Wolf

The English "wolf" stems from the Old English , which is itself derived from the Proto-Germanic . The Proto-Indo-European root is also the source of the Latin word for the animal lupus (from Proto-Italic ). The name "grey wolf" refers to the greyish colour of the species. Since pre-Christian times, Germanic peoples such as the Anglo-Saxons took on wulf as a prefix or suffix in their names. Examples include Wulfhere ("Wolf Army"), Cynewulf ("Royal Wolf"), Cēnwulf ("Bold Wolf"), Wulfheard ("Wolf-hard"), Earnwulf ("Eagle Wolf"), Wulfstān ("Wolf Stone") Æðelwulf ("Noble Wolf"), Wolfhroc ("Wolf-Frock"), Wolfhetan ("Wolf Hide"), Scrutolf ("Garb Wolf"), Wolfgang ("Wolf Gait") and Wolfdregil ("Wolf Runner"). == Taxonomy ==

}} }} In 1758, the Swedish botanist and zoologist Carl Linnaeus published in his Systema Naturae the binomial nomenclature. and under this genus he listed the doglike carnivores including domestic dogs, wolves, and jackals. He classified the domestic dog as Canis familiaris, and the wolf as Canis lupus. Remains from Cripple Creek Sump in Alaska may be considerably older, around 1 million years old, or Beringia. While some sources have suggested that this was a consequence of a population bottleneck, other studies have suggested that this a result of gene flow homogenising ancestry. A 2016 genomic study suggests that Old World and New World wolves split around 12,500 years ago followed by the divergence of the lineage that led to dogs from other Old World wolves around 11,100–12,300 years ago. An extinct Late Pleistocene wolf may have been the ancestor of the dog, with the dog's similarity to the extant wolf being the result of genetic admixture between the two. The dingo, Basenji, Tibetan Mastiff and Chinese indigenous breeds are basal members of the domestic dog clade. The divergence time for wolves in Europe, the Middle East, and Asia is estimated to be fairly recent at around 1,600 years ago. Among New World wolves, the Mexican wolf diverged around 5,400 years ago. Admixture with other canids in the wild animal park at Kadzidłowo, Poland. Left: product of a male wolf and a female spaniel; right: from a female wolf and a male West Siberian Laika In the distant past, there was gene flow between African wolves, golden jackals, and grey wolves. The African wolf is a descendant of a genetically admixed canid of 72% wolf and 28% Ethiopian wolf ancestry. One African wolf from the Egyptian Sinai Peninsula showed admixture with Middle Eastern grey wolves and dogs. There is evidence of gene flow between golden jackals and Middle Eastern wolves, less so with European and Asian wolves, and least with North American wolves. This indicates the golden jackal ancestry found in North American wolves may have occurred before the divergence of the Eurasian and North American wolves. The common ancestor of the coyote and the wolf is admixed with a ghost population of an extinct unidentified canid. This canid was genetically close to the dhole and evolved after the divergence of the African hunting dog from the other canid species. The basal position of the coyote compared to the wolf has been proposed to be due to the coyote retaining more of the mitochondrial genome of this unidentified canid. Similarly, a museum specimen of a wolf from southern China collected in 1963 showed a genome that was 12–14% admixed from this unknown canid. In North America, some coyotes and wolves show varying degrees of past genetic admixture. In more recent times, some male Italian wolves originated from dog ancestry, which indicates female wolves will breed with male dogs in the wild. In the Caucasus Mountains, ten percent of dogs including livestock guardian dogs, are first generation hybrids. Although mating between golden jackals and wolves has never been observed, evidence of jackal-wolf hybridization was discovered through mitochondrial DNA analysis of jackals living in the Caucasus Mountains and in Bulgaria. In 2021, a genetic study found that the dog's similarity to the extant grey wolf was the result of substantial dog-into-wolf gene flow, with little evidence of the reverse. == Description ==

