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() Kingdom: Phylum: Class: Order: Suborder: Family: Genus: Species: C. lupus
Canis lupus, Numerous and disputed, see Historical (red + green) and modern
(green) range of wild subspecies of C. Lupus The gray wolf or grey wolf ( Canis
lupus), also known as the timber wolf or western wolf, is a native to the
wilderness and remote areas of and. It is the largest member of its family, with
males averaging 43–45 kg (95–99 lb) and females 36–38.5 kg (79–85 lb). Like the,
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. (3rd ed., 2005),
a standard reference work in zoology, recognises 38 of C. The gray wolf is the
second most member of the Canis, after the, as demonstrated by its adaptations
to hunting 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,, and, to produce fertile. It is the only species of Canis
to have a range encompassing both the and, and originated in Eurasia during the,
colonizing North America on at least three separate occasions during the.

It is a, travelling in consisting of a, accompanied by the pair's adult
offspring. The gray wolf is typically an throughout its range, with only and
posing a serious threat to it. Homer The Odyssey Robert Fagles Pdf Files. It
feeds primarily on large, though it also eats smaller animals, livestock,
carrion, and garbage. The gray wolf is one of the world's best-known and
most-researched animals, with probably more books written about it than any
other wildlife species.

It has a long history of association with humans, having been despised and
hunted in most communities because of its attacks on livestock, while conversely
being respected in some and 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. 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. Main article: The species Canis lupus was first recorded by in his
publication in 1758, with the Latin classification translating into the English
words 'dog wolf'.

The thirty-seven subspecies of Canis lupus are listed under the designated
common name of 'wolf' in third edition that was published in 2005. The is the (
Canis lupus lupus), also known as the common wolf.

The subspecies includes the,, and, but lists and C. Communis as of. However, the
classification of several as either species or subspecies has recently.
Evolution and relationship with the dog. Artist's impression of a The evolution
of the wolf occurred over a of 800,000 years, transforming the first wolf
specimen that is recognized as being similar to Canis lupus into today's,, and
gray wolf.

Ecological factors including habitat type, climate, prey specialization and
predatory competition will greatly influence the wolf's and cranio-dental.
Wolves went through a 20,000 (YBP), which indicates that many wolf populations
had gone extinct at a time that coincided with the and the expansion of modern
humans worldwide with their technology for capturing large game.

The domestic dog is the most widely abundant large carnivore, and a descendant
of one of those now-extinct wolf populations. Population structure In 2013, a
genetic study found that the wolf population in Europe was divided along a
north-south axis and formed five major clusters. Bodybuilding Anatomija Knjiga
O. Three clusters were identified occupying southern and central Europe in
Italy, the Dinaric-, the.

Another two clusters were identified occupying 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 formed the
Dinaric-Balkans cluster. Wolves from Finland, Latvia, Belarus, Poland and Russia
formed 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 were more similar to
the wolves from the than they were to wolves from north-central Europe. These
clusters may have been the result of expansion from glacial, an adaptation to
local environments, and landscape fragmentation and the killing of wolves in
some areas by humans.

In 2016, two genetic studies of North American gray wolves found that they
formed six – genetically and ecologically distinct populations separated from
other populations by their different type of habitat. These six wolf 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. These findings were in accord with
previous studies that precipitation influenced morphology and that vegetation
and habitat type influenced wolf differences. The local adaptation of a wolf
ecotype most likely reflects a wolf's preference to remain in the type of
habitat that it was born into. Hybridization with other Canis.

From Wildlife Park Kadzidlowo, Poland. The first is the product of a male wolf
and a female, while the latter comes from a female wolf and a male It was once
thought that and gray wolves did not voluntarily interbreed in the wild, though
they can produce fertile offspring.

In 2010, a study of 74 male lineages found that 5 of them originated from dog
ancestry, indicating that female wolves will breed with stray male dogs in the
wild. In North America, acquired their coloration from wolf-dog hybridization,
which occurred 10,000–15,000 years ago. Like pure wolves, hybrids breed once
annually, though their mating season occurs three months earlier, with pups
mostly being born in the winter period, thus lessening their chances of
survival. However, one genetic study undertaken in the showed that as many as
10% of dogs in the area, including, are first generation hybrids. The of hybrids
has proliferated in the USA, with 300,000 such animals being present there.

The gray wolf has interbred extensively with the producing a hybrid population
termed. Unlike the red and eastern wolf, the gray wolf does not readily
interbreed with coyotes. Nevertheless, coyote genetic markers have been found in
some wild isolated gray wolf populations in the southern United States. Gray
wolf have also been found in Texan coyote haplotypes.

In tests performed on a Texan canid of ambiguous species, mtDNA analysis showed
that it was a coyote, though subsequent tests revealed that it was a coyote–gray
wolf hybrid sired by a male Mexican gray wolf. In 2013, a captive breeding
experiment in Utah between gray wolves and western coyotes produced six hybrids
through, making this the first hybridization case between pure coyotes and. At
six months of age, the hybrids were closely monitored and were shown to display
both physical and behavioral characteristics from both species. Although has
never been observed, evidence of such occurrences was discovered through mtDNA
analysis on jackals in Bulgaria.

