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Subspecies of Canis lupus

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Skull of a European wolf
Skull of a Canadian wolf
Original distribution of subspecies
Present distribution of subspecies (excluding dingos and domestic dogs)

Canis lupus, the gray wolf, 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.

Geographical variations

Wolves show a great deal of polymorphism geographically, though they can interbreed. The Zoological Gardens of London for example once successfully managed to mate a male European wolf to an Indian female, resulting in a pup bearing an almost exact likeness to its sire.[2]

Europe

European wolves tend to have fur with less soft wool intermixed than American wolves. Their heads are narrower, their ears longer, higher placed and somewhat closer to each other. Their loins are more slender, their legs longer, their feet narrower, and their tails more thinly clothed with fur.[3] Pelt color in European wolves ranges from white, cream, red, grey and black, sometimes with all colors combined. Wolves in central Europe tend to be more richly colored than those in Northern Europe. Eastern European wolves tend to be shorter and more heavily built than Northern Russian ones.[4]

North America

North American wolves are generally the same size as European wolves, but have shorter legs, larger, rounder heads, broader, more obtuse muzzles, and a sensible depression at the union of nose and forehead, which is more arched and broad. Their ears are shorter and have a more conical form. They typically lack the black mark on the forelegs, as is the case in European races. They have long and comparatively fine fur, mixed with a shorter wooly hair, and are more robust.[3] Fur color in American wolves ranges from white, black, red, yellow, brown, grey, and grizzled skins, and others representing every shade between, although usually each locality has its prevailing tint. There are pronounced differences in North American wolves of different localities; wolves from Texas and New Mexico are comparatively slim animals with small teeth.[5] Mexican wolves in particular resemble some European wolves in stature, though their heads are usually broader, their necks thicker, their ears longer and their tails shorter.[6] Wolves of the central and northern chains of the Rocky Mountains and coastal ranges are more formidable animals than the more southern plains wolves, and resemble Russian and Scandinavian wolves in size and proportions.[5]

List of subspecies

Canis lupus subspecies

Subspecies as of 2005:[7]

Subspecies Authority Description Range Synonyms
Eurasian wolf
Canis lupus lupus (nominate subspecies)

Linnaeus 1758[8] Generally a large subspecies measuring 105–160 cm in length and weighing 40–80 kg. The pelt is usually a mix of rusty ocherous and light grey.[9] 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)[10]
Tundra wolf
Canis lupus albus

Kerr 1792[11] A large subspecies, with adults measuring 112–137 cm, and weighing 36.6–52 kg. The fur is very long, dense, fluffy and soft and is usually very light and grey in color. The lower fur is lead-grey and the upper fur is reddish-grey.[9] 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[9] dybowskii (Domaniewski, 1926), kamtschaticus (Dybowski, 1922),

turuchanensis (Ognev, 1923)[12]

Kenai Peninsula wolf
Canis lupus alces
Goldman 1941[13] A large wolf measuring over 200 cm in length and weighing 45–90 kg. It is thought that its large size was an adaptation to hunting the extremely large moose of the Kenai Peninsula.[14] Kenai Peninsula
Arabian wolf
Canis lupus arabs

Pocock 1934[15] A small, "desert adapted" wolf that is around 66 cm tall and weighs, on average, about 18 kg.[16] Its fur coat varies from short in the summer and long in the winter, possibly because of solar radiation.[17] Southern Israel, Southern and western Iraq, Oman, Yemen, Jordan, Saudi Arabia, and probably some parts of the Sinai Peninsula
Arctic wolf
Canis lupus arctos

Pocock 1935[18] A medium-sized wolf that is between 64 and 79 cm tall and 89 to 189 cm long, weighing between 35 and 45 kg on average, though there have been specimens found weighing up to 68 kg.[19][20] Canadian Arctic, Alaska and northern Greenland
Mexican wolf
Canis lupus baileyi

Nelson and Goldman 1929[21] A small subspecies which weighs 25–45 kg and measures 140–170 cm in total length (nose to tip of tail), and 72–80 cm in shoulder height. The pelt contains a mix of grey, black, brown, and rust colors in a characteristic pattern, with white underparts[22] Northern Mexico, western Texas, southern New Mexico, and southeastern and central Arizona[22]
Newfoundland wolf
Canis lupus beothucus

