Golden jackal

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Golden jackal
Temporal range: Late Pleistocene – Recent
Golden jackals howling
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Carnivora
Family: Canidae
Genus: Canis
Species:
C. aureus
Binomial name
Canis aureus
Subspecies
  • C. a. aureus
  • C. a. cruesemanni
  • C. a. ecsedensis
  • C. a. indicus
  • C. a. moreoticus
  • C. a. naria
  • C. a. syriacus
Range of the golden jackal

The golden jackal (Canis aureus) is a Eurasian canine that is similar in appearance to a small gray wolf. Compared with the gray wolf, the golden jackal is smaller in size and lighter in weight, and has shorter legs, a shorter tail, a more elongated torso, a less-prominent forehead, and a narrower and more pointed muzzle. Male golden jackals weigh 6–14 kg (13–31 lb) compared with the smallest wolf, the Arabian wolf, which weighs on average 20 kg (44 lb). Its winter fur also differs from a wolf's by its more fulvous-reddish colour. It is native to Southeast Europe, Southwestern Asia, South Asia, and regions of Southeast Asia. In Europe it is now expanding into the Baltic states and Central Europe. It is listed as Least Concern on the IUCN Red List due to its widespread range in areas with optimum food and shelter.

The distant ancestor of the golden jackal is believed to be the extinct Arno river dog that lived in Mediterranean Europe 1.9 million years ago. It is described as being a small, jackal-like canine. Genetic studies indicate that the golden jackal expanded from India at the end of the last ice age around 20,000 years ago. The oldest golden jackal fossil is 20,000 years old and was found at the Ksar Akil rock shelter near Beirut, Lebanon. The oldest golden jackal fossils in Europe were found in Greece and are 7,000 years old. There are seven sub-species of the golden jackal. Despite its name, the golden jackal is not closely related to the African black-backed or side-striped jackals, being instead more closely related to the gray wolf, coyote, African golden wolf, and Ethiopian wolf. It is capable of producing fertile hybrids with both the gray wolf and the African golden wolf. Dog-jackal hybrids called Sulimov dogs are in service at the Sheremetyevo Airport near Moscow where they are deployed by the Russian airline Aeroflot for scent-detection.

Golden jackals are abundant in valleys and beside rivers and their tributaries, canals, lakes, and seashores. They are rare in foothills and low mountains, and they avoid waterless deserts and snow areas. The golden jackal is a social species, the basic social unit of which consists of a breeding pair and any young offspring. It is very adaptable, with the ability to exploit food ranging from fruit and insects to small ungulates. They will attack domestic fowl and domestic mammals up to the size of domestic water buffalo calves. The jackal's competitors are the red fox, wolf, jungle cat, forest wildcat, and the raccoon in the Caucausus, and the steppe wildcat in Central Asia. In Europe, jackals will not occupy the same areas as wolves, with the jackal's expansion being attributed to their occupying those areas where wolves are few or non-existent.

Etymology

The word "jackal" appeared in the English language around 1600. It derives from the Turkish word çakal, which originates from the Persian word šagāl.[3]

Taxonomy

Phylogenetic tree of the extant wolf-like canids
Caninae 3.5 Ma
Phylogenetic relationships between the extant wolf-like clade of canids based on nuclear DNA sequence data taken from the cell nucleus,[4][5] except for the Himalayan wolf, based on mitochondrial DNA sequences.[5][6] Timing in millions of years.[5]

The golden jackal (Canis aureus, "golden dog"), also known as the common jackal, Asiatic jackal,[1][7] or Eurasian golden jackal,[5] was first recorded by the Swedish zoologist Carl Linnaeus in his 1758 publication Systema Naturae.[2] The mammalogist W. Christopher Wozencraft listed thirteen subspecies under C. aureus in the third edition of Mammal Species of the World that was published in 2005.[8] Mitochondrial DNA (mDNA) passes along the maternal line and can date back thousands of years.[9] Therefore, phylogenetic analysis of mDNA sequences within a species provides a history of maternal lineages that can be represented as a phylogenetic tree.[10][11] Results from two recent studies of mDNA from golden jackals indicate that those specimens from Africa are genetically closer to the gray wolf than are the specimens from Eurasia.[12][13]

In 2015 a major DNA study of golden jackals proposes that the six C. aureus subspecies found in Africa should be reclassified under the new species C. anthus (African golden wolf),[5][14] which would reduce the number of golden jackal subspecies to seven. The phylogenetic tree generated from this study shows the golden jackal diverging from the wolf/coyote lineage 1.9 million years ago and the African golden wolf diverging 1.3 million years ago. The study found that the golden jackal and the African golden wolf shared a very similar skull and body morphology and that this had confused taxonomists into regarding these as one species. The study proposes that the very similar skull and body morphology is due to both species having originated from a larger common ancestor.[5]

Evolution

Three golden jackal–dog hybrids from Croatia[15]

The Arno river dog (Canis arnensis) is an extinct species of canine that was endemic to Mediterranean Europe during the Early Pleistocene around 1.9 million years ago. It is described as a small jackal-like dog and probably the ancestor of modern jackals.[16] Its anatomy and morphology relate it more to the modern golden jackal than to the two African jackal species,[17][18] the black-backed jackal (C. mesomelas) and the side-striped jackal (Canis adustus).