Although there is no genetic evidence of gray wolf-jackal hybridization in the
Caucasus Mountains, there have been cases where otherwise genetically pure
golden jackals have displayed remarkably gray wolf-like, to the point of being
mistaken for wolves by trained biologists. Wolf mandible diagram showing the
names and positions of the teeth. The gray wolf is the largest extant member of
the, excepting certain large breeds of domestic dog. Gray wolf weight and size
can vary greatly worldwide, tending to increase proportionally with latitude as
predicted by, with the large wolves of and Canada sometimes weighing 3–6 times
more than their and cousins.

On average, adult wolves measure 105–160 cm (41–63 in) in length and 80–85 cm
(31–33 in) in shoulder height. The tail measures 29–50 cm (11–20 in) in length.
The ears are 90–110 mm (3.5–4.3 in) in height, and the hind feet are 220–250 mm
(8.7–9.8 in). The mean body mass of the extant gray wolf is 40 kg (88 lb), with
the smallest specimen recorded at 12 kg (26 lb) and the largest at 80 kg
(180 lb).

Gray wolf weight varies geographically; 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). Females in any given wolf population typically weigh
5–10 lb (2.3–4.5 kg) less than males. Wolves weighing over 54 kg (119 lb) are
uncommon, though exceptionally large individuals have been recorded in Alaska,
Canada, and the forests of western. The heaviest recorded gray wolf in North
America was killed on 70 Mile River in east-central Alaska on July 12, 1939 and
weighed 79.4 kg (175 lb).

Compared to its closest wild cousins (the and ), the gray wolf is larger and
heavier, with a broader snout, shorter ears, a shorter torso and longer tail. It
is a slender, powerfully built animal with a large, deeply descending, a sloping
back and a heavily muscled neck. 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. The ears
are relatively small and triangular.

Females tend to have narrower muzzles and foreheads, thinner necks, slightly
shorter legs and less massive shoulders than males. The gray wolf usually
carries its head at the same level as the back, raising it only when alert. It
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), and 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 gray 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. Skull and dentition The gray wolf's head is large and heavy, with a
wide forehead, strong jaws and a long, blunt muzzle. The skull averages
230–280 mm (9.1–11.0 in) in length, and 130–150 mm (5.1–5.9 in) wide.

The teeth are heavy and large, being better suited to crushing bone than those
of other extant canids, though not as specialised as those found in. Its have a
flat chewing surface, but not to the same extent as the coyote, whose diet
contains more vegetable matter.

The gray wolf's jaws can exert a crushing pressure of perhaps 10,340 kPa
(1,500 psi) compared to 5,200 kPa (750 psi) for a. This force is sufficient to
break open most bones.

A study of the estimated bite force at the canine teeth of a large sample of
living and fossil mammalian predators when adjusted for the body mass found that
for mammals, the bite force at the canines (in /kilogram of body weight) was
greatest in the extinct dire wolf (163), then followed among the extant by the
four hypercarnivores that often prey on animals larger than themselves: the
African hunting dog (142), the gray wolf (136), the dhole (112), and the dingo
(108). Black and white-furred gray wolves The gray wolf has very dense and
fluffy winter fur, with short underfur and long, coarse. Most of the underfur
and some of the guard hairs are shed in the spring and grow back in the autumn
period. 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 down
to the. The winter fur is highly resistant to cold; wolves in northern climates
can rest comfortably in open areas at −40° by placing their muzzles between the
rear legs and covering their faces with their tail.

Wolf fur provides better insulation than dog fur, and does not collect ice when
warm breath is condensed against it. In warm climates, the fur is coarser and
scarcer than in northern wolves. Female wolves tend to have smoother furred
limbs than males, and generally develop the smoothest overall coats as they age.
Older wolves generally have more white hairs in the tip of the tail, along the
nose and on the forehead. The winter fur is retained longest in lactating
females, though with some hair loss around their nipples. Hair length on the
middle of the back is 60–70 mm (2.4–2.8 in). Hair length of 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).

Coat color ranges from almost pure white through various shades of blond, cream,
and ochre to grays, browns, and blacks, with variation in fur color tending to
increase in higher latitudes. Differences in coat color between sexes are
largely absent, though females may have redder tones. In North America inherited
the K b responsible for from past interbreeding with dogs, while the mutation
was found to be naturally occurring in wolves from Iran. Black specimens are
more common in North America than in Eurasia, with about half the wolves in
Yellowstone National Park being black. Gray wolf pack The gray wolf is a social
animal, whose basic social unit consists of a mated pair, accompanied by the
pair's adult offspring. The average consists of a family of 5–11 animals (1–2
adults, 3–6 juveniles and 1–3 yearlings), or sometimes two or three such
families, with exceptionally large packs consisting of up to 42 wolves being
known. In ideal conditions, the mated pair produces pups every year, with such
offspring typically staying in the pack for 10–54 months before dispersing.

Triggers for dispersal include the onset of and competition within the pack for
food. 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. 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.
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 (1–3 years of age) 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. During times of ungulate abundance
(migration, calving etc.), different wolf packs may temporarily join forces.
Wolves are highly animals, and generally establish territories far larger than
they require to survive in order to assure 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
or when the pups reach the age of 6 months, thus having the same nutritional
needs as adults. 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 km 2 (14 sq mi), in which they spend 50% of
their time.

