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The Tibetan wolf (Canis lupus filchneri) is a subspecies of the gray wolf that is native to China in the regions of Gansu, Qinghai, and the Tibetan Autonomous Region. It is distinguished by its genetic markers, with whole genome sequencing indicating that it is the most genetically divergent wolf population,[3] and mitochondrial DNA sequencing indicating that it is genetically the same wolf as the Himalayan wolf, is genetically basal to the Holarctic grey wolf,[4][5][6] and has an association with the African golden wolf (Canis anthus).[5][6]

Tibetan wolf
Tibetan Wolf By Stanzin (Stakpa) cropped.jpg
Scientific classification e
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Carnivora
Family: Canidae
Genus: Canis
Species:
Subspecies:
C. l. filchneri
Trinomial name
Canis lupus filchneri
Range of Canis lupus filchneri in China.jpg
Map showing the Chinese provinces (in red) in which C. l. filchneri is found
Synonyms

TaxonomyEdit

In 1847, the British naturalist Brian Houghton Hodgson classified a female wolf from Tibet as Canis lupus laniger.[2]

In 1907, the German zoologist Paul Matschie classified a wolf skin from Xining in the Qinghai province of China as Canis lupus filchneri.[1] Wilhelm Filchner was a German explorer who led an expedition from China to Tibet in 1903-1905[7] and the wolf was named by Matschie in his honor. In 1938, the American zoologist Glover Morrill Allen classified Canis lupus filchneri and Canis lupus laniger as taxonomic synonyms for Canis lupus chanco (Mongolian wolf).[8] In the third edition of Mammal Species of the World published in 2005, the American mammalogist W. Christopher Wozencraft listed under the wolf Canis lupus the subspecies Canis lupus filchneri, under which he included as a taxonomic synonym the taxon laniger Hodgson 1847.[9]

The NCBI/Genbank lists Canis lupus laniger[10] as the Tibetan wolf, and separately Canis lupus chanco [11] as the Mongolian wolf. Additionally, there are recent academic works that still refer to the Tibetan wolf as C. l. laniger.[12][13][14][15]

LineageEdit

Phylogenetic tree of the extant wolf-like canids with timing in millions of years[a]
Caninae 3.5 Ma
3.0
2.5
2.0
0.96
0.60
0.38
0.25
?
0.08
0.031

Dog  

Holarctic grey wolf  

Late Pleistocene wolf 

Indian plains wolf  

Himalayan wolf/Tibetan wolf  

Coyote  

0.11

African golden wolf northwestern Africa 

African golden wolf eastern Africa 

Golden jackal  

Ethiopian wolf  

Dhole  

African wild dog  

2.6

Side-striped jackal  

Black-backed jackal  

Blue shading represents the species Canis lupus

Studies using mitochondrial DNA (mDNA) indicate the Tibetan wolf to be genetically basal to the Holarctic gray wolf.[4][5][6] Its MT-ND4L gene commences with the base pairs GTG, whereas all other canids commence with ATG.[12] A whole genome study found it to be the most genetically divergent wolf population.[3]

In 2016, a whole genome study found that the Tibet population had shown decline over the past 25,000 years, however the Qinghai population had shown growth. The wolf's lineage had suffered a historical population bottleneck having only recently recolonized the Tibetan Plateau. Glaciation during the Last Glacial Maximum may have caused its habitat loss, genetic isolation then local adaptation, and there was evidence of ancient inbreeding. The study proposes that these are the reasons that the wolves from Tibet were the most genetically distinct. The Qinghai population showed gene flow from Chinese indigenous dogs of 16%. Gene flow from the Tibetan wolf forms 2% of the dingo's genome,[3] which likely represents ancient admixture in eastern Eurasia.[16][17][3]

Studies based on mDNA,[4] and on both mDNA and DNA taken from the cell nucleus,[5][6] indicate that the Himalayan wolf is genetically the same wolf as the Tibetan wolf. This wolf contrasts with the wolves found in the lower altitudes of Inner Mongolia, Mongolia, and the Xinjiang province of China (Mongolian wolf).[6] A further analysis based on single-nucleotide polymorphisms indicates that this wolf possesses a genetic adaptation to help it cope with living in low-oxygen high altitude habitats. This adaptation could not be found in the Holarctic grey wolf.[6][18] Some specimens found from as far away as China and Mongolia also fell within a Himalayan/Tibetan wolf clade, indicating a common maternal ancestor and a wider genetic distribution of this wolf.[4] There was evidence of hybridization with the grey wolf at Sachyat-Ertash in the Issyk-Kul region of Kyrgyzstan, and of introgression from either the grey wolf or the dog into the Himalayan wolf in Nepal.[6]

