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Himalayan wolf

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Himalayan wolf
In the Upper Mustang region of the Annapurna Conservation Area, Nepal
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Carnivora
Family: Canidae
Genus: Canis
Species:
C. lupus
Subspecies:
C. l. filchneri
Trinomial name
Canis lupus filchneri

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 the new Himalayan wolf clade is disputed, with the separate species Canis himalayensis proposed based on two studies limited to mtDNA. The model produced by the first study only had 50% likelihood, which provides weak support for the proposal. The second study used more specimens and a larger number of genetic sequences; its model had a 92–99% likelihood, and showed a close relationship with the African golden wolf.[1]

Distribution

The proposed Himalayan wolf is found in northern India in the Ladakh region of eastern Kashmir[2] and the Lahaul and Spiti region in the northeastern part of Himachal Pradesh.[3][4] It is also found in Nepal in the Upper Dolpa[5] and Upper Mustang regions.[6][3] A population of them can also be found in Tibet.[3]

The Indian population consists of 350 wolves with a range of 70,000 km2 (27,000 sq mi).[7] In 2004, a group of 33 Himalayan wolves were spotted in the Spiti Valley in the northeastern part of Himachal Pradesh.[8] In the same year, the wolf was spotted for the first time in Nepal in the Upper Mustang region.[6]

Description

Morphological appearance of the wolves from different parts of India shows certain dissimilarities. Skulls of the two males from Chumar, Ladakh were measured by Allen (234 and 236 mm), which are the largest for wolves in India, but smaller compared to North American wolves, which can measure up to 290 mm.

The wolf from peninsular India appears smaller in size and more brownish in colour, whereas wolves from the Himalayan regions are large and whitish. Peninsular wolf weighs 25 kg on an average, which may be the lowest among all wolves, whereas wolves from the Himalayan region weigh about 35 kg, similar to Tibetan wolves.[9]

The wolves from Upper Mustang, Nepal are characterized by their "distinct white coloration around the throat, chest, belly, and inner part of the legs; woolliness of body fur; stumpy legs; unusual elongation of the muzzle, a muzzle arrayed with closely-spaced black speckles which extend below the eye on to the upper cheeks and ears; and smaller size compared to the European wolf."[10]

Canis lupus pallipes has the smallest skull length, measuring maximum up to 220 mm. Zygomatic widths of the skull of wolves from Ladakh (129 and 128 mm) were also comparatively larger than those of peninsular wolves from India (90.2–109 mm). Upper cheek teeth, i.e. canine to last molar of two wolves from Ladakh measured 105 and 98.4 mm, which is larger compared to those of peninsular wolves and Arabian wolves (93.6–97 mm and 81.3–93 mm respectively)14–16.

A comparative study of Himalayan wolves with other grey wolf subspecies howls demonstrated that the Himalayan wolf howls typically had lower frequencies and were shorter in duration. The study found that Himalayan and North African wolves showed the most acoustically distinct howls and differed significantly from each other and to the other wolf subspecies.[11]

Taxonomy

The cladogram, below, shows genetic relationships between the extant species of wolf-like canids, based on DNA sequence data taken from the cell nucleus,[12][13] except for the Himalayan wolf, whose placement in the diagram is based only on mitochondrial DNA sequences.[13][1]

Phylogenetic tree of the extant wolf-like canids
Caninae 3.5 Ma
Phylogenetic tree of extant wolf-like canids based on nuclear DNA,[12][13] with Himalayan wolf placement only from mtDNA sequences.[13][1] Numbers on the branches are estimated divergence times, in millions of years.[13]

The wolves of Tibet were first described by British zoologist Brian Houghton Hodgson in 1847 as Canis laniger.[14] In 1907, the German zoologist Paul Matschie described a type specimen of a wolf that lived in the Gansu and Qinghai regions of China, which he named Canis filchneri Matschie (1907).[15] In the third edition of Mammal Species of the World published in 2005, the mammalogist W. Christopher Wozencraft listed under the wolf Canis lupus the taxonomic synonyms for the subspecies Canis lupus filchneri, under which he included filchneri Matschie (1907) and laniger Hodgson (1847).[16]

In 1941, Reginald Pocock had referred to the Tibetan wolf as Canis lupus laniger and classified it as a synonym under C. l. chanco.[4] There are some researchers who still refer to Pocock's classification of the Tibetan wolf as C. l. chanco, which has caused taxonomic confusion.

