The dhole is a highly social animal, living in large clans without rigid dominance hierarchies[11] and containing multiple breeding females.[12] Such clans usually consist of 12 individuals, but groups of over 40 are known.[5] It is a diurnal pack hunter which preferentially targets medium and large sized ungulates.[13] In tropical forests, the dhole competes with tigers and leopards, targeting somewhat different prey species, but still with substantial dietary overlap.[14]
It is listed as Endangered by the IUCN as populations are decreasing and are estimated at fewer than 2,500 adults. Factors contributing to this decline include habitat loss, loss of prey, competition with other species, persecution due to livestock predation and disease transfer from domestic dogs.[2]
An Ussuri dhole (Cuon alpinus alpinus
Etymology and naming
Discovery, taxonomy, and evolution
Phylogenetic tree of the extant wolf-like canids | ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
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Phylogenetic relationships between the extant wolf-like clade of canids based on nuclear DNA sequence data taken from the cell nucleus,[7][42] except for the Himalayan wolf, based on mitochondrial DNA sequences.[42][43] Timing in millions of years.[42] |
Subspecies
Historically, up to 10 subspecies of dholes have been recognised.[44] As of 2005, only three subspecies are recognised by MSW3.[1]Subspecies | Image | Trinomial authority | Common names | Description | Range | Synonyms |
---|---|---|---|---|---|---|
C. a. alpinus (Nominate subspecies) | Pallas, 1811 | Indian wild dog Southern dhole[45] Ussuri dhole[9] |
Large subspecies with bright red coat and narrow skull.[9] | Far eastern Russia, Mongolia, China, Nepal, Indian subcontinent, Bhutan, Burma, Indochina and Java. | adustus (Pocock, 1941), antiquus (Matthew & Granger, 1923), clamitans (Heude, 1892), dukhunensis (Sykes, 1831), fumosus (Pocock, 1936), grayiformis (Hodgson, 1863), infuscus (Pocock, 1936), javanicus (Desmarest, 1820), laniger (Pocock, 1936), lepturus (Heude, 1892), primaevus (Hodgson, 1833), rutilans (Müller, 1839) | |
C. a. hesperius | Afanasjev and Zolotarev, 1935 | Northern dhole[45] Tien Shan dhole[9] |
Smaller than C. a. alpinus, with wider skull and lighter-coloured winter fur[9] | Altai, Tien Shan and possibly Pamir and Kashmir | jason (Pocock, 1936) | |
C. a. sumatrensis | Hardwicke, 1821 | Sumatran dhole | Has short, coarse fur with no woolly underfur, and much black on the back[45] | Sumatra |
Characteristics
Dholes produce whistles resembling the calls of red foxes, sometimes rendered as coo-coo. How this sound is produced is unknown, though it is thought to help in coordinating the pack when travelling through thick brush. When attacking prey, they emit screaming KaKaKaKAA sounds.[54] Other sounds include whines (food soliciting), growls (warning), screams, chatterings (both of which are alarm calls) and yapping cries.[55] In contrast to wolves, dholes do not howl or bark.[9] Dholes have a complex body language. Friendly or submissive greetings are accompanied by horizontal lip retraction and the lowering of the tail, as well as licking. Playful dholes open their mouths with their lips retracted and their tails held in a vertical position whilst assuming a play bow. Aggressive or threatening dholes pucker their lips forward in a snarl and raise the hairs on their backs, as well as keep their tails horizontal or vertical. When afraid, they pull their lips back horizontally with their tails tucked and their ears flat against the skull.[56]
Distribution and habitat
The dhole might still be present in the Tunkinsky National Park in extreme southern Siberia near Lake Baikal.[57] It possibly still exists in the Primorsky Krai province in far-eastern Russia, where it was considered a rare and endangered species in 2004, with unconfirmed reports in the Pikthsa-Tigrovy Dom protected forest area; no sighting was reported in other areas such as the Mataisky Zakaznik forest since the late 1970s.[58]
Currently, no other recent reports are confirmed of dhole being present in Russia, with no recent reports from Mongolia, Kazakhstan, Kyrgyzstan or Tajikistan, though one specimen was caught in southern China's Jiangxi district. Also, in 2011 to 2013, local government officials and herders reported the presence of several packs at altitudes of 2,000 to 3,500 m near the Taxkorgan Reserve in the Karakoram/Pamir Mountains region of the Xinjiang Autonomous Region, on China's border with Pakistan, Tajikistan, and Afghanistan.[59]