G. M. Allen and Barbour 1937 A white colored subspecies, extinct in 1911, typically measuring 180 cm in length and weighing 45 kg[14] Newfoundland
Bernard's wolf
Canis lupus bernardi
Anderson 1943 This subspecies became extinct in 1934. It was described as "white with black-tipped hair along the ridge of the back".[23] Limited to Banks and Victoria Islands in the arctic banksianus (Anderson, 1943)[24]
Steppe wolf
Canis lupus campestris

File:MSU V2P1a - Canis lupus campestris.png

Dwigubski 1804 A wolf of average size with short, coarse and sparse fur. The fur is light grey on the sides and rusty, brownish grey on the back[9] Northern Ukraine, southern Kazakhstan, Caucasus and Trans-Caucasus[9] bactrianus (Laptev, 1929), cubanenesis (Ognev, 1923), desertorum (Bogdanov, 1882)[25]
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[9] Central Asia from Turkestan, Tien Shan throughout Tibet to Mongolia, Northern China, Shensi, Sichuan, Yunnan, the Western Himalayas in Kashmir from Chitral to Lahul.[26] Also occurs in the Korean peninsula[27] coreanus (Abe, 1923), dorogostaiskii (Skalon, 1936), ekloni (Przewalski, 1883), filchneri (Matschie, 1907), karanorensis (Matschie, 1907), laniger (Hodgson, 1847), niger (Sclater, 1874), tschiliensis (Matschie, 1907)[28]
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, it is generally greyish-white or white in fur color. It is a very social subspecies and can usually be found roaming in packs of five to thirty-five individuals.[29] Vancouver Island, British Columbia Cascade Mountain wolf
Canis lupus fuscus
Richardson 1839 A cinnamon colored wolf measuring 165 cm and weighing 36–49 kg[14] Cascade Range
Gregory's wolf
Canis lupus gregoryi
Goldman 1937[30] A medium-sized subspecies, though slender and tawny, its coat contains a mixture of various colors, including black, grey, white, and cinnamon.[30] In and around the lower Mississippi River basin gigas (Townsend, 1850)[31]
Manitoba wolf
Canis lupus griseoalbus
Baird 1858 North Alberta, Saskatchewan, and Manitoba knightii (Anderson, 1945)[32]
Hokkaidō wolf
Canis lupus hattai

Kishida 1931 Hokkaidō rex (Pocock, 1935)[33]
Honshū wolf
Canis lupus hodophilax

Temminck 1839 Honshū, Shikoku, and Kyūshū hodopylax (Temminck, 1844), japonicus (Nehring, 1885)[34]
Hudson Bay wolf
Canis lupus hudsonicus
Goldman 1941 Northern Manitoba and the Northwest Territories
Northern Rocky Mountains wolf
Canis lupus irremotus
Goldman 1937[30][35] This subspecies generally weighs 70–135 pounds (32–61 kg), making it one of the largest subspecies of the gray wolf in existence.[36] It is a lighter colored animal than its southern brethren, the Southern Rocky Mountains Wolf, with a coat that includes far more white and less black. In general, the subspecies favors lighter colors, with black mixing in among them.[30][37] Northern Rocky Mountains
Labrador wolf
Canis lupus labradorius

Goldman 1937[30] Labrador and northern Quebec; recent confirmed sightings on Newfoundland[38][39]
Alexander Archipelago wolf
Canis lupus ligoni
Goldman 1937[30] Alexander Archipelago Mackenzie River wolf
Canis lupus mackenzii
Anderson 1943 Northwest Territories
Baffin Island wolf
Canis lupus manningi
Anderson 1943 Baffin Island
Mogollon Mountain wolf
Canis lupus mogollonensis
Goldman 1937[30] A dark colored wolf measuring 135–150 cm in length, and weighing 27–36 kg[14] Arizona and New Mexico
Texas wolf
Canis lupus monstrabilis
Goldman 1937[30] Similar in size and color to C. lupus mogollonensis[14] Texas and New Mexico niger (Bartram, 1791)[40]
Great Plains wolf
Canis lupus nubilus

