Subspecies of ''Canis lupus''

Canis lupus subspecies
Temporal range: Middle Pleistocene – present (700,000-0 YBP)
Skulls of various gray wolf subspecies from North America
Scientific classification
Kingdom:Animalia
Phylum:Chordata
Class:Mammalia
Order:Carnivora
Family:Canidae
Tribe:Canini
Genus:Canis
Species: C. lupus
Binomial name
Canis lupus
Subspecies

Numerous and disputed

Present range of wild subspecies of C. lupus

Canis lupus has 38 subspecies listed in the taxonomic authority Mammal Species of the World, 2005 edition. These subspecies were named over the past 250 years, and since naming a number of them have gone extinct. 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,[2] 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.

Taxonomy

The species Canis lupus was first recorded by Carl Linnaeus in his publication Systema Naturae in 1758,[1] with the Latin classification translating into the English words "dog wolf".

A subspecies is the taxonomic rank below species.[3] 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.[4] The thirty-eight subspecies of Canis lupus are listed in Mammal Species of the World (third edition) that was published in 2005,[5][6] and in the Catalogue of Life.[7] The nominate subspecies is the Eurasian wolf (Canis lupus lupus),[6] also known as the common wolf.[8] The subspecies includes the domestic dog, dingo, eastern wolf and red wolf.[5] However, the classification of several as either species or subspecies has recently been challenged.

List of extant subspecies

The living subspecies recognized by MSW3 as of 2005[9] and divided into Old World and New World:[10]

Eurasia and Australia

For Eurasia, in 1995 mammalogist Robert Nowak recognized five subspecies based on skull morphology, these being: C. l. lupus, C. l. albus, C. l. pallipes, C. l. cubanensis and C. l. communis.[11] In 2003, Nowak also recognized the distinctiveness of C. l. arabs, C. l. hattai, and C. l. hodophilax.[12] In 2005, MSW3 included C. l. filchneri.[9] In 2003, two forms were distinguished in southern China and Inner Mongolia as being separate from C. l. chanco and C. l. filchneri and have yet to be named.[13][14]

Subspecies of Canis lupus
Subspecies Image Authority Description Range Synonyms
C. l. albus
Tundra wolf		 Kerr, 1792[15] A large, light-furred subspecies.[16] 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[16] dybowskii Domaniewski, 1926, kamtschaticus Dybowski, 1922, turuchanensis Ognev, 1923[17]
C. l. arabs
Arabian wolf		 Pocock, 1934[18] A small, "desert adapted" wolf that is around 66 cm tall and weighs, on average, about 18 kg.[19] Its fur coat varies from short in the summer and long in the winter, possibly because of solar radiation.[20] Southern Israel, Southern and western Iraq, Oman, Yemen, Jordan, Saudi Arabia, and probably some parts of the Sinai Peninsula
C. l. campestris
Steppe wolf		 Dwigubski, 1804 A wolf of average size with short, coarse and sparse fur.[21] Northern Ukraine, southern Kazakhstan, Caucasus and Trans-Caucasus[21] bactrianus Laptev, 1929, cubanenesis Ognev, 1923, desertorum Bogdanov, 1882[22]
C. l. chanco
Mongolian wolf		 Gray, 1863[23] The fur fulvous, 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] Mongolia,[24] northern and central China,[13][14] Korea,[25] and the Ussuri region of Russia.[26] chanco Gray, 1863, coreanus Abe, 1923, dorogostaiskii Skalon, 1936, karanorensis Matschie, 1907, niger Sclater, 1874, tschiliensis Matschie, 1907
C. l. dingo
–Dingo and New Guinea Singing Dog		 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.[27] Fur color is mostly sandy to reddish brown, but can include tan patterns and be occasionally black, light brown or white.[28] Australia 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, hallstromi Troughton, 1957, harappensis Prashad, 1936[29]
C. l. familiaris
Domestic dog		 Linnaeus, 1758 The dog is a divergent subspecies of the gray wolf and was derived from a now-extinct population of Late Pleistocene wolves.[10][30][31] 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.[32] 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[33]

Proposed as the species Canis familiaris, but debated.[34]

