The Asiatic Wild Dog or Dhole
Distribution Habitat/Territory Names Subspecies Description Propagation Pack Life Bibliography
This member of the dog family is nearly unknown to the western world. It is rather dog-like in its outward appearance and was once (however fleetingly) thought to be the ancestor of the domesticated dog. In reality, some of the features of its skull and dentition are so different from those of Canis familiaris that it is unlikely that the Asiatic Wild Dog played any role in the evolution of the domestic dog. At present, it is accorded the status of an independent single-species genus. In the early 19th century, when the first reports and descriptions appeared in print, the dhole was so common and numerous in India that it was considered vermin and was persecuted mercilessly. From 1912 to 1972, there was an Indian government program that offered a bounty for killed dholes. (see ‘Dhole in Danger’) Today, the drastically reduced dhole populations are legally Ueno Zoo, Tokyo--Photo © Jim Davis protected everywhere.
The vast range of historic Cuon alpinus encompasses central Asia east of longitude 70o E and south of latitude 50o N (i.e., southern Siberia and parts of China and Mongolia), the Indian subcontinent, Indo-China, and the Indonesian islands of Java, Panaitan and Sumatra. There are no credible reports of sightings of Cuon alpinus in Pakistan or Japan or in Borneo or Sri Lanka (formerly Ceylon).
With the exception of peninsular India, little is known about the dhole’s present distribution. Writing in 1967, Igor Sosnovskii underlined the rarity of the species in the then Soviet Union by stating that "not one specimen has been delivered during the past 20 years to the purchasing posts of Zoobycdinenie"; but he also allows that the dhole’s Russian habitat is "virtually inaccessible to scientists" and has effectively precluded "adequate study." Wilson/Reeder, without documentation, report Cuon alpinus hesperius (= the Russian subspecies from the Altai Mountains) as extinct.
There seem to be no accounts on the current dhole populations in China, Indo-China or Malaysia. In 1997, the World Conservation Monitoring Centre (WCMC) called for, among other conservation steps, the "assessment of the status of the species in Myanmar, the former USSR and China." (see ‘World Conservation Monitoring Centre – The Socialist Republic of Viet Nam’, p. 12). In his 1985 study, Johnsingh passes on a personal communication from H. G. Hundley, according to whom "dholes [= Cuon alpinus infuscus and Cuon alpinus adustus] are still common in most of the forested tracts of Burma," a note of hope that may have caused the WCMC to include Myanmar among the regions where an assessment of the dhole population is needed. There is no current information on whether any progress has been made. The Centre’s overall pessimistic prediction: "In the long-term, populations outside Protected Areas are unlikely to survive." (ibid.)
For India we have A. J. T. Johnsingh’s 1985 study that shows Cuon alpinus dukhunensis still to be common, especially in the wildlife parks and reserves of the Central Indian Highlands and in the southern states of India. And this situation appears to be stable. In an e-mail to the author, dated July 1999, Dr. Johnsingh states: "The conservation status of the dhole is as good as it was 20 years ago." In East India, with the exception of the Caro Hills area of Meghalaya (Assam) where they are reportedly still common, dholes are either "very rare" (Johnsingh, 1985: 204) or "extinct" (Alderton, 1998: 162).
The prehistoric distribution is detailed by E.Thenius (1954). The oldest fossil records for Cuon were found in China near Beijing and date from the early Pleistocene epoch. They were first described and named Cuon dubius (‘doubtful Cuon’) by Teilhard de Chardin in 1940.
Somewhat younger, but still belonging to the early Pleistocene, are Cuon dubius stehlini (‘Stehlin’s doubtful Cuon’) from Rosières (France), Cuon alpinus fossilis (‘fossil Cuon’, Nehring, 1890) from the Heppenloch (= Gutenberg Cave in Southern Germany), and Cuon priscus (‘archaic Cuon’, Thenius, 1954) from Hundsheim (Austria).
Cuon alpinus europaeus is the name given to a number of fossil finds from the European late Pleistocene, with Cuon alpinus sardous (‘Cuon alpinus of Sardinia’) being a smaller, more delicate Mediterranean form of this.
These data suggest a paneurasian distribution of pre-historic Cuon. [Back to the beginning]
Habitat and Home Territory
The Asiatic Wild Dog lives primarily in alpine regions (hence the name Cuon alpinus). The first dhole specimens of the Moscow zoo were trapped in the province of Chinghai in China at an elevation of approximately 3000 m (Sosnovskii, 1967: 120) and, in India, they "have been sighted at altitudes up to 2100 m" (Cohen, 1978: 2). They prefer dense forests (Russia, Indonesia) and thick scrub jungle (India) but are (were?) also found in open country (Tibet). R. Keller describes the terrain where the members of his expedition observed a pack of dholes for about 20 days in August/September 1972 in the Kanha National Park (central India) as the "Kanha-Wiesen" (meadows) and as a "durch offene Salwälder unterbroche Savanne" (a savannah interspersed with open stands of Sal trees, 1973: 129). He adds, however, that dholes are not permanent residents of the park. -- Dholes shy away from human habitation with fear of man and the more likely presence of prey animals in remoter areas being equally plausible suppositions.