The oldest golden jackal fossil was found at the Ksar Akil rock shelter located 10 km (6.2 mi) northeast of Beirut, Lebanon. The fragment of a single tooth is dated to the end of the Last Glacial Maximum, approximately 20,000 years before present (or YBP).[19] The oldest golden jackal fossils found in Europe are from Delphi and Kitsos in Greece and are dated 7,000–6,500 YBP.[20] An unusual fossil found in Azokh Cave, Republic of Artsakh in Transcaucasia, dates to the Middle Pleistocene and is described as probably belonging to the golden jackal; however, its classification is not clear.[18] The absence of clearly identified golden jackal fossils in the Caucasus region and Transcaucasia, areas where the species currently resides, indicates that the species is a relatively recent arrival.[21]

A haplotype is a group of genes found in an organism that is inherited from one of its parents.[22][23] A haplogroup is a group of similar haplotypes that share a single mutation inherited from their common ancestor.[9] The mDNA haplotypes of the golden jackal form two haplogroups: the oldest haplogroup is formed by golden jackals from India, and the other, younger, haplogroup diverging from this includes golden jackals from all of the other regions.[24] Indian golden jackals exhibit the highest genetic diversity, and those from northern and western India are the most basal, which indicates that India was the center from which golden jackals spread. The extant golden jackal lineage commenced expanding its population in India 37,000 years ago. During the Last Glacial Maximum, 25,000 to 18,000 years ago, the warmer regions of India and Southeast Asia provided a refuge when compared with the surrounding areas. At the end of the Last Glacial Maximum and the beginning of the warming cycles, the golden jackal lineage expanded out of India and into Eurasia to reach the Middle East and Europe.[25]

Outside of India, golden jackals in the Caucasus and Turkey indicate the next highest genetic diversity,[24] while those in Europe indicate low genetic diversity,[26][27] which confirms their more recent expansion into Europe.[28] Genetic data indicates that the golden jackals of the Peloponnese Peninsula in Greece and the Dalmatian coast in Croatia may represent two ancient populations that have survived into modern times. Golden jackals in the Balkans originated from those expanding from southeast Europe and from the Caucasus. Golden jackals from both the Balkans and the Caucasus are expanding into the Baltic. In the Middle East, golden jackals from Israel have a higher genetic diversity than European jackals. This is thought to be due to Israeli jackals having hybridized with dogs, gray wolves, and African golden wolves,[28] creating a hybrid zone in Israel.[5]

All species within the wolf-like canids possess 78 chromosomes and therefore can potentially interbreed.[29] Genetic analysis reveals that hybridization takes place between gray wolves and jackals,[30] creating a jackal-wolf hybrid. Hybridization also occurs between female golden jackals and male dogs, which produces fertile offspring,[15] a jackal–dog hybrid. There was 11%–13% of ancient gene flow into the golden jackal from the population that was ancestral to wolves and dogs, and an additional 3% contributed from extant wolf populations.[31][32] Up to 15% of the Israeli wolf genome is derived from admixure with golden jackals in ancient times.[31]

Subspecies

Physical description

Skull, as illustrated by Charles Knight
Gray wolf and golden jackal exhibit at The Museum of Zoology, St. Petersburg – note the jackal's smaller size and narrower muzzle.

The golden jackal is very similar to the gray wolf but is distinguished by its smaller size, lighter weight, more elongated torso, less-prominent forehead, shorter legs and tail, and a muzzle that is narrower and more pointed. Males measure 71–85 cm (28–33 in) in body length and females 69–73 cm (27–29 in). Males weigh 6–14 kg (13–31 lb) and females weigh 7–11 kg (15–24 lb). The shoulder height is 45–50 cm (18–20 in) for both.[49] In comparison, the smallest wolf is the Arabian wolf (Canis lupus arabs), which weighs on average 20 kg (44 lb).[50] The legs of the golden jackal are long in relation to its body, and the feet are slender with small pads. Females have four pairs of teats.[7]

The skull is most like that of the dingo, and is closer to that of the coyote (C. latrans) and the gray wolf (C. lupus) than to that of the black-backed jackal (C. mesomalas), the side-striped jackal (C. adustus), and the Ethiopian wolf (C. simensis).[51] Compared with the wolf, the skull of the golden jackal is smaller and less massive, with a lower nasal region and shorter facial region; the projections of the skull are prominent but weaker than those of the wolf; the canine teeth are large and strong but relatively thinner; and its carnassial teeth are weaker.[49] The golden jackal is a less specialized species than the gray wolf, and these skull features relate to the jackal's diet of small birds, rodents, small vertebrates, insects, carrion,[52] fruit, and some vegetable matter.[51] Occasionally, the golden jackal develops a horny growth on the skull referred to as a "jackal's horn", which usually measures 1.3 cm (0.51 in) in length and is concealed by fur. This feature was once associated with magical powers by the people of Sri Lanka.[53]

