Dhole (Cuon alpinus)

The adult dhole is characterized by a rusty red coat with a pale underside. Their bellies, chest, and feet are usually white, but not always. Dholes also have dark, almost always black, bushy fox-like tails. Depending on the region, a dhole’s pelage may vary from light brownish-charcoal gray, to sandy beige, or an iconic uniform red coat.

Dholes are so uniformly marked that there is usually no way to tell individuals apart. It can even be difficult to distinguish between sexes. Oftentimes, researchers and zookeepers must rely on behavioral differences more than physical differences to distinguish individual dholes from one another.

Male dholes tend to be larger and heavier than females. Females average 22-29 pounds, or 10-13 kilograms, while males normally weigh 33-44 pounds, or 15-20 kilograms.

The dhole is an average-sized canine, normally about the size of a German shepherd dog (Canis lupus familiaris). They are often seen as surprisingly small, scaling at 30-50% the size of an average wolf. They can grow up to 3 feet, almost 1 meter, in length and stand at about 20 inches, or 50 centimeters, at the shoulder. Their tails are often 15-18 inches, or 40-45 centimeters, long.

The dhole is one of the most extreme canids given its unique morphological and behavioral traits, such as having specialized teeth for hypercarnivory. It is one of only three canid species with specialized dental adaptations for the exclusively hypercarnivorous diet. It is set apart from other canids in that it has an unusually thick muzzle, short jaw, and one less molar tooth on each side of its lower jaw. Dholes have only two molars on each side of their lower jaw, instead of three. Other members of the family Canidae have a total of 42 teeth.

SEXUAL DIMORPHISM

Larger, Heavier Males

SHOULDER HEIGHT

50 cm. / 20 in.

BODY LENGTH

88-92 cm. / 35-36 in.

TAIL LENGTH

40-45 cm. / 15-18 in.

BODY MASS

9-12 kg. / 22-45 lb.

DENTAL FORMULA

I ³⁄3, C ¹⁄1, P ⁴⁄₄, M ³⁄2, ×2 = 40

LIFESPAN

10-16 yr.

GENERATION LENGTH

5 yr.

LOCOMOTION

Digitigrade

The canid’s scientific name is Cuon alpinus, and they are the only species in the Cuon genus. In other words, if we lose the dhole we lose an entire evolutionary genus.

Of the canids, African wild dogs (Lycaon pictus) are arguably most similar to dholes. Both species have specialized dental morphology for obligate hypercarnivory, which is rather unique among canids. Phylogenetic research shows both species are in the same monophyletic group, and otherwise both species are similar in body size, reproduction, social behavior, and feeding behavior. Dholes and African wild dog also live social lives, in that they engage in obligate cooperative group hunting and group care of young. Subordinate pack members are reproductively suppressed and instead help care for the young of the dominant alpha pair.

The dhole was originally distinguished as two separate species of Cuon, the northern dhole, (Cuon alpinus,) and the southern dhole, (Cuon javanicus,) based on differences in body size and the second upper and second lower molars.

Dholes were later recognized as a single species, but separated into as many as 11 subspecies based on differences in size, coat length, and color. As of 2005, the subspecies were simplified to 3.

The validity of many of these subspecies is doubtful as genetics studies have found no clear subspecies distinctions. Examination of the auditory bulla showed at least one northern subspecies, from the Tian Shan Mountains, to be morphologically distinct from other subspecies. The genetic distinctiveness between northern and southern dholes remains unknown. Further research is needed to clarify the morphological and genetic differences between them, especially because conservation efforts would need to employ different strategies for the two groups, as they occupy vastly different habitats and prey on different species.

The putative northern-dhole group historically occurred throughout East Asia to as far south as the Himalayan Mountains and the Yangtze River, and would include the following five subspecies: C. a. alpinus, C. a. primaevus, C. a. laniger, C. a. hesperius, and C. a. fumosus. The putative southern-dhole group would include the remaining six subspecies: C. a. lepturus, C. a. dukhumensis, C. a. adjustus, C. a. infuscus, C. a. sumatrensis, and C. a. javanicus.

