Temporal range: Middle Pleistocene-Recent
|Captive C. alpinus, Zooparc de Trégomeur, Brittany, France|
The dhole (Cuon alpinus) is a canid native to Central and Southeast Asia. It is genetically close to species within the genus Canis,(Fig. 10) though its skull is convex rather than concave in profile, it lacks a third lower molar, and the upper molars sport only a single cusp as opposed to 2-4. During the Pleistocene, the dhole ranged throughout Asia, Europe and North America, but became restricted to its historical range 12,000-18,000 years ago.
The dhole is a highly social animal, living in large clans without rigid dominance hierarchies and containing multiple breeding females. Such clans usually consist of 12 individuals, but groups of over 40 are known. It is a diurnal pack hunter which preferentially targets medium and large sized ungulates. In tropical forests, the dhole competes with tigers and leopards, targeting somewhat different prey species, but still with substantial dietary overlap.
It is listed as Endangered by the IUCN, as populations are decreasing and estimated at less than 2,500 adults. Factors contributing to this decline include habitat loss, loss of prey, competition with other species, persecution, and disease transfer from domestic dogs.
- 1 Characteristics
- 2 Distribution and habitat
- 3 Ecology and behaviour
- 4 Threats
- 5 Conservation
- 6 Discovery, taxonomy and evolution
- 7 In culture and literature
- 8 Etymology and naming
- 9 See also
- 10 References
- 11 External links
In appearance, the dhole has been variously described as combining the physical characteristics of the grey wolf and red fox, and as being "cat-like" on account of its long backbone and slender limbs. It has a wide and massive skull with a well-developed sagittal crest, and its masseter muscles are highly developed compared to other canid species, giving the face an almost hyena-like appearance. The rostrum is shorter than that of domestic dogs and most other canids. The species 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 2-4, as is usual in canids, an adaptation thought to improve shearing ability, thus allowing it to compete more successfully with kleptoparasites. Adults may weigh over 18 kg (40 lb), with females usually weighing 4.5 kg (9.9 lb) less than males. It stands 17-22 inches at the shoulder and measures three feet in body length. Like the African wild dog, its ears are rounded rather than pointed. It has 6-7 teats, sometimes eight.
The general tone of the fur is reddish, with the brightest hues occurring in winter. In the winter coat, 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-ocherous colour, with a dark brown 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.
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.
Distribution and habitat
There are currently no confirmed recent reports of dhole being present in Russia, Mongolia, Kazakhstan, Kyrgyzstan or Tajikistan, though one specimen was caught in southern China's Jiangxi district. It is unknown if dholes continue to inhabit Tien Shan, though they possibly occur in small numbers in Gansu Province, with one pack being sighted in the Qilian Mountains within that province in 2006. Dholes still occur in Tibet, and may still inhabit North Korea. Although they have not been recorded in Pakistan, they once occurred in the alpine steppes extending into Kashmir. They occur in most of India south of the Ganges, particularly in the Central Indian Highlands and the Western and Eastern Ghats of the southern states. In north-east India, they inhabit Arunachal Pradesh, Assam, Meghalaya, and West Bengal. The situation of dholes in the Himalaya and north-west India is precarious, and populations fragmented. They may occur in Kashmir's Ladakh area.
Dholes once occurred in the Indo-Gangetic Plain's Terai region. In 2011, dhole packs were recorded by camera traps in the Chitwan National Park. Their presence was confirmed in the Kanchenjunga Conservation Area in 2011 by camera traps.
It is unknown whether the species still lives in Bangladesh, where it once inhabited the forested areas of the Chittagong and Sylhet District. 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.
Their range is highly fragmented in the Malaysian Peninsula, Sumatra, Java, Vietnam and Thailand. A camera trapping survey in the Khao Ang Rue Nai Wildlife Sanctuary during January 2008 to February 2010 revealed at least one healthy dhole pack.
In Central Asia, dholes primarily inhabit mountainous areas; in the western half of its range, they live mostly in alpine meadows and high-montane steppes high above sea level, while in the east, they mainly ranges in montane taigas, though may appear along coastlines. In India, Myanmar, Indochina, Indonesia and China, they prefer forested areas in alpine zones, and occasionally also in plains regions.
Ecology and behaviour
Social and territorial behaviour
Dholes are more social than grey wolves, and have less of a dominance hierarchy, as seasonal scarcity of food is not a serious concern for them. In this manner, they closely resemble African wild dogs in social structure. 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. 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. 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.
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.
Reproduction and development
In India, 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 copulatory tie characteristic of other canids 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 4-6 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.
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.
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. 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.
Enemies and competitors
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. 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.
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. There is at least one record of a lone wolf associating with a pair of dholes in Debrigarh Wildlife Sanctuary. They infrequently associate in mixed groups with golden jackals. Domestic dogs may kill dholes, though they will feed alongside them on occasion.