The size of the home territory, or hunting area, of a dhole pack is difficult to determine because type of terrain and availability of prey animals tend to affect it; "Large area" is a term used repeatedly in the literature to describe it (see Cohen, 1977: 146). However, more detailed estimates suggest that the dhole’s home range is actually rather small: Keller (1973: 129) estimated that the pack that his expedition observed, hunted an area of 16 km2. In its "First Report from the Field – January 1999" (http://www.dcpbase.demon.co.uk/dcp/fieldrep.htm), the Dhole Conservation Project (dcp) reports the minimum home range areas of the two packs under observation in a National Park in central India as 17 and 23 km2. More detailed information is provided by Johnsingh (1982: 443) who found that the pack he studied in the Bandipur Tiger Reserve, Karnataka, covered an area of up to 40 km2 (= home range), half of it more intensively (= core area); during the denning season the hunting area was reduced from more than 15 km2 to 11 km2. Fox (1984: 41f.) postulates a correlation between the size of the area hunted and the seasons of the year: larger during the time of the rains and more restricted during the dry season (March to early May), which, of course, largely coincides with the denning season.
The literature has no reports of territorial marking or defense of territory by a dhole pack except, perhaps, Johnsingh’s observation (1982: 448 f.) that neighboring dhole packs did not invade "his" pack’s core area even when it had left to hunt somewhere else. The absence (or weak development) of territoriality may facilitate interactions between different family groups and enhance exogamy (Fox, 1984: 92).
There are indications, such as abandoned dens, that dhole packs do not necessarily "stay put" in any given area but rather move around as human presence becomes threatening or as the prey animals change location. [Back to the beginning]
The Many Names of Cuon alpinus
The only generally accepted designation for the species is the Greek/Latin combination Cuon alpinus (‘mountain dog’). To be sure, Peter Simon Pallas, who was the first to describe it (at least for the western world), named it Canis alpinus and a number of other early naturalists also attributed it to the genus Canis. But ever since H. B. Hodgson established Cuon as a separate genus in 1838, these early classifications must be considered inaccurate and obsolete. Still, one does find them occasionally, mostly in non-zoological contexts, and lest someone think that they refer to a canid other than Cuon alpinus, they are listed here. Canis javanicus, Canis sumatrensis, and Canis (also Cuon) rutilans (‘Red dog’) all refer to the Indonesian and Canis indicus to the Indian form of Cuon alpinus. Canis primaevus was Hodgson’s earlier name (1833) for Cuon alpinus primaevus, which presently also includes his Cuon grayiformes of Sikhim (‘Gray’s cuon’, in Gray, 1863). Cuon javanicus is Mivart’s (1890) and, in his footsteps, Pocock’s (1936) unsuccessful designation for the species; today, the adjective javanicus is applied only to the Javanese race of the Asiatic Wild Dog – Cuon alpinus javanicus.
The 11 proposed subspecies/races of Cuon alpinus (Cuon alpinud alpinus, Cuon alpinis adustus, Cuon alpinus. dukhunensis, Cuon alpinus fumosus, Cuon alpinus hesperius, Cuon alpinus infuscus, Cuon alpinus javanicus, Cuon alpinus laniger, Cuon alpinus lepturus, Cuon alpinus primaevus, Cuon alpinus sumatrensis,) are discussed in more detail in the section on subspecies.
Cuon alpinus europaeus, Cuomn alpinus sardous, Cuon alpinus fossilis, Cuon priscus, Cuon dubius, Cuon dubius stehlini refer to fossil forms of the genus Cuon and are discussed briefly in the section on prehistoric distribution of Cuon alpinus.
Note:Ever so often, one encounters the term Cyon; this is the philologically preferable but entirely unsuccessful emendation of Cuon proposed by Blanford (1888).
a) Asiatic Wild Dog: Descriptive but too encompassing as large parts of western and northern Asia lie outside of the range of Cuon alpinus.
b) Red dog: A designation, although frequent in local names, best avoided since Cuon alpinus is neither a dog nor red in all its subspecies.
c) Dhole: In the original and narrower sense, dhole refers to the Indian wild dog of the Dekkan (Cuon alpinus dukhunensis), and that is the definition most often found in English language dictionaries. In the wider sense, it also stands for all of Cuon alpinus and is used in such a manner, for example, by the Dhole Conservation Project. The first attested use of the word dhole occurred in 1827 and is attributed to Col. C. Hamilton Smith: "the Dhole or Wild Dog of the East Indies." The etymology of the word is considered unexplained (cf. Oxford English Dictionary, sub verbo). There is, however, the Kannada word tôla with its definition "a wolf" (F. Kittle. A Kannada-English Dictionary. 1894, p. 759). Kannada, or Kanarese, is the principal Dravidian language of the Karnataka district of Mysore in southwest India, home of Bandipur National Park where dholes can be observed to this day. The phonetics seem to fit: All dentals are unavoidably somewhat aspirated (whether or not reflected by spelling), and the t/d shift is unremarkable (cf. German Tor and English door) and even could be explained as the result of imperfect oral communication. The English rendering "a wolf" in Kittle’s dictionary suggests that he took tôla to be the name of a specific form of wolf rather than having the generic meaning "wolf" (not unlike the definition "dhole: a wild dog"). In other words: "a wolf" is Kittle’s interpretation of a native word and as such should not disqualify tôla as source for dhole.
d) Indian Wild Dog, Deccan Wild Dog, Burmese Wild Dog, Nepal Wild Dog, Java Wild Dog, Siberian Wild Dog, Sumatra Wild Dog: These are all names of various geographic subspecies/races of Cuon alpinus and are sorted out in the section on the subspecies.
