Temporal range: Post-Pleistocene-Recent
The dhole (//), Cuon alpinus, also called the Asiatic wild dog or Indian wild dog, is a species of canid native to South and Southeast Asia. It is the only extant member of the genus Cuon, which differs from Canis by the reduced number of molars and greater number of teats. The dholes are classed as endangered by the IUCN, due to ongoing habitat loss, depletion of its prey base, competition from other predators, persecution and possibly diseases from domestic and feral dogs.
The dhole is a highly social animal, living in large clans which occasionally split up into small packs to hunt. It primarily preys on medium-sized ungulates, which it hunts by tiring them out in long chases, and kills by disemboweling them. Unlike most social canids (but similar to African wild dogs), dholes let their pups eat first at a kill. Though fearful of humans, dhole packs are bold enough to attack large and dangerous animals such as wild boar, water buffalo, and even tigers.
- 1 Naming and etymology
- 2 Evolution and taxonomy
- 3 Physical description
- 4 Behaviour
- 5 Communication
- 6 Diseases and parasites
- 7 Range
- 8 Relationships with humans
- 9 References
- 10 External links
Naming and etymology
Since these canids are called dholes only in English, the etymology remains unclear, but it may have come from Kannada: tōḷa (‘wolf’). Some 19th-century authors connected this word with Turkish: deli ‘mad, crazy’, and erroneously compared the Turkish word with Old Saxon: dol and Dutch: dol (cf. also English: dull; German: toll), which are in fact from Proto-Germanic *dwalaz ‘foolish, stupid’.
Vernacular names include lal rakshasa (red devil) and rakshur kukur (devil dog) in India, and jangli rakshasa (jungle devil) or hounds of Kali in Assam. In the Himalayas, they are variously known as Bhaosa, Bhansa and Buansu.
- Assamese: raang-kukur
- Bengali: রাম কুত্তা (ram kut ta), জংলি কুকুর (jongli-kukur), বুনো কুকুর (būno-kukūr)
- Hindi/Urdu: jangli-kutta, ram-kutta, son-kutta or ban-kutta
- Kannada: ಸೀಳು ನಾಯಿ (sīḷu nāyi), ಕಾಡು ನಾಯಿ (kāḍu nāyi)
- Malayalam: കാട്ടുനായ (Kāttunāya) or കാട്ടുപട്ടി (kāttu patti)
- Marathi: kolsun, kolasna, kolasra or kolsa
- Mongol: улаан чоно
- Tamil: செந்நாய் Sen Naai (Red dog)
- Telugu: resu-kukka or adavi-kukka అడవి కుక్క, రేసు కుక్క
- Gondi: eram-naiko
- Ho: tani
- Kashmiri: ram-hun
- Ladakhi: siddaki
- Tibetan: hazi or phara
- Bhotia: paoho
- Lepcha: sa-tum
- Chinese: 豺 chái
- Burmese: tau-khwe
- Indonesia: ajag
- Malay: serigala or anjing hutan
- Nepali: वन कुकूर (wan kukūr)
- Russian: Красный волк (krasnyy volk) ‘red wolf’
- Korean: 승냥이 seungnyangi
- Gujarati: jangli-kutra
- Baltistan-Pakistan: jangli-kuta
- Thai: ma nai (หมาใน)
- Lao: ma nai (ໝາໃນ)
Evolution and taxonomy
Dholes are post-Pleistocene in origin, and are more closely related to jackals than they are to wolves. One theory has dholes becoming social animals as an adaptation to living with tigers and Indian leopards.
George Gaylord Simpson placed dholes under the subfamily Symocyoninae along with the African wild dog and bush dog on account of shared anatomical features, namely the reduction of postcarnassial molars. Many have questioned this classification, arguing that these shared features are due to convergent evolution. Juliet Clutton-Brock concluded from comparing the morphological, behavioural and ecological characteristics of 39 different canid species that with the exception of skull and dentition, dholes more closely resembled canids of the genus Canis, Dusicyon and Vulpes/Alopex than to African wild dogs and bush dogs. A comparative study on dhole and other canid mtDNA in 1997 showed dholes diverged from the Lupus lupus lineage before the black-backed jackal and the golden jackal diverged, a few million years before the domestication of the dog.
