Wolf

The wolf (Canis lupus), also known as the

gray/grey wolf, is a canine native to the
wilderness and remote areas of Eurasia
and North America. It is the largest extant
member of its family, with males
averaging 43–45 kg (95–99 lb) and
females 36–38.5 kg (79–85 lb). It is
distinguished from other Canis species by
its larger size and less pointed features,
particularly on the ears and muzzle. Its
winter fur is long and bushy and
predominantly a mottled gray in color,
although nearly pure white, red and brown
to black also occur. Mammal Species of
the World (3rd ed., 2005), a standard
reference work in zoology, recognises 38
subspecies of C. lupus.
 Wolf
Temporal range: Middle Pleistocene – present
(810,000–0 years BP[1]

Eurasian wolf (Canis lupus lupus).

0:00

Wolf pack howling

Conservation status

Least Concern (IUCN 3.1)[2]

Scientific classification

Kingdom: Animalia
Phylum: Chordata
Class: Mammalia

Order: Carnivora

Family: Canidae

Genus: Canis

Species: C. lupus

Binomial name

Canis lupus
Linnaeus, 1758[3]

Subspecies

refer Subspecies of Canis lupus

 Historical (red + green) and modern (green)
range of wild subspecies of C. lupus. Recent
evidence suggests the wolf range extends
across all of mainland China, [4][5][6] and in the
US down the Cascade Range from British
Columbia through Washington State , Oregon
State , and northern California to Lassen
Peak.

The wolf is the most specialized member

of the genus Canis in the direction of
cooperative big game hunting, as
demonstrated by its morphological
adaptations to tackling large prey, its more
gregarious nature, and its highly advanced
expressive behavior. It is nonetheless
closely related enough to smaller Canis
species, such as the coyote and golden
jackal, to produce fertile hybrids. It is the
only species of Canis to have a range
encompassing both Eurasia and North
America, and originated in Eurasia during
the Pleistocene, colonizing North America
on at least three separate occasions
during the Rancholabrean. It is a social
animal, travelling in nuclear families
consisting of a mated pair, accompanied
by the pair's adult offspring. The wolf is
typically an apex predator throughout its
range. It feeds primarily on large wild
ungulates, though it also eats smaller
animals, livestock, carrion, and garbage. A
seven-year-old wolf is considered to be
relatively old, and the maximum lifespan is
about 16 years.

The global wolf population is estimated to

be 300,000. The wolf is one of the world's
best-known and most-researched animals,
with probably more books written about it
than any other wild species. It has a long
history of association with humans, having
been despised and hunted in most
pastoral communities because of its
attacks on livestock, while conversely
being respected in some agrarian and
hunter-gatherer societies. Although the
fear of wolves is pervasive in many human
societies, the majority of recorded attacks
on people have been attributed to animals
suffering from rabies. Non-rabid wolves
have attacked and killed people, mainly
children, but this is rare, as wolves are
relatively few, live away from people, and
have developed a fear of humans from
hunters and shepherds.

Etymology
The English 'wolf' stems from the Old
English wulf, which is itself thought to be
derived from the Proto-Germanic *wulfaz.
The Latin lupus is a Sabine loanword.[7]
Both derive from the Proto-Indo-European
root *wĺ ̥kʷos or *lúkʷos.[8]

Taxonomy
The species Canis lupus was named by the
Swedish botanist and zoologist Carl
Linnaeus in his publication Systema
Naturae in 1758.[3] The nominate
subspecies is the Eurasian wolf (Canis
lupus lupus), based on the type specimen
that Linnaeus studied in Sweden. The 38
subspecies of Canis lupus are listed under
the biological common name of "wolf" in
Mammal Species of the World (3rd edition)
that was published in 2005. The domestic
dog is listed as a subspecies, and the
dingo, together with the New Guinea
singing dog, form another subspecies.[9]

Evolution

Artist's impression of a Beringian wolf

The earliest fossils of Canis 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 but could date 1 million YBP.
Considerable morphological diversity
existed among grey wolves by the Late
Pleistocene. These are regarded as having
been more cranio-dentally robust than
modern grey wolves, often with a
shortened rostrum, the 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 that is similar to that seen
in the extinct dire wolf. This suggests that
these either often processed carcasses, or
that they competed with other carnivores
and needed to quickly consume their prey.
The frequency and location of tooth
fractures found in these wolves, compared
with the modern spotted hyena, indicates
that these wolves were habitual bone
crackers.[10]

This continuous wolf presence contrasts

with genomic studies, which suggest that
all modern wolves and dogs descend from
a common ancestral wolf
population[11][12][13] that existed as recently
as 20,000 years ago.[11] These studies
indicate 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 that the original wolf populations
were out-competed by a new type of wolf
which replaced them.[14] However, the
geographic origin of this radiation is not
known.

Population structure

The wolf is a highly adaptable species that

is able to exist in a range of environments
and which possesses a wide distribution
across the Holarctic. Studies of modern
wolves have identified distinct sub-
populations that live in close proximity to
each other.[15][16] This variation in sub-
populations is closely linked to differences
in habitat – precipitation, temperature,
vegetation, and prey specialization –
which affect cranio-dental
plasticity.[17][18][19][20]

The wolf population in Europe is divided

along a north-south axis and forms five
major clusters. Three clusters occupy Italy,
the Dinaric-Balkans, and the Carpathians.
Another two clusters occupy north-central
Europe and the Ukrainian steppe. The
Italian wolf consisted of an isolated
population with low genetic diversity.
Wolves from Croatia, Bulgaria, and Greece
form the Dinaric-Balkans cluster. Wolves
from Finland, Latvia, Belarus, Poland and
Russia form the north-central Europe
cluster, with wolves from the Carpathians
cluster coming from a mixture of wolves
from the north-central cluster and the
Dinaric-Balkans cluster. The wolves from
the Carpathians are more similar to the
wolves from the Pontic-Caspian Steppe
than they are to wolves from north-central
Europe. These clusters may be the result
of their expansions from glacial refugia, an
adaptation to local environments, or
landscape fragmentation and the killing of
wolves in some areas by humans.[21]

The wolf population in North American

forms six ecotypes – genetically and
ecologically distinct populations
separated from other populations by their
different type of habitat. These six
ecotypes were named West Forest, Boreal
Forest, Arctic, High Arctic, Baffin, and
British Columbia. The studies found that
precipitation and mean diurnal
temperature range were the most
influential variables on the formation of
these ecotypes.[16][22] The local adaptation
of a wolf ecotype reflects a wolf's
preference to remain in the type of habitat
that it was born into.[16]

Admixture with other canids

Wolf-dog hybrids. The first is the product of a male

wolf and a female spaniel, the second from a female
wolf and a male West Siberian Laika

The wolf-like canids are a group of large

carnivores that are closely related because
their chromosomes number 78,[23][24][25]
and therefore they can potentially
interbreed.[26] In the distant past there has
been gene flow between African golden
wolves, golden jackals, and wolves. The
African golden wolf is a descendant of a
genetically admixed canid of 72% wolf and
28% Ethiopian wolf ancestry. One African
golden wolf from the Egyptian Sinai
Peninsula shows admixture with the
Middle Eastern 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 that 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 has admixed with a ghost population
of an extinct unidentified canid. This canid
is genetically close to the dhole and has
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.[27] In
North America, most coyotes and wolves
show varying degrees of past genetic
admixture.[28] Black-colored wolves in
North America acquired their coloration
from wolf-dog admixture after the first
arrival of dogs.[29]

In more recent times some male Italian

wolves originated from dog ancestry,
which indicates that female wolves will
breed with male dogs in the wild.[30] 10
percent of dogs in the Caucasus
Mountains, including livestock guardian
dogs, are first generation hybrids.[31]
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.[31] and in Bulgaria.[32]
Description

Differences between the wolf and coyote

Wolf skeleton
The wolf is the largest member of the
Canidae family.[33] It is similar to its
closest wild cousins the coyote and the
golden jackal but is distinguished by being
larger and heavier, with a broader snout,
shorter ears, a shorter torso and a longer
tail.[34][33] It is slender and powerfully built
with a large, deeply descending rib cage, a
sloping back, and a heavily muscled
neck.[35] The wolf's legs are moderately
longer than those of other canids, which
enables the animal to move swiftly, and
allows it to overcome the deep snow that
covers most of its geographical range.[36]
The ears are relatively small and
triangular.[35] Females tend to have
narrower muzzles and foreheads, thinner
necks, slightly shorter legs and less
massive shoulders than males.[37]

On average, 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 (3.5–4.3 in) in
height, and the hind feet are 220–250 mm
(8.7–9.8 in).[35] Wolf weight and size
increase proportionally with latitude in
accord with Bergmann's rule.[35] The mean
body mass of the wolf is 40 kg (88 lb), with
the smallest specimen recorded at 12 kg
(26 lb) and the largest at 79.4 kg
(175 lb).[38][35] On average, European
wolves may weigh 38.5 kg (85 lb), North
American wolves 36 kg (79 lb), and Indian
and Arabian wolves 25 kg (55 lb).[39]
Females in any given wolf population
typically weigh 5–10 lb (2.3–4.5 kg) less
than males.[40] Wolves weighing over
54 kg (119 lb) are uncommon, though
exceptionally large individuals have been
recorded in Alaska, Canada,[40] and the
forests of western Russia.[35] The heaviest
wolf to be taken by the United States Fish
and Wildlife Service was killed on 70 Mile
River in east-central Alaska on July 12,
1939 and weighed 79.4 kg (175 lb).[41]
The wolf usually travels at a loping pace,
placing its paws one directly in front of the
other. This gait can be maintained for
hours at a rate of 8–9 km/h (5.0–
5.6 mph),[42] which allows the wolf to
cover great distances. On bare paths, a
wolf can quickly achieve speeds of 50–
60 km/h (31–37 mph). The wolf has a
running gait of 55–70 km/h (34–43 mph),
can leap 5 m (16 ft) horizontally in a single
bound, and can maintain rapid pursuit for
at least 20 minutes.[43]

The wolf usually carries its head at the

same level as the back, raising it only
when alert.[35] Its head is large and heavy,
with a wide forehead, strong jaws and a
long, blunt muzzle.[35] The skull averages
230–280 mm (9.1–11.0 in) in length and
130–150 mm (5.1–5.9 in) in width.[44] The
teeth are heavy and large, which are better
suited to crushing bone than those of
other canids, although not as specialised
as those found in hyenas.[45][46] Its molars
have a flat chewing surface but not to the
same extent as the coyote, whose diet
contains more vegetable matter.[47]

Wolf mandible diagram showing the names and

Wolf mandible diagram showing the names and
positions of the teeth.