Regional / local names
Note:The following terms were found here and there in the literature. Many could not be verified. Their transliterations from non-latin alphabets are, in all likelihood, neither uniform in method nor accurate throughout. Their ‘literal meanings’, due to inadequate dictionary support, can not be warranted.
ajag (with variant spellings adjag, ajak): Indonesian, ‘forest dog’
anjing hutan (kutar?): Indonesian, Malay, ‘forest dog’
aNTikam: Tamil, ‘(a) wild dog’
ban-kuta: Hindustani, Kashmiri; ban kutta: Bengali, ‘forest dog’
ban-kukur: Bengali, Gurkhali, ‘forest dog’
búánsú (with variant spellings bhansa, buansua): Kashmiri, ‘wolf’
bun-secta: Korku (a Munda language of Central India)
bwanso: Nepali / Nepalese, ‘wolf’
chennai (with variant spellings ken-nai, chen-nai): Tamil/Canarese, ‘red dog’
chó lu?:, Vietnamese
chó røng: Vietnamese, ‘wild dog,’ ‘jungle dog’
hahmasai-kuta: Malayalam (Drawidian language of SW India)
jungli-kuta: Hindustani, Kashmiri, ‘forest dog’
kokku: Tamil, ‘wild dog’
kolarsi: Malayalam (Drawidian language of SW India)
kolsa: Malayalam (Drawidian language of SW India)
kolsun: Marathi (principal language in the Indian State of Maharashtra)
kotsun: Malayalam (Drawidian language of SW India)
kuang-kukur: Assami / Assamese
kyi-kwa-lam: Kachin (a language of northernmost Burma)
maa paa: Thai
phara: Bhutanese, Tibetan
rang-kukur: Assami / Assamese, ‘forest dog’
rasa-kuta: Hindi, ‘wary dog’
sa-tun: Lepcha (language spoken in Sikkim)
serigala (with variant sirgala) : Indonesian, Malay, ‘wolf.’ (Note: Canis lupus is not indigenous to Indonesia.)
son kutta (with variants son-kuta, sona-kuta): Hindi, ‘golden dog’
vanna-kooka: Telegu (a Drawidian language of Central India), ‘jungle/forest dog’
Cuon alpinus in some European languages other than English
Dutch: de alpenhond ‘mountain dog’, de wilde hond van India ‘Indian wild dog’
French: le dhole, le loup de Java ‘wolf of Java’
German: der Rothund, der Rotwolf ‘red dog or wolf’
Italian: il cane rosso ‘red dog’, il dhole
Russian: volk krasni (or kracni) ‘red wolf’
Spanish: salvaje hindú (‘wild Hindu’?), el dhole
Note:One scientific (?) name found in the literature could not be verified: The reference for Anurocyon clamitans (Heude, 1892: 102) reported by Cohen (1978:1) and Pocock (1936: 33 with footnote) could not be found on the information provided, and one hesitates to accept the name on trust –‘shouting tailless dog’ sounds so very ‘un-dhole-like’. And Pocock’s footnote that the designation is based on a specimen "which accidentally had lost half its tail" is not credible: Who would name a "new species" after an accidental character? And if it were to be true it would tend to undermine Heude’s standing as a scientist (which may well have been Pocock’s intent). Pocock himself suggests (ibid., 41) that Anurocyon is but a seasonal variation of Cuon lepturus (Heude, 1888, see below), an identification that is repeated without additional debate by Ellerman/Morrison-Scott (1966: 233). Whatever the case, the designation Anurocyon clamitans be best forgotten – at least in regard to Cuon alpinus.
[Back to the beginning]
Subspecies / Races
There are only two true subspecies, i.e., subspecies that are defined not only by their locality but also by some differentiating physical character: Cuon alpinus alpinus and Cuon alpinus hesperius. The various other subspecies designations refer to regional forms, or races, of Cuon alpinus alpinus that differ only in coat color and coat texture.
Several of these classifications are based on exceedingly small numbers of specimens; for example, fumosus, infuscus, laniger and lepturus are each derived from one single specimen/skin. One cannot help but wonder if the taxonomists were not somewhat overenthusiastic.
The following table is abstracted chiefly from information provided by the web site of the Dhole Conservation Project (dcp) and is supplemented with other pertinent data from the literature and/or expert informants.
Subspecies Cuon alpinus alpinus
Notes: (1) This group, excepting the nominate (sub)species Cuon alpinus alpinus, also has been called Southern Dhole (Mivart, 1890: 177-188); Pocock, 1936: 35).
