Subspecies of Canis lupus

Canis lupus species Temporal range: 0.7–0 Ma PreЄ Є O S D C P T J K Pg N ↓ Middle Pleistocene – Recent
Skulls of various gray wolf subspecies from North America
Conservation status
Least Concern  (IUCN 3.1)[1]
Scientific classification
Kingdom:	Animalia
Phylum:	Chordata
Class:	Mammalia
Order:	Carnivora
Family:	Canidae
Subfamily:	Caninae
Tribe:	Canini
Genus:	Canis
Species:	C. lupus
Binomial name
Canis lupus Linnaeus, 1758[2]
Subspecies
Numerous and disputed
Historical range of wild subspecies of C. lupus

Canis lupus has 40 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,^[1] 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.

In 2010, a study found that there were 75 different gray wolf mDNA haplotypes that include 23 in Europe, 30 in Asia, 18 in North America, 3 in both Europe and Asia, and 1 in both Europe and North America.^[3]

Taxonomy

Canis lupus was recorded by Carl Linnaeus in his publication Systema Naturae in 1758.^[4] The Latin classification translates into English as "dog wolf". The subspecies of Canis lupus are listed in Mammal Species of the World.^[5][6] The nominate subspecies is the Eurasian wolf (Canis lupus lupus),^[6] also known as the common wolf.^[7] As of 2005,^[5] 37 subspecies of C. lupus are recognised by MSW3, however the classification of several as either species or subspecies has recently been challenged.

Evolution

Ancestry

Feliforms and caniforms emerged within the super-family Carnivoramorpha 43 million YBP.^[8] The caniforms included the fox-like Leptocyon genus whose various species existed from 34 million YBP before branching 11.9 million YBP into vulpes (foxes) and canini (canines). The Eucyon genus diverged 6.2 million YBP towards Canis ferox, which diverged 5 million YBP towards Canis lepophagus, which diverged 3.5 million YBP towards the wolf-like canids.^[9]:174–5 If the geological attribution of the material is correct then the earliest identifiable C. lupus remains date 800,000 YBP^[9]:5 (Middle Pleistocene) with wolves similar to the living species^[9]:150 occurring at both the Olyor fauna (Siberia) and in the Cripple Creek Sump fauna (Alaska), which points to an origin of these wolves in Beringia.^[9]:181 In 2010, a study found that the diversity of the Canis group decreased by the end of the Early Pleistocene to Middle Pleistocene and was limited to the small wolves of the Canis mosbachensis–Canis variabilis group and the large hypercarnivorous Canis (Xenocyon) lycaonoides in Eurasia. The true gray wolves made their appearance at the end of the Middle Pleistocene at about 0.5–0.3 million YBP.^[10] Based on morphology from China, the Pliocene wolf Canis chihliensis may have been the ancestor for both Canis armbrusteri and Canis lupus before their migration into North America.^[9]:181 At the end of the most recent glacial retreat 30,000 YBP, warming melted the glacial barriers across northern Canada allowing arctic mammals to extend their range into mid-latitude North America, including elk, caribou, bison, and the gray wolf.^[11]

Distant Canis lupus ancestry - relationships and timeline (Tedford & Wang 2009)^[12]

Below is a cladogram illustrating the currently known phylogenetic relationships between extant wolf-like canids:^[13][14]

	Side-striped jackal Black-backed jackal African golden wolf Dog Grey wolf Coyote Ethiopian wolf Golden jackal Dhole African wild dog

Haplogroups

Canis divergence

Canis Golden jackal 1.9 million YBP[14] Coyote 1.1 million YBP[14] Canis lupus Himalayan wolf 630,000 YBP[15] Indian grey wolf 270,000 YBP[15] Haplogroup-2 Italian wolf (Apennine Peninsula only)[3] clade B (pockets of Eurasia only)[3] clade C (pockets of Eurasia only)[3] Dog 40,000 YBP[16] Haplogroup-1 Holarctic Gray wolf 40,000 YBP[16]

Divergence times and phylogenetic mDNA relationships. Haplogroup 2 basal to Haplogroup 1

