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From Wikipedia, the free encyclopedia

This article is about the wolf within the species Canis lupus. For other species of wolf and other uses,
see Wolf (disambiguation).
"Grey Wolf" and "Gray Wolf" redirect here. For other uses, see Grey Wolf (disambiguation).

Wolf

Temporal range: 
Middle Pleistocene –
present (810,000–0 YBP)[1]

Eurasian wolf (Canis lupus

lupus) at Polar Park in Bardu,
Norway

Conservation status
 Least Concern (IUCN 3.1)[2]

CITES Appendix II (CITES)[3][a]

Scientific classification

Kingdom: Animalia

Phylum: Chordata

Class: Mammalia

Order: Carnivora

Family: Canidae

Genus: Canis

Species: C. lupus

Binomial name

Canis lupus

Linnaeus, 1758[4]

Subspecies

See Subspecies of Canis lupus

Global wolf range based on

 IUCN's 2018 assessment.[2]

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.
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.

Etymology
See also: Wolf (name)
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 *wĺ̥kʷos may also be the source of the Latin word for
the animal lupus (*lúkʷos).[5][6] The name "gray wolf" refers to the grayish colour of the species.[7]
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").[8]

Taxonomy
Canine phylogeny with ages of
divergence

3.50 m 3.06 my 2.74 my 1.92 my 1.62 my

 ya

Cladogram and divergence of the gray

wolf (including the domestic dog) among
its closest extant relatives[9]
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]
Subspecies
Main article: Subspecies of Canis lupus
Further information: Pleistocene wolf
In the third edition of Mammal Species of the World published in 2005, the mammalogist W. Christopher
Wozencraft listed under C. lupus 36 wild subspecies, and proposed two additional
subspecies: familiaris (Linnaeus, 1758) and dingo (Meyer, 1793). Wozencraft included hallstromi—
the New Guinea singing dog—as a taxonomic synonym for the dingo. Wozencraft referred to a
1999 mitochondrial DNA (mtDNA) study as one of the guides in forming his decision, and listed the
38 subspecies of C. lupus under the biological common name of "wolf", the nominate subspecies being
the Eurasian wolf (C. l. lupus) based on the type specimen that Linnaeus studied in Sweden.[12] Studies
using paleogenomic techniques reveal that the modern wolf and the dog are sister taxa, as modern
wolves are not closely related to the population of wolves that was first domesticated.[13] In 2019, a
workshop hosted by the IUCN/Species Survival Commission's Canid Specialist Group considered the New
Guinea singing dog and the dingo to be feral Canis familiaris, and therefore should not be assessed for
the IUCN Red List.[14]
Evolution
Main article: Evolution of the wolf
Further information: Origin of the domestic dog

Life restoration of Canis mosbachensis, the wolf's immediate

ancestor
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]
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.[19] An extinct Late Pleistocene wolf may have been the ancestor of the dog,[23]
[16] with the dog's similarity to the extant wolf being the result of genetic admixture between the two.
[16] 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.[19]
Admixture with other canids
Main article: Canid hybrid

Wolf–dog hybrids 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 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]
The common ancestor of the coyote and the wolf 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 is proposed to be due to the coyote retaining more of the mitochondrial genome of this
unidentified canid.[24] 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.[26] In North America, some
coyotes and wolves show varying degrees of past genetic admixture.[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 almost negligible wolf-into-dog gene flow since the dog's
domestication. Some gray wolves were related to all ancient and modern dogs.[30]

Description
 A North American wolf
The wolf is the largest extant member of the Canidae family,[31] and is further distinguished from
coyotes and jackals by a broader snout, shorter ears, a shorter torso and a longer tail.[32][31] It is
slender and powerfully built, with a large, deeply descending rib cage, a sloping back, and a heavily
muscled neck.[33] The wolf's legs are moderately longer than those of other canids, which enables the
animal to move swiftly, and to overcome the deep snow that covers most of its geographical range in
winter.[34] The ears are relatively small and triangular.[33] The wolf's head is large and heavy, with a
wide forehead, strong jaws and a long, blunt muzzle.[35] The skull is 230–280 mm (9–11 in) in length and
130–150 mm (5–6 in) in width.[36] 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.[37]
[38] Its molars have a flat chewing surface, but not to the same extent as the coyote, whose diet
contains more vegetable matter.[39] Females tend to have narrower muzzles and foreheads, thinner
necks, slightly shorter legs, and less massive shoulders than males.[40]