(2) Blanford (1888:143-147) separated out Cuon alpinus dukhunensis as a distinguishable form, thus arriving at three (sub)species, a division that lives on in M. Burton (1968: 147)
|Scientific and Common Names||Local Name(s)||Distribution||Description||Comments|
|Cuon alpinus alpinus
Eastern Asiatic wild dog
|volk krasni ‘red wolf’||middle reaches of the Amur
river and Ussiriland , or:
South East Russia: from the Eastern Sayans and the Trans-Baikal area to the Far East
thick tawny red coat (bright rusty-red in winter), grayish neck, and ochre muzzle
observed by Pallas in vicinity of Udskoi (Udskii) ostrog ‘U. settlement’; present-day Udskoya on the Uda river ?
Cuon alpinus ad- ustus ‘tan/brown dog’
Burmese Wild Dog Wild dog of Indo-China
|Kyi-kwa-lam, Kachin||northern Myanmar (Burma) and Indo-China||reddish brown coat||
same as Cuon rutilans (Müller 1839) border to C..a. infuscus in Southern Burma is undefined
| Cuon alpinus
dukhu-nensis ‘C..a. of the Dekkan’
Indian Wild Dog
|chennai, Tamil, ‘red
kolsun, Marathison kutta, Hindi, ‘wild dog’
rang kukur, Assaami‘forest dog’
peninsular India south of Ganges River
dense forests and thick scrub jungle, at altitudes to 2100 m
|reddish coat, short hair
between pads, black whiskers.
For more details see sections ‘Measure-ments’ and ‘Descrip-tion’
|Cuon alpinus fumosus ‘sooty C.a.’||nyar, Chinese||western Szechwan, China and Mongolia||luxuriant yellowish red coat with dark back and gray neck||
|Cuon alpinus infuscus
‘brown C. a.’
Burmese wild dog
| chó røng, Vietnamese, ‘wild dog, jungle dog’
Sirgala. Ajing-kutar Malay
|southern Myanmar (Burma), Thailand, Malaysia, Vietnam||
dark brown coat, with distinctive cranial features:
"the muzzle above the anterior half of pm4 being exceptionally wide and prominent and the frontals and nasals exceptionally narrow as compared with the length of the skull" (Pocock, 1936: 38 f.)
Border to C.a. adustus in northern Burma is undefined.
|Cuon alpinus javanicus
Wild Dog of Java
| ajag (also
ajak, adjag), Indonesian, ‘forest dog’
anjing hutan, Indo-nesian, ‘forest dog’)
serigala, Indonesian, ‘wolf’
Islands of Java and Panatia (Indonesia)
|short bright-red coat with regional color variations||Desmarest, 1820
same as Canis rutilans (Müller, 1839) ‘red dog’,
not included in Eller-man/Morrison-Scott
|Cuon alpinus laniger
ram-hun, ban-kuta, bhansa all Kashmiri
|Kashmir and southern Tibet (Lhasa)||
full yellowish gray coat; distinguished from primaevus by its much fuller, softer coat and much paler color
Johnsingh (1985: 207): "no information [ on population] available"
| Cuon alpinus
Poyang Lake, south of the Yangtze River. Kiangsi, China
uniformly red above and on flanks, lower neck brown mixed with dirty white
Canis lepturus (Heude 1888)
Pocock (1936: 41): "of doubtful identity"
| Cuon alpinus
Wild Dog of Nepal
sa-tun, Lepcha (Sikkim)
|Himalayan regions of Kumaon, Nepal, Sikkim and Bhutan||longer redder coat than dukhunensis, long hair between pads, black whiskers||
Johnsingh (1985: 207): "very rare"
|Cuon alpinus suma- trensis Wild Dog of Sumatra||ajag, (also ajak,
adjag), Indonesian, ‘forest dog’
anjing hutan, Indo-nesian, ‘forest dog’
serigala, sirgalaa, Indonesian/Malay,‘wolf’,
Island of Sumatra (Indonesia)
|short bright-red coat with dark whiskers, long hairs between pads||
not included in Eller-man/Morrison-Scott
b) Subspecies Cuon alpinus hesperius:
Pocock (1936: 51-52) named this subspecies Cuon javanicus jason (a designation that did not stick) and distinguished it from all other forms of the Asiatic Wild Dog as follows:
"[T]he upper cheek-teeth, apart from the canine, are individually appreciably larger than those of the other available skulls of the northern thick-coated races, m2 being exceptionally well developed in width, a little wider than the basal length of the canine. The skull itself also shows a combination of minor differences. The nasals are wider posteriorly, their narrowed proximal portion is short, forming a nearly equilateral instead of an isosceles triangle, and, owing to the shortness of the nasal processes of the frontals, the junction of the nasals with the maxillae is longer, 19 mm. as compared with 13 mm. in the Ussuri skull (alpinus), 12 and 9 in the Kashmir skull (laniger), and from 7 to 13 in skulls assigned to primaevus, in which, as in the Kashmir skull, the length may vary considerably on the two sides. The skull is also broad in its interorbital region and across the muzzle above pm4, and the mandible has a quadrate coronoid process, unlike that of any other skull of the genus."