C. l. filchneri
Tibetan wolf		 Matschie, 1907[35] Long sharp face, elevated brows, broad head, large pointed ears, thick woolly pilage, and very full brush of medial length. Above, dull earthy-brown; below, with the entire face and limbs yellowish-white.[36] China in the regions of Gansu, Qinghai, and Xichang (Tibet),[13][14] and northern India in the Ladakh region of Jammu and Kashmir[37] and the Lahoul region of Himachal Pradesh.[38] filchneri Matschie, 1907, laniger Hodgson, 1847
C. l. lupus
Eurasian wolf
(nominate subspecies)		 Linnaeus, 1758[39] Generally a large subspecies with rusty ocherous or light gray fur.[40] 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[41]
C. l. pallipes
Indian wolf		 Sykes, 1831 A small wolf with pelage shorter than that of northern wolves, and with little to no underfur.[42] 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.[43] India, Pakistan, Iran, Turkey, Saudi Arabia and southern Israel

North America

North American gray wolf subspecies distribution according to Goldman (1944). These subspecies are included in MSW3 2005.

For North America, in 1944 the zoologist Edward Goldman recognized as many as 23 subspecies based on morphology.[44] In 1959, E. Raymond Hall proposed that there had been 24 subspecies of lupus in North America.[45] 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.[46] The 24 subspecies were accepted by many authorities in 1981 and these were based on morphological or geographical differences or a unique history.[47] In 1995, the American mammologist 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 gray wolf. These include a large toothed arctic wolf named C. l. arctos, a large wolf from Alaska and western Canada named C. l. occidentalis, a small wolf from southeast Canada named C. l. lycaon, a small wolf from the southwest named C. l. baileyi, and a moderate-sized wolf that was originally found from Texas to Hudson Bay and from Oregon to New Foundland named C. l. nubilus.[48][49] This proposal was not reflected in the taxonomic classification of Canis lupus subspecies in Mammal Species of the World (2005).[9]

Subspecies of Canis lupus
Subspecies Image Authority Description Range Synonyms
C. l. arctos
Arctic wolf		 Pocock, 1935[50] A medium-sized, almost completely white subspecies.[51] Melville Island (Northwest Territories and Nunavut), Ellesmere Island and Alaska
C. l. baileyi
Mexican wolf		 Nelson and Goldman, 1929[52] Smallest of North America's gray wolves, with dark fur.[53] Presently found in southeastern New Mexico and southeastern Arizona
C. l. columbianus
British Columbia wolf		 Goldman, 1941 Yukon, British Columbia, and Alberta
C. l. crassodon
Vancouver Island wolf		 Hall, 1932 A medium-sized subspecies with grayish fur.[54] Vancouver Island, British Columbia
C. l. gregoryi
Gregory's wolf
but refer Synonyms		 Goldman, 1937[55] A medium-sized subspecies, though slender and tawny; its coat contains a mixture of various colors, including black, gray, white and cinnamon.[55] In and around the lower Mississippi River basin Proposed as a subspecies of Canis rufus[56] but debated
C. l. hudsonicus
Hudson Bay wolf		 Goldman, 1941 A light-colored subspecies similar to occidentalis, but smaller.[57] Northern Manitoba and the Northwest Territories
C. l. irremotus
Northern Rocky Mountain wolf		 Goldman, 1937[55][58] A medium to large-sized subspecies with pale fur.[59] Northern Rocky Mountains
C. l. labradorius
Labrador wolf		 Goldman, 1937[55] A light-colored, medium-sized subspecies.[60] Labrador and northern Quebec; recent confirmed sightings on Newfoundland[61][62]
C. l. ligoni
Alexander Archipelago wolf		 Goldman, 1937[55] A medium-sized, dark-colored subspecies.[63] Alexander Archipelago, Alaska
C. l. lycaon
Eastern wolf
but refer Synonyms		 Schreber, 1775 A small, dark-colored form.[64] 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[65]