Prey density tends to be much higher in the territory's surrounding areas,
though wolves tend to avoid hunting in the fringes of their range unless
desperate, because of the possibility of fatal encounters with neighboring
packs. The smallest territory on record was held by a pack of six wolves in
northeastern Minnesota, which occupied an estimated 33 km 2 (13 sq mi), while
the largest was held by an Alaskan pack of ten wolves encompassing a 6,272 km 2
(2,422 sq mi) area. Wolf packs are typically settled, and usually only leave
their accustomed ranges during severe food shortages. Wolves defend their
territories from other packs through a combination of, direct attacks and
howling (see ). Scent marking is used for territorial advertisement, and
involves urination, defecation and ground scratching. Scent marks are generally
left every 240 m (260 yd) throughout the territory on regular travelways and
junctions.

Such markers can last for 2–3 weeks, and are typically placed near rocks,
boulders, trees, or the skeletons of large animals. Territorial fights are among
the principal causes of wolf mortality, with one study concluding that 14–65% of
wolf deaths in Minnesota and the were due to predation by other wolves.
Reproduction and development. Gray wolves mating The gray wolf is generally,
with mated pairs usually remaining together for life. Upon the death of one
mated wolf, pairs are quickly re-established. Since males often predominate in
any given wolf population, unpaired females are a rarity. If a dispersing male
gray wolf is unable to establish a territory or find a mate, he mates with the
daughters of already established breeding pairs from other packs.

Such gray wolves are termed ' wolves' and, unlike males from established packs,
they do not form with the females they mate with. Some gray wolf packs may have
multiple breeding females this way, as is the case in. Gray wolves also
practice, in which a wolf pair may adopt the pup or pups of another.

This might take place if the original parents die or are for some reason
separated from them. In addition to heterosexual behavior, homosexual behavior
has been observed in gray wolves. Male gray wolves often when the highest
ranking female in the pack comes into heat. Illustration of various gray wolf
growth stages The age of first breeding in gray 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. This is further demonstrated by the fact that captive wolves
have been known to breed as soon as they reach 9–10 months, while the youngest
recorded breeding wolves in the wild were 2 years old. Females are capable of
producing pups every year, with one annually being the average. Unlike the
coyote, the gray wolf never reaches.

Typically occurs in late winter, with older, multiparous females entering estrus
2–3 weeks earlier than younger females. 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.

Old females usually in the den of their previous litter, while younger females
typically den near their birthplace. The lasts 62–75 days, with pups usually
being born in the summer period. Wolves bear relatively large pups in small
litters compared to other canid species. The average litter consists of 5–6
pups, with litter sizes tending to increase in areas where prey is abundant,
though exceptionally large litters of 14–17 pups occur only 1% of the time. Pups
are usually born in spring, coinciding with a corresponding increase in prey
populations.

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 9–12 days. The
milk canines erupt after one month. Pups first leave the den after 3 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 3–4
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.

Wolf pups begin play fighting at the age of 3 weeks, though unlike young foxes
and coyotes, their bites are inhibited. Actual fights to establish hierarchy
usually occur at 5–8 weeks of age. This is in contrast to young foxes and
coyotes, which may begin fighting even before the onset of play behavior.

By autumn, the pups are mature enough to accompany adults on hunts for large
prey. Gray wolf trotting. The gray wolf generally places its hind paws in the
tracks made by the front paws. 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, and
unaided. The gray wolf's is relatively weakly developed when compared to that of
some 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 is acute enough to be able to hear up to a
frequency of 26 kHz, which is sufficient to register the fall of leaves in the
autumn period. A gray 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. • The stalk: The wolves
attempt to conceal themselves as they approach. 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. •
The encounter: Once the prey detects the wolves, it can either approach the
wolves, stand its ground, or flee. Large prey, such as moose,, 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. If the targeted animal stands its
ground, the wolves either ignore it, or try to intimidate it into running. • 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. 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. •
The chase: A continuation of the rush, the wolves attempt to catch up with their
prey and kill it. 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). 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. Killing a 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 area, causing massive. 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. With medium-sized prey such as or, wolves kill by biting the throat,
severing nerve tracks and the, thus causing the animal to die within a few
seconds to a minute.

With small, -like prey, wolves leap in a high arc and immobilize it with their
forepaws. When prey is vulnerable and abundant, wolves may occasionally. 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) or during the denning
period, when wolves require a ready supply of meat when denbound.

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. Two gray wolves eating a white-tailed deer 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. 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.

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,, and lining.
The and are eaten once they are exposed, followed by the. A single wolf can eat
15–19% of its body weight in a single feeding.

See also: Visual The gray wolf's expressive behavior is more complex than that
of the coyote and golden jackal, as necessitated by its group living and hunting
habits. While less gregarious canids generally possess simple repertoires of
visual signals, wolves have more varied signals that subtly inter grade in
intensity. When neutral, the legs are not stiffened, the tail hangs down
loosely, the face is smooth, the lips untensed, and the ears point in no
particular direction. Postural communication in wolves consists of a variety of
facial expressions, tail positions and. Aggressive, or self-assertive wolves are
characterized by their slow and deliberate movements, high body and raised,
while submissive ones carry their bodies low, sleeken their fur and lower their
ears and tail.

When a breeding male encounters a subordinate family member, it may stare at it,
standing erect and still with the tails horizontal to its spine. Two forms of
submissive behavior are recognized: passive and active. Passive submission
usually occurs as a reaction to the approach of a, and consists of the
submissive wolf lying partly on its back and allowing the dominant wolf to sniff
its. Active submission occurs often as a form of greeting, and involves the
submissive wolf approaching another in a low posture, and licking the other
wolf's face. When wolves are together, they commonly indulge in behaviors such
as nose pushing, jaw wrestling, cheek rubbing and facial licking. The mouthing
of each other's muzzles is a friendly gesture, while clamping on the muzzle with
bared teeth is a dominance display. Similar to humans, gray wolves have facial
color patterns in which the gaze direction can be easily identified, although
this is often not the case in other canid species.

