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The dhole /doʊl/ (Cuon alpinus) is a canid native to Central, South and Southeast Asia. Other English names for the species include Asiatic wild dog,[3] Indian wild dog,[4] whistling dog, red dog,[5] and mountain wolf.[6] It is genetically close to species within the genus Canis,[7](Fig. 10) though its skull is convex rather than concave in profile, it lacks a third lower molar[8] and the upper molars sport only a single cusp as opposed to two to four.[9] During the Pleistocene, the dhole ranged throughout Asia, Europe
Europe
and North America but became restricted to its historical range 12,000–18,000 years ago.[10] The dhole is a highly social animal, living in large clans without rigid dominance hierarchies[11] and containing multiple breeding females.[12] Such clans usually consist of 12 individuals, but groups of over 40 are known.[5] It is a diurnal pack hunter which preferentially targets medium and large sized ungulates.[13] In tropical forests, the dhole competes with tigers and leopards, targeting somewhat different prey species, but still with substantial dietary overlap.[14] It is listed as Endangered
Endangered
by the IUCN
IUCN
as populations are decreasing and are estimated at fewer than 2,500 adults. Factors contributing to this decline include habitat loss, loss of prey, competition with other species, persecution due to livestock predation and disease transfer from domestic dogs.[2]

Contents

1 Etymology
Etymology
and naming

1.1 Local and indigenous names

2 Discovery, taxonomy, and evolution

2.1 Subspecies

3 Characteristics 4 Distribution and habitat 5 Ecology and behaviour

5.1 Social and territorial behaviour 5.2 Denning 5.3 Reproduction and development 5.4 Hunting behaviour 5.5 Feeding ecology 5.6 Enemies and competitors 5.7 Diseases and parasites

6 Threats 7 Conservation 8 In culture and literature

8.1 Tameability

9 See also 10 References

10.1 Bibliography

11 External links

Etymology
Etymology
and naming[edit]

Captive Indian wild dogs resting

The etymology of "dhole" is unclear. The possible earliest 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.[15] In 1827, Charles Hamilton Smith claimed that it was derived from a language spoken in 'various parts of the East'.[16] 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),[17] which are in fact from the Proto-Germanic *dwalaz ‘foolish, stupid’.[18] Richard Lydekker
Richard Lydekker
wrote nearly 80 years later that the word was not used by the natives living within the species' range.[4] The Merriam-Webster
Merriam-Webster
Dictionary theorises that it may have come from the Kannada: tōḷa (‘wolf’).[19] Local and indigenous names[edit]

It has been suggested that this section be merged into Wiktionary:dhole#Translations. (Discuss) Proposed since December 2016.

Indigenous names for Cuon alpinus

Linguistic group or area Indigenous name

Assamese kuang-kukur[13] rang kukur[13]

Bengali bon-kutta[13] bon-kukur[13]

Bhutanese phara[13] phou[13]

Burmese tan-kwe[13]

Buryat Ӡурби (zurbi)[13]

Chinese 豺 (chai)[20] 豺蜀 (chai shu)[20] 豺狼 (chailang)[21]

Gilyak tschoramlatsch[22]

Gondi eram-naiko[4]

Gujarati કુત્ર (kutra)[13]

Gurkha ban-kukur[13]

Hindi adivi-kutta[13] son-kuta[13] sona-kutta[13] rasa-kutta[13] jangli-kutta[4]

Ho Kol tani[4]

Indonesian ajak[23] ajag[13] anjing hutan[13]

Javanese asu alas[13]

Kachin kyi-kwa-lam[13]

Kannada kadu nai[13] korku[13] bun-seeta[13] 'Kennai(ಕೆನ್ನಾಯಿ)[13] '

Kashmiri jangli-kuta[24] ram-hun[13] ban-kuta[13] bhansa[13]

Kazakh Чуе (chue)[13]

Khmer ឆ្កែព្រៃ (chkai prey)[13]

Korean 승냥이 (seungnyang-i)

Ladakh farra[13] siddaki[4]

Lao ໝາໃນ (ma nai)[13]

Lepcha sa-tun[13]

Malayalam vatai-karau[4]

Malay serigala[25] anjing hutan[25]

Meitei huithou[13]

Marathi kolsun[13] kolasna[4] kolasra[4] kolsa[4]

Mongolian Дшергул (dshergul)[13]

Nepali bwaso[13]

Odia balia kukura[13]

Russian Красный волк (krasnyi volk)[13] Ди́кая собака (dikaya sobaka)[13] Чикалка (chikalka)[13]

Tamil செந்நாய் (chen naai)[13]

Telugu రేచు కుక్క (rechu kukka)[13] reza-kutta[4] అడవి కుక్క (adavi-kukka)[4]

Thai หมาใน (ma nai)[13]

Tibetan farra[13] hazi[4]

Tungus Дергил (dzergil)[13]

Urdu jangli kutta[13]

Vietnamese chó sói lửa[13]

Discovery, taxonomy, and evolution[edit]

A Tiger
Tiger
Hunted by Wild Dogs (1807) by Samuel Howitt: This is one of the first illustrations of the species, featured in Thomas Williamson's Oriental Field Sports. The depiction, though, is based on Williamson's description of the animal as resembling the Indian pariah dog.

The species was first described in European 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
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 and in the region of the upper Lena River, though he wrote that it also occurred around the Yenisei
Yenisei
River, and that it occasionally crossed into China.[26][27] This northern Russian range reported by this "nearly impeccable" author Pallas, during the 18th and 19th centuries, is "considerably north" of where this species occurs today.[28] The British naturalist Brian Hodgson gave the dhole the binomial name Canis
Canis
primaevus and proposed that it was the progenitor of the domestic dog.[29] Hodgson later took note of the dhole's physical distinctiveness from the genus Canis
Canis
and assigned it to a new genus Cuon.[30]

Illustration (1859) by Leopold von Schrenck, one of the first accurate depictions of the species, based on a single skin purchased in the village of Dshare on the Amur[22]