The wolf is the largest extant member of the family Canidae, The ears are relatively small and triangular. The wolf's head is large and heavy, with a wide forehead, strong jaws and a long, blunt muzzle. The skull is in length and in width. The teeth are heavy and large, making them better suited to crushing bone than those of other canids, though they are not as specialized as those found in hyenas. Its molars have a flat chewing surface, but not to the same extent as the coyote, whose diet contains more vegetable matter. Females tend to have narrower muzzles and foreheads, thinner necks, slightly shorter legs, and less massive shoulders than males. , Italy Adult wolves measure in length and at shoulder height. The tail measures in length, the ears in height, and the hind feet are . The size and weight of the modern wolf increases proportionally with latitude in accordance with Bergmann's rule. The mean body mass of the wolf is , the smallest specimen recorded at and the largest at . On average, European wolves weigh , North American wolves , and Indian and Arabian wolves . Females in any given wolf population typically weigh less than males. Wolves weighing over are uncommon, though exceptionally large individuals have been recorded in Alaska and Canada. In central Russia, exceptionally large males can reach a weight of . Pelage Zoo, France The wolf has very dense and fluffy winter fur, with a short undercoat and long, coarse guard hairs. Most of the undercoat and some guard hairs are shed in spring and grow back in autumn. The longest hairs occur on the back, particularly on the front quarters and neck. Especially long hairs grow on the shoulders and almost form a crest on the upper part of the neck. The hairs on the cheeks are elongated and form tufts. The ears are covered in short hairs and project from the fur. Short, elastic and closely adjacent hairs are present on the limbs from the elbows down to the calcaneal tendons. The winter fur is highly resistant to the cold. Wolves in northern climates can rest comfortably in open areas at by placing their muzzles between the rear legs and covering their faces with their tail. Wolf fur provides better insulation than dog fur and does not collect ice when warm breath is condensed against it. In cold climates, the wolf can reduce the flow of blood near its skin to conserve body heat. The warmth of the foot pads is regulated independently from the rest of the body and is maintained at just above tissue-freezing point where the pads come in contact with ice and snow. In warm climates, the fur is coarser and scarcer than in northern wolves. Female wolves tend to have smoother furred limbs than males and generally develop the smoothest overall coats as they age. Older wolves generally have more white hairs on the tip of the tail, along the nose, and on the forehead. Winter fur is retained longest by lactating females, although with some hair loss around their teats. Hair length on the middle of the back is , and the guard hairs on the shoulders generally do not exceed , but can reach . A wolf's coat colour is determined by its guard hairs. Wolves usually have some hairs that are white, brown, grey and black. The coat of the Eurasian wolf is a mixture of ochreous (yellow to orange) and rusty ochreous (orange/red/brown) colours with light grey. The muzzle is pale ochreous grey, and the area of the lips, cheeks, chin, and throat is white. The top of the head, forehead, under and between the eyes, and between the eyes and ears is grey with a reddish film. The neck is ochreous. Long, black tips on the hairs along the back form a broad stripe, with black hair tips on the shoulders, upper chest and rear of the body. The sides of the body, tail, and outer limbs are a pale dirty ochreous colour, while the inner sides of the limbs, belly, and groin are white. Apart from those wolves which are pure white or black, these tones vary little across geographical areas, although the patterns of these colours vary between individuals. In North America, the coat colours of wolves follow Gloger's rule, wolves in the Canadian arctic being white and those in southern Canada, the U.S., and Mexico being predominantly grey. In some areas of the Rocky Mountains of Alberta and British Columbia, the coat colour is predominantly black, some being blue-grey and some with silver and black. Differences in coat colour between sexes is absent in Eurasia; females tend to have redder tones in North America. Black-coloured wolves in North America acquired their colour from wolf-dog admixture after the first arrival of dogs across the Bering Strait 12,000 to 14,000 years ago. Research into the inheritance of white colour from dogs into wolves has yet to be undertaken. == Ecology ==