A wolf skeleton housed in the Wolf Museum, Abruzzo National

Park, Italy
Adult wolves measure 105–160 cm (41–63 in) in length and 80–85 cm (31–33 in) at shoulder height.
[35] The tail measures 29–50 cm (11–20 in) in length, the ears 90–110 mm (3+1⁄2–4+3⁄8 in) in height, and
the hind feet are 220–250 mm (8+5⁄8–9+7⁄8 in).[41] The size and weight of the modern wolf increases
proportionally with latitude in accord with Bergmann's rule.[42] The mean body mass of the wolf is 40 kg
(88 lb), the smallest specimen recorded at 12 kg (26 lb) and the largest at 79.4 kg (175 lb).[43][35] On
average, European wolves weigh 38.5 kg (85 lb), North American wolves 36 kg (79 lb), and Indian
and Arabian wolves 25 kg (55 lb).[44] Females in any given wolf population typically weigh 2.3–4.5 kg (5–
10 lb) less than males. Wolves weighing over 54 kg (119 lb) are uncommon, though exceptionally large
individuals have been recorded in Alaska and Canada.[45] In central Russia, exceptionally large males can
reach a weight of 69–79 kg (152–174 lb).[41]
Pelage
 Wolf in Spiti Valley, northern India
The wolf has very dense and fluffy winter fur, with a short undercoat and long, coarse guard hairs.
[35] Most of the undercoat and some guard hairs are shed in spring and grow back in autumn.[44] 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.[35] The
winter fur is highly resistant to the cold. Wolves in northern climates can rest comfortably in open areas
at −40 °C (−40 °F) 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.[44]
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.[46] In warm climates, the
fur is coarser and scarcer than in northern wolves.[35] 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.[40] Hair length on the
middle of the back is 60–70 mm (2+3⁄8–2+3⁄4 in), and the guard hairs on the shoulders generally do not
exceed 90 mm (3+1⁄2 in), but can reach 110–130 mm (4+3⁄8–5+1⁄8 in).[35]

Wolves in the La Boissière-du-Doré Zoo, France

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]

Ecology
Distribution and habitat
Main article: Wolf distribution

An Italian wolf 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 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]
Diet

A wolf carrying a caribou hindquarter, Denali National Park, Alaska

Like all land mammals that are pack hunters, the wolf feeds predominantly on wild herbivorous hoofed
mammals that can be divided into large size 240–650 kg (530–1,430 lb) and medium size 23–130 kg (51–
287 lb), and have a body mass similar to that of the combined mass of the pack members.[53][54] The
wolf specializes in preying on the vulnerable individuals of large prey,[39] with a pack of 15 able to bring
down an adult moose.[55] The variation in diet between wolves living on different continents is based on
the variety of hoofed mammals and of available smaller and domesticated prey.[56]
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] 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.[59] A well-fed wolf stores fat under the skin,
around the heart, intestines, kidneys, and bone marrow, particularly during the autumn and winter.[60]
Nonetheless, wolves are not fussy eaters. 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, frogs, and large insects when
available.[61] Wolves in some areas may consume fish and even marine life.[62][63][64] 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.[65] 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.[61]
In times of scarcity, wolves will readily eat carrion.[61] In Eurasian areas with dense human activity,
many wolf populations are forced to subsist largely on livestock and garbage.[57] 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.[66] 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.[61][67][68]
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.
[69] 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.[70] Interactions have been observed in Eurasia between wolves
and golden jackals, the latter's numbers being comparatively small in areas with high wolf densities.[35]
[69][71] Wolves also kill red, Arctic and corsac foxes, usually in disputes over carcasses, sometimes eating
them.[35][72]
 A wolf, a bear, coyotes and ravens compete
over a kill
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.[73] 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.[74] 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.[75]
Wolves may interact and compete with felids, such as the Eurasian lynx, which may feed on smaller prey
where wolves are present[76] and may be suppressed by large wolf populations.[77] 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). In winter, when snow accumulation forces their prey into valleys, interactions
between the two species become more likely. Wolves in packs usually dominate cougars and can steal
their kills or even kill them,[78] while one-to-one encounters tend to be dominated by the cat. There are
several documented cases of cougars killing wolves.[79] Wolves more broadly affect cougar population
dynamics and distribution by dominating territory and prey opportunities and disrupting the feline's
behaviour.[80] 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. Only human
depletion of tiger numbers appears to protect wolves from competitive exclusion from them. With
perhaps only four proven records of tigers killing wolves, these cases are rare; attacks appear to be
competitive rather than predatory in nature.[81][76]
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,[82] but wolf packs can drive off
single or outnumbered hyenas.[83][84] There is at least one case in Israel of a hyena associating and
cooperating with a wolf pack. It is proposed that the hyena could benefit from the wolves' superior
ability to hunt large, agile prey. The wolves could benefit from the hyena's superior sense of smell, to
locate and dig out tortoises, to crack open large bones, and to tear open discarded food containers like
tin cans.[85]