In 2010, a study compared the mitochondrial DNA haplotypes of 24 ancient wolf specimens from western Europe dated between 44,000-1,200 YBP with those of modern gray wolves. The phylogenetic tree indicated that the haplotypes represented two haplogroups that were separated by 5 mutational steps. Haplogroup 1 formed a monophyletic clade (indicating a common ancestor). All other haplotypes were basal in the tree and these formed 2-3 smaller clades that were assigned to haplogroup 2. Haplogroups 1 and 2 could be found spread across Eurasia but only haplogroup 1 could be found in North America. The ancient wolf samples from western Europe all belonged to haplogroup 2, indicating haplogroup 2 predominance in this region for over 40,000 years before and after the Last Glacial Maximum. A comparison of current and past frequencies indicated that in Europe haplogroup 2 became outnumbered by haplogroup 1 over the past several thousand years but in North America haplogroup 2 became extinct and was replaced by haplogroup 1 after the Last Glacial Maximum.^[3] Access into North America was available between 20,000-11,000 years ago, after the Wisconsin glaciation had retreated but before the Bering land bridge became inundated by the sea.^[17] Therefore, haplogroup 1 was able to enter into North America during this period.

Analysis of stable isotopes, which offer conclusions about the diet and therefore the ecology of the extinct wolf populations, suggest that the Pleistocene wolves from haplogroup 2 found in Beringia and Belgium preyed mainly on Pleistocene megafauna,^[3][18][19] which became rare at the beginning of the Holocene 12,000 years ago.^[3][20] The Pleistocene Eurasian wolves have been found to be morphologically and genetically comparable to the Pleistocene eastern-Beringian wolves,^[21] with some of the ancient European and Beringian wolves sharing a common haplotype (a17),^[3][18] which makes ecological similarity likely.^[3] It has been proposed that Pleistocene wolves across northern Eurasia and northern North America represented a continuous and almost panmictic population that was genetically and probably also ecologically distinct from the wolves living in this area today.^[3][22] The specialized Pleistocene wolves, thus, did not contribute to the genetic diversity of modern wolves. Rather, modern wolf populations across the Holarctic are likely be the descendants of wolves from populations that came from more southern refuges.^[22] Haplogroup 2 did not become extinct in Europe, and if before the Last Glacial Maximum haplogroup 2 was exclusively associated with the wolf ecomorph specialized in preying on megafauna, it would mean that in Europe it was capable of adapting to changing prey.^[3]

These 2 haplogroups exclude the older-lineage Himalayan wolf and the Indian gray wolf.^[3][15][23]

Wolf population differences

Gray wolves have a wide, natural distribution across the Holarctic that includes many different habitats, which can vary from the high arctic to dense forests, open steppe and deserts. The genetic differences between different populations of gray wolves is tightly linked to the type of habitat in which they live.^[24] Differences in genetic markers among the Scandinavian wolf population has arisen in only just over a decade due to their small population size,^[24][25] which indicates that these differences are not dependent on a long time spent in isolation and that larger population patterns can evolve in just a few thousand years.^[24] These differences can also include fur color and density, and body size.^[24][26][27] The differences can also include behavior, as coastal wolves eat fish^[24][26] and tundra wolves migrate.^[24][27] These differences have been observed between two wolf populations that are living in close proximity. It has been shown that mountain wolves do not interbreed with nearby coastal wolves, and the Alps of France and Switzerland have been repopulated with wolves from the mountains of nearby Italy^[24][28] and from the far away mountains of Croatia^[24][29] rather than from the nearer lowlands, which indicates that distance is not the driving force in differences between the two ecomorphs.^[24]

Ecological factors including habitat type, climate, prey specialization and predatory competition will greatly influence their genetic population structure and cranio-dental plasticity.^{[24][27][30][31][32][33][34][35][36]} Therefore, within the Pleistocene gray wolf population the variations between local environments would have encouraged a range of wolf ecotypes that were genetically, morphologically and ecologically distinct from one another.^[36]

During the Last Glacial Maximum 20,000 YBP, the Pleistocene steppe stretched across northern and central Eurasia and through Beringia into North America. The Pleistocene wolves of Beringia, and perhaps those across the steppe, were adapted to this habitat. Their tooth and skull morphology indicates that they specialized in preying on now-extinct Pleistocene megafauna, and their tooth wear indicates that their behavior was different to modern wolves.^{[18][24][37][38]} This gray wolf ecomorph became extinct at the end of the glaciation, along with the horse and other species on which it depended, and was replaced by wolves from southern North America. This indicates that specialized wolf ecomorphs can become extinct when their environment changes even though the habitat may still support other wolves.^[24] Wolves went through a population bottleneck 20,000 YBP that coincides with the Last Glacial Maximum,^{[39][24][36][40]} which indicates that many wolf populations may have gone extinct at the same time as the Beringian wolves.^[24]