In 2014, a study compared the facial color pattern across 25 canid species. The
results suggested that the facial color pattern of canid species is related to
their gaze communication, and that especially gray wolves use the gaze signal in
conspecific communication. Facial expressions (, 1952). Bottom to top:
increasing fear (ears back); left to right: increasing aggression (snarl); top
right: maximum of both. Problems playing this file? Gray 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 unfamiliar territory and to
communicate across great distances. Wolf howls can under certain conditions be
heard over areas of up to 130 km 2 (50 sq mi).

Wolf howls are generally indistinguishable from those of large dogs. Male wolves
give voice through an, passing to a deep with a stress on ' O', while females
produce a modulated nasal with stress on ' U'. Pups almost never howl, while
yearling wolves produce howls ending in a series of dog-like yelps. Howling
consists of a fundamental frequency that may lie between 150 and 780 Hz, and
consists of up to 12 harmonically related overtones. The pitch usually remains
constant or varies smoothly, and may change direction as many as four or five
times.

Howls used for calling pack mates to a kill are long, smooth sounds similar to
the beginning of the cry of a. When pursuing prey, they emit a higher pitched
howl, vibrating on two notes. When closing in on their prey, they emit a
combination of a short bark and a howl. When howling together, wolves harmonize
rather than chorus on the same note, thus creating the illusion of there being
more wolves than there actually are.

Lone wolves typically avoid howling in areas where other packs are present.
Wolves from different geographic locations may howl in different fashions: the
howls of European wolves are much more protracted and melodious than those of
North American wolves, whose howls are louder and have a stronger emphasis on
the first syllable. The two are however, as North American wolves have been
recorded to respond to European-style howls made by biologists.

Other vocalisations of wolves are usually divided into three categories: growls,
barks and whines. Barking has a fundamental frequency between 320–904 Hz, and is
usually emitted by startled wolves. Wolves do not bark as loudly or continuously
as dogs do, but bark a few times and retreat from perceived danger. Growling has
a fundamental frequency of 380–450 Hz, and is usually emitted during food
challenges. Pups commonly growl when playing. One variation of the howl is
accompanied by a high pitched whine, which precedes a lunging attack. Whining is
associated with situations of anxiety, curiosity, inquiry and intimacy such as
greeting, feeding pups and playing.

Gray wolf marking its territory with urine is probably the wolf's most acute
sense, and plays a fundamental role in communication. The wolf has a large
number of on the face, lips, back, and between the toes. The odor produced by
these glands varies according to the individual wolf's and diet, giving each a
distinct 'odor fingerprint'. A combination of apocrine and on the feet allows
the wolf to deposit its scent whilst scratching the ground, which usually occurs
after urine marking and defecation during the.

The present on the guard hairs from the wolf's back have clusters of apocrine
and at their bases. As the skin on the back is usually folded, this provides a
for bacterial propagation around the glands. During piloerection, the guard
hairs on the back are raised and the skin folds spread, thus releasing scent.
The may play a role in expressing aggression, as combative wolves raise the base
of their tails whilst drooping the tip, thus positioning the at the highest
point.

The wolf possesses a pair of beneath the rectum, which contain both apocrine and
sebaceous glands. The components of anal sac secretions vary according to season
and gender, thus indicating that the secretions provide information related to
and reproductive state. The secretions of the may advertise hormonal condition
or social position, as dominant wolves have been observed to stand over
subordinates, apparently presenting the genital area for investigation, which
may include genital licking. During the breeding season, female wolves secrete,
which communicate the females' reproductive state, and can be detected by males
from long distances. Urine marking is the best-studied means of in wolves. Its
exact function is debated, though most researchers agree that its primary
purpose is to establish boundaries.

Wolves urine mark more frequently and vigorously in unfamiliar areas, or areas
of intrusion, where the scent of other wolves or canids is present. So-called
(RLU) is more common in male wolves than in females, and may serve the purpose
of maximizing the possibility of detection by conspecifics, as well as reflect
the height of the marking wolf. Only dominant wolves typically use RLU, with
subordinate males continuing to use the juvenile standing posture throughout
adulthood. RLU 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.

Ecology Habitat. Gray wolf carrying hindquarter, The gray wolf is a habitat
generalist, and can occur in,, and. Habitat use by gray wolves is strongly
correlated with the abundance of prey, snow conditions, absence or low livestock
densities, road densities, human presence and. In cold climates, the gray 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 point where the pads come in contact with ice and snow.

Gray wolves use different places for their diurnal rest: places with cover are
preferred during cold, damp and windy weather, while wolves in dry, calm and
warm weather readily rest in the open. During the autumn-spring period, when
wolves are more active, they willingly lie out in the open, whatever their
location. Actual 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 1–3
openings. The den is usually constructed not more than 500 m (550 yd) away from
a water source, and typically faces southwards, thus ensuring enough sunlight
exposure, keeping the denning area relatively snow free. 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 and. 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, paved and. Diet Globally, gray wolf diet is predominantly composed of
large (240–650 kg (530–1,430 lb)) and medium-sized (23–130 kg (51–287 lb)) wild,
with local population variations due to the mix of wild ungulates, smaller prey
and domestic species consumed.