Behaviour
See also: Dog behaviour
Social structure
See also: Pack (canine) § Pack behavior in grey wolves
 Indian wolves at the Mysore Zoo
The wolf is a social animal.[35] 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.[86] The wolf's
basic social unit is the nuclear family consisting of a mated pair accompanied by their offspring.[35] The
average pack size in North America is eight wolves and in Europe 5.5 wolves.[42] The average pack
across Eurasia consists of a family of eight wolves (two adults, juveniles, and yearlings),[35] or
sometimes two or three such families,[39] with examples of exceptionally large packs consisting of up to
42 wolves being known.[87] Cortisol levels in wolves rise significantly when a pack member dies,
indicating the presence of stress.[88] During times of prey abundance caused by calving or migration,
different wolf packs may join together temporarily.[35]
Offspring typically stay in the pack for 10–54 months before dispersing.[89] Triggers for dispersal include
the onset of sexual maturity and competition within the pack for food.[90] The distance travelled by
dispersing wolves varies widely; some stay in the vicinity of the parental group, while other individuals
may travel great distances of upwards of 206 km (128 mi), 390 km (240 mi), and 670 km (420 mi) from
their natal (birth) packs.[91] A new pack is usually founded by an unrelated dispersing male and female,
travelling together in search of an area devoid of other hostile packs.[92] Wolf packs rarely adopt other
wolves into their fold and typically kill them. In the rare cases where other wolves are adopted, the
adoptee is almost invariably an immature animal of one to three years old, and unlikely to compete for
breeding rights with the mated pair. This usually occurs between the months of February and May.
Adoptee males may mate with an available pack female and then form their own pack. In some cases, a
lone wolf is adopted into a pack to replace a deceased breeder.[87]
Wolves are territorial and generally establish territories far larger than they require to survive assuring a
steady supply of prey. Territory size depends largely on the amount of prey available and the age of the
pack's pups. They tend to increase in size in areas with low prey populations,[93] or when the pups reach
the age of six months when they have the same nutritional needs as adults.[94] Wolf packs travel
constantly in search of prey, covering roughly 9% of their territory per day, on average 25 km/d
(16 mi/d). The core of their territory is on average 35 km2 (14 sq mi) where they spend 50% of their
time.[93] Prey density tends to be much higher on the territory's periphery. Except out of desperation,
wolves tend to avoid hunting on the fringes of their range to avoid fatal confrontations with
neighbouring packs.[95] The smallest territory on record was held by a pack of six wolves in northeastern
Minnesota, which occupied an estimated 33 km2 (13 sq mi), while the largest was held by an Alaskan
pack of ten wolves encompassing 6,272 km2 (2,422 sq mi).[94] Wolf packs are typically settled, and
usually leave their accustomed ranges only during severe food shortages.[35] Territorial fights are among
the principal causes of wolf mortality, one study concluding that 14–65% of wolf deaths in Minnesota
and the Denali National Park and Preserve were due to other wolves.[96]
Communication
Main article: Wolf communication
Wolves howling
0:29

Rallying cry
0:19

Problems playing these files? See media help.

Wolves communicate using vocalizations, body postures, scent, touch, and taste.[97] The phases of the
moon have no effect on wolf vocalization, and despite popular belief, wolves do not howl at the moon.
[98] Wolves howl to assemble the pack usually before and after hunts, to pass on an alarm particularly at
a den site, to locate each other during a storm, while crossing unfamiliar territory, and to communicate
across great distances.[99] Wolf howls can under certain conditions be heard over areas of up to
130 km2 (50 sq mi).[39] Other vocalizations include growls, barks and whines. Wolves do not bark as
loudly or continuously as dogs do in confrontations, rather barking a few times and then retreating from
a perceived danger.[100] Aggressive or self-assertive wolves are characterized by their slow and
deliberate movements, high body posture and raised hackles, while submissive ones carry their bodies
low, flatten their fur, and lower their ears and tail.[101]
Scent marking involves urine, feces, and anal gland scents. This is more effective at advertising territory
than howling and is often used in combination with scratch marks. Wolves increase their rate of scent
marking when they encounter the marks of wolves from other packs. Lone wolves will rarely mark, but
newly bonded pairs will scent mark the most.[39] These marks are generally left every 240 m (260 yd)
throughout the territory on regular travelways and junctions. Such markers can last for two to three
weeks,[94] and are typically placed near rocks, boulders, trees, or the skeletons of large animals.
[35] Raised leg urination is considered to be one of the most important forms of scent communication in
the wolf, making up 60–80% of all scent marks observed.[102]
Reproduction
See also: Canine reproduction

Korean wolves mating in the Tama Zoological Park, Japan

Wolves are monogamous, mated pairs usually remaining together for life. Should one of the pair die,
another mate is found quickly.[103] With wolves in the wild, inbreeding does not occur where
outbreeding is possible.[104] Wolves become mature at the age of two years and sexually mature from
the age of three years.[103] The age of first breeding in wolves depends largely on environmental
factors: when food is plentiful, or when wolf populations are heavily managed, wolves can rear pups at
younger ages to better exploit abundant resources. Females are capable of producing pups every year,
one litter annually being the average.[105] Oestrus and rut begin in the second half of winter and lasts
for two weeks.[103]