Dholes have been also recently reported from the Altyn-Tagh (Altun) Mountains in the southern portion of the Xinjiang Autonomous Region close to Tibet, as well.[60] It is unknown if dholes continue to inhabit Tien Shan, though they occur in small numbers in Gansu Province, with one pack being sighted in the Qilian Mountains within that province in 2006;[61] Camera-trap surveys in the Yanchiwan National Nature Reserve in the northern edge of this Gansu Province in 2013-2014 confirmed the continued presence of several packs and a female adult with pups in this area at altitudes around 2,500 to 4,000 m.[59]
Dholes still occur in Tibet, and possibly also in North Korea. They once occurred in the alpine steppes extending into Kashmir to the Ladakh area, but have not been recorded in Pakistan.[2]
They occur in most of India south of the Ganges, particularly in the Central Indian Highlands and the Western and Eastern Ghats. In northeast India, it is present in Arunachal Pradesh, Assam, Meghalaya, and West Bengal and in the Indo-Gangetic Plain's Terai region. Dhole populations in the Himalaya and northwest India are fragmented.[2]
In 2011, dhole packs were recorded by camera traps in the Chitwan National Park.[62] Its presence was confirmed in the Kanchenjunga Conservation Area in 2011 by camera-traps.[63]
In Bhutan, dholes have recovered from a poisoning campaign during the 1970s, and became re-established in the 1990s.[64] Today they occur in the Jigme Dorji National Park.[65]
Dholes still occur in northeastern Bangladesh's forest reserves in the Sylhet area, as well the Chittagong Hill Tracts in the southeast. These zones are unlikely to contain viable populations, considering most sightings involve small groups or solitary specimens, and they are likely decreasing in number due to the lack of prey.[2]
The presence of dholes in Myanmar was confirmed by camera-trapping in 11 areas, and alongside leopards, have apparently replaced tigers as the country's top predators.[2] In 2015, dholes and tigers were recorded by camera-traps for the first time in the hill forests of Karen State.[66]
Their range is highly fragmented in the Malaysian Peninsula, Sumatra, Java, Vietnam, and Thailand.[2] In 2014, camera trap videos in the montane tropical forests at 2,000 m in the Kerinci Seblat National Park in Sumatra revealed the continued presence of this species.[67] A camera trapping survey in the Khao Ang Rue Nai Wildlife Sanctuary in Thailand from January 2008 to February 2010 revealed at least one healthy dhole pack.[68] In northern Laos, dholes have been studied (2012) in protected areas.[69]
A disjunct population of this species has been reported from the area of Trabzon and Rize in northeastern Turkey near the border with Georgia in the 1990s by two Turkish zoologists.[70] Some authorities have accepted this report,[71] but others considered it to be unreliable.[2] Also, one single individual was claimed to have been shot in 2013 in the nearby Kabardino-Balkaria Republic (a subject republic of Russia immediately north of Georgia in the Central Caucasus); its remains (including a skull) were analyzed by a biologist from the Kabardino-Balkarian State University in May 2015, who concluded the skull was from a dhole.[72] Recently, in August 2015, researchers from the National Museum of Natural History from Sofia, Bulgaria (including Dr. Nikolai Spassov, the current director of this museum) and the Karadeniz Technical University began an expedition to track and document this possible Turkish population of dhole.[73] On October 12, 2015, this research team reported the preliminary conclusion that no real evidence exists of a living population of the dhole in Turkey (or in the Kabardino-Balkaria Republic), pending DNA analysis of samples from the original 1994 Serez-Eroglu skins.[74]
Ecology and behaviour
Social and territorial behaviour
Denning
Four kinds of den have been described; simple earth dens with one entrance (usually remodeled striped hyena or porcupine dens); complex cavernous earth dens with more than one entrance; simple cavernous dens excavated under or between rocks; and complex cavernous dens with several other dens in the vicinity, some of which are interconnected. Dens are typically located under dense scrub or on the banks of dry rivers or creeks. The entrance to a dhole den can be almost vertical, with a sharp turn three to four feet down. The tunnel opens into an antechamber, from which extends more than one passage. Some dens may have up to six entrances leading up to 100 feet (30 m) of interconnecting tunnels. These "cities" may be developed over many generations of dholes, and are shared by the clan females when raising young together.[79] Like African wild dogs and dingoes, dholes will avoid killing prey close to their dens.[80]Reproduction and development
Hunting behaviour