Proposed as the species Canis lycaon[56] but debated
C. l. mackenzii
Mackenzie River wolf		 Anderson, 1943 A subspecies with variable fur and intermediate in size between occidentalis and manningi.[66] Northwest Territories
C. l. manningi
Baffin Island wolf		 Anderson, 1943 The smallest gray wolf of the Arctic, with white, buffy fur.[67] Baffin Island
C. l. occidentalis
Northwestern wolf		 Richardson, 1829 A very large, usually light-colored subspecies.[68] Alaska, Yukon, Northwest Territories, British Columbia, Alberta, Saskatchewan, and Northwestern United States ater Richardson, 1829, sticte Richardson, 1829[69]
C. l. orion
Greenland wolf		 Pocock, 1935 Greenland and Queen Elizabeth Islands[70]
C. l. pambasileus
Yukon wolf		 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.[71] Alaska Interior and Yukon, save for the tundra region of the Arctic Coast.[72]
C. l. rufus
Red wolf
but refer Synonyms		 Audubon and Bachman, 1851 Has a brownish or cinnamon pelt, with gray 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 gray wolves.[73] Eastern North Carolina[74] Proposed as the species Canis rufus[56] but debated
C. l. tundrarum
Alaskan tundra wolf		 Miller, 1912 A large, white-colored wolf closely resembling C. l. pambasileus, though lighter in color.[75] Barren Grounds of the Arctic Coast region from near Point Barrow eastward toward Hudson Bay and probably northwards to the Arctic Archipelago[76]

List of historically extinct subspecies

The subspecies recognized by MSW3 as of 2005 and which have gone extinct over the past 150 years:[9]

Extinct subspecies of Canis lupus
Subspecies Image Authority Description Range Synonyms
† C. l. alces
Kenai Peninsula wolf		 Goldman 1941[77] One of the largest subspecies, similar to pambasileus.Its fur color is unknown.[78] Kenai Peninsula
† C. l. beothucus
Newfoundland wolf		 G. M. Allen and Barbour 1937 A medium-sized, white-furred subspecies.[79] Newfoundland
† C. l. bernardi
Bernard's wolf		 Anderson 1943 A large, slender subspecies with a narrow muzzle and large carnassials.[80] Limited to Banks and Victoria Islands in the Arctic banksianus Anderson, 1943[81]
† C. l. floridanus
Florida black wolf		 Miller 1912 A jet black wolf that is described as being extremely similar to the red wolf in both size and weight.[82] This subspecies became extinct in 1908.[83] Florida
† C. l. fuscus
Cascade Mountains wolf		 Richardson 1839 A cinnamon-colored wolf similar to columbianus and irremotus, but darker in color.[84] Cascade Range gigas Townsend, 1850[85]
† C. l. griseoalbus
Manitoba wolf		 Baird 1858 North Alberta, Saskatchewan, and Manitoba knightii Anderson, 1945[86]
† C. l. hattai
Hokkaidō wolf		 Kishida 1931 Similar in size and related to the gray wolves of North America.[87] Hokkaido and Sakhalin islands,[88][89]:p42 the Kamchatka peninsula, and Iturup and Kunashir islands just to the east of Hokkaido in the Kuril archipelago.[89]:p42 rex Pocock, 1935[90]
† C. l. hodophilax
Japanese wolf		 Temminck 1839 Smaller in size compared to other gray wolves except for the Arabian wolf (Canis lupus arabs).[89]:p53 Japanese islands of Honshū, Shikoku, and Kyūshū (but not Hokkaido)[91][92] japonicus Nehring, 1885[93]
† C. l. mogollonensis
Mogollon mountain wolf		 Goldman 1937[55] A small, dark-colored subspecies, intermediate in size between youngi and baileyi.[94] Arizona and New Mexico
† C. l. monstrabilis
Texas wolf		 Goldman 1937[55] Similar in size and color to C. lupus mogollonensis.[95] Texas and New Mexico niger Bartram, 1791[96]
† C. l. nubilus
Great Plains wolf		 Say 1823 A light-furred, medium-sized subspecies.[97] Throughout the Great Plains from southern Manitoba and Saskatchewan southward to northern Texas.[98] variabilis Wied-Neuwied, 1841[99]
† C. l. youngi
Southern Rocky Mountain wolf		 Goldman 1937[55] A light-colored, medium-sized subspecies closely resembling C. l. nubilus, though larger, with more blackish-buff hairs on the back.[100] Southeastern Idaho, southwestern Wyoming, northeastern Nevada, Utah, western and central Colorado, northwestern Arizona and northwestern New Mexico

Disputed subspecies and species

Skull of a European wolf
Skull of a Canadian wolf

Eurasia

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

The Apennine wolf (Italian wolf) was first recognised as a distinct subspecies Canis lupus italicus in 1921 by zoologist Giuseppe Altobello.[101] Altobello's classification was later rejected by several authors, including Reginald Innes Pocock, who synonymised C. l. italicus with C. l. lupus.[102] In 2002, the noted paleontologist R.M. Nowak reaffirmed the morphological distinctiveness of the Italian wolf and recommended the recognition of Canis lupus italicus.[102] A number of DNA studies have found the Italian wolf to be genetically distinct.[103][104] 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.[105] 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.[106]