Iberian wolf pups stimulating their mother to regurgitate some food

Dens are usually constructed for pups during the summer period. When building dens, females make use
of natural shelters like fissures in rocks, cliffs overhanging riverbanks and holes thickly covered by
vegetation. Sometimes, the den is the appropriated burrow of smaller animals such as foxes, badgers or
marmots. An appropriated den is often widened and partly remade. On rare occasions, female wolves
dig burrows themselves, which are usually small and short with one to three openings. The den is usually
constructed not more than 500 m (550 yd) away from a water source. It typically faces southwards
where it can be better warmed by sunlight exposure, and the snow can thaw more quickly. Resting
places, play areas for the pups, and food remains are commonly found around wolf dens. The odor of
urine and rotting food emanating from the denning area often attracts scavenging birds
like magpies and ravens. Though they mostly avoid areas within human sight, wolves have been known
to nest near domiciles, paved roads and railways.[106] During pregnancy, female wolves remain in a den
located away from the peripheral zone of their territories, where violent encounters with other packs are
less likely to occur.[107]
The gestation period lasts 62–75 days with pups usually being born in the spring months or early
summer in very cold places such as on the tundra. Young females give birth to four to five young, and
older females from six to eight young and up to 14. Their mortality rate is 60–80%.[108] Newborn wolf
pups look similar to German Shepherd Dog pups.[109] They are born blind and deaf and are covered in
short soft grayish-brown fur. They weigh 300–500 g (10+1⁄2–17+3⁄4 oz) at birth and begin to see after nine
to 12 days. The milk canines erupt after one month. Pups first leave the den after three weeks. At one-
and-a-half months of age, they are agile enough to flee from danger. Mother wolves do not leave the
den for the first few weeks, relying on the fathers to provide food for them and their young. Pups begin
to eat solid food at the age of three to four weeks. They have a fast growth rate during their first four
months of life: during this period, a pup's weight can increase nearly 30 times.[108][110] Wolf pups
begin play-fighting at the age of three weeks, though unlike young coyotes and foxes, their bites are
gentle and controlled. Actual fights to establish hierarchy usually occur at five to eight weeks of age. This
is in contrast to young coyotes and foxes, which may begin fighting even before the onset of play
behaviour.[111] By autumn, the pups are mature enough to accompany the adults on hunts for large
prey.[107]
Hunting and feeding
Main article: Hunting behavior of gray wolves
 Wolves pursuing a bull elk
Single wolves or mated pairs typically have higher success rates in hunting than do large packs; single
wolves have occasionally been observed to kill large prey such as moose, bison and muskoxen unaided.
[112][113] This contrasts with the commonly held belief that larger packs benefit from cooperative
hunting to bring down large game.[113] The size of a wolf hunting pack is related to the number of pups
that survived the previous winter, adult survival, and the rate of dispersing wolves leaving the pack. The
optimal pack size for hunting elk is four wolves, and for bison a large pack size is more successful.[114]
Wolves move around their territory when hunting, using the same trails for extended periods. After
snowfalls, wolves find their old trails and continue using them. These follow the banks of rivers, the
shorelines of lakes, ravines overgrown with shrubs, plantations, or roads and human paths.[115] Wolves
are nocturnal predators. During the winter, a pack will commence hunting in the twilight of early evening
and will hunt all night, traveling tens of kilometres. Sometimes hunting large prey occurs during the day.
During the summer, wolves generally tend to hunt individually, ambushing their prey and rarely giving
pursuit.[116]
When hunting large gregarious prey, wolves will try to isolate an individual from its group.[117] If
successful, a wolf pack can bring down game that will feed it for days, but one error in judgement can
lead to serious injury or death. Most large prey have developed defensive adaptations and behaviours.
Wolves have been killed while attempting to bring down bison, elk, moose, muskoxen, and even by one
of their smallest hoofed prey, the white-tailed deer. With smaller prey like beaver, geese, and hares,
there is no risk to the wolf.[118] Although people often believe wolves can easily overcome any of their
prey, their success rate in hunting hoofed prey is usually low.[119]