Distribution and habitat in a mountainous habitat in the Apennines in Sassoferrato, Italy Wolves occur across Eurasia and North America. However, deliberate human persecution because of livestock predation and fear of attacks on humans has reduced the wolf's range to about one-third of its historic range; the wolf is now extirpated (locally extinct) from much of its range in Western Europe, the United States and Mexico, and completely in the British Isles and Japan. In modern times, the wolf occurs mostly in wilderness and remote areas. The wolf can be found between sea level and . Wolves live in forests, inland wetlands, shrublands, grasslands (including Arctic tundra), pastures, deserts, and rocky peaks on mountains. Wolves can digest their meal in a few hours and can feed several times in one day, making quick use of large quantities of meat. A well-fed wolf stores fat under the skin, around the heart, intestines, kidneys, and bone marrow, particularly during the autumn and winter. Nonetheless, wolves are not limited to large prey. Smaller-sized animals that may supplement their diet include rodents, hares, insectivores and smaller carnivores. They frequently eat waterfowl and their eggs. When such foods are insufficient, they prey on lizards, snakes, and frogs, when available, and have even been known to feed on grasshoppers. Wolves in some areas may consume fish and even marine life. Wolves also consume some plant material. In Europe, they eat apples, pears, figs, melons, berries and cherries. In North America, wolves eat blueberries and raspberries. They also eat grass, which may provide some vitamins, but is most likely used mainly to induce vomiting to rid themselves of intestinal parasites or long guard hairs. They are known to eat the berries of mountain-ash, lily of the valley, bilberries, cowberries, European black nightshade, grain crops, and the shoots of reeds. In times of scarcity, wolves will readily eat carrion. In Eurasian areas with dense human activity, many wolf populations are forced to subsist largely on livestock and garbage. As prey in North America continue to occupy suitable habitats with low human density, North American wolves eat livestock and garbage only in dire circumstances. Cannibalism is not uncommon in wolves during harsh winters, when packs often attack weak or injured wolves and may eat the bodies of dead pack members. Interactions with other predators Wolves typically dominate other canid species in areas where they both occur. In North America, incidents of wolves killing coyotes are common, particularly in winter, when coyotes feed on wolf kills. Wolves may attack coyote den sites, digging out and killing their pups, though rarely eating them. There are no records of coyotes killing wolves, though coyotes may chase wolves if they outnumber them. According to a press release by the U.S. Department of Agriculture in 1921, the infamous Custer Wolf relied on coyotes to accompany him and warn him of danger. Though they fed from his kills, he never allowed them to approach him. Interactions have been observed in Eurasia between wolves and golden jackals, the latter's numbers being comparatively small in areas with high wolf densities. Wolves also kill red, Arctic and corsac foxes, usually in disputes over carcasses, sometimes eating them. Brown bears typically dominate wolf packs in disputes over carcasses, while wolf packs mostly prevail against bears when defending their den sites. Both species kill each other's young. Wolves eat the brown bears they kill, while brown bears seem to eat only young wolves. Wolf interactions with American black bears are much rarer because of differences in habitat preferences. Wolves have been recorded on numerous occasions actively seeking out American black bears in their dens and killing them without eating them. Unlike brown bears, American black bears frequently lose against wolves in disputes over kills. Wolves also dominate and sometimes kill wolverines, and will chase off those that attempt to scavenge from their kills. Wolverines escape from wolves in caves or up trees. Wolves may interact and compete with felids, such as the Eurasian lynx, which may feed on smaller prey where wolves are present and may be suppressed by large wolf populations. Wolves encounter cougars along portions of the Rocky Mountains and adjacent mountain ranges. Wolves and cougars typically avoid encountering each other by hunting at different elevations for different prey (niche partitioning). This is more difficult during winter. Wolves in packs usually dominate cougars and can steal their kills or even kill them, while one-to-one encounters tend to be dominated by the cat, who likewise will kill wolves. Wolves