Dhole

Dhole[1]
Temporal range: Post-Pleistocene-Recent
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)
Dhole range

The dhole (Cuon alpinus) 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.[2]

The dhole is a highly social animal, living in large clans which occasionally split up into small packs to hunt.[3] 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.[4][5][6] Though fearful of humans, dhole packs are bold enough to attack large and dangerous animals such as wild boar, water buffalo, and even tigers.

Contents

Naming and etymology

Since these canids are called dholes only in English,[7] the etymology remains unclear,[8] but it probably may come from Kannada: tōḷa (‘wolf’).[9] Some 19th-century authors[10] 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’.[11]

Other names for the species include wild dogs,[12] whistling dogs, chennai,[13] red wolves[14] (not to be confused with Canis [lupus] rufus), red dogs[15] and mountain wolves.[16]

Vernacular names

Vernacular names include lal rakshas (red devil) and rakshur kukur (devil dog) in India, and jungli rakshas or hounds of Kali in Assam.[17] In the Himalayas, they are variously known as Bhaosa, Bhansa and Buansu.[7]

Evolution and taxonomy

Dholes are post-Pleistocene in origin, and are more closely related to jackals than they are to wolves.[19] It has been theorised that dholes became social animals as an adaptation to living with tigers and leopards.[20]

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 post-carnassial 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 Alopex than to African wild dogs and bush dogs.[19] A comparative study on dhole and other Canid mtDNA in 1997 showed that dholes diverged from the Lupus lupus lineage before the black-backed Jackal and the golden jackal diverged, a couple of million years before the domestication of the dog.[21]

Subspecies

Subspecies Trinomial authority Description Range Synonyms
Eastern[22] or Ussuri dhole[23]
Cuon a. alpinus		 Pallas, 1811 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.[23] Russian Far East, China, India, Nepal, Sikkim, Bhutan, Kashmir, Tibet, Mongolia, Myanmar, Thailand, Laos, Cambodia, Vietnam, Java and Indonesia adustus (Pocock, 1941)

antiquus (Matthew and 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)
†Late Pleistocene dhole
Cuon a. europaeus		 Bourguignat, 1875 The earliest form to evolve a single cusped, sharply trenchant tooth in place of the lower tubercular molar[24] 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[24] Heppenloch, Germany
Western[22] or Tien Shan dhole[25]
Cuon a. hesperius		 Afanasjev and Zolotarev, 1935 A small subspecies, with 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 surface of the limbs is sandy-yellow, while the flanks and inner sides of the limbs have little to no yellowish tint.[25] Transoxiana, Eastern Russia and China jason (Pocock, 1936)
†Early Middle Pleistocene dhole
Cuon a. priscus		 Thenius, 1954
Sumatran dhole
Cuon a. sumatrensis		 Hardwicke, 1821 A small subspecies, measuring only two feet in length, and standing 14 inches (360 mm) 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.[26] Sumatra and Indonesia

Physical description

Anatomy

Dholes have relatively short, heavy and massive skulls, with shortened facial regions, widely separated zygomatic arches and well-developed sagittal crests.[27] The frontal bone is inflated, and passes down onto the snout, giving the animals a convex rather than concave profile.[28] The masseter muscles are highly developed compared to other canid species, giving the face an almost hyena-like appearance.[29] The skull is broader, and has a shorter rostrum than that of domestic dogs and most other canids.[15] The dental formula is Upper: 3.1.4.2, lower: 3.1.4.2

The species uniquely has 6 rather than 7 lower molars.[30] The upper molars are weak, being 2-3 times smaller than those of wolves, and have only one cusp as opposed to 2-4 as is usual in canids,[27] an adaptation thought to improve shearing ability, thus allowing it to compete more successfully with kleptoparasites.[31] The canine teeth are slightly curved and short.[27]