Cuon a. dukhunensis
|Sykes, 1831||Peninsular India, south of Ganges River|
Cuon a. primaevus
|Hodgson, 1833||Gentically distinct||Himalaya and northern regions of the Indian subcontinent: Khangchendzonga Biosphere Reserve, Sikkim, India|
Cuon a. laniger
|Pocock, 1936||Kashmir, Lhasa|
Cuon a. lepturus
|Heude, 1892||Southern China||grayiformis (Hodgson, 1863)
clamitans (Heude, 1892) rutilans (Müller, 1839)
Cuon a. infuscus
|Pocock, 1936||Myanmar, Thailand, Laos, Cambodia, Vietnam||adustus (Pocock, 1941)
Cuon a. sumatrensis
|Hardwicke, 1821||A small subspecies, it measures only two feet in length, and stands 360 mm (14 in) high at the shoulder. It has a pointed, black, fox-like muzzle with long, dark whiskers. The nose and lips are foxy brown mixed with black. The general colour is foxy ferraginous red, with lighter shades on the belly and inner sides of the legs.||Sumatra, Indonesia|
Cuon a. javanicus
|Desmarest, 1820||Java, Indonesia|
|Eastern or Ussuri dhole
Cuon a. alpinus
|Pallas, 1811||This is the largest subspecies, with a long, narrow face and a skull measuring 189 mm long on average. The winter fur's general tone is intense rusty-red. The top of the head and the outer ears are brownish-rusty with black-brown highlights. The shoulders and upper surface of the back is brownish-rusty with black-brown highlights. The outer sides of the legs are rusty brown, while the inner sides of the legs and lower sides of the body are yellowish.||Russian Far East, China, Tibet, Mongolia||fumosus (Pocock, 1936)
|Western or Tien Shan dhole
Cuon a. hesperius
|Afanasjev and Zolotarev, 1935||A small subspecies, it has a short, wide face and a skull measuring 180 mm long on average. The general tone of the winter fur is lighter-coloured than C. a. alpinus, with weakly developed rusty-red tints. The top of the head and outer sides of the ears are reddish-straw coloured. The upper surface of the neck is dirty-white, with a narrow, sandy-yellow-coloured band running along the upper surface of the back from the ears to the shoulders. The outer surfaces of the limbs are sandy-yellow, while the flanks and inner sides of the limbs have little to no yellowish tint.||Transoxiana, Eastern Russia and China||jason (Pocock, 1936)|
|†Late Pleistocene dhole
Cuon a. europaeus
|Bourguignat, 1875||The earliest form to evolve a singlely cusped, sharply trenchant tooth in place of the lower tubercular molar||Czech Republic, Hungary, Austria, Switzerland and French Riviera|
|†Late Middle Pleistocene dhole
Cuon a. fossilis
|Nehring, 1890||An intermediate form between Cuon a. priscus and Cuon a. europaeus||Heppenloch, Germany|
|†Early Middle Pleistocene dhole
Cuon a. priscus
Dholes have relatively short, heavy and massive skulls, with shortened facial regions, widely separated zygomatic arches and well-developed sagittal crests. The frontal bone is inflated, and passes down onto the snout, giving the animals a convex rather than concave profile. The masseter muscles are highly developed compared to other canid species, giving the face an almost hyena-like appearance. The skull is broader, and has a shorter rostrum than that of domestic dogs and most other canids. The dental formula is 22.214.171.124
The species uniquely has six rather than seven lower molars. The upper molars are weak, being one-third to one-half the size of those of wolves, and have only one cusp as opposed to two to four, as is usual in canids, an adaptation thought to improve shearing ability, thus allowing it to compete more successfully with kleptoparasites. The canine teeth are slightly curved and short.