The wolves living in the Trans-Himalaya region have unique haplotypes, do not cluster phylogenetically with other gray wolves, were basal to all other wolves and are closer to the jackal. This indicates that these are the descendants of an ancient wolf distribution and a new taxonomic classification of Canis himalayensis is proposed.[3][17][18][19] The taxonomic reference Mammal Species of the World does not recognize Canis himalayensis, however NCBI/Genbank does list Canis lupus himalayensis.[20]

There are now academic works that refer to the Himalayan wolf as Canis himalayensis,[5][21][17][3] and the Tibetan wolf as Canis lupus laniger.[22][23]

See also: Tibetan wolf § Taxonomy

Lineage

A study in 2004 compared sequences of 582 base pairs in length from the mtDNA control region[a] for 27 wolves from the Himalayas and the Tibetan Plateau. The phylogenetic tree generated from the sequences indicated that there were 5 related haplotypes which formed a clade that is basal to all other wolves.[b] This clade included one sequence from Ladakh in eastern Kashmir, nine from the Spiti Valley in Himachal Pradesh, four from Nepal, and two from Tibet. Seven wolves also from Kashmir did not fall into this clade.

Hodgson (1847) described the Tibetan wolf as C. laniger. A study of the mitochondrial control region of the specimen that Hodgson collected (labelled BM58.6.24.61) was found to fall within the proposed Himalayan wolf clade.[3] However, the 50% likelihood provided by the model on which this study rested provides only weak support for this proposal.[9][17][3] A criticism of this study is that it was based on zoo specimens and it is known that all zoo specimens have been captive-bred, descended from only two females.[a] Therefore, the study did not provide a representative sample.[10][9] Additionally, the wolf population in the Kashmir valley is known to have recently arrived in that area.[9]

Himalayan wolf profile
Pin valley within in the Spiti valley, Himachal Pradesh showing the environment of the wolves

In 2007, a study compared sequences of 1,300 base pairs in length from the mtDNA[a] control region for 18 wolves from the Padmaja Naidu Himalayan Zoological Park near Darjeeling in West Bengal, northern India. The study found that the wolves were basal[b] to all other wolves and formed one haplotype, indicating that they shared a common female ancestor. The study did not ascertain if the zoo specimens were all related. The conclusion supported the 2004 study that the Himalayan wolf differed from other Tibetan wolves.[17] A criticism of this study is that it was based on limited data with no samples collected from the Kashmir valley population, despite suggesting that Kashmir is an area of potential contact of the closely related wolf clades. Instead, the samples were collected from Indian zoos or museum specimens. Additionally, the areas under study are part of the same landscape, and the question of what ecological or behavioral barriers could be facilitating such strict divergence, particularly when no striking morphological differences occur between the wolves from Tibet and Indian Trans-Himalaya, remains unanswered.[9]

In April 2009, the Latin binomen Canis himalayensis was proposed for this clade as a separate species of wolf through the Nomenclature Specialist on the CITES Animals Committee. The proposal was based on one study that relied on only a limited number of museum and zoo samples that may not have been representative of the wild population.[9][5] The committee recommended against this proposal but suggested that the name be entered into the species database as a synonym for Canis lupus.[25][26] The committee stated that the classification was for conservation purposes only, and did not "reflect the latest state of taxonomic knowledge".[26] Further fieldwork was called for.[9]

Other researchers have questioned this proposed taxonomic classification, claiming that recent genetic studies have not provided a complete picture. The 2007 argues that the Himalayan wolf is different to the wolves from Tibet. As these areas are part of the same landscape, the question of what ecological or behavioural barriers could be facilitating such strict divergence, particularly when no striking morphological differences occur between the wolves from Tibet and Indian Trans-Himalaya, remains unanswered. Another problem is related to limited data as none of the studies have collected samples from the Kashmir valley population, despite suggesting it as the area of potential contact of the closely related wolf clades. Instead, the samples have been collected from Indian zoos or museum specimens.[9]

