| Himalayan wolf | |
|---|---|
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| Himalayan wolf in theUpper Mustang region ofAnnapurna Conservation Area in Nepal | |
Scientific classification | |
| Domain: | Eukaryota |
| Kingdom: | Animalia |
| Phylum: | Chordata |
| Class: | Mammalia |
| Order: | Carnivora |
| Family: | Canidae |
| Genus: | Canis |
| Species: | |
| Subspecies: | C. l. chanco |
| Trinomial name | |
| Canis lupus chanco | |
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| Himalayan wolf distribution (red dots in highlands) compared with the holarctic grey wolf (blue dots in lowlands)[3] | |
TheHimalayan wolf (Canis lupus chanco) is acanine of debated taxonomy.[3] It is distinguished by itsgenetic markers, withmitochondrial DNA indicating that it is geneticallybasal to theHolarcticgrey wolf, genetically the same wolf as the Tibetan andMongolian wolf,[4][5][3] and has an association with theAfrican wolf (Canis lupaster).[6][5][3] No striking morphological differences are seen between the wolves from the Himalayas and those from Tibet.[7] The Himalayan wolf lineage can be found living inLadakh in theHimalayas, theTibetan Plateau,[8][9] and themountains of Central Asia[9] predominantly above 4,000 m (13,000 ft) in elevation because it has adapted to a low-oxygen environment, compared with other wolves that are found only at lower elevations.[8]
Some authors have proposed the reclassification of this lineage as a separate species.[10][11] In 2019, a workshop hosted by theIUCN/SSC Canid Specialist Group noted that the Himalayan wolf's distribution included the Himalayan range and the Tibetan Plateau. The group recommends that this wolf lineage be known as the "Himalayan wolf" and be classified asCanis lupus chanco until a genetic analysis of theholotypes is available. The Himalayan wolf lacks a proper morphological analysis.[12] The wolves in India and Nepal are listed onCITES Appendix I as endangered due to international trade.[13]
Canis chanco was thescientific name proposed byJohn Edward Gray in 1863, who described a skin of a wolf that was shot inChinese Tartary.[2] This specimen was classified as a wolfsubspeciesCanis lupus chanco bySt. George Jackson Mivart in 1880.[14] In the 19th and 20th centuries, severalzoological specimens were described:
In 1938,Glover Morrill Allen classified these specimens assynonyms forC. l. chanco.[19] In 1941,Reginald Pocock corroborated this assessment after reviewing wolf skins and skulls in the collection of theNatural History Museum, London.[20] In 2005,W. Christopher Wozencraft also listedC. l. niger,C. l. filchneri,C. l. karanorensis, andC. l. tschiliensis as synonyms forC. l. chanco.[21]
Canis himalayensis was proposed by Aggarwalet al. in 2007 for wolf specimens from the IndianHimalayas that differed inmitochondrial DNA from specimens collected in other parts of India.[10] In April 2009,Canis himalayensis was proposed as a distinct wolf species through the Nomenclature Specialist on theCITES 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.[7][22] The committee recommended against this proposal, but suggested that the name be entered into the CITES species database as a synonym forCanis lupus. The committee stated that the classification was for conservation purposes only, and did not "reflect the latest state of taxonomic knowledge".[23][24] Further fieldwork was called for.[7] Thisgenetic lineage shows a 3.9% divergence in the mDNAcytochrome b gene when compared with the Holarctic grey wolf, which may justify it being classified as a distinct species.[3] In 2019, a workshop hosted by theIUCN/SSC Canid Specialist Group noted that the Himalayan wolf's distribution included the Himalayan range and the Tibetan Plateau. The group determined that the earliest available Latin name isCanis chanco Gray, 1863, but the geographic location of theholotype is unclear. The group recommends that this wolf lineage be known as the "Himalayan wolf" and classified asCanis lupus chanco until a genetic analysis of the holotypes is available.[12]
In 2020, more recent research on the Himalayan wolf genome indicates that it warrants species-level recognition under the Unified Species Concept, the Differential Fitness Species Concept, and the Biological Species Concept. It was identified as anevolutionary significant unit that warranted assignment onto theIUCN Red List for its protection.[8]
