Edward Goldman recognized as many as 23 subspecies in North America, based on morphology alone. In 1959, E. Raymond Hall proposed that there had been 24 subspecies of lupus in North America. In 1970, L David Mech proposed that there was "probably far too many sub specific designations...in use" as most did not exhibit enough points of differentiation to be classified as a separate subspecies. The 24 subspecies were accepted by many authorities in 1981 and these were based on morphological or geographical differences or a unique history. However, in 1996, Ronald M. Nowak and Nick E. Federoff challenged Hall's 24 subspecies and proposed that based on detailed skull comparisons there had only been five: C. l. occidentalis, C. l. nubilus, C. l. arctos, C. l. baileyi and C. l. lycaon. Both classifications are accepted in North America.
History of classification
Eastern and red wolves
In 2000, a genetic study of canids from Algonquin Provincial Park indicated that C. l. lycaon was a separate species from C. lupus, more closely related to C. rufus. In a monograph prepared within the United States Fish and Wildlife Service (USF&WS), Chambers et al. (2012) reviewed many genetic studies and concluded that the eastern wolf and red wolf are separate species from the gray wolf, having originated in North America 150,000–300,000 years ago from the same line as coyotes. The Chambers review concluded that the subspecific status of C. l. arctos is doubtful, as Arctic wolf populations do not possess unique haplotypes. However, the Chambers review became controversial, forcing the USF&WS to commission a peer review of it, known as NCAES (2014). This peer review concluded that "virtually all taxonomic conclusions from Chambers et al. are accepted uncritically." Director of RESOLVE's Science Program, Steven Courtney, who was in charge of the peer review, had also noted that the Chambers conclusion that the eastern wolf should be listed outside the species limits of the gray wolf was based primarily on two non-recombining markers – those being mtDNA and sex chromosome – which the other panelists agreed unanimously "is insufficient to determine the existence of a species and specifically is completely insufficient for ruling out the alternative hypothesis that the pattern is explained by ancient (and recent) hybridization between C. lupus and C. latrans". Evolutionary biologist Dr. Robert Wayne of the UCLA Department of Ecology and Evolutionary Biology further elaborated that the Chambers review on the taxonomy of the eastern wolf not only suffered from insufficient sampling but that it was also biased in terms of attributing the presences of the gray wolf Y chromosomes in the modern day eastern wolves to two unfounded hypotheses: that C. lupus was historically absent in the eastern USA by C. lycaon followed with the suggestion of the former's invasion into the eastern third of North America and later introgressing into the latter's gene pool, and that the gray wolf Y chromosomes discovered in the Algonquin Provincial Park's eastern wolf samples originated from domestic dogs.
Two subsequent reviews of updated research based on the 2013 and 2014 reviews, one commissioned to the Wildlife Management Institute by the USFWS, and one journal review, concluded that historically there were four unique canid species in North America, gray wolf, eastern wolf, coyote, and dog, and that "the red wolf may be conspecific with the eastern wolf". This view consistent with the idea that the coyote and gray wolf did not historically range into the southeastern United States. These reviews and a 2015 genetics study, the most comprehensive to date, led the Committee on the Status of Endangered Wildlife in Canada (COSEWIC) in May, 2015 to change the designation of the eastern wolf back into a distinct species, Canis lycaon. However, the previous assertion that gray wolves did not occur in the eastern third of the United States is still heavily ill-founded by the newer genetic study's lead authors, such as Dr. Linda Y. Rutledge, who noted in the conclusions that "the recognition of the eastern wolf as a separate species does not exclude the possibility that a grey wolf × eastern wolf hybrid animal (previously identified as Canis lupus lycaon, boreal/Ontario-type), similar to a Great Lakes boreal wolf currently located in the Great Lakes states and across Manitoba, northern Ontario, and northern Quebec, historically inhabited the northeastern United States alongside eastern wolves, and there is some evidence to support the historical presence of both Canis types." The study also suggests that the coyote markers present in the Canis lupus populations that currently occupy the western Great Lakes states and western Ontario may have been historically circuited into the population by the eastern wolves since pure gray wolves in the wild rarely hybridize with coyotes whereas eastern wolves have a history of hybridizing with both species.