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.
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.
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. The dhole has been protected in Russia since 1974, though it is vulnerable to poison let 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.
Discovery, taxonomy and evolution
The species was first described in literature in 1794 by an explorer named Pesteref, who encountered dholes during his travels in far eastern Russia. He described the animal as being a regular pack hunter of Alpine ibex, and of bearing many similarities with the golden jackal. It was given the binomial name Canis alpinus in 1811 by Peter Pallas, who described its range as encompassing the upper levels of Udskoi Ostrog in Amurland, towards the eastern side of the Lena River, though he wrote that it also occurred around the Yenisei, and that it occasionally crossed into China. The British naturalist Brian Hodgson gave the dhole the binomial name Canis primaevus, assuming that it is the progenitor of the domestic dog. Hodgson later took note of the dhole's physical distinctiveness from the genus Canis and assigned it to a new genus Cuon.
The first study on the origins of the species was conducted by paleontologist Erich Thenius, who concluded that the dhole was a post-Pleistocene descendant of a golden jackal-like ancestor. The earliest known member of the genus Cuon is the Chinese C. majori of the Villafranchian period. It resembled Canis in its physical form more than the modern species, which has greatly reduced molars, whose cusps have developed into sharply trenchant points. By the Middle Pleistocene, C. majori had lost the last lower molar altogether. C. alpinus itself arose during the late Middle Pleistocene, by which point the transformation of the lower molar into a single cusped, slicing tooth had been completed. Late Middle Pleistocene dholes were virtually indistinguishable from their modern descendants, save for their greater size, which closely approached that of the grey wolf. The dhole became extinct in much of Europe during the late Würm period, though it may have survived up until the early Holocene in the Iberian Peninsula and at Riparo Fredian in northern Italy The fossil record indicates that the species also occurred in North America, with remains being found in Beringia and Mexico.
The dhole's distinctive morphology has been a source of much confusion in determining the species' systematic position among the canidae. George Simpson placed the dhole in the subfamily Simocyoninae alongside the African wild dog and the bush dog, on account of all three species' similar dentition. Subsequent authors, including Juliet Clutton-Brock, noted greater morphological similarities to canids of the genera Canis, Dusicyon and Alopex than to either Speothos or Lycaon, with any resemblance to the latter two being due to convergent evolution. Subsequent studies on the canid genome revealed that the dhole and African wild dog are closely related to members of the genus Canis, and that both are more closely related to grey wolves, coyotes, golden jackals and Ethiopian wolves than the more basal black-backed and side-striped jackals are. This closeness to Canis may have been confirmed in a menagerie in Madras where, according to zoologist Reginald Pocock, a dhole interbred with a golden jackal.
|Subspecies||Image||Trinomial authority||Common names||Description||Range||Synonyms|
|C. a. alpinus||Pallas, 1811||Southern dhole
|Large subspecies with bright red coat and narrow skull.||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), sumatrensis (Hodgson, 1863)|
|C. a. hesperius||Afanasjev and Zolotarev, 1935||Northern dhole
Tien Shan dhole
|Smaller than C. a. alpinus, with wider skull and lighter coloured winter fur.||Altai, Tien Shan and possibly Pamir and Kashmir||jason (Pocock, 1936)|
|C. a. sumatrensis||Hardwicke, 1821||Has short, coarse fur with no woolly underfur, and much black on the back.||Sumatra|
However, studies on dhole mtDNA and microsatellite genotype showed that there are no clear subspecific distinctions. Nevertheless, two major phylogeographic groupings were discovered in dholes of the Asian mainland, which likely diverged during a glaciation event. One population extends from South, Central, and North India (south of the Ganges) into Burma, and the other extends from India north of the Ganges into northeastern India, Burma, Thailand and the Malaysian Peninsula. The origin of dholes in Sumatra and Java is, as of 2005, unclear, as they show greater relatedness to dholes in India, Burma and China rather than with those in nearby Malaysia. In the absence of further data, the researchers involved in the study speculated that Javan and Sumatran dholes could have been introduced to the islands by humans.
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. 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". 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. 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.
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.
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.
Etymology and naming
The etymology of 'dhole' is unclear. The earliest possible written use of the word in English occurred in 1808 by soldier Thomas Williamson, who encountered the animal in Ramghur district. He stated that 'dhole' was a common local name for the species. In 1827, Charles Hamilton Smith claimed that it was derived from a language spoken in 'various parts of the East'. Two years later, Smith connected this word with Turkish: deli ‘mad, crazy’, and erroneously compared the Turkish word with Old Saxon: dol and Dutch: dol (cfr. also English: dull; German: toll), which are in fact from the Proto-Germanic *dwalaz ‘foolish, stupid’. Richard Lydekker wrote nearly 80 years later that the word was not used by the natives living within the species' range. The Merriam-Webster Dictionary theorises that it may have come from the Kannada: tōḷa (‘wolf’).