The taxonomic reference Mammal Species of the World (2005) does not recognize Canis lupus italicus, however NCBI/Genbank publishes research papers under that name.[107]

Iberian wolf

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.[108][109] The taxonomic reference Mammal Species of the World (2005) does not recognize Canis lupus signatus, however NCBI/Genbank does list it.[110]

Himalayan wolf

Divergence times

 

 

 

Golden jackal 1.9 million YBP[111]

 
 

 

African golden wolf 1.3 million YBP[111]

 
 

 

Coyote 1.1 million YBP[111]

 
 

 

Himalayan wolf 630,000 YBP[112]

 
 

 

Indian gray wolf 270,000 YBP[112]

 
 

 

Eurasian gray wolf 80,000 YBP (extant)[10]

 
 

Dog 40,000 YBP[113]

 
 
 
 
 
 
 
 
Lineage and divergence times based on DNA

The Himalayan wolf is a proposed clade within the Tibetan wolf (Canis lupus filchneri) that is distinguished by its mitochondrial DNA, which is basal to all other wolves including other Tibetan wolves. The taxonomic status of this wolf clade is disputed, with the separate species Canis himalayensis being proposed based on two limited DNA studies.[112][114] The proposal has not 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.[115] In 2017, a study of mitochondrial DNA, X-chromosome (maternal lineage) markers and Y-chromosome (male lineage) markers found that the Himalayan wolf is genetically basal to the holarctic grey wolf.[116] The taxonomic reference Mammal Species of the World (2005) does not recognize Canis himalayensis, however NCBI/Genbank lists a new subspecies Canis lupus himalayensis.[117]

Indian gray wolf

The Indian gray 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 proposed Himalayan wolf. The taxonomic status of this wolf clade is disputed, with the separate species Canis indica being proposed based on two limited DNA studies.[112][114] The proposal has not 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.[115] The taxonomic reference Mammal Species of the World (2005) does not recognize Canis indica, however NCBI/Genbank lists a new subspecies Canis lupus indica.[118]

North America

Coastal wolves

A study of the three coastal wolves indicates 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.[119] 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.[120][121]

Eastern wolf

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.[122] The taxonomic reference Mammal Species of the World (2005) does not recognize Canis lycaon, however NCBI/Genbank lists it.[123]

Red wolf

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.[122] The taxonomic reference Mammal Species of the World (2005) does not recognize Canis rufus, however NCBI/Genbank lists it.[124]