Their limbs are moderately long, and their thoraces are proportional. Along with African wild dogs, dholes are often referred to as "cat-like" canids, due to their long, fine limbs and backbones. They have great jumping and leaping abilities, being able to jump 3.0–3.5 m (10–12 ft) high, and leap 5– to 6-m (17– to 20-ft) distances in one leap with a running start. Their tails measure 16–17 in long, and are almost half the length of their bodies, nearly touching the ground when in full winter fur. They are smaller than African wild dogs. Weights range from 10 to 25 kg (22 to 55 lb), with males averaging about 4.5 kg (9.9 lb) heavier than females. This dog is 88 to 113 cm (35 to 44 in) long from the snout to the base of the tail, with the tail averaging 45 cm (18 in) in length. Shoulder height is 42 to 55 cm (17 to 22 in). Like African wild dogs, their ears are rounded rather than pointed. However, unlike the former species, male dholes do not have a clearly visible prepuce, thus making the sexing of individuals difficult even at close proximity. Unlike members of the Canis genus, females have 12–14 teats rather than 10. They are not as odorous as wolves, jackals and foxes, having a smaller number of anal scent glands. Their stomachs have been estimated to hold 6.5 lb (2.9 kg) of food.
The general tone of the fur is reddish, with the brightest hues occurring in winter. When in their winter fur, the back is clothed in a saturated rusty-red to reddish colour with brownish highlights along the top of the head, neck and shoulders. The throat, chest, flanks, belly and the upper parts of the limbs are less brightly coloured, and are more yellowish in tone. The lower parts of the limbs are whitish, with dark brownish bands on the anterior sides of the forelimbs. The muzzle and forehead are greyish-reddish. The tail is very luxuriant and fluffy, and is mainly of a reddish-ochre colour, with a dark brown or blackish tip. The summer coat is shorter, coarser and darker. The dorsal and lateral guard hairs in adults measure 20–30 mm in length. Dholes in the Moscow Zoo moult once a year from March to May.
Social and territorial behaviours
Dholes are more social than wolves, and have less of a dominance hierarchy, as seasonal scarcity of food is not a serious concern for them as it is with wolves. In this sense, they closely resemble African wild dogs in social structure. Dominant dholes are hard to identify, as they do not engage in dominance displays as wolves do, though other clan members will show submissive behaviour toward them. Intragroup fighting is rarely observed. They live in clans rather than packs, as the latter term refers to a group of animals that always hunt together. In contrast, dhole clans frequently break into small packs of 3–5 animals, particularly during the spring season, as this is the optimal number for catching fawns. Dholes are far less territorial than wolves, with pups from one clan often joining another without trouble once they mature sexually. Clans typically number 5-12 individuals in India, though clans of 40 have been reported. In Thailand, clans rarely exceed three individuals. Unlike other canids, there is no evidence of dholes using urine to mark their territories or travel routes. They may defecate in conspicuous places, though a territorial function is unlikely, as faeces are mostly deposited within the clan's territory rather than the periphery. Faeces are often deposited in what appear to be communal latrines. They do not scrape the earth with their feet as other canids do to mark their territories.
Reproduction and development
Among Indian dholes, the mating season occurs between mid-October and January, while captive dholes in the Moscow Zoo breed mostly in February. Unlike wolf packs, dhole clans may contain more than one breeding female. More than one female dhole may den and rear their litters together in the same den. During mating, the female assumes a crouched, cat-like position. There is no "tug of war" characteristic of other canids[further explanation needed] when the male dismounts. Instead, the pair lie on their sides facing each other in a semicircular formation. The gestation period lasts 60–63 days, with litter sizes averaging four to six pups. Their growth rate is much faster than that of wolves, being similar in rate to that of coyotes. Pups are suckled at least 58 days. During this time, the pack feeds the mother at the den site. Dholes do not use rendezvous sites to meet their pups as wolves do, though one or more adults will stay with the pups at the den while the rest of the pack hunts. Once weaning begins, the adults of the clan will regurgitate food for the pups until they are old enough to join in hunting. They remain at the den site 70–80 days. By the age of six months, pups accompany the adults on hunts, and will assist in killing large prey such as sambar by the age of eight months. Maximum longevity in captivity is 15–16 years.
