Dhole

• 1. Introduction

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  The dhole, a reddish wild dog of Asia, cannot howl and does not bark. Its voice is a whistle that resembles the call of a red fox, sometimes written as coo-coo. Scientists know the animal as Cuon alpinus, a canid native to South, East and Southeast Asia. Today the IUCN lists it as Endangered, with fewer than 2,500 mature individuals left. Why does a dog with such unusual teeth baffle the experts who try to place it on the canid family tree? How did an animal that once ranged from Europe across Asia end up scattered in small pockets of forest? And what does a clan of whistling hunters do that wolves do not?

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• 2. Teeth and Fur

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  The dhole is the only member of the genus Cuon, and its skull is the first clue to why. In profile the skull curves outward, where the skulls of Canis species curve inward. It is wide and massive, with a well-developed sagittal crest, the ridge along the top where jaw muscles attach. The masseter muscles are large even by canid standards, giving the face an almost hyena-like look. The snout is shorter than in domestic dogs and most other canids.

  The teeth are stranger still. A dhole has six lower molars rather than seven, because it lacks the third lower molar. Its upper molars are weak, only one third to one half the size of a wolf's. Each has a single cusp, where most canids have between two and four. This shape is thought to improve shearing and so speed up how fast prey is eaten. That speed may help dholes compete with kleptoparasites, animals that try to steal a kill.

  The body is slight for such a determined hunter. Dholes average about 88-113 cm in length, not counting a tail of 41-50 cm, and stand 42-50 cm at the shoulder. Adult females weigh 10-17 kg, while the slightly larger males weigh 15-21 kg. Observers have described the animal as a mix of gray wolf and red fox. Others call it cat-like, because of its long backbone and slender limbs.

  The coat is brightest in winter. The back turns a saturated rusty-red, with brownish highlights on the head, neck and shoulders. The throat, chest, flanks and belly are more yellowish, and the lower legs are whitish. The tail is fluffy, mostly reddish-ocher, with a dark brown tip. The summer coat is shorter, coarser and darker, and dholes at the Moscow Zoo moult once a year from March to May. Not every dhole fits this palette, since a melanistic individual was recorded in the northern Coimbatore Forest Division in Tamil Nadu.

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• 3. Names and Kin

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  In 1808 a soldier named Thomas Williamson wrote down the word dhole after meeting the animal in the Ramghur district of India. His is possibly the earliest use of the word in English, and he called it a common local name. Nobody has settled where it came from. In 1827 Charles Hamilton Smith said it came from a language spoken in 'various parts of the East'. Two years later he tied it to a word for 'mad, crazy', then made a comparison that proved wrong, since the look-alike words descend from Proto-Germanic *dwalaz, meaning 'foolish, stupid'. Richard Lydekker wrote nearly 80 years later that people living within the animal's range did not use the word at all. The Merriam-Webster Dictionary theorises that it may come from the word written ತೋಳ.

  The animal has collected many other English names. They include Asian wild dog, Indian wild dog, whistling dog, red dog and red wolf.

  Scientific names came just as untidily. In 1811 Peter Simon Pallas called it Canis alpinus and placed it around the upper Lena River, the Yenisei River and the upper Udskoi Ostrog in Amurland. That is considerably north of where the species lives now. In 1833 Brian Houghton Hodgson named it Canis primaevus, believing it a primitive dog and the ancestor of the domestic dog. He later noticed how physically distinct it was and proposed the genus Cuon.

  The teeth then sent later scholars in the wrong direction. George Simpson grouped the dhole with the African wild dog and the bush dog in the subfamily Simocyoninae, because all three have similar dentition. Juliet Clutton-Brock and others countered that any resemblance to those two came from convergent evolution. The dhole looked closer to Canis, and genome studies agree. Zoologist Reginald Innes Pocock recorded a dhole in a menagerie in Madras that bred with a golden jackal. In 1955 the paleontologist Erich Thenius had proposed a golden jackal-like ancestor.

  In 2018 whole genome sequencing found strong evidence of ancient mixing between the dhole and the African wild dog. The two live far apart today. The study suggested their ancestors may have met in the Middle East and mixed in North Africa, though no fossil evidence places dholes in either region. The extinct Sardinian dhole, Cynotherium sardous, turned out to be the living dhole's closest relative. The two lineages split around 900,000 years ago, and both carry African wild dog ancestry, though modern dholes carry far more.

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• 4. Ice Age Wanderers

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  A jaw fragment and some teeth found in San Josecito Cave in northeast Mexico may be the only trace of a dhole in North America. They date to roughly 27,000-11,000 years ago. Other researchers call the evidence insufficient, or say it needs corroboration before anyone can name the species with confidence.

  Europe has a much firmer record. In his 1968 book Pleistocene Mammals of Europe, Bjorn Kurten described a primitive form, Canis majori, found in Valdarno, Italy, and in China. It was almost indistinguishable from Canis. Early dholes appear at Hundsheim and Mosbach in Germany. By the Late Pleistocene the European dhole, C. a. europaeus, looked modern and had a single-cusped slicing tooth, though it was as large as a wolf. It vanished from Europe at the end of the late Würm period. It may have lingered in the Iberian Peninsula into the early Holocene, and remains at Riparo Fredian in northern Italy are 10,800 years old.

