Himalayan wolf

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In the Upper Mustang region of the Annapurna Conservation Area, Nepal

The Himalayan wolf (Canis lupus) is a canine of debated taxonomy.[1] It is distinguished by its genetic markers, with mitochondrial DNA sequencing indicating that it is genetically basal to the Holarctic grey wolf, genetically the same wolf as the Tibetan wolf and has an association with the African golden wolf (Canis anthus).[2][3][1] The 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.

Taxonomy[edit]

The wolves of the Himalayas have been referred to as Canis laniger Hodgson 1847, Canis chanco Gray 1863, Canis filchneri Matschie 1907, and Canis himalayensis Aggawal 2003.[1]

A female wolf from Tibet was described by Brian Houghton Hodgson in 1847 as Lupus laniger.[4] In 1907, the German zoologist Paul Matschie described a wolf skin from Xining in China as Lupus filchneri.[5] In 1941, Reginald Innes Pocock referred to the Tibetan wolf as Canis lupus laniger and classified it as a taxonomic synonym of C. l. chanco.[6] 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).[7]

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[8][9] or Canis lupus himalayensis.[10] are proposed.

The taxonomic reference Mammal Species of the World does not recognize Canis himalayensis, however NCBI/Genbank does list Canis lupus himalayensis.[11]

Lineage[edit]

Phylogenetic tree of the extant wolf-like canids
Caninae 3.5 Ma
3.0
2.7
1.9
1.6
1.3
1.1
0.8

Dog Tibetan mastiff (white background).jpg

Holarctic grey wolf Dogs, jackals, wolves, and foxes (Plate I).jpg

Himalayan wolf Dogs, jackals, wolves, and foxes (Plate I).jpg

Coyote Dogs, jackals, wolves, and foxes (Plate IX).jpg

African golden wolf Dogs, jackals, wolves, and foxes (Plate XI).jpg

Ethiopian wolf Dogs, jackals, wolves, and foxes (Plate VI).jpg

Golden jackal Dogs, jackals, wolves, and foxes (Plate X).jpg

Dhole Dogs, jackals, wolves, and foxes (Plate XLI).jpg

African wild dog Dogs, jackals, wolves, and foxes (Plate XLIV).jpg

2.6

Side-striped jackal Dogs, jackals, wolves, and foxes (Plate XIII).jpg

Black-backed jackal Dogs, jackals, wolves, and foxes (Plate XII).jpg

Phylogenetic relationships between the extant wolf-like canids based on DNA taken from the cell nucleus,[12][13] except for the Himalayan wolf that is based on mitochondrial DNA sequences[13][3] plus X chromosome and Y chromosome sequences.[3][1] Timing in millions of years.[13]

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.[15]

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.[15] However, the 50% likelihood provided by the model on which this study rested provides only weak support for this proposal.[16][8][15] 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.[17][16] Additionally, the wolf population in the Kashmir valley is known to have recently arrived in that area.[16]

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.[8] 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.[16]

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.[16][18] The committee recommended against this proposal but suggested that the name be entered into the species database as a synonym for Canis lupus.[19][20] The committee stated that the classification was for conservation purposes only, and did not "reflect the latest state of taxonomic knowledge".[20] Further fieldwork was called for.[16]

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.[16]

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.[21][22][23] 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.[18] 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.[17]

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,[2] indicating a common maternal ancestor[a] and a wider genetic distribution.[2] In 2017 and 2018, another two studies found the Himalayan wolf to be the same wolf as the Tibetan wolf.[3][1]

Relationship with the African golden wolf[edit]

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

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.[24][13] 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.[3]

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.[3][25][16][18][20][19] 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.[17]

In 2018, whole genome sequencing was used to compare members of the genus Canis. The study found evidence of gene flow between African golden wolves, golden jackals, and grey wolves (from Saudi Arabia and Syria). One African golden wolf from the Egyptian Sinai Peninsula showed high admixture with the Middle Eastern grey wolves and dogs, highlighting the role of the land bridge between the African and Eurasian continents in canid evolution. The African golden wolf was found to be the descendant of a genetically admixed canid of 72% grey wolf and 28% Ethiopian wolf ancestry.[26]

Description[edit]

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

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.[4]

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.[27]

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).[28] Black wolves in Tibet are known locally as "chanko nagpo", and these are considered bolder and more aggressive than the pale coloured variety.[27]

Himalayan wolf profile

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.[16]

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.[17]

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.[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.[29]

Distribution[edit]

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

The wolf is found in northern India in the Ladakh region of eastern Kashmir[27] and the Lahaul and Spiti region in the northeastern part of Himachal Pradesh.[15][6] It is also found in Nepal in the Upper Dolpa[18] and Upper Mustang regions.[30]

The Indian population consists of 350 wolves with a range of 70,000 km2 (27,000 sq mi).[31] In 2004, a group of 33 Himalayan wolves were spotted in the Spiti Valley in the northeastern part of Himachal Pradesh.[32] In the same year, the wolf was spotted for the first time in Nepal in the Upper Mustang region.[30] 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.[33] In 2013, the media reported that a Tibetan wolf was photographed by a camera trap installed at about 3,500 m (11,500 ft) altitude near the Sunderdhunga Glacier in Bageshwar district, Uttarakhand, India.[34]

Conservation[edit]

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.[16] 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.[25]

It has been suggested by several biologists in India for recognition as a critically endangered species of canid.[15] 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.[31]

See also[edit]

Notes[edit]

  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."[14]
  3. ^ For the purpose of this study the "Himalayas" classification included specimens from Tibet as well as from the Himalayas.

References[edit]

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