Two later studies compared sequences from the wolves of the Himalayas against worldwide wolf sequences and confirmed their basal position, however these studies did not include wolves from Tibet.[27][28][29] In 2012, a study compared sequences of 300 base pairs in length from the mtDNA control region of the scats of 2 wolves from remote and widely separated areas in the Upper Dolpa, Nepal and found that these sequences matched the Himalayan wolf.[5] In 2016, study compared sequences of 220 base pairs in length from the mtDNA control region from the fecal remains of 4 wild wolves from the Upper Mustang region of the Annapurna Conservation Area, Nepal. The study showed that they fell within the Himalayan wolf clade but formed a separate haplotype to those previously studied.[10]

Later in 2016, a study compared wolf mitochondrial DNA sequences of 582 base pairs in length. The phylogenetic tree generated from the sequences revealed that the two most-basal[b] clades were formed by specimens from India and the Himalayas.[c] Within the Himalayan / Tibetan wolf clade also fell some specimens from China and Mongolia,[30] indicating a common maternal ancestor[a] and a wider genetic distribution.[30]

See also: Tibetan wolf § Lineage

Relationship with the African golden wolf

In 2015, an mDNA study showed that the sequences from 3 wolf specimens from the Himalayas and Tibet clustered with those of the African golden wolf rather than the gray wolf.[13] In 2017, a study of mitochondrial DNA and other DNA sequences found that the Himalayan wolf formed a basal distinct monophyletic clade relative to the holarctic gray wolf.

The wolves at the Padmaja Naidu Himalayan Zoological Park, Darjeeling were found by a 2007 study to fall into the Himalayan wolf clade

Its degree of divergence to the holarctic gray wolf is similar to the degree of divergence of the African golden wolf from the holarctic gray wolf. The Himalayan wolf possesses a unique X-chromosome haplotype but shares a Y-chromosome haplotype with the African golden wolf. The study's samples show the range of the Himalayan wolf to include the Himalayas then north across the Tibetan plateau to the Qinghai Lake in Qinghai Province, China.[1]

Various authorities have called for a study to collect and analyse the genetic samples from wolves from all areas in the Himalayas, in order to provide wide representation and hence more reliable results of genetic relatedness among the different wolf-like canids.[1][31][9][5][26][25]

The recognition of a separate species or subspecies is pending on more DNA evidence from nuclear markers (taken from the cell nucleus rather than from the cell mitochondria).[10]

Conservation

Wolves at the Padmaja Naidu Himalayan Zoological Park, Darjeeling

The Himalayan wolf is listed as an endangered species in certain areas of Jammu and Kashmir, Himachal Pradesh, and Uttarakhand. A large portion of the wolf population in these areas exists outside of the protected area network, which is alarming for the initiatives of their conservation and suggests that management for conservation in these areas should equally consider the area outside protected areas.[9] Their scarce populations and evolutionary uniqueness have been underlined in some recent studies. Lack of information about their basic ecology in this landscape is a severe hindrance towards a sound conservation plan for these animals.[31]

It has been suggested by several biologists in India for recognition as a critically endangered species of canid.[3] Although the Indian government has added the Himalayan wolf to its endangered species list in 1998, it still lacks legal protection in Tibet.

Captive breeding

Eighteen Himalayan wolves are being bred in captivity. They were captured in the wild and are now being preserved in the trans-Himalayan region of India, at the Padmaja Naidu Himalayan Zoological Park in Shiwalik Hills on the lower range of the Himalaya in West Bengal, and in the Kufri Zoo with Kufri Himalayan National Park located in Himachal Pradesh province.[7]

See also

Notes

  1. ^ a b c d DNA sequences can be mapped to reveal a phylogenetic tree that represents evolutionary relationships, with each branch point representing the proposed divergence of two lineages from a common ancestor. An animal's mitochondrial DNA (mtDNA) is inherited only from its mother.
  2. ^ a b c The term basal taxon refers to a lineage that diverges early in the history of the group and lies on a branch that originates near the common ancestor of the group."[24]
  3. ^ For the purpose of this study the "Himalayas" classification included specimens from Tibet as well as from the Himalayas.