The first study on the origins of the species was conducted by paleontologist Erich Thenius, who concluded that the dhole was a post- Pleistocene
Pleistocene
descendant of a golden jackal-like ancestor.[31] The earliest known member of the genus Cuon is the Chinese Cuon majori of the Villafranchian period. It resembled Canis
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
Europe
during the late Würm period,[32] though it may have survived up until the early Holocene
Holocene
in the Iberian Peninsula.[33] and at Riparo Fredian in northern Italy[34] The vast Pleistocene
Pleistocene
range of this species also included numerous islands in Asia
Asia
that this species no longer inhabits, such as Sri Lanka, Borneo, and possibly Palawan
Palawan
in the Philippines.[35][36][37][38][39] The fossil record indicates that the species also occurred in North America, with remains being found in Beringia
Beringia
and Mexico.[40]

Skeletal remains of a European dhole
European dhole
dating back to upper Würm period from Cova Negra de Játiva, Valencia, Spain

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
African wild dog
and the bush dog, on account of all three species' similar dentition.[41] Subsequent authors, including Juliet Clutton-Brock, noted greater morphological similarities to canids of the genera Canis, Dusicyon, and Alopex than to either Speothos
Speothos
or Lycaon, with any resemblance to the latter two being due to convergent evolution.[8] Some authors consider the extinct Canis subgenus named Xenocyon
Xenocyon
as ancestral of both genus Lycaon and genus Cuon.[42][43][44][45]:p149 Subsequent studies on the canid genome revealed that the dhole and African wild dog
African wild dog
are closely related to members of the genus Canis.[7] This closeness to Canis
Canis
may have been confirmed in a menagerie in Madras, where according to zoologist Reginald Pocock, a dhole interbred with a golden jackal.[46]

Phylogenetic tree
Phylogenetic tree
of the extant wolf-like canids

Caninae 3.5 Ma

3.0

2.7

1.9

1.6

1.3

1.1

Dog
Dog

Gray wolf
Gray wolf

Himalayan wolf
Himalayan wolf

Coyote
Coyote

African golden wolf
African golden wolf

Ethiopian wolf
Ethiopian wolf

Golden jackal
Golden jackal

Dhole
Dhole

African wild dog
African wild dog

2.6

Side-striped jackal
Side-striped jackal

Black-backed jackal
Black-backed jackal

Phylogenetic relationships between the extant wolf-like clade of canids based on nuclear DNA sequence data taken from the cell nucleus,[7][47] except for the Himalayan wolf, based on mitochondrial DNA sequences.[47][48] Timing in millions of years.[47]

Subspecies[edit] Historically, up to 10 subspecies of dholes have been recognised.[49] As of 2005[update], only three subspecies are recognised by MSW3.[1]

Subspecies Image Trinomial authority Common names Description Range Synonyms

C. a. alpinus (Nominate subspecies)

Pallas, 1811 Indian wild dog Southern dhole[50] Ussuri dhole[9] Large subspecies with bright red coat and narrow skull.[9] Far eastern Russia, Mongolia, China, Nepal, Indian subcontinent, Bhutan, Burma, Indochina
Indochina
and Java. adustus (Pocock, 1941), antiquus (Matthew & Granger, 1923), clamitans (Heude, 1892), dukhunensis (Sykes, 1831), fumosus (Pocock, 1936), grayiformis (Hodgson, 1863), infuscus (Pocock, 1936), javanicus (Desmarest, 1820), laniger (Pocock, 1936), lepturus (Heude, 1892), primaevus (Hodgson, 1833), rutilans (Müller, 1839)

C. a. hesperius

Afanasjev and Zolotarev, 1935 Northern dhole[50] Tien Shan
Tien Shan
dhole[9] Smaller than C. a. alpinus, with wider skull and lighter-coloured winter fur[9] Altai, Tien Shan
Tien Shan
and possibly Pamir and Kashmir jason (Pocock, 1936)

C. a. sumatrensis

Hardwicke, 1821 Sumatran dhole Has short, coarse fur with no woolly underfur, and much black on the back[50] Sumatra

However, studies on dhole mtDNA and microsatellite genotype showed 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
Thailand
and the Malaysian Peninsula. The origin of dholes in Sumatra
Sumatra
and Java
Java
is, as of 2005, unclear, as they show greater relatedness to dholes in India, Burma and China
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.[51] Characteristics[edit]

Dhole
Dhole
skull and molars illustrated by St. George Mivart
St. George Mivart
(1890)

In appearance, the dhole has been variously described as combining the physical characteristics of the grey wolf and red fox,[9] and as being "cat-like" on account of its long backbone and slender limbs.[31] It has a wide and massive skull with a well-developed sagittal crest,[9] and its masseter muscles are highly developed compared to other canid species, giving the face an almost hyena-like appearance.[52] The rostrum is shorter than that of domestic dogs and most other canids.[5] The species has six rather than seven lower molars.[53] 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,[9] an adaptation thought to improve shearing ability, thus allowing it to compete more successfully with kleptoparasites.[13] Adult females can weigh from 10 to 17 kg (22 to 37 lb), while the slightly larger male may weigh from 15 to 21 kg (33 to 46 lb). The mean weight of adults from three small samples was 15.1 kg (33 lb).[13][54][55][56] Occasionally, dholes may be sympatric with the Indian wolf
Indian wolf
(Canis lupus pallipes), which is one of the smallest races of gray wolf but is still approximately 25% heavier on average.[57][58] It stands 17–22 in (43–56 cm) at the shoulder and measures 3.0 ft (0.91 m) in body length. Like the African wild dog, its ears are rounded rather than pointed.[53] It has six or seven pairs of teats, sometimes eight.[9]

Subadult

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, and 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.[9] The dorsal and lateral guard hairs in adults measure 20–30 mm in length. Dholes in the Moscow Zoo
Moscow Zoo
moult once a year from March to May.[5] 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.[59] Other sounds include whines (food soliciting), growls (warning), screams, chatterings (both of which are alarm calls) and yapping cries.[60] In contrast to wolves, dholes do not howl or bark.[9] Dholes have a complex body language. Friendly or submissive greetings are accompanied by horizontal lip retraction and the lowering of the tail, as well as licking. Playful dholes open their mouths with their lips retracted and their tails held in a vertical position whilst assuming a play bow. Aggressive or threatening dholes pucker their lips forward in a snarl and raise the hairs on their backs, as well as keep their tails horizontal or vertical. When afraid, they pull their lips back horizontally with their tails tucked and their ears flat against the skull.[61]