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. Like African wild dogs and dingoes, dholes will avoid killing prey close to their dens.
Diet, hunting and feeding behaviours
Prey animals in India include chital, sambar, muntjac, mouse deer, swamp deer, wild boar, gaur, water buffalo, banteng, cattle, nilgai, goats, Indian hares, Himalayan field rats and langurs. There is one record of a pack bringing down an Indian elephant calf in Assam, despite desperate defense of the mother resulting in numerous losses to the pack. In Kashmir, they may hunt markhor, and thamin in Burma. Javan rusas are hunted in Java. In the Tien Shan and Tarbagatai Mountains, dholes prey on Siberian ibexes, arkhar, roe deer, maral and wild boar. In the Altai and Sayan Mountains, they prey on musk deer and reindeer. In eastern Siberia, they prey on roe deer, Manchurian wapiti, wild boar, musk deer, and reindeer, while in Primorye they feed on sika deer and goral, too. In Mongolia, they prey on argali and rarely Siberian ibex. Like African wild dogs, but unlike wolves, dholes are not known to attack people. Dholes eat fruit and vegetable matter more readily than other canids. In captivity, they eat various kinds of grasses, herbs and leaves, seemingly for pleasure rather than just when ill. In summertime in the Tien Shan Mountains, dholes eat large quantities of mountain rhubarb. Bael fruits are also eaten. Although opportunistic, dholes have a seeming aversion to hunting cattle and their calves. Livestock predation by dholes has been a problem in Bhutan since the late 1990s, as domestic animals are often left outside to graze in the forest, sometimes for weeks at a time. Livestock stall-fed at night and grazed near homes are never attacked. Oxen are killed more often than cows are, probably because they are given less protection.
Before embarking on a hunt, clans go through elaborate prehunt social rituals involving nuzzling, body rubbing and homo- and heterosexual mounting. Dholes are primarily diurnal hunters, hunting in the early hours of the morning. They rarely hunt nocturnally, except on moonlit nights, indicating they greatly rely on sight when hunting. Though not as fast as jackals and foxes, they can chase their prey for many hours. During a pursuit, one or more dholes may take over chasing their prey, while the rest of the pack keeps up at a steadier pace behind, taking over once the other group tires. Most chases are short, lasting only 500 m. When chasing fleet-footed prey, they run at a pace of 30 mph. Dholes frequently drive their prey into water bodies, where the targeted animal's movements are hindered.
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. They occasionally blind their prey by attacking the eyes. 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. 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. 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. 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. They are generally tolerant of scavengers at their kills.
Both mother and young are provided with regurgitated food by other pack members.
Relationships with other predators
In some areas, dholes are sympatric to tigers and leopards. Competition between these species is mostly avoided through differences in prey selection, although there is still substantial dietary overlap Along with leopards, dholes typically target animals in the 30–175 kg range (mean weights of 35.3 kg for dhole and 23.4 kg for leopard), while tigers selected for prey animals heavier than 176 kg (but their mean prey weight was 65.5 kg). Also, other characteristics of the prey, such as sex, arboreality, and aggressiveness, may play a role in prey selection. For example, dholes preferentially select male chital, whereas leopards kill both sexes more evenly (and tigers prefer larger prey altogether), dholes and tigers kill langurs rarely compared to leopards due to the leopards' greater arboreality, while leopards kill wild boar infrequently due to the inability of this relatively light predator to tackle aggressive prey of comparable weight. On some rare 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. Tigers are extremely dangerous opponents for dholes, as they have sufficient strength to kill a single dhole with one paw strike. Even a successful tiger kill is usually accompanied by losses to the pack. Dhole packs may steal leopard kills, while leopards may kill dholes if they encounter them singly or in pairs. Because leopards are smaller than tigers, and are more likely to hunt dholes, dhole packs tend to react more aggressively toward them than they do with tigers. There are numerous records of leopards being treed by dholes. Dholes sometimes drive tiger, leopards, and bears (see below) from their kills. 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.