Note: The author was unable to obtain a copy of the 1935 article by Afanasiev/Zolotarey which, reportedly, establishes the same two true subspecies.
|Scientific and Common Names||Local Name(s)||Distribution||Description||Comments|
Western Asiatic Wild Dog
Siberian Red Dog
|volk krasni ‘red wolf’||Aksai district of
Semiryechensk region, Eastern Russian Turkestan
(or: Altai and Tian Shan mountains)
|long bright yellow ticked coat, white underside, pale whiskers; smaller than Eastern Asiatic wild dog||
same as Pocock’s (1936: 51 f.) Cuon javanicus jason;
reported as extinct in the Altai and Tian Shan mountains by Wilson/Reeder
[Back to the beginning]
Cuon vs. Canis
The genus Cuon differs from most members of the genus Canis in the greater number of mammae (six or seven pairs vs. the typical five), the more rounded external ears, and the following cranial and dental characters (taken from Pocock, 1936: 34) of which only the first has a direct bearing on the outward appearance of the animal:
the relatively broader premolar region of the muzzle, i.e., the muzzle begins to widen noticeably immediately beyond the canine teeth [cf. figure 2 in Cohen (1979)],
the longer anterior palatine foramina,
the expansion of the nasals in their proximal half,
the presence of one instead of two cusps in the heel of the lower carnassial (m1),
the reduction in size of m2 and m2,
the suppression of m3.
Note: Items d-f are discussed in slightly more detail below under "Teeth."
Note: The details in the following descriptions are based on the Indian form of the Asiatic Wild Dog because it is the (only) one that has been studied with any degree of thoroughness. In the literature, quantitative measurements vary considerably; the numbers given here represent the extreme values rather than averages of those measurements. All data regarding relative/proportional measurements have been taken from the article by Juliet Clutton-Brock et al. -- For pictures/photos Google search dhole and, on the Results page, click on Images.
Height at shoulders: 42 to 55 cm (The literature does not differentiate between males and females; presumably, the latter occupy the lower end of the range given.)
Weight: 15 to 20 kg (males), 10 to 13 kg (females)
Length of skull: 17.4 to 18.8 cm (measured ventrally from occipital condyle to point of "chin")
Zygomatic width: 10.3 to 11.8 cm (= widest part of head)
Length of rostrum: 7.4 to 8.0 cm (measured from stop (juncture of forehead and rostrum) to tip of nose)
Length of head and body: 88 to 113 cm
Length of tail: 28 to 50 cm
Number of chromosomes: 78 (i.e., same as Canis and Lycaon)
Life span: 12+ years in captivity (best guess)
General appearance: Cuon alpinus is a medium-sized, dog-like canid (larger than the jackal but smaller than the wolf, substantially longer than high, with a somewhat spitz-like head and a long bushy tail. Coat color varies with regions and ranges from brownish gray to mahogany red, often with extensive areas of lighter to white fur.
Expression: alert and intelligent and, with its light eyes, somewhat fierce.
Head: Triangular; skull broad between ears with well developed interparietal crest; foreface very short in relation to skull width; bones of forehead and upper jaws are "swollen," producing a concave ("dish-faced") profile. Noticeable but not pronounced stop. Nose leather black.
Eyes: Hooded (i.e., with heavy upper eyelid); round pupil, iris amber or light brown.
Ears: Set on high; erect, rounded (similar to Lycaon) with white hair on the periphery; heavily furred on inside and outside.
Teeth: I 3/3, C 1/1, PM 4/4, M 2/2: i.e., there are three incisors, one canine, four premolars and two molars in each quadrant (=one-half of both the upper and lower jaws). This is basically the "normal" dentition of the canids with the exception that the "usual" lower third molar (m3) is missing on both sides so that the total number of teeth is reduced from the usual 42 to 40. The crushing and grinding capability of the dhole’s dentition is diminished further by the relative small size of the post-carnassial molars and, more importantly, by a modification of the lower carnassial (= Dens sectorius, first molar, m1): "The inner cusp of the talonid is missing, so that this part of the tooth does not form a basin but a subsidiary blade" (Ewer, 1967: 41). These features, according to Ewer, "suggest a highly predacious habit with diminished importance of vegetable foods".-- Due to the shortness of the muzzle, the overall position of the teeth is rather crowded.
Neck: Short in relation to combined length of thorax and lumbar region (35%); of the canids, only the Arctic Fox (Alopex lagopus) has a shorter neck (34%).
Forequarters: In comparison to other canids, forelegs fairly short in relation to body length (60% of length of cervical to lumbar vertebrae). Shoulders well laid back with a short straight upper arm.
Body: Level topline, chest broad and brisket above elbows. No marked tuckup. Ribs well sprung.
Hindquarters: Hind legs fairly short in relation to length of body spine (70%, well towards the lower end among the canids). Upper thigh (femur) longer than lower thigh (tibia), a feature shared only with the Dingo and Bush Dog (Speothos venaticus). Heavily muscled with enormous drive/thrust.