All terrestrial mammalian social predators feed predominantly on terrestrial
herbivorous mammals that have a body mass similar to that of the combined mass
of the social group members. The gray wolf generally specializes in predating on
the vulnerable individuals of large prey, with pack of timber wolves capable of
bringing down a 500 kg (1,100 lb) moose.

Only takes a few hours, thus wolves can feed several times in one day, making
quick use of large quantities of meat. Although wolves primarily feed on medium
to large sized, they are not fussy eaters. Smaller sized animals that may
supplement the diet of wolves include,,,,,,,, and other, as well as. They
frequently eat and their. When such foods are insufficient, they prey on,,,
rarely and large as available. In times of scarcity, wolves readily eat,
visiting cattle burial grounds and. Is not uncommon in wolves: during harsh
winters, packs often attack weak or injured wolves, and may eat the bodies of
dead pack members.

Wolf packs in hunt on the coastline and some wolf packs in Alaska and Western
Canada have been observed to feed on. Are rarely, but occasionally preyed upon.
Other occasionally taken by wolves include in Nepal and in Saudi Arabia. In
Eurasia, many gray wolf populations are forced to subsist largely on and in
areas with dense human activity, though wild ungulates such as,, and are still
the most important food sources in Russia and the more mountainous regions of
Eastern Europe. Other prey species include,,,,,,,, and. 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. Animals preferred as prey by North American wolves
include moose,,,,,,, and.

Wolves supplement their diet with fruit and vegetable matter. They willingly eat
the of,,, and. Other fruits include, and. They readily visit fields during the
summer months. A well-fed wolf stores under the skin, around the heart,
intestines, kidneys, and bone marrow, particularly during the autumn and winter.
Gray wolves confronting over carcass (1919), Gray wolves typically dominate
other species in areas where they both occur.

In North America, incidents of gray wolves killing 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. Near identical interactions have been observed in Eurasia
between gray wolves and, with the latter's numbers being comparatively small in
areas with high wolf densities.

Gray wolves are the most important predator of, killing large numbers of them in
the spring and summer periods. Wolves also kill, and, usually in disputes over
carcasses, sometimes eating them. In Asia, they may compete with, though there
is at least one record of a lone wolf associating with a pair of dholes in.
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.

Wolf interactions with are much rarer than with brown bears, because of
differences in habitat preferences. The majority of 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 black bears
in their dens and kill them without eating them. Unlike brown bears, black bears
frequently lose against wolves in disputes over kills. While encounters with
brown and black bears appear to be common, are rarely encountered by wolves,
though there are two records of wolf packs killing polar bear cubs. Wolves also
kill the cubs of. Wolves may encounter 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-to-one,
hyenas dominate wolves, and may prey on them, but wolf packs can drive off
single or outnumbered hyenas.

However, there was a case of a female striped hyena dominating 12 Arabian
wolves. Two cases are known however from southern Israel, where wolves and
striped hyenas associated closely with each other in an apparently amicable way.
Large wolf populations limit the numbers of small to medium-sized.

Wolves encounter along portions of the 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 kills. They have been reported
killing mothers and their kittens. Wolves hunt, and may scavenge from kills.
Wolves may also reduce populations.

Wolves may kill lynxes by running them down, or killing them before they can
escape into trees. Similar reports of encounters between wolves and have been
documented. Leftovers of wolf kills are sometimes scavenged. Wolverines usually
wait until the wolves are done feeding, but have been known to drive away wolves
from kills. However, there have been confirmed reports of wolf packs killing
wolverines. Other than humans, appear to be the only serious predators of
wolves.

Wolf and tiger interactions are well documented in, where tigers depress wolf
numbers, either to the point of or to such low numbers as to make them a
functionally insignificant component of the. Wolves appear capable of escaping
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. Range and conservation. See also:,, and The gray wolf
was once one of the world's most widely distributed mammals, living throughout
the northern hemisphere north of 15°N latitude in North America and 12°N in
India. However, deliberate human persecution has reduced the species' range to
about one third, because of livestock predation and fear over attacks on humans.

The species is now extinct in much of, in Mexico and much of the United States.
In modern times, the gray wolf occurs mostly in wilderness and remote areas,
particularly in Canada, and northern USA,, and from about 75°N to 12°N. 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 species. Despite these threats, because of the gray wolf's relatively
widespread range and stable population, it is classified as by the. Europe
Decline.

Of a gray wolf,, France Although wolf-dog hybridization in Europe has raised
concern among conservation groups fearing for the gray wolf's purity, genetic
tests show that introgression of dog genes into European gray wolf populations
does not pose a significant threat. Also, as wolf and dog do not fully coincide,
the likelihood of wild wolves and dogs mating and producing surviving offspring
is small. The extermination of 's wolves first became an organized effort during
the, and continued until the late 1800s. In, wolf persecution was enforced by,
and the last wolf was killed in the early sixteenth century during the reign of.
Wolves lasted longer in, where they sheltered in vast tracts of forest, which
were subsequently burned down.

Wolves managed to survive in the forests of and until 1684. The extirpation of
wolves in followed a similar course, with the last wolf believed to have been
killed in 1786. A wolf bounty was introduced in in 1647, after the extermination
of moose and reindeer forced wolves to feed on livestock. The extirpated wolves
in northern Sweden in organized drives. By 1960, few wolves remained in Sweden,
because of the use of snowmobiles in hunting them, with the last specimen being
killed in 1966. The gray wolf was exterminated in Denmark in 1772 and Norway's
last wolf was killed in 1973. The species was decimated in 20th century Finland,
despite regular dispersals from Russia.