Distribution and habitat[edit]

Dhole
Dhole
feeding on sambar carcass, Khao Yai National Park

In Central Asia, dholes primarily inhabit mountainous areas; in the western half of their range, they live mostly in alpine meadows and high-montane steppes high above sea level, while in the east, they mainly range 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.[9] The dhole might still be present in the Tunkinsky National Park
Tunkinsky National Park
in extreme southern Siberia
Siberia
near Lake Baikal.[62] It possibly still exists in the Primorsky Krai
Primorsky Krai
province in far-eastern Russia, where it was considered a rare and endangered species in 2004, with unconfirmed reports in the Pikthsa-Tigrovy Dom protected forest area; no sighting was reported in other areas such as the Mataisky Zakaznik forest since the late 1970s.[63] Currently, no other recent reports are confirmed of dhole being present in Russia, with no recent reports from Mongolia, Kazakhstan, Kyrgyzstan
Kyrgyzstan
or Tajikistan, though one specimen was caught in southern China's Jiangxi
Jiangxi
district. Also, in 2011 to 2013, local government officials and herders reported the presence of several packs at altitudes of 2,000 to 3,500 m near the Taxkorgan Reserve
Taxkorgan Reserve
in the Karakoram/ Pamir Mountains
Pamir Mountains
region of the Xinjiang Autonomous Region, on China’s border with Pakistan, Tajikistan, and Afghanistan.[64] Dholes have been also recently reported from the Altyn-Tagh
Altyn-Tagh
(Altun) Mountains in the southern portion of the Xinjiang Autonomous Region close to Tibet, as well.[65] It is unknown if dholes continue to inhabit Tien Shan, though they occur in small numbers in Gansu Province, with one pack being sighted in the Qilian Mountains
Qilian Mountains
within that province in 2006;[66] Camera-trap surveys in the Yanchiwan National Nature Reserve in the northern edge of this Gansu Province
Gansu Province
in 2013-2014 confirmed the continued presence of several packs and a female adult with pups in this area at altitudes around 2,500 to 4,000 m.[64] Dholes still occur in Tibet, and possibly also in North Korea. They once occurred in the alpine steppes extending into Kashmir
Kashmir
to the Ladakh
Ladakh
area, but have not been recorded in Pakistan.[2] They occur in most of India south of the Ganges, particularly in the Central Indian Highlands and the Western and Eastern Ghats. In northeast India, it is present in Arunachal Pradesh, Assam, Meghalaya, and West Bengal
West Bengal
and in the Indo-Gangetic Plain's Terai
Terai
region. Dhole populations in the Himalaya
Himalaya
and northwest India are fragmented.[2] In 2011, dhole packs were recorded by camera traps in the Chitwan National Park.[67] Its presence was confirmed in the Kanchenjunga Conservation Area in 2011 by camera-traps.[68] In Bhutan, dholes have recovered from a poisoning campaign during the 1970s, and became re-established in the 1990s.[69] Today they occur in the Jigme Dorji National Park.[70] Dholes still occur in northeastern Bangladesh's forest reserves in the Sylhet area, as well the Chittagong Hill Tracts in the southeast. These zones are unlikely to contain viable populations, considering most sightings involve small groups or solitary specimens, and they are likely decreasing in number due to the lack of prey.[2] The presence of dholes in Myanmar
Myanmar
was confirmed by camera-trapping in 11 areas, and alongside leopards, have apparently replaced tigers as the country's top predators.[2] In 2015, dholes and tigers were recorded by camera-traps for the first time in the hill forests of Karen State.[71] Their range is highly fragmented in the Malaysian Peninsula, Sumatra, Java, Vietnam, and Thailand.[2] In 2014, camera trap videos in the montane tropical forests at 2,000 m in the Kerinci Seblat National Park in Sumatra
Sumatra
revealed the continued presence of this species.[72] A camera trapping survey in the Khao Ang Rue Nai Wildlife Sanctuary
Khao Ang Rue Nai Wildlife Sanctuary
in Thailand
Thailand
from January 2008 to February 2010 revealed at least one healthy dhole pack.[73] In northern Laos, dholes have been studied (2012) in protected areas.[74] A disjunct population of this species has been reported from the area of Trabzon
Trabzon
and Rize
Rize
in northeastern Turkey
Turkey
near the border with Georgia in the 1990s by two Turkish zoologists.[75] Some authorities have accepted this report,[76] but others considered it to be unreliable.[2] Also, one single individual was claimed to have been shot in 2013 in the nearby Kabardino-Balkaria
Kabardino-Balkaria
Republic (a subject republic of Russia
Russia
immediately north of Georgia in the Central Caucasus); its remains (including a skull) were analyzed by a biologist from the Kabardino-Balkarian State University in May 2015, who concluded the skull was from a dhole.[77] Recently, in August 2015, researchers from the National Museum of Natural History from Sofia, Bulgaria
Bulgaria
(including Dr. Nikolai Spassov, the current director of this museum) and the Karadeniz Technical University
Karadeniz Technical University
began an expedition to track and document this possible Turkish population of dhole.[78] On October 12, 2015, this research team reported the preliminary conclusion that no real evidence exists of a living population of the dhole in Turkey
Turkey
(or in the Kabardino-Balkaria Republic), pending DNA analysis of samples from the original 1994 Serez-Eroglu skins.[79] Ecology and behaviour[edit] Social and territorial behaviour[edit]

Dholes playing, Pench Tiger
Tiger
Reserve

Dholes are more social than grey wolves,[9] 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.[11] 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.[80] 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.[12] Intragroup fighting is rarely observed.[81] Dholes are far less territorial than wolves, with pups from one clan often joining another without trouble once they mature sexually.[82] Clans typically number 5-12 individuals in India, though clans of 40 have been reported. In Thailand, clans rarely exceed three individuals.[5] Unlike other canids, there is no evidence of dholes using urine to mark their territories or travel routes. When urinating, dholes, especially males, may raise one hind leg or both to result in a handstand. Handstand urination is also seen in bush dogs (Speothos venaticus).[83] 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.[61] Denning[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.[84] Like African wild dogs and dingoes, dholes will avoid killing prey close to their dens.[85] Reproduction and development[edit]