Though usually antagonistic toward wolves, they may hunt and feed alongside one another. They infrequently associate in mixed groups with golden jackals. Domestic dogs may kill dholes, though they will feed alongside them on occasion.
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. Other sounds include whines (food soliciting), growls (warning), screams, chatterings (both of which are alarm calls) and yapping cries. In contrast to wolves, dholes do not howl or bark.
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 will open their mouths with their lips retracted and their tails held in a vertical position whilst assuming a play bow. Aggressive or threatening dholes will 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.
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.
Dholes once ranged throughout most of South, East and Southeast Asia, extending from the Tien Shan of Kyrgyzstan and China, the eastern Pamir Mountains of Tajikistan, and the Altai Mountains and the Primorsky Krai region of Russia southward through Mongolia, Korea, China, Tibet, Nepal, Pakistan-controlled Kashmir territory, India, and south-eastwards into Myanmar and Indochina, Thailand, Malaysia, Sumatra and Java. A report  of a population in northeastern Turkey has been noted by some authorities, but considered to be unreliable by others.
During the last glacial period, they ranged across most of Eurasia, and are known to have once inhabited North America from a single fossil find in the Gulf of Mexico. This range extended as far west as Western Europe and included several islands that this species no longer inhabits, such as Sri Lanka, Borneo, and possibly Palawan (in the Philippines). A canid called the Sardinian dhole (Cynotherium sardous) lived on the Mediterranean island of Sardinia during the Pleistocene, but it is not as closely related to the living species as its name would imply.
Dholes have not been reported recently in Russia, Mongolia, Kazakhstan, Kyrgyzstan and Tajikistan. One capture was reported in southern China's Jiangxi province. In 2006, one pack was observed in the Qilian Shan mountains in the Gansu province of northwestern China. Dholes still occur in Tibet, particularly in southeast Tibet. They may still be present in North Korea. They still occur in India south of the Ganges River, especially in the central Indian Highlands and the Western and Eastern Ghats. Dholes also occur in northeast India's states of Arunachal Pradesh, Assam, Meghalaya and West Bengal and in Ladakh. They have a precarious, fragmented distribution in Himalaya and northwest India. They are occasionally reported in the Ladakh area of Kashmir, contiguous with the Tibetan highlands and China. In Bhutan, dholes have since recovered from a government-sponsored poisoning campaign started in the 1970s, with reports of livestock predation occurring in the lower Kheng region. It is uncertain if they still occur in Bangladesh. Camera trapping has confirmed dholes still occur in 11 survey areas in Myanmar, where they have replaced tigers as main predators. Dhole populations are highly fragmented in Thailand and Indochina, particularly in Vietnam. They are known to occur in four sites in northern and central Malaysia. In Java, they appear to be most common in the island's protected eastern and western ends. They are also known to occur in Sumatra's protected areas in the southern, central and northern areas.
Relationships with humans
Hunting and persecution
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. 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. 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. In Indochina, dholes suffer heavily from nonselective hunting techniques such as snaring.
The fur trade does not pose a significant threat to dholes. The people of India do not eat dhole flesh, and their fur is not considered overly valuable. 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.
Dholes in folklore, mythology, literature and popular culture
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. 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. The dangerous wild canids mentioned by Scaliger as having lived in the forests of Montefalcone could have been 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.
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. They would reappear in two animated television adaptations: Jungle Book Shōnen Mowgli and Jungle Cubs.
In an episode called "Alpha" in season six of The X-Files, a cryptid dhole from China is blamed for multiple killings. This dhole (Wanshang dhole) is also mentioned in the Angel episode That Vision Thing. Both episodes were written by Jeffrey Bell.