Bite force adjusted for body weight in Newtons per kilogram[48]

Canid Carnassial Canine

Wolf 131.6 127.3

Dhole 130.7 132.0

African wild dog 127.7 131.1

Greenland dog and Dingo 117.4 114.3

Coyote 107.2 98.9

Side-striped jackal 93.0 87.5

Golden jackal 89.6 87.7

Black-backed jackal 80.6 78.3

Black and white captive wolves in France

The wolf has very dense and fluffy winter
fur, with short underfur and long, coarse
guard hairs.[35] Most of the underfur and
some of the guard hairs are shed in the
spring and grow back in the autumn
period.[39] The longest hairs occur on the
back, particularly on the front quarters and
neck. Especially long hairs are 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, which
strongly 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, and 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.[39] 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 in the tip
of the tail, along the nose and on the
forehead. The winter fur is retained
longest by lactating females, although
with some hair loss around their
nipples.[37] Hair length on the middle of the
back is 60–70 mm (2.4–2.8 in),and the
guard hairs on the shoulders generally
does not exceed 90 mm (3.5 in) but can
reach 110–130 mm (4.3–5.1 in).[35]

Coat color ranges from almost pure white

through various shades of blond, cream
and ochre to grays, browns and blacks,[49]
with variation in fur color tending to
increase in higher latitudes.[50] Differences
in coat color between sexes are largely
absent,[35] although females may have
redder tones.[51]
Adaptation
Distribution and habitat

Wolf in mountainous habitat in Himachal Pradesh's

Spiti Valley, India

Originally, wolves occurred across Eurasia

above the 12th parallel north and across
North America above the 15th parallel
north. However, deliberate human
persecution has reduced the wolf's range
to about one-third of what it once was
because of livestock predation and fear of
attacks on humans. The wolf is now
extirpated (made locally extinct) in much
of Western Europe, in Mexico and much of
the United States. In modern times, the
wolf occurs mostly in wilderness and
remote areas, particularly in Canada,
Alaska and the northern United States,
Europe and Asia from about 75°N to
12°N.[2] The wolf can be found between
sea level and 3,000 metres (9,800 ft). It
lives in forests, inland wetlands,
shrublands, grasslands including arctic
tundra, pastures, deserts, and rocky peaks
on mountains.[2] Habitat use by wolves is
strongly correlated with the abundance of
prey, snow conditions, absence or low
livestock densities, road densities, human
presence and topography.[47] In cold
climates, the wolf can reduce the flow of
blood near its skin to conserve body heat.
The warmth of the footpads is regulated
independently of the rest of the body, and
is maintained at just above tissue-freezing
point where the pads come in contact with
ice and snow.[52]

Diet
Wolf carrying caribou hindquarter, Denali National
Park, Alaska

Like all land mammals that are pack

hunters, the wolf predominantly feeds on
herbivorous mammals that 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,[47] with a pack of
timber wolves being capable of bringing
down a 500 kg (1,100 lb) moose.[55]
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.[56] A well-fed wolf stores fat
under the skin, around the heart,
intestines, kidneys, and bone marrow,
particularly during the autumn and
winter.[57]

Across their range, wolves predominantly

feed on wild ungulates that can be divided
into large size 240–650 kg (530–1,430 lb)
and medium size 23–130 kg (51–287 lb).
The variation in diet between wolves living
on different continents is based on the
different varieties of ungulate species and
of smaller and domestic prey that are
available. In North America, the wolf's diet
is dominated by wild large ungulates and
medium-sized mammals. In Asia and
Europe, their diet is dominated by wild
medium sized ungulates and domestic
species. The wolf depends on wild
ungulates, and if these are not readily
available, as in Asia, the wolf is more
reliant on domestic species.[58] Across
Eurasia, wolves prey mostly on moose, red
deer, roe deer and wild boar.[59] In North
America, important prey rangewide are elk,
moose, caribou, white-tailed deer and
mule deer.[60]
Although wolves primarily feed on medium
to large sized ungulates, they are not fussy
eaters. Smaller sized animals that may
supplement the diet of wolves 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 as available.[61]
Wolves in northern Minnesota prey on
northern pike in freshwater streams.[62]
The diet of coastal wolves in Alaska
includes 20 percent salmon,[63] while
coastal wolves in British Columbia
includes 25 percent marine sources and
those on the nearby islands 75 percent.[64]
In Europe, wolves eat apples, pears, figs,
mellons, berries and cherries. In North
America, wolves eat blueberries and
raspberries. Wolves also eat grass, which
may provide some vitamins.[65] They are
known to eat the berries of mountain ash,
lily of the valley, bilberries, cowberry,
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.[59] The prey animals of North
American wolves have largely continued to
occupy suitable habitats with low human
density, and cases of wolves subsisting
largely on garbage or livestock are
exceptional.[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]

Behavior

Social behavior
Italian wolf pack resting in shade

The wolf is a social animal, whose basic

social unit consists of a mated pair
accompanied by their adult offspring. The
average pack consists of a family of five to
eleven wolves (two adults, three to six
juveniles, and one to three yearlings),[69] or
sometimes two or three such families,[47]
with exceptionally large packs consisting
of up to forty-two wolves being known.[70]
In ideal conditions, the mated pair
produces pups every year, with such
offspring typically staying in the pack for
10–54 months before dispersing.[71]
Triggers for dispersal include the onset of
sexual maturity and competition within the
pack for food.[72] 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 206 km (128 mi), 390 km
(240 mi), and 670 km (420 mi) from their
natal packs.[43] 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.[73]
Cortisol levels in wolves rise significantly
when a pack member dies, indicating the
presence of stress.[74] 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 of age and unlikely to
compete for breeding rights with the
mated pair. In some cases, a lone wolf is
adopted into a pack to replace a deceased
breeder.[70] During times of ungulate
abundance caused through calving or
migration, different wolf packs may
temporarily join together.[69]
Wolves are highly territorial and generally
establish territories far larger than they
require in order to survive, which assures a
steady supply of prey. Territory size
depends largely on the amount of prey
available and the age of the pack's pups,
tending to increase in size in areas with
low prey populations,[75] or when the pups
reach the age of six months when they
have the same nutritional needs as
adults.[76] Wolf packs travel constantly in
search of prey, covering roughly 9% of their
territory per day (average 25 km/d
(16 mi/d)). The core of their territory is on
average 35 km2 (14 sq mi), in which they
spend 50% of their time.[75] Prey density
tends to be much higher in their territory's
surrounding areas, although wolves tend
to avoid hunting on the fringes of their
range unless desperate because of the
possibility of fatal encounters with
neighboring packs.[77] 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 a 6,272 km2 (2,422 sq mi)
area.[76] Wolf packs are typically settled,
and usually only leave their accustomed
ranges during severe food shortages.[69]
Wolves advertise their territories to other
packs through howling and scent marking.
Scent marking involves urine, feces, and
anal gland scents. Scent marking 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 scent marks of wolves from other
packs. Lone wolves will rarely scent mark,
but newly bonded pairs will scent mark the
most.[47] Scent 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,[76] and are typically placed
near rocks, boulders, trees, or the
skeletons of large animals.[69] Territorial
fights are among the principal causes of
wolf mortality, with one study concluding
that 14–65% of wolf deaths in Minnesota
and the Denali National Park and Preserve
were due to predation by other wolves.[78]

Facial expressions (Konrad Lorenz, 1952). Bottom to

top: increasing fear (ears back); left to right: increasing
aggression (snarl); top right: maximum of both.
Wolves communicate to anticipate what
their packmates or other wolves might do
next.[79] It includes the use of vocalization,
body posture, scent, touch, and taste.[80]
The phases of the moon have no effect on
wolf vocalisation, and despite popular
belief wolves do not howl at the moon.[81]
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 or while
crossing unfamiliar territory, and to
communicate across great distances.[82]
Other vocalisations include growls, barks
and whines.[83] Wolves do not bark as
loudly or continuously as dogs do but they
bark a few times and then retreat from a
perceived danger.[83] 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,
sleeken their fur, and lower their ears and
tail.[84] 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.[85]

Reproduction
Korean wolves mating

The foundation of a wolf pack is the

mated pair. Known uncommon variations
include a mature male and two mature
females, a mature male and his mate
along with his yearling son from a previous
mating, and a mature female with a new
mate and his younger brother. One unusual
pack from Yellowstone was formed by
twelve dispersers from four other packs.
Packs may include offspring from up to
four years of breeding, with one pack
consisting of forty-two wolves.[86] The wolf
is monogamous, with mated pairs usually
remaining together for life. Should one of
the pair die, another mate is quickly found.
Males form a majority in the wolf pack,
with unpaired females being rare.[87] In
favorable conditions, there are sometimes
more than one breeding pair within the
natal pack. This pair may later move into
an area adjacent to, or overlapping with,
the pack territory. Some wolves may leave
the pack but remain in the pack's territory,
waiting for one of the breeding parents to
perish before they can breed. Most foreign
mature wolves are killed by the pack
unless the pack needs to replace a
breeder. Sometimes a foreign wolf will be
adopted and provided with alloparental
care by a pack that already has a breeding
pair, and this usually occurs between the
months of February and May. These
adopted wolves are usually male and
between the age of one to three years old.
Adoptee males may mate with an
available pack female to then form their
own pack.[88] With wolves in the wild,
inbreeding does not occur where
outbreeding is possible.[89]