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The Himalayan wolf has a thick, woolly fur that is dull earthy-brown on the back and tail, and yellowish-white on the face, tummy, and limbs. It has closely spaced black speckles on the muzzle, below the eyes, and on the upper cheeks and ears.[25]
Larger than theIndian andcommon European wolves,[26][27] it reaches 110 to 180 cm (45 to 70 in) in length, 68 to 76 cm (27 to 30 in) tall at the shoulder, and weighs around 30 to 55 kg (66 to 121 lb) on average.[7][20][28]
The heart of the Himalayan wolf can withstand the low oxygen level at high elevations. It has a strong selection forRYR2, a gene that initiates cardiac excitation.[29]
The mitochondrial DNA of 27 wolves from theHimalayas and theTibetan Plateau was compared in 2004. Results indicate that five relatedhaplotypes formed aclade that isbasal to all other wolves. This clade included one sample fromLadakh, nine from theSpiti Valley inHimachal Pradesh, four fromNepal, and two fromTibet. The Himalayan wolf clade diverged from other canids 800,000 years ago. Seven wolves fromKashmir did not fall into this clade.[30] The mtDNA of 18 captive wolves in thePadmaja Naidu Himalayan Zoological Park was analysed in 2007. Results showed that they shared a common female ancestor.[10] As this study was based on captive-bred zoo specimens that had descended from only two females, these samples were not considered to be representative. Additionally, the wolf population in the Kashmir Valley is known to have recently arrived in that area.[7][25] Subsequent genetic research showed that wolf samples from Tibet aregenetically basal to the Holarctic gray wolf.[31][32][33][34] ItsMT-ND4Lgene commences with thebase pairsGTG, whereas all othercanids commence withATG.[35] Results ofwhole genome sequencing showed that it is the mostgenetically divergent wolf.[36]
Analysis of scat samples from two wolves collected in upperDolpo in Nepal matched the Himalayan wolf.[22] Fecal remains of four wolves collected in theupper Mustang region of theAnnapurna Conservation Area also fell within the Himalayan wolf clade but formed a separate haplotype from those previously studied.[25]
The Himalayan wolf population in Tibet declined over the past 25,000 years and suffered a historicalpopulation bottleneck. Glaciation during theLast Glacial Maximum may have caused habitat loss, genetic isolation, and ancient inbreeding. The population in Qinghai had grown, though, showing agene flow of 16% from Chinese indigenous dogs and 2% of the dingo's genome. It probably recolonised the Tibetan Plateau.[36] The Himalayan wolf contrasts with the wolves living at lower elevations inInner Mongolia,Mongolia, andXinjiang province. Some wolves in China and Mongolia also fall within the Himalayan wolf clade, indicating a common maternal ancestor and a wide distribution.[4] There was evidence ofhybridization with the grey wolf at Sachyat-Ertash in theIssyk-Kul region ofKyrgyzstan, and ofintrogression from either the grey wolf or the dog into the Himalayan wolf in Nepal.[3]
A genomic study on China's wolves included museum specimens of wolves from southern China that were collected between 1963 and 1988. The wolves in the study formed three clades: north Asian wolves that included those from northern China and eastern Russia, wolves from the Tibetan Plateau, and a unique population from southern China. One specimen located as far southeast asJiangxi province shows evidence of being admixed between Tibetan-related wolves and other wolves in China.[37]
| Phylogenetic tree ofCanis lupus with timing in years[a] | |||||||||||||||||||||
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DNA sequences can be mapped to reveal aphylogenetic tree that represents evolutionary relationships, with each branch point representing the divergence of two lineages from a common ancestor. On this tree, the term“basal” is used to describe a lineage that forms a branch diverging nearest to the common ancestor.[38]
In 2021, a study compared both themitochondrial DNA and thenuclear DNA (from the cell nucleus) from the wolves of the Himalayas with those of the wolves from the lowlands of theIndian subcontinent. The genomic analyses indicate that the Himalayan wolf and the Indian lowland wolf were genetically distinct from one another. These wolves were also genetically distinct from – and genetically basal to – the other wolf populations across the northern hemisphere. These other wolves form a single mitochondrialclade, indicating that they originated from a single expansion from one region within the last 100,000 years. However, the study indicated that the Himalayan wolf had separated from this lineage 496,000 years ago, and the Indian lowland wolf 200,000 years ago.[39]