Dhole
Dhole
pup, Kolmården Wildlife Park

In India, the mating season occurs between mid-October and January, while captive dholes in the Moscow Zoo
Moscow Zoo
breed mostly in February.[5] Unlike wolf packs, dhole clans may contain more than one breeding female.[12] More than one female dhole may den and rear their litters together in the same den.[81] 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.[86] The gestation period lasts 60–63 days, with litter sizes averaging 4–6 pups.[5] 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.[85] Maximum longevity in captivity is 15–16 years.[81] Hunting behaviour[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.[87] 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.[88] Though not as fast as jackals and foxes, they can chase their prey for many hours.[9] 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.[89] When chasing fleet-footed prey, they run at a pace of 30 mph.[9] Dholes frequently drive their prey into water bodies, where the targeted animal's movements are hindered.[90] 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.[91] They occasionally blind their prey by attacking the eyes.[92] Serows are among the only ungulate species capable of effectively defending themselves against dhole attacks, due to their thick, protective coats and short, sharp horns capable of easily impaling dholes.[4] They will tear open their prey's flanks and disembowel it, eating the heart, liver, lungs and some sections of the intestines. The stomach and rumen are usually left untouched.[93] 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.[94] Unlike wolf packs, in which the breeding pair monopolises food, dholes give priority to the pups when feeding at a kill, allowing them to eat first.[12] They are generally tolerant of scavengers at their kills.[95] Both mother and young are provided with regurgitated food by other pack members.[81]

Feeding ecology[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.[5][46][96] There is one record of a pack bringing down an Indian elephant
Indian elephant
calf in Assam, despite desperate defense of the mother resulting in numerous losses to the pack.[6] In Kashmir, they prey on markhor,[46] and thamin in Myanmar,[5] Malayan tapir, Sumatran serow
Sumatran serow
in Sumatra
Sumatra
and Malay Peninsula, and Javan rusa
Javan rusa
in Java.[13] In the Tien Shan
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 pig, musk deer, and reindeer, while in Primorye
Primorye
they feed on sika deer and goral, too. In Mongolia, they prey on argali and rarely Siberian ibex.[9] Like African wild dogs, but unlike wolves, dholes are not known to attack people.[9][46] 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.[50] In summertime in the Tien Shan
Tien Shan
Mountains, dholes eat large quantities of mountain rhubarb.[9] Although opportunistic, dholes have a seeming aversion to hunting cattle and their calves.[97] Livestock predation by dholes has been a problem in Bhutan
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
Oxen
are killed more often than cows, probably because they are given less protection.[98] Enemies and competitors[edit]

Dhole
Dhole
killed and cached in a tree by a leopard, India

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.[14] On some occasions, dholes may attack tigers. When confronted by dholes, tigers will seek refuge in trees or stand with their backs to a tree or bush, where they may be mobbed for lengthy periods before finally attempting escape. Escaping tigers are usually killed, while tigers which stand their ground have a greater chance of survival.[46] Tigers are dangerous opponents for dholes, as they have sufficient strength to kill a dhole with a single paw strike.[6] Dhole
Dhole
packs may steal leopard kills, while leopards may kill dholes if they encounter them singly or in pairs.[46] 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.[99] There are numerous records of leopards being treed by dholes.[81] Dholes sometimes drive tigers, leopards, and bears (see below) from their kills.[81] Dholes were once thought to be a major factor in reducing Asiatic cheetah
Asiatic cheetah
populations, though this is doubtful, as cheetahs live in open areas as opposed to forested areas favoured by dholes.[100] Dhole
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.[46] Though usually antagonistic toward wolves,[9] they may hunt and feed alongside one another.[101] There is at least one record of a lone wolf associating with a pair of dholes in Debrigarh Wildlife Sanctuary.[102] They infrequently associate in mixed groups with golden jackals. Domestic dogs may kill dholes, though they will feed alongside them on occasion.[103] Diseases and parasites[edit] 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
Toxocara canis
are present in their faeces. They may suffer from rabies, canine distemper, mange, trypanosomiasis, canine parvovirus, and endoparasites such as cestodes and roundworms.[13] Threats[edit] 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.[81] 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
British Raj
did so under the conviction that dholes were responsible for drops in game populations. Persecution of dholes still occurs with varying degrees of intensity according to region.[13] Bounties paid for dholes used to be 25 rupees, though this was reduced to 20 in 1926 after the number of presented dhole carcasses became too numerous to maintain the established reward.[104] In Indochina, dholes suffer heavily from nonselective hunting techniques such as snaring.[13] The fur trade does not pose a significant threat to dholes.[13] The people of India do not eat dhole flesh, and their fur is not considered overly valuable.[50] 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
Ussuriysk
during the late 1860s for a few silver rubles. In the early 20th century, dhole pelts reached eight rubles in Manchuria. In Semirechye, fur coats made from dhole skin were considered the warmest, but were very costly.[9] Conservation[edit] The dhole is protected under Schedule 2 of the Wildlife Protection Act, 1972. The creation of reserves under Project Tiger
Tiger
provided some protection for dhole populations sympatric with tigers. In 2014, the Indian government sanctioned its first dhole conservation breeding centre at the Indira Gandhi Zoological Park
Indira Gandhi Zoological Park
(IGZP) in Visakhapatnam.[105] The dhole has been protected in Russia
Russia
since 1974, though it is vulnerable to poison left out for wolves. In China, the animal is listed as a category II protected species under the Chinese wildlife protection act of 1988. In Cambodia, the dhole is protected from all hunting, while conservation laws in Vietnam
Vietnam
limit extraction and utilization.[2] In 2016, the Korean company Sooam Biotech was reported to be attempting to clone the dhole using dogs as surrogate mothers to help conserve the species.[106] In culture and literature[edit]

