|In the Upper Mustang region of the Annapurna Conservation Area, Nepal|
|Subspecies:||C. l. filchneri|
|Canis lupus filchneri|
The Himalayan wolf is a canine of unresolved taxonomy. It is currently classified as Canis lupus filchneri (Tibetan wolf). It is distinguished by its mitochondrial DNA and other genetic markers, which show it to be genetically basal to the Holarctic grey wolf and indicate an association with the African golden wolf (Canis anthus). Several studies propose this wolf as the separate species Canis himalayensis.
|Phylogenetic tree of the extant wolf-like canids|
|Phylogenetic relationships between the extant wolf-like canids based on DNA taken from the cell nucleus, except for the Himalayan wolf (Canis lupus filchneri) that is based on mitochondrial DNA sequences plus X chromosome and Y chromosome sequences. Timing in millions of years.|
The wolves of Tibet were first described by British zoologist Brian Houghton Hodgson in 1847 as Canis laniger. 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). 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).
In 1941, Reginald Pocock had referred to the Tibetan wolf as Canis lupus laniger and classified it as a synonym under C. l. chanco. 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 grey 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. The taxonomic reference Mammal Species of the World does not recognize Canis himalayensis, however NCBI/Genbank does list Canis lupus himalayensis.
A 2004 study 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. The study proposed that the Himalayan wolf clade diverged from other canids 800,000 years ago. 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 BM126.96.36.199) was found to fall within the proposed Himalayan wolf clade. However, the 50% likelihood provided by the model on which this study rested provides only weak support for this proposal. 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. Additionally, the wolf population in the Kashmir valley is known to have recently arrived in that area.
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. 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.
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. The committee recommended against this proposal but suggested that the name be entered into the species database as a synonym for Canis lupus. The committee stated that the classification was for conservation purposes only, and did not "reflect the latest state of taxonomic knowledge". Further fieldwork was called for.
Other researchers have questioned this proposed taxonomic classification, claiming that recent genetic studies have not provided a complete picture. The 2007 study proposes that the Himalayan wolf is genetically 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.
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. 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. 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.
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, indicating a common maternal ancestor[a] and a wider genetic distribution.
Relationship with the African golden wolf
Between 2011 and 2015, two mDNA studies found that the Himalayan wolf and Indian grey wolf were closer to the African golden wolf than they were to the Holarctic grey wolf. 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. Its degree of divergence from the holarctic grey wolf is similar to the degree of divergence of the African golden wolf from the holarctic grey wolf. The Himalayan wolf shares a maternal lineage with the African golden wolf. It possesses a unique paternal lineage that falls between the grey wolf and 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. The study called for the Himalayan wolf to be recognised as the species Canis himalayensis or at least as the subsepcies Canis lupus himalayensis.
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. The recognition of a separate species or subspecies is pending on more DNA evidence from the additional inclusion of genetic markers taken from the cell nucleus.
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.
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.
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.
The proposed Himalayan wolf is found in northern India in the Ladakh region of eastern Kashmir and the Lahaul and Spiti region in the northeastern part of Himachal Pradesh. It is also found in Nepal in the Upper Dolpa and Upper Mustang regions, then north across the Tibetan plateau to the Qinghai Lake in Qinghai Province, China.
The Indian population consists of 350 wolves with a range of 70,000 km2 (27,000 sq mi). In 2004, a group of 33 Himalayan wolves were spotted in the Spiti Valley in the northeastern part of Himachal Pradesh. In the same year, the wolf was spotted for the first time in Nepal in the Upper Mustang region.
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. 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.
It has been suggested by several biologists in India for recognition as a critically endangered species of canid. Although the Indian government has added the Himalayan wolf to its endangered species list in 1998, it still lacks legal protection in Tibet.
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.
- 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.
- 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."
- For the purpose of this study the "Himalayas" classification included specimens from Tibet as well as from the Himalayas.
- 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. Bibcode:2017RSOS....470186W. doi:10.1098/rsos.170186.
- 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.
- 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)
- 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.
- Koepfli, K.-P.; Pollinger, J.; Godinho, R.; Robinson, J.; Lea, A.; Hendricks, S.; Schweizer, R. M.; Thalmann, O.; Silva, P.; Fan, Z.; Yurchenko, A. A.; Dobrynin, P.; Makunin, A.; Cahill, J. A.; Shapiro, B.; Álvares, F.; Brito, J. C.; Geffen, E.; Leonard, J. A.; Helgen, K. M.; Johnson, W. E.; O'Brien, S. J.; Van Valkenburgh, B.; Wayne, R. K. (2015-08-17). "Genome-wide Evidence Reveals that African and Eurasian Golden Jackals Are Distinct Species". Current Biology. 25 (16): 2158–65. doi:10.1016/j.cub.2015.06.060. PMID 26234211.