Local and indigenous names
|Linguistic group or area||Indigenous name|
豺蜀 (chai shu)
|Khmer||ចឰ បែ (chkai prey)|
|Lao||ໝາໃນ (ma nai)|
|Malayalam/Tamil||செந்நாய் (chen nai)
|Russian||Красный волк (krasnyi volk)
Ди́кая собака (dikaya sobaka)
|Telugu||అడవి కుక్క (resu kukka)
రేసు కుక్క (adavi-kutta)
|Thai||หมาใน (maa nay)|
|Vietnamese||chó sói lửa|
- Wozencraft, W. C. (2005). "Order Carnivora". In Wilson, D. E.; Reeder, D. M. Mammal Species of the World (3rd ed.). Johns Hopkins University Press. 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 2014.3. International Union for Conservation of Nature.
- Lindblad-Toh, K.; Wade, C. M.; Mikkelsen, T. S.; Karlsson, E. K.; Jaffe, D. B.; Kamal, M.; Clamp, M.; Chang, J. L. 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.
- Clutton-Brock, J., Corbet, G. G., and Hills, M. (1976). "A review of the family Canidae, with a classification by numerical methods". Bulletin of the British Museum of Natural History 29: 179–180.
- 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., pp. 566-86, ISBN 1-886106-81-9
- Zhang, H. & Chen, L. (2010). "The complete mitochondrial genome of dhole Cuon alpinus: phylogenetic analysis and dating evolutionary divergence within Canidae". Molecular Biology Reports 38 (3): 1651–1660.
- Fox 1984, p. 85
- Fox 1984, pp. 86–7
- Cohen, J. A. (1978). Cuon alpinus. Mammalian Species 100: 1-3.
- Durbin, L.S., Venkataraman, A., Hedges, S. & Duckworth, W. (2004). Dhole Cuon alpinus (Pallas 1811), in Sillero-Zubiri, C., Hoffmann, M. & Macdonald, D.W. (eds.) Canids: Foxes, Wolves, Jackals and Dogs: Status Survey and Conservation Action Plan. IUCN/SSC Canid Specialist Group. Gland, Switzerland and Cambridge, UK. x + pp. 210-219
- 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.
- (German) Thenius, E. (1955). "Zur Abstammung der Rotwölfe (Gattung Cuon Hodgson) [On the origins of the dholes (Genus Cuon Hodgson)]". Österreichische Zoologische Zeitschrift 5: 377–388.
- Fox 1984, pp. 61–2
- Fox 1984, pp. 41
- Fox 1984, p. 93
- Fox 1984, p. 95
- Fox 1984, p. 97
- 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. and Subedi, N. (2013). "First camera trap record of pack hunting dholes in Chitwan National Park, Nepal". Canid Biology & Conservation 16 (2): 4–7.
- Khatiwada, A. P., Awasthi, K. D., Gautam, N. P., Jnawali, S. R., Subedi, N., Aryal, A. (2011). "The Pack Hunter (Dhole): Received Little Scientific Attention". The Initiation 4: 8–13.
- Wangchuk, T. (2004). Predator-prey dynamics: the role of predators in the control of problem species. J. Bhutan Studies 10: 68–89.
- Thinley, P., Kamler, J. F., Wang, S. W., Lham, K., Stenkewitz, U. (2011). "Seasonal diet of dholes (Cuon alpinus) in northwestern Bhutan". Mammalian Biology 76: 518–520.
- Jenks, K. E., Songsasen, N. and P. Leimgruber (2012). "Camera trap records of dholes in Khao Ang Rue Nai Wildlife Sanctuary, Thailand". Canid News. online: 1–5.
- Fox 1984, pp. 81–2
- Walker, E. P., Nowak, R. M., Warnick, F. (1983). Walker's Mammals of the World. 4th ed. Baltimore: Johns Hopkins University Press.
- Fox 1984, p. 92
- Fox 1984, pp. 43–49
- Fox 1984, p. 80
- Fox 1984, p. 79
- 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, R. (1907). The game animals of India, Burma, Malaya, and Tibet. London: R. Ward Limited.
- Fox 1984, p. 63
- Fox 1984, p. 70
- Fox 1984, p. 51
- Pocock, R. I. (1941), Fauna of British India: Mammals volume 2, Taylor & Francis, pp. 146-63
- Fox 1984, pp. 58–60
- Perry, R. (1964). The World of the Tiger. London: Cassell.
- Mivart, G. (1890), Dogs, Jackals, Wolves and Foxes: A Monograph of the Canidæ, London : R.H. Porter : Dulau, pp. 177-88
- 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.