Admixture with the Tibetan mastiffEdit

The Tibetan Mastiff was able to adapt to the extreme highland conditions of the Tibetan Plateau very quickly compared to other mammals such as the yak, Tibetan antelope, snow leopard, and the wild boar. The Tibetan Mastiff's ability to avoid hypoxia in high altitudes, due to its higher hemoglobin levels compared to low-altitude dogs, was due to prehistoric interbreeding with the Tibetan wolf.[19][20]

Physical descriptionEdit

Hodgson described his type specimen as follows:

Lupus laniger. The Changu of the Tibetans. Hab. Tibet. Wolf with 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. No marks on the limbs. Tail concolorous with the body, that is brownish above and yellowish below, and no dark tip. Length 45 in (110 cm). Height 30 in (76 cm). This animal is found all over Tibet.[2]

The British naturalist Richard Lydekker wrote in 1900:

In order to withstand the intense winter cold of the bleak altitudes of which it dwells, and at the same time to harmonise with its physical surroundings, the wolf of Ladak and Tibet has developed a woolly character in its fur, and at the same time has become a much paler animal than ordinary examples of the European race.[21]

The British zoologist Thomas C. Jerdon wrote in 1874 that the Tibetan wolf is larger than the Indian Wolf and known as "chankodi" among the people who live near the Niti Pass on the Tibet/Indian border (Kumaon District, India).[22] Black wolves in Tibet are known locally as "chanko nagpo", and these are considered bolder and more aggressive than the pale coloured variety.[21]

The British naturalist William Thomas Blanford, writing in his The Fauna of British India, Including Ceylon and Burma in 1888, described the variety of wolf found in Ladakh in the Indian Himalayas and in Tibet as very pale-coloured, with woolly fur, and has generally been distinguished as Canis laniger.[23] There are no striking morphological differences between the wolves from the Indian Trans-Himalayas and those from Tibet.[24]

Distribution and habitatEdit

 
Tibetan wolf distribution (red dots in highlands) compared with the holarctic grey wolf (blue dots in lowlands)[6]

The Tibetan wolf is native to China in the regions of Gansu, Qinghai, and the Tibetan Autonomous Region.[25][26] Between 2005 and 2008, sightings and scat of Tibetan wolves were recorded in the alpine meadows above the tree line north-east of the Nanda Devi National Park in Uttarakhand, India.[27] In 2013, the media reported that a Tibetan wolf was photographed by a camera trap installed around 3,500 m (11,500 ft) altitude near the Sunderdhunga Glacier in Bageshwar district, Uttarakhand, India.[28]

Tibetan wolves, which generally occupy territories up to 3,000 above sea level, have evolved hearts that withstand the low oxygen levels. Specifically, these wolves have a strong selection for RYR2, a gene that initiates cardiac excitation.[29]

Relationship with humansEdit

 
A Tibetan wolf killed during the 1938 German expedition into Tibet. Note the proportionately short legs.

Historical sources indicate that wolves occasionally killed children in Ladakh and Lahoul.[30] Within the proposed Gya-Miru Wildlife Sanctuary in Ladakh, the intensity of livestock depredation was assessed in three villages. The assessment found that Tibetan wolves were the most important predators accounting for 60% of the total livestock losses, followed by the snow leopard and Eurasian lynx. The most frequent prey were domestic goats (32%), followed by sheep (30%), yaks (15%), and horses (13%). The wolves killed horses significantly more, and goats less, than would be expected from their relative abundance.[31]

ConservationEdit

China's wolf population largely lives in areas where little human-influenced change has occurred - the Qinghai–Tibet Plateau, the Mongolia Plateau, and the northeast Plain. In 2003, an estimated 12,500 wolves were living in China. In 2015, wolves were listed as a vulnerable species in the Red List of China’s Vertebrates, with all hunting being banned for this legally protected animal.[32][33]

NotesEdit

  1. ^ For a full set of supporting references refer to the note (a) in the phylotree at Evolution of the wolf#Wolf-like canids

