Canis lupus the Wolves

The wolf (Canis lupus), also known as the gray wolf or grey wolf, is a large canine native to Eurasia and North America. More than thirty subspecies of Canis lupus have been recognized, and gray wolves, as popularly understood, comprise wild subspecies. The wolf is the largest extant member of the family Canidae. It is also distinguished from other Canis species by its less pointed ears and muzzle, as well as a shorter torso and a longer tail. The wolf is nonetheless related closely enough to smaller Canis species, such as the coyote and the golden jackal, to produce fertile hybrids with them. The banded fur of a wolf is usually mottled white, brown, gray, and black, although subspecies in the arctic region may be nearly all white.

The Himalayan wolf (Canis lupus chanco) is a canine of debated taxonomy. It is distinguished by its genetic markers, with mitochondrial DNA indicating that it is genetically basal to the Holarctic gray wolf, genetically the same wolf as the Tibetan and Mongolian wolf, and has an association with the African wolf (Canis lupaster). No striking morphological differences are seen between the wolves from the Himalayas and those from Tibet. The Himalayan wolf lineage can be found living in Ladakh in the Himalayas, the Tibetan Plateau, and the mountains of Central Asia 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.

Some authors have proposed the reclassification of this lineage as a separate species. In 2019, a workshop hosted by the IUCN/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 as Canis lupus chanco until a genetic analysis of the holotypes is available. The Himalayan wolf lacks a proper morphological analysis. The wolves in India and Nepal are listed on CITES Appendix I as endangered due to international trade.

Canis himalayensis was proposed by Aggarwal et al. in 2007 for wolf specimens from the Indian Himalayas that differed in mitochondrial DNA from specimens collected in other parts of India. In April 2009, Canis himalayensis was proposed as a distinct wolf species 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 CITES 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. This genetic lineage shows a 3.9% divergence in the mDNA cytochrome b gene when compared with the Holarctic grey wolf, which may justify it being classified as a distinct species. In 2019, a workshop hosted by the IUCN/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 is Canis chanco Gray, 1863, but the geographic location of the holotype is unclear. The group recommends that this wolf lineage be known as the "Himalayan wolf" and classified as Canis lupus chanco until a genetic analysis of the holotypes is available.

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 an evolutionary significant unit that warranted assignment onto the IUCN Red List for its protection.

The mitochondrial DNA of 27 wolves from the Himalayas and the Tibetan Plateau was compared in 2004. Results indicate that five related haplotypes formed a clade that is basal to all other wolves. This clade included one sample from Ladakh, nine from the Spiti Valley in Himachal Pradesh, four from Nepal, and two from Tibet. The Himalayan wolf clade diverged from other canids 800,000 years ago. Seven wolves from Kashmir did not fall into this clade. The mtDNA of 18 captive wolves in the Padmaja Naidu Himalayan Zoological Park was analysed in 2007. Results showed that they shared a common female ancestor. 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. Subsequent genetic research showed that wolf samples from Tibet are genetically basal to the Holarctic gray wolf. Its MT-ND4L gene commences with the base pairs GTG, whereas all other canids commence with ATG. Results of whole genome sequencing showed that it is the most genetically divergent wolf.

Analysis of scat samples from two wolves collected in upper Dolpo in Nepal matched the Himalayan wolf. Fecal remains of four wolves collected in the upper Mustang region of the Annapurna Conservation Area also fell within the Himalayan wolf clade but formed a separate haplotype from those previously studied.

The Himalayan wolf population in Tibet declined over the past 25,000 years and suffered a historical population bottleneck. Glaciation during the Last Glacial Maximum may have caused habitat loss, genetic isolation, and ancient inbreeding. The population in Qinghai had grown, though, showing a gene flow of 16% from Chinese indigenous dogs and 2% of the dingo's genome. It probably recolonised the Tibetan Plateau. The Himalayan wolf contrasts with the wolves living at lower elevations in Inner Mongolia, Mongolia, and Xinjiang province. Some wolves in China and Mongolia also fall within the Himalayan wolf clade, indicating a common maternal ancestor and a wide distribution. 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.