List of subspecies
Eurasia and Australia
Canis lupus lupus
|Linnaeus 1758||Generally a large subspecies with rusty ocherous or light grey fur.||Has the largest range among wolf subspecies and is the most common in Europe and Asia, ranging through Western Europe, Scandinavia, Caucasus, Russia, China, Mongolia, and the Himalayan Mountains. Habitat overlaps with Indian wolf in some regions of Turkey.||altaicus (Noack, 1911), argunensis (Dybowski, 1922), canus (Sélys Longchamps, 1839), communis (Dwigubski, 1804), deitanus (Cabrera, 1907), desertorum (Bogdanov, 1882), flavus (Kerr, 1792), fulvus (Sélys Longchamps, 1839), italicus (Altobello, 1921), kurjak (Bolkay, 1925), lycaon (Trouessart, 1910), major (Ogérien, 1863), minor (Ogerien, 1863), niger (Hermann, 1804), orientalis (Wagner, 1841), orientalis (Dybowski, 1922), signatus (Cabrera, 1907)|
Canis lupus albus
|Kerr 1792||A large, light-furred subspecies.||Northern tundra and forest zones in the European and Asian parts of Russia and Kamchatka. Outside Russia, its range includes the extreme north of Scandinavia||dybowskii (Domaniewski, 1926), kamtschaticus (Dybowski, 1922),
turuchanensis (Ognev, 1923)
Canis lupus arabs
|Pocock 1934||A small, "desert adapted" wolf that is around 66 cm tall and weighs, on average, about 18 kg. Its fur coat varies from short in the summer and long in the winter, possibly because of solar radiation.||Southern Israel, Southern and western Iraq, Oman, Yemen, Jordan, Saudi Arabia, and probably some parts of the Sinai Peninsula|
Canis lupus campestris
|Dwigubski 1804||A wolf of average size with short, coarse and sparse fur.||Northern Ukraine, southern Kazakhstan, Caucasus and Trans-Caucasus||bactrianus (Laptev, 1929), cubanenesis (Ognev, 1923), desertorum (Bogdanov, 1882)|
Canis lupus chanco
|Gray 1863||A small subspecies rarely exceeding 45 kg in weight. It is of a light, whitish-grey color, with an admixture of brownish tones on the upper part of the body||Central Asia from Turkestan, Tien Shan throughout Tibet to Mongolia, Northern China, Shensi, Sichuan, Yunnan, the Western Himalayas in Kashmir from Chitral to Lahul. Also occurs in the Korean peninsula||coreanus (Abe, 1923), dorogostaiskii (Skalon, 1936), ekloni (Przewalski, 1883), filchneri (Matschie, 1907), karanorensis (Matschie, 1907), laniger (Hodgson, 1847), niger (Sclater, 1874), tschiliensis (Matschie, 1907)|
Canis lupus dingo
|Meyer 1793||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. Fur color is mostly sandy to reddish brown, but can include tan patterns and be occasionally black, light brown, or white||Australia, ancient India, Indonesia, and New Guinea||antarticus (Kerr, 1792) [suppressed ICZN O451:1957], australasiae (Desmarest, 1820), australiae (Gray, 1826), dingoides (Matschie, 1915), macdonnellensis (Matschie, 1915), novaehollandiae (Voigt, 1831), papuensis (Ramsay, 1879), tenggerana (Kohlbrugge, 1896), harappensis (Prashad, 1936), hallstromi (Troughton, 1957)|
Canis lupus familiaris
|Linnaeus 1758||The dog is a divergent subspecies of the gray wolf and was derived from a now-extinct population of Late Pleistocene wolves. Through selective pressure and selective breeding, the dog has developed into hundreds of varied breeds, and shows more behavioral and morphological variation than any other land mammal.||Worldwide||
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), vertegus (Gmelin, 1792)
Canis lupus hattai
||||Kishida 1931||Similar in size and related to the gray wolves of North America.||Hokkaido and Sakhalin islands,:p42 the Kamchatka peninsula, and Iturup and Kunashir islands just to the east of Hokkaido in the Kuril archipelago.:p42||rex (Pocock, 1935)|