Say 1823 Minnesota, Michigan, and Wisconsin. Single wolves have been reported in the Dakotas and as far south as Nebraska variabilis (Wied-Neuwied, 1841)[41] labradorius, irremotus, youngi (Goldman, 1937)
Northwestern wolf
Canis lupus occidentalis

Richardson 1829 Western Canada sticte (Richardson, 1829), ater (Richardson, 1829),[42] 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 Alaska and Yukon Alaskan tundra wolf
Canis lupus tundrarum

Miller 1912 Has heavier dentition than pambasileus Alaska
Southern Rocky Mountains wolf
Canis lupus youngi
Goldman 1937[30] A medium-size wolf that weighed around 90 lbs on average.[43][44] It is considered to have been the "second largest wolf in the United States".[45] The coloring of the subspecies tended toward black, with lighter areas on the edges of its fur and white in various small patches.[30] Southern Rocky Mountains

Disputed subspecies and species

Italian (Apennine) wolf from the National Park of Abruzzo, Lazio e Molise
Iberian wolves

Two 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 Italian and Iberian peninsulas have morphologically distinct features from other Eurasian wolves and each are considered by their researchers to represent their own subspecies.[46][47][48]

The genetic distinction of the Italian wolf subspecies was recently 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.[49]

Recent genetic research suggests that the Indian wolf populations in the Indian subcontinent may represent a distinct species from their conspecifics. Similar results were obtained for the Himalayan wolf, which is traditionally placed under the Tibetan wolf (Canis lupus chanco).[50]

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.[51] 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.[52] The eastern wolf is also considered a distinct species by the USFWS.[51] However, this clasification is still controversial and a review stated that it was "not well-supported by best available science."[53]

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.[54] Later, research made in the early 2010s 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.[55][56]