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 as Jiangxi province shows evidence of being admixed between Tibetan-related wolves and other wolves in China.

DNA sequences can be mapped to reveal a phylogenetic 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.

Relationship to the Indian lowland wolf

In 2021, a study compared both the mitochondrial DNA and the nuclear DNA (from the cell nucleus) from the wolves of the Himalayas with those of the wolves from the lowlands of the Indian 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 mitochondrial clade, 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.

Admixture with an unknown wolf-like canid

The Tibetan mastiff breed was able to adapt to the extreme highland conditions of the Tibetan Plateau very quickly, comparably to other mammals such as the yak, the Tibetan antelope, the snow leopard, and the wild boar. The Tibetan mastiff's ability to avoid hypoxia in high elevations due to its higher hemoglobin levels compared to low-altitude dogs, was due to prehistoric interbreeding with the wolves of Tibet.

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 grey wolves, dogs, and a ghost 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 the EPAS1 allele which can be found in both Himalayan wolves and dogs which allows them to live in high altitudes.

Domestic dogs exhibit diverse coat colours and patterns. In many mammals, different colour patterns are the result of the regulation of the Agouti 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 is agouti, 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.

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 genetic introgression into the Late Pleistocene grey wolf from a ghost 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.

Relationship with the African golden wolf

In 2011, the Indian wolf, Himalayan wolf, and African wolf 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.

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 were radiocarbon dated and stratigraphically dated, and together with DNA sequences, a time-based phylogenetic tree was generated for wolves. The study inferred that the most recent common ancestor for all other Canis lupus specimens – modern and extinct – was 80,000 years before present. 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.

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. From 2017, two studies based on mDNA, and X-chromosome and Y-chromosome markers taken from the cell 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. The results of these two studies imply that the Himalayan wolf distribution range extends from the Himalayan range north across the Tibetan Plateau up to the Qinghai Lake region in China’s Qinghai Province.

In 2018, whole genome sequencing was used to compare members of the genus Canis. The African golden wolf was found to be the descendant of a genetically admixed canid of 72% gray wolf and 28% Ethiopian wolf ancestry. 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 other single-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.

Distribution and habitat

Pin Valley National Park located in Himachal Pradesh In China, the Himalayan wolf lives on the Tibetan Plateau in the provinces of Gansu, Qinghai, Tibet,[48][49] and western Sichuan. In northern India, it occurs in the Union Territory of Ladakh and in the Lahaul and Spiti region in northeastern Himachal Pradesh. 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). Between 2005 and 2008, it was sighted in the alpine meadows above the treeline northeast of Nanda Devi National Park in Uttarakhand.[50] 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's Bageshwar district. In Nepal, it was recorded in Api Nampa Conservation Area, Upper Dolpa, Humla, Manaslu, Upper Mustang, and the Kanchenjunga Conservation Area.[52][53] The Nepal Himalayas provide an important habitat refuge for the Himalayan wolf.

Behaviour and ecology

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.

Prey

Himalayan wolves prefer wild over domestic prey. They prefer the smaller Tibetan gazelle over the larger white-lipped deer, and they prefer the plains-dwelling Tibetan gazelle over the cliff-dwelling bharal. Supplementary food includes the small Himalayan marmot, woolly hare, and pikas. 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. Kiang, Siberian roe deer, Siberian ibex, Przewalski's horse, wild yak, argali, urial, markhor, Bactrian deer, Yarkand deer, and Tibetan red deer have also been recorded as prey species of Himalayan wolves.

Historical sources indicate that wolves occasionally killed children in Ladakh and Lahaul.[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 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.

 Tibetan Wolf (Canis lupus chanco) - seen in many locations in Qinghai Province on the Tibetan Plateau

Himalayan Wolf (Canis lupus himalayensis) - seen several in Hemis, Ladakh India. Pure breeding individuals and clear hybrids were seen in this region as well. At the current time, there seems to be a bit of confusion on these wolves. Clearly there is hybridization seen in some individuals in the below gallery. Ladakhi experts claimed these were different subspecies than the Tibetan wolves in the above gallery and a quick observation can make this obvious. I am keeping them with this nomenclature for now until there is more known about the true genetics and relationships between these very different populations.

