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This page is about the species of wild dog. For H. P. Lovecraft's fictional monster, see Dhole (Cthulhu Mythos). For the town in France, see Cuon, Maine-et-Loire.
Dhole[1]
Temporal range: Post-Pleistocene-Recent
Not a fox.png
Captive C. alpinus, Zooparc de Trégomeur, Brittany, France
Conservation status
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Carnivora
Family: Canidae
Subfamily: Caninae
Genus: Cuon
Hodgson, 1838
Species: C. alpinus
Binomial name
Cuon alpinus
(Pallas, 1811)
Cuon-alpinus-map.png
Dhole range


The dhole[a] (Cuon alpinus), also known as the red dog or Indian wild dog, is a monospecific canid native to Central and Southeast Asia. Although genetically close to dogs of the genus Canis,[9](Fig. 10) it is morphologically distinguished by its distinctive skull and teeth.[10] 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.[11]

[12]


[13]

[14]


Discovery, taxonomy and evolution[edit]

Illustration (1859) by Leopold von Schrenck, one of the first authentic depictions of the species, based on a single skin purchased in the village of Dshare on Amur.[15]
Skeletal remains of a dhole dating back to upper Würm period from Cova Negra de Játiva, Valencia, Spain.

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, 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.[16]

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.[17] 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 tubercular teeth, 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 tubercular tooth 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,[18] though it may have survived in the Iberian Peninsula up until the early Holocene.[19] The fossil record indicates that the species also occurred in North America, with remains being found in Beringia and Mexico.[20]

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.[21] 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.[10] 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.[9] 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.[22]





Side-striped jackal



Black-backed jackal








Golden jackal





Dog



Grey wolf




Coyote





Ethiopian wolf




Dhole





African wild dog






Subspecies[edit]

Historically, up to ten subspecies of dhole have been recognised.[23] As of 2005,[24] only three subspecies are recognised by MSW3. However, studies on dhole mtDNA and microsatellite genotype showed that there were no clear subspecies 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 display lower genetic distances with 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.[25]

Subspecies Trinomial authority Common names Description Range Synonyms
C. a. alpinus

(Nominate subspecies)

Pallas, 1811 Ussuri dhole[14]
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 Tien Shan dhole[14] Smaller than C. a. alpinus, with wider skull and lighter coloured winter fur.[14] Altai, Tien Shan and possibly Pamir and Kashmir jason (Pocock, 1936)
C. a. sumatrensis Hardwicke, 1821 Sumatra

Physical description[edit]

In appearance, the dhole has been variously described as combining the physical characteristics of the grey wolf and red fox,[14] and as being "cat-like" on account of its long backbone and slender limbs.[17] It has a wide and massive skull with a well-developed sagittal crest,[14] and its masseter muscles are highly developed compared to other canid species, giving the face an almost hyena-like appearance.[26] The rostrum is shorter than that of domestic dogs and most other canids.[13] The species has six rather than seven lower molars.[27] 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,[14] an adaptation thought to improve shearing ability, thus allowing it to compete more successfully with kleptoparasites.[12] Adults may weigh over 40 lbs, with females usually weighing 10 lbs 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.[27] It has 6-7 teats, sometimes eight.[14]

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.[14] 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.[13]

Behaviour[edit]

Social and territorial behaviours[edit]

Dholes playing, Pench Tiger Reserve.

Dholes are more social than grey wolves,[14] 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.[28] 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.[29] 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.[30] Intragroup fighting is rarely observed.[31] Dholes are far less territorial than wolves, with pups from one clan often joining another without trouble once they mature sexually.[32] Clans typically number 5-12 individuals in India, though clans of 40 have been reported. In Thailand, clans rarely exceed three individuals.[13] 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.[33]

Reproduction and development[edit]

In India, the mating season occurs between mid-October and January, while captive dholes in the Moscow Zoo breed mostly in February.[34] Unlike wolf packs, dhole clans may contain more than one breeding female.[30] More than one female dhole may den and rear their litters together in the same den.[35] 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.[36] The gestation period lasts 60–63 days, with litter sizes averaging 4-6 pups.[13] 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.[37] Maximum longevity in captivity is 15-16 years.[38]

Hunting and feeding behaviours[edit]

Dholes attacking a sambar, Bandipur National Park

Before embarking on a hunt, clans go through elaborate prehunt social rituals involving nuzzling, body rubbing and homo- and heterosexual mounting.[39] 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.[40] Though not as fast as jackals and foxes, they can chase their prey for many hours.[41] 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.[42] When chasing fleet-footed prey, they run at a pace of 30 mph.[14] Dholes frequently drive their prey into water bodies, where the targeted animal's movements are hindered.[43]