Tameability and possible relation to the dog
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. 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. A dhole may have been presented as a gift to Ibbi-Sin as tribute.
- Wozencraft, W. C. (2005). "Order Carnivora". In Wilson, D. E.; Reeder, D. M. Mammal Species of the World (3rd ed.). Johns Hopkins University Press. pp. 532–628. ISBN 978-0-8018-8221-0. OCLC 62265494.
- Durbin, L. S., Hedges, S., Duckworth, J. W., Tyson, M., Lyenga, A. & Venkataraman, A. (IUCN SSC Canid Specialist Group – Dhole Working Group) (2008). "Cuon alpinus". IUCN Red List of Threatened Species. Version 2014.2. International Union for Conservation of Nature.
- Fox 1984, p. 85
- Fox 1984, pp. 58–60
- Fox 1984, p. 63
- Fox 1984, pp. 86–7
- Lydekker 1907, p. 360
- dhole. Oxford dictionary on-line.
- dhole. Merriam-Webster Dictionary.
- Smith & Jardine 1839, p. 179
- Orel, Vladimir (2003), A Handbook of Germanic Etymology, Leiden, Boston: Brill, p. 81, ISBN 90-04-12875-1
- Lydekker 1907
- Fox 1984
- Heptner & Naumov 1998
- Cohen, James A. (1978). "Cuon alpinus". Mammalian Species 100 (100): 1–3. doi:10.2307/3503800.
- Perry 1965, p. 147
- Perry 1965, p. 145
- Shretha 1997, p. 121
- Sillero-Zubiri, Hoffman & MacDonald 2004, p. 210
- Venkataraman, A. (1995). "Do dholes (Cuon alpinus) live in packs in response to competition with or predation by large cats?". Current Science 11: 934–936.
- Wayne, Robert K.; Geffen, Eli; Girman, Derek J.; Koepfli, Klaus P.; Lau, Lisa M. and Marshall, Charles R. (1997). "Molecular Systematics of the Canidae". Systematic Biology 46 (4): 622–653. doi:10.1093/sysbio/46.4.622. PMID 11975336.
- Iyengar, A., et al. Phylogeography, genetic structure, and diversity in the dhole (Cuon alpinus). Molecular Ecology 14.8 (2005): 2281-2297.
- Bashir, T., Bhattacharya, T., Poudyal, K., Roy, M., Sathyakumar, S. (2013).Precarious status of the Endangered dhole Cuon alpinus in the high elevation Eastern Himalayan habitats of Khangchendzonga Biosphere Reserve, Sikkim, India. Oryx 48: 125–132.
- Smith & Jardine 1839, pp. 186–7
- Fox 1984, p. 40
- Heptner & Naumov 1998, p. 578
- Heptner & Naumov 1998, p. 579
- Kurtén 1968, pp. 112–14
- Heptner & Naumov 1998, p. 567
- Pocock 1941, p. 149
- Fox 1984, pp. 61–2
- Fox 1984, p. 41
- Sillero-Zubiri, Hoffman & MacDonald 2004, p. 214
- Heptner & Naumov 1998, p. 568
- Sosnovskii, Igor P. "Breeding the Red dog or dhole Cuon alpinus at Moscow Zoo". In: International Zoo Yearbook 7, 1967, 120–122.
- Heptner & Naumov 1998, p. 571
- Fox 1984, p. 8
- Boitani, Luigi, Simon & Schuster's Guide to Mammals. Simon & Schuster/Touchstone Books (1984), ISBN 978-0-671-42805-1
- Dogs and Hyenas – Relative success in different climatic zones. askabiologist.org.uk (29 August 2009).
- Dhole, red dog, Asiatic wild dog, Indian wild dog. BBC (April 2012).