Two wolves feeding on a white-tailed deer

The wolf must give chase and gain on its fleeing prey, slow it down by biting through thick hair and hide,
and then disable it enough to begin feeding.[118] After chasing and then confronting a large prey animal,
the wolf makes use of its 6 cm (2+1⁄2 in) fangs and its powerful masseter muscles to deliver a bite force of
28 kg/cm2 (400 lbf/in2), which is capable of breaking open the skulls of many of its prey animals. The
wolf leaps at its quarry and tears at it. One wolf was observed being dragged for dozens of metres
attached to the hind leg of a moose; another was seen being dragged over a fallen log while attached to
a bull elk's nose.[114] Wolves may wound large prey and then lie around resting for hours before killing
it when it is weaker due to blood loss, thereby lessening the risk of injury to themselves.[120] With
medium-sized prey, such as roe deer or sheep, wolves kill by biting the throat, severing nerve tracks and
the carotid artery, thus causing the animal to die within a few seconds to a minute. With
small, mouselike prey, wolves leap in a high arc and immobilize it with their forepaws.[121]
Once prey is brought down, wolves begin to feed excitedly, ripping and tugging at the carcass in all
directions, and bolting down large chunks of it.[122] The breeding pair typically monopolizes food to
continue producing pups. When food is scarce, this is done at the expense of other family members,
especially non-pups.[123] The breeding pair typically eats first. They usually work the hardest at killing
prey, and may rest after a long hunt and allow the rest of the family to eat undisturbed. Once the
breeding pair has finished eating, the rest of the family tears off pieces of the carcass and transports
them to secluded areas where they can eat in peace. Wolves typically commence feeding by gorging on
the larger internal organs, like the heart, liver, lungs, and stomach lining. The kidneys and spleen are
eaten once they are exposed, followed by the muscles.[124] A wolf can eat 15–19% of its body weight in
one sitting.[60]

Infections
Viral and bacterial
Footage of a wolf taken from Abruzzo Natural Park showing advanced signs of canine distemper
Viral diseases carried by wolves include: rabies, canine distemper, canine parvovirus, infectious canine
hepatitis, papillomatosis, and canine coronavirus.[125] Wolves are a major host for rabies in Russia,
Iran, Afghanistan, Iraq and India.[126] In wolves, the incubation period is eight to 21 days, and results in
the host becoming agitated, deserting its pack, and travelling up to 80 km (50 mi) a day, thus increasing
the risk of infecting other wolves. Infected wolves do not show any fear of humans, most documented
wolf attacks on people being attributed to rabid animals. 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. Wolves may catch infectious canine hepatitis from dogs, though there
are no records of wolves dying from it. Papillomatosis has been recorded only once in wolves, and likely
does not cause serious illness or death, though it may alter feeding behaviours. The canine coronavirus
has been recorded in Alaskan wolves, infections being most prevalent in winter months.[125]
Bacterial diseases carried by wolves include: brucellosis, Lyme disease, leptospirosis, tularemia, bovine
tuberculosis,[127] listeriosis and anthrax.[126] Wolves can catch Brucella suis from wild and domestic
reindeer. While adult wolves tend not to show any clinical signs, it can severely weaken the pups of
infected females. 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 living
near farms are more vulnerable to the disease than those living in the wilderness, probably because of
prolonged contact with infected domestic animal waste. Wolves may catch tularemia
from lagomorph prey, though its effect on wolves is unknown. Although bovine tuberculosis is not
considered a major threat to wolves, it has been recorded to have killed two wolf pups in Canada.[127]
Parasitic
 In Yellowstone National Park
Wolves carry ectoparasites and endoparasites; those in the former Soviet Union have been recorded to
carry at least 50 species.[126] Most of these parasites infect wolves without adverse effects, though the
effects may become more serious in sick or malnourished specimens.[128] Parasitic infection in wolves is
of particular concern to people. Wolves can spread them to dogs, which in turn can carry the parasites to
humans. In areas where wolves inhabit pastoral areas, the parasites can be spread to livestock.[126]
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),[128] though they
rarely develop full-blown mange, unlike foxes.[35] Lice, such as Trichodectes canis, may cause sickness in
wolves, but rarely death. Ticks of the genus Ixodes can infect wolves with Lyme disease and Rocky
Mountain spotted fever.[128] The tick Dermacentor pictus also infests wolves. Other ectoparasites
include chewing lice, sucking lice and the fleas Pulex irritans and Ctenocephalides canis.[35]
Endoparasites known to infect wolves
include: protozoans and helminths (flukes, tapeworms, roundworms and thorny-headed worms). Of
30,000 protozoan species, only a few have been recorded to infect
wolves: Isospora, Toxoplasma, Sarcocystis, Babesia, and Giardia.[128] Some wolves carry Neospora
caninum, which can be spread to cattle and is correlated with bovine miscarriages.[129] Among flukes,
the most common in North American wolves is Alaria, which infects small rodents and amphibians which
are eaten by wolves. Upon reaching maturity, Alaria migrates to the wolf's intestine, but does little
harm. Metorchis conjunctus, which enters wolves through eating fish, infects the wolf's liver or gall
bladder, causing liver disease, inflammation of the pancreas, and emaciation. Most other fluke species
reside in the wolf's intestine, though Paragonimus westermani lives in the lungs. Tapeworms are
commonly found in wolves, as their primary hosts are ungulates, small mammals, and fish, which wolves
feed upon. Tapeworms 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. Infections by the
tapeworm Echinococcus granulosus in ungulate populations tend to increase in areas with high wolf
densities, as wolves can shed Echinoccocus eggs in their feces onto grazing areas.[128]
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. Ancylostoma caninum attaches itself on the
intestinal wall to feed on the host's blood, and can cause hyperchromic anemia, emaciation, diarrhea,
and possibly death. Toxocara canis, a hookworm known to infect wolf pups in the uterus, can cause
intestinal irritation, bloating, vomiting, and diarrhea. Wolves may catch Dioctophyma renale from minks,
which infects the kidneys, and can grow to lengths of 100 cm (40 in). D. renale causes the complete
destruction of the kidney's functional tissue and can be fatal if both kidneys are infected. Wolves can
tolerate low levels of Dirofilaria immitis for many years without showing any ill effects, though high levels
can kill wolves through cardiac enlargement and congestive hepatopathy. Wolves probably become
infected with Trichinella spiralis by eating infected ungulates. Although T. spiralis is not known to
produce clinical signs in wolves, it can cause emaciation, salivation, and crippling muscle pains in dogs.
Thorny-headed worms rarely infect wolves, though three species have been identified in Russian
wolves: Nicolla skrjabini, Macracanthorhynchus catulinus, and Moniliformis moniliformis.[128]