See also

References

  1. ^ Template:IUCN2011.1
  2. ^ The Living Age, published by Littell, Son and Co., 1851
  3. ^ a b Richardson, J., Swainson, W., Kirby, W. (1829) Fauna Boreali-americana, Or, The Zoology of the Northern Parts of British America: Containing Descriptions of the Objects of Natural History Collected on the Late Northern Land Expeditions, Under Command of Captain Sir John Franklin, R.N. J. Murray, London book preview
  4. ^ Hutchinson's animals of all countries: the living animals of the world in picture and story. Volume I. 1923. p. 384.
  5. ^ a b Hunting the Grisly and Other Sketches by Theodore Roosevelt - Full Text Free Book (Part 3/3)
  6. ^ The Natural History of Dogs: Canidæ Or Genus Canis of Authors. Including Also the Genera Hyæna and Proteles by Charles Hamilton Smith, contributor William Home Lizars, Samuel Highley, W. Curry, Junr. & Co, Published by W.H. Lizars, ... S. Highley, ... London; and W. Curry, jun. and Co. Dublin., 1839
  7. ^ Wozencraft, W. C. (2005). "Order Carnivora". In Wilson, D. E.; Reeder, D. M. (eds.). Mammal Species of the World: A Taxonomic and Geographic Reference (3rd ed.). Johns Hopkins University Press. ISBN 978-0-8018-8221-0. OCLC 62265494.
  8. ^ "Canis lupus lupus Linnaeus, 1758". Integrated Taxonomic Information System.
  9. ^ a b c d e f Mammals of the Soviet Union Vol.II Part 1a, SIRENIA AND CARNIVORA (Sea cows; Wolves and Bears), V.G Heptner and N.P Naumov editors, Science Publishers, Inc. USA. 1998. ISBN 1-886106-81-9
  10. ^ Wozencraft, W. C. (2005). "Order Carnivora". In Wilson, D. E.; Reeder, D. M. (eds.). Mammal Species of the World: A Taxonomic and Geographic Reference (3rd ed.). Johns Hopkins University Press. ISBN 978-0-8018-8221-0. OCLC 62265494.
  11. ^ "Canis lupus albus Kerr, 1792". Integrated Taxonomic Information System.
  12. ^ Wozencraft, W. C. (2005). "Order Carnivora". In Wilson, D. E.; Reeder, D. M. (eds.). Mammal Species of the World: A Taxonomic and Geographic Reference (3rd ed.). Johns Hopkins University Press. ISBN 978-0-8018-8221-0. OCLC 62265494.
  13. ^ "Canis lupus alces Goldman, 1941". Integrated Taxonomic Information System.
  14. ^ a b c d e The Encyclopedia of Vanished Species by David Day, Universe Books ltd. 1981. ISBN 0-947889-30-2
  15. ^ "Canis lupus arabs Pocock, 1934". Integrated Taxonomic Information System.
  16. ^ Cite error: The named reference Lopez was invoked but never defined (see the help page).
  17. ^ Fred H. Harrington, Paul C. Paquet (1982). Wolves of the World: Perspectives of Behavior, Ecology, and Conservation. p. 474. ISBN 0-8155-0905-7.
  18. ^ "Canis lupus arctos Pocock, 1935". Integrated Taxonomic Information System.
  19. ^ "White wolf of the North", World Wide Fund for Nature
  20. ^ "Arctic wolf", Toronto Zoo
  21. ^ "Canis lupus baileyi Nelson and Goldman, 1929". Integrated Taxonomic Information System.
  22. ^ a b "Green Fire" Returns to the Southwest: Reintroduction of the Mexican Wolf- Author(s): David R. Parsons. Source: Wildlife Society Bulletin, Vol. 26, No. 4, Commemorative Issue Celebrating the 50th Anniversary of "A Sand County Almanac" and the Legacy of Aldo Leopold (Winter, 1998), pp. 799–807. Published by: Allen Press
  23. ^ "Bernard, P. and J.", The Eponym Dictionary of Mammals by Bo Beolens, Michael Watkins and Michael Grayson, JHU Press, 2009, Pg. 40
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  25. ^ Wozencraft, W. C. (2005). "Order Carnivora". In Wilson, D. E.; Reeder, D. M. (eds.). Mammal Species of the World: A Taxonomic and Geographic Reference (3rd ed.). Johns Hopkins University Press. ISBN 978-0-8018-8221-0. OCLC 62265494.
  26. ^ Fauna of British India: Mammals Volume 2 by R. I. Pocock, printed by Taylor and Francis, 1941
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  29. ^ "Preliminary Investigations of the Vancouver Island Wolf", Wolves of the world: perspectives of behavior, ecology, and conservation by Fred H. Harrington and Paul C. Paquet, William Andrew, 1982, Pg. 54
  30. ^ a b c d e f g h i j "The Wolves of North America", E. A. Goldman, Journal of Mammalogy, Vol. 18, No. 1 (Feb., 1937), pp. 37–45 Cite error: The named reference "Goldman" was defined multiple times with different content (see the help page).
  31. ^ Wozencraft, W. C. (2005). "Order Carnivora". In Wilson, D. E.; Reeder, D. M. (eds.). Mammal Species of the World: A Taxonomic and Geographic Reference (3rd ed.). Johns Hopkins University Press. ISBN 978-0-8018-8221-0. OCLC 62265494.