- 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.
- Finn, F. (1929). Sterndale's Mammalia of India. London: Thacker, Spink & Co.
- Shrestha, T. J. (1997). Mammals of Nepal: (with reference to those of India, Bangladesh, Bhutan and Pakistan). Kathmandu: Bimala Shrestha. ISBN 0-9524390-6-9.
- Nair, M. V. & Panda, S. K. (2013). Just Friends. Sanctuary Asia, Vol. XXXIII, No. 3
- Humphrey, S. R., Bain, J. R. (1990). Endangered Animals of Thailand. Gainesville: Sandhill Crane Press. ISBN 1-877743-07-0.
- Fox 1984, p. 109
- Pallas, P. S. (1811), Zoographia Rosso-Asiatica : sistens omnium animalium in extenso Imperio Rossico, et adjacentibus maribus observatorum recensionem, domicilia, mores et descriptiones, anatomen atque icones plurimorum, Petropoli : In officina Caes. Acadamiae Scientiarum Impress. MDCCCXI, pp. 34-5
- Hodgson, B. H. (1833). "Description and Characters of the Wild Dog of the Himalaya (Canis primævus)". Asiatic Researches. XVIII, Part 2: 221–237.
- Hodgson, B. H. (1842). "European Notices of Indian Canines, with Further Illustrations of the New Genus Cuon vel Chrysæus". Calcutta Journal of Natural History II: 205–209.
- (German) Schrenk, L. v. (1859), Reisen und forschungen im Amur-lande in den jahren 1854-1856, St. Petersburg : K. Akademie der Wissenschaften, pp. 48-50
- Kurtén, Björn (1968), Pleistocene mammals of Europe, Weidenfeld and Nicolson, pp. 111-114
- Ripoll, P. et al. (2010), Presence of the genus Cuon in upper Pleistocene and initial Holocene sites of the Iberian Peninsula: new remains identified in archaeological contexts of the Mediterranean region, Journal of Archaeological Science, 37: 437-450
- Petrucci, Mauro; Romiti, Serena; and Sardella, Raffaele. 2012. "The Middle-Late Pleistocene Cuon Hodgson, 1838 (Carnivora, Canidae) from Italy." Bollettino della Società Paleontologica Italiana, 51 (2); p 146. Article (in English) accessed at: http://paleoitalia.org/media/u/archives/07.Petrucci_et_al._2012_BSPI_512_1.pdf
- Kurtén, Björn (1980), Pleistocene mammals of North America, Columbia University Press, p. 172
- Simpson, G.G. (1945), The principles of classification and a classification of mammals, Bulletin of the American Museum of Natural History, 85:1–350
- Ellerman, J.R. & Morrison-Scott, T.C.S. (1966). Checklist of Palaearctic and Indian mammals, British Museum (Natural History), London, UK.
- Iyengar, A., Babu, V. N., Hedges, S., Venkataraman, A. B., Maclean, N. and P. A. Morin (2005). "Phylogeography, genetic structure, and diversity in the dhole (Cuon alpinus)". Molecular Ecology 14 (8): 2281–2297.
- van der Geer, A. A. E. (2008), Animals in stone: Indian mammals sculptured through time, BRILL, p. 188, ISBN 90-04-16819-2
- Smith, C. H. & Jardine, W. (1839). The natural history of dogs : canidae or genus canis of authors ; including also the genera hyaena and proteles, Vol. I. Edinburgh: W.H. Lizars.
- Skabelund, A. H. (2011). Empire of Dogs: Canines, Japan, and the Making of the Modern Imperial World. Cornell University Press, p. 85, ISBN 0801463246
- McIntosh, J. (2008). The ancient Indus Valley: new perspectives, p. 130, ABC-CLIO, ISBN 1-57607-907-4
- Williamson, T. (1808). Oriental field sports: being a complete, detailed, and accurate description of the wild sports of the East. Volume II. London: Orme.
- Smith, C. H. (1827). The class Mammalia. London: Geo. B. Whittaker.
- Orel, V. (2003), A Handbook of Germanic Etymology, Leiden, Boston: Brill, p. 81, ISBN 90-04-12875-1
- dhole. Merriam-Webster Dictionary.
- Fox 1984
- Andrew T. Smith, Yan Xie, Robert S. Hoffmann, Darrin Lunde, John MacKinnon, Don E. Wilson, W. Chris Wozencraft (2010). A Guide to the Mammals of China. Princeton University Press. pp. 418-19. ISBN 1400834112
- Fox, M. W. (1984). The Whistling Hunters: Field Studies of the Asiatic Wild Dog (Cuon Alpinus). Albany: State University of New York Press. ISBN 0-9524390-6-9.
|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