The jackal's fur is coarse and relatively short,[51] with the base color golden, varying seasonally from a pale creamy yellow to a dark tawny. The fur on the back is composed of a mixture of black, brown, and white hairs, sometimes giving the appearance of the dark saddle like that seen on the black-backed jackal. The underparts are a light pale ginger to cream color. Individual specimens can be distinguished by their unique light markings on the throat and chest.[7] The coats of jackals from high elevations tend to be more buff-colored than those of their lowland counterparts[42] while those of jackals in rocky, mountainous areas may exhibit a grayer shade. The bushy tail has a tan to black tip.[7] Melanism can cause a dark-colored coat in some golden jackals, a coloring once fairly common in Bengal.[54] Unlike melanistic wolves and coyotes that received their dark pigmentation from interbreeding with domestic dogs, melanism in golden jackals probably stems from an independent mutation that could be an adaptive trait.[55] What is possibly an albino specimen was photographed in southeastern Iran during 2012.[56]

The jackal moults twice a year, in spring and in autumn. In Transcaucasia and Tajikistan, the spring moult begins at the end of winter. If the winter has been warm, the spring moult starts in the middle of February; if the winter has been cold, it begins in the middle of March. The spring moult lasts for 60–65 days; however, if the animal is sick, it loses only half of its winter fur. The spring moult commences with the head and limbs, extends to the flanks, chest, belly and rump, and ends at the tail. Fur on the underparts is absent. The autumn moult occurs from mid-September with the growth of winter fur; the shedding of the summer fur occurs at the same time. The development of the autumn coat starts with the rump and tail and spreads to the back, flanks, belly, chest, limbs and head, with full winter fur being attained at the end of November.[57]

Behavior

Golden jackals exhibit flexible social organization depending on the availability of food. The breeding pair is the basic social unit, and they are sometimes accompanied by their current litter of pups and older siblings. In India, their distributions are a single jackal, 31%, two jackals, 35%, three jackals, 14%, and more than three jackals, 20%.[7] Family groups of up to 4–5 individuals have been recorded.[58] Scent marking through urination and defecation is common around golden jackal den areas and on the trails they most often use. Scent marking is thought to assist in territorial defense. The hunting ranges of several jackals can overlap. Jackals can travel up to 12–15 km (7.5–9.3 mi) during a single night in search of either food or more suitable habitat. Non-breeding members of a pack may stay near a distant food source, such as a carcass, for up to several days before returning to their home range. Home range sizes can vary between 1–20 km2 (0.39–7.72 sq mi), depending on the available food.[7]

Social interactions such as greetings, grooming, and group howling are common in jackals. Howling is more frequent between December and April when pair bonds are being formed and breeding occurs, which suggests howling has a role in the delineation of territory and for defense.[7] Adult jackals howl standing and the young or subordinate jackals howl sitting.[59] Jackals are easily induced to howl and a single howl may solicit replies from several jackals in the vicinity. Howling begins with 2–3 low-pitched calls that rise to high-pitched calls.[7] The howl consists of a wail repeated 3–4 times on an ascending scale, followed by three short yelps that have been described as sounding similar to "Dead hindoo, where, where, where".[46] Jackals typically howl at dawn and in the evening, and sometimes at midday. Adults may howl to accompany the ringing of church bells, with their young responding to sirens or the whistles of steam engines and boats. When there is a change in the weather they will produce a long and continuous chorus.[60] Golden jackals give a warning call that is very different from their normal howling when they detect the presence of large carnivores such as wolves and tigers.[7][46] This howl is described as sounding similar to "pheal", "phion" or "phnew".[46] When hunting in a pack, the dominant jackal initiates an attack by repeatedly emitting a sound similar to "okkay!".[53]

Reproduction

Syrian jackal (C. a. syriacus) pup at the entrance to its den, park Yarkon, Israel

Golden jackals are monogamous and will remain with the one partner until death.[61] Female jackals have only one breeding cycle each year. Breeding occurs from October to March in Israel and from February to March in India, Turkmenistan,[7] Bulgaria, and Transcaucasia, with the mating period lasting up to 26–28 days. Females undergoing their first estrus are often pursued by several males that may quarrel among themselves.[61] Mating results in a copulatory tie that lasts for several minutes, as it does with all other canids. Gestation lasts 63 days, and the timing of the births coincides with the annual abundance of food.[7]