The gray wolf was only present in the eastern and northern parts of Finland by
1900, though its numbers increased after. In, wolves were dramatically reduced
in number during the early nineteenth century, because of organized hunts and
reductions in ungulate populations. In, the last wolf was killed in 1847, and
had disappeared from the regions by 1899 In, wolves were extinct in the
twentieth century; they are naturally coming back from Italy since the 1990s. In
1934, became the first state in modern history to place the wolf under
protection, though the species was already extirpated in Germany at this point.
The last free-living wolf to be killed on the soil of present-day before 1945
was the so-called ', which was shot near, (then ) in 1904.

Today, wolves have returned to the area. Wolf hunting in France was first
institutionalized by between 800–813, when he established the louveterie, a
special corps of wolf hunters. The louveterie was abolished after the in 1789,
but was re-established in 1814. In 1883, up to 1,386 wolves were killed, with
many more by poison.

Last wolf to be killed in central Finland (, 1911) In, wolves were never fully
exterminated, because of the area's contiguity with Asia and its large forested
areas. However, Eastern European wolf populations were reduced to very low
numbers by the late nineteenth century. Wolves were extirpated in during the
first decade of the twentieth century and, by the mid-twentieth century, could
only be found in a few forested areas in eastern Poland. Wolves in the eastern
benefitted from the region's contiguity with the and large areas of plains,
mountains and farmlands.

Wolves in Hungary occurred in only half the country around the start of the 20th
century, and were largely restricted to the. Wolf populations in Romania
remained largely substantial, with an average of 2,800 wolves being killed
annually out of a population of 4,600 from 1955–1965. An all-time low was
reached in 1967, when the population was reduced to 1,550 animals. The
extermination of wolves in Bulgaria was relatively recent, as a previous
population of about 1,000 animals in 1955 was reduced to about 100–200 in 1964.
In Greece, the species disappeared from the southern in 1930. Despite periods of
intense hunting during the eighteenth century, wolves never disappeared in the
western Balkans, from to the. Organized persecution of wolves began in
Yugoslavia in 1923, with the setting up of the Wolf Extermination Committee
(WEC) in,.

The WEC was successful in reducing wolf numbers in the. In, wolf extermination
was not as complete as in Northern Europe, because of greater cultural tolerance
of the species. Wolf populations only began declining in the in the early
19th-century, and was reduced by a half of its original size by 1900. Wolf
bounties were regularly paid in Italy as late as 1950.

Wolves were exterminated in the by 1800, and numbered only 100 by 1973,
inhabiting only 3–5% of their former Italian range. Range of Canis lupus in
Europe, based on Canis lupus European regional assessment, IUCN and Large
Carnivore Initiative for Europe. Lime color stands for population lower than
800, while green is above 800. The recovery of European wolf populations began
after the 1950s, when traditional pastoral and rural economies declined and thus
removed the need to heavily persecute wolves.

By the 1980s, small and isolated wolf populations expanded in the wake of
decreased human density in rural areas and the recovery of wild prey
populations. The gray wolf has been fully protected in Italy since 1976, and now
holds a population of over 1,269-1,800.

Italian wolves entered France's in 1993, and at least fifty wolves were
discovered in the western in 2000. By 2013 the 250 wolves in the imposed a
significant burden on traditional and husbandry with a loss of over 5,000
animals in 2012. There are approximately 2,000 wolves inhabiting the, of which
150 reside in northeastern. In, the species occurs in,, and.

Although hundreds of Iberian wolves are illegally killed annually, the
population has expanded south across the river and east to the and. In, Germany
In 1978, wolves began recolonising central Sweden after a twelve-year absence,
and have since expanded into southern Norway. As of 2005, the total number of
Swedish and Norwegian wolves is estimated to be at least one hundred, including
eleven. The gray wolf is fully protected in Sweden and partially controlled in
Norway. The Scandinavian wolf populations owe their continued existence to
neighbouring Finland's contiguity with the, which houses a large population of
wolves. Wolves in Finland are protected only in the southern third of the
country, and can be hunted in other areas during specific seasons, though
poaching remains common, with 90% of young wolf deaths being due to human
predation, and the number of wolves killed exceeds the number of hunting
licenses, in some areas by a factor of two. Furthermore, the decline in the
moose populations has reduced the wolf's food supply.

Since 2011, the Netherlands, Belgium and Denmark have also reported wolf
sightings presumably by natural migration from adjacent countries. In 2016, a
female wolf tracked 550 kilometers from a region southwest of Berlin to settle
in Jutland, Denmark where male wolves had been reported in 2012 for the first
time in 200 years.

Wolves have also commenced breeding in Lower Austria's Waldviertel region for
the first time in over 130 years. Wolf populations in Poland have increased to
about 800–900 individuals since being classified as a game species in 1976.
Poland plays a fundamental role in providing routes of expansion into
neighbouring countries. In the east, its range overlaps with populations in
Lithuania,,, and. A population in western Poland expanded into eastern Germany
and in 2000 the first pups were born on German territory.