Canis lupus hodophilax
|Temminck 1839||Smaller in size compared to other gray wolves except for the Arabian wolf (Canis lupus arabs):p53||Japanese islands of Honshū, Shikoku, and Kyūshū (but not Hokkaido)||hodopylax (Temminck, 1844), japonicus (Nehring, 1885)|
Canis lupus pallipes
|Sykes 1831||A small wolf with pelage shorter than that of northern wolves, and with little to no underfur. Fur color ranges from greyish 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.||India, Pakistan, Iran, Turkey, Saudi Arabia and southern Israel|
|† Kenai Peninsula wolf
Canis lupus alces
|Goldman 1941||A very large subspecies similar to pambasileus.||Kenai Peninsula|
Canis lupus arctos
|Pocock 1935||A medium-sized, almost completely white subspecies.||Melville Island (Northwest Territories and Nunavut) and Ellesmere Island|
Canis lupus baileyi
|Nelson and Goldman 1929||Smallest of North America's gray wolves, with dark fur.||Northern Mexico, western Texas, southern New Mexico, and southeastern and central Arizona|
|† Newfoundland wolf
Canis lupus beothucus
|G. M. Allen and Barbour 1937||A medium-sized, white furred subspecies.||Newfoundland|
|† Bank's Island Tundra wolf
Canis lupus bernardi
|Anderson 1943||A large, slender subspecies with a narrow muzzle and large carnassials.||Limited to Banks and Victoria Islands in the arctic||banksianus (Anderson, 1943)|
|British Columbia wolf
Canis lupus columbianus
|Goldman 1941||Yukon, British Columbia, and Alberta|
|Vancouver Island wolf
Canis lupus crassodon
|Hall 1932||A medium-sized subspecies with grayish fur.||Vancouver Island, British Columbia|
|† Florida black wolf
Canis lupus floridanus
|Miller 1912||A jet black wolf that is described as being extremely similar to the red wolf in both size and weight. This subspecies became extinct in 1908.||Florida|
|† Cascade mountain wolf
Canis lupus fuscus
|Richardson 1839||A cinnamon colored wolf similar to columbianus and irremotus, but darker in color.||Cascade Range|
|† Gregory's wolf
Canis lupus gregoryi
|Goldman 1937||A medium-sized subspecies, though slender and tawny, its coat contains a mixture of various colors, including black, grey, white, and cinnamon.||In and around the lower Mississippi River basin||gigas (Townsend, 1850)|
Canis lupus griseoalbus
|Baird 1858||North Alberta, Saskatchewan, and Manitoba||knightii (Anderson, 1945)|
|Hudson Bay wolf
Canis lupus hudsonicus
|Goldman 1941||A light-colored subspecies similar to occidentalis, but smaller.||Northern Manitoba and the Northwest Territories|
|Northern Rocky Mountains wolf
Canis lupus irremotus
|Goldman 1937||A medium to large-sized subspecies with pale fur.||Northern Rocky Mountains|
Canis lupus labradorius
|Goldman 1937||A light-colored, medium-sized subspecies.||Labrador and northern Quebec; recent confirmed sightings on Newfoundland|
|Alexander Archipelago wolf
Canis lupus ligoni
|Goldman 1937||A medium-sized, dark colored subspecies.||Alexander Archipelago, Alaska|
|Eastern (timber) wolf
Canis lupus lycaon
|Schreber 1775||A small, dark-colored form.||Mainly occupies the area in and around Algonquin Provincial Park in Ontario, and also ventures into adjacent parts of Quebec, Canada. It also may be present in Michigan, Wisconsin, Minnesota and Manitoba||canadensis (de Blainville, 1843), ungavensis (Comeau, 1940)|
|Mackenzie River wolf
Canis lupus mackenzii
|Anderson 1943||A subspecies with variable fur and intermediate in size between occidentalis and manningi.||Northwest Territories|