Wolves become mature at the age of two

years, and sexually mature from the age of
three years.[87] 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 in
order to better exploit abundant resources.
Females are capable of producing pups
every year, with one litter annually being
the average. Unlike the coyote, the wolf
never reaches menopause.[90] Estrus
commences in female wolves and the rut
begins in the second half of winter, when
the pack disperses. Adults disperse first,
then yearlings, then juveniles. The pack
later reunites but stays away from the
breeding pair. Estrus lasts for two
weeks.[87]
Illustrated growth stages of the northwestern wolf:
newborn, three weeks old, two months old, and one-
year-old adult

Dens are usually constructed for pups

during the summer period. When building
dens, females make use of natural
shelters such as 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,
and typically faces southwards where it
can be better warmed by sunlight
exposure, and the snow more-quickly
thawing. Resting places, play areas for the
pups and food remains are commonly
found around wolf dens. The odour of
urine and rotting food emanating from the
denning area often attracts scavenging
birds such as magpies and ravens. As
there are few convenient places for
burrows, wolf dens are usually occupied
by animals of the same family. Though
they mostly avoid areas within human
sight, wolves have been known to nest
near domiciles, paved roads and
railways.[91] During pregnancy, female
wolves remain in a den located away from
the peripheral zone of their territories,
where violent encounters with other packs
are more likely to occur.[92]

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 with 4-5 young, and
older females from 6-8 young and up to 14
young. Their mortality rate is 60-80
percent.[93] Pups are born blind and deaf,
and are covered in short soft grayish-
brown fur. They weigh 300–500 g (11–
18 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 1.5 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. Pups have a fast
growth rate during their first four months
of life: during this period, a pup's weight
can increase nearly 30 times.[93][94] 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 behavior.[95] By autumn, the
pups are mature enough to accompany
the adults on hunts for large prey.[92]

Foraging
American bison standing its ground against a pack of
wolves, thus increasing its chances of survival

Iberian wolf trotting in summer fur. They generally

place their hind paws in the tracks made by the front
paws.[36]

Although social animals, single wolves or

mated pairs typically have higher success
rates in hunting than do large packs, with
single wolves having occasionally been
observed to kill large prey such as moose,
bison and muskoxen unaided.[96] The
wolf's sense of smell is relatively weakly
developed when compared to that of
some hunting dog breeds, being able to
detect carrion upwind no farther than 2–3
kilometres (1.2–1.9 mi). Because of this, it
rarely manages to capture hidden hares or
birds, though it can easily follow fresh
tracks. Its auditory perception is acute
enough to be able to hear up to a
frequency of 26 kHz,[97] which is sufficient
to register the fall of leaves in the autumn
period.[69] A wolf hunt can be divided into
five stages:
Locating prey: The wolves travel in
search of prey through their power of
scent, chance encounter, and tracking.
Wolves typically locate their prey by
scent, though they must usually be
directly downwind of it. When a breeze
carrying the prey's scent is located, the
wolves stand alert, and point their eyes,
ears and nose towards their target. In
open areas, wolves may precede the
hunt with group ceremonies involving
standing nose-to-nose and wagging
their tails. Once concluded, the wolves
head towards their prey.[98]
The stalk: The wolves attempt to
conceal themselves as they
approach.[99] As the gap between the
wolves and their prey closes, the wolves
quicken their pace, wag their tails, and
peer intently, getting as close to their
quarry as possible without making it
flee.[100]
The encounter: Once the prey detects
the wolves, it can either approach the
wolves, stand its ground, or flee. Large
prey, such as moose, elk, and muskoxen,
usually stand their ground. Should this
occur, the wolves hold back, as they
require the stimulus of a running animal
to proceed with an attack.[101] If the
targeted animal stands its ground, the
wolves either ignore it, or try to
intimidate it into running.[96]
The rush: If the prey attempts to flee,
the wolves immediately pursue it. This
is the most critical stage of the hunt, as
wolves may never catch up with prey
running at top speed.[102] If their prey is
travelling in a group, the wolves either
attempt to break up the herd, or isolate
one or two animals from it.[99]
The chase: A continuation of the rush,
the wolves attempt to catch up with
their prey and kill it.[103] When chasing
small prey, wolves attempt to catch up
with their prey as soon as possible,
 while with larger animals, the chase is
prolonged, in order to wear the selected
prey out. Wolves usually give up chases
after 1–2 km (0.62–1.3 mi), though one
wolf was recorded to chase a deer for
21 km (13 mi).[96] Both Russian and
North American wolves have been
observed to drive prey onto crusted ice,
precipices, ravines, slopes and steep
banks to slow them down.[104]

Two wolves killing a moose in typical fashion: biting

the hindquarters
The actual killing method varies according
to prey species. With large prey, mature
wolves usually avoid attacking frontally,
instead focusing on the rear and sides of
the animal. Large prey, such as moose, is
killed by biting large chunks of flesh from
the soft perineum area, causing massive
blood loss. Such bites can cause wounds
10–15 cm (3.9–5.9 in) in length, with three
such bites to the perineum usually being
sufficient to bring down a large deer in
optimum health.[104] 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, mouse-like prey,
wolves leap in a high arc and immobilize it
with their forepaws.[105] When prey is
vulnerable and abundant, wolves may
occasionally surplus kill. Such instances
are common in domestic animals, but rare
in the wild. In the wild, surplus killing
primarily occurs during late winter or
spring, when snow is unusually deep (thus
impeding the movements of prey)[106] or
during the denning period, when wolves
require a ready supply of meat when
denbound.[107] Medium-sized prey are
especially vulnerable to surplus killing, as
the swift throat-biting method by which
they are killed allows wolves to quickly kill
one animal and move on to another.[105]

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.[108] The breeding
pair typically monopolizes food in order to
continue producing pups. When food is
scarce, this is done at the expense of
other family members, especially non-
pups.[109] The breeding pair typically eats
first, though as it is they who usually work
the hardest in killing prey, they may rest
after a long hunt and allow the rest of the
family to eat unmolested. Once the
breeding pair has finished eating, the rest
of the family tears off pieces of the
carcass and transport them to secluded
areas where they can eat in peace. Wolves
typically commence feeding by consuming
the larger internal organs of their prey,
such as the heart, liver, lungs and stomach
lining. The kidneys and spleen are eaten
once they are exposed, followed by the
muscles.[110] A single wolf can eat 15–
19% of its body weight in a single
feeding.[57]

Competition
 

Wolves attacking a mother grizzly bear with cubs

Wolves confronting coyotes over a pronghorn carcass

(1919), Louis Agassiz Fuertes

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.[111][112]
Near-identical interactions have been
observed in Eurasia between wolves and
golden jackals, with the latter's numbers
being comparatively small in areas with
high wolf densities.[69][111][113] Wolves are
the most important predator of raccoon
dogs, killing large numbers of them in the
spring and summer periods.[69] Wolves
also kill red, Arctic and corsac foxes,
usually in disputes over carcasses,
sometimes eating them.[69][114]

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
only eat young wolves.[115] Wolf
interactions with American black bears are
much rarer than with brown bears,
because of differences in habitat
preferences. The majority of American
black bear encounters with wolves occur
in the species' northern range, with no
interactions being recorded in Mexico.
Wolves have been recorded on numerous
occasions to actively seek out American
black bears in their dens and kill them
without eating them. Unlike brown bears,
American black bears frequently lose
against wolves in disputes over kills.[116]

Large wolf populations limit the numbers

of small to medium-sized felines, including
Eurasian lynx and bobcat.[117][118] Wolves
encounter cougars along portions of the
Rocky Mountains and adjacent mountain
ranges. Wolves and cougars typically
avoid encountering each other by hunting
on different elevations. In winter, however,
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. They have been reported
killing mothers and their kittens.[119][120]
Other than humans, tigers appear to be the
only serious predators of
wolves.[69][121][122][123][124] Wolf and tiger
interactions are well documented in
Sikhote-Alin, where tigers depress wolf
numbers, either to the point of localized
extinction or to such low numbers as to
make them a functionally insignificant
component of the ecosystem. Wolves
appear capable of escaping competitive
exclusion from tigers only when human
persecution decreases tiger numbers.
Proven cases of tigers killing wolves are
rare and attacks appear to be competitive
rather than predatory in nature, with at
least four proven records of tigers killing
wolves without consuming them.[125]

Wolves may encounter striped hyenas in

Israel, Central Asia and India, usually in
disputes over carcasses. Striped hyenas
feed extensively on wolf-killed carcasses
in areas where the two species interact.
One-on-one, hyenas dominate wolves, and
may prey on them,[126] but wolf packs can
drive off single or outnumbered
hyenas.[127][128] However, there was a case
of a female striped hyena dominating 12
Arabian wolves.[129]

Cooperation

In 1994 near Eilat in southern Israel,

researchers discovered the tracks of three
wolves and one striped hyena which
indicated that they were moving across
the landscape together. In 1998 near the
same location, three researchers sighted a
pack consisting of 4 adult and 3 sub-adult
wolves and one striped hyena. The hyena
was moving in the middle of the pack and
not following behind it. 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 such as tin cans.[130]

According to a press release by the US

Department of Agriculture in 1921, the
Custer Wolf moved across the landscape
with two coyotes far out on his flanks
which gave him warning of ambush or
danger. They fed from his kills, however he
would never allow them to approach
him.[131]
Diseases and parasites
Viral and bacterial infections

Play media
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, canine coronavirus,[132]
and foot and mouth disease. Wolves are a
major host for rabies in Russia, Iran,
Afghanistan, Iraq and India.[133] 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
kilometres (50 mi) a day, thus increasing
the risk of infecting other wolves. Infected
wolves do not show any fear of humans,
with 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 behaviors. The canine
coronavirus has been recorded in Alaskan
wolves, with infections being most
prevalent in winter months.[132]