more broadly affect cougar population dynamics and distribution by dominating territory and prey opportunities and disrupting the feline's behaviour. Wolf and Siberian tiger interactions are well-documented in the Russian Far East, where tigers significantly depress wolf numbers, sometimes to the point of localized extinction. In Israel, Palestine, Central Asia and India wolves may encounter striped hyenas, usually in disputes over carcasses. Striped hyenas feed extensively on wolf-killed carcasses in areas where the two species interact. One-to-one, hyenas dominate wolves, and may prey on them, but wolf packs can drive off single or outnumbered hyenas. There is at least one case in Israel of a hyena associating and cooperating with a wolf pack. Infections Viral diseases carried by wolves include: rabies, canine distemper, canine parvovirus, infectious canine hepatitis, papillomatosis, and canine coronavirus. In wolves, the incubation period for rabies is eight to 21 days, and results in the host becoming agitated, deserting its pack, and travelling up to a day, thus increasing the risk of infecting other wolves. Although canine distemper is lethal in dogs, it has not been recorded to kill wolves, except in Canada and Alaska. The canine parvovirus, which causes death by dehydration, electrolyte imbalance, and endotoxic shock or sepsis, is largely survivable in wolves, but can be lethal to pups. Bacterial diseases carried by wolves include: brucellosis, Lyme disease, leptospirosis, tularemia, bovine tuberculosis, listeriosis and anthrax. Although lyme disease can debilitate individual wolves, it does not appear to significantly affect wolf populations. Leptospirosis can be contracted through contact with infected prey or urine, and can cause fever, anorexia, vomiting, anemia, hematuria, icterus, and death. Wolves are often infested with a variety of arthropod exoparasites, including fleas, ticks, lice, and mites. The most harmful to wolves, particularly pups, is the mange mite (Sarcoptes scabiei), though they rarely develop full-blown mange, unlike foxes. Endoparasites known to infect wolves include: protozoans and helminths (flukes, tapeworms, roundworms and thorny-headed worms). Most fluke species reside in the wolf's intestines. Tapeworms are commonly found in wolves, which they get though their prey, and generally cause little harm in wolves, though this depends on the number and size of the parasites, and the sensitivity of the host. Symptoms often include constipation, toxic and allergic reactions, irritation of the intestinal mucosa, and malnutrition. Wolves can carry over 30 roundworm species, though most roundworm infections appear benign, depending on the number of worms and the age of the host. == Behaviour ==

Social structure The wolf is a social animal. Its populations consist of packs and lone wolves, most lone wolves being temporarily alone while they disperse from packs to form their own or join another one. The wolf's basic social unit is a mated pair accompanied by their offspring. The average pack size in North America is eight wolves and 5.5 in Europe. == Status and conservation ==

The global wild wolf population in 2003 was estimated at 300,000. Wolf population declines have been arrested since the 1970s. This has fostered recolonization and reintroduction in parts of its former range as a result of legal protection, changes in land use, and rural human population shifts to cities. Competition with humans for livestock and game species, concerns over the danger posed by wolves to people, and habitat fragmentation pose a continued threat to the wolf. Despite these threats, the IUCN classifies the wolf as Least Concern on its Red List due to its relatively widespread range and stable population. The species is listed under AppendixII of the Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES), meaning international trade in the species (including parts and derivatives) is regulated. However, populations of Bhutan, India, Nepal and Pakistan are listed in AppendixI which prohibits commercial international trade in wild-sourced specimens. Wolves have been persecuted in Europe for centuries, having been exterminated in Great Britain by 1684, in Ireland by 1770, in Central Europe by 1899, in France by the 1930s, and in much of Scandinavia by the early 1970s. They continued to survive in parts of Finland, Eastern Europe and Southern Europe. Since 1980, European wolves have rebounded and expanded into parts of their former range. The decline of the traditional