  Asia's islands once held dholes too. Fossils come from Sri Lanka and Borneo, and possibly Palawan in the Philippines. Middle Pleistocene bones lie in Matsukae Cave in northern Kyushu and in the Lower Kuzuu fauna in Tochigi Prefecture on Honshu. Luobi Cave on Hainan Island in south China yielded Late Pleistocene fossils dated to about 10,700 years ago. Canid fossils from the Dajia River in Taichung County, Taiwan, may also belong to dholes. Overall the animal was confined to its historical range only 12,000-18,000 years ago.

  In 2021 researchers studied mitochondrial genomes from two dholes found in the Jáchymka cave in the Czech Republic, dated to 35,000-45,000 years old. These European animals sat at the base of the modern dhole family tree, and they carried much greater genetic diversity than living dholes.

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• 5. Life in the Clan

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  In India a dhole clan typically holds 5 to 12 animals, though clans of 40 have been reported. In Thailand clans rarely exceed three individuals. The word clan is deliberate, since a pack means animals that always hunt together, and dholes often split. In spring a clan breaks into small packs of three to five, the best number for catching fawns.

  Dholes are more social than gray wolves and keep a looser hierarchy, because seasonal food shortage is not a serious worry for them. In this they resemble African wild dogs. Dominant animals are hard to spot, since dholes do not stage dominance displays, though others behave submissively toward them. Fighting within the group is rarely seen. They are far less territorial than wolves, and pups often move into another clan once they mature sexually.

  There is no evidence that dholes mark territory with urine, and they do not scrape the ground with their feet. Males especially may raise one or both hind legs and urinate in a handstand, as bush dogs and domestic dogs do. They leave droppings in communal latrines, mostly inside the clan's territory rather than at its edge.

  Their body language is rich. A friendly dhole retracts its lips horizontally, lowers its tail and licks. A playful one opens its mouth, holds its tail up and drops into a play bow. A threatening one puckers its lips into a snarl and raises the hair on its back. A frightened one pulls its lips back, tucks its tail and flattens its ears.

  Six entrances and up to 30 m of interconnected tunnels mark the largest dens, which the source describes as cities. Four kinds are known, from simple earth dens with one entrance, often remodeled from striped hyena or porcupine dens, to complex cavernous ones with several linked dens nearby. They sit under dense scrub or on dry river banks. Several females share a den to raise young together, and the clan avoids killing prey nearby, as African wild dogs and dingoes do.

  In India the mating season runs from mid-October to January. The female crouches in a cat-like position, and afterward there is no copulatory tie. The pair simply lie on their sides, facing each other in a semicircle. Gestation lasts 60-63 days and litters average four to six pups. The pack feeds the mother at the den while the pups suckle for at least 58 days. Adults then regurgitate food for the pups, which stay at the den for 70-80 days. By six months the pups join hunts, and by eight months they help kill large prey such as sambar. In captivity a dhole may live 15-16 years.

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• 6. The Chase

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  Before a hunt, a dhole clan goes through a ritual of nuzzling, body rubbing and mounting. The hunt itself usually comes in the early hours of the morning. Dholes rarely go out at night except under a moon, which shows how much they rely on sight. They can chase prey for many hours, with one or more dholes taking the lead while the rest follow at a steadier pace and take over when the leaders tire. Most chases are short, covering only 500 m, and the dholes reach about 30 mph over open ground. They often drive prey into water, where its movement is hindered. When attacking they scream KaKaKaKAA.

  Once they catch a large animal, one dhole grabs its nose while the others pull it down by the flanks and hindquarters. There is no killing bite to the throat. They sometimes blind prey by attacking the eyes. Serows can fight back, thanks to thick coats and short, sharp horns that can impale a dhole. Prey under 50 kg usually dies within two minutes, though a large stag may take 15 minutes. The dholes tear open the flanks and eat the heart, liver, lungs and parts of the intestines. The stomach and rumen usually go untouched.

  The menu spans the continent. In India it includes chital, sambar, wild boar, gaur and langurs, and there is one record of a pack bringing down an Indian elephant calf in Assam despite the mother's desperate defense. In Kashmir they take markhor, and in Sumatra the Sumatran serow. In the Altai and Sayan Mountains they take musk deer and reindeer, and in Mongolia argali. Dholes also eat insects, lizards, fruit and vegetable matter, and in the summer in the Tian Shan Mountains they eat large quantities of mountain rhubarb. They show a seeming aversion to cattle, and unlike wolves are not known to hunt people actively.

  In forests shared with tigers and leopards, differences in prey keep the peace, though the diets still overlap substantially. Dholes and leopards usually take animals of 30-175 kg, with mean prey weights of 35.3 kg for dholes and 23.4 kg for leopards. Tigers favor animals heavier than 176 kg. Tigers can kill a dhole with a single paw strike, and dhole packs are smaller where tigers are dense. Because tigers steal kills, dholes prefer smaller carcasses they can eat more of before a tiger arrives. Dhole packs also steal leopard kills, and there are many records of leopards treed by dholes. Packs occasionally attack Asiatic black bears, snow leopards and sloth bears, lacerating the hindquarters so the bears cannot reach caves.