ReferencesEdit

  1. ^ a b lupus filchneri Matschie. Wiss Ergebn Exped. Filchner mach China u. Tibet, 10 1, p153. Si-ning to the east from Kukunor (Siningfu, Kansu, China)
  2. ^ a b c Description of the wild ass and wolf of Tibet. Hodgson BH. (1847) Calcutta Journal of Natural History 7: 469–477
  3. ^ a b c d 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.
  4. ^ a b c d 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.
  5. ^ a b c d 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. PMC 5493914. PMID 28680672.
  6. ^ a b c d e f g h Werhahn, Geraldine; Senn, Helen; Ghazali, Muhammad; Karmacharya, Dibesh; Sherchan, Adarsh Man; Joshi, Jyoti; Kusi, Naresh; López-Bao, José Vincente; Rosen, Tanya; Kachel, Shannon; Sillero-Zubiri, Claudio; MacDonald, David W. (2018). "The unique genetic adaptation of the Himalayan wolf to high-altitudes and consequences for conservation". Global Ecology and Conservation. 16: e00455. doi:10.1016/j.gecco.2018.e00455.
  7. ^ Wissenschaftliche Ergebnisse Der Expedition Filchner Nach China Und Tibet Band.4 (1903-1905). by Filchner, Wilhelm. Published 1913
  8. ^ Allen, G.M. (1938). "Canis lupus chanco". The Mammals of China and Mongolia. XI (First ed.). The American Museum of Natural History, New York. p. 340. ISBN 978-1330410516.
  9. ^ Wozencraft, C. W. (2005). "Order Carnivora". In Wilson, D. E.; Reader, D. M. (eds.). Mammal Species of the World: A Taxonomic and Geographic Reference. 1 (3 ed.). Johns Hopkins University Press. ISBN 978-0-8018-8221-0.
  10. ^ "Canis lupus laniger".
  11. ^ "Canis lupus chanco".
  12. ^ a b 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.
  13. ^ Zhang, Honghai; Chen, Lei (2010). "The complete mitochondrial genome of dhole Cuon alpinus: Phylogenetic analysis and dating evolutionary divergence within canidae". Molecular Biology Reports. 38 (3): 1651–60. doi:10.1007/s11033-010-0276-y. PMID 20859694.
  14. ^ Li, Yinxia; Li, Qifa; Zhao, Xingbo; Xie, Zhuang; Xu, Yinxue (2010). "Complete sequence of the Tibetan Mastiff mitochondrial genome and its phylogenetic relationship with other Canids (Canis, Canidae)". Animal. 5 (1): 18–25. doi:10.1017/S1751731110001370. PMID 22440697.
  15. ^ 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.
  16. ^ Freedman, Adam H.; Gronau, Ilan; Schweizer, Rena M.; Ortega-Del Vecchyo, Diego; Han, Eunjung; Silva, Pedro M.; Galaverni, Marco; Fan, Zhenxin; Marx, Peter; Lorente-Galdos, Belen; Beale, Holly; Ramirez, Oscar; Hormozdiari, Farhad; Alkan, Can; Vilà, Carles; Squire, Kevin; Geffen, Eli; Kusak, Josip; Boyko, Adam R.; Parker, Heidi G.; Lee, Clarence; Tadigotla, Vasisht; Siepel, Adam; Bustamante, Carlos D.; Harkins, Timothy T.; Nelson, Stanley F.; Ostrander, Elaine A.; Marques-Bonet, Tomas; Wayne, Robert K.; et al. (2014). "Genome Sequencing Highlights the Dynamic Early History of Dogs". PLOS Genetics. 10 (1): e1004016. doi:10.1371/journal.pgen.1004016. PMC 3894170. PMID 24453982.
  17. ^ Wang, Guo-Dong; Zhai, Weiwei; Yang, He-Chuan; Wang, Lu; Zhong, Li; Liu, Yan-Hu; Fan, Ruo-Xi; Yin, Ting-Ting; Zhu, Chun-Ling; Poyarkov, Andrei D; Irwin, David M; Hytönen, Marjo K; Lohi, Hannes; Wu, Chung-I; Savolainen, Peter; Zhang, Ya-Ping (2015). "Out of southern East Asia: The natural history of domestic dogs across the world". Cell Research. 26 (1): 21–33. doi:10.1038/cr.2015.147. PMC 4816135. PMID 26667385.
