About Cuon alpinus (Pallas, 1811)
Cuon alpinus (Pallas, 1811), commonly called the dhole, has characteristic fur with an overall reddish base tone, which is brightest during winter. In winter coat, the back is a deep rusty-red to reddish shade, with brownish highlights along the top of the head, neck, and shoulders. The throat, chest, flanks, belly, and upper limbs are paler, with a more yellowish tone. The lower limbs are whitish, with dark brown bands on the front sides of the forelimbs. The muzzle and forehead are greyish-reddish. The tail is very full and fluffy, primarily reddish-ochre with a dark brown tip. The summer coat is shorter, coarser, and darker. Adult dorsal and lateral guard hairs measure 20–30 mm (0.79–1.18 in) long. Dholes held at the Moscow Zoo moult once per year between March and May. One melanistic dhole individual has been recorded in the northern Coimbatore Forest Division of Tamil Nadu. The dhole has a wide, massive skull with a well-developed sagittal crest, and highly developed masseter muscles compared to other canid species, which gives its face an almost hyena-like look. Its rostrum is shorter than that of domestic dogs and most other canids. It has six lower molars, rather than the seven seen in most canids. Its upper molars are weak, measuring one third to one half the size of a wolf’s upper molars, and only have one cusp, unlike the two to four cusps typical for canids. This tooth structure is thought to be an adaptation that improves shearing ability and speeds up prey consumption, which may help dholes compete more successfully against kleptoparasites. In terms of body size, dholes have an average head-body length of 88–113 cm (35–44 in), not counting their 41–50 cm (16–20 in) long tail, and stand 42–50 cm (17–20 in) tall at the shoulder. Adult females weigh 10–17 kg (22–37 lb), while slightly larger males weigh 15–21 kg (33–46 lb). The average weight of adult dholes from three small study samples was 15.1 kg (33 lb). Dholes have been described as combining physical traits of gray wolves and red foxes, and as having a "cat-like" build due to their long backbone and slender limbs. Historically, dholes lived in Singapore and across Central Asia, including Afghanistan, Kyrgyzstan, Kazakhstan, Mongolia, Tajikistan, and Uzbekistan, but they are now regionally extinct in all these areas. Historical records from the Veritable Records of the Joseon Dynasty show dholes once lived in Yangju, Gyeonggi Province, South Korea, but the species is now extinct in South Korea; the last known captures were reported in 1909 and 1921 from Yeoncheon, Gyeonggi Province. The current presence of dholes in North Korea and Pakistan is considered uncertain. Dholes once inhabited the alpine steppes stretching from Kashmir into the Ladakh area, and have disappeared from 60% of their historic range in India over the past century. In India, Myanmar, Indochina, Indonesia, and China, dholes prefer forested areas in alpine zones, and are occasionally seen in plains regions. Genetic samples collected in 2019 from Bek-Tosot Conservancy, southern Kyrgyzstan, indicated dholes are likely present there; this was the first record of dholes in the country in almost 30 years. Dholes may still occur in Tunkinsky National Park, located in extreme southern Siberia near Lake Baikal. They may also still live in Primorsky Krai, far eastern Russia, where they were classified as a rare endangered species in 2004, with unconfirmed reports from the Pikthsa-Tigrovy Dom protected forest area; no confirmed sightings have been reported from other areas of the region since the late 1970s. Currently, there are no other confirmed recent reports of dholes in Russia, so the IUCN classifies them as extinct in Russia. However, dholes may be present in the eastern Sayan Mountains and the Transbaikal region; they have been sighted in Tofalaria (Irkutsk Oblast), the Republic of Buryatia, and Zabaykalsky Krai. One dhole pack was sighted in the Qilian Mountains in 2006. From 2011 to 2013, local government officials and herders reported several dhole packs living at elevations of 2,000 to 3,500 m (6,600 to 11,500 ft) near Taxkorgan Nature Reserve in China’s Xinjiang Autonomous Region. Camera traps also recorded several packs and an adult female with pups at elevations of roughly 2,500 to 4,000 m (8,200 to 13,100 ft) in Yanchiwan National Nature Reserve, northern Gansu Province, China, between 2013 and 2014. Dholes have also been reported in the Altyn-Tagh Mountains. In China’s Yunnan Province, dholes were recorded in Baima Xueshan Nature Reserve between 2010 and 2011. Dhole samples were collected in Jiangxi Province in 2013. Since 2008, camera traps have confirmed dhole presence in southern and western Gansu province, southern Shaanxi province, southern Qinghai province, southern and western Yunnan province, western Sichuan province, southern Xinjiang Autonomous Region, and the Southeastern Tibet Autonomous Region of China. Historical records of dholes on China’s Hainan Island date from 1521 to 1935, but the species is no longer present there and is estimated to have gone extinct around 1942. Dholes occur across most of India south of the Ganges, particularly in the Central Indian Highlands and the Western and Eastern Ghats. They are also present in Arunachal Pradesh, Assam, Meghalaya, West Bengal, and the Terai region of the Indo-Gangetic Plain. Dhole populations in the Himalayas and northwest India are fragmented. Camera traps recorded dhole packs in Chitwan National Park in 2011. Their presence was confirmed in the Kanchenjunga Conservation Area in 2011 via camera traps. In February 2020, dholes were sighted in Vansda National Park, India, and camera traps confirmed the presence of two individuals in May of that year; this was the first confirmed dhole sighting in Gujarat since 1970. In Bhutan, dholes are present in Jigme Dorji National Park. In Bangladesh, dholes inhabit forest reserves in the Sylhet area and the Chittagong Hill Tracts in the southeast. 