Bacterial diseases carried by wolves

include brucellosis, lyme disease,
leptospirosis, tularemia, bovine
tuberculosis,[134] listeriosis and
anthrax.[133] 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 have any significant
effect on 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 once killed two wolf pups in
Canada.[134]

Parasitic infections

Wolves carry ectoparasites and

endoparasites, with wolves in the former
Soviet Union having been recorded to carry
at least 50 species.[133] Most of these
parasites infect wolves without adverse
effects, though the effects may become
more serious in sick or malnourished
specimens.[135] Parasitic infection in
wolves is of particular concern to people,
as 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.[133]

Wolf with sarcoptic mange

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),[135] though they
rarely develop full-blown mange, unlike
foxes.[69] 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.[135] The tick
Dermacentor pictus also infests wolves.
Other ectoparasites include chewing lice,
sucking lice and the fleas Pulex irritans
and Ctenocephalides canis.[69]
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.[135]
Wolves may carry Neospora caninum,
which is of particular concern to farmers,
as the disease can be spread to livestock,
with infected animals being three to 13
times more likely to miscarry than those
not infected.[136][137] Among flukes, the
most common in North American wolves
is Alaria, which infects small rodents and
amphibians that are eaten by wolves.
Upon reaching maturity, Alaria migrates to
the wolf's intestine, but harms it little.
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. 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 utero, 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. 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, Macrocantorhynchus catulinus,
and Moniliformis moniliformis.[135]

Conservation

 
Face of a wolf

Wolf population declines have been

arrested since the 1970s, and have
fostered recolonization and reintroduction
in parts of its former range due to 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 wolf's relatively widespread range and
stable population allows it to be classified
as Least Concern on the IUCN Red List.[2]
The species is included in CITES Appendix
II, except populations from Bhutan, India,
Nepal and Pakistan, which are listed on
Appendix I. The species is strictly
protected under the 1979 Berne
Convention on the Conservation of
European Wildlife and Natural Habitats
(Appendix II) and the 1992 European Union
Council Directive 92/43/EEC on the
Conservation of Natural Habitats and of
Wild Fauna and Flora (Annex II and IV), and
there is extensive legal protection in many
European countries; however, there are
national exceptions, and enforcement is
variable and often non-existent. The wolf
occurs in many protected areas across its
range, breeds well in captivity and is
common in many zoological gardens.[2]
The wolf received Endangered Species Act
(ESA) protection in Minnesota, Wisconsin,
and Michigan in 1974, and was
reclassified from endangered to
threatened in 2003. Reintroduced Mexican
wolves in Arizona and New Mexico are
protected under the ESA and, as of late
2002, number 28 individuals in eight
packs.[138]

Relationships with humans

In culture
In personal names

Old English literature contains several

instances of Anglo-Saxon kings and
warriors taking on wulf as a prefix or suffix
in their names. Examples include
Wulfhere, Cynewulf, Ceonwulf, Wulfheard,
Earnwulf, Wulfmǣr, Wulfstān and
Æthelwulf. Wolf-related names were also
common among pre-Christian Germanic
warriors: Wolfhroc (Wolf-Frock), Wolfhetan
(Wolf Hide), Isangrim (Grey Mask), Scrutolf
(Garb Wolf), Wolfgang (Wolf Gait) and
Wolfdregil (Wolf Runner).[139]

In mythology and folklore

 

Capitoline Wolf, sculpture of the mythical she-wolf

feeding the twins Romulus and Remus, from the
legend of the founding of Rome, Italy.

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.[140]
Romulus and Remus, the legendary
founders of Rome, were said to have been
raised and suckled by a she-wolf.[141]
Norse mythology includes the feared giant
wolf Fenrir, eldest child of Loki and
Angrboda,[142] and Geri and Freki, Odin's
faithful pets.[143] In the Pawnee creation
myth, the wolf was the first animal brought
to earth and when humans killed it, they
were punished with death, destruction and
the loss of immortality.[144]

The concept of people turning into wolves

has been present in many cultures. One
Greek myth tells of Lycaon of Arcadia
being transformed into a wolf by Zeus as
punishment for his evil deeds.[145] The
legend of the werewolf has been
widespread in European folklore and
involves people willingly turning into
wolves to attack and kill others.[146] The
Navajo have traditionally believed that
witches would turn into wolves by donning
wolf skins and would kill people and raid
graveyards.[147] The Dena’ina believed
wolves were once men, and viewed them
as brothers.[140]

For the Pawnee, Sirius was the "wolf star"

and its disappearance and reappearance
signified the wolf moving to and from the
spirit world. Both the Pawnee and
Blackfoot called the Milky Way the "wolf
trail".[148] In Chinese astronomy, the wolf
represents Sirius as the "blue beast" and
the star itself is called the "heavenly
wolf".[149]

In fable and literature

Little Red Riding Hood (1883), Gustave Doré

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 behavior, his
portrayals added to the wolf's image as a
deceitful and dangerous animal.[150] The
Bible contains 13 references to wolves,
usually as metaphors for greed and
destructiveness. In the New Testament,
Jesus is quoted to have used wolves as
illustrations to the dangers his followers,
whom he represents as sheep, would face
should they follow him (Matthew 7:15 ,
Matthew 10:16 , Acts 20:29 ).[151]
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.
Although portrayed as loyal, honest and
moral, Isengrim is forever the victim of
Reynard's wit and cruelty, often dying at
the end of each story.[152]
The tale of Little Red Riding Hood, first
written in 1697 by Charles Perrault, is
largely considered to have had more
influence than any other source of
literature in forging the wolf's negative
reputation in the western world. The wolf
in this story is portrayed as a potential
rapist, capable of imitating human
speech.[153] Similar wolf characters appear
in The Three Little Pigs and The Wolf and
the Seven Young Goats.[154] The hunting of
wolves, and their attacks on humans and
livestock feature prominently in Russian
literature, and are included in the works of
Tolstoy, Chekhov, Nekrasov, Bunin,
Sabaneyev, and others. Tolstoy's War and
Peace and Chekhov's Peasants both
feature scenes in which wolves are hunted
with hounds and Borzois.[155] Wolves are
among the central characters of Rudyard
Kipling's The Jungle Book, and his
portrayal of wolves has been praised
posthumously by wolf biologists for his
depiction of them: rather than being
villanous 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. The wolf's motto of "For
the strength of the Wolf is the Pack, and
the strength of the Pack is the Wolf" has
also been confirmed in field studies, which
show that success rates in hunting and
repelling intruders are dependent on pack
size and the number of adult males or old
wolves.[156] Farley Mowat's largely fictional
1963 memoir Never Cry Wolf is largely
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.[157][158][159]
In heraldry and symbolism

Coat of arms of the Elvange family, featuring a wolf

charge rampant

The wolf is a frequent charge in English

armory. It is illustrated as a supporter on
the shields of Lord Welby, Rendel, and
Viscount Wolseley, and can be found on
the coat of arms of Lovett and the vast
majority of the Wilsons and Lows. The
demi-wolf is a common crest, appearing in
the arms and crests of members of many
families, including that of the Wolfes,
whose crest depicts a demi-wolf holding a
crown in its paws, in reference to the
assistance the family gave to Charles II
during the battle of Worcester. Wolf heads
are common in Scottish heraldry,
particularly in the coats of Clan Robertson
and Skene. The wolf is the most common
animal in Spanish heraldry, and is often
depicted as carrying a lamb in its mouth,
or across its back.[160] The wolf is featured
on the flags of the Confederated Tribes of
the Colville Reservation, the Oneida Nation
of Wisconsin and the Pawnee.[161] The
Chechen wolf has been a symbol of the
Chechen Nation.[162] In modern times, the
wolf is widely used as an emblem for
military and paramilitary groups. It is the
unofficial symbol of the spetsnaz, and
serves as the logo of the Turkish Grey
Wolves. During the Yugoslav Wars, several
Serb paramilitary units adopted the wolf
as their symbol, including the White
Wolves and the Wolves of Vučjak.[163]

Conflicts

Waiting for a Chinook (c. 1900), Charles Marion

Waiting for a Chinook (c. 1900), Charles Marion
Russell

Human presence appears to stress

wolves, as seen by increased cortisol
levels in instances such as snowmobiling
near their territory.[164]

Wolf predation on livestock

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.[165]
Some nations help offset economic losses
to wolves through compensation
programmes or state insurance.[166]
Domesticated animals are easy prey for
wolves, as they have evolved under
constant human protection, and are thus
unable to defend themselves very well.[167]
Wolves typically resort to attacking
livestock when wild prey is depleted: 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.[165]
The majority of losses occur during the
summer grazing period, with untended
livestock in remote pastures being the
most vulnerable to wolf predation.[168] The
most frequently targeted livestock species
are sheep (Europe), domestic reindeer
(northern Scandinavia), goats (India),
horses (Mongolia), cattle and turkeys
(North America).[165] 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.[169] Wolves mainly
attack livestock when the animals are
grazing, though they occasionally break
into fenced enclosures.[104] In some cases,
wolves do not need to physically attack
livestock to negatively affect it: the stress
livestock experiences in being vigilant for
wolves may result in miscarriages, weight
loss and a decrease in meat quality.[136]

Conflicts with dogs

Being the most abundant carnivores, free-

ranging dogs have the greatest potential
to compete with wolves. A review of the
studies in the competitive effects of dogs
on sympatric carnivores did not mention
any research on competition between
dogs and wolves.[170][171] Competition
would favor the wolf, which is known to kill
dogs, however wolves tend to live in pairs
or in small packs in areas where they are
highly persecuted, giving them a
disadvantage facing large groups of
dogs.[171][172]

Wolves kill dogs on occasion, with some

wolf populations relying 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 behavior when attacking dogs
accompanied by people, sometimes
ignoring nearby humans.[165] Wolf attacks
on dogs may occur both in house yards
and in forests.[173] Wolf attacks on hunting
dogs are considered a major problem in
Scandinavia and Wisconsin.[165][173] The
most frequently killed hunting breeds in
Scandinavia are harriers, with 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.[173]