See also

References

  1. 1 2 Linnæus, Carl (1758). Systema naturæ per regna tria naturæ, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I (in Latin) (10th ed.). Holmiæ (Stockholm): Laurentius Salvius. pp. 39–40. Retrieved November 23, 2012.
  2. ↑ Mech, L.D.; Boitani, L. & IUCN SSC Wolf Specialist Group (2010). "Canis lupus". The IUCN Red List of Threatened Species. 2010: e.T3746A10049204. doi:10.2305/IUCN.UK.2010-4.RLTS.T3746A10049204.en. Retrieved 23 December 2017.
  3. ↑ International Code of Zoological Nomenclature. "ICZN Glossary". International Commission on Zoological Nomenclature.
  4. ↑ Peter J. Russell; Paul E. Hertz; Beverly McMillan (2011). "21-Speciation". Biology: The Dynamic Science. Brooks/Cole California. p. 456. ISBN 978-1133418849.
  5. 1 2 Wozencraft, W.C. (2005). "Order Carnivora". In Wilson, D.E.; Reeder, D.M. Mammal Species of the World: A Taxonomic and Geographic Reference (3rd ed.). Johns Hopkins University Press. pp. 532–628. ISBN 978-0-8018-8221-0. OCLC 62265494.
  6. 1 2 Smithsonian - Animal Species of the World database. "Canis lupus".
  7. ↑ Canis lupus
  8. ↑ Mech, L. David (1981). The Wolf: The Ecology and Behaviour of an Endangered Species, University of Minnesota Press, p. 354, ISBN 0-8166-1026-6
  9. 1 2 3 4 Wozencraft, W.C. (2005). "Order Carnivora". In Wilson, D.E.; Reeder, D.M. Mammal Species of the World: A Taxonomic and Geographic Reference (3rd ed.). Johns Hopkins University Press. pp. 575–577. ISBN 978-0-8018-8221-0. OCLC 62265494. url=https://books.google.com/books?id=JgAMbNSt8ikC&pg=PA576
  10. 1 2 3 Fan, Zhenxin; Silva, Pedro; Gronau, Ilan; Wang, Shuoguo; Armero, Aitor Serres; Schweizer, Rena M.; Ramirez, Oscar; Pollinger, John; Galaverni, Marco; Ortega Del-Vecchyo, Diego; Du, Lianming; Zhang, Wenping; Zhang, Zhihe; Xing, Jinchuan; Vilà, Carles; Marques-Bonet, Tomas; Godinho, Raquel; Yue, Bisong; Wayne, Robert K. (2016). "Worldwide patterns of genomic variation and admixture in gray wolves". Genome Research. 26 (2): 163–73. doi:10.1101/gr.197517.115. PMC 4728369. PMID 26680994.
  11. ↑ Nowak, R. M. (1995). Another look at wolf taxonomy. pp. 375-397 in L. N. Carbyn, S. H. Fritts and D. R. Seip (eds), Ecology and conservation of wolves in a changing world: proceedings of the second North American symposium on wolves, Edmonton, Canada.
  12. ↑ Mech & Boitani 2003, pp. 246
  13. 1 2 3 Andrew T. Smith; Yan Xie; Robert S. Hoffmann; Darrin Lunde; John MacKinnon; Don E. Wilson; W. Chris Wozencraft, eds. (2008). A Guide to the Mammals of China. Princeton University press. pp. 416–418. ISBN 978-0691099842.
  14. 1 2 3 Wang, Yingxiang (2003). A Complete Checklist of Mammal Species and Subspecies in China (A Taxonomic and Geographic Reference). China Forestry Publishing House, Beijing, China. ISBN 978-7503831317.
  15. ↑ "Canis lupus albus Kerr, 1792". Integrated Taxonomic Information System.
  16. 1 2 Heptner, V. G. & Naumov, N., P. (1998) Mammals of the Soviet Union Vol.II Part 1a, SIRENIA AND CARNIVORA (Sea cows; Wolves and Bears), Science Publishers, Inc., USA, pp. 182-184, ISBN 1-886106-81-9
  17. ↑ Wozencraft, W.C. (2005). "Order Carnivora". In Wilson, D.E.; Reeder, D.M. Mammal Species of the World: A Taxonomic and Geographic Reference (3rd ed.). Johns Hopkins University Press. ISBN 978-0-8018-8221-0. OCLC 62265494.
  18. ↑ "Canis lupus arabs Pocock, 1934". Integrated Taxonomic Information System.
  19. ↑ Lopez, Barry (1978). Of wolves and men. New York: Scribner Classics. p. 320. ISBN 978-0-7432-4936-2.
  20. ↑ Fred H. Harrington; Paul C. Paquet (1982). Wolves of the World: Perspectives of Behavior, Ecology, and Conservation. p. 474. ISBN 978-0-8155-0905-9.
  21. 1 2 Heptner, V. G. & Naumov, N., P. (1998) Mammals of the Soviet Union Vol.II Part 1a, SIRENIA AND CARNIVORA (Sea cows; Wolves and Bears), Science Publishers, Inc., USA, pp. 188-89, ISBN 1-886106-81-9
  22. ↑ Wozencraft, W.C. (2005). "Order Carnivora". In Wilson, D.E.; Reeder, D.M. Mammal Species of the World: A Taxonomic and Geographic Reference (3rd ed.). Johns Hopkins University Press. ISBN 978-0-8018-8221-0. OCLC 62265494.
  23. 1 2 Gray, J. E. (1863). Notice of the Chanco or Golden Wolf (Canis chanco) from Chinese Tartary. Proceedings of the Zoological Society of London: 94
  24. ↑ Canis lupus var. chanco Mivart Monograph of Canidae, p. 8, pi. 3, 1890.
  25. ↑ Abe Yoshio, "Nukutei ni tisuit" (On Nuketei) Dobutsugaku zasshi (Zoological Magazine) 35 (1923): 320-86
  26. ↑ Heptner, V.G. and Naumov, N.P. (1998). Mammals of the Soviet Union Vol.II Part 1a, SIRENIA AND CARNIVORA (Sea cows; Wolves and Bears), Science Publishers, Inc. USA., pp. 164-270, ISBN 1-886106-81-9
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