In India, the golden jackal will take over the dens of the Bengal fox and the Indian crested porcupine, and will use abandoned gray wolf dens.[7] Most breeding pairs are spaced well apart and maintain a core territory around their dens. Den excavations commence from late April to May in India, with dens located in scrub areas. Rivulets, gullies, and road and check-dam embankments are prime denning habitats. Drainage pipes and culverts have been used as dens. Dens are 2–3 m (6.6–9.8 ft) long and 0.5–1 m (1.6–3.3 ft) deep, with between 1–3 openings. Young pups can be moved between 2–4 dens.[7] The male helps with digging the den and raising the pups.[61] In the Caucasus and Transcaucasia, the burrow is located either in thick shrub, on the slopes of gullies, or on flat surfaces. In Dagestan and Azerbaijan, litters are sometimes located within the hollows of fallen trees, among tree roots, and under stones on river banks. In Central Asia, the golden jackal does not dig burrows but constructs lairs in dense tugai thickets. Jackals in the tugais and cultivated lands of Tajikistan construct lairs in long grass, shrubs, and reed openings.[58]

In Transcaucasia, golden jackal pups are born from late March to late April,[61] and in northeastern Italy during late April;[21] they can be born at any time of year in Nepal.[42] The number of pups born in a single litter varies geographically. Jackals in Transcaucasia give birth to 3–8 pups, Tajikistan 3–7 pups, Uzbekistan 2–8 pups, and Bulgaria 4–7 pups; in India the average is four pups.[61] The pups are born with closed eyes that open after 8–11 days, with the ears erecting after 10–13 days.[57] Their teeth erupt at 11 days after birth,[7] and the eruption of adult dentition is completed after five months. Pups are born with soft fur that ranges in color from light gray to dark brown. At the age of one month, their fur is shed and replaced with a new reddish-colored pelt with black speckles. The pups have a fast growth rate and weigh 0.201–0.214 kg (0.44–0.47 lb) at two days of age, 0.560–0.726 kg (1.23–1.60 lb) at one month, and 2.700–3.250 kg (5.95–7.17 lb) at four months.[57] Lactation lasts for up to 8–10 weeks.[7] The pups begin to eat meat at the age of 15–20 days.[57]

Dog pups show unrestrained fighting with their siblings from 2 weeks of age, with injury avoided only due to their undeveloped jaw muscles. This fighting gives way to play-chasing with the development of running skills at 4–5 weeks. Wolf pups possess more-developed jaw muscles from 2 weeks of age, when they first show signs of play-fighting with their siblings; serious fighting occurs during 4–6 weeks of age.[62] Compared to wolf and dog pups, golden jackal pups develop aggression at the age of 4–6 weeks, when play-fighting frequently escalates into uninhibited biting intended to harm. This aggression ceases by 10–12 weeks when a hierarchy has formed.[63] Once the lactation period concludes, the female drives off the pups. Pups born late remain with their mother until early autumn, at which time they leave either singly or in groups of two to four individuals. Females reach sexual maturity after 10–11 months and males at 21–22 months.[57]

Foraging

Pair of Sri Lankan jackals (C. a. naria) in Udawalawe National Park

The golden jackal often hunts alone, and sometimes in pairs, but rarely hunts in a pack. When hunting alone, it will trot around an area and occasionally stop to sniff and listen. Once prey is located, it will conceal itself, quickly approach, then pounce.[60] Single jackals hunt rodents, hares, and birds. They hunt rodents in grass by locating them with their hearing before leaping into the air and pouncing on them. In India, they can dig Indian gerbils out from their burrows, and they can hunt young, old, and infirm ungulates up to 4–5 times their body weight. Jackals search for hiding blackbuck calves throughout the day during the calving period. The peak times for their searches are the early morning and the late evening. When hunting in pairs or packs, jackals run parallel to their prey and overtake it in unison. When hunting aquatic rodents or birds, they will run along both sides of narrow rivers or streams and drive their prey from one jackal to another.[60]

Pack-hunting of langurs is recorded in India. Packs of between 5 and 18 jackals scavenging on the carcasses of large ungulates is recorded in India and Israel.[7] Packs of 8–12 jackals consisting of more than one family have been observed in the summer periods in Transcaucasia.[60] In India, the Montagu's harrier and the Pallid harrier roost in their hundreds in grasslands during their winter migration. Jackals stalk close to these roosting harriers and then rush at them, attempting to catch one before the harriers can take off or gain sufficient height to escape. In some regions of Bangladesh and India, jackals subsist by scavenging on carrion and garbage, and will cache extra food by burying it. Golden jackals cause damage to melon, peanut, grape, coffee, maize, and sugarcane crops. They sometimes prey on lambs, weak sheep, kids, goats, and poultry. The jackal will scavenge off the kills made by the lion, tiger, leopard, dhole, and gray wolf.[7]