Although the numbers of dogs killed each

year are relatively low, it induces a fear of
wolves entering villages and farmyards to
take dogs. In many cultures, there are
strong social and emotional bonds
between humans and their dogs that can
be seen as family members or working
team members. The loss of a dog can lead
to strong emotional responses with
demands for more liberal wolf hunting
regulations.[171]

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.[171][174] Shepherd dogs are not
particularly aggressive, but they can
disrupt potential wolf predation by
displaying what is to the wolf ambiguous
behaviors, such as barking, social greeting,
invitation to play or aggression.[171][175]
The historical use of shepherd dogs
across Eurasia has been effective against
wolf predation,[171][175] especially when
confining sheep in the presence of several
livestock guardian dogs.[171][176] However,
shepherd dogs are sometimes killed by
wolves.[171][177]

Wolf attacks on humans

 

Small farmers 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.[178] 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.[179]
Although wolves may react aggressively
under provocation, such attacks are
mostly limited to quick bites on
extremities, and the attacks are not
pressed.[178]

Predatory attacks (attacks by wolves

treating humans as food) 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,[178] though cases of non-rabid
wolf attacks in winter have been recorded
in Belarus, Kirov and Irkutsk oblasts,
Karelia and Ukraine.[69] Also, wolves with
pups experience greater food stresses
during this period.[69] 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.[178]
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, Middle East and Central
Asia. Wolves apparently develop the
"furious" phase of rabies to a very high
degree which, coupled with their size and
strength, makes rabid wolves perhaps the
most dangerous of rabid animals,[178] with
bites from rabid wolves being 15 times
more dangerous than those of rabid
dogs.[69] 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 only occur on a single day. The
victims are chosen at random, though the
majority of cases involve adult men.
During the 50 years to 2002, there were
eight fatal attacks in Europe and Russia,
and more than 200 in southern Asia.[178]

Human hunting of wolves

 

Carcasses of hunted wolves in Russia

Theodore Roosevelt stated that wolves are

difficult to hunt because of their
elusiveness, sharp senses, high
endurance, and ability to quickly
incapacitate and kill hunting dogs.[180]
Historic methods include killing of spring-
born litters in their dens, coursing with
dogs (usually combinations of
sighthounds, bloodhounds and fox
terriers), poisoning with strychnine, and
trapping.[181][182] 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 smell. Some hunters
are able to 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,[182] but is
controversial.[183] Several types of dog,
including the Borzoi and Kyrgyz Tajgan,
have been specifically bred for wolf
hunting.[171]

As pets and working animals

Wild wolves are sometimes kept as exotic

pets and, in some rarer occasions, as
working animals. Although closely related
to domesticated dogs, wolves do not show
the same tractability as dogs in living
alongside humans, and generally, much
more work is required in order to obtain
the same amount of reliability. Wolves also
need much more space than dogs, about
26–39 square kilometres (10–15 sq mi),
so they can exercise.[184]