- Smith & Jardine 1839, pp. 168–9
- Fox 1984, p. 59
- Heptner & Naumov 1998, p. 572
- Nowak, Ronald M; and Paradiso, John L. Walker's Mammals of the World. 4th ed. Baltimore: Johns Hopkins University Press; 1983. p963
- Fox 1984, pp. 81–2
- Fox 1984, p. 92
- Fox 1984, p. 97
- Nowak, Ronald M; and Paradiso, John L. Walker's Mammals of the World. 4th ed. Baltimore: Johns Hopkins University Press; 1983. p962
- Fox 1984, p. 79
- Fox 1984, p. 80
- Fox 1984, pp. 43–49
- Pocock 1941, p. 161
- Heptner & Naumov 1998, p. 583
- Pocock 1941, p. 162
- Heptner & Naumov 1998, p. 584
- Mivart 1890, p. 181
- Shrestha 1996, p. 122
- Fox 1984, p. 71
- Johnsingh, A.J.T., Yonten, Deki & Wangchuck, Sangay (2007). "Livestock-Dhole Conflict in Western Bhutan". J. Bombay Nat. Hist. Soc. 104 (2): 201.
- Fox 1984, pp. 100–1
- Fox 1984, p. 50
- Fox 1984, p. 73
- Fox 1984, p. 67
- Grassman, L. I., Jr., M. E. Tewes, N. J. Silvy, and K. Kreetiyutanont (2005). "Spatial ecology and diet of the dhole Cuon alpinus (Canidae, Carnivora) in north central Thailand". Mammalia 69 (1): 11–20. doi:10.1515/mamm.2005.002.
- Lydekker 1907, p. 363-2
- Fox 1984, p. 70
- Fox 1984, p. 51
- Karanth K. U. and Sunquist M. E. (1995). "Prey selection by tiger, leopard and dhole in tropical forests". Journal of Animal Ecology 64 (4): 439–450. doi:10.2307/5647. JSTOR 5647.
- Perry 1968, p. 149
- Perry 1968, p. 150
- Finn 1929, p. 120
- Heptner & Naumov 1998, p. 585
- Humphrey & Bain 1990, p. 572
- Fox 1984, p. 93
- Fox 1984, p. 95
- Heptner & Naumov 1998, p. 586
- Sillero-Zubiri, Hoffman & MacDonald 2004, p. 216
- Sillero-Zubiri, Hoffman & MacDonald 2004, pp. 212–3
- Serez, Mehment; Eroglu, Mahmut, 1994: A new threatened wolf species, Cuon alpinus hesperius Afanasiev and Zolatarev, 1935 in Turkey. Council of Europe Environmental Encounters Series 17: 103-106
- Nowak, Ronald M. Walker's Carnivores of the World. Baltimore: Johns Hopkins University Press; 2005. pp110-111 f=false
- Wang, Xiaoming & Tedford, Richard H. Dogs: their fossil relatives and evolutionary history p. 64, Columbia University Press, 2008, ISBN 0-231-13528-9
- Nowak, Ronald M. Walker's Carnivores of the World. Baltimore: Johns Hopkins University Press; 2005. pp110-111
- Ochoa, Janine; Paz, Victor; Lewis, Helen; Carlos, Jane; Robles, Emil; Noel Amano2, Maria Rebecca Ferreras, Myra Lara, Benjamin Vallejo, Jr.5, Gretchen Velarde, Villaluz, Sarah Agatha; Ronquillo, Wilfredo; and Solheim II, Wilhelm. The archaeology and palaeobiological record of Pasimbahan-Magsanib Site, northern Palawan, Philippines. Philipine Science Letters. 2004; 7 (1):22-36. p31
- Lyras, G.A.; Van Der Geer, A.E.; Dermitzakis, M.; De Vos, J. (2006), "Cynotherium sardous, an insular canid (Mammalia: Carnivora) from the Pleistocene Of Sardinia (Italy), and its origin", Journal of Vertebrate Paleontology 26 (3): 735–745, doi:10.1671/0272-4634(2006)26[735:CSAICM]2.0.CO
- Harris, R. B. 2006. Attempted predation on blue sheep Pseudois nayaur by dholes Cuon alpinus. Journal of the Bombay Natural History Society 103: 95-97.