There are a small number of Canis remains that have been found at Goyet Cave, Belgium (36,500 YBP)^[41] Razboinichya Cave, Russia (33,500 YBP)^[42] Kostenki 8, Russia (33,500-26,500 YBP)^[43] Predmosti, Czech Republic (31,000 YBP)^[44] and Eliseevichi 1, Russia (17,000 YBP).^[45] Based on cranial morphometric study of the characteristics thought to be associated with the domestication process, these have been proposed as early Paleolithic dogs.^[43] These characteristics of shortened rostrum, tooth crowding, and absence or rotation of premolars have been documented in both ancient and modern wolves.^{[18][35][36][46][47][48]} Rather than representing early dogs, these specimens may represent "a morphologically distinct local, now extinct, population of wolves".^[36][49]

Domestic dog

Gray wolf divergence

wolf/dog ancestor New World clade Mexico North America/Hokkaido Old World clade Dog Old World wolves Asian highland Asian lowland Middle East Europe

Whole-genome phylogenetic tree - extant gray wolf populations[50]

The domestic dog (Canis lupus familiaris) is the most widely abundant large carnivore.^[50][51][52] Over the past million years, numerous wolf-like forms existed but their turnover has been high, and modern wolves are not the lineal ancestors of dogs.^{[39][50][52][53]} Although research had suggested that dogs and wolves were genetically very close relatives,^[54][55][56] later phylogenetic analysis strongly supported the hypothesis that dogs and wolves are reciprocally monophylic taxa that form two sister clades,^[39][54][57] suggesting that none of the modern wolf populations are related to the wolves that were first domesticated.^[39][57] Recent mitochondrial DNA analyses of ancient and modern gray wolf specimens supports a pattern of population reduction and turnover.^[3][18][52]

In 2016, a study investigated for the first time the population subdivisions, demography, and the relationships of gray wolves based on their whole-genome sequences. The study indicated that the dog was a divergent subspecies of the gray wolf and was derived from a now-extinct ghost population of Late Pleistocene wolves,^[39][50][52] and the dog and the dingo are not separate species.^[50] The genome-wide phylogenetic tree indicated a genetic divergence between New World and Old World wolves, which was then followed by a divergence between the dog and Old World wolves 27,000YBP^[16][50] - 29,000 YBP.^[50] The dog forms a sister taxon with Eurasian gray wolves but not North American wolves. The dog had considerable pre-ancestry after its divergence from the Old World wolves before it separated into distinct lineages that are nearly as distinct from one another as they are from wolves.^[50] The study suggested that previous datings based on the divergence between wolves and coyotes of one million years ago using fossils of what appeared to be coyote-like specimens may not reflect the ancestry of the modern forms.^{[14][16][39][50]}

The study indicated that the Mexican wolf was also a divergent form of gray wolf, suggesting that may have been part of an early invasion into North America.^[50][53] The Tibetan wolf was found to be the most highly-divergent of the Old World wolves, had suffered a historical population bottleneck and had only recently recolonized the Tibetan Plateau. Glaciation may have caused its habitat loss, genetic isolation then local adaption.^[50]

The study indicated that there has been extensive genetic admixture between domestic dogs and wolves, with up to 25% of the genome of Old World wolves showing signs of dog ancestry, possibly as the result of gene flow from dogs into wolves that were ancestral to all modern wolves. There was evidence of significant gene flow between the European wolves plus the Israeli wolf with the basenji and boxer, which suggests admixture between the lineages ancestral to these breeds and wolf populations.^[39][50] For the lowland Asian wolves: the Central Russian and East Russian wolves and all of the lowland Chinese wolves had significant gene flow with the Chinese indigenous dogs, the Tibetan Mastiff and the dingo. For the highland Asian wolves: The Tibetan wolves did not show significant admixture with dogs, however the Qinghai wolves had gene flow with the dingo and one of them had gene flow with the Chinese dogs. The New World wolves did not show any gene flow with the boxer, dingo or Chinese indigenous dogs but there was indication of gene flow between the Mexican wolf and the African basenji.^[50] All species within the Canis genus, the wolf-like canids, are phylogenetically closely related with 78 chromosomes and can potentially interbreed.^[56] There was indication of flow into the golden jackal from the population ancestral to all wolves and dogs (11.3%–13.6%) and much lower rates (up to 2.8%) from extant wolf populations.^[39][50]