In 2012, an estimated 14 wolf packs were living in Germany (mostly in the east
and north) and a pack with pups has been sighted within 15 miles of; the number
increased to 46 packs in 2016. The gray wolf is protected in Slovakia, though an
exception is made for wolves killing livestock. A few Slovakian wolves disperse
into the Czech Republic, where they are afforded full protection.

Wolves in Slovakia, Ukraine and may disperse into Hungary, where the lack of
cover hinders the buildup of an autonomous population. Although wolves have
special status in Hungary, they may be hunted with a year-round permit if they
cause problems. Romania has a large population of wolves, numbering 2,500
animals. The wolf has been a protected animal in Romania since 1996, although
the law is not enforced. The number of wolves in and is largely unknown, despite
the importance the two countries have in linking wolf populations from Greece to
those of and Croatia.

Although protected, sometimes wolves are still illegally killed in Greece, and
their future is uncertain. Wolf numbers have declined in Bosnia and Herzegovina
since 1986, while the species is fully protected in neighbouring Croatia and.

Asia Historical range and decline. Monument to the last Japanese gray wolf,,
Japan During the 19th century, gray wolves were widespread in many parts of the
east and west of the. However, they decreased considerably in number between
1964 and 1980, largely because of persecution by farmers. The species was not
considered common in northern and central during the 19th century, with most
early publications involving animals either from southwestern, northern rocky
areas bordering, or areas surrounding. The gray wolf's range in the encompassed
nearly the entire territory of the country, being absent only on the,,, and the,
and.

The species was exterminated twice in, once after the, and again after.
Following the two world wars, Soviet wolf populations peaked twice. 30,000
wolves were harvested annually out of a population of 200,000 during the 1940s,
with 40,000–50,000 harvested during peak years. Soviet wolf populations reached
a low around 1970, disappearing over much of.

The population increased again by 1980 to about 75,000, with 32,000 being killed
in 1979. Wolf populations in northern declined during the 1940s, primarily
because of poaching of, the wolf's main prey. In, wolves were heavily persecuted
because of their attacks on sheep, goats and children. In 1876, 2,825 wolves
were bountied in the (NWP) and. By the 1920s, wolf extermination remained a
priority in the NWP and. Overall, over 100,000 wolves were killed for bounties
in British India between 1871 and 1916.

Wolves in Japan became extinct during the period, an extermination known as
ōkami no kujo. The wolf was deemed a threat to ranching, which the Meiji
government promoted at the time, and targeted via a bounty system and a direct
chemical extermination campaign inspired by the similar contemporary American
campaign. The last Japanese wolf was a male killed on January 23, 1905 near
Washikaguchi (now called Higashi Yoshiro). The now extinct Japanese wolves were
descended from large Siberian wolves, which colonized the and Japan, before it
separated from mainland Asia, 20,000 years ago during the Pleistocene. During
the, the widened and isolated from, thus causing climatic changes leading to the
extinction of most large bodied ungulates inhabiting the archipelago. Japanese
wolves likely underwent a process of 7,000–13,000 years ago in response to these
climatological and ecological pressures.

(formerly native to Hokkaidō) was significantly larger than its southern cousin,
as it inhabited higher elevations and had access to larger prey, as well as a
continuing genetic interaction with dispersing wolves from Siberia. Modern
range. Gray wolf near, Turkey.

Although Turkish wolves have no legal protection, they may number about 7000
individuals. In 2008, an authoritative reference stated that the gray wolf could
be found across mainland China. In 2017, a comprehensive study found that the
gray wolf was present across all of mainland China, both in the past and today.
It exists in southern China, which refutes claims made by some researchers in
the that the wolf had never existed in southern China.

There is little reliable data on the status of wolves in the, save for those in
Israel and, though their numbers appear to be stable, and are likely to remain
so. Israel's conservation policies and effective law enforcement maintain a
moderately sized wolf population, which radiates into neighbouring countries,
while Saudi Arabia has vast tracts of desert, where about 300–600 wolves live
undisturbed.

The wolf survives throughout most of its historical range in Saudi Arabia,
probably because of a lack of pastoralism and abundant human waste. Turkey may
play an important role in maintaining wolves in the region, because of its
contiguity with Central Asia. The mountains of Turkey have served as a refuge
for the few wolves remaining in. A small wolf population occurs in the, and is
well protected by the military activities there.

Wolves living in the southern desert are contiguous with populations living in
the Egyptian and. Throughout the Middle East, the species is only protected in
Israel. Elsewhere, it can be hunted year-round. Little is known of current wolf
populations in Iran, which once occurred throughout the country in low densities
during the mid-1970s. The northern regions of and Pakistan are important
strongholds for the wolf.

It has been estimated that there are about 300 wolves in approximately 60,000 km
2 (23,000 sq mi) of in northern India, and 50 more in. Overall, India supports
about 800-3,000 wolves, scattered among several remnant populations. Although
protected since 1972, Indian wolves are classed as endangered, with many
populations lingering in low numbers or living in areas increasingly used by
humans.

Although present in and, there is no information of wolves occurring there. Wolf
populations throughout and are largely unknown, but are estimated in the
hundreds of thousands based on annual harvests. Since the, continent-wide
extermination of wolves has ceased, and wolf populations have increased to about
25,000–30,000 animals throughout the. In China and, wolves are only protected in
reserves. Mongolian populations have been estimated at 10,000–30,000, while the
status of wolves in China is more fragmentary. The north has a declining
population of an estimated 400 wolves, while and hold about 10,000 and 2,000
respectively.