- Thapa, K., Kelly, M. J., Karki, J. B., & Subedi, N. (2013). First camera trap record of pack hunting Dholes in Chitwan National Park, Nepal. Canid Biology and Conservation 16 (2): 4−7.
- Fox 1984, p. 109
- Mivart 1890, p. 187
- van der Geer, Alexandra Anna Enrica Animals in stone: Indian mammals sculptured through time, p. 188, BRILL, 2008, ISBN 90-04-16819-2
- Smith & Jardine 1839, p. 178
- Lydekker 1907, p. 365
- McIntosh, Jane The ancient Indus Valley: new perspectives, p. 130, ABC-CLIO, 2008 ISBN 1-57607-907-4
- Lindblad-Toh, K.; Wade, CM; Mikkelsen, TS; Karlsson, EK; Jaffe, DB; Kamal, M; Clamp, M; Chang, JL et al. (2005). "Genome sequence, comparative analysis and haplotype structure of the domestic dog". Nature 438 (7069): 803–819. doi:10.1038/nature04338. PMID 16341006.
- Blandford, W. T. (1888). The Fauna of British India including Ceylon and Burma: Mammalia. London: Taylor and Francis.
- Finn, F. (1929). Sterndale's Mammalia of India. London: Thacker, Spink & Co.
- Fox, M. W. (1984). The Whistling Hunters: Field Studies of the Indian Wild Dog (Cuon Alpinus). Steven Simpson Books. ISBN 0-9524390-6-9.
- Heptner, V. G.; Naumov, N. P. (1998). Mammals of the Soviet Union Vol.II Part 1a, SIRENIA AND CARNIVORA (Sea cows; Wolves and Bears). Science Publishers, Inc. USA. ISBN 1-886106-81-9.
- Humphrey, S. R.; Bain, J. R. (1990). Endangered Animals of Thailand. Gainesville, USA: Sandhill Crane Press. ISBN 1-877743-07-0.
- Jerdon, T. C. (1874). The mammals of India; a natural history of all the animals known to inhabit continental India. London: J. Wheldon.
- Kurtén, Björn (1968). Pleistocene mammals of Europe. Weidenfeld and Nicolson.
- Lydekker, R. (1907). The great and small game of India, Burma, and Tibet. Rowland Ward Ltd.
- MacDonald, David W.; Sillero-Zubiri, Claudio (2004). The biology and conservation of wild canids. Oxford University Press. ISBN 0-19-851556-1.
- Mivart, George (1890). Dogs, Jackals, Wolves and Foxes: A Monograph of the Canidæ.
- Perry, Richard (1965). The World of the Tiger. Cassell & Company ltd. ASIN B0007DU2IU.
- Pocock, R. I. (1941). Fauna of British India: Mammals Volume 2. London: Taylor and Francis.
- Shretha, Tej Kumar (1997). Mammals of Nepal: (with reference to those of India, Bangladesh, Bhutan and Pakistan). Steven Simpson Books. ISBN 0-9524390-6-9.
- Sillero-Zubiri, C.; Hoffman, M.; MacDonald, D. W. (2004). Canids: Foxes, Wolves, Jackals and Dogs – 2004 Status Survey and Conservation Action Plan. IUCN/SSC Canid Specialist Group. ISBN 2-8317-0786-2.
- Smith, C. H.; Jardine, W. (1839). The natural history of dogs : canidae or genus canis of authors; including also the genera hyaena and proteles. Edinburgh: W. H. Lizars.
|Wikispecies has information related to: Cuon alpinus|
|Wikimedia Commons has media related to Cuon alpinus.|
- Dhole Home Page
- mapping the dhole in south east asia
- ARKive – images and movies of the dhole
- Photos of dhole in Bandipur