|Baffin Island wolf
Canis lupus manningi
|Anderson 1943||The smallest gray wolf of the arctic, with white, buffy fur.||Baffin Island|
|† Mogollon mountain wolf
Canis lupus mogollonensis
|Goldman 1937||A small, dark-colored subspecies, intermediate in size between youngi and baileyi.||Arizona and New Mexico|
|†Texas (gray) wolf
Canis lupus monstrabilis
|Goldman 1937||Similar in size and color to C. lupus mogollonensis.||Texas and New Mexico||niger (Bartram, 1791)|
|Great Plains wolf
Canis lupus nubilus
|Say 1823||A light-furred, medium-sized subspecies.||Presently found in Ontario, Minnesota, Michigan, and Wisconsin. Single wolves have been reported in the Dakotas and as far south as Nebraska||variabilis (Wied-Neuwied, 1841) labradorius, irremotus, youngi (Goldman, 1937)|
Canis lupus occidentalis
|Richardson 1829||A very large, usually light-colored subspecies.||Alaska, Yukon, Northwest Territories, British Columbia, Alberta, Saskatchewan, and Northwestern United States||sticte (Richardson, 1829), ater (Richardson, 1829), alces (Goldman, 1941), ater (Richardson, 1829), mackenzii (Anderson, 1943 (1908), pambasileus (Elliot, 1905), sticte (Richardson, 1829), tundrarum (Miller, 1912)|
Canis lupus orion
|Pocock 1935||Greenland and Queen Elizabeth Islands|
Canis lupus pambasileus
|Elliot 1905||Larger in skull and tooth proportions than C. l. occidentalis, with fur that is black to white or a mix of both in color.||Alaska Interior and Yukon, save for the tundra region of the Arctic Coast.|
formerly Canis lupus rufus now Canis rufus
|Audubon and Bachman 1851||Has a brownish or cinnamon pelt, with grey and black shading on the back and tail. Generally intermediate in size between other American wolf subspecies and coyotes. Like other wolves, it has almond-shaped eyes, a broad muzzle and a wide nosepad, 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 wolves||Eastern North Carolina|
|Alaskan tundra wolf
Canis lupus tundrarum
|Miller 1912||A large, white-colored wolf closely resembling C. l. pambasileus, though lighter in color.||Barren grounds of the Arctic Coast region from near Point Barrow eastward toward Hudson Bay and probably northwards to the Arctic Archipelago|
|†Southern Rocky Mountains wolf
Canis lupus youngi
|Goldman 1937||A light-colored, medium-sized subspecies closely resembling C. l. nubilus, though larger, with more blackish-buff hairs on the back.||Southeastern Idaho, southwestern Wyoming, northeastern Nevada, Utah, western and central Colorado, northwestern Arizona and northwestern New Mexico|
Disputed subspecies and species
The Italian wolf was first recognised as a distinct subspecies Canis lupus italicus in 1921 by zoologist Giuseppe Altobello. Altobello's classification was later rejected by several authors, including Reginald Innes Pocock, who synonymised C. l. italicus with C. l. lupus. In 2002, the noted paleontologist R.M. Nowak reaffirmed the morphological distinctiveness of the Italian wolf and recommended the recognition of Canis lupus italicus. A number of DNA studies have found the Italian wolf to be genetically distinct. In 2004, the genetic distinction of the Italian wolf subspecies was supported by analysis which consistently assigned all the wolf genotypes of a sample in Italy to a single group. This population also showed a unique mitochondrial DNA control-region haplotype, the absence of private alleles and lower heterozygosity at microsatellite loci, as compared to other wolf populations. 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.