Status and conservation

Further information: List of gray wolf populations by country
The global wild wolf population in 2003 was estimated at 300,000.[130] 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 Appendix II 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 Appendix I which prohibits commercial international
trade in wild-sourced specimens.[2]
North America

Captive Mexican wolf at Sevilleta National Wildlife Refuge in New

Mexico, as part of reintroduction
In Canada, 50,000–60,000 wolves live in 80% of their historical range, making Canada an important
stronghold for the species.[39] Under Canadian law, First Nations people can hunt wolves without
restrictions, but others must acquire licenses for the hunting and trapping seasons. As many as 4,000
wolves may be harvested in Canada each year.[131] The wolf is a protected species in national parks
under the Canada National Parks Act.[132] In Alaska, 7,000–11,000 wolves are found on 85% of the
state's 1,517,733 km2 (586,000 sq mi) area. Wolves may be hunted or trapped with a license; around
1,200 wolves are harvested annually.[133]
In the contiguous United States, wolf declines were caused by the expansion of agriculture, the
decimation of the wolf's main prey species like the American bison, and extermination campaigns.
[39] Wolves were given protection under the Endangered Species Act (ESA) of 1973, and have since
returned to parts of their former range thanks to both natural recolonizations and reintroductions in
Yellowstone and Idaho.[134] The repopulation of wolves in Midwestern United States has been
concentrated in the Great Lakes states of Minnesota, Wisconsin and Michigan where wolves number
over 4,000 as of 2018.[135] Wolves also occupy much of the northern Rocky Mountains region and the
northwest, with a total population over 3,000 as of the 2020s.[136] In Mexico and parts of the
southwestern United States, the Mexican and U.S. governments collaborated from 1977 to 1980 in
capturing all Mexican wolves remaining in the wild to prevent their extinction and established captive
breeding programs for reintroduction.[137] As of 2023, the reintroduced Mexican wolf population
numbers over 200 individuals.[138]
Eurasia

Europe, excluding Russia, Belarus and Ukraine, has 17,000 wolves in more than 28 countries.[139] In
many countries of the European Union, the wolf is strictly protected under the 1979 Berne Convention
on the Conservation of European Wildlife and Natural Habitats (Appendix II) and the 1992 Council
Directive 92/43/EEC on the Conservation of Natural Habitats and of Wild Fauna and Flora (Annex II
and IV). There is extensive legal protection in many European countries, although there are national
exceptions.[2][140]
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.
[141] 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.[131] 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.[139]
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.[142] In Russia, the wolf is regarded as a pest because of its
attacks on livestock, and wolf management means controlling their numbers by destroying them
throughout the year. Russian history over the past century shows that reduced hunting leads to an
abundance of wolves.[143] The Russian government has continued to pay bounties for wolves and
annual harvests of 20–30% do not appear to significantly affect their numbers.[144]
 A wolf in southern Israel
In the Middle East, only Israel and Oman give wolves explicit legal protection.[145] Israel has protected
its wolves since 1954 and has maintained a moderately sized population of 150 through effective
enforcement of conservation policies. These wolves have moved into neighboring countries.
Approximately 300–600 wolves inhabit the Arabian Peninsula.[146] The wolf also appears to be
widespread in Iran.[147] Turkey has an estimated population of about 7,000 wolves.[148] Outside of
Turkey, wolf populations in the Middle East may total 1,000–2,000.[145]
In southern Asia, the northern regions of Afghanistan and Pakistan are important strongholds for wolves.
The wolf has been protected in India since 1972.[149] The Indian wolf is distributed across the states
of Gujarat, Rajasthan, Haryana, Uttar Pradesh, Madhya Pradesh, Maharashtra, Karnataka and Andhra
Pradesh.[150] As of 2019, it is estimated that there are around 2,000–3,000 Indian wolves in the country.
[151] 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.[152] 2017 evidence suggests that wolves range
across all of mainland China.[153] Wolves have been historically persecuted in China[154] but have been
legally protected since 1998.[155] The last Japanese wolf was captured and killed in 1905.[156]