Three races of the dhole exist in India alone: Trans-Himalayan, Himalayan, and Peninsular.

KINGDOM

Animalia

PHYLUM

Chordata

CLASS

Mammalia

ORDER

Carnivora

FAMILY

Canidae

GENUS

Cuon

SPECIES

alpinus

SUBSPECIES (3)

C. a. alpinus (Chinese/Eastern Asiatic/Indian/Southern/Ussuri), C. a. hesperius (Siberian/Northern/Tian Shan/Western Asiatic), C. a. sumatrensis (Javan/Malayan/Sumatran)

Although little-known, dholes have a plethora of names: the Asiatic wild dog, Indian wild dog, mountain wolf, red wolf, red dog, and whistling dog among them, along with some 40 or so indigenous names. However, the origins of the name dhole remain obscure.

Due to their similarity to African wild dogs (Lycaon pictus), dholes are often called Asiatic wild dogs or Indian wild dogs.

Dholes are known as whistling dogs, among other names, because of their vocal nature and bird-like whistling calls.

ALTERNATE

Asiatic Wild Dog, Indian Wild Dog, Mountain Wolf, Red Dog, Red Wolf, Whistling Dog

GROUP

Pack

YOUNG

Pup

From the Altai Mountains in Manchuria in Central and Eastern Asia, the dhole’s range spreads southwards through the forest tracts of India, Burma, and the Malayan Archipelago. The dog is found throughout most of northeastern region of India, particularly in the states of Assam, Meghalaya, Arunachal Pradesh, and West Bengal. They are also found in various regions in the country south of the Ganges River, especially in the Central Indian highlands, and the Eastern and Western Ghats of the southern states of the country.

However, experts believe that the species’ historical range could have included all or most of the Malaysian peninsular as well as the Indonesian islands of Java and Sumatra.

Dholes have disappeared from most of their historical range. Recent population analysis by the International Union for Conservation of Nature (IUCN)’s Dhole Working Group has revealed that the area of occupancy of this species has significantly declined close to 50% since the last assessment in 2008. Populations are still declining in most areas due to several main threats which include depletion of prey base, habitat loss, persecution due to livestock predation, disease transmission from domestic dogs, and possibly interspecific competition.

EXTANT

Bangladesh, Bhutan, Cambodia, China, India, Indonesia, Lao, Malaysia, Myanmar, Nepal, Thailand

POSSIBLY EXTINCT

Vietnam

EXTINCT

Afghanistan, Kazakhstan, Korea, Kyrgyzstan, Mongolia, Russia, Singapore, Tajikistan, Uzbekistan

UNCERTAIN

Korea, Pakistan

The dhole is a habitat generalist and can occur in a wide variety of vegetation types, including: primary, secondary, and degraded forms of tropical dry and moist deciduous forests; evergreen and semi-evergreen forests; temperate deciduous forests; boreal forests; dry thorn forests; grassland–scrub–forest mosaics; temperate steppe; and alpine steppe. Consequently, their elevation range is from sea level to as high as 5,300 m asl in Ladakh.

Dholes like open spaces and can often be found on jungle roads, river beds, jungle clearings, and paths, where they rest during the day. Their dens are usually constructed near streambeds or among rocks. They have never been recorded in desert regions, though.

The dhole has the largest land requirements of any Asian species. The dhole’s hunting range is about 40 square kilometers, or 15 square miles, but can range from 23-199 square kilometers in India and 60-80 kilometers in Thailand.

Due to the demands imposed by hypercarnivory, sufficient numbers of ungulate prey are the dhole’s major habitat requirements. In India, tropical dry and moist deciduous forest may represent optimal habitats, based on the areas thought to hold the largest dhole populations. Ungulate biomass, particularly that of cervid species, is highest in these habitat types when compared to other habitats in the same region.