Once large prey is caught, one dhole will grab the prey's nose, while the rest of the pack pulls the animal down by the flanks and hindquarters. They do not use a killing bite to the throat.[86] They occasionally blind their prey by attacking the eyes.[87] Serows are among the only ungulate species capable of effectively defending themselves against dhole attacks, due to their thick, protective coats and short, sharp horns capable of easily impaling dholes.[4] They will tear open their prey's flanks and disembowel it, eating the heart, liver, lungs and some sections of the intestines. The stomach and rumen are usually left untouched.[88] Prey weighing less than 50 kg is usually killed within two minutes, while large stags may take 15 minutes to die. Once prey is secured, dholes will tear off pieces of the carcass and eat in seclusion.[89] Unlike wolf packs, in which the breeding pair monopolises food, dholes give priority to the pups when feeding at a kill, allowing them to eat first.[12] They are generally tolerant of scavengers at their kills.[90] Both mother and young are provided with regurgitated food by other pack members.[76]
Feeding ecology
Enemies and competitors
On some occasions, dholes may attack tigers. When confronted by dholes, tigers will seek refuge in trees or stand with their backs to a tree or bush, where they may be mobbed for lengthy periods before finally attempting escape. Escaping tigers are usually killed, while tigers which stand their ground have a greater chance of survival.[41] Tigers are dangerous opponents for dholes, as they have sufficient strength to kill a dhole with a single paw strike.[6] Dhole packs may steal leopard kills, while leopards may kill dholes if they encounter them singly or in pairs.[41] Since leopards are smaller than tigers and more likely hunt dholes, dhole packs tend to react more aggressively toward them than they do towards tigers.[94]
There are numerous records of leopards being treed by dholes.[76] Dholes sometimes drive tigers, snow leopards, leopards, and bears (see below) from their kills.[76] Dholes were once thought to be a major factor in reducing Asiatic cheetah populations, though this is doubtful, as cheetahs live in open areas as opposed to forested areas favoured by dholes.[95]
Dhole packs occasionally attack Asiatic black bears, snow leopards, and sloth bears. When attacking bears, dholes will attempt to prevent them from seeking refuge in caves, and lacerate their hindquarters.[41]
Although usually antagonistic toward wolves,[9] they may hunt and feed alongside one another.[96] There is at least one record of a lone wolf associating with a pair of dholes in Debrigarh Wildlife Sanctuary.[97] They infrequently associate in mixed groups with golden jackals. Domestic dogs may kill dholes, though they will feed alongside them on occasion.[98]
Diseases and parasites
Dholes are vulnerable to a number of different diseases, particularly in areas where they are sympatric with other canid species. Infectious pathogens such as Toxocara canis are present in their faeces. They may suffer from rabies, canine distemper, mange, trypanosomiasis, canine parvovirus, and endoparasites such as cestodes and roundworms.[13]Threats
The dhole only rarely takes domestic livestock. Certain people, such as the Kurumbas and some Mon Khmer-speaking tribes will appropriate dhole kills; some Indian villagers welcome the dhole because of this appropriation of dhole kills.[76] Dholes were persecuted throughout India for bounties until they were given protection by the Wildlife Protection Act of 1972. Methods used for dhole hunting included poisoning, snaring, shooting and clubbing at den sites. Native Indian people killed dholes primarily to protect livestock, while British sporthunters during the British Raj did so under the conviction that dholes were responsible for drops in game populations. Persecution of dholes still occurs with varying degrees of intensity according to region.[13] Bounties paid for dholes used to be 25 rupees, though this was reduced to 20 in 1926 after the number of presented dhole carcasses became too numerous to maintain the established reward.[99] In Indochina, dholes suffer heavily from nonselective hunting techniques such as snaring.[13]The fur trade does not pose a significant threat to dholes.[13] The people of India do not eat dhole flesh, and their fur is not considered overly valuable.[45] Due to their rarity, dholes were never harvested for their skins in large numbers in the Soviet Union, and were sometimes accepted as dog or wolf pelts (being labeled as "half wolf" for the latter). The winter fur was prized by the Chinese, who bought dhole pelts in Ussuriysk during the late 1860s for a few silver rubles. In the early 20th century, dhole pelts reached eight rubles in Manchuria. In Semirechye, fur coats made from dhole skin were considered the warmest, but were very costly.[9]