Russian ruble

Kazakhstani tenge

Three dhole-like animals are featured on the coping stone of the Bharhut
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.[107] The animal's fearsome reputation in India is reflected by the number of pejorative names it possesses in Hindi, which variously translate as "red devil", "devil dog", "jungle devil", or "hound of Kali".[6] According to zoologist and explorer Leopold von Schrenck, he had trouble obtaining dhole specimens during his exploration of Amurland, as the local Gilyaks greatly feared the species. This fear and superstition was not however shared by neighbouring Tungusic peoples. Von Schrenk speculated that this differing attitude towards dholes was due to the Tungusic people's more nomadic, hunter-gatherer lifestyle.[22] 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
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
Scaliger
as having lived in the forests of Montefalcone could have been based on dholes, as they were described as unlike wolves in habits, voice and appearance. The Montefalcone family's coat of arms had a pair of red dogs as supporters.[17] Dholes appear in Rudyard Kipling's Red Dog, where they are portrayed as aggressive and bloodthirsty animals which descend from the Deccan Plateau into the Seeonee Hills
Seeonee Hills
inhabited by Mowgli
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
Shere Khan
and Hathi
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
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.[108] A fictional version of the Dhole, imbued with supernatural abilities, appears in the Season 6 episode of The X-Files
The X-Files
titled Alpha. Dholes also appear as enemies in the game Far Cry 4, alongside other predators such as the Bengal tiger, honey badger, snow leopard, clouded leopard, Tibetan wolf
Tibetan wolf
and Asian black bear. They can be found hunting the player and other NPC's across the map, but are easily killed, being one of the weakest enemies in the game. They once again appear in the game Far Cry Primal, where they play similar roles as their counterparts in the previous game, but can now also be tamed and used in combat by Takkar, the main protagonist of the game. Tameability[edit] Brian Houghton Hodgson
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.[50] According to Richard Lydekker, adult dholes are nearly impossible to tame, though pups are docile and can even be allowed to play with domestic dog pups until they reach early adulthood.[4] A dhole may have been presented as a gift to Ibbi-Sin
Ibbi-Sin
as tribute.[109] See also[edit]

Wild Dog
Dog
Diaries

References[edit]

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1984, pp. 58–60 ^ Fox
Fox
1984, p. 71 ^ Johnsingh, A.J.T., Yonten, D. & Wangchuck, S. (2007). "Livestock- Dhole
Dhole
Conflict in Western Bhutan". Journal of the Bombay Natural History Society. 104 (2): 201–202. CS1 maint: Uses authors parameter (link) ^ 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
Bhutan
and Pakistan). Kathmandu: Bimala Shrestha. ISBN 0-9524390-6-9.  ^ Nair M. V., Panda S. K. (2013). "Just Friends". Sanctuary Asia. XXXIII: 3.  ^ Humphrey, S. R.; Bain, J. R. (1990). Endangered
Endangered
Animals of Thailand. Gainesville: Sandhill Crane Press. ISBN 1-877743-07-0.  ^ Fox
Fox
1984, p. 109 ^ Zoo to have conservation breeding centre for ‘dhole’, The Hindu (August 18, 2014) ^ Zastrow, Mark (8 February 2016). "Inside the cloning factory that creates 500 new animals a day". New Scientist. Retrieved 23 February 2016.  ^ van der Geer, A. A. E. (2008), Animals in stone: Indian mammals sculptured through time, BRILL, p. 188, ISBN 90-04-16819-2 ^ 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

Bibliography[edit]

Fox, M. W. (1984). The Whistling Hunters: Field Studies of the Asiatic Wild Dog
Dog
(Cuon Alpinus). Albany: State University of New York Press. ISBN 0-9524390-6-9. 

External links[edit]

Wikispecies
Wikispecies
has information related to Cuon alpinus

Wikimedia Commons has media related to Cuon alpinus.

Dhole
Dhole
Home Page ARKive
ARKive
– images and movies of the dhole Saving the dhole: The forgotten 'badass' Asian dog more endangered than tigers, The Guardian (25 June 2015) Photos of dhole in Bandipur

v t e

Extant Carnivora
Carnivora
species

Kingdom: Animalia Phylum: Chordata Class: Mammalia Infraclass: Eutheria Superorder: Laurasiatheria

Suborder Feliformia

Nandiniidae

Nandinia

African palm civet
African palm civet
(N. binotata)

Herpestidae (Mongooses)

Atilax

Marsh mongoose
Marsh mongoose
(A. paludinosus)

Bdeogale

Bushy-tailed mongoose
Bushy-tailed mongoose
(B. crassicauda) Jackson's mongoose
Jackson's mongoose
(B. jacksoni) Black-footed mongoose
Black-footed mongoose
(B. nigripes)

Crossarchus

Alexander's kusimanse
Alexander's kusimanse
(C. alexandri) Angolan kusimanse
Angolan kusimanse
(C. ansorgei) Common kusimanse
Common kusimanse
(C. obscurus) Flat-headed kusimanse
Flat-headed kusimanse
(C. platycephalus)

Cynictis

Yellow mongoose
Yellow mongoose
(C. penicillata)

Dologale

Pousargues's mongoose
Pousargues's mongoose
(D. dybowskii)

Galerella

Angolan slender mongoose
Angolan slender mongoose
(G. flavescens) Black mongoose
Black mongoose
(G. nigrata) Somalian slender mongoose
Somalian slender mongoose
(G. ochracea) Cape gray mongoose
Cape gray mongoose
(G. pulverulenta) Slender mongoose
Slender mongoose
(G. sanguinea)

Helogale

Ethiopian dwarf mongoose
Ethiopian dwarf mongoose
(H. hirtula) Common dwarf mongoose
Common dwarf mongoose
(H. parvula)

Herpestes

Short-tailed mongoose
Short-tailed mongoose
(H. brachyurus) Indian gray mongoose
Indian gray mongoose
(H. edwardsii) Indian brown mongoose
Indian brown mongoose
(H. fuscus) Egyptian mongoose
Egyptian mongoose
(H. ichneumon) Small Asian mongoose
Small Asian mongoose
(H. javanicus) Long-nosed mongoose
Long-nosed mongoose
(H. naso) Collared mongoose
Collared mongoose
(H. semitorquatus) Ruddy mongoose
Ruddy mongoose
(H. smithii) Crab-eating mongoose
Crab-eating mongoose
(H. urva) Stripe-necked mongoose
Stripe-necked mongoose
(H. vitticollis)