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• 7. The Shrinking Map

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  The last known captures of dholes in South Korea were reported in 1909 and 1921, at Yeoncheon in Gyeonggi Province. Records in the Veritable Records of the Joseon Dynasty show the species once lived in Yangju. Hainan Island has dhole records from 1521-1935 and the species is estimated to have died out there around 1942. The Tien Shan subspecies, C. a. hesperius, is considered extinct since 1946. The dhole has also vanished from Central Asian countries including Afghanistan, Kazakhstan and Mongolia, and in India it disappeared from 60% of its historic range over the past century.

  Russia shows the same retreat. Dholes were considered rare and endangered in Primorsky Krai in 2004, and no sighting has been reported in other areas since the late 1970s. The IUCN considers the animal extinct in Russia, although it might survive in the eastern Sayan Mountains and the Transbaikal region.

  Still, camera traps keep turning up proof of life. Packs were recorded in Chitwan National Park and the Kanchenjunga Conservation Area in 2011. In 2014 videos from Kerinci Seblat National Park in Sumatra showed dholes at 2000 m. In 2015 dholes and tigers were recorded together for the first time in Karen State, Myanmar. In China, camera traps since 2008 have confirmed dholes in Gansu, Shaanxi, Qinghai, Yunnan, Sichuan, Xinjiang and Tibet. In February 2020 dholes were sighted in Vansda National Park, and camera traps confirmed two individuals there in May. It was the first confirmed sighting in Gujarat since 1970. Genetic samples collected in 2019 in southern Kyrgyzstan suggested dholes were likely present, the first record there in almost three decades.

  Some supposed sightings fail on inspection. A skull from a single animal claimed to have been shot in 2013 in Kabardino-Balkaria in the central Caucasus was first judged to be a dhole. Photographs studied in 2015 showed it was a grey wolf. That same year two skins from Turkey were judged probably to belong to dogs.

  Persecution did much of the damage. In India dholes were hunted for bounties until the Wildlife Protection Act of 1972 protected them. Methods included poisoning, snaring, shooting and clubbing at den sites. British sporthunters during the British Raj believed dholes were cutting game numbers. The bounty was 25 rupees, then cut to 20 in 1926 because the carcasses presented became too numerous to pay for. Fur was never the main threat. The Chinese bought pelts in Ussuriysk in the late 1860s for a few silver rubles, and in the early 20th century pelts in Manchuria reached eight rubles.

  Other pressures keep building. Fragmented populations suffer inbreeding and the Allee effect, which threaten long-term viability. Diseases include rabies, canine distemper, mange and canine parvovirus. In Bhutan, livestock grazing unattended in the forest for weeks has led to predation since the late 1990s, whereas stock kept in stalls at night is never attacked.

  Protection is patchy but real. India protects the dhole under Schedule 2 of its 1972 act, and Project Tiger reserves shelter some populations. In 2014 India approved its first dhole conservation breeding centre at the Indira Gandhi Zoological Park in Visakhapatnam. Russia has protected the animal since 1974, but it remains vulnerable to poison left out for wolves. China listed it as a category II protected species under its 1988 wildlife law. In 2016 the Korean company Sooam Biotech was reported to be trying to clone the dhole using dogs as surrogate mothers.

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• 8. The Red Devil in Story

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  Carved on the coping stone of the Bharhut stupa, dated to 100 BC, three dhole-like animals wait by a tree while a woman or spirit is trapped up it. The scene resembles dholes treeing tigers. In Tamil Sangam literature of 100 BC to 200 AD the dhole recurs as a character. Ainkurunuru 323 tells of a pregnant dhole craving wild pig, so her mate goes hunting for pig.

  In Hindi the animal's fearsome reputation shows in its names, which translate as "red devil", "devil dog", "jungle devil" and "hound of Kali". Rudyard Kipling is not named in the source, but The Second Jungle Book features dholes, called red dogs, in the chapter Red Dog. There Mowgli and the Seeonee Wolf Pack must fight a pack of them that has invaded their jungle.

  Fear varied by people. Leopold von Schrenck had trouble getting specimens in Amurland because the local Gilyaks dreaded the species. Neighbouring Tungusic peoples did not share the superstition. It was speculated this was because of their more nomadic, hunter-gatherer way of life.

  In 1901 the Japanese author Uchida Roan wrote about the declining popularity of indigenous dog breeds, which he held to be descended from the dhole. In China the dhole appears throughout history and mythology, and the Nine sons of the dragon included a creature that was part dhole and part dragon. The modern Chinese word for dhole is often confused with 'jackal' or 'wolf', so the animal is often mistranslated. An inscription may even record a dhole sent as a gift to the Akkadian king Ibbi-Sin, referred to as the "red dog of Meluhha", a name for the Indus Valley Civilisation.

  Hodgson kept captured dholes and found that, with one exception, they stayed shy and vicious even after 10 months. Adults are nearly impossible to tame. Pups, however, are docile and can play with domestic dog pups until they reach early adulthood.

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