Cooperation

In Southeastern Asia, golden jackals have been known to hunt alongside dhole packs.[35] They have been observed in the Blackbuck National Park, Velavadar, India, following Indian wolves (Canis lupus pallipes) when these are on a hunt, and they will scavenge off wolf kills without any hostility shown from these wolves.[7] In India, lone jackals expelled from their pack have been known to form commensal relationships with tigers. These solitary jackals, known as kol-bahl, will associate themselves with a particular tiger, trailing it at a safe distance to feed on the big cat's kills. A kol-bahl will even alert a tiger to prey with a loud pheal. Tigers have been known to tolerate these jackals, with one report describing how a jackal confidently walked in and out between three tigers walking together.[64][65] Golden jackals and wild boar can occupy the same territory.[44]

Ecology

Habitat

The golden jackal's omnivorous diet allows it to eat a large range of foods and to live in different habitats. The jackal's long legs in conjunction with its lithe body allows it to trot over great distances in search of food. It is able to go without water for extended periods and has been observed on islands that have no fresh water.[7] Jackals are abundant in valleys, beside rivers and their tributaries, canals, lakes, and seashores, but they are rare in foothills and low mountains. They avoid waterless deserts but can be found at their edges or in oases. They are found in dense thickets of prickly bushes, reed flood-lands and forests. They have been known to ascend over 1,000 m (3,300 ft) up the slopes of the Himalayas; they can withstand temperatures as low as −25 °C (−13 °F) and sometimes −35 °C (−31 °F). They are not adapted to snow, and in snow country they must travel along paths made by larger animals or humans. In India, they will occupy the surrounding foothills above arable areas,[66] entering human settlements at night to feed on garbage, and have established themselves around hill stations at 2,000 m (6,600 ft) height above mean sea level.[7] Although they generally avoid mountainous forests, they may enter alpine and sub-alpine areas during dispersal. In Turkey, the Caucasus, and Transcaucasia they have been observed at heights of up to 1,000 m (3,300 ft) above mean sea level, particularly in areas where the climate supports shrublands in high elevations.[21]

Diet

... It is more noisy in its pursuits even than the dog, and more voracious than the wolf.

The golden jackal is both a predator and a scavenger,[68] and an omnivorous and opportunistic forager with a diet that varies according to its habitat and the season. In Bharatpur, India, over 60% of its diet was measured to consist of rodents, birds, and fruit. In the Kanha Tiger Reserve, 80% of its diet consists of rodents, reptiles and fruit. Vegetable matter forms part of the jackal diet, and in India they feed intensively on the fruits of Buckthorn (Ziziphus), Dogbane (Carissa carvanda), Java plum (Syzygium cumini), and the pods of mesquite (Prosopis juliflora) and the golden rain tree Cassia fistula.[7]

... Although the species of the wolf approaches very near to that of the dog, yet the jackal seems to be placed between them; to the savage fierceness of the wolf it adds the impudent familiarity of the dog ... It is more noisy in its pursuits even than the dog, and more voracious than the wolf.

In the Caucasus and Transcaucasia, golden jackals primarily hunt hares and mouse-like rodents, and also pheasants, francolins, ducks, coots, moorhens, and passerines. Vegetable matter eaten by jackals in these areas includes fruits, such as pears, hawthorn, dogwood, and the cones of common medlars. The jackal is implicated in the destruction of grape, watermelon, muskmelon, and nut crops. Near the Vakhsh River, their spring diet consists almost exclusively of plant bulbs and the roots of wild sugar cane, while during winter they feed on wild stony olives. Around the edges of the Karakum Desert, jackals feed on gerbils, lizards, snakes, fish, muskrats, the fruits of wild stony olives, mulberry, dried apricots, watermelons, muskmelons, tomatoes, and grapes.[68]

Indian jackal (C. a. indicus) feeding on chital carcass, Pench National Park

In Dalmatia, the golden jackal's diet consists of mammals, fruits, vegetables, insects, birds and their eggs, grasses and leaves.[69] Jackals change their diet to more readily available foods. In Serbia, their diet is primarily livestock carcasses that are increasingly prevalent due to a lack of removal, and this may have led to the expansion of their population.[70] In Hungary, 55% of their diet is composed of common voles and bank voles, and 41% is composed of wild boar carcasses.[71] Information on the diet of the golden jackal in northeastern Italy is scant, but it is known to prey on small roe deer and hares.[21] In Israel, golden jackals are significant predators of snakes; during a poisoning campaign against golden jackals there was an increase in human snakebite reports, but a decrease when the poisoning ceased.[72]