The data indicated that all wolves shared similar population trajectories, followed by population decline that coincided with the expansion of modern humans worldwide and their technology for capturing large game.^[50][58] Late Pleistocene carnivores would have been social living in large prides, clans and packs in order to hunt the larger game available at that time, and these larger groups would have been more conspicuous targets for human persecutors.^[58] Large dogs accompanying the humans may have accelerated the rate of decline of carnivores that competed for game,^[50][59] therefore humans expanded across Eurasia, encountered wolves, domesticated some and possibly caused the decline of others.^[50]

See further: Wolves compared with dogs

History of classification

Old World gray wolf subspecies

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.^[60] In 2003, Nowak also recognized the distinctiveness of C. l. arabs, C. l. hattai, C. l. hodophilax and C. l. lupaster.^[61] Furthermore, genetic studies on gray wolves in Italy revealed that, unlike several European gray wolf populations, Italian wolves do not share haplotypes with either other gray wolves or domestic dogs, and are morphologically distinct enough to be classed as a separate subspecies; C. l. italicus.^[62][63][64]

Rueness et al. (2014) showed that wolves in the Caucasus Mountains of the putative Caucasian subspecies, C. l. cubanensis, are not genetically distinct enough to be considered a subspecies, but may represent a local ecomorph of C. l. lupus.^[65]

In 2015, a study of mitochondrial and nuclear genomes on putative wolves and golden jackals in Africa revealed that they both represented differing ecomorphs of the same species, Canis anthus, which is distinct from the gray wolf.^[14]

New World gray wolf subspecies

C. lupus colonized North America during the late Rancholabrean era through the Bering land bridge in at least three separate invasions, with each wave being represented by one or more different Eurasian gray wolf clades.^[66] Among the first to enter was a broad-skulled, hypercarnivorous ecomorph which never expanded its range below the Wisconsin ice sheet, likely because of competitive exclusion by C. dirus populations in the south, with both dying out during the Quaternary extinction event without leaving any modern descendants.^[18] The first gray wolves to permanently enter North America were the ancestors of C. l. baileyi, though these were followed and displaced by C. l. nubilus and pushed southwards. C. l. nubilus was in turn displaced from its northern range by C. l. occidentalis, likely during the Holocene, a process which may have continued into historic times.^[66]

Skull of a European wolf

Skull of a Canadian wolf

In 1944, American zoologist Edward Goldman recognized as many as 23 subspecies in North America, based on morphology alone.^[67] In 1995, mammalogist Ronald Nowak disputed these classifications, based on his comparison of numerous wolf skulls from throughout the continent. He concluded that there are only five North American subspecies: C. l. occidentalis, C. l. nubilus, C. l. arctos, C. l. baileyi and C. l. lycaon. Wilson et al. (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.^[68]

The taxonomy of wolves in the coastal rainforests of British Columbia and southeastern Alaska has also followed a variable path, regarding the putative Vancouver Island wolf (C. l. crassodon) and Alexander Archipelago wolf (C. l. ligoni), respectively. Based on skull morphometrics, C. l. ligoni was recognized by Goldman (1944), Hall (1981) and Pedersen (1982) as a distinct population possibly warranting subspecific classification; however, Nowak (1996) considered it to be an isolated population of C. l. nubilis.^[69] From 2005 to 2014 several studies and the NCAES (2014) peer review have found the pacific coastal wolves to have a phenotypically distinct genotype.^{[70][71][72][73][74]} The putative British Columbian Raincoast Island wolves became the central discussion in a 2015 National Geographic documentary that introduced them as sea wolves due in part to the morphologically distinct skeletal structures of these wolves compared to inland North American wolves, which gives them the ability to swim longer distances between the islands than those on the mainland continent.^[75] The Raincoast wolves are also known to have a unique dietary preference for fish, orcas, mollusks, seals, and various small prey endemic to the islands, as opposed to their inland cousins who subsist on medium-sized ungulates. In the 2014 NCAES peer review, Dr. Robert Wayne suggested that centuries of adaptation to these remote islands coupled with the unique dietary habits may have played a critical role in uniforming the modern day Raincoast wolves' genetic distinction from Vancouver island wolves and inland wolves, thus warranting a new subspecific classification distinct from Nowak's previous C. l. nubilus classification.

