Cuon alpinus (Pallas, 1811)

Don E. Wilson & Russell A. Mittermeier, 2009, Canidae, Handbook of the Mammals of the World – Volume 1 Carnivores, Barcelona: Lynx Edicions, pp. 352-446 : 423

8.

Dhole

Cuon alpinus

French: Dhole / German: Asiatischer Wildhund / Spanish: Cuén

Other common names: Asiatic Wild Dog

Taxonomy. Canis alpinus Pallas, 1811 .

Amur region, former USSR.

The genus Cuon is post-Pleistocene in origin, and related more closely to the extant jackals than to wolves. Although initially placed in the subfamily Simocyoninae , together with Lycaon pictus and Speothos venaticus , analysis of morphological, ecological and behavioral characteristics across 39 canid species showed that Cuon was actually more similar to Canis , Dusicyon , and even Alopex , the resemblance with Speothos or Lycaon being based solely on skull and dental characteristics. Analysis of sequences from mitochondrial DNA provided further evidence for the taxonomic distinctiveness among Speothos , Cuon , and Lycaon . Eleven subspecies have been recognized, although many of those forms are doubtful. Three subspecies recongnized here.

Subspecies and Distribution.

C. a. alpinus Pallas, 1811 — C Russia and W China southward through India to Bhutan and Bangladesh.

C. a. hesperius Afanas’ev & Zolotarev, 1935 — E Russia, China, and SE Asia.

C. a. sumatrensis Hardwicke, 1821 — Sumatra and Java.

Descriptive notes. Head-body 88-1355 cm and tail 32-50 cm; weight 15-20 kg for males and 10-13 kg for females. Large canids, usually with a reddish or brown coat and a darker, bushy tail (sometimes with a white tip). Sexual dimorphism is not very distinct. Ears triangular, about half the length of the face, with rounded tips. The pinnae are usually whitish-fawn on the inside and reddish-brown on the outside. The muzzle is brown, relatively short, and slightly convex in profile. Nose black and eyes slightly hooded;irises are amber. The dorsal and lateral pelage is red to brown and the foreneck, chest, and undersides are often whitish or pale ginger colored. In the south and south-west of the Dhole’s range, their fur is shorter and rusty-red colored. In the north and north-east, the fur is longer, brownish-red, or yellowish-brown. The legs are notably shorter in some alpine regions and the coat is a yellowish-gray color in Himalayan regions. In Thailand, the coat is more uniformly brown, lacking the paler throat and chest. The toes are red, brown, and/or white; the hairless foretoe pads (on all feet) are joined at the base (near the main pad) unlike most domestic dogs. Dentition is unique within the Canidae , having one fewer lower molar teeth (13/3, C 1/1, PM 4/4, M 2/2 = 40), with the heel of the lower carnassial M, crested and with a single cusp (all other canids within the range of Dhole have two cusps). There are usually six or seven pairs of mammae.

Habitat. Dholes are found in a wide variety of vegetation types, including primary, secondary, and degraded forms of tropical dry and moist deciduous forest; evergreen and semi-evergreen forests; dry thorn forests; grassland—scrub—forest mosaics; and alpine steppe (above 3000 m). They are not recorded from desert regions. In India, tropical dry and moist deciduous forest may represent optimal habitats, based on the regions thought to hold the largest Dhole populations. Ungulate biomass, particularly that of cervid species, is highest in these vegetation types when compared to others in the same region. Important factors that may influence habitat selection include the availability of medium to large ungulate prey species, water, the presence of other large carnivore species, human population levels, and suitability of breeding sites (i.e. in terms of proximity to water, presence of suitable boulder structures, and sufficient prey).

Food and Feeding. Diet includes beetles, rodents, birds, and occasionally grass and other plants like many other canids. However, Dholes hunt mainly vertebrate prey, with a preference for medium to large ungulates. They are also known to scavenge and occasionally have been observed eating carrion (e.g. Asian Elephant (Elephas maximus) and Gaur (Bos frontalis) carcasses). It has been suggested that such scavenging only occurs during periods of prey scarcity, particularly during the dry season. Grass is ingested, but may serve an anti-helminthic function rather than a nutritional one. Dholes are communal hunters, occasionally forming packs of over 30 animals, but are more often found in hunting groups of fewer than ten animals. Depending on prey availability, Dholes may also hunt alone or in pairs, taking smaller prey such as deer fawns or hares. During hunts, some Dholes may lie in ambush while others drive prey towards them. Dholes often drive deer into water, where they swim out to surround and capture them. It is common for certain individuals to take particular roles in the hunt, such as leading the chase or taking the first grab at the prey.

Activity patterns. Primarily a crepuscular forager, but can hunt at any time of the day or night.