Competition

Painting of golden jackals and striped hyenas at a kill

In the past, the tiger and the leopard were enemies of the jackal, which fed on the remains of their kills and in one case on a dead tiger. Leopards once hunted jackals; however, today the tiger is extinct in the jackal's range, and the leopard is rare. The jackal's competitors are the red fox, wolf, jungle cat, forest wildcat, and raccoon in the Caucasus, and the steppe wildcat in Central Asia.[57] Wolves dominate jackals, and jackals dominate foxes. In Europe, the range of wolves and jackals is mutually exclusive, with jackals abandoning their territory with the arrival of a wolf pack. One experiment used loudspeakers to broadcast the calls of jackals, and this attracted wolves at a trotting pace to chase away the perceived competitors. Dogs responded to these calls in the same way while barking aggressively. Unleashed dogs have been observed to immediately chase away jackals when the jackals were detected.[44] The jackal's recent expansion throughout eastern and western Europe has been attributed to the extermination of the local wolf populations. The present diffusion of the jackal into the northern Adriatic hinterland is in areas where the wolf is absent or very rare.[59][73]

Foxes can be found only at the fringes of jackal territory.[44] Red foxes and golden jackals share similar diets, with the jackal being three times bigger than the red fox. The red fox will avoid close physical proximity with jackals, and fox populations decrease where jackals are abundant.[74] Striped hyenas prey on golden jackals, and three jackal carcasses were found in one hyena den.[7]

Distribution and legal status

Range map of golden jackal subspecies

In South Asia the golden jackal inhabits Afghanistan,[8] Bangladesh,[8] Bhutan,[7] India,[7] Nepal,[7] Pakistan,[8] and Sri Lanka.[8] In Central Asia it inhabits Tadjikistan, Turkmenistan, and Uzbekistan.[8] In Southeast Asia it inhabits Myanmar and Thailand.[8] Although there have been two reported sightings from Cambodia, three from southern Laos, and two from Vietnam – each sighting made only in lowland, open deciduous forest – no specimens were presented.[75] In Southwestern Asia it inhabits Iran,[8] Iraq,[8] Israel,[8] Jordan,[8] Kuwait,[7] Lebanon,[8] Oman,[8] Saudi Arabia,[8] Qatar,[7] Syria,[8] Turkey,[8] United Arab Emirates,[8] and Yemen.[8] In Europe it inhabits Albania,[8] Armenia,[76] Austria,[7] Azerbaijan,[76] the Baltic states,[73] Bosnia and Herzegovina,[7] Bulgaria,[7] Croatia,[8] Georgia,[76] Greece,[8] Hungary,[73] Italy,[8] Kosovo,[76] Macedonia,[8] Moldova,[76] Montenegro,[76] Poland,[73] Romania,[73] the Russian Federation,[76] Serbia,[73] Slovakia,[77] Slovenia,[8] Switzerland,[73] Turkey,[8] and the Ukraine.[73] It has been sighted in Belarus,[76] the Czech Republic,[78] and Germany,[73] reported in the media in Denmark,[79][80] and reported in the media in the Netherlands, but it is unclear if this jackal was an escapee from a private zoo.[81]

In Europe golden jackals are not listed under the 1973 Convention on International Trade in Endangered Species of Wild Fauna and Flora nor the 1979 Convention on the Conservation of Migratory Species of Wild Animals. However, golden jackals in Europe fall under a number of international legal instruments. These include the 1979 Berne Convention on the Conservation of European Wildlife and Natural Habitats, the 1992 Convention on Biological Diversity, and the 1992 European Union Council Directive 92/43/EEC on the Conservation of Natural Habitats and of Wild Fauna and Flora. The Council Directive provides both guidance and limits on what participating governments can do when responding to the arrival of expanding jackals. These legislative instruments aim to contribute to conserving native wildlife; however, some governments argue that the golden jackal is not native wildlife but an invading species.[76]

The known legal status of golden jackals in European countries is: Albania, fully protected; Austria, variable (depending on the part of the country); Belarus, unprotected (with hunting not forbidden); Bosnia and Herzegovina, hunted (within the law); Bulgaria, hunted; Croatia, hunted; Czech Republic, unprotected; Estonia, unprotected; Germany, fully protected; Greece, unprotected; Hungary, hunted; Italy, fully protected; Kosovo, hunted; Latvia, hunted; Lithuania, hunted; Macedonia, fully protected; Moldova, unknown; Montenegro, hunted; Poland, fully protected; Romania, hunted; Serbia, hunted; Slovakia, hunted; Slovenia, hunted; Switzerland, fully protected; Turkey, variable; and the Ukraine, hunted.[76]

In India, the golden jackal occurs in all of India's protected areas apart from those in the higher areas of the Himalayas. It is included in CITES Appendix III, and is listed in the Wildlife Protection Act, 1972, under Schedule III, thus receiving legal protection at the lowest level to help control the trade of pelts and tails in India.[1]