2016 camera trap photos from Chittagong confirmed dholes continue to live there. These areas likely do not support a viable dhole population, as mostly small groups or solitary individuals have been sighted. In Myanmar, dholes are present in multiple protected areas. In 2015, camera traps recorded dholes and tigers for the first time in the hill forests of Karen State, Myanmar. Dhole range is highly fragmented in the Malaysian Peninsula, Sumatra, Java, Vietnam, and Thailand; the Vietnamese population is considered possibly extinct. In 2014, camera trap videos from montane tropical forests at 2,000 m (6,600 ft) in Sumatra’s Kerinci Seblat National Park confirmed dholes continue to occur there. A camera trapping survey conducted in Thailand’s Khao Ang Rue Nai Wildlife Sanctuary from January 2008 to February 2010 documented one healthy dhole pack. In northern Laos, dholes have been studied in Nam Et-Phou Louey National Protected Area, and camera trap surveys from 2012 to 2017 recorded dholes in this same protected area. In Vietnam, dholes were only sighted in Pu Mat National Park in 1999, Yok Don National Park in 2003 and 2004, and Ninh Thuan Province in 2014. A separate disjunct dhole population was reported in the Trabzon and Rize area of northeastern Turkey near the Georgian border in the 1990s; this report is not considered reliable. A single dhole individual was claimed to have been shot in 2013 in the nearby Kabardino-Balkaria Republic of Russia, in the central Caucasus. A biologist from Kabardino-Balkarian State University analyzed the remains in May 2015, and concluded the skull belonged to a dhole. In August 2015, researchers from the National Museum of Natural History and Karadeniz Technical University launched an expedition to track and document a possible Turkish dhole population. In October 2015, they concluded two skins claimed to be from dholes in Turkey most likely belonged to dogs, pending further DNA analysis of skin samples. After analyzing photos of the claimed dhole skull from Kabardino-Balkaria Republic, Russia, they concluded it was actually a gray wolf skull. In terms of ecology and behaviour, dholes produce whistles that resemble the calls of red foxes, sometimes described as coo-coo. It is unknown how this sound is produced, but it is thought to help packs coordinate when moving through thick brush. When attacking prey, dholes emit screaming KaKaKaKAA sounds. Other sounds dholes make include whines (used when soliciting food), growls (used as a warning), screams, chatterings (both screams and chatterings function as alarm calls), and yapping cries. Unlike wolves, dholes do not howl or bark. Dholes have complex body language. Friendly or submissive greetings involve horizontal lip retraction, lowering the tail, and licking. Playful dholes open their mouths with retracted lips, hold their tails vertically, and perform a play bow. Aggressive or threatening dholes push their lips forward into a snarl, raise the hairs on their backs, and hold their tails horizontally or vertically. When afraid, dholes pull their lips back horizontally, tuck their tails, and flatten their ears against their skull. In India, the dhole mating season falls between mid-October and January, while captive dholes at the Moscow Zoo mostly breed in February. Unlike wolf packs, dhole clans can include more than one breeding female. Multiple female dholes may share a den and raise their litters together in the same space. During mating, the female assumes a crouched, cat-like position. There is no copulatory tie typical of other canids after the male dismounts; instead, the pair lie on their sides facing each other in a semicircular shape. The gestation period lasts 60–63 days, and litter sizes average four to six pups. A study of hormone metabolites from five male and three female captive dholes in Thai zoos found breeding males have increased testosterone levels from October to January. Captive females have increased oestrogen levels for roughly two weeks in January, followed by increased progesterone, and display sexual behaviours during the oestrogen peak. Pups are suckled for at least 58 days. During this period, the pack brings food to the mother at the den site. Unlike wolves, dholes do not use rendezvous sites to meet their pups; instead, one or more adults stay with the pups at the den while the rest of the pack hunts. Once weaning begins, adult clan members regurgitate food for the pups until the pups are old enough to join hunts. Pups remain at the den site for 70–80 days. By six months of age, pups accompany adults on hunts, and can assist in killing large prey such as sambar by eight months of age. The maximum recorded longevity of captive dholes is 15–16 years. In terms of feeding ecology, dhole prey in India includes chital, sambar deer, muntjac, mouse deer, barasingha, wild boar, gaur, water buffaloes, banteng, cattle, nilgai, goats, Indian hares, Himalayan field rats, and langurs. There is one recorded case of a dhole pack killing an Indian elephant calf in Assam, despite a desperate defensive effort from the mother that caused numerous injuries to pack members. In Kashmir, dholes prey on markhor; in Myanmar they prey on thamin; in Sumatra and the Malay Peninsula they prey on Malayan tapir and Sumatran serow; and in Java they prey on Javan rusa. In the Tian Shan and Tarbagatai Mountains, dholes prey on Siberian ibexes, arkhar, roe deer, Caspian red deer, 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 they feed on sika deer and goral. In Mongolia, they prey on argali and rarely on Siberian ibex. Like African wild dogs, but unlike wolves, dholes are not known to actively hunt humans. Dholes are known to eat insects and lizards. They eat fruit and plant matter more readily than other canids. In captivity, dholes eat various kinds of grasses, herbs, and leaves, apparently for pleasure rather than only when ill. In summer in the Tian Shan Mountains, dholes eat large amounts of mountain rhubarb. Although dholes are opportunistic hunters, they appear to avoid hunting cattle and their calves. Livestock predation by dholes has been a problem in Bhutan since the late 1990s, because domestic animals are often left to graze unsupervised in the forest for weeks at a time. Livestock that are stall-fed at night and graze near human homes are never attacked. Oxen are killed more often than cows, likely because oxen receive less protection.