FOREST

Boreal, Temperate, Subtropical/Tropical Dry, Subtropical/Tropical Moist Lowland, Subtropical/Tropical Moist Montane

SHRUBLAND

Subtropical/Tropical Moist

GRASSLAND

Temperate, Subtropical/Tropical Dry, Subtropical/Tropical High Altitude

Dholes are highly social, pack-driven animals that like to keep an eye on each other. They rarely separate from one another as they live in close-knit packs averaging 5-12. They interact with other dholes outside of their own group, but the original pack rarely exceeds 20. Pack sizes differ depending on the ecosystem and prey availability. In the past, researchers have reported packs as big as 40 and there are rumors of packs even reaching 100.

Pack sizes differ depending on the ecosystem and prey availability. In India, dholes form relatively large packs to efficiently hunt large numbers of prey, as well as to protect litters, which are usually large. However, in tropical evergreen forests of Southeast Asia, dholes appear to persist in smaller packs and presumably have smaller litters, probably due the low prey biomass and small size of ungulate prey in these habitats.

Dholes have clear hierarchy in their packs with each group containing a dominant monogamous breeding alpha pair. While alpha dhole pairs are dominant, they are not excessively aggressive towards the beta pack members. Subordinate pack members are reproductively suppressed and instead help care for the young of the dominant pair.

Dholes feed in order of dominance to avoid conflicts. Unlike wolves, they allow pups to eat before any of the other pack members. Dholes compete for food, not by fighting, but by how fast they can eat. An adult dhole can eat up to 4 kilograms, or 8.8 pounds, of meat in one hour.

Dholes are docile animals. Gentle and communal, they are described as more egalitarian than wolves. Aside from cubs play fighting, there is rarely any evidence of aggressiveness among pack members, and there is almost never any bullying. They are known to have high levels of amicable behavior within packs, even among adults, which includes playing and face licking. Dholes seem to be playful and not have such antagonistic encounters between pack members as you might see in wolves.

Dholes do not attack human beings and usually retreat at the sight of a person. They are super-sensitive animals, skittish to the point of potentially overheating under stress.

Dholes communicate with each other using body language and vocal sounds.

Dholes can be quite vocal, calling to keep in touch over short or long distances, which is handy when they are cooperatively hunting. They also communicate in the form of whistles, barks, growls, alarm calls, and chatter.

The dhole’s greeting includes snapping at each other—an expression of endearment for them, but not so endearing for humans. Just as domestic dogs, they also wag their tails.

Dholes are intelligent canines and can be trained to follow commands. The zookeepers at the San Diego Zoo recognize the etiquette of dhole pack dynamics while working with their dhole individuals for husbandry purposes. To maximize animal welfare and healthcare and minimize stress to the animals, each dhole—Sanuk, Lucius, Beni, Jetsan, Katsu, Torma, Kono, and Yoshi—are schooled in practical behaviors like station (staying in a particular spot), down, sit, target, rise up, present paw, and open mouth. This training is accomplished through positive reinforcement with rewards, which deepens the animals’ relationship with their keepers and keeps the sessions upbeat and constructive.

Dholes are great diggers. The vast outdoor area of the dhole habitat at the San Diego Zoo Safari Park needed a few adjustments, in order to ensure the safety of the animals. Because dholes are great diggers, the underground, three-foot barrier along the fenceline was closely inspected and reinforced before the animals were introduced to the habitat.

Dholes are very fond of water and pools tend to be favorite hang-out spots. After meals, dholes will race to a water site. If the water is near their kill, they will leave their food for a small drink of water. Dholes have also been spotted sitting in shallow pools of water regardless of the temperature and often build their dens near streambeds.

CIRCADIAN RHYTHM

Diurnal

The dhole is a hypercarnivore, meaning its diet is made up of at least 70% meat. They require relatively high prey numbers in order to raise their sizable litters and sustain their large pack sizes. They will hunt both, small and large prey, and even scavenge.

The majority of the dhole’s prey base consists of deer, such as sambar (Rusa unicolor), and other hoofed animals, such as gaur (Bos gaurus) with a body mass of 40-60 kilograms. Packs of dholes feast on mammals ranging from rodents to deer, but individual dholes will also eat wild berries, insects, and lizards. Some of the dhole’s favorites include wild boar (Sus scrofa), hares, wild goats, sheep (Ovis aries), and occasionally a monkey. Although this occurs on rare occasions, dholes will also prey on domestic livestock.