Diseases and parasites

Adult heartworm in the right ventricle of the heart of a golden jackal

Some golden jackals carry diseases and parasites harmful to human health. These include rabies, and Donovan's Leishmania that is harmless to jackals but may cause leishmaniasis in people. Jackals in southwestern Tajikistan can carry up to 16 species of parasitic cestodes (flatworm), roundworms, and acanthocephalans (thorny-headed worms), these being: Sparganum mansoni, Diphyllobothrium mansonoides, Taenia hydatigena, T. pisiformis, Taenia ovis, Hydatigera taeniaeformis, Dipylidium caninum, Mesocestoides lineatus, Ancylostoma caninum, Uncinaria stenocephala, Dioctophyma renale, Toxocara canis, Toxascaris leonina, Dracunculus medinensis, Filariata and Macracanthorhynchus catulinum. Jackals infected with Dracunculus medinensis can infect bodies of water with their eggs, which cause dracunculiasis in people who drink from them. Jackals may also play a large part in spreading coenurosis in sheep and cattle, and canine distemper in dogs. In Tajikistan, jackals carry at least 12 tick species (which include Ixodes, Rhipicephalus turanicus, R. leporis, R. rossicus, R. sanguineus, R. pumilio, R. schulzei, Hyalomma anatolicum, H. scupense and H. asiaticum), four flea species (Pulex irritans, Xenopsylla nesokiae, Ctenocephanlides canis and C. felis), and one species of louse (Trichodectes canis).[82]

In Iran, some golden jackals carry intestinal worms (helminths)[83] and Echinococcus granulosus.[84] In Israel, some jackals carry intestinal helminths[85] and Leishmania tropica infection.[86] In Romania, a jackal was found to be carrying Trichinella britovi.[87] In northeastern Italy, the jackal is a carrier of the tick species Ixodes ricinus and Dermacentor reticulatus, and the smallest human fluke Metagonimus yokogawai that can be caught from ingesting infected raw fish, which can lead to metagonimiasis.[88] In Hungary, some jackals carry dog heartworm Dirofilaria immitis,[89] and some have provided the first record in Hungary of Trichinella spiralis and the first record in Europe of Echinococcus multilocularis. The jackal is dispersing across Europe through rivers and valleys, bringing parasites into regions where these did not previously exist.[84]

Relationships with humans

In folklore, mythology and literature

Tabaqui (left) torments Father Wolf and his family, as illustrated in the 1895 edition of Rudyard Kipling's The Two Jungle Books.

Golden jackals appear in Indian folklore and in two ancient texts, the Jakatas and the Panchatantra, where they are portrayed as intelligent and wily creatures.[7] One popular Indian saying describes the jackal as "the sharpest among beasts, the crow among birds, and the barber among men". For a person embarking on an early morning journey, hearing a jackal howl was considered to be a sign of impending good fortune, as was seeing a jackal crossing a road from the left side.[90] In Hinduism, the jackal is portrayed as the familiar of several deities with the most common being Chamunda, the emaciated, devouring goddess of the cremation grounds. Another deity associated with jackals is Kali, who inhabits the cremation ground and is surrounded by millions of jackals. According to the Tantrasara scripture, when offered animal flesh, Kali appears in the form of a jackal. The goddess Shivatudi is depicted with a jackal's head. In Buddhist tales, the jackal is regarded as being cunning in a way similar to the fox in European tales.[91] According to the flood myth of the Kamar people in Raipur district, India, the god Mahadeo (Shiva) caused a deluge to dispose of a jackal who had offended him.[92]

In Rudyard Kipling's Mowgli stories collected in The Jungle Book, the character Tabaqui is a jackal despised by the Sioni wolf pack, due to his mock cordiality, his scavenging habits, and his subservience to Shere Khan the tiger.

Livestock, game, and crop predation

The golden jackal can be a harmful pest that attacks domestic animals such as turkeys, lambs, sheep, goats, domestic water buffalo calves, and valuable game species like newborn roe deer, hares, coypu, pheasants, francolins, grey partridges, bustards and waterfowl. It destroys grapes and eats watermelons, muskmelons, and nuts.[93] In Greece, golden jackals are not as damaging to livestock as wolves and red foxes although they can become a serious nuisance to small stock when in great numbers. In southern Bulgaria, over 1,000 attacks on sheep and lambs were recorded between 1982 and 1987, along with some damage to newborn deer in game farms. However, the damage by jackals in Bulgaria was minimal when compared to the livestock losses due to wolves.[44] Approximately 1.5%–1.9% of calves born on the Golan Heights die due to predation, mainly by jackals.[94] The high predation rate by jackals in both Bulgaria and Israel is attributable to the lack of preventative measures in those countries and the availability of food in illegal garbage dumps, leading to jackal population explosions.[44] There are no known attacks on humans in Europe.[28]

Golden jackals are extremely harmful to fur-bearing rodents, such as coypu and muskrats. Coypu can be completely extirpated in shallow water bodies. During the winter of 1948–1949 in the Amu Darya, muskrats constituted 12.3% of jackal faeces contents, and 71% of muskrat houses were destroyed by jackals. Jackals also harm the fur industry by eating muskrats caught in traps or taking skins left out to dry.[93]

Hunting

Hunting Jackals by Samuel Howitt, illustrating a group of jackals rushing to the defence of a fallen packmate