Eastern and red wolves

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.^[66] However, the Chambers review became controversial, forcing the USF&WS to commission a peer review of it, known as NCAES (2014).^[76] 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".^[77][78] This view consistent with the idea that the coyote and gray wolf did not historically range into the southeastern United States.^[77] These reviews and a 2015 genetics study, the most comprehensive to date,^[79] 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

Subspecies recognized by MSW3 as of 2005^[80] and divided into Old World and New World:^[50]

Old World

Subspecies	Image	Authority	Description	Range	Synonyms
Eurasian wolf Canis lupus lupus (nominate subspecies)		Linnaeus 1758[81]	Generally a large subspecies with rusty ocherous or light grey fur.[82]	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)[83]
Tundra wolf Canis lupus albus		Kerr 1792[84]	A large, light-furred subspecies.[85]	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[85]	dybowskii (Domaniewski, 1926), kamtschaticus (Dybowski, 1922), turuchanensis (Ognev, 1923)[86]
Arabian wolf Canis lupus arabs		Pocock 1934[87]	A small, "desert adapted" wolf that is around 66 cm tall and weighs, on average, about 18 kg.[88] Its fur coat varies from short in the summer and long in the winter, possibly because of solar radiation.[89]	Southern Israel, Southern and western Iraq, Oman, Yemen, Jordan, Saudi Arabia, and probably some parts of the Sinai Peninsula
Steppe wolf Canis lupus campestris		Dwigubski 1804	A wolf of average size with short, coarse and sparse fur.[90]	Northern Ukraine, southern Kazakhstan, Caucasus and Trans-Caucasus[90]	bactrianus (Laptev, 1929), cubanenesis (Ognev, 1923), desertorum (Bogdanov, 1882)[91]
Tibetan wolf 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[92]	Central Asia from Turkestan, Tien Shan throughout Tibet to Mongolia, Northern China, Shensi, Sichuan, Yunnan, the Western Himalayas in Kashmir from Chitral to Lahul.[93] Also occurs in the Korean peninsula[94]	coreanus (Abe, 1923), dorogostaiskii (Skalon, 1936), ekloni (Przewalski, 1883), filchneri (Matschie, 1907), karanorensis (Matschie, 1907), laniger (Hodgson, 1847), niger (Sclater, 1874), tschiliensis (Matschie, 1907)[95]
Dingo 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.[96] Fur color is mostly sandy to reddish brown, but can include tan patterns and be occasionally black, light brown, or white[97]	Australia, ancient India, Indonesia, and New Guinea	antarcticus (Kerr, 1792), 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)[98]
Domestic dog Canis lupus familiaris		Linnaeus 1758	The dog was a divergent subspecies of the gray wolf and was derived from a now-extinct population of Late Pleistocene wolves.[39][50][52] 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.[99]	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)[100]
†Hokkaidō wolf Canis lupus hattai		Kishida 1931	Similar in size and related to the gray wolves of North America.[101]	Hokkaido and Sakhalin islands,[102][103]^:p42 the Kamchatka peninsula, and Iturup and Kunashir islands just to the east of Hokkaido in the Kuril archipelago.[103]^:p42	rex (Pocock, 1935)[104]
†Japanese wolf Canis lupus hodophilax		Temminck 1839	Smaller in size compared to other gray wolves except for the Arabian wolf (Canis lupus arabs)[103]^:p53	Japanese islands of Honshū, Shikoku, and Kyūshū (but not Hokkaido)[105][106]	hodopylax (Temminck, 1844), japonicus (Nehring, 1885)[107]
Indian wolf Canis lupus pallipes		Sykes 1831	A small wolf with pelage shorter than that of northern wolves, and with little to no underfur.[108] 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.[109]	India, Pakistan, Iran, Turkey, Saudi Arabia and southern Israel