Dhole have also been witnessed stealing kills from tigers (Panthera tigris) and even killing the big cat.

Seasonal changes occur in the dhole’s diet, reflecting seasonal changes in availability and numbers of prey.

Dholes are quite effective as predators when hunting in packs and have been referred to as natural born killers and dogs that kill better than cats. One of the keys to the dholes’ hunting ability is its pack size and flexibility. Dhole hunting parties are known to employ a variety of tactics to bring down their prey, including splitting into small scouting groups. While a few dholes chase the animal, others break away in a flanking movement to run ahead of their quarry and ambush it as it dashes past them.

Dholes have the ability to tackle large and dangerous prey like sambar, wild boar, and gaurs – wild bovines that can weigh 50 times more than a dhole. Sambar are extraordinarily hardy, yet are literally eaten alive by these dogs before being killed. Two to three dholes can kill a 50 kilogram, (110 pound) deer in less than two minutes.

Unlike many other canines, the dhole seldom kills by biting the throat. When successful, dholes don’t suffocate prey with a bite to the throat like a tiger (Panthera tigris) or leopard (Panthera pardus), but bring their prey down and begin eating. Smaller prey is caught by any part of the body and killed by a swift blow to the head. Larger mammals are attacked from the rear by the flankers, as they bite out the animal’s eyes, disembowel it, and hamstring and emasculate it in their efforts to bring it down. Once brought to the ground, the prey is immediately disemboweled and the pack begins to feed on it before it’s dead. Larger prey rarely dies from the attack, itself, but from blood loss and shock as its intestines, heart, liver, and eyes are feasted upon. Sometimes, prey can trail its intestines behind itself up to 20 feet, trying to protect itself from the canines.

Dholes feed in order of dominance to avoid conflicts. Unlike wolves, they allow pups to eat before any of the other pack members. Dholes compete for food, not by fighting, but by how fast they can eat. An adult dhole can eat up to 4 kilograms, or 8.8 pounds, of meat in one hour.

PREFERENCE

Generalist

STYLE

Pursuit

Dholes have clear hierarchy in their packs with each group containing a dominant monogamous breeding alpha pair. Subordinate pack members are reproductively suppressed and instead help care for the young of the dominant pair.

The dhole’s gestation period is 60-62 days and the pups are born throughout the end of fall, winter, and the first spring months, usually between November and March.

The mother usually gives birth to 3-10 pups at a time, but can have up to 16 young. This large number of offspring, along with the fact that female dholes have more teats than other canid species suggests their ability to take care of large litters. A female dhole can have up to 16 mammae, allowing the dhole to have larger litters and pack sizes.

After the dominant female dhole has given birth, a few other adults take part in feeding the mother, as well as the pups. The pups, as early as the tender age of three weeks, and the mother are fed regurgitated meat.

A dhole is born with a sooty brown color, acquiring an adult color at three months of age. The pups reach sexual maturity at about a year.

BREEDING SEASON

September-February

PARENTAL INVESTMENT

Cooperative Breeding, Maternal, Paternal

GESTATION

60-62 Days

BIRTHING SEASON

November-March

LITTER

3-16

MAMMAE

16

WEANING

2 Months

SEXUAL MATURITY

1 Year

Dholes are remarkable in that they thrive – and tackle big prey – in regions inhabited by much larger predators, such as tigers (Panthera tigirs), snow leopards (Panthera uncia), bears, grey wolves (Canis lupus), and leopards (Panthera pardus). Competition for prey and space can bring dholes into encounters with these other apex carnivores.

Dholes have been witnessed stealing kills from tigers and hounding, and harassing the big cat. There are even unconfirmed observations of dholes killing the panther. Because, no verified stories of dholes actually killing a tiger exist, though, such tales may be more legend than reality. Dholes have been documented to have been killed by tigers and attacked by leopards, indicating both larger carnivores may be behaviorally dominant over dholes.

During the Pleistocene, dholes were found across Europe and North America, surviving in a land of saber-toothed cats and giant bear dogs.