Movements, Home range and Social organization. Dholes usually live in packs of 5-10 individuals, but groups of as many as 25 (including juveniles) have been recorded on a regular basis. Their ability to regurgitate small quantities of meat at will and thus transport food to pack-mates and neonates facilitates communal breeding, providing food for the pups, the mother, and other adult helpers that remain at the natal den. Group size and composition may vary under different environmental conditions, but most of the current data are from India. Packs have been reported to contain significantly more males than females, perhaps a reflection of female-biased dispersal. Pack members regularly play together, engaging in mock-fights, rolling, and allogrooming. Social rank is established by pushing and holding, but rarely by aggressive biting. Groups have a strong hierarchical structure, with a dominant male and female who are the main breeders. Pack members over-mark each other’s feces and urine on latrines throughout the group’s range. The ranges of neighboring packs often exhibit little overlap, though interactions between groups occur and can be either friendly or hostile. In India, reported home range size ranged from 40 to 83 km ®. In Thailand, two radio-collared adult male Dholes occupied home range sizes of 12 km * and 49-5 km?. Dholes have a broad and unusual vocal repertoire that includes whines, mews, squeaks, growls, growl-barks, chattering calls, and screams. This large range of alarm calls may have evolved to alert pack members to danger from predators (e.g. Leopard, Tiger, Humans), and could serve to intimidate adversaries. A repetitive whistle-like contact call may allow dispersed pack members to identify one another and to regroup. Maintaining group cohesion in this way is likely to be highly adaptive in areas with other large predators.

Breeding. Dholes give birth once a year and have a gestation period of about nine weeks. Mating periods vary in different locations, occurring between November and April (dry season) in India, and January to May (end of the wet season) in East Java. Females are seasonally polyestrous with a cycle of 4-6 weeks. The dominant pair engages in vigorous play and marking, culminating in a copulatory tie. Usually only the dominant female breeds, but exceptions have been noted. Subordinate males sometimes show sexual interest in the alpha female and may contribute to the paternity of the litter. Litter sizes vary dramatically, even within the same pack in different years. The largest litter size recorded is twelve, with only one lactating female in the group. In captivity, newborn pups can weigh 200-350 g, although by the age of ten days their body weight can double, and they have a total body length of about 340 mm. Pups suckle from the mother until they are about three weeks old, when they start to receive regurgitated meat from other pack members. Weaning occurs by about 6-7 weeks, although in captivity this can happen as late as 8-9 weeks. All adults take part in guarding, feeding, grooming, and playing with the pups. By about three months, the pups accompany the adults on hunts; however, the pack may not be fully mobile until about eight months. Dholes reach adult size by about 15 months. In captivity, Dholes of both sexes can reproduce at two years of age. Den types range from earthen burrows to rocky caverns.

Status and Conservation. CITES Appendix II. Listed as Endangered on The IUCN Red List. Latest estimates state that fewer than 2500 mature individuals remain in the wild. Current wildlife decrees in Cambodia give the Dhole protection from all hunting. A new forestry law is under preparation, and a proposalto list the species for full protection is under discussion. India’s 1972 Wildlife Act affords legal protection, specifying that permission is required to kill any individual unless in self-defense or if an individual is a man-killer. The creation of Project Tiger Reserves in India has provided some protection for Dhole populations. In the former Soviet Union, Dholes received the status of “protected animal” in 1974; however, the poisoning and shooting of Gray Wolves may inadvertently affect any remnant Dhole populations. The Dhole is legally protected in Vietnam. Knowledge of Dhole abundance is limited to estimates of the number of packs within a few protected areas in southern and central India, where Dholes are generally thought to be abundant. Abundance is relatively lower in West Bengal, Assam, and Arunachal Pradesh. In the rest of north-eastern India, Dholes are currently extinct or close to extinction. No remotely comparable information on density is available for any part of South-east Asia, and there are no empirical data on trends in this region. Threats include depletion of prey base, habitat loss and transformation, persecution by cattle grazers, and disease.

Bibliography. Clutton-Brock et al. (1976), Cohen (1977, 1978), Davidar (1973, 1975), Durbin (1998), Durbin et al. (2004), Fox (1984), IUCN (2008), Johnsingh (1979, 1982, 1983), Paulraj et al. (1992), Thenius (1954), Venkataraman (1998), Venkataraman et al. (1995), Wayne et al. (1997).

Figures

8. Dhole (Cuon alpinus), 9. African Wild Dog (Lycaon pictus), 10. Maned Wolf (Chrysocyon brachyurus), 11. Bush Dog (Speothos venaticus), 12. Crab-eating Fox (Cerdocyon thous), 13. Short-eared Dog (Atelocynus macrotis)

Subspecies and Distribution. C. a. alpinus Pallas, 1811 — C Russia and W China southward through India to Bhutan and Bangladesh . C. a. hesperius Afanas’ev & Zolotarev, 1935 — E Russia , China , and SE Asia. C. a. sumatrensis Hardwicke, 1821 — Sumatra and Java.