New World

Subspecies	Image	Authority	Description	Range	Synonyms
† Kenai Peninsula wolf Canis lupus alces		Goldman 1941[110]	A very large subspecies similar to pambasileus.[111]	Kenai Peninsula
Arctic wolf Canis lupus arctos		Pocock 1935[112]	A medium-sized, almost completely white subspecies.[113]	Mellville Island and Ellesmere Island
Mexican wolf Canis lupus baileyi		Nelson and Goldman 1929[114]	Smallest of North America's gray wolves, with dark fur.[115]	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.[116]	Newfoundland
† Bank's Island Tundra wolf Canis lupus bernardi		Anderson 1943	A large, slender subspecies with a narrow muzzle and large carnassials.[117]	Limited to Banks and Victoria Islands in the arctic	banksianus (Anderson, 1943)[118]
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.[119]	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.[120] This subspecies became extinct in 1908.[121]	Florida
† Cascade mountain wolf Canis lupus fuscus		Richardson 1839	A cinnamon colored wolf similar to columbianus and irremotus, but darker in color.[122]	Cascade Range
† Gregory's wolf Canis lupus gregoryi		Goldman 1937[123]	A medium-sized subspecies, though slender and tawny, its coat contains a mixture of various colors, including black, grey, white, and cinnamon.[123]	In and around the lower Mississippi River basin	gigas (Townsend, 1850)[124]
†Manitoba wolf Canis lupus griseoalbus		Baird 1858	North Alberta, Saskatchewan, and Manitoba	knightii (Anderson, 1945)[125]
Hudson Bay wolf Canis lupus hudsonicus		Goldman 1941	A light-colored subspecies similar to occidentalis, but smaller.[126]	Northern Manitoba and the Northwest Territories
Northern Rocky Mountains wolf Canis lupus irremotus		Goldman 1937[123][127]	A medium to large-sized subspecies with pale fur.[128]	Northern Rocky Mountains
Labrador wolf Canis lupus labradorius		Goldman 1937[123]	A light-colored, medium-sized subspecies.[129]	Labrador and northern Quebec; recent confirmed sightings on Newfoundland[130][131]
Alexander Archipelago wolf Canis lupus ligoni		Goldman 1937[123]	A medium-sized, dark colored subspecies.[132]	Alexander Archipelago
Eastern (timber) wolf Canis lupus lycaon		Schreber 1775	A small, dark-colored form.[133]	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 Minnesota and Manitoba	canadensis (de Blainville, 1843), ungavensis (Comeau, 1940)[134]
Mackenzie River wolf Canis lupus mackenzii		Anderson 1943	A subspecies with variable fur and intermediate in size between occidentalis and manningi.[135]	Northwest Territories
Baffin Island wolf Canis lupus manningi		Anderson 1943	The smallest gray wolf of the arctic, with white, buffy fur.[136]	Baffin Island
† Mogollon mountain wolf Canis lupus mogollonensis		Goldman 1937[123]	A small, dark-colored subspecies, intermediate in size between youngi and baileyi.[137]	Arizona and New Mexico
†Texas (gray)wolf Canis lupus monstrabilis		Goldman 1937[123]	Similar in size and color to C. lupus mogollonensis.[138]	Texas and New Mexico	niger (Bartram, 1791)[139]
Great Plains wolf Canis lupus nubilus		Say 1823	A light-furred, medium-sized subspecies.[140]	Minnesota, Michigan, and Wisconsin. Single wolves have been reported in the Dakotas and as far south as Nebraska	variabilis (Wied-Neuwied, 1841)[141] labradorius, irremotus, youngi (Goldman, 1937)
Northwestern wolf Canis lupus occidentalis		Richardson 1829	A very large, usually light-colored subspecies.[142]	Western Canada	sticte (Richardson, 1829), ater (Richardson, 1829),[143] alces (Goldman, 1941), ater (Richardson, 1829), mackenzii (Anderson, 1943 (1908), pambasileus (Elliot, 1905), sticte (Richardson, 1829), tundrarum (Miller, 1912)
Greenland wolf Canis lupus orion		Pocock 1935		Greenland
Yukon wolf 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.[144]	Alaska Interior and Yukon, save for the tundra region of the Arctic Coast.[145]
Red wolf 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[146]	Eastern North Carolina[147]
Alaskan tundra wolf Canis lupus tundrarum		Miller 1912	A large, white-colored wolf closely resembling C. l. pambasileus, though lighter in color.[148]	Barren grounds of the Arctic Coast region from near Point Barrow eastward toward Hudson Bay and probably northwards to the Arctic Archipelago[149]
†Southern Rocky Mountains wolf Canis lupus youngi		Goldman 1937[123]	A light-colored, medium-sized subspecies closely resembling C. l. nubilus, though larger, with more blackish-buff hairs on the back.[150]	Southeastern Idaho, southwestern Wyoming, northeastern Nevada, Utah, western and central Colorado, northwestern Arizona and northwestern New Mexico

Disputed subspecies and species

Giuseppe Altobello's 1925 comparative illustration of the skulls and dentition of C. l. lupus (a) and C. l. italicus (b). The distinct status of the latter is currently unrecognised by MSW3.