References
1. Tedford, Richard H.; Wang, Xiaoming;
Taylor, Beryl E. (2009). "Phylogenetic
Systematics of the North American
Fossil Caninae (Carnivora: Canidae)".
Bulletin of the American Museum of
Natural History. 325: 1–218.
doi:10.1206/574.1 . hdl:2246/5999 .
2. Boitani, L., Phillips, M. & Jhala, Y.
(2018). "Canis lupus" . The IUCN Red
List of Threatened Species. IUCN.
2018. e.T3746A119623865.
doi:10.2305/IUCN.UK.2018-
2.RLTS.T3746A119623865.en .
3. Linnæus, Carl (1758). "Canis Lupus" .
Systema naturæ per regna tria naturæ,
secundum classes, ordines, genera,
species, cum characteribus,
differentiis, synonymis, locis. Tomus I
(in Latin) (10 ed.). Holmiæ
(Stockholm): Laurentius Salvius.
pp. 39–40.
4. Wozencraft, W.C. (2008). "Family
Canidae" . In Smith, A.T.; Xie, Y.; et al.
(eds.). A Guide to the Mammals of
China. Princeton University Press.
pp. 416–418. ISBN 978-0691099842.
5. Wang, L; Ma, Y. P.; Zhou, Q. J.; Zhang, Y.
P.; Savolaimen, P.; Wang, G. D. (2016).
"The geographical distribution of grey
wolves (Canis lupus) in China: A
systematic review" . Zoological
Research. 37 (6): 315–326.
doi:10.13918/j.issn.2095-
8137.2016.6.315 . PMC 5359319 .
PMID 28105796 .
6. Larson, Greger; Larson, Greger (2017).
"Reconsidering the distribution of gray
wolves" . Zoological Research. 38 (3):
115–116. doi:10.24272/j.issn.2095-
8137.2017.021 . PMC 5460078 .
PMID 28585433 .
7. Boitani, L. (1995). "Ecological and
cultural diversities in the evolution of
wolf–human relationships" (PDF). In
Carbyn, L. N.; Fritts, S. H.; Seip, D. R.
(eds.). Ecology and conservation of
wolves in a changing world.
Edmonton: Canadian Circumpolar
Institute. pp. 3–12.
8. Harper, Douglas. "wolf" . Online
Etymology Dictionary.
9. Wozencraft, C. W. (2005). "Order
Carnivora" . In Wilson, D. E.; Reader, D.
M. (eds.). Mammal Species of the
World: A Taxonomic and Geographic
Reference. 1 (3rd ed.). Johns Hopkins
University Press. pp. 575–577.
ISBN 978-0-8018-8221-0.
10. Thalmann, Olaf; Perri, Angela R.
(2018). "Paleogenomic Inferences of
Dog Domestication". In Lindqvist, C.;
Rajora, O. (eds.). Paleogenomics.
Population Genomics. Springer, Cham.
pp. 273–306.
doi:10.1007/13836_2018_27 .
ISBN 978-3-030-04752-8.
11. Freedman, Adam H.; Gronau, Ilan;
Schweizer, Rena M.; Ortega-Del
Vecchyo, Diego; Han, Eunjung; et al.
(2014). "Genome Sequencing
Highlights the Dynamic Early History
of Dogs" . PLOS Genetics. 10 (1).
e1004016.
doi:10.1371/journal.pgen.1004016 .
PMC 3894170 . PMID 24453982 .
12. Skoglund, Pontus; Ersmark, Erik;
Palkopoulou, Eleftheria; Dalén, Love
(2015). "Ancient Wolf Genome Reveals
an Early Divergence of Domestic Dog
Ancestors and Admixture into High-
Latitude Breeds". Current Biology. 25
(11): 1515–1519.
doi:10.1016/j.cub.2015.04.019 .
PMID 26004765 .
13. Fan, Zhenxin; Silva, Pedro; Gronau, Ilan;
Wang, Shuoguo; Armero, Aitor Serres;
Schweizer, Rena M.; Ramirez, Oscar;
Pollinger, John; Galaverni, Marco;
Ortega Del-Vecchyo, Diego; Du,
Lianming; Zhang, Wenping; Zhang,
Zhihe; Xing, Jinchuan; Vilà, Carles;
Marques-Bonet, Tomas; Godinho,
Raquel; Yue, Bisong; Wayne, Robert K.
(2016). "Worldwide patterns of
genomic variation and admixture in
gray wolves" . Genome Research. 26
(2): 163–73.
doi:10.1101/gr.197517.115 .
PMC 4728369 . PMID 26680994 .
14. Koblmüller, Stephan; Vilà, Carles;
Lorente-Galdos, Belen; Dabad, Marc;
Ramirez, Oscar; Marques-Bonet,
Tomas; Wayne, Robert K.; Leonard,
Jennifer A. (2016). "Whole
mitochondrial genomes illuminate
ancient intercontinental dispersals of
grey wolves (Canis lupus)". Journal of
Biogeography. 43 (9): 1728–1738.
doi:10.1111/jbi.12765 .
hdl:10261/153364 .
15. Musiani M, Leonard JA, Cluff H, Gates
CC, Mariani S, et al. (2007).
"Differentiation of tundra/taiga and
boreal coniferous forest wolves:
genetics, coat colour and association
with migratory caribou". Mol. Ecol. 16
(19): 4149–70. doi:10.1111/j.1365-
294x.2007.03458.x . PMID 17725575 .
16. Schweizer, Rena M.; Vonholdt, Bridgett
M.; Harrigan, Ryan; Knowles, James C.;
Musiani, Marco; Coltman, David;
Novembre, John; Wayne, Robert K.
(2016). "Genetic subdivision and
candidate genes under selection in
North American grey wolves".
Molecular Ecology. 25 (1): 380–402.
doi:10.1111/mec.13364 .
PMID 26333947 .
17. Geffen, ELI; Anderson, Marti J.; Wayne,
Robert K. (2004). "Climate and habitat
barriers to dispersal in the highly
mobile grey wolf". Molecular Ecology.
13 (8): 2481–90. doi:10.1111/j.1365-
294X.2004.02244.x .
PMID 15245420 .
18. Pilot, Malgorzata; Jedrzejewski,
Wlodzimierz; Branicki, Wojciech;
Sidorovich, Vadim E.; Jedrzejewska,
Bogumila; Stachura, Krystyna; Funk,
Stephan M. (2006). "Ecological factors
influence population genetic structure
of European grey wolves". Molecular
Ecology. 15 (14): 4533–53.
doi:10.1111/j.1365-
294X.2006.03110.x .
PMID 17107481 .
19. Flower, Lucy O.H.; Schreve, Danielle C.
(2014). "An investigation of
palaeodietary variability in European
Pleistocene canids". Quaternary
Science Reviews. 96: 188–203.
Bibcode:2014QSRv...96..188F .
doi:10.1016/j.quascirev.2014.04.015 .
20. Leonard, Jennifer (2014). "Ecology
drives evolution in grey wolves" (PDF).
Evolution Ecology Research. 16: 461–
473.
21. Stronen, Astrid V.; Jä™Drzejewska,
Bogumiå'a; Pertoldi, Cino; Demontis,
Ditte; Randi, Ettore; Niedziaå'Kowska,
Magdalena; Pilot, MaÅ'Gorzata;
Sidorovich, Vadim E.; Dykyy, Ihor;
Kusak, Josip; Tsingarska, Elena;
Kojola, Ilpo; Karamanlidis, Alexandros
A.; Ornicans, Aivars; Lobkov, Vladimir
A.; Dumenko, Vitalii; Czarnomska,
Sylwia D. (2013). "North-South
Differentiation and a Region of High
Diversity in European Wolves (Canis
lupus)" . PLOS ONE. 8 (10). e76454.
Bibcode:2013PLoSO...876454S .
doi:10.1371/journal.pone.0076454 .
PMC 3795770 . PMID 24146871 .
22. Schweizer, Rena M.; Robinson,
Jacqueline; Harrigan, Ryan; Silva,
Pedro; Galverni, Marco; Musiani,
Marco; Green, Richard E.; Novembre,
John; Wayne, Robert K. (2016).
"Targeted capture and resequencing of
1040 genes reveal environmentally
driven functional variation in grey
wolves". Molecular Ecology. 25 (1):
357–79. doi:10.1111/mec.13467 .
PMID 26562361 .
23. Wayne, R. (1993). "Molecular evolution
of the dog family". Trends in Genetics.
9 (6): 218–24. doi:10.1016/0168-
9525(93)90122-X . PMID 8337763 .
24. Wayne, R. K.; Benvenista, R. E.;
Janczewski, D. N.; O'Brien, S. J. (1989).
"17-Molecular and biomechanical
evolution of the Carnivora" . In
Gittleman, J. L. (ed.). Carnivore
Behavior, Ecology, and Evolution.
Springer. pp. 477–478.
ISBN 9780412343605.
25. Vila, C.; Leonard, J. (2012). "1-Canid
Phylogeny and the Origin of the
Domestic Dog" . In Ostrander, E. A.;
Ruvinski, A. (eds.). Genetics of the Dog
(2 ed.). CABI. p. 3.
ISBN 9781845939403.
26. Wayne, R. & Ostrander, Elaine A.
(1999). "Origin, genetic diversity, and
genome structure of the domestic
dog". BioEssays. 21 (3): 247–57.
doi:10.1002/(SICI)1521-
1878(199903)21:3<247::AID-
BIES9>3.0.CO;2-Z . PMID 10333734 .
27. Gopalakrishnan, Shyam; Sinding,
Mikkel-Holger S.; Ramos-Madrigal,
Jazmín; Niemann, Jonas; Samaniego
Castruita, Jose A.; Vieira, Filipe G.;
Carøe, Christian; Montero, Marc de
Manuel; Kuderna, Lukas; Serres, Aitor;
González-Basallote, Víctor Manuel; Liu,
Yan-Hu; Wang, Guo-Dong; Marques-
Bonet, Tomas; Mirarab, Siavash;
Fernandes, Carlos; Gaubert, Philippe;
Koepfli, Klaus-Peter; Budd, Jane;
Rueness, Eli Knispel; Heide-Jørgensen,
Mads Peter; Petersen, Bent; Sicheritz-
Ponten, Thomas; Bachmann, Lutz;
Wiig, Øystein; Hansen, Anders J.;
Gilbert, M. Thomas P. (2018).
"Interspecific Gene Flow Shaped the
Evolution of the Genus Canis" .
Current Biology. 28 (21): 3441–
3449.e5.
doi:10.1016/j.cub.2018.08.041 .
PMC 6224481 . PMID 30344120 .
28. Sinding, Mikkel-Holger S.;
Gopalakrishan, Shyam; Vieira, Filipe G.;
Samaniego Castruita, Jose A.;
Raundrup, Katrine; Heide Jørgensen,
Mads Peter; Meldgaard, Morten;
Petersen, Bent; Sicheritz-Ponten,
Thomas; Mikkelsen, Johan Brus;
Marquard-Petersen, Ulf; Dietz, Rune;
Sonne, Christian; Dalén, Love;
Bachmann, Lutz; Wiig, Øystein;
Hansen, Anders J.; Gilbert, M. Thomas
P. (2018). "Population genomics of
grey wolves and wolf-like canids in
North America" . PLOS Genetics. 14
(11): e1007745.
 doi:10.1371/journal.pgen.1007745 .
PMC 6231604 . PMID 30419012 .
29. Anderson, T. M.; Vonholdt, B. M.;
Candille, S. I.; Musiani, M.; Greco, C.;
Stahler, D. R.; Smith, D. W.;
Padhukasahasram, B.; Randi, E.;
Leonard, J. A.; Bustamante, C. D.;
Ostrander, E. A.; Tang, H.; Wayne, R. K.;
Barsh, G. S. (2009). "Molecular and
Evolutionary History of Melanism in
North American Gray Wolves" .
Science. 323 (5919): 1339–1343.
Bibcode:2009Sci...323.1339A .
doi:10.1126/science.1165448 .
PMC 2903542 . PMID 19197024 .
30. Iacolina, Laura; Scandura, Massimo;
Gazzola, Andrea; Cappai, Nadia;
Capitani, Claudia; Mattioli, Luca;
Vercillo, Francesca; Apollonio, Marco
(2010). "Y-chromosome microsatellite
variation in Italian wolves: A
contribution to the study of wolf-dog
hybridization patterns". Mammalian
Biology - Zeitschrift für
Säugetierkunde. 75 (4): 341.
doi:10.1016/j.mambio.2010.02.004 .
31. Kopaliani, N.; Shakarashvili, M.;
Gurielidze, Z.; Qurkhuli, T.;
Tarkhnishvili, D. (2014). "Gene Flow
between Wolf and Shepherd Dog
Populations in Georgia (Caucasus)".
Journal of Heredity. 105 (3): 345–53.
doi:10.1093/jhered/esu014 .
PMID 24622972 .
32. Moura, A. E.; Tsingarska, E.;
Dąbrowski, M. J.; Czarnomska, S. D.;
Jędrzejewska, B. A.; Pilot, M. G.
(2013). "Unregulated hunting and
genetic recovery from a severe
population decline: The cautionary
case of Bulgarian wolves".
Conservation Genetics. 15 (2): 405–
417. doi:10.1007/s10592-013-0547-y .
33. Mech, D. L. (1974). "Canis lupus" .
Mammalian Species. 37 (37): 1–6.
doi:10.2307/3503924 .
JSTOR 3503924 . Retrieved July 30,
2019.
34. Heptner & Naumov 1998, pp. 129-132.
35. Heptner & Naumov 1998, pp. 164-270.
36. Mech 1981, p. 13.
37. Lopez 1978, p. 23.
38. Macdonald, D.W.; Norris, S. (2001).
Encyclopedia of Mammals . Oxford
University Press. p. 45. ISBN 978-0-
7607-1969-5.
39. Lopez 1978, p. 19.
40. Lopez 1978, p. 18.
41. Young & Goldman 1944, p. 69.
42. Mech & Boitani 2003, p. 32.
43. Nowak, R.M.; Paradiso, J.L. (1983).
"Carnivora;Canidae". Walker's
Mammals of the World . 2 (4th ed.).
Johns Hopkins University Press.
p. 953. ISBN 9780801825255.
44. Mech 1981, p. 14.
45. Therrien, F. O. (2005). "Mandibular
force profiles of extant carnivorans
and implications for the feeding
behaviour of extinct predators".
Journal of Zoology. 267 (3): 249–270.
doi:10.1017/S0952836905007430 .
46. Mech & Boitani 2003, p. 112.
47. Paquet, P.; Carbyn, L.W. (2003). "Ch23:
Gray wolf Canis lupus and allies". In
Feldhamer, G.A.; Thompson, B.C.;
Chapman, J.A. (eds.). Wild Mammals
of North America: Biology,
Management, and Conservation (2
ed.). John Hopkins University Press.
pp. 482–510. ISBN 0-8018-7416-5.
48. Christiansen, Per; Wroe, Stephen
(2007). "Bite Forces and Evolutionary
Adaptations to Feeding Ecology in
Carnivores". Ecology. 88 (2): 347–358.
doi:10.1890/0012-
9658(2007)88[347:bfaeat]2.0.co;2 .
PMID 17479753 .
49. Lopez 1978, p. 21.
50. Mech 1981, pp. 16–17.
51. Lopez 1978, p. 22.
52. Lopez 1978, pp. 19–20.
53. Earle, M (1987). "A flexible body mass
in social carnivores". American
Naturalist. 129 (5): 755–760.
doi:10.1086/284670 .
54. Sorkin, Boris (2008). "A biomechanical
constraint on body mass in terrestrial
mammalian predators". Lethaia. 41
(4): 333–347. doi:10.1111/j.1502-
3931.2007.00091.x .
55. Mech, L. David (1966). The Wolves of
Isle Royale . Fauna Series 7. Fauna of
the National Parks of the United
States. p. 76. ISBN 978-1-4102-0249-9.
56. Mech 1981, p. 172.
57. Mech & Boitani 2003, pp. 201.
58. Newsome, Thomas M.; Boitani, Luigi;
Chapron, Guillaume; Ciucci, Paolo;
Dickman, Christopher R.; Dellinger,
Justin A.; López-Bao, José V.;