During British rule in India, sportsmen conducted golden jackal hunting on horseback with hounds, with jackal coursing a substitute for the fox hunting of their native England. Although not considered as beautiful as English red foxes, golden jackals were esteemed for their endurance in the chase with one pursuit lasting 3+12 hours. India's weather and terrain added further challenges to jackal hunters that were not present in England: the hounds of India were rarely in the same good condition as English hounds, and although the golden jackal has a strong odour, the terrain of northern India was not good in retaining scent.[95] Also, unlike foxes, jackals sometimes feigned death when caught and could be ferociously protective of their captured packmates.[54]

Jackals were hunted in three ways: with greyhounds, with foxhounds, and with mixed packs. Hunting jackals with greyhounds offered poor sport because greyhounds were too fast for jackals, and mixed packs were too difficult to control.[95] From 1946 in Iraq, British diplomats and Iraqi riders conducted jackal coursing together. They distinguished three types of jackal: the "city scavenger", which was described as being slow and so smelly that dogs did not like to follow them; the "village jack", which was described as being faster, more alert, and less odorous; and the "open-country jack", which was described as being the fastest, cleaner, and providing better sport.[96]

Some indigenous people of India, such as the Kolis and Vaghirs of Gujarat and Rajasthan and the Narikuravas in Tamil Nadu, hunt and eat golden jackals, but the majority of South Asian cultures consider the animal to be unclean. The orthodox dharma texts forbid the eating of jackals because they have five nails.[91] In the area of the former Soviet Union, jackals are not actively hunted and are usually captured only incidentally during the hunting of other animals by means of traps or shooting during drives. In Transcaucasia, jackals are captured with large fishing hooks baited with meat and suspended 75–100 cm (30–39 in) from the ground with wire. The jackals can only reach the meat by jumping, and are then hooked by the lip or jaw.[93]

Fur use

In Russia and the other nations of the former Soviet Union, golden jackals are considered furbearers of low quality because of their sparse, coarse, and monotonously colored fur.[93] Jackal hairs have very little fur fibre; therefore, their pelts have a flat appearance. The jackals of Asia and the Middle East produce the coarsest pelts, though this can be remedied during the dressing process. Elburz in northern Iran produces the softest furs.[97] Jackal skins are not graded to a fur standard, and are made into collars, women's coats, and fur coats. During the 1880s, 200 jackals were captured annually in Mervsk and in the Zakatal area of the Transcaucasus, with 300 jackals being captured there during 1896. In this same period, a total of 10,000 jackals were taken within Russia and their furs sent exclusively to the Nizhegorod fair. In the early 1930s there were 20,000–25,000 jackal skins tanned annually in the Soviet Union, but these could not be utilized within the country, and so the majority were exported to the United States. Commencing from 1949, they were all used within the Soviet Union.[93]

Sulimov dog

European jackal undergoing training at Sheremetyevo Airport, Russia

The golden jackal may have once been tamed in Neolithic Turkey 11,000 years ago, as there is a sculpture of a man cradling a jackal found in Göbekli Tepe.[98] French explorers during the 19th century noted that people in the Levant kept golden jackals in their homes.[99] The Kalmyk people near the Caspian Sea were known to frequently cross their dogs with jackals,[99] and Balkan shepherds once crossed their sheepdogs with jackals.[21]

The Russian military established the Red Star kennels in 1924 to improve the performance of working dogs and to conduct military dog research. The Red Star kennel developed "Laikoid" dogs, which were a cross-breed of Spitz-type Russian Laikas with German Shepherds. By the 1980s, the ability of Russia's bomb and narcotic detection dogs were assessed as being inadequate. Klim Sulimov, a research scientist with the DS Likhachev Scientific Research Institute for Cultural Heritage and Environmental Protection, began cross-breeding dogs with their wild relatives in an attempt to improve their scent-detection abilities. The researchers assumed that during domestication dogs had lost some of their scent-detection ability because they no longer had to detect prey. Sumilov crossed European jackals with Laikas, and also with fox terriers to add trainability and loyalty to the mix. He used the jackal because he believed that it was the wild ancestor of the dog, that it had superior scent-detecting ability, and, because it was smaller with more endurance than the dog, it could be housed outdoors in the Russian climate. Sulimov favored a mix of one quarter jackal and three-quarters dog. Sulimov's program continues today with the use of the hybrid Sulimov dogs at the Sheremetyevo Airport near Moscow by the Russian airline Aeroflot.[100]

The hybrid program has been criticized, with one of Sulimov's colleagues pointing out that in other tests the Laika performed just as well as the jackal hybrids. The assumption that dogs have lost some of their scent-detection ability may be incorrect, in that dogs need to be able to scent-detect and identify the many humans that they come into contact with in their domesticated environment. Another researcher crossed German Shepherds with wolves and claimed that this hybrid had superior scent-detection abilities. The scientific evidence to support the claims of hybrid researchers is minimal, and more research has been called for.[100]

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