  32. ^ Wozencraft, W. C. (2005). "Order Carnivora". In Wilson, D. E.; Reeder, D. M. (eds.). Mammal Species of the World: A Taxonomic and Geographic Reference (3rd ed.). Johns Hopkins University Press. ISBN 978-0-8018-8221-0. OCLC 62265494.
  33. ^ Wozencraft, W. C. (2005). "Order Carnivora". In Wilson, D. E.; Reeder, D. M. (eds.). Mammal Species of the World: A Taxonomic and Geographic Reference (3rd ed.). Johns Hopkins University Press. ISBN 978-0-8018-8221-0. OCLC 62265494.
  34. ^ Wozencraft, W. C. (2005). "Order Carnivora". In Wilson, D. E.; Reeder, D. M. (eds.). Mammal Species of the World: A Taxonomic and Geographic Reference (3rd ed.). Johns Hopkins University Press. ISBN 978-0-8018-8221-0. OCLC 62265494.
  35. ^ "Canis lupus irremotus Goldman, 1937". Integrated Taxonomic Information System.
  36. ^ Nelson King (2007). "Wolves in Yellowstone: A Short History". Yellowstone Insider.
  37. ^ B. J. Verts & Leslie N. Carraway (1998). "Canis lupus Linnaeus, 1758". Land Mammals of Oregon. University of California Press. pp. 360–363. ISBN 978-0-520-21199-5.
  38. ^ "Wolf in Newfoundland probably made it to island on ice, experts say". The Telegram. 25 May 2012. Retrieved 26 August 2012.
  39. ^ "Genetic Retesting of DNA Confirms Second Wolf on Island of Newfoundland". Department of Environment and Conservation, Newfoundland and Labrador. 23 August 2012. Retrieved 26 August 2012.
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  41. ^ Wozencraft, W. C. (2005). "Order Carnivora". In Wilson, D. E.; Reeder, D. M. (eds.). Mammal Species of the World: A Taxonomic and Geographic Reference (3rd ed.). Johns Hopkins University Press. ISBN 978-0-8018-8221-0. OCLC 62265494.
  42. ^ Wozencraft, W. C. (2005). "Order Carnivora". In Wilson, D. E.; Reeder, D. M. (eds.). Mammal Species of the World: A Taxonomic and Geographic Reference (3rd ed.). Johns Hopkins University Press. ISBN 978-0-8018-8221-0. OCLC 62265494.
  43. ^ "Letters to the Editor" - The Idaho Statesman
  44. ^ "The Company of Wolves" - Google Books
  45. ^ "The wolves of North America" - Google Books
  46. ^ The wolf in Spain
  47. ^ Canis lupus italicus
  48. ^ J. Vos: Food habits and livestock depredation of two Iberian wolf packs (Canis lupus signatus) in the north of Portugal. Journal of Zoology (2000), 251: 457-462 Cambridge University Press. online abstract
  49. ^ V. LUCCHINI, A. GALOV and E. RANDI Evidence of genetic distinction and long-term population decline in wolves (Canis lupus) in the Italian Apennines. Molecular Ecology (2004) 13, 523–536. abstract online
  50. ^ R. K. Aggarwal, T. Kivisild, J. Ramadevi, L. Singh: Mitochondrial DNA coding region sequences support the phylogenetic distinction of two Indian wolf species. Journal of Zoological Systematics and Evolutionary Research, Volume 45 Issue 2 Page 163-172, May 2007 online
  51. ^ a b Chambers, Steven M.; Fain, Steven R.; Fazio, Bud; Amaral, Michael (2012). "An account of the taxonomy of North American wolves from morphological and genetic analyses". North American Fauna. 77: 1–67. doi:10.3996/nafa.77.0001.
  52. ^ "DNA profiles of the eastern Canadian wolf and the red wolf provide evidence for a common evolutionary history independent of the gray wolf". Canadian Journal of Zoology. 78 (12): 2156–2166. December 2000. doi:10.1139/z00-158. {{cite journal}}: |access-date= requires |url= (help)
  53. ^ "Review of Proposed Rule Regarding Status of the Wolf Under the Endangered Species Act" (PDF). January 2014. Retrieved June 6, 2015. {{cite web}}: line feed character in |title= at position 48 (help)
  54. ^ Wozencraft, W. C. (2005). "Order Carnivora". In Wilson, D. E.; Reeder, D. M. (eds.). 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.
  55. ^ Gaubert P, Bloch C, Benyacoub S, Abdelhamid A, Pagani P; et al. (2012). "Reviving the African Wolf Canis lupus lupaster in North and West Africa: A Mitochondrial Lineage Ranging More than 6,000 km Wide". PLoS ONE. 7 (8): e42740. doi:10.1371/journal.pone.0042740. {{cite journal}}: Explicit use of et al. in: |author= (help)CS1 maint: multiple names: authors list (link) CS1 maint: unflagged free DOI (link)
  56. ^ Gaubert P, Bloch C, Benyacoub S, Abdelhamid A, Pagani P; et al. (2012). "Reviving the African Wolf Canis lupus lupaster in North and West Africa: A Mitochondrial Lineage Ranging More than 6,000 km Wide". PLoS ONE. 7 (8): e42740. doi:10.1371/journal.pone.0042740.{{cite journal}}: CS1 maint: multiple names: authors list (link) CS1 maint: unflagged free DOI (link)