Their limbs are moderately long, and their thoraxes proportional.[32] Along with African wild dogs, dholes are often referred to as "cat-like" canids, due to their long fine limbs and backbones.[30] They have great jumping and leaping abilities, being able to jump 3-3.5 m (10–12 ft) high, and leap 5–6 m (17–20 ft) long distances in one leap with a running start.[33] Their tails measure 16-17 inches long,[30] and are almost half the length of their bodies, nearly touching the ground when in full winter fur.[34] They are smaller than African wild dogs.[35] Weight ranges from 10 to 25 kg (22 to 55 lb), with males averaging about 4.5 kg (9.9 lb) heavier. 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.[36][37] Shoulder height is 42 to 55 cm (17 to 22 in).[38] 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 family, females have 12-14 teats rather than 10.[30] They are not as odorous as wolves, jackals and foxes, having a smaller number of anal scent glands.[39] It has been estimated that their stomachs can hold 6.5 lb (2.9 kg) of food.[40]

Fur

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-ocherous colour, with a dark brown tip. The summer coat is shorter, coarser and darker.[41] 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.[15]

Behaviour

Social and territorial behaviours

Dholes are more social than wolves,[18] 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.[3] 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.[6] 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.[42] Dholes are far less territorial than wolves, with pups from one clan often joining another without trouble once they mature sexually.[43] Clans typically number 5-12 individuals in India, though clans of 40 have been reported. In Thailand, clans rarely exceed three individuals.[15] 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.[44]

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.[15] Unlike wolf packs, dhole clans may contain more than one breeding female.[6] During mating, the female assumes a crouched, cat-like position. There is no "tug of war" characteristic of other canids when the male dismounts. Instead, the pair lie on their sides facing each other in a semicircular formation.[45] The gestation period lasts 60–63 days, with litter sizes averaging 4-6 pups.[15] Their growth rate is much faster than that of wolves, being similar in speed to that of coyotes. Pups are suckled until at least the age of 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 till the age of 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 8 months.[46]

Denning behaviour

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 3–4 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.[47] Like African wild dogs and dingoes, dholes will avoid killing prey close to their dens.[46]

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.[4][15][48] 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.[16] In Kashmir, they may hunt markhor,[48] and thamin in Burma.[15] Javan rusas are hunted in Java.[31] 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.[49] Like African wild dogs, but unlike wolves, dholes are not known to attack people.[50][51] 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.[52] In summertime in the Tien Shan Mountains, dholes eat large quantities of mountain rhubarb.[49] Bael fruits are also eaten.[53] Although opportunistic, dholes have a seeming aversion to hunting cattle and their calves.[54] 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.[55]

Before embarking on a hunt, clans go through elaborate prehunt social rituals involving nuzzling, body rubbing and homo and heterosexual mounting.[56] Dholes are primarily diurnal hunters, hunting in the early hours of the morning. They rarely hunt nocturnally, except on moonlit nights, indicating that they greatly rely on sight when hunting.[57] Though not as fast as jackals and foxes, they can chase their prey for many hours.[49] 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 metres.[58] When chasing fleet-footed prey, they run at a pace of 30 mph.[53] Dholes frequently drive their prey into water bodies, where the targeted animal's movements are hindered.[59]

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 hind quarters. They do not use a killing bite to the throat.[29] They occasionally blind their prey by attacking the eyes.[60] 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.[61] 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.[5] 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.[62] 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.[6] They are generally tolerant of scavengers at their kills.[63]

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.[64] 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).[65] 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 leopard kill both sexes more evenly (and tigers prefer larger prey altogether), dholes and tigers kill langurs rarely compared to leopards due to the leopard's greater arboreality, while leopards kill wild boar infrequently due to the inability of this relatively light predator to tackle aggressive prey of comparable weight.[65] 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.[50][66] 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.[67] Dhole packs may steal leopard kills, while leopards may kill dholes if they encounter them singly or in pairs.[48] 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.[20] It was once thought that dholes were 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.[68]

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 hind quarters.[48]

Though usually antagonistic toward wolves,[69] they may hunt and feed alongside one another.[53] They infrequently associate in mixed groups with golden jackals. Domestic dogs may kill dholes, though they will feed alongside them on occasion.[70]