Note: The Dhole’s jumping and leaping abilities, as reported by Sosnovskii (1967: 121) for his adult Moscow Zoo specimens, are truly astounding. Long jump from standing start: 3 to 3.5 m (10 to 12 ft), with running start: 5 to 6 m (17 to 20 ft.), high jump: 3 to 3.5 m (10 to 12 ft).
Paws: Digits 5/4, i.e., there are five toes (four fully developed, one, the dewclaw, as vestigial "thumb") in front and four in back (as in all canids with the exception of Lycaon); with hair growing between toes ("fins") and pads.
Tail: For canids, moderately long in relation to total length of head and body (41%, more than the wolf and less than most foxes). Of medium bushiness, darker than body coat as the black overlay increases with distance from the body. Tip entirely black.
Coat: Overall extensive red color; the amount of black tipping of the hair on the back can vary widely from one individual to the next affecting the apparent brightness of the coat; outer coat coarse and either short (approximately 11-13 mm on back, 27-30 mm on neck) and without pile (in summer) or longish (21-30 mm on back, 32-50 mm on neck) with fairly dense undercoat (in winter). Ventral pelage white or paler red than rest of body. White fur may be present or absent on lips, ears, throat, chest, belly, paws and inside of limbs; muzzle either faded black mask or self-colored. The coat of subadults is a dull tawny brown. [Back to the beginning]
In the Moscow Zoo, the mating season for the Asiatic Wild Dog is February (Sosnovskii, 1967), i.e., in the middle of the Russian winter. In the Himalayan country of Bhutan, mating occurs mainly in November/December (late autumn, early winter). The corresponding data for India vary widely between September and January (see table in Cohen, 1977: 148), i.e., from the middle of the rainy season through the cool season of the Northeastern monsoon.
Nothing is known (or reported) about how the mating partners choose each other. Does the female select her mate? Is the alpha male the only one to cover the females, or may other males "get lucky," too? Fox’s statement (1984: 79) "the breeding male was not the largest member in the pack", begs the question because size is only one criterion of several that make a dog the leader of a pack. Reports of several females raising their individual liters together in one den indicate that breeding is not restricted to the alpha female alone. There is, however, Fox’s (1984:42) ‘weasel-worded’ statement "It may also be deduced from his [= Johnsingh’s, 1982] observations that dominance among females results in only one pregnancy and litter." Perhaps these conflicting observations would be resolved if a further variable, food supply, could be shown to be a factor: several litters in times of plenty, only one when food is scarce?
The gestation period is 60 to 62 days (Moscow Zoo), "ca. 63 days" in India. Thus the "Russian" young are born in April, i.e., in late winter, the Bhutan cubs in January/February (mid to late winter) and their "Indian cousins" some time between November and March. The literature does not attempt to correlate breeding seasons and climate, but it appears that whelping is timed in such a manner that, when the pups are ready to abandon the den [at age 70 to 80 days, (Johnsingh, 1978)], the weather is at its most propitious for their survival.
Dhole Propagation and Climate in Bandipur Tiger Reserve Karnataka, India (Data abstracted from Johnsingh, 1982):
* Time of maximum humidity (83%).
† Time of minimum humidity (20%) and maximum temperature during the day (32oC).
†† Time of minimum temperature during the day (19oC).
** Part of the pack emigrates early in the month.
Litter size is usually stated as "4 to 6"; Johnsingh, however, gives the "average" litter size as eight (Macdonald, 1995) and "around 8" (1982: 460). Cohen (1978: 2) reports two instances in which postmortem examination of pregnant females showed eight and nine fetuses respectively (cf. also Fox,1984: 81)..
The moderate litter size of the dhole appears at odds with the (for canids) exceptionally large number of mammary glands (up to 16). Because nothing is known about the evolutionary pressure(s) that may have produced this adaptation, all attempts at explaining the feature (unrealized reproductive potential, adaptation to communal breeding and nursing, both Cohen, 1977: 148, survival trait in that one nursing female can foster the pups of another that cannot nurse her own, etc.) can be but speculation.
There seem to be no data about the mortality rate of dhole pups.
The pups develop at a rate comparable to that of domestic dogs; Johnsingh (1982: 453) observed a litter actively playing with their mother outside the den at age eight weeks; one week later, the pups ventured out of the den on their own; one week after that, now ca. ten weeks old, the pups left the den for good; and at approximately seven months they were able to assist an adult male in making a kill.
There are no "normal" or "typical" dhole dens; they range in construction from simple one-entrance, one-cavern burrows to elaborate, complex multi-entrance and multi-cavern structures that may have taken generations of dogs to dig (Johnsingh, 1982; Fox, 1984: 43-49). However, there are no reported observations of dholes actually digging their own den while the use of dens dug by other animals, e.g. porcupines, is seen. The dens have ready access to water and provide a clear view of the surroundings. They are well hidden among the scrubs on the banks of dry streambeds or amidst rock outcrops, and the immediate surroundings are kept remarkably clean of feces and remains of kills. Near the den, a "playground" for the pups may be found, an area of 2 to 3 m in diameter where pups (and adults) play and practice their hunting skills. [Back to the beginning]
Pack and Pack Life
Note: Fox (1984: 82 ff.) proposes that the term ‘pack’ be reserved for hunting parties and that the groups of dogs that live together, be called ‘clans’. This in itself is a useful distinction that is, however, difficult to maintain against the established usage (Fox himself speaks of "the pack he studied" when he means "clan") and has produced some confusion (e.g., Alderton, 1998:160: "Packs may come together and form larger units, described as clans, ..."). For these reasons, it seems prudent to avoid the term ‘clan’ altogether.The pack size usually is stated as 5 to 12 dogs with males outnumbering females at the rate of approximately 2 to 1. After the denning season, when the new litter(s) grow towards maturity, the pack may have 20 or more members. Just before the next denning season (i.e., halfway into the gestation period, i.e., early November, in Bandipur Tiger Reserve), part of the pack may leave or "emigrate," to use Johnsingh’s (1982: 446) term, reducing pack size again to the normal (lower) number.