TheTibetan mastiff breed was able to adapt to the extreme highland conditions of the Tibetan Plateau very quickly, comparably to other mammals such as theyak,Tibetan antelope,snow leopard, andwild boar. The Tibetan mastiff's ability to avoidhypoxia in high elevations due to its higherhemoglobin levels compared to low-altitude dogs, was due to prehistoricinterbreeding with the wolves of Tibet.[40][41]
In 2020, a genomic analysis indicates that the wolves of the Himalayas and the Tibetan plateau are closely related. These wolves have an admixed history which includes gray wolves, dogs, and aghost population of an unknown wolf-like canid. This ghost population is deeply-diverged from modern Holarctic wolves and dogs, has contributed 39% to the Himalayan wolf's nuclear genome, and contributed theEPAS1 allele which can be found in both Himalayan wolves and dogs which allows them to live in high altitudes.[42]
Domestic dogs exhibit diversecoat colours and patterns. In many mammals, different colour patterns are the result of the regulation of theAgouti gene, which can cause hair follicles to switch from making black or brown pigments to yellow or nearly white pigments. The most common coat pattern found in modern wolves isagouti, in which the upperside of the body has banded hairs and the underside exhibits lighter shading. The colour yellow is dominant to the colour black and is found in dogs across much of the world and the dingo in Australia.[43]
In 2021, a study of whole genome sequences taken from dogs and wolves focused on the genetic relationships between them based on coat colour. The study found that most dog colour haplotypes were similar to most wolf haplotypes, however dominant yellow in dogs was closely related to white in arctic wolves from North America. This result suggests a common origin for dominant yellow in dogs and white in wolves but without recent gene flow, because this clade was found to be basal to the golden jackal and genetically distinct from all other canids. The most recent common ancestor of the golden jackal and the wolf lineage dates back to 2 million YBP. The study proposes that 35,000 YBP there was geneticintrogression into the Late Pleistocene grey wolf from aghost population of an extinct canid which had diverged from the grey wolf lineage over 2 million YBP. This colour diversity could be found 35,000 YBP in wolves and 9,500 YBP in dogs. A closely related haplotype exists among those wolves of Tibet which possess yellow shading in their coats. The study explains the colour relationships between modern dogs and wolves, white wolves from North America, yellow dogs, and yellowish wolves from Tibet. The study concludes that during the Late Pleistocene, natural selection laid the genetic foundation for modern coat colour diversity in dogs and wolves.[43]
In 2011, the Himalayan, Indian and African wolves were proposed to represent ancient wolf lineages, with the African wolf having colonised Africa prior to the Northern Hemisphere radiation of the Holarctic gray wolf.[33]
Two studies of the mitochondrial genome of both modern and extinct gray wolves (Canis lupus) have been conducted, but these excluded the genetically divergent lineages of the Himalayan wolf and the Indian wolf. The ancient specimens wereradiocarbon dated andstratigraphically dated, and together with DNA sequences, a time-basedphylogenetic tree was generated for wolves. The study inferred that themost recent common ancestor for all otherCanis lupus specimens – modern and extinct – was 80,000 years before present.[44][45] An analysis of the Himalayan wolf mitochondrial genome indicates that the Himalayan wolf diverged between 740,000 and 691,000 years ago from the lineage that would become the Holarctic gray wolf.[3]
Between 2011 and 2015, two mDNA studies found that the Himalayan wolf and Indian gray wolf were genetically closer to the African golden wolf than they were to the Holarctic gray wolf.[33][6] From 2017, two studies based on mDNA, and X-chromosome and Y-chromosome markers taken from thecell nucleus, indicate that the Himalayan wolf is genetically basal to the Holarctic gray wolf. Its degree of divergence from the Holarctic gray wolf is similar to the degree of divergence of the African wolf from the Holarctic wolf. The Himalayan wolf shares a maternal lineage with the African wolf. It possesses a unique paternal lineage that falls between the gray wolf and the African wolf.[5][3] The results of these two studies imply that the Himalayanwolf distribution range extends from the Himalayan range north across the Tibetan Plateau up to theQinghai Lake region in China’s Qinghai Province.[5]