Communication

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.[71] Other sounds include whines (food soliciting), growls (warning), screams, chatterings (both of which are alarm calls) and yapping cries.[72] In contrast to wolves, dholes do not howl.[73]

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.[44]

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. It is known that they may suffer from rabies, canine distemper, mange, trypanosomiasis, canine parvovirus, and endoparasites like cestodes and roundworms.[74]

Range

Historical

Dholes once ranged throughout most of South, East and Southeast Asia, extending from the Tien Shan and Altai Mountains and the Primorsky Krai southward through Mongolia, Korea, China, Tibet, Nepal, India, and south-eastwards into Myanmar and Indochina, Thailand, Malaysia, Sumatra and Java.[75]

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.[76] 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.[77]

Current

There are currently no reports of dholes in Russia, Mongolia, Kazakhstan, Kyrgystan and Tajikistan. There is one report of a dhole being captured in southern China's Jiangxi province. Dholes still occur in Tibet, particularly in south-east 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 north-west India. They are occasionally reported in the Ladakh area of Kashmir, contiguous with the Tibetan highlands and China. In Nepal, dholes were formerly recorded in Terai, including the Royal Chitwan National Park. Dholes were reported in the Dhorpatan Hunting Reserve in the late 1990s. 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 that 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. Dholes 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.[75]

Relationships with humans

Hunting and persecution

Certain people, such as the Kurumbas and some Mon Khmer speaking tribes will appropriate 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.[74] 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.[78] In Indochina, dholes suffer heavily from non-selective hunting techniques such as snaring.[74]

The fur trade does not pose a significant threat to dholes.[74] The people of India do not eat dhole flesh, and their fur is not considered overly valuable.[79] 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 3-4 silver rubles. In the early 20th century, dhole pelts reached 8 rubles in Manchuria. In Semirechye, fur coats made from dhole skin were considered the warmest, but were very costly.[73]

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 B.C.. They are shown waiting by a tree, with a woman or spirit trapped up it, a scene reminiscent of dholes treeing tigers.[80] 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.[39] It has been suggested that 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.[81]

Dholes appear in Rudyard Kipling's Red Dog, where they are portrayed as aggressive and bloodthirsty animals who 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 that with the exception of one animal, they remained shy and vicious even after 10 months.[52] 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.[82] A dhole may have been presented as a gift to Ibbi-Sin as tribute.[83]

Although genetic distance and mitochondrial DNA studies have proven that dogs descended from wolves, the dhole is still put forward by some researchers as a more likely candidate. Points raised in favour of the dhole ancestry cite the general domestic dog-like passiveness of the dhole in having its kills taken by humans, which contrasts with the behaviour of sympatric wolves, which will defend their prey aggressively against humans. It is also claimed that dhole skulls bear more similarities to dog skulls than wolves do, with the glaring exception of the dentition.[84]

References

Notes

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  2. ^ a b 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. Downloaded on 22 March 2009. Database entry includes justification for why this species is endangered
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  5. ^ a b Fox 1984, p. 63
  6. ^ a b c d Fox 1984, pp. 86–7
  7. ^ a b c d e f g h i j k l m n o p q r Lydekker 1907, p. 360
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  43. ^ Fox 1984, p. 92
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  45. ^ Fox 1984, p. 79
  46. ^ a b Fox 1984, p. 80
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  48. ^ a b c d Pocock 1941, p. 161
  49. ^ a b c Heptner & Naumov 1998, p. 583
  50. ^ a b Pocock 1941, p. 162
  51. ^ Heptner & Naumov 1998, p. 584
  52. ^ a b Mivart 1890, p. 181
  53. ^ a b c Shrestha 1996, p. 122
  54. ^ Fox 1984, p. 71
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  56. ^ Fox 1984, pp. 100–1
  57. ^ Fox 1984, p. 50
  58. ^ Fox 1984, p. 73
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Bibliography

External links