pastoral and rural economies seems to have ended the need to exterminate the wolf in parts of Europe. As of 2016, estimates of wolf numbers include: 4,000 in the Balkans, 3,460–3,849 in the Carpathian Mountains, 1,700–2,240 in the Baltic states, 1,100–2,400 in the Italian Peninsula, and around 2,500 in the northwest Iberian peninsula as of 2007. In the former Soviet Union, wolf populations have retained much of their historical range despite Soviet-era large scale extermination campaigns. Their numbers range from 1,500 in Georgia, to 20,000 in Kazakhstan and up to 45,000 in Russia. Wolf populations in Pakistan have suffered population declines and range contraction. They are now confined to remote, barren, mountainous regions and extensive deserts. Numerous factors are thought to be responsible for their decline. The expansion of agricultural practices and land conversion has caused habitat loss. The wolf has been protected in India since 1972. The Indian wolf is distributed across the states of Gujarat, Rajasthan, Haryana, Uttar Pradesh, Madhya Pradesh, Maharashtra, Karnataka and Andhra Pradesh. As of 2019, it is estimated that there are around 2,000–3,000 Indian wolves in the country. In East Asia, Mongolia's population numbers 10,000–20,000. In China, Heilongjiang has roughly 650 wolves, Xinjiang has 10,000 and Tibet has 2,000. 2017 evidence suggests that wolves range across all of mainland China. Wolves have been historically persecuted in China but have been legally protected since 1998. The last Japanese wolf was captured and killed in 1905. == Relationships with humans ==

In culture In folklore, religion and mythology '', sculpture of the mythical she-wolf feeding the twins Romulus and Remus, from the legend of the founding of Rome, Italy, 13th century AD. (The twins are a 15th-century addition.) The wolf is a common motif in the mythologies and cosmologies of peoples throughout its historical range. The Ancient Greeks associated wolves with Apollo, the god of light and order. The Ancient Romans connected the wolf with their god of war and agriculture Mars, and believed their city's founders, Romulus and Remus, were suckled by a she-wolf. Norse mythology includes the feared giant wolf Fenrir, In Eurasia, a large part of the diet of some wolf populations consists of livestock, while such incidents are rare in North America, where healthy populations of wild prey have been largely restored. The majority of losses occur during the summer grazing period, untended livestock in remote pastures being the most vulnerable to wolf predation. The most frequently targeted livestock species are sheep (Europe), domestic reindeer (northern Scandinavia), goats (India), horses (Mongolia), cattle and turkeys (North America). The number of animals killed in single attacks varies according to species: most attacks on cattle and horses result in one death, while turkeys, sheep and domestic reindeer may be killed in surplus. Wolves mainly attack livestock when the animals are grazing, though they occasionally break into fenced enclosures. Competition with dogs A review of the studies on the competitive effects of dogs on sympatric carnivores did not mention any research on competition between dogs and wolves. Competition would favour the wolf, which is known to kill dogs; however, wolves usually live in pairs or in small packs in areas with high human persecution, giving them a disadvantage when facing large groups of dogs. Wolves kill dogs on occasion, and some wolf populations rely on dogs as an important food source. In Croatia, wolves kill more dogs than sheep, and wolves in Russia appear to limit stray dog populations. Wolves may display unusually bold behaviour when attacking dogs accompanied by people, sometimes ignoring nearby humans. Wolf attacks on dogs may occur both in house yards and in forests. Wolf attacks on hunting dogs are considered a major problem in Scandinavia and Wisconsin. Although the number of dogs killed each year by wolves is relatively low, it induces a fear of wolves entering villages and farmyards to prey on them. In many cultures, dogs are seen as family members, or at least working team members, and losing one can lead to strong emotional responses such as demanding more liberal hunting regulations. Dogs that are employed to guard sheep help to mitigate human–wolf conflicts, and are often proposed as one of the non-lethal tools in the conservation of wolves. Shepherd dogs are not particularly aggressive, but they can disrupt potential wolf predation by displaying