Peterson, Rolf O.; Shores, Carolyn R.;
Wirsing, Aaron J.; Ripple, William J.
(2016). "Food habits of the world's
grey wolves". Mammal Review. 46 (4):
255. doi:10.1111/mam.12067 .
59. Mech & Boitani 2003, p. 107.
60. Mech & Boitani 2003, pp. 109-110.
61. Heptner & Naumov 1998, pp. 213-231.
62. Gable, T. D.; Windels, S. K.; Homkes, A.
T. (2018). "Do wolves hunt freshwater
fish in spring as a food source?".
Mammalian Biology. 91: 30–33.
doi:10.1016/j.mambio.2018.03.007 .
63. Woodford, Riley. "Alaska's Salmon-
Eating Wolves" .
Wildlifenews.alaska.gov. Retrieved
July 25, 2019.
64. McAllister, I. (2007). The Last Wild
Wolves: Ghosts of the Rain Forest .
University of California Press. p. 144.
ISBN 978-0520254732.
65. Fuller, T.K. (2019). "Ch3-What wolves
eat". Wolves: Spirit of the Wild .
Chartwell Crestline. p. 53. ISBN 978-
0785837381.
66. Mech & Boitani 2003, p. 109.
67. Mech 1981, p. 180.
68. Klein, D. R. (1995). "The introduction,
increase, and demise of wolves on
Coronation Island, Alaska". In Carbyn,
L. N., Fritts, S. H., Seip, D. R. (eds.).
Ecology and conservation of wolves in
a changing world. Canadian
Circumpolar Institute, Occasional
Publication No. 35. pp. 275–280.
69. Heptner, V.G. & Naumov, N.P. (1998).
Mammals of the Soviet Union Vol.II
Part 1a, SIRENIA AND CARNIVORA
(Sea cows; Wolves and Bears) .
Science Publishers, Inc. USA. pp. 164–
270. ISBN 1-886106-81-9.
70. Mech & Boitani 2003, pp. 2–3, 28.
71. Mech & Boitani 2003, pp. 1–2.
72. Mech & Boitani 2003, pp. 12–13.
73. Mech & Boitani 2003, p. 38.
74. Molnar, B., Fattebert, J., Palme, R.,
Ciucci, P., Betschart, B., Smith, D.W., &
Diehl, P. (2015). "Environmental and
intrinsic correlates of stress in free-
ranging wolves" . PLOS ONE. 10 (9).
e0137378.
Bibcode:2015PLoSO..1037378M .
doi:10.1371/journal.pone.0137378 .
PMC 4580640 . PMID 26398784 .
75. Jędrzejewski, W. O.; Schmidt, K.;
Theuerkauf, J. R.; Jędrzejewska, B. A.;
Kowalczyk, R. (2007). "Territory size of
wolves Canis lupus: Linking local
(Białowieża Primeval Forest, Poland)
and Holarctic-scale patterns".
Ecography. 30: 66–76.
doi:10.1111/j.0906-
7590.2007.04826.x .
76. Mech & Boitani 2003, pp. 19–26.
77. Mech, L. D. (1977). "Wolf-Pack Buffer
Zones as Prey Reservoirs" . Science.
198 (4314): 320–321.
Bibcode:1977Sci...198..320M .
doi:10.1126/science.198.4314.320 .
PMID 17770508 .
78. Mech, D.L.; Adams, L.G.; Meier, T.J.;
Burch, J.W.; Dale, B.W. (2003). "Ch.8-
The Denali Wolf-Prey System". The
Wolves of Denali . University of
Minnesota Press. p. 163. ISBN 0-8166-
2959-5.
79. Mech & Boitani 2003, p. 67.
80. Mech & Boitani 2003, p. 66-103.
81. Busch 2007, p. 59.
82. Lopez 1978, p. 38.
83. Lopez 1978, pp. 39–41.
84. Mech & Boitani 2003, p. 90.
85. Peters, R. P.; Mech, L. D. (1975).
"Scent-marking in wolves". American
Scientist. 63 (6): 628–637.
Bibcode:1975AmSci..63..628P .
PMID 1200478 .
86. Mech & Boitani 2003, p. 1.
87. Heptner & Naumov 1998, pp. 248.
88. Mech & Boitani 2003, p. 2-3.
89. Mech & Boitani 2003, p. 5.
90. Mech & Boitani 2003, p. 175.
91. Heptner & Naumov 1998, pp. 234-238.
 92. Mech & Boitani 2003, pp. 42–46.
93. Heptner & Naumov 1998, pp. 249-254.
94. Mech & Boitani 2003, pp. 46–49.
95. Fox, M.W. (1978). The Dog: Its
Domestication and Behavior . Garland
STPM. p. 33. ISBN 978-0894642029.
96. Mech & Boitani 2003, pp. 119–121.
97. Lopez 1978, p. 43.
98. Mech 1981, pp. 196–199.
99. Graves 2007, p. 43.
100. Mech 1981, pp. 199–200.
101. Mech 1981, pp. 200–201.
102. Mech 1981, pp. 201–202.
103. Mech 1981, pp. 202–203
104. Graves 2007, p. 45.
105. Zimen 1981, pp. 217–218.
106. Mech & Boitani 2003, p. 144.
107. Lopez 1978, pp. 54–55.
108. Mech 1981, p. 185.
109. Mech & Boitani 2003, p. 58.
110. Mech & Boitani 2003, pp. 122–5.
111. Mech & Boitani 2003, pp. 266–68.
112. Robbins, Jim (1998). "Weaving A New
Web: Wolves Change An Ecosystem" .
Smithsonian National Zoological Park.
Archived from the original on January
24, 2009. Retrieved August 10, 2007.
113. Giannatos G. (April 2004).
"Conservation Action Plan for the
golden jackal Canis aureus L. in
Greece" (PDF). WWF Greece. pp. 1–
47. Retrieved 2006-11-15.
114. Mech & Boitani 2003, p. 269.
115. Mech & Boitani 2003, pp. 261–63.
116. Mech & Boitani 2003, pp. 263–64.
117. Sunquist, Melvin E. & Sunquist, Fiona
(2002). Wild cats of the world .
University of Chicago Press. p. 167.
ISBN 0-226-77999-8.
118. "Conservation Groups Challenge Bush
Administration Wolf Killing Plan — "It's
going to be open season on wolves,"
says Natural Resources Defense
Council" . Yellowstone National
Park.com. July 25, 2019. Retrieved
2019-07-16.
119. Mech & Boitani 2003, pp. 264–65
120. Grooms, Steve (2010). "Cougar Wolf
Interactions: It's a Lot Like Cats and
Dogs" . International Wolf. 20 (2): 8–
11.
121. Perry, R. (1964). The World of the
Tiger. Cassell & Company. p. 148.
122. Lopez 1978, p. 29.
123. Mech & Boitani 2003, p. 265.
124. Heptner, V.G. & Sludskii, A.A. (1992).
Mammals of the Soviet Union:
Carnivora (hyaenas and cats), Volume
2. BRILL. pp. 313, 492. ISBN 90-04-
08876-8.
125. Miquelle, D.G., Stephens, P.A., Smirnov,
E.N., Goodrich, J.M., Zaumyslova,
O.Yu. & Myslenkov, A.I. (2005). "Tigers
and Wolves in the Russian Far East:
Competitive Exclusion, Functional
Redundancy and Conservation
Implications". In Ray, J.C., Berger, J.,
Redford, K.H. & Steneck, R. (eds.).
Large Carnivores and the Conservation
of Biodiversity . Island Press. pp. 179–
207. ISBN 1-55963-080-9.
126. Monchot, H. & Mashkour, H. (2010).
"Hyenas around the cities. The case of
Kaftarkhoun (Kashan- Iran)" . Journal
of Taphonomy. 8 (1): 17–32..
127. Mills, M. G. L.; Mills, Gus; Hofer,
Heribert (1998). Hyaenas: status
survey and conservation action plan .
IUCN. pp. 24–25. ISBN 978-2-8317-
0442-5.
128. Nayak, S.; Shah, S.; Borah, J. (2015).
"Going for the kill: an observation of
wolf-hyaena interaction in Kailadevi
Wildlife Sanctuary, Rajasthan, India".
Canid Biology & Conservation. 18 (7):
27–29.
129. "BBC Two – Wild Arabia, The Jewel of
Arabia, Wolves vs hyena" . BBC.
130. Dinets, Vladimir; Eligulashvili,
Beniamin (2016). "Striped Hyaenas
(Hyaena hyaena) in Grey Wolf (Canis
lupus) packs: Cooperation,
commensalism or singular
aberration?". Zoology in the Middle
East. 62: 85–87.
doi:10.1080/09397140.2016.1144292
.
131. Merrit, Dixon (January 7, 1921).
"World's Greatest Animal Dead" (PDF).
US Department of Agriculture Division
of Publications. p. 2. Retrieved July 26,
2019.
132. Mech & Boitani 2003, pp. 208–211.
133. Graves 2007, pp. 77–85.
134. Mech & Boitani 2003, pp. 211–213.
135. Mech & Boitani 2003, pp. 202–208.
136. "Effects of Wolves and Other
Predators on Farms in Wisconsin:
Beyond Verified Losses" (PDF).
Wisconsin Department of Natural
Resources. Archived from the original
(PDF) on March 19, 2009. Retrieved
January 25, 2013.
137. Dubey, J. P.; Jenkins, M. C.; Rajendran,
C.; Miska, K.; Ferreira, L. R.; Martins, J.;
Kwok, O. C. H.; Choudhary, S. (2011).
"Gray wolf (Canis lupus) is a natural
definitive host for Neospora
caninum" . Veterinary Parasitology.
181 (2–4): 382–387.
doi:10.1016/j.vetpar.2011.05.018 .
PMID 21640485 .
138. Mech & Boitani 2003, pp. 321–324
139. Marvin 2012, pp. 74–75.
140. Mech & Boitani 2003, p. 292.
141. Marvin 2012, pp. 128.
142. Simek, Rudolf (2007). Dictionary of
Northern Mythology. translated by
Angela Hall. D.S. Brewer. p. 81.
ISBN 978-0-85991-513-7.
143. Marvin 2012, pp. 78.
144. Lopez 1978, p. 133.
145. Marvin 2012, p. 47.
146. Marvin 2012, p. 50.
147. Lopez 1978, p. 123.
148. Busch 2007, p. 110.
149. Brosch, Noah (2008). Sirius Matters.
Springer. p. 29. ISBN 1-4020-8318-1.
150. Marvin 2012, pp. 38–40.
151. Bright, Michael (2006). Beasts of the
Field: The Revealing Natural History of
Animals in the Bible. pp. 115–116.
ISBN 1-86105-831-4.
152. Lopez 1978, p. 259.
153. Marvin 2012, pp. 64–70.
154. Lopez 1978, p. 263.
155. Graves 2007, pp. 21, 123.
156. Cassidy, K, DW Smith, LD Mech, DR
MacNulty, DR Stahler, MC Metz (2006).
"Territoriality and interpack aggression
in wolves: Shaping a social carnivore's
life history. Rudyard Kipling's Law of
the Jungle Meets Yellowstone's Law of
the Mountains" . Yellowstone Science.
24 (1): 37–41.
157. Mech & Boitani 2003, p. 294.
158. Jones, Karen (2001). "Never Cry Wolf:
Science, Sentiment, and the Literary
Rehabilitation of Canis Lupus" . The
Canadian Historical Review. 84.
159. Grooms, Steve (2008). "The Mixed
Legacy of Never Cry Wolf" (PDF).
International Wolf. 18 (3): 11–13.
Archived from the original (PDF) on
June 21, 2010.
160. Davies, Arthur Charles Fox & Johnston,
Graham (2004). A Complete Guide to
Heraldry . Kessinger Publishing.
pp. 196–197. ISBN 1-4179-0630-8.
161. Healy, Donald T. & Orenski, Peter J.
(2003). Native American Flags .
University of Oklahoma Press. pp. 48,
153, 166. ISBN 0-8061-3556-5.
162. Jaimoukha, Amjad (2005). The
Chechens: A Handbook, Caucasus
World: Peoples of the Caucasus.
Routledge. pp. 146–147. ISBN 978-
0415323284.
163. Marvin 2012, pp. 78–79.
164. Creel, S., Fox, J.E., Hardy, A., Sands, J.,
Garrott, B., & Peterson, R.O. (2002).
"Snowmobile activity and
glucocorticoid stress responses in
wolves and elk". Conservation Biology.
16 (3): 809–814. doi:10.1046/j.1523-
1739.2002.00554.x .
165. Mech & Boitani 2003, p. 305.
166. Mech & Boitani 2003, p. 309
167. Mech 1981, p. 173
168. Mech & Boitani 2003, p. 307
169. Mech & Boitani 2003, p. 306
170. Vanak, A.T., Dickman, C.R., Silva-
Rodriguez, E.A., Butler, J.R.A., Ritchie,
E.G. (2014). "Top-dogs and under-
dogs: competition between dogs and
sympatric carnivores". In Gompper,
M.E. (ed.). Free-Ranging Dogs and
Wildlife Conservation. Oxford
University Press. pp. 69–93. ISBN 978-
0199663217.
171. Lescureux, Nicolas; Linnell, John D.C.
(2014). "Warring brothers: The
complex interactions between wolves
(Canis lupus) and dogs (Canis
familiaris) in a conservation context".
Biological Conservation. 171: 232–
245.
doi:10.1016/j.biocon.2014.01.032 .
172. Boitani, L. (1983). "Wolf and dog
competition in Italy". Acta Zoologica
Fennica (174): 259–264.
173. Backeryd, J. (2007). "Wolf attacks on
dogs in Scandinavia 19952005 – Will
wolves in Scandinavia go extinct if dog
owners are allowed to kill a wolf
attacking a dog?" (PDF).
Examensarbete, Institutionen för
ekologi, Grimsö forskningsstation.
Sveriges Lantbruksuniversitet.
Retrieved 2019-07-17..
174. Shivik, John A. (2006). "Tools for the
Edge: What's New for Conserving
Carnivores". BioScience. 56 (3): 253.
doi:10.1641/0006-
3568(2006)056[0253:TFTEWN]2.0.CO;
2.
175. Coppinger, R. & Schneider, R. (1995).
"Evolution of working dogs" . In
Serpell, J. (ed.). The Domestic Dog: Its
Evolution, Behaviour and Interactions
With People. University Press,
Cambridge. pp. 21–47.
176. Espuno, Nathalie; Lequette, Benoit;
Poulle, Marie-Lazarine; Migot, Pierre;
Lebreton, Jean-Dominique (2004).
"Heterogeneous response to
preventive sheep husbandry during
wolf recolonization of the French
Alps". Wildlife Society Bulletin. 32 (4):
1195–1208. doi:10.2193/0091-
7648(2004)032[1195:HRTPSH]2.0.CO;
2.
177. Mertens, A., Schneider, H., 2005. What
is wrong with Romanian livestock
guarding dogs? A discussion.
Carnivore Damage Prevent. News 9:
9–14
178. Linnell, J. D. C. (2002). The Fear of
Wolves: A Review of Wolf Attacks on
Humans (PDF). Norsk Institutt for
Naturforskning (NINA). ISBN 978-82-
426-1292-2.
179. Mech & Boitani 2003, pp. 300–304.
180. Roosevelt, T. (1909). Hunting the grisly
and other sketches; an account of the
big game of the United States and its
chase with horse, hound, and rifle . G.
P. Putnam's sons. pp. 179–207.
181. Lopez 1978, p. 108.
182. Graves 2007, pp. 121–40.
183. Lopez 1978, pp. 159–60
184. Tucker, P. & Weide, B. (1998). "Can You
Turn a Wolf into a Dog" (PDF). Wild
Sentry. Archived from the original
(PDF) on 2015-12-08.

