The wolf (Canis lupus;[b] pl.: wolves), also known as the gray wolf or grey wolf, is a large canine native to Eurasia and North America. More than thirty subspecies of Canis lupus have been recognized, including the dog and dingo, though gray wolves, as popularly understood, only comprise naturally-occurring wild subspecies. The wolf is the largest extant member of the family Canidae, and is further distinguished from other Canis species by its less pointed ears and muzzle, as well as a shorter torso and a longer tail. The wolf is nonetheless related closely enough to smaller Canis species, such as the coyote and the golden jackal, to produce fertile hybrids with them. The wolf's fur is usually mottled white, brown, gray, and black, although subspecies in the arctic region may be nearly all white.
Of all members of the genus Canis, the wolf is most specialized for cooperative game hunting as demonstrated by its physical adaptations to tackling large prey, its more social nature, and its highly advanced expressive behaviour, including individual or group howling. It travels in nuclear families consisting of a mated pair accompanied by their offspring. Offspring may leave to form their own packs on the onset of sexual maturity and in response to competition for food within the pack. Wolves are also territorial, and fights over territory are among the principal causes of mortality. The wolf is mainly a carnivore and feeds on large wild hooved mammals as well as smaller animals, livestock, carrion, and garbage. Single wolves or mated pairs typically have higher success rates in hunting than do large packs. Pathogens and parasites, notably the rabies virus, may infect wolves.
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The global wild wolf population was estimated to be 300,000 in 2003 and is considered to be of Least Concern by the International Union for Conservation of Nature (IUCN). Wolves have a long history of interactions with humans, having been despised and hunted in most pastoral communities because of their attacks on livestock, while conversely being respected in some agrarian and hunter-gatherer societies. Although the fear of wolves exists in many human societies, the majority of recorded attacks on people have been attributed to animals suffering from rabies. Wolf attacks on humans are rare because wolves are relatively few, live away from people, and have developed a fear of humans because of their experiences with hunters, farmers, ranchers, and shepherds.
The English "wolf" stems from the Old English wulf, which is itself thought to be derived from the Proto-Germanic *wulfaz. The Proto-Indo-European root *wkos may also be the source of the Latin word for the animal lupus (*lkos).[5][6] The name "gray wolf" refers to the grayish colour of the species.[7]
In 1758, the Swedish botanist and zoologist Carl Linnaeus published in his Systema Naturae the binomial nomenclature.[4] Canis is the Latin word meaning "dog",[10] 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.[4] Linnaeus considered the dog to be a separate species from the wolf because of its "cauda recurvata" (upturning tail) which is not found in any other canid.[11]
The phylogenetic descent of the extant wolf C. lupus from C. etruscus through C. mosbachensis is widely accepted.[15] The earliest fossils of C. lupus were found in what was once eastern Beringia at Old Crow, Yukon, Canada, and at Cripple Creek Sump, Fairbanks, Alaska. The age is not agreed upon but could date to one million years ago. Considerable morphological diversity existed among wolves by the Late Pleistocene. They had more robust skulls and teeth than modern wolves, often with a shortened snout, a pronounced development of the temporalis muscle, and robust premolars. It is proposed that these features were specialized adaptations for the processing of carcass and bone associated with the hunting and scavenging of Pleistocene megafauna. Compared with modern wolves, some Pleistocene wolves showed an increase in tooth breakage similar to that seen in the extinct dire wolf. This suggests they either often processed carcasses, or that they competed with other carnivores and needed to consume their prey quickly. Compared with those found in the modern spotted hyena, the frequency and location of tooth fractures in these wolves indicates they were habitual bone crackers.[16]
Genomic studies suggest modern wolves and dogs descend from a common ancestral wolf population[17][18][19] that existed 20,000 years ago.[17] A 2021 study found that the Himalayan wolf and the Indian plains wolf are part of a lineage that is basal to other wolves and split from them 200,000 years ago.[20] Other wolves appear to have originated in Beringia in an expansion that was driven by the huge ecological changes during the close of the Late Pleistocene.[21] A study in 2016 indicates that a population bottleneck was followed by a rapid radiation from an ancestral population at a time during, or just after, the Last Glacial Maximum. This implies the original morphologically diverse wolf populations were out-competed and replaced by more modern wolves.[22]
In the distant past, there was gene flow between African wolves, golden jackals, and gray 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 gray wolves and dogs.[24] 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.[25]
In more recent times, some male Italian wolves originated from dog ancestry, which indicates female wolves will breed with male dogs in the wild.[27] In the Caucasus Mountains, ten percent of dogs including livestock guardian dogs, are first generation hybrids.[28] 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[28] and in Bulgaria.[29] In 2021, a genetic study found that the dog's similarity to the extant gray wolf was the result of substantial dog-into-wolf gene flow, with little evidence of the reverse.[30]
A wolf's coat colour is determined by its guard hairs. Wolves usually have some hairs that are white, brown, gray and black.[47] The coat of the Eurasian wolf is a mixture of ochreous (yellow to orange) and rusty ochreous (orange/red/brown) colours with light gray. The muzzle is pale ochreous gray, 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 gray 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.[48]
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 gray. In some areas of the Rocky Mountains of Alberta and British Columbia, the coat colour is predominantly black, some being blue-gray and some with silver and black.[47] Differences in coat colour between sexes is absent in Eurasia;[49] females tend to have redder tones in North America.[50] 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.[51] Research into the inheritance of white colour from dogs into wolves has yet to be undertaken.[52]
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 3,000 m (9,800 ft). Wolves live in forests, inland wetlands, shrublands, grasslands (including Arctic tundra), pastures, deserts, and rocky peaks on mountains.[2] Habitat use by wolves depends on the abundance of prey, snow conditions, livestock densities, road densities, human presence and topography.[39]
In North America, the wolf's diet is dominated by wild large hoofed mammals (ungulates) and medium-sized mammals. In Asia and Europe, their diet is dominated by wild medium-sized hoofed mammals and domestic species. The wolf depends on wild species, and if these are not readily available, as in Asia, the wolf is more reliant on domestic species.[56] Across Eurasia, wolves prey mostly on moose, red deer, roe deer and wild boar.[57] In North America, important range-wide prey are elk, moose, caribou, white-tailed deer and mule deer.[58] Prior to their extirpation from North America, wild horses were among the most frequently consumed prey of North American wolves.[59] 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.[60] A well-fed wolf stores fat under the skin, around the heart, intestines, kidneys, and bone marrow, particularly during the autumn and winter.[61] 152ee80cbc
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