Dholes are more social than wolves,[18] and have less of a dominance heirarchy, as seasonal scarcity of food is not a serious concern for them as it is with wolves. In this sense, they closely resemble African wild dogs in social structure.[3] Dominant dholes are hard to identify, as they do not engage in dominance displays as wolves do, though other clan members will show submissive behaviour toward them.[6] They live in clans rather than packs, as the latter term refers to a group of animals that always hunt together. In contrast, dhole clans frequently break into small packs of 3-5 animals, particularly during the spring season, as this is the optimal number for catching fawns.[39] Dholes are far less territorial than wolves, with pups from one clan often joining another without trouble once they mature sexually.[40] Clans typically number 5-12 individuals in India, though clans of 40 have been reported. In Thailand, clans rarely exceed three individuals.[15] Unlike other canids, there is no evidence of dholes using urine to mark their territories or travel routes. They may defecate in conspicuous places, though a territorial function is unlikely, as faeces are mostly deposited within the clan's territory rather than the periphery. Faeces are often deposited in what appear to be communual latrines. They do not scrape the earth with their feet as other canids do to mark their territories.[41]
Reproduction and development[]
Among Indian dholes, the mating season occurs between mid-October and January, while captive dholes in the Moscow Zoo breed mostly in February.[15] Unlike wolf packs, dhole clans may contain more than one breeding female.[6] During mating, the female assumes a crouched, cat-like position. There is no "tug of war" characteristic of other canids when the male dismounts. Instead, the pair lie on their sides facing each other in a semicircular formation.[42] The gestation period lasts 60–63 days, with litter sizes averaging 4-6 pups.[15] Their growth rate is much faster than that of wolves, being similar in speed to that of coyotes. Pups are suckled until at least the age of 58 days. During this time, the pack feeds the mother at the den site. Dholes do not use rendezvous sites to meet their pups as wolves do, though one or more adults will stay with the pups at the den while the rest of the pack hunts. Once weaning begins, the adults of the clan will regurgitate food for the pups until they are old enough to join in hunting. They remain at the den site till the age of 70–80 days. By the age of six months, pups accompany the adults on hunts, and will assist in killing large prey such as sambar by the age of 8 months.[43]
Denning behaviour[]
Four kinds of den have been described; simple earth dens with one entrance (usually remodeled striped hyena or porcupine dens); complex cavernous earth dens with more than one entrance; simple cavernous dens excavated under or between rocks; and complex cavernous dens with several other dens in the vicinity, some of which are interconnected. Dens are typically located under dense scrub or on the banks of dry rivers or creeks. The entrance to a dhole den can be almost vertical, with a sharp turn 3–4 feet down. The tunnel opens into an antechamber, from which extends more than one passage. Some dens may have up to six entrances leading up to 100 feet (30 m) of interconnecting tunnels. These "cities" may be developed over many generations of dholes, and are shared by the clan females when raising young together.[44] Like African wild dogs and dingoes, dholes will avoid killing prey close to their dens.[43]
Diet, hunting and feeding behaviours
Prey animals in India include chital, sambar, muntjac, mouse deer, swamp deer, wild boar, gaur, water buffalo, banteng, cattle, nilgai, goats, Indian hares, Himalayan field rats and langurs.[4][15][45] There is one record of a pack bringing down an Indian elephant calf in Assam, despite desperate defense of the mother resulting in numerous losses to the pack.[16] In Kashmir, they may hunt markhor,[45] and thamin in Burma.[15] Javan rusas are hunted in Java.[31] In the Tien Shan and Tarbagatai Mountains, dholes prey on Siberian ibexes, arkhar, roe deer, maral and wild boar. In the Altai and Sayan Mountains, they prey on musk deer and reindeer. In eastern Siberia, they prey on roe deer, Manchurian wapiti, wild boar, musk deer, and reindeer, while in Primorye they feed on sika deer and goral too. In Mongolia, they prey on argali and rarely Siberian ibex.[46] Like African wild dogs, but unlike wolves, dholes are not known to attack people.[47][48] 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.[49] In summertime in the Tien Shan Mountains, dholes eat large quantities of mountain rhubarb.[46] Bael fruits are also eaten.[50] Although opportunistic, dholes have a seeming aversion to hunting cattle and their calves.[51] Livestock predation by dholes has been a problem in Bhutan since the late 1990s, as domestic animals are often left outside to graze in the forest, sometimes for weeks at a time. Livestock stall-fed at night and grazed near homes are never attacked. Oxen are killed more often than cows are, probably because they are given less protection.[52]
Before embarking on a hunt, clans go through elaborate prehunt social rituals involving nuzzling, body rubbing and homo and heterosexual mounting.[53] Dholes are primarily diurnal hunters, hunting in the early hours of the morning. They rarely hunt nocturnally, except on moonlit nights, indicating that they greatly rely on sight when hunting.[54] Though not as fast as jackals and foxes, they can chase their prey for many hours.[46] During a pursuit, one or more dholes may take over chasing their prey, while the rest of the pack keeps up at a steadier pace behind, taking over once the other group tires. Most chases are short, lasting only 500 metres.[55] When chasing fleet-footed prey, they run at a pace of 30 mph.[50] Dholes frequently drive their prey into water bodies, where the targeted animal's movements are hindered.[56]