  18. ^ Zhang, Wenping; Fan, Zhenxin; Han, Eunjung; Hou, Rong; Zhang, Liang; Galaverni, Marco; Huang, Jie; Liu, Hong; Silva, Pedro; Li, Peng; Pollinger, John P.; Du, Lianming; Zhang, Xiuyyue; Yue, Bisong; Wayne, Robert K.; Zhang, Zhihe (2014). "Hypoxia Adaptations in the Grey Wolf (Canis lupus chanco) from Qinghai-Tibet Plateau". PLOS Genetics. 10 (7): e1004466. doi:10.1371/journal.pgen.1004466. PMC 4117439. PMID 25078401.
  19. ^ Miao, Benpeng; Wang, Zhen; Li, Yixue (2016). "Genomic Analysis Reveals Hypoxia Adaptation in the Tibetan Mastiff by Introgression of the Grey Wolf from the Tibetan Plateau" (PDF). Molecular Biology and Evolution: msw274. doi:10.1093/molbev/msw274.
  20. ^ Signore, Anthony V.; Yang, Ying-Zhong; Yang, Quan-Yu; Qin, Ga; Moriyama, Hideaki; Ge, Ri-Li; Storz, Jay F. (2019). "Adaptive Changes in Hemoglobin Function in High-Altitude Tibetan Canids Were Derived via Gene Conversion and Introgression". Molecular Biology and Evolution. doi:10.1093/molbev/msz097. PMID 31362306.
  21. ^ a b Lydekker, R. (1900). The Tibetan Wolf. Pages 339–340 in: The great and small game of India, Burma, and Tibet. R. Ward, London.
  22. ^ Jerdon, T.C. (1874). The Indian wolf. Pages 140–141 in: The mammals of India: a natural history of all the animals known to inhabit continental India. John Wheldon, London.
  23. ^ Blanford, W. T. (1888). "Carnivora". In Blanford, W. T. (ed.). The Fauna of British India, Including Ceylon and Burma. Mammalia (1 ed.). Taylor and Francis. pp. 135–140.
  24. ^ 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.
  25. ^ 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.CS1 maint: uses editors parameter (link)
  26. ^ 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.
  27. ^ Bhattacharya, T. Sathyakumar (2010). "Sighting of Tibetan Wolf Canis lupus chanko in the Greater Himalayan range of Nanda Devi Biosphere Reserve, Uttarakhand, India: a new record" (PDF). Journal of Threatened Taxa. 2 (12): 1345–1348. doi:10.11609/jott.o2423.1345-8. Archived from the original (PDF) on 2012-11-01. NOTE: It was not chanco, it was the Tibetan wolf filchneri.
  28. ^ "Snow Leopard, Tibetan Wolf sighted". The Pioneer, 15 February 2014.
  29. ^ Zhang, W., Fan, Z., Han, E., Hou, R., Zhang, L., Galaverni, M., Liu, H., Silva, P., Li, P., Pollinger, J.P., Du, L. Zhang, X., Yue, B., Wayne, R.K., & Zhang, Z. (2014). "Hypoxia adaptations in the Grey wolf (Canis lupus chanco) from Qinghai-Tibet Plateau". PLOS Genetics. 10 (7): e1004466. doi:10.1371/journal.pgen.1004466. PMC 4117439. PMID 25078401.CS1 maint: uses authors parameter (link)
  30. ^ Pocock, R. I. (1941). Canis lupus chanco Pages 86–90 in: Fauna of British India: Mammals Volume 2. Taylor and Francis, London
  31. ^ Namgail, T.; Fox, J.L.; Bhatnagar, Y.V. (2007). "Carnivore-Caused Livestock Mortality in Trans-Himalaya". Environmental Management. 39 (4): 490–496. doi:10.1007/s00267-005-0178-2. PMID 17318699.
  32. ^ Xu, Yu; Yang, Biao; Dou, Liang (2015). "Local villagers' perceptions of wolves in Jiuzhaigou County, western China". PeerJ. 3: e982. doi:10.7717/peerj.982. PMC 4465947. PMID 26082870.
  33. ^ Fang, Hongxia; Ping, Xaiong (2016). "Red List of China's Vertebrates". Biodiversity Science. 24 (5): 500–551. doi:10.17520/biods.2016076.Canis lupus status=NT (Near Threatened)

External linksEdit