Ichneumia

White-tailed mongoose
White-tailed mongoose
(I. albicauda)

Liberiictus

Liberian mongoose
Liberian mongoose
(L. kuhni)

Mungos

Gambian mongoose
Gambian mongoose
(M. gambianus) Banded mongoose
Banded mongoose
(M. mungo)

Paracynictis

Selous' mongoose
Selous' mongoose
(P. selousi)

Rhynchogale

Meller's mongoose
Meller's mongoose
(R. melleri)

Suricata

Meerkat
Meerkat
(S. suricatta)

Hyaenidae (Hyenas)

Crocuta

Spotted hyena
Spotted hyena
(C. crocuta)

Hyaena

Brown hyena
Brown hyena
(H. brunnea) Striped hyena
Striped hyena
(H. hyaena)

Proteles

Aardwolf
Aardwolf
(P. cristatus)

Felidae

Large family listed below

Viverridae

Large family listed below

Eupleridae

Small family listed below

Family Felidae

Felinae

Acinonyx

Cheetah
Cheetah
(A. jubatus)

Caracal

Caracal
Caracal
(C. caracal) African golden cat
African golden cat
(C. aurata)

Catopuma

Bay cat
Bay cat
(C. badia) Asian golden cat
Asian golden cat
(C. temminckii)

Felis

European wildcat
European wildcat
(F. silvestris) African wildcat
African wildcat
(F. lybica) Jungle cat
Jungle cat
(F. chaus) Black-footed cat
Black-footed cat
(F. nigripes) Sand cat
Sand cat
(F. margarita) Chinese mountain cat
Chinese mountain cat
(F. bieti) Domestic cat (F. catus)

Leopardus

Ocelot
Ocelot
(L. pardalis) Margay
Margay
(L. wiedii) Pampas cat
Pampas cat
(L. colocola) Geoffroy's cat
Geoffroy's cat
(L. geoffroyi) Kodkod
Kodkod
(L. guigna) Andean mountain cat
Andean mountain cat
(L. jacobita) Oncilla
Oncilla
(L. tigrinus) Southern tigrina
Southern tigrina
(L. guttulus)

Leptailurus

Serval
Serval
(L. serval)

Lynx

Canadian lynx (L. canadensis) Eurasian lynx
Eurasian lynx
(L. lynx) Iberian lynx
Iberian lynx
(L. pardinus) Bobcat
Bobcat
(L. rufus)

Otocolobus

Pallas's cat
Pallas's cat
(O. manul)

Pardofelis

Marbled cat
Marbled cat
(P. marmorata)

Prionailurus

Fishing cat
Fishing cat
(P. viverrinus) Leopard
Leopard
cat (P. bengalensis) Sundaland leopard cat (P. javanensis) Flat-headed cat
Flat-headed cat
(P. planiceps) Rusty-spotted cat
Rusty-spotted cat
(P. rubiginosus)

Puma

Cougar
Cougar
(P. concolor)

Herpailurus

Jaguarundi
Jaguarundi
(H. yagouaroundi)

Pantherinae

Panthera

Lion
Lion
(P. leo) Jaguar
Jaguar
(P. onca) Leopard
Leopard
(P. pardus) Tiger
Tiger
(P. tigris) Snow leopard
Snow leopard
(P. uncia)

Neofelis

Clouded leopard
Clouded leopard
(N. nebulosa) Sunda clouded leopard
Sunda clouded leopard
(N. diardi)

Family Viverridae
Viverridae
(includes Civets)

Paradoxurinae

Arctictis

Binturong
Binturong
(A. binturong)

Arctogalidia

Small-toothed palm civet
Small-toothed palm civet
(A. trivirgata)

Macrogalidia

Sulawesi palm civet
Sulawesi palm civet
(M. musschenbroekii)

Paguma

Masked palm civet
Masked palm civet
(P. larvata)

Paradoxurus

Golden wet-zone palm civet (P. aureus) Asian palm civet
Asian palm civet
(P. hermaphroditus) Jerdon's palm civet (P. jerdoni) Golden palm civet
Golden palm civet
(P. zeylonensis)

Hemigalinae

Chrotogale

Owston's palm civet
Owston's palm civet
(C. owstoni)

Cynogale

Otter civet
Otter civet
(C. bennettii)

Diplogale

Hose's palm civet
Hose's palm civet
(D. hosei)

Hemigalus

Banded palm civet
Banded palm civet
(H. derbyanus)

Prionodontinae (Asiatic linsangs)

Prionodon

Banded linsang
Banded linsang
(P. linsang) Spotted linsang
Spotted linsang
(P. pardicolor)

Viverrinae

Civettictis

African civet
African civet
(C. civetta)

Genetta (Genets)

Abyssinian genet
Abyssinian genet
(G. abyssinica) Angolan genet
Angolan genet
(G. angolensis) Bourlon's genet
Bourlon's genet
(G. bourloni) Crested servaline genet
Crested servaline genet
(G. cristata) Common genet
Common genet
(G. genetta) Johnston's genet
Johnston's genet
(G. johnstoni) Rusty-spotted genet
Rusty-spotted genet
(G. maculata) Pardine genet
Pardine genet
(G. pardina) Aquatic genet
Aquatic genet
(G. piscivora) King genet
King genet
(G. poensis) Servaline genet
Servaline genet
(G. servalina) Haussa genet
Haussa genet
(G. thierryi) Cape genet
Cape genet
(G. tigrina) Giant forest genet
Giant forest genet
(G. victoriae)

Poiana

African linsang
African linsang
(P. richardsonii) Leighton's linsang
Leighton's linsang
(P. leightoni)

Viverra

Malabar large-spotted civet
Malabar large-spotted civet
(V. civettina) Large-spotted civet
Large-spotted civet
(V. megaspila) Malayan civet
Malayan civet
(V. tangalunga) Large Indian civet
Large Indian civet
(V. zibetha)

Viverricula

Small Indian civet
Small Indian civet
(V. indica)

Family Eupleridae

Euplerinae

Cryptoprocta

Fossa (C. ferox)

Eupleres

Eastern falanouc
Eastern falanouc
(E. goudotii) Western falanouc (E. major)

Fossa

Malagasy civet
Malagasy civet
(F. fossana)