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.[26] They occasionally blind their prey by attacking the eyes.[44] 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.Cite error: A <ref> tag is missing the closing </ref> (see the help page). 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.[45] 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.[30] They are generally tolerant of scavengers at their kills.[46]

Both mother and young are provided with regurgitated food by other pack members.[47]

Communication[edit]

Ecology[edit]

Habitat[edit]

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.[48] Like African wild dogs and dingoes, dholes will avoid killing prey close to their dens.[37]

Diet[edit]

Dholes feeding on a chital, Bandipur National Park

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.[49][13][50] 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.[51] In Kashmir, they may hunt markhor,[22] and thamin in Burma.[13] Javan rusas are hunted in Java.[12] 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.[14] Like African wild dogs, but unlike wolves, dholes are not known to attack people.[22]Cite error: A <ref> tag is missing the closing </ref> (see the help page). In summertime in the Tien Shan Mountains, dholes eat large quantities of mountain rhubarb.[14] Although opportunistic, dholes have a seeming aversion to hunting cattle and their calves.[52] 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.[53]

Enemies and competitors[edit]

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[54] 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).[54] 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.[54] 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.[55][56] 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.[57] Dhole packs may steal leopard kills, while leopards may kill dholes if they encounter them singly or in pairs.[50] 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.[58] There are numerous records of leopards being treed by dholes.[59] Dholes sometimes drive tiger, leopards, and bears (see below) from their kills.[60] 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.[61]

Dhole packs occasionally attack Asiatic black bears and sloth bears. When attacking bears, dholes will attempt to prevent them from seeking refuge in caves, and lacerate their hindquarters.[50]

Though usually antagonistic toward wolves,[62] they may hunt and feed alongside one another.[63] They infrequently associate in mixed groups with golden jackals. Domestic dogs may kill dholes, though they will feed alongside them on occasion.[64]

Range and conservation[edit]

Relationships with humans[edit]

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.[65] 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, which they called tschoramlatsch. This fear and superstition was not however shared by neighbouring Tungusic peoples, who called it dshargul. Von Schrenk speculated that this differing attitude towards dholes was due to the Tungusic people's more nomadic, hunter-gatherer lifestyle.[15] 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.[5]

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.

Notes[edit]

  1. ^ The etymology of 'dhole' is unclear. The earliest possible written use of the word occurred in 1808 by soldier Thomas Williamson, who encountered the animal in West Bengal's Ramghur district. He stated that 'dhole' was a common local name for the species.[3] In 1827, Charles Hamilton Smith claimed that it was derived from a language spoken in 'various parts of the East'.[4] 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)[5], which are in fact from the Proto-Germanic *dwalaz ‘foolish, stupid’.[6] Richard Lydekker wrote nearly 80 years later that the word was not used by the natives living within the species' range.[7] The Merriam-Webster Dictionary theorises that it may have come from the Kannada: tōḷa (‘wolf’).[8]

References[edit]