References

  1. ^ a b c d e Werhahn, G.; Senn, H.; Kaden, J.; Joshi, J.; Bhattarai, S.; Kusi, N.; Sillero-Zubiri, C.; MacDonald, D. W. (2017). "Phylogenetic evidence for the ancient Himalayan wolf: Towards a clarification of its taxonomic status based on genetic sampling from western Nepal" (PDF). Royal Society Open Science. 4 (6): 170186. doi:10.1098/rsos.170186.
  2. ^ Lydekker, R. (1900). The Tibetan Wolf. Pages 339–340 in: The great and small game of India, Burma, and Tibet. R. Ward, London.
  3. ^ a b c d e f g h Sharma, D. K.; Maldonado, J. E.; Jhala, Y. V.; Fleischer, R. C. (2004). "Ancient wolf lineages in India". Proceedings of the Royal Society B: Biological Sciences. 271 (Supplement 3): S1–S4. doi:10.1098/rsbl.2003.0071. PMC 1809981. PMID 15101402.
  4. ^ a b Pocock, R. I. (1941). Canis lupus chanco Pages 86–90 in: Fauna of British India: Mammals Volume 2. Taylor and Francis, London
  5. ^ a b c d e Subba, S.A. (2012). "Assessing the genetic status, distribution, prey selection and conservation issues of Himalayan wolf (Canis himalayensis) in Trans-Himalayan Dolpa, Nepal" (PDF). Rufford Small Grants Foundation.
  6. ^ a b Arnold, Carrie (28 April 2016). "Woolly Wolf Spotted in Nepal Is Likely a New Species". National Geographic. Retrieved 29 April 2016.
  7. ^ a b Aggarwal, R. K.; Ramadevi, J.; L. Singh (2003). "Ancient origin and evolution of the Indian wolf: evidence from mitochondrial DNA typing of wolves from Trans-Himalayan region and Pennisular India". Genome Biology. 4: 6. doi:10.1186/gb-2003-4-6-p6. Retrieved 28 August 2013.{{cite journal}}: CS1 maint: unflagged free DOI (link) Note: this is a "deposited article" which means a first copy not peer reviewed. The final published work was Aggarwal (2007)
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  12. ^ a b Lindblad-Toh, K.; Wade, C. M.; Mikkelsen, T. S.; Karlsson, E. K.; Jaffe, D. B.; Kamal, M.; Clamp, M.; Chang, J. L.; Kulbokas, E. J.; Zody, M. C.; Mauceli, E.; Xie, X.; Breen, M.; Wayne, R. K.; Ostrander, E. A.; Ponting, C. P.; Galibert, F.; Smith, D. R.; Dejong, P. J.; Kirkness, E.; Alvarez, P.; Biagi, T.; Brockman, W.; Butler, J.; Chin, C. W.; Cook, A.; Cuff, J.; Daly, M. J.; Decaprio, D.; et al. (2005). "Genome sequence, comparative analysis and haplotype structure of the domestic dog". Nature. 438 (7069): 803–819. Bibcode:2005Natur.438..803L. doi:10.1038/nature04338. PMID 16341006.
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  14. ^ Hodgson BH. (1847) Description of the wild ass and wolf of Tibet. Calcutta Journal of Natural History 7: 469–477
  15. ^ lupus filchneri Matschie. Wiss Ergebn Exped. Filchner mach China u. Tibet, 10 1, p153. Si-ning to the east from Kukunor (Siningfu, Kansu, China)
  16. ^ 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. 575–577. ISBN 978-0-8018-8221-0. OCLC 62265494. url=https://books.google.com/books?id=JgAMbNSt8ikC&pg=PA576
  17. ^ a b c d Aggarwal, R. K.; Kivisild, T.; Ramadevi, J.; Singh, L. (2007). "Mitochondrial DNA coding region sequences support the phylogenetic distinction of two Indian wolf species". Journal of Zoological Systematics and Evolutionary Research. 45 (2): 163–172. doi:10.1111/j.1439-0469.2006.00400.x. Note: this is the final, peer-reviewed version of Aggarwal (2003)