Note: There seems to be no further information available about these émigrés, only questions: Do they leave as a group or individually? Are they all males? Last year’s "crop"? Do they rejoin later? Or form their own packs? Or join other packs?
Persecuted by man, dholes have become very elusive. Their mainly crepuscular and nocturnal life (i.e., from dusk to dawn) makes it difficult to fully observe and understand their communal activities. They most frequently are seen when hunting in the early morning hours; but it is unclear whether this is indeed the preferred time to hunt because there are many reports of observers hearing kills being made during the night or of finding the remains of such kills the next day. The size of the observed hunting parties varies from one solitary animal to up to 12 hunters, and one may wonder if there might not be a correlation between the size of the "dawn patrol" and the success (or lack thereof) of the nocturnal hunt. We know nothing about these night hunts, but field workers (Keller, Johnsingh, Fox) have observed and described a number of daylight hunts.
In contrast to most canids, dholes lack a killing bite, i.e., the capability of dispatching a prey animal swiftly and efficiently, and the act of killing can be a messy and lengthy affair (2 to 15 minutes). The dholes attack the prey’s head region (especially face, lips, muzzle) and abdomen; and the animal is more likely to die from pain-induced shock than any one fatal wound. It is this mode of killing (head attack and evisceration) that has "earned" the dhole the reputation of a cruel and bloody killer. A frequently observed part of the hunt is driving the prey animal into (deep) water where exhaustion overcomes the will to fight more quickly and where the water prevents the animal from using its sharp hoofs in its defense.
In India, especially in Bandipur National Park, the only region for which such information is available, dholes hunt (in order of frequency as described by Johnsingh, 1980, and Fox 1984: 60): chital (Axis axis), Indian hare (Lepus nigricollis), sambar (Cervus unicolor), microtine rodents, cattle, barking deer, langur, wild pig, and gaur,with the first three being the primary prey animals: Remains of one or the other of them were found in nearly 80% of the 509 dhole scats examined by Johnsingh.
In the Moscow zoo, the dholes are fed once a day. Adults eat about 1.2 to 1.5 kg (~ 2.6 to 3.3 pounds) of raw meat (Sosnovskii, 1967: 121). Fox estimates that an empty dhole stomach can hold "about 6.5 pounds of prey" (1984: 59). (The not inconsiderable difference in the stated amounts of food may, at least partially, be rationalized by the fact that the wild dholes, in contrast to their captive brethren, must expend much more energy in hunting their dinner and, from time to time, may have to go without any). -- A hare may feed one or two dholes, a chital perhaps ten to twelve, and a sambar may provide meat for as many as 40. It has been speculated that the decline of the sambar population has led to today’s smaller dhole packs of about twelve members or less, down from the forty in older reports. There is very little dhole predation on domestic cattle, especially so when the beasts are brought into the village overnight.
As the term ‘denning season’ (Johnsingh) indicates, dholes do not live in caves year-round; rather the den serves as a safe haven for whelping and the first ten to twelve weeks of a cub’s life. The scientific literature is silent about where the dholes spend their days (perhaps it is too obvious to warrant comment). Rudyard Kipling, however, tells us that Mowgli "had seen the fearless dholes sleeping and playing and scratching themselves in the little hollows and tussocks that they use for lairs."
Dholes have command of a wide range of vocalizations. There is the whining (whimpering) and the growling and the barking and the barking and growling in combination and the screaming that all dog owners are familiar with and that convey pretty much the same message that our domesticated companions convey: soliciting food, warning, aggressive warning, expressing fear or pain, etc. But the dholes also have one vocalization all of their own: whistling. This sound, described as "a repetitive, whistlelike, high pitched howl" (Fox, 1984: 94) of "medium range" (Durbin, 1998), serves to reassemble the dispersed pack, e.g., after an unsuccessful hunt, and also to coordinate hunting attacks in dense cover where the hunting pack members cannot see each other. Acoustic analysis of the whistling sounds produced by a captive dhole population (which allowed visual identification of the whistler) suggests that the "whistles can be individually distinctive" (Durbin, 1998). Whereas human hearing lacks the acuity to differentiate between individual whistles, it is quite likely that the much more powerful dhole ear allows the animals to know each other by their whistling.
Arun Venkataraman observed a young female help a mother dhole move her pups from one den to another.