Relationships with humans

In culture
In folklore, religion and mythology
Main article: Wolves in folklore, religion and mythology
See also: Wolves in heraldry

The Capitoline Wolf, 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.[157] The Ancient
Romans connected the wolf with their god of war and agriculture Mars,[158] and believed their city's
founders, Romulus and Remus, were suckled by a she-wolf.[159] Norse mythology includes the feared
giant wolf Fenrir,[160] and Geri and Freki, Odin's faithful pets.[161]
In Chinese astronomy, the wolf represents Sirius and guards the heavenly gate. In China, the wolf was
traditionally associated with greed and cruelty and wolf epithets were used to describe negative
behaviours such as cruelty ("wolf's heart"), mistrust ("wolf's look") and lechery ("wolf-sex"). In
both Hinduism and Buddhism, the wolf is ridden by gods of protection. In Vedic Hinduism, the wolf is a
symbol of the night and the daytime quail must escape from its jaws. In Tantric Buddhism, wolves are
depicted as inhabitants of graveyards and destroyers of corpses.[160]
In the Pawnee creation myth, the wolf was the first animal brought to Earth. When humans killed it, they
were punished with death, destruction and the loss of immortality.[162] For the Pawnee, Sirius is the
"wolf star" and its disappearance and reappearance signified the wolf moving to and from the spirit
world. Both Pawnee and Blackfoot call the Milky Way the "wolf trail".[163] The wolf is also an
important crest symbol for clans of the Pacific Northwest like the Kwakwakaʼwakw.[160]
The concept of people turning into wolves, and the inverse, has been present in many cultures.
One Greek myth tells of Lycaon being transformed into a wolf by Zeus as punishment for his evil deeds.
[164] The legend of the werewolf has been widespread in European folklore and involves people
willingly turning into wolves to attack and kill others.[165] The Navajo have traditionally believed
that witches would turn into wolves by donning wolf skins and would kill people and raid graveyards.
[166] The Dena'ina believed wolves were once men and viewed them as brothers.[157]
In fable and literature
See also: List of fictional wolves
Aesop featured wolves in several of his fables, playing on the concerns of Ancient Greece's settled,
sheep-herding world. His most famous is the fable of "The Boy Who Cried Wolf", which is directed at
those who knowingly raise false alarms, and from which the idiomatic phrase "to cry wolf" is derived.
Some of his other fables concentrate on maintaining the trust between shepherds and guard dogs in
their vigilance against wolves, as well as anxieties over the close relationship between wolves and dogs.
Although Aesop used wolves to warn, criticize and moralize about human behaviour, his portrayals
added to the wolf's image as a deceitful and dangerous animal. The Bible uses an image of a wolf lying
with a lamb in a utopian vision of the future. In the New Testament, Jesus is said to have used wolves as
illustrations of the dangers his followers, whom he represents as sheep, would face should they follow
him.[167]

Little Red Riding Hood (1883), Gustave Doré

Isengrim the wolf, a character first appearing in the 12th-century Latin poem Ysengrimus, is a major
character in the Reynard Cycle, where he stands for the low nobility, whilst his adversary, Reynard the
fox, represents the peasant hero. Isengrim is forever the victim of Reynard's wit and cruelty, often dying
at the end of each story.[168] The tale of "Little Red Riding Hood", first written in 1697 by Charles
Perrault, is considered to have further contributed to the wolf's negative reputation in the Western
world. The Big Bad Wolf is portrayed as a villain capable of imitating human speech and disguising itself
with human clothing. The character has been interpreted as an allegorical sexual predator.
[169] Villainous wolf characters also appear in The Three Little Pigs and "The Wolf and the Seven Young
Goats".[170] The hunting of wolves, and their attacks on humans and livestock, feature prominently
in Russian literature, and are included in the works of Leo Tolstoy, Anton Chekhov, Nikolay
Nekrasov, Ivan Bunin, Leonid Pavlovich Sabaneyev, and others. Tolstoy's War and Peace and
Chekhov's Peasants both feature scenes in which wolves are hunted with hounds and Borzois.[171] The
musical Peter and the Wolf involves a wolf being captured for eating a duck, but is spared and sent to a
zoo.[172]
Wolves are among the central characters of Rudyard Kipling's The Jungle Book. His portrayal of wolves
has been praised posthumously by wolf biologists for his depiction of them: rather than being villainous
or gluttonous, as was common in wolf portrayals at the time of the book's publication, they are shown as
living in amiable family groups and drawing on the experience of infirm but experienced elder pack
members.[173] Farley Mowat's largely fictional 1963 memoir Never Cry Wolf is widely considered to be
the most popular book on wolves, having been adapted into a Hollywood film and taught in several
schools decades after its publication. Although credited with having changed popular perceptions on
wolves by portraying them as loving, cooperative and noble, it has been criticized for its idealization of
wolves and its factual inaccuracies.[174][175][176]
Conflicts
Human presence appears to stress wolves, as seen by increased cortisol levels in instances such as
snowmobiling near their territory.[177]
Predation on livestock