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
Enterprises. ISBN 978-1-55059-332-7.
Heptner, V. G.; Naumov, N. P. (1998).
Mammals of the Soviet Union Vol. II Part
1a, Sirenia 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 978-0-7432-4936-2.
Marvin, Garry (2012). Wolf . Reaktion
Books Ldt. ISBN 978-1-86189-879-1.
Mech, L. David (1981). The Wolf: The
Ecology and Behaviour of an Endangered
Species . University of Minnesota Press.
ISBN 978-0-8166-1026-6.
Mech, L. David; Boitani, Luigi, eds.
(2003). Wolves: Behaviour, Ecology and
Conservation . University of Chicago
Press. ISBN 978-0-226-51696-7.
Young, Stanley P.; Goldman, Edward A.
(1944). The Wolves of North America,
Part I . New York, Dover Publications,
Inc.
 Zimen, Erik (1981). The Wolf: His Place
in the Natural World . Souvenir Press.
ISBN 978-0-285-62411-5.

Further reading
Apollonio, Marco; Mattioli, Luca (2006).
Il Lupo in Provincia di Arezzo (in Italian).
Editrice Le Balze. ISBN 978-88-7539-
123-2.
Bibikov, D. I. (1985). "Volk:
Proiskhozhdenie, sistematika,
morfologia, ekologia [The Wolf: History,
Systematics, Morphology and Ecology]"
(in Russian). Nauka, Moscow, USSR.
ASIN B001A1TKK4 .
Coleman, Jon T. (2006). Vicious: Wolves
and Men in America. Yale University
Press. ISBN 978-0-300-11972-5.
Dutcher, Jim; Dutcher, Jamie (2003).
Wolves at Our Door: The Extraordinary
Story of the Couple Who Lived with
Wolves. William Andrew. ISBN 978-0-
7434-0049-7.
Fischer, Hank (1995). Wolf Wars .
Falcon. ISBN 978-1-56044-352-0.
Fuller, Todd K. (2004). Wolves of the
World. Voyageur Press. ISBN 978-0-
89658-640-6.
Grooms, Steve (1999). Return of the
Wolf. Northword Press. ISBN 978-1-
55971-717-5.
Hampton, Bruce (1997). The Great
American Wolf. Holt Paperbacks.
ISBN 978-0-8050-5528-3.
Harrington, Fred H.; Paquet, Paul C.
(1982). Wolves of the world: perspectives
of behavior, ecology, and conservation.
Simon & Schuster. ISBN 978-0-8155-
0905-9.
McIntyre, Rick (1996). A Society of
Wolves: National Parks and the Battle
over the Wolf. Voyageur Press.
ISBN 978-0-89658-325-2.
McNamee, Thomas (1998). The Return
of the Wolf to Yellowstone . Holt
Paperbacks. ISBN 978-0-8050-5792-8.
Mech, L. David (1966). Wolves of Isle
Royale . U.S. Department of the Interior,
Park Service.
Mech, L. David (1993). The Way of the
Wolf. Voyageur Press. ISBN 978-0-
89658-179-1.
Murie, Adolph (1944). Wolves of Mount
McKinley . U.S. Department of the
Interior, Park Service.
Musiani, Marco; Boitani, Luigi; Paquet,
Paul C. (2010). The World of Wolves:
New Perspectives on Ecology, Behaviour,
and Management. University of Calgary
Press. ISBN 978-1-55238-269-1.
 Nie, Martin (2003). Beyond Wolves: The
Politics of Wolf Recovery and
Management. University of Minnesota
Press. ISBN 978-0-300-11972-5.
Peterson, Rolf Olin (1977). Wolf Ecology
and Prey Relationships on Isle Royale .
National Park Service Scientific
Monograph Series.
Weaver, John (1978). Wolves of
Yellowstone . U.S. Department of the
Interior, Park Service.

External links

Wikimedia Commons has media

related to Canis lupus.
Wikispecies has information related to
Canis lupus

Wikiquote has quotations related to:

Wolf

Dog (mammal) at the Encyclopædia

Britannica
Gray wolf (mammal) at the
Encyclopædia Britannica
Wolf (mammal) at the Encyclopædia
Britannica
California Wolf Center
The International Wolf Center
 Staying Safe in Wolf Country , ADFG
(January 2009)
UK Wolf Conservation Trust
Watch Death of a Legend and Cry of the
Wild by Bill Mason

Retrieved from
"https://en.wikipedia.org/w/index.php?
title=Wolf&oldid=912651106"

Last edited 3 days ago by InternetAr…

Content is available under CC BY-SA 3.0 unless

otherwise noted.