R. Keller saw a pack of dholes work together to pull a chital buck from the water of a river (where they had chased and then killed the exhausted animal) onto dry land.
It is also Keller who reports that, while feeding, ever so often a (different) member of the pack would ‘leave the table’ and scan, with head held high and ears erect, the surrounding area for possible threats, standing on his/her hind legs or even pronging to see over the high grass.
Rudyard Kipling ("Red Dog" , in The Second Jungle Book, 1897) tells a story of how Mowgli and his Seeonee Pack of 40 wolves fought and annihilated a pack of 200 dholes that had come north from the Dekkan where game had grown scarce. The story is fiction and anthropomorphizes to the extreme but most likely reflects accurately the then attitude of the common people and of "naturalists" alike towards the dhole: fear and hatred. [Back to the beginning]
(Note: The starred titles could not be found.)
Cohen, J[ames] A. "A Review of the Biology of the Dhole or Asiatic Wild Dog (Cuon alpinus, Pallas)." In: Animal Regulation Studies 1, 1977, 141-158.
Cohen, James A. "Cuon alpinus." In: Mammalian Species 100, 1978, 1-3.
Fox, Michael W. The Whistling Hunters. Field Studies of the Asiatic Wild Dog. (SUNY Series in Animal Behavior) State University of New York Press: Albany, N.Y. 1984.
*Hodgson, B[rian] H[oughton]. "Description of the wild dog of the Himalaya (Canis primaevus)". In: Asiatick Researches 18(2), 1833, 221-237.
*Hodgson, B[rian] H[oughton. [title unknown] In: The Annals and Magazine of Natural History. Series 1, vol. 1, 1838, 151-153.
* Johnsingh, A.J.T. Ecology and Behaviour of the wild dog (Cuon alpinus). PhD. dissertation, Madurai Kamarajar University, Maduraicity, India, 1980.
Johnsingh, A.J.T. "Reproductive and social behaviour of the Dhole, Cuon alpinus (Canidae)." In: Journal of Zoology (London), 198, 1982, 443-463.
Johnsingh, A,J.T. "Dhole." In: Macdonald, David (Ed.). Encyclopedia of Mammals. 1984, pp. 80-81.
Johnsingh, A.J.T. "Distribution and Status of Dhole Cuon alpinus, Pallas 1811, in South Asia." In: Mammalia 49, 1985, 203-208.
Keller, R. "Einige Beobachtungen zum Verhalten des Dekkan-Rothundes (Cuon alpinus dukhunensis Sykes) im Kanha-Nationalpark." In: Vierteljahrsschrift der Naturforschenden Gesellschaft in Zürich 118, 1973, 129-135.
Kipling, Rudyard. The Second Jungle Book. 1897. Chapter "Red Dog."
*Pallas, Peter Simon. Zoographia Russo-Asiatica. Vol. 1 (1811) p. 33-35.
Pocock, R. I. "The Asiatic Wild Dog or Dhole (Cuon javanicus)." In: Proceedings of the Zoological Society of London 1936, 33-55.
Sosnovskii, Igor P. "Breeding the Red dog or dhole Cuon alpinus at Moscow Zoo". In: International Zoo Yearbook 7, 1967, 120-122.
Thenius, E[rich]. "Zur Abstammung der Rotwölfe (Gattung Cuon HODGSON)". In: Österr. zoologische Zeitschrift 5, 1954, 377-387.
Dhole: www.blarg.net/~critter/articles/canines/dhole (= Johnsingh, A.J.T. "Dhole." In: David Macdonald, Ed. The Encyclopedia of Mammals. 1984: 80-81.)
Dhole Conservation Project: Dhole Home Page: www.dcpbase.demon.co.uk/dcp
Dholes in Danger: www.dcpbase,demon.so.uk/dcp/fieldrep.htm
Durbin, Leon S.. "Individuality in the Whistle Call of the Asiatic Wild Dog." www.dcpbase.demon.co.uk/dcp/whispapsum.htm
Johnsingh, A., N. Ovsyanikov, G. Schaller, P.Stewart. "Dhole (Cuon alpinus)."www.canids.org/SPPACCTS/dhole.htm (Last update: 27 June 2000)
Kappeler, Markus. "Rothund Cuon alpinus. 1997. www.markuskappeler.ch/tex/texs/rothund2.html
Lioncrusher’s Dhole Information Page: www.geocities.com/RainForest/Vines/8591/dhole
The Living Edens:Bhutan Animal Archive-Dhole,Jackal,Wolf: web- cr02.pbs.org/edens/bhutan/a_wolf.htm
Venkataraman, Arun. "The Social Life of the Dhole.": www.thewildones.org/Animals/dhole.html
World Conservation Monitoring Centre – The Republic of Viet Nam: www.wcmc.org.uk/infoserv/countryp/vietnam/app4.html
The Living Edens. ANAMALAI: India’s Elephant Mountain. PBS Video [no date, 2000?]. Contains several brief sequences featuring dholes.
[Back to the beginning]
Distribution Habitat/Territory Names Subspecies Description Propagation Pack Life Bibliography
Comments, additions and/or corrections are welcome at Kingswayhb@aol.com