The Iberian wolf was first recognised as a distinct subspecies (Canis lupus signatus) in 1907 by zoologist Ángel Cabrera. The wolves of Iberian peninsula have morphologically distinct features from other Eurasian wolves and each are considered by their researchers to represent their own subspecies. The taxonomic reference Mammal Species of the World (2005) does not recognize Canis lupus signatus, however NCBI/Genbank does list it.
Himalayan wolf and Indian gray wolf
|Lineage and divergence times - southern Asian wolves|
The Himalayan wolf is formed by one haplotype that currently falls within the Tibetan wolf (Canis lupus chanco) subspecies, but based on mDNA sequencing has been proposed as a separate species Canis himalayensis. The Indian gray wolf is formed by two closely related haplotypes:116 that fall within the Indian wolf (Canis lupus pallipes) subspecies, but based on mDNA sequencing has been proposed as a separate species Canis indica. Neither proposal has been endorsed because they 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. Based on a fossil record estimate that the divergence time between the coyote and the wolf lineages occurred one million years ago and with an assumed wolf mutation rate, one study estimated the time of divergence of the Himalayan wolf and the Indian gray from the wolf/dog ancestor to be 800,000 years and 400,000 years ago respectively. Another study, which expressed some concerns with the earlier study, gave an estimate of 630,000 ago years and 270,000 ago years respectively. During Pleistocene glaciations these wolf lineages were isolated in refuges. In 2007, a study found that the Indian gray wolf was basil to all extant gray wolves and that the Himalayan wolf belonged to a different clade. In 2010, a study found that these two wolves formed a separate clade being 6 mutations distant from the extant gray wolf, which indicated distinct (i.e. different) lineages. In 2012, a limited genetic analysis of the scats of 2 Himalayan wolves from remote and widely separated areas reconfirmed their basal lineage.
The taxonomic reference Mammal Species of the World (2005) does not recognize Canis himalayensis nor Canis indica, however NCBI/Genbank lists a new subspecies Canis lupus himalayensis as separate from Canis lupus chanco, and a new subspecies Canis lupus indica as separate from Canis lupus pallipes.
In 2009, the Tibetan wolf was found to be genetically distinct enough to propose a separate species. In 2011, another genetic study found that the Tibetan wolf might be an archaic pedigree within the wolf subspecies, however the study defined Canis lupus laniger as the Tibetan wolf distinct from Canis lupus chanco the Mongolian wolf. In 2013, a major genetic study of dogs and wolves included the DNA sequences of 2 Tibetan wolves but then "excluded two aberrant modern wolf sequences from this analysis since their phylogenetic positioning suggests only a distant relationship to all extant gray wolves and their taxonomic classification as a member of Canis lupus or a separate sub-species is a matter of debate.":Sup The taxonomic reference Mammal Species of the World (2005) does not recognize Canis lupus laniger, however NCBI/Genbank lists it as the Tibetan wolf, and separately Canis lupus chanco  as the Mongolian wolf.
A study of the three coastal wolves indicated a close phylogenetic relationship across regions that are geographically and ecologically contiguous, and the study proposed that Canis lupus ligoni (Alexander Archipelago wolf), Canis lupus columbianus (British Columbia wolf), and Canis lupus crassodon (Vancouver Island wolf) should be recognized as a single subspecies of Canis lupus. They share the same habitat and prey species, and form one study's 6 identified North American ecotypes - a genetically and ecologically distinct population separated from other populations by their different type of habitat.
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. The taxonomic reference Mammal Species of the World (2005) does not recognize Canis lycaon, however NCBI/Genbank lists it.
The red wolf is an enigmatic taxon of which there are two proposals over its origin. One is that the red wolf was 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 admixture between coyotes and gray wolves, due to the gray wolf population being eliminated by humans. The taxonomic reference Mammal Species of the World (2005) does not recognize Canis rufus, however NCBI/Genbank lists it.
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