Most of the dholes that live outside or in the fringes of core protected areas are highly vulnerable to conflict with humans.

During a radio-tracking study in the buffer zone of the Kanha Tiger Reserve in central India in 2000, the surveyors discovered that at least 16 out of the 24 dholes in a particular pack died all of a sudden from lethal poisoning, indicating conflict with humans.

Dholes have become an indirect food source for the residents of the jungles in Asia. Dholes do not attack human beings and usually retreat at the sight of a person. They are super-sensitive skittish, docile, animals. Taking advantage of this fact, human residents of the jungle follow dholes when they are hunting in order to steal their kills. After a dhole has taken down its prey, humans will scare it away and steal the carcass.

Historically, dholes have had a poor reputation, falling into the stereotypical category of the big bad wolf and being referred to as pests of the jungle. Although this occurs on rare occasions, dholes can attack livestock at the cost of the owner. Due to this, they have suffered heavily from local perceptions as problem predators and livestock killers. If a dhole is found to have killed livestock, herders poison the carcasses, which can unintentionally wipe out an entire pack and kill other carnivores and vital scavengers, like vultures. Retaliatory killing is a primary factor limiting dhole conservation.

Estimates range from 949 to 2,215 breeding dholes left in the wild, less than the world’s breeding tigers (Panthera tigris), with a total of 4,500-10,500 individuals in total.

Little is known about dhole ecology as they’re extremely difficult to study.

Outside of India, dholes remain primarily in pockets of remote areas in Southeast Asia and China. In these areas, they tend to occur in low densities in thick forests that cover mountainous or rugged terrain.

There is almost no quantitative information on dhole population trends through their distribution. Population estimates of dholes are not available for any country and the size of subpopulations of dholes has not been reported anywhere. The only estimates of local dhole densities come from a few protected areas in India.

In order to better understand territorial requirements for packs, researchers rely on GPS collaring of wild dholes. Understanding the dholes’ space helps in estimating the number of packs that live in protected areas. The collars are linked to satellites, giving scientists an exact location of each collared dhole. Collaring any wild carnivore is problematic but dholes may be among the trickiest.

MATURE INDIVIDUALS

949-2,215

FRAGMENTATION

Severely Fragmented

Threats include loss of habitat, inter-species competition, depletion of prey base consisting mostly of deer and other hoofed animals, indiscriminate persecution, and also possible disease transfer from feral and domestic dogs (Canis lupus familiaris).

Dholes inhabit some of the most threatened, degraded and disconnected forest landscapes on the planet. The rainforests of Southeast Asia have seen unprecedented destruction over the past fifty years for palm oil, paper, rubber, timber, mining and other commodities. Where forests haven’t been destroyed, they have been fragmented by booming human populations, roads, and ever-expanding development projects.

Overhunting and snaring has decimated many prey species across southern Asia. In fact, the region is known for so-called “empty forests syndome”. Here, forests are largely emptied of any large-to-medium-sized mammal or bird, wiped out by hunting both for food and the Traditional Chinese Medicine industry.

Even where forests still stand and prey remains abundant, dholes face human persecution.

RESIDENTIAL & COMMERCIAL DEVELOPMENT

Housing & Urban Areas

AGRICULTURE & AQUACULTURE

Annual & Perennial Non-Timber Crops

BIOLOGICAL RESOUCE USE

Hunting & Trapping Terrestrial Animals

INVASIVE & OTHER PROBLEMATIC SPECIES, GENES, & DISEASES

Invasive Non-Native/Alien Species/Diseases, Problematic Native Species/Diseases, Viral/Prion-Induced Diseases

In order to help re-populate the endangered dhole population, the San Diego Zoo Wildlife Alliance has been breeding dholes since 2001 in an off-exhibit breeding site. 20 pups have been born there to four different mothers. The newest litter of dholes was born in January 2015, bringing the current off-exhibit pack population to twelve. Because of the breeding success, the animal care staff was able to form a non-breeding group that has taken up residence in an Asian Savanna exhibit. Besides the San Diego Safari Park, only two other facilities in the U.S. have dholes: The Wilds in Ohio and the Minnesota Zoo.