Galidiinae

Galidia

Ring-tailed mongoose
Ring-tailed mongoose
(G. elegans)

Galidictis

Broad-striped Malagasy mongoose
Broad-striped Malagasy mongoose
(G. fasciata) Grandidier's mongoose
Grandidier's mongoose
(G. grandidieri)

Mungotictis

Narrow-striped mongoose
Narrow-striped mongoose
(M. decemlineata)

Salanoia

Brown-tailed mongoose
Brown-tailed mongoose
(S. concolor) Durrell's vontsira (S. durrelli)

Suborder Caniformia
Caniformia
(cont. below)

Ursidae (Bears)

Ailuropoda

Giant panda
Giant panda
(A. melanoleuca)

Helarctos

Sun bear
Sun bear
(H. malayanus)

Melursus

Sloth bear
Sloth bear
(M. ursinus)

Tremarctos

Spectacled bear
Spectacled bear
(T. ornatus)

Ursus

American black bear
American black bear
(U. americanus) Brown bear
Brown bear
(U. arctos) Polar bear
Polar bear
(U. maritimus) Asian black bear
Asian black bear
(U. thibetanus)

Mephitidae

Conepatus (Hog-nosed skunks)

Molina's hog-nosed skunk
Molina's hog-nosed skunk
(C. chinga) Humboldt's hog-nosed skunk
Humboldt's hog-nosed skunk
(C. humboldtii) American hog-nosed skunk
American hog-nosed skunk
(C. leuconotus) Striped hog-nosed skunk
Striped hog-nosed skunk
(C. semistriatus)

Mephitis

Hooded skunk
Hooded skunk
(M. macroura) Striped skunk
Striped skunk
(M. mephitis)

Mydaus

Sunda stink badger
Sunda stink badger
(M. javanensis) Palawan
Palawan
stink badger (M. marchei)

Spilogale (Spotted skunks)

Southern spotted skunk
Southern spotted skunk
(S. angustifrons) Western spotted skunk
Western spotted skunk
(S. gracilis) Eastern spotted skunk
Eastern spotted skunk
(S. putorius) Pygmy spotted skunk
Pygmy spotted skunk
(S. pygmaea)

Procyonidae

Bassaricyon (Olingos)

Eastern lowland olingo
Eastern lowland olingo
(B. alleni) Northern olingo
Northern olingo
(B. gabbii) Western lowland olingo
Western lowland olingo
(B. medius) Olinguito
Olinguito
(B. neblina)

Bassariscus

Ring-tailed cat
Ring-tailed cat
(B. astutus) Cacomistle
Cacomistle
(B. sumichrasti)

Nasua (Coatis inclusive)

White-nosed coati
White-nosed coati
(N. narica) South American coati
South American coati
(N. nasua)

Nasuella (Coatis inclusive)

Western mountain coati (N. olivacea) Eastern mountain coati (N. meridensis)

Potos

Kinkajou
Kinkajou
(P. flavus)

Procyon

Crab-eating raccoon
Crab-eating raccoon
(P. cancrivorus) Raccoon
Raccoon
(P. lotor) Cozumel raccoon
Cozumel raccoon
(P. pygmaeus)

Ailuridae

Ailurus

Red panda
Red panda
(A. fulgens)

Suborder Caniformia
Caniformia
(cont. above)

Otariidae (Eared seals) (includes fur seals and sea lions) ( Pinniped
Pinniped
inclusive)

Arctocephalus

South American fur seal
South American fur seal
(A. australis) Australasian fur seal (A. forsteri) Galápagos fur seal
Galápagos fur seal
(A. galapagoensis) Antarctic fur seal
Antarctic fur seal
(A. gazella) Juan Fernández fur seal
Juan Fernández fur seal
(A. philippii) Brown fur seal
Brown fur seal
(A. pusillus) Guadalupe fur seal
Guadalupe fur seal
(A. townsendi) Subantarctic fur seal
Subantarctic fur seal
(A. tropicalis)

Callorhinus

Northern fur seal
Northern fur seal
(C. ursinus)

Eumetopias

Steller sea lion
Steller sea lion
(E. jubatus)

Neophoca

Australian sea lion
Australian sea lion
(N. cinerea)

Otaria

South American sea lion
South American sea lion
(O. flavescens)

Phocarctos

New Zealand sea lion
New Zealand sea lion
(P. hookeri)

Zalophus

California sea lion
California sea lion
(Z. californianus) Galápagos sea lion
Galápagos sea lion
(Z. wollebaeki)

Odobenidae ( Pinniped
Pinniped
inclusive)

Odobenus

Walrus
Walrus
(O. rosmarus)

Phocidae (Earless seals) ( Pinniped
Pinniped
inclusive)

Cystophora

Hooded seal
Hooded seal
(C. cristata)

Erignathus

Bearded seal
Bearded seal
(E. barbatus)

Halichoerus

Gray seal (H. grypus)

Histriophoca

Ribbon seal
Ribbon seal
(H. fasciata)

Hydrurga

Leopard
Leopard
seal (H. leptonyx)

Leptonychotes

Weddell seal
Weddell seal
(L. weddellii)

Lobodon

Crabeater seal
Crabeater seal
(L. carcinophagus)

Mirounga (Elephant seals)

Northern elephant seal
Northern elephant seal
(M. angustirostris) Southern elephant seal
Southern elephant seal
(M. leonina)

Monachus

Mediterranean monk seal
Mediterranean monk seal
(M. monachus) Hawaiian monk seal
Hawaiian monk seal
(M. schauinslandi)

Ommatophoca

Ross seal
Ross seal
(O. rossi)

Pagophilus

Harp seal
Harp seal
(P. groenlandicus)

Phoca

Spotted seal
Spotted seal
(P. largha) Harbor seal
Harbor seal
(P. vitulina)

Pusa

Caspian seal
Caspian seal
(P. caspica) Ringed seal
Ringed seal
(P. hispida) Baikal seal
Baikal seal
(P. sibirica)

Canidae

Large family listed below

Mustelidae

Large family listed below

Family Canidae
Canidae
(includes dogs)

Atelocynus

Short-eared dog
Short-eared dog
(A. microtis)