Once large prey is caught, one dhole will grab the prey's nose, while the rest of the pack pulls the animal down by the flanks and hind quarters. They do not use a killing bite to the throat.[29] They occasionally blind their prey by attacking the eyes.[57] 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.[58] They will tear open their prey's flanks and disembowel it, eating the heart, liver, lungs and some sections of the intestines. The stomach and rumen are usually left untouched.[5] Prey weighing less than 50 kg is usually killed within two minutes, while large stags may take 15 minutes to die. Once prey is secured, dholes will tear off pieces of the carcass and eat in seclusion.[59] Unlike wolf packs, in which the breeding pair monopolises food, dholes give priority to the pups when feeding at a kill, allowing them to eat first.[6] They are generally tolerant of scavengers at their kills.[60]
Relationships with other predators[]
In some areas, dholes are sympatric to tigers and leopards. Competition between these species is mostly avoided through differences in prey selection, although there is still substantial dietary overlap.[61] 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).[62] Also, other characteristics of the prey, such as sex, arboreality, and aggressiveness, may play a role in prey selection. For example, dholes preferentially select male chital whereas leopard kill both sexes more evenly (and tigers prefer larger prey altogether), dholes and tigers kill langurs rarely compared to leopards due to the leopard's greater arboreality, while leopards kill wild boar infrequently due to the inability of this relatively light predator to tackle aggressive prey of comparable weight.[62] On some rare occasions, dholes may attack tigers. When confronted by dholes, tigers will seek refuge in trees or stand with their backs to a tree or bush, where they may be mobbed for lengthy periods before finally attempting escape. Escaping tigers are usually killed, while tigers which stand their ground have a greater chance of survival.[47][63] Tigers are extremely dangerous opponents for dholes, as they have sufficient strength to kill a single dhole with one paw strike. Even a successful tiger kill is usually accompanied by losses to the pack.[64] Dhole packs may steal leopard kills, while leopards may kill dholes if they encounter them singly or in pairs.[45] Because leopards are smaller than tigers, and are more likely to hunt dholes, dhole packs tend to react more aggressively toward them than they do with tigers.[20] It was once thought that dholes were a major factor in reducing Asiatic cheetah populations, though this is doubtful, as cheetahs live in open areas as opposed to forested areas favoured by dholes.[65]
Dhole packs occasionally attack Asiatic black bears and sloth bears. When attacking bears, dholes will attempt to prevent them from seeking refuge in caves, and lacerate their hind quarters.[45]
Though usually antagonistic toward wolves,[66] they may hunt and feed alongside one another.[50] They infrequently associate in mixed groups with golden jackals. Domestic dogs may kill dholes, though they will feed alongside them on occasion.[67]/
Communication[]
Dholes produce whistles resembling the calls of red foxes, sometimes rendered as "coo-coo". How this sound is produced is unknown, though it is thought to help in coordinating the pack when travelling through thick brush. When attacking prey, they emit screaming "KaKaKaKAA" sounds.[68] Other sounds include whines (food soliciting), growls (warning), screams, chatterings (both of which are alarm calls) and yapping cries.[69] In contrast to wolves, dholes do not howl.[70]
Dholes have a complex body language. Friendly or submissive greetings are accompanied by horizontal lip retraction and the lowering of the tail, as well as licking. Playful dholes will open their mouths with their lips retracted and their tails held in a vertical position whilst assuming a play bow. Aggressive or threatening dholes will pucker their lips forward in a snarl and raise the hairs on their backs, as well as keep their tails horizontal or vertical. When afraid, they pull their lips back horizontally with their tails tucked and their ears flat against the skull.[41]/
Range[]
Historical[]
Dholes once ranged throughout most of South, East and Southeast Asia, extending from the Tien Shan and Altai Mountains and the Primorsky Krai southward through Mongolia, Korea, China, Tibet, Nepal, India, and south-eastwards into Myanmar and Indochina, Thailand, Malaysia, Sumatra and Java.[72]
During the last glacial period, they ranged across most of Eurasia, and are known to have once inhabited North America from a single fossil find in the Gulf of Mexico.[73] A canid called the Sardinian Dhole (Cynotherium sardous) lived on the Mediterranean island of Sardinia during the Pleistocene, but it is not as closely related to the living species as its name would imply.[74]
Current[]
There are currently no reports of dholes in Russia, Mongolia, Kazakhstan, Kyrgystan and Tajikistan. There is one report of a dhole being captured in southern China's Jiangxi district. Dholes still occur in Tibet, particularly in the Ladakh region bordering India and south-east Tibet. They may still be present in North Korea. They still occur in India south of the Ganges River, especially in the Central Indian Highlands and the Western and Eastern Ghats. Dholes also occur in northeast India's states of Arunachal Pradesh, Assam, Meghalaya and West Bengal. They have a precarious, fragmented distribution in Himalaya and north-west India. They are occasionally reported in the Ladakh area of Kashmir, contiguous with the Tibetan highlands and China. In Nepal, dholes were formerly recorded in Terai, including the Royal Chitwan National Park. Dholes were reported in the Dhorpatan Hunting Reserve in the late 1990s. In Bhutan, dholes have since recovered from a government sponsored poisoning campaign started in the 1970s, with reports of livestock predation occurring in the lower Kheng region. It is uncertain if they still occur in Bangladesh. Camera trapping has confirmed that dholes still occur in 11 survey areas in Myanmar, where they have replaced tigers as main predators. Dhole populations are highly fragmented in Thailand and Indochina, particularly in Vietnam. Dholes are known to occur in four sites in northern and central Malaysia. In Java, they appear to be most common in the island's protected eastern and western ends. They are also known to occur in Sumatra's protected areas in the southern, central and northern areas.[72]