  18. ^ Spotte, Stephen (2012). "1-What makes a wolf?". Societies of Wolves and Free-ranging Dogs. Cambridge University Press, Cambridge, UK. p. 4. ISBN 978-1-107-01519-7. {{cite book}}: Cite has empty unknown parameter: |editors= (help)
  19. ^ Aggarwal, Ramesh K. (2007). "7-Molecular Genetic Studies on highly Endangered Species". You Deserve, We Conserve: A Biotechnological Approach to Wildlife Conservation. I K International Publishing House, New Dehli. pp. 54–57. ISBN 978-81-89866-24-2. {{cite book}}: Unknown parameter |editors= ignored (|editor= suggested) (help)
  20. ^ "Canis lupus himalayensis". NCBI.NLM.NIH.gov. National Center for Biotechnology Information, U.S. National Institutes of Health.
  21. ^ Joshi, L.R., 2011. Canis himalayensis (Wolf). Wildlife Biology.
  22. ^ Zhao, Chao; Zhang, Honghai; Zhang, Jin; Chen, Lei; Sha, Weilai; Yang, Xiufeng; Liu, Guangshuai (2014). "The complete mitochondrial genome sequence of the Tibetan wolf (Canis lupus laniger)". Mitochondrial DNA. 27 (1): 7–8. doi:10.3109/19401736.2013.865181. PMID 24438245.
  23. ^ Meng, Chao; Zhang, Honghai; Meng, Qingcheng (2009). "Mitochondrial genome of the Tibetan wolf". Mitochondrial DNA. 20 (2–3): 61–3. doi:10.1080/19401730902852968. PMID 19347764.
  24. ^ "26-Phylogeny and the tree of life". Campbell Biology Australian and New Zealand version (10 ed.). Pierson Australia. 2015. pp. 561–562. ISBN 9781486007042. {{cite book}}: Unknown parameter |authors= ignored (help)
  25. ^ a b CITES. "Canis lupus". Checklist of CITES species. CITES.
  26. ^ a b c CITES (2009). "Nomenclatural Matters: Twenty-fourth meeting of the Animals Committee Geneva, (Switzerland), 20–24 April 2009, AC24 Doc. 13 Rev. 1 Annex 1" (PDF). p. 3.
  27. ^ Miklosi, Adam (2015). Dog Behaviour, Evolution, and Cognition. Oxford Biology (2 ed.). Oxford University Press. pp. 106–107. ISBN 978-0199545667.
  28. ^ Pilot, Małgorzata; Branicki, Wojciech; Jędrzejewski, Włodzimierz; Goszczyński, Jacek; Jędrzejewska, Bogumiła; Dykyy, Ihor; Shkvyrya, Maryna; Tsingarska, Elena (2010). "Phylogeographic history of grey wolves in Europe". BMC Evolutionary Biology. 10: 104. doi:10.1186/1471-2148-10-104. PMC 2873414. PMID 20409299.{{cite journal}}: CS1 maint: unflagged free DOI (link)
  29. ^ Leonard, Jennifer A.; Vilà, Carles; Fox-Dobbs, Kena; Koch, Paul L.; Wayne, Robert K.; Van Valkenburgh, Blaire (2007). "Megafaunal Extinctions and the Disappearance of a Specialized Wolf Ecomorph" (PDF). Current Biology. 17 (13): 1146. doi:10.1016/j.cub.2007.05.072. PMID 17583509.
  30. ^ a b Ersmark, Erik; Klütsch, Cornelya F. C.; Chan, Yvonne L.; Sinding, Mikkel-Holger S.; Fain, Steven R.; Illarionova, Natalia A.; Oskarsson, Mattias; Uhlén, Mathias; Zhang, Ya-Ping; Dalén, Love; Savolainen, Peter (2016). "From the Past to the Present: Wolf Phylogeography and Demographic History Based on the Mitochondrial Control Region". Frontiers in Ecology and Evolution. 4. doi:10.3389/fevo.2016.00134.{{cite journal}}: CS1 maint: unflagged free DOI (link)
  31. ^ a b Habib, B.; Shrotriya, S.; Jhala, Y. V. (January 2013). "Ecology and Conservation of Himalayan Wolf". Technical Report No. TR – 2013/01. Wildlife Institute of India: 46. doi:10.13140/RG.2.2.36012.87685.
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