The Arabian wolf (Canis lupus arabs) is a subspecies of gray wolf native to the Arabian Peninsula, the Negev Desert, the Sinai Peninsula, and Jordan. It is the smallest wolf subspecies, and a desert-adapted subspecies that normally lives in small groups. It is omnivorous, eating small to medium-sized prey.

Once thought to be synonymous with C. l. pallipes (the Indian wolf), the Arabian wolf was designated Canis lupus arabs by the British zoologist Reginald Innes Pocock in 1934. Pocock noted its smaller skull and smaller size. 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 subspecies Canis lupus arabs. A 2014 study suggests that genetically the Arabian wolf is closer to C. l. lupus than it is to C. l. pallipes and supports the subspecies designation C. l. arabs. There has been admixture with domestic dogs, but it is unclear whether or not this is why this wolf is genetically closer to C. l. lupus. This raises a concern of extinction by hybridization as Arabian wolves are more adapted to desert life than wolf/dog hybrids.

In Israel and Palestine, there is some disagreement as to the exact taxonomic status of wolves. Some scientists hold that two subspecies of wolf are present- C. l. pallipes in the northern parts, and C. l. arabs in the south. They point out that the southern wolves are smaller than the northern wolves which are also darker and have longer fur. Other scientists consider the wolf in the area to be C. l. arabs, with no real distinction between northern and southern wolves. As in other countries, there is interbreeding with feral dogs, which adds an element of uncertainty.

Admixture with other Canis species

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.

The Arabian wolf is small for a wolf. It stands on average 25–26 inches (64–66 cm) at shoulder height and the adult weighs an average of 45 pounds (20.41 kg).[13] The cranial length of the adult Arabian wolf measures on average 200.8 mm (0.659 ft), which is smaller than most wolves. Along with the Indian wolf, it is probably smaller than other wolves to help it adapt to life in a hot, dry climate. This is an example of Bergmann's rule, where mammal size varies by the warmth of their environment. Its ears are proportionally larger in relation to its body size when compared to other sub-species of Canis lupus, an adaptation probably developed to help disperse body heat (Allen's Rule). Arabian wolves do not usually live in large packs, and instead hunt in pairs or in groups of about three or four animals. They have a short thin coat which is usually a grayish beige color, "... a mixture of black and slightly buffy grey" according to Pocock.[5] Similar to other canines, the Arabian wolf does not have sweat glands and so it must control its body temperature by rapid panting, which causes evaporation from the lungs.

Arabian wolves are mainly carnivorous, but also omnivorous and in some areas largely dependent on human garbage and excess products. They also feed on hares, rodents, small ungulates, cats, sweet fruits, roadkill and other carrion. Opportunistically almost any small animal including fish, snails, baby baboons can be part of their diet. Because Arabian wolves can attack and eat any domestic animals up to the size of a goat, Bedouins and farmers will not hesitate to shoot, poison, or trap them.

The Arabian wolf was once found throughout the Arabian Peninsula, but now lives only in small pockets in southern Israel, southern and western Iraq, Oman, Yemen, Jordan, Saudi Arabia, and some parts of the Sinai Peninsula in Egypt. It is rare throughout most of its range because of human persecution. In Oman, wolf populations have increased because of a ban on hunting, and they may naturally re-establish themselves in certain places within the region in the relatively near term. In Israel, between 100 and 150 Arabian wolves are found across the Negev and the Arava. The United Arab Emirates and Egypt both have a captive breeding program, and the wolf is protected in Oman and Israel, but elsewhere, its future is uncertain. In Saudi Arabia, the wolf is protected in places and still exists in places with sparse human activity.

Arabian Wolf (Canis lupus arabs) - Seen wild in Jordan (first three images in the below gallery) at rest stop and captive at Shaumari Reserve

Mexican Wolf (Canis lupus baileyi) - captive pair seen in the Sonoran-Desert Museum in Tucson, Arizona

Domestic Dogs (Canis lupus familiaris) - Tibetan Mastiff, Hokkaido Dog, a smaller Japanese breed and a couple third-world mutts