  1. ^ 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. 
  2. ^ 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. In: IUCN 2008. IUCN Red List of Threatened Species. Retrieved 22 March 2009. Database entry includes justification for why this species is endangered
  3. ^ Williamson, T. (1808), Oriental field sports: being a complete, detailed, and accurate description of the wild sports of the East, Vol. II, Orme, pp. 1-12
  4. ^ Smith, C. H. (1827), The class Mammalia (1827), London : Printed for Geo. B. Whittaker, p. 326
  5. ^ a b 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, pp. 167-87
  6. ^ Orel, Vladimir (2003), A Handbook of Germanic Etymology, Leiden, Boston: Brill, p. 81, ISBN 90-04-12875-1 
  7. ^ Lydekker, R. (1907), The game animals of India, Burma, Malaya, and Tibet, London, R. Ward, limited, pp. 360-65
  8. ^ dhole. Merriam-Webster Dictionary.
  9. ^ a b 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. 
  10. ^ a b Clutton-Brock, J., Corbet, G.G., and Hills, M. (1976). "A review of the family Canidae, with a classification by numerical methods." Bull. Brit. Mus. Nat. Hist. 29, 179–180.
  11. ^ 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.
  12. ^ a b c 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
  13. ^ a b c d e f g Cohen, J. A. (1978), Cuon alpinus. Mammalian Species 100: 1-3.
  14. ^ a b c d e f g h i j k l m 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
  15. ^ a b Schrenk, L. v. (1859), Reisen und forschungen im Amur-lande in den jahren 1854-1856, St. Petersburg : K. Akademie der Wissenschaften, pp. 48-50
  16. ^ 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
  17. ^ a b Thenius, E. (1954), On the origins of the dholes, Osterreich Zoologie Zietshcrift, 5, 377–388
  18. ^ Kurtén, Björn (1968), Pleistocene mammals of Europe, Weidenfeld and Nicolson, pp. 111-114
  19. ^ 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
  20. ^ Kurtén, Björn (1980), Pleistocene mammals of North America, Columbia University Press, p. 172
  21. ^ Simpson, G.G. (1945), The principles of classification and a classification of mammals, Bulletin of the American Museum of Natural History, 85:1–350
  22. ^ a b c Pocock, R. I. (1941), Fauna of British India: Mammals volume 2, Taylor & Francis, pp. 146-63
  23. ^ Ellerman, J.R. & Morrison-Scott, T.C.S. (1966), Checklist of Palaearctic and Indian mammals, British Museum (Natural History), London, UK.
  24. ^ 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. 
  25. ^ Iyengar, A., et al. (2005), Phylogeography, genetic structure, and diversity in the dhole (Cuon alpinus), Molecular Ecology 14.8 (2005): 2281-2297
  26. ^ a b Fox 1984, pp. 61–2
  27. ^ a b Fox 1984, pp. 41
  28. ^ Fox 1984, p. 85
  29. ^ Fox 1984, pp. 81–2
  30. ^ a b c Fox 1984, pp. 86–7
  31. ^ Nowak, Ronald M; and Paradiso, John L. Walker's Mammals of the World. 4th ed. Baltimore: Johns Hopkins University Press; 1983. p. 963
  32. ^ Fox 1984, p. 92
  33. ^ Cite error: The named reference f97 was invoked but never defined (see the help page).
  34. ^ Cite error: The named reference cohen was invoked but never defined (see the help page).
  35. ^ Nowak, Ronald M; and Paradiso, John L. Walker's Mammals of the World. 4th ed. Baltimore: Johns Hopkins University Press; 1983. p962
  36. ^ Fox 1984, p. 79
  37. ^ a b Fox 1984, p. 80
  38. ^ Nowak, Ronald M; and Paradiso, John L. Walker's Mammals of the World. 4th ed. Baltimore: Johns Hopkins University Press; 1983. p. 962
  39. ^ Fox 1984, pp. 100–1
  40. ^ Fox 1984, p. 50
  41. ^ Cite error: The named reference h583 was invoked but never defined (see the help page).
  42. ^ Fox 1984, p. 73
  43. ^ Fox 1984, p. 67
  44. ^ 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. 
  45. ^ Fox 1984, p. 70
  46. ^ Fox 1984, p. 51
  47. ^ Nowak, Ronald M; and Paradiso, John L. Walker's Mammals of the World. 4th ed. Baltimore: Johns Hopkins University Press; 1983. p963
  48. ^ Fox 1984, pp. 43–49
  49. ^ Fox 1984, pp. 58–60
  50. ^ Cite error: The named reference r161 was invoked but never defined (see the help page).
  51. ^ Perry 1968, p. 147
  52. ^ Fox 1984, p. 71
  53. ^ Johnsingh, A.J.T., Yonten, Deki & Wangchuck, Sangay (2007). "Livestock-Dhole Conflict in Western Bhutan". J. Bombay Nat. Hist. Soc. 104 (2): 201. 
  54. ^ a b c 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. 
  55. ^ Pocock 1941, p. 162
  56. ^ Perry 1968, p. 149
  57. ^ Perry 1968, p. 150
  58. ^ Cite error: The named reference venkataraman was invoked but never defined (see the help page).
  59. ^ Nowak, Ronald M; and Paradiso, John L. Walker's Mammals of the World. 4th ed. Baltimore: Johns Hopkins University Press; 1983. p962
  60. ^ Nowak, Ronald M; and Paradiso, John L. Walker's Mammals of the World. 4th ed. Baltimore: Johns Hopkins University Press; 1983. p963
  61. ^ Finn 1929, p. 120
  62. ^ Heptner & Naumov 1998, p. 585
  63. ^ Shrestha 1996, p. 122
  64. ^ Humphrey & Bain 1990, p. 572
  65. ^ van der Geer, A. A. E. (2008), Animals in stone: Indian mammals sculptured through time, BRILL, p. 188, ISBN 90-04-16819-2

Bibliography[edit]

External links[edit]

Category:Canines Category:Mammals of Southeast Asia Category:Mammals of South Asia Category:Mammals of China Category:Megafauna of Eurasia Category:Carnivora of Malaysia