In 2018, whole genome sequencing was used to compare members of the genusCanis. The African golden wolf was found to be the descendant of a geneticallyadmixed canid of 72% gray wolf and 28% Ethiopian wolf ancestry.[46] The Ethiopian wolf does not share the single-nucleotide polymorphisms that confer hypoxia adaptation with the Himalayan wolf. The adaptation of the Ethiopian wolf to living in high elevations may occur at othersingle-nucleotide polymorphism locations. This indicates that the Ethiopian wolf's adaptation has not been inherited by descent from a common ancestor shared with the Himalayan wolf.[3]
In China, the Himalayan wolf lives on the Tibetan Plateau in the provinces ofGansu,Qinghai, Tibet,[47][48] and westernSichuan.[8]
In northernIndia, it occurs in the Union Territory ofLadakh and in theLahaul and Spiti region in northeasternHimachal Pradesh.[20] In 2004, the Himalayan wolf population in India was estimated to consist of 350 individuals ranging across an area of about 70,000 km2 (27,000 sq mi).[30]Between 2005 and 2008, it was sighted in the alpine meadows above the treeline northeast ofNanda Devi National Park inUttarakhand.[49] In 2013, a wolf was photographed by a camera trap installed at an elevation around 3,500 m (11,500 ft) near the Sunderdhunga Glacier in Uttarakhand'sBageshwar district.[50]
In Nepal, it was recorded inApi Nampa Conservation Area, UpperDolpa,Humla,Manaslu,Upper Mustang, and theKanchenjunga Conservation Area.[51][52] The Nepal Himalayas provide an important habitat refuge for the Himalayan wolf.[3]
The howls of the Himalayan wolf have lower frequencies, unmodulated frequencies, and are shorter in duration compared to Holarctic wolf howls. The Himalayan and North African wolves have the most acoustically distinct howls and differ significantly from each other and the Holarctic wolves.[53]
Himalayan wolves prefer wild over domestic prey. It usually prefers the smallerTibetan gazelle over the largerwhite-lipped deer, and the plains-dwelling Tibetan gazelle over the cliff-dwellingblue sheep. Supplementary diet includes the smallHimalayan marmot,big-eared pika andwoolly hare. Himalayan wolves avoid livestock where wild prey is available, but habitat encroachment and the depletion of wild prey populations is expected to lead to conflict with herders. To protect them, securing healthy wild prey populations through setting aside wildlife habitat reserves and refuges is essential.[54] Other recorded prey species areBactrian deer,Yarkand deer,Tibetan red deer,Siberian roe deer,Siberian ibex,Tibetan wild ass,Przewalski's horse,wild yak,markhor,argali andurial.[55]
Historical sources indicate that wolves occasionally killed children in Ladakh andLahaul.[20] Within the proposed Gya-Miru Wildlife Sanctuary in Ladakh, the intensity of livestock depredation assessed in three villages found that Tibetan wolves were the most prevalent predators, accounting for 60% of the total livestock losses, followed by the snow leopard and Eurasian lynx. The most frequent prey weredomestic goats (32%), followed by sheep (30%),yaks (15%), andhorses (13%). The wolves killed horses significantly more, and goats less, than would be expected from their relative abundance.[56]
The wolf in Bhutan, India, Nepal, and Pakistan is listed onCITES Appendix I.[13]In India, the wolf is protected under Schedule I of theWildlife Protection Act, 1972, which prohibits hunting; a zoo needs a permission from the government to acquire a wolf. It is listed as endangered inJammu and Kashmir, Himachal Pradesh, and Uttarakhand, where a large portion of the wolf population lives outside the protected area network.[7] Lack of information about its basic ecology in this landscape is an obstacle for developing a conservation plan.[57] In Nepal, it is protected under Schedule I of theNational Parks and Wildlife Conservation Act, 2029 (1973) prohibiting hunting it.[58] In China, the wolf is listed as vulnerable in theRed List of China’s Vertebrates, and hunting it is banned.[59][60]
In 2007, 18 Himalayan wolves were kept for breeding in two Indian zoos. They were captured in the wild and were kept at the Padmaja Naidu Himalayan Zoological Park inWest Bengal, and in theKufri Zoo in Himachal Pradesh.[10]
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