Conservation
In India, the dhole is protected under Schedule 2 of the Wildlife Protection Act, 1972. The creation of reserves under Project Tiger provided some protection for dhole populations sympatric with tigers. In 2014, the Indian government sanctioned its first dhole conservation breeding centre at the Indira Gandhi Zoological Park (IGZP) in Visakhapatnam.[100] The dhole has been protected in Russia since 1974, though it is vulnerable to poison left out for wolves. In China, the animal is listed as a category II protected species under the Chinese wildlife protection act of 1988. In Cambodia, the dhole is protected from all hunting, while conservation laws in Vietnam limit extraction and utilization.[2]In 2016, the Korean company Sooam Biotech was reported to be attempting to clone the dhole using dogs as surrogate mothers to help conserve the species.[101]
In culture and literature
Three dhole-like animals are featured on the coping stone of the Bharhut stupa dating from 100 BC. They are shown waiting by a tree, with a woman or spirit trapped up it, a scene reminiscent of dholes treeing tigers.[102] The animal's fearsome reputation in India is reflected by the number of pejorative names it possesses in Hindi, which variously translate as "red devil", "devil dog", "jungle devil", or "hound of Kali".[6] According to zoologist and explorer Leopold von Schrenck, he had trouble obtaining dhole specimens during his exploration of Amurland, as the local Gilyaks greatly feared the species. This fear and superstition was not however shared by neighbouring Tungusic peoples. Von Schrenk speculated that this differing attitude towards dholes was due to the Tungusic people's more nomadic, hunter-gatherer lifestyle.[25] Dhole-like animals are described in numerous old European texts, including the Ostrogoth sagas, where they are portrayed as hell hounds. The demon dogs accompanying Hellequin in Mediaeval French passion plays, as well as the ones inhabiting the legendary forest of Brocéliande, have been attributed to dholes. According to Charles Hamilton Smith, the dangerous wild canids mentioned by Scaliger as having lived in the forests of Montefalcone could have been based on dholes, as they were described as unlike wolves in habits, voice and appearance. The Montefalcone family's coat of arms had a pair of red dogs as supporters.[17]Dholes appear in Rudyard Kipling's Red Dog, where they are portrayed as aggressive and bloodthirsty animals which descend from the Deccan Plateau into the Seeonee Hills inhabited by Mowgli and his adopted wolf pack to cause carnage among the jungle's denizens. They are described as living in packs numbering hundreds of individuals, and that even Shere Khan and Hathi make way for them when they descend into the jungle. The dholes are despised by the wolves because of their destructiveness, their habit of not living in dens and the hair between their toes. With Mowgli and Kaa's help, the Seeonee wolf pack manages to wipe out the dholes by leading them through bee hives and torrential waters before finishing off the rest in battle.
Japanese author Uchida Roan wrote 犬物語 (Inu monogatari; A dog's tale) in 1901 as a nationalistic critique of the declining popularity of indigenous dog breeds, which he asserted were descended from the dhole.[103] A fictional version of the Dhole, imbued with supernatural abilities, appears in the Season 6 episode of The X-Files titled Alpha.
Dholes also appear as enemies in the game Far Cry 4, alongside other predators such as the Bengal tiger, honey badger, snow leopard, clouded leopard, Tibetan wolf and Asian black bear. They can be found hunting the player and other NPC's across the map, but are easily killed, being one of the weakest enemies in the game. They once again appear in the game Far Cry Primal, where they play similar roles as their counterparts in the previous game, but can now also be tamed and used in combat by Takkar, the main protagonist of the game.
Tameability
Brian Houghton Hodgson kept captured dholes in captivity, and found, with the exception of one animal, they remained shy and vicious even after 10 months.[45] According to Richard Lydekker, adult dholes are nearly impossible to tame, though pups are docile and can even be allowed to play with domestic dog pups until they reach early adulthood.[4] A dhole may have been presented as a gift to Ibbi-Sin as tribute.[104]Cuon alpinus
(Pallas, 1811)
Dhole range
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