A 1905 postcard of the Hexham wolf, an escaped wolf shot for

killing livestock in England
Livestock depredation has been one of the primary reasons for hunting wolves and can pose a severe
problem for wolf conservation. As well as causing economic losses, the threat of wolf predation causes
great stress on livestock producers, and no foolproof solution of preventing such attacks short of
exterminating wolves has been found.[178] Some nations help offset economic losses to wolves through
compensation programs or state insurance.[179] Domesticated animals are easy prey for wolves, as they
have been bred under constant human protection, and are thus unable to defend themselves very well.
[180] Wolves typically resort to attacking livestock when wild prey is depleted.[181] 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.[178]
The majority of losses occur during the summer grazing period, untended livestock in remote pastures
being the most vulnerable to wolf predation.[182] The most frequently targeted livestock species are
sheep (Europe), domestic reindeer (northern
Scandinavia), goats (India), horses (Mongolia), cattle and turkeys (North America).[178] 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.[183] Wolves mainly attack
livestock when the animals are grazing, though they occasionally break into fenced enclosures.[184]
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.[185][186] 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.[186][187]
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.[178][188] The most
frequently killed hunting breeds in Scandinavia are Harriers, older animals being most at risk, likely
because they are less timid than younger animals, and react to the presence of wolves differently. Large
hunting dogs such as Swedish Elkhounds are more likely to survive wolf attacks because of their better
ability to defend themselves.[188]
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.[186]
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.[186][189] 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,[186][190] especially
when confining sheep in the presence of several livestock guardian dogs.[186][191] Shepherd dogs are
sometimes killed by wolves.[186]
Attacks on humans
Main articles: Wolf attack and List of wolf attacks
 Country children surprised by a wolf (1833) by François Grenier de
Saint-Martin
The fear of wolves has been pervasive in many societies, though humans are not part of the wolf's
natural prey.[192] 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.[193] Although wolves may react aggressively when provoked, such attacks are mostly limited
to quick bites on extremities, and the attacks are not pressed.[192]
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.[192] 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.[35] 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.[192] 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.[194]
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.[192] Bites from rabid wolves are 15 times more dangerous than those of rabid dogs.
[195] 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.[192]
Human hunting of wolves
Main articles: Wolf hunting and Wolf hunting with dogs
See also: Human uses of hunted wolves
 Carcasses of hunted wolves in Volgograd Oblast, 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.[196] 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.[197][198]
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 with eagles
and 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.
[198] Several types of dog, including the Borzoi and Kyrgyz Tajgan, have been specifically bred for wolf
hunting.[186]
As pets and working animals
Main article: Wolves as pets and working animals
Wolves and wolf-dog hybrids are sometimes kept as exotic pets. Although closely related to domestic
dogs, 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. A person is more likely to be fatally
mauled by a pet wolf or wolf-dog hybrid than by a dog.[199]

Notes

1. ^ Populations of Bhutan, India, Nepal, and Pakistan are included in Appendix I. Excludes
domesticated form and dingo, which are referenced as Canus lupus familiaris and Canus
lupus dingo.
2. ^ Domestic and feral dogs are included in the phylogenetic but not colloquial definition
of 'wolf', and thus not in the scope of this article.

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Bibliography
  Busch, R. H. (2007). Wolf Almanac, New and Revised: A Celebration Of Wolves And Their
World (3 ed.). Rowman & Littlefield. ISBN 978-1-59921-069-8.
 Graves, Will (2007). Wolves in Russia: Anxiety throughout the ages. Detselig
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and Carnivora (Sea cows; Wolves and Bears). Science Publishers, Inc. USA. ISBN 978-1-
886106-81-9.
 Lopez, Barry H. (1978). Of Wolves and Men. J. M. Dent and Sons Limited. ISBN 978-0-7432-
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External links

Wikimedia Commons has media related to Canis lupus.

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 The International Wolf Center

 Staying Safe in Wolf Country, ADFG (January 2009)
 Watch Death of a Legend and Cry of the Wild by Bill Mason

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 Wikidata: Q18498
 Wikispecies: Canis lupus
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