- Hodgson BH. (1847) Description of the wild ass and wolf of Tibet. Calcutta Journal of Natural History 7: 469–477
- lupus filchneri Matschie. Wiss Ergebn Exped. Filchner mach China u. Tibet, 10 1, p153. Si-ning to the east from Kukunor (Siningfu, Kansu, China)
- 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
- Pocock, R. I. (1941). Canis lupus chanco Pages 86–90 in: Fauna of British India: Mammals Volume 2. Taylor and Francis, London
- 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.
- Aggarwal, Ramesh K. (2007). "7-Molecular Genetic Studies on highly Endangered Species". In M. W. Pandit, S. Shivaji, Lalji Singh. 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.
- "Canis lupus himalayensis". NCBI.NLM.NIH.gov. National Center for Biotechnology Information, U.S. National Institutes of Health.
- 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.
- Joshi, L.R., 2011. Canis himalayensis (Wolf). Wildlife Biology.
- Sarkar, Mriganka Shekhar; Segu, Harika; Bhaskar, J. V; Jakher, Rajendra; Mohapatra, Swati; Shalini, K; Shivaji, S; Reddy, P. Anuradha (2017). "Ecological preferences of large carnivores in remote, high-altitude protected areas: Insights from Buxa Tiger Reserve, India". Oryx. 52: 66. doi:10.1017/S0030605317000060.
- 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.
- 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.
- Jane B. Reece, Noel Meyers, Lisa A. Urry, Michael L. Cain, Steven A. Wasserman, Peter V. Minorsky, Robert B. Jackson, Bernard N. Cooke (2015). "26-Phylogeny and the tree of life". Campbell Biology Australian and New Zealand version (10 ed.). Pierson Australia. pp. 561–562. ISBN 9781486007042.
- Shrotriya, S.; Lyngdoh, S.; Habib, B. (2012). "Wolves in Trans-Himalayas: 165 years of taxonomic confusion" (PDF). Current Science. 103 (8). Retrieved June 27, 2014.
- Chetri, Madhu; Jhala, Yadvendradev; Jnawali, Shant Raj; Subedi, Naresh; Dhakal, Maheshwar; Yumnam, Bibek (21 Apr 2016). "Ancient Himalayan wolf (Canis lupus chanco) lineage in Upper Mustang of the Annapurna Conservation Area, Nepal". ZooKeys. 582: 143–156. doi:10.3897/zookeys.582.5966. PMC 4857050. PMID 27199590.
- CITES. "Canis lupus". Checklist of CITES species. CITES.
- 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.
- 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.
- 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.
- Miklosi, Adam (2015). Dog Behaviour, Evolution, and Cognition. Oxford Biology (2 ed.). Oxford University Press. pp. 106–107. ISBN 978-0199545667.
- 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.
- Rueness, Eli Knispel; Asmyhr, Maria Gulbrandsen; Sillero-Zubiri, Claudio; MacDonald, David W; Bekele, Afework; Atickem, Anagaw; Stenseth, Nils Chr (2011). "The Cryptic African Wolf: Canis aureus lupaster is Not a Golden Jackal and is Not Endemic to Egypt". PLoS ONE. 6 (1): e16385. Bibcode:2011PLoSO...616385R. doi:10.1371/journal.pone.0016385. PMC 3027653. PMID 21298107.
- 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.
- Hennelly, Lauren; Habib, Bilal; Root-Gutteridge, Holly; Palacios, Vicente; Passilongo, Daniela (2017). "Howl variation across Himalayan, North African, Indian, and Holarctic wolf clades: tracing divergence in the world's oldest wolf lineages using acoustics". Current Zoology. 63 (3): 341–348. doi:10.1093/cz/zox001.
- Lydekker, R. (1900). The Tibetan Wolf. Pages 339–340 in: The great and small game of India, Burma, and Tibet. R. Ward, London.
- Arnold, Carrie (28 April 2016). "Woolly Wolf Spotted in Nepal Is Likely a New Species". National Geographic. Retrieved 29 April 2016.
- 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. Note: this is a "deposited article" which means a first copy not peer reviewed. The final published work was Aggarwal (2007)
- "Indian wolves are world's oldest". BBC News. April 17, 2004.