Canis

Side-striped jackal
Side-striped jackal
(C. adustus) African golden wolf
African golden wolf
(C. anthus) Golden jackal
Golden jackal
(C. aureus) Coyote
Coyote
(C. latrans) Gray wolf
Gray wolf
(C. lupus) Black-backed jackal
Black-backed jackal
(C. mesomelas) Red wolf
Red wolf
(C. rufus) Ethiopian wolf
Ethiopian wolf
(C. simensis)

Cerdocyon

Crab-eating fox
Crab-eating fox
(C. thous)

Chrysocyon

Maned wolf
Maned wolf
(C. brachyurus)

Cuon

Dhole
Dhole
(C. alpinus)

Lycalopex

Culpeo
Culpeo
(L. culpaeus) Darwin's fox
Darwin's fox
(L. fulvipes) South American gray fox
South American gray fox
(L. griseus) Pampas fox
Pampas fox
(L. gymnocercus) Sechuran fox
Sechuran fox
(L. sechurae) Hoary fox
Hoary fox
(L. vetulus)

Lycaon

African wild dog
African wild dog
(L. pictus)

Nyctereutes

Raccoon
Raccoon
dog (N. procyonoides)

Otocyon

Bat-eared fox
Bat-eared fox
(O. megalotis)

Speothos

Bush dog
Bush dog
(S. venaticus)

Urocyon

Gray fox
Gray fox
(U. cinereoargenteus) Island fox
Island fox
(U. littoralis)

Vulpes (Foxes)

Bengal fox
Bengal fox
(V. bengalensis) Blanford's fox
Blanford's fox
(V. cana) Cape fox
Cape fox
(V. chama) Corsac fox
Corsac fox
(V. corsac) Tibetan sand fox
Tibetan sand fox
(V. ferrilata) Arctic fox
Arctic fox
(V. lagopus) Kit fox
Kit fox
(V. macrotis) Pale fox
Pale fox
(V. pallida) Rüppell's fox
Rüppell's fox
(V. rueppelli) Swift fox
Swift fox
(V. velox) Red fox
Red fox
(V. vulpes) Fennec fox
Fennec fox
(V. zerda)

Family Mustelidae

Lutrinae (Otters)

Aonyx

African clawless otter
African clawless otter
(A. capensis) Oriental small-clawed otter
Oriental small-clawed otter
(A. cinerea)

Enhydra

Sea otter
Sea otter
(E. lutris)

Hydrictis

Spotted-necked otter
Spotted-necked otter
(H. maculicollis)

Lontra

North American river otter
North American river otter
(L. canadensis) Marine otter
Marine otter
(L. felina) Neotropical otter
Neotropical otter
(L. longicaudis) Southern river otter
Southern river otter
(L. provocax)

Lutra

Eurasian otter
Eurasian otter
(L. lutra) Hairy-nosed otter
Hairy-nosed otter
(L. sumatrana)

Lutrogale

Smooth-coated otter
Smooth-coated otter
(L. perspicillata)

Pteronura

Giant otter
Giant otter
(P. brasiliensis)

Mustelinae (including badgers)

Arctonyx

Hog badger
Hog badger
(A. collaris)

Eira

Tayra
Tayra
(E. barbara)

Galictis

Lesser grison
Lesser grison
(G. cuja) Greater grison
Greater grison
(G. vittata)

Gulo

Wolverine
Wolverine
(G. gulo)

Ictonyx

Saharan striped polecat
Saharan striped polecat
(I. libyca) Striped polecat
Striped polecat
(I. striatus)

Lyncodon

Patagonian weasel
Patagonian weasel
(L. patagonicus)

Martes (Martens)

American marten
American marten
(M. americana) Yellow-throated marten
Yellow-throated marten
(M. flavigula) Beech marten
Beech marten
(M. foina) Nilgiri marten
Nilgiri marten
(M. gwatkinsii) European pine marten
European pine marten
(M. martes) Japanese marten
Japanese marten
(M. melampus) Sable
Sable
(M. zibellina)

Pekania

Fisher (P. pennanti)

Meles

Japanese badger
Japanese badger
(M. anakuma) Asian badger
Asian badger
(M. leucurus) European badger
European badger
(M. meles)

Mellivora

Honey badger
Honey badger
(M. capensis)

Melogale (Ferret-badgers)

Bornean ferret-badger
Bornean ferret-badger
(M. everetti) Chinese ferret-badger
Chinese ferret-badger
(M. moschata) Javan ferret-badger
Javan ferret-badger
(M. orientalis) Burmese ferret-badger
Burmese ferret-badger
(M. personata)

Mustela (Weasels and Ferrets)

Amazon weasel
Amazon weasel
(M. africana) Mountain weasel
Mountain weasel
(M. altaica) Stoat
Stoat
(M. erminea) Steppe polecat
Steppe polecat
(M. eversmannii) Colombian weasel
Colombian weasel
(M. felipei) Long-tailed weasel
Long-tailed weasel
(M. frenata) Japanese weasel
Japanese weasel
(M. itatsi) Yellow-bellied weasel
Yellow-bellied weasel
(M. kathiah) European mink
European mink
(M. lutreola) Indonesian mountain weasel
Indonesian mountain weasel
(M. lutreolina) Black-footed ferret
Black-footed ferret
(M. nigripes) Least weasel
Least weasel
(M. nivalis) Malayan weasel
Malayan weasel
(M. nudipes) European polecat
European polecat
(M. putorius) Siberian weasel
Siberian weasel
(M. sibirica) Back-striped weasel
Back-striped weasel
(M. strigidorsa) Egyptian weasel
Egyptian weasel
(M. subpalmata)

Neovison (Minks)

American mink
American mink
(N. vison)

Poecilogale

African striped weasel
African striped weasel
(P. albinucha)

Taxidea

American badger
American badger
(T. taxus)

Vormela

Marbled polecat
Marbled polecat
(V. peregusna)

Taxon identifiers

Wd: Q132585 ARKive: cuon-alpinus BioLib: 1882 EoL: 328688 Fossilworks: 45383 GBIF: 2434317 iNaturalist: 42101 ITIS: 183831 IUCN: 5953 MSW: 14000793 NCBI: 68

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