what is to the wolf ambiguous behaviours, such as barking, social greeting, invitation to play or aggression. The historical use of shepherd dogs across Eurasia has been effective against wolf predation, especially when confining sheep in the presence of several livestock guardian dogs. Shepherd dogs are sometimes killed by wolves. Attacks on humans The fear of wolves has been pervasive in many societies, though humans are not part of the wolf's natural prey. How wolves react to humans depends largely on their prior experience with people: wolves lacking any negative experience of humans, or which are food-conditioned, may show little fear of people. Although wolves may react aggressively when provoked, such attacks are mostly limited to quick bites on extremities, and the attacks are not pressed. Predatory attacks may be preceded by a long period of habituation, in which wolves gradually lose their fear of humans. The victims are repeatedly bitten on the head and face, and are then dragged off and consumed unless the wolves are driven off. Such attacks typically occur only locally and do not stop until the wolves involved are eliminated. Predatory attacks can occur at any time of the year, with a peak in the June–August period, when the chances of people entering forested areas (for livestock grazing or berry and mushroom picking) increase. Cases of non-rabid wolf attacks in winter have been recorded in Belarus, Kirov and Irkutsk oblasts, Karelia and Ukraine. Also, wolves with pups experience greater food stresses during this period. The majority of victims of predatory wolf attacks are children under the age of 18 and, in the rare cases where adults are killed, the victims are almost always women. Indian wolves have a history of preying on children, a phenomenon called "child-lifting". They may be taken primarily in the spring and summer periods during the evening hours, and often within human settlements. Cases of rabid wolves are low when compared to other species, as wolves do not serve as primary reservoirs of the disease, but can be infected by animals such as dogs, jackals and foxes. Incidents of rabies in wolves are very rare in North America, though numerous in the eastern Mediterranean, the Middle East and Central Asia. Wolves apparently develop the "furious" phase of rabies to a very high degree. This, coupled with their size and strength, makes rabid wolves perhaps the most dangerous of rabid animals. Bites from rabid wolves are 15 times more dangerous than those of rabid dogs. Rabid wolves usually act alone, travelling large distances and often biting large numbers of people and domestic animals. Most rabid wolf attacks occur in the spring and autumn periods. Unlike with predatory attacks, the victims of rabid wolves are not eaten, and the attacks generally occur only on a single day. The victims are chosen at random, though most cases involve adult men. During the fifty years up to 2002, there were eight fatal attacks in Europe and Russia, and more than two hundred in southern Asia. Human hunting of wolves , Russia Theodore Roosevelt said wolves are difficult to hunt because of their elusiveness, sharp senses, high endurance, and ability to quickly incapacitate and kill hunting dogs. Historic methods included killing of spring-born litters in their dens, coursing with dogs (usually combinations of sighthounds, Bloodhounds and Fox Terriers), poisoning with strychnine, and trapping. A popular method of wolf hunting in Russia involves trapping a pack within a small area by encircling it with fladry poles carrying a human scent. This method relies heavily on the wolf's fear of human scents, though it can lose its effectiveness when wolves become accustomed to the odor. Some hunters can lure wolves by imitating their calls. In Kazakhstan and Mongolia, wolves are traditionally hunted using eagles and large falcons, though this practice is declining, as experienced falconers are becoming few in number. Shooting wolves from aircraft is highly effective, due to increased visibility and direct lines of fire. Several types of dog, including the Borzoi and Kyrgyz Tajgan, have been specifically bred for wolf hunting. Wolves as pets and working animals Wolves and wolf-dog hybrids are sometimes kept as exotic pets, wolves do not show the same tractability as dogs in living alongside humans, being generally less responsive to human commands and more likely to act aggressively. Humans are more likely to be fatally mauled by a pet wolf or wolf-dog hybrid than by a dog. == Notes ==