Apennine wolf and Iberian wolf

Two proposed subspecies not mentioned in the list above are the Italian wolf (Canis lupus italicus) and the Iberian wolf (Canis lupus signatus). The wolves of the Apennine (Italian) and Iberian peninsulas have morphologically distinct features from other Eurasian wolves and each are considered by their researchers to represent their own subspecies.^[151][152] 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.^[153] 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.^[3] The taxonomic reference Mammal Species of the World (2005) does not recognize Canis lupus signatus nor Canis lupus italicus, however NCBI/Genbank does list Canis lupus signatus^[154] and publishes research papers under that name Canis lupus italicus.^[155]

Himalayan wolf and Indian gray wolf

Divergence times

Golden jackal 1.9 million YBP[14] African golden wolf 1.3 million YBP[14] Coyote 1.1 million YBP[14] Himalayan wolf 630,000 YBP[15] Indian gray wolf 270,000 YBP[15] Eurasian gray wolf Dog

Lineage and divergence times - southern Asian wolves

The Himalayan wolf is formed by one haplotype^[15] that currently falls within the Tibetan wolf (Canis lupus chanco) subspecies, but based on mDNA sequencing has be proposed as a separate species Canis himalayensis.^[15][23] The Indian gray wolf is formed by two closely related haplotypes^[15]:116 that fall within the Indian wolf (Canis lupus pallipes) subspecies, but based on mDNA sequencing has be proposed as a separate species Canis indica.^[15][23] Neither proposal has been endorsed because they relied on a limited number of museum and zoo samples that may not have be representative of the wild population, and a call for further fieldwork has been made.^[156] 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.^[23] Another study, which expressed some concerns with the earlier study, gave an estimate of 630,000 ago years and 270,000 ago years respectively.^[15] During Pleistocene glaciations these wolf lineages were isolated in refuges.^[23] 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.^[18] 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.^[3] In 2012, a limited genetic analysis of the scats of 2 Himalayan wolves from remote and widely separated areas reconfirmed their basal lineage.^[157]

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^[158] as separate from Canis lupus chanco, and a new subspecies Canis lupus indica^[159] as separate from Canis lupus pallipes.

Red wolf

The red wolf is listed above, despite being considered a distinct species by many other authorities, including the US Fish and Wildlife Service, the main government authority of red wolves.^[160] Some studies have concluded that red wolves, along with eastern wolves, evolved in North America between 150,000 and 300,000 years ago independent from gray wolves in Eurasia.^[161] The taxonomic reference Mammal Species of the World (2005) does not recognize Canis rufus, however NCBI/Genbank lists it.^[162]

Eastern wolf

In 2011, the US government recognized the Eastern wolf Canis lupus lycaon as a separate species Canis lycaon.^[163] In 2015, the Canadian government recognized the Eastern wolf as Canis species cf lycaon (Canis species believed to be lycaon).^[164][165] The taxonomic reference Mammal Species of the World (2005) does not recognize Canis lycaon, however NCBI/Genbank lists it.^[166]

African golden wolf

As of 2005, the African wolf is listed as a subspecies of the golden jackal by the third and current edition of Mammal Species of the World.^[167] In the early 2010s research disputed this, as mtDNA of wolf-like canids in various locations in northern, western, and eastern Africa more closely resemble those of gray wolves than golden jackals.^[168][169] In 2015, a series of analyses on the species' mitochondrial DNA and nuclear genome indicated that it was distinct from both the golden jackal and the grey wolf, and more closely related to grey wolves and coyotes. ^[170]

Tibetan wolf

In 2009, the Tibetan wolf was found to be genetically distinct enough to propose a separate species.^[171][172] 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.^[173] 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."^[174]:Sup The taxonomic reference Mammal Species of the World (2005) does not recognize Canis lupus laniger, however NCBI/Genbank lists it^[175] as the Tibetan wolf, and separately Canis lupus chanco ^[176] as the Mongolian wolf.

See also

• California Wolf Center
• List of gray wolf populations by country

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External links

• Canis lupus on the ITIS (Integrated Taxonomic Information System)
• Citations for Mammal Species of the World, as a PDF
• Ancient origin and evolution of the Indian wolf including discussion on naming.
• Wolf subspecies list with photos and maps
• Animal Corner, UK list of wolves