North America. 100 lb (45 kg) gray wolf killed in Montana, 1928. Before they
were extirpated around 1930, Montana's wolves could be very large. Wolves
recolonized the state from Canada beginning in the 1970s. Historical range and
decline Originally, the gray wolf occupied all of North America north of about
20°N. It occurred all over the mainland, save for the southeastern United
States, west of the, and the tropical and subtropical areas of Mexico. Large
continental islands occupied by wolves included,, southeastern Alaskan islands,
and throughout the and.

While Lohr and Ballard postulated that the gray wolf had never been present on,:
392 analysis of references to the island's native fauna in unpublished and
published historical records has found that gray wolves were resident there at
the time of the first French settlement in 1720. In his 6 November 1721 letter
to the French Minister of the Marine, Louis Denys de La Ronde reported that the
island was home to wolves 'of a prodigious size', and sent a wolf pelt back to
France to substantiate his claim. As the island was cleared for settlement, the
gray wolf population may have been extirpated, or relocated to the mainland
across the winter ice: the few subsequent wolf reports date from the
mid-nineteenth century and describe the creatures as transient visitors from
across the.: 386 The decline of North American wolf populations coincided with
increasing human populations and the expansion of agriculture. By the start of
the 20th century, the species had almost disappeared from the eastern USA,
excepting some areas of the and the northwestern. In Canada, the gray wolf was
extirpated in and between 1870 and 1921, and in Newfoundland around 1911. It
vanished from the southern regions of and between 1850 and 1900. The gray wolf's
decline in the prairies began with the extermination of the and other ungulates
in the 1860s–70s.

From 1900–1930, the gray wolf was virtually eliminated from the western USA and
adjoining parts of Canada, because of intensive predator control programs aimed
at eradicating the species. The gray wolf was exterminated by federal and state
governments from all of the USA by 1960, except in Alaska and northern. The
decline in North American wolf populations was reversed from the 1930s to the
early 1950s, particularly in southwestern Canada, because of expanding ungulate
populations resulting from improved regulation of big game hunting. This
increase triggered a resumption of wolf control in western and northern Canada.

Thousands of wolves were killed from the early 1950s to the early 1960s, mostly
by poisoning. This campaign was halted and wolf populations increased again by
the mid-1970s. Modern range. Captive The species' modern range in North America
is mostly confined to Alaska and Canada, with populations also occurring in
northern Minnesota, northern and 's, and small portions of,, northern, and. A
functional wolf population should exist in California by 2024 according to
estimates by state wildlife officials.

Canadian wolves began to naturally re-colonize northern Montana around in 1979,
and the first wolf den in the western U.S. In over half a century was documented
there in 1986. The wolf population in northwest Montana initially grew as a
result of natural reproduction and dispersal to about 48 wolves by the end of
1994. From 1995–1996, wolves from and were relocated to and Idaho. In addition,
the Mexican wolf ( Canis lupus baileyi) was reintroduced to and in 1998.

The gray wolf is found in approximately 80% of its historical range in Canada,
thus making it an important stronghold for the species. Canada is home to about
52,000–60,000 wolves, whose legal status varies according to.

Residents may hunt wolves without restriction, and some provinces require
licenses for residents to hunt wolves while others do not. In, wolves on private
land may be baited and hunted by the landowner without requiring a license, and
in some areas, wolf hunting bounty programs exist. Large-scale wolf population
control through poisoning, trapping and aerial hunting is also presently
conducted by government-mandated programs in order to support populations of
endangered prey species such as. In Alaska, the gray wolf population is
estimated at 6,000–7,000, and can be legally harvested during hunting and
trapping seasons, with and other restrictions. As of 2002, there are 250 wolves
in 28 packs in Yellowstone, and 260 wolves in 25 packs in Idaho.

The gray wolf received (ESA) protection in Minnesota, Wisconsin, and Michigan in
1974, and was re-classed from to in 2003. Reintroduced in Arizona and New Mexico
are protected under the ESA and, as of late 2002, number 28 individuals in eight
packs. A female wolf shot in 2013 in by a hunter was the first gray wolf seen in
Kentucky in modern times. DNA analysis by Fish and Wildlife laboratories showed
genetic characteristics similar to those of wolves in the Great Lakes Region.
Main article: The wolf is a common motif in the foundational mythologies and
cosmologies of peoples throughout and (corresponding to the historical extent of
the habitat of the gray wolf). The obvious attribute of the wolf is its nature
of a, and correspondingly it is strongly associated with danger and destruction,
making it the symbol of the on one hand, and that of the on the other.

The modern trope of the is a development of this. The wolf holds great
importance in the cultures and religions of the nomadic peoples, both of the and
of the. In many cultures, the identification of the warrior with the wolf ()
gave rise to the notion of, the mythical or ritual identification of man and
wolf. In fable and literature. (1883), featured wolves in several of his,
playing on the concerns of 's settled, sheep-herding world. His most famous is
the fable of, which is directed at those who knowingly raise false alarms, and
from which the idiomatic phrase ' 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.
Isengrim the wolf, a character first appearing in the 12th-century Latin poem,
is a major character in the 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. The tale of, first written in
1697 by, 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, capable of imitating human
speech. The hunting of wolves, and their attacks on humans and livestock feature
prominently in, and are included in the works of,,,,, and others.

Tolstoy's and Chekhov's Peasants both feature scenes in which wolves are hunted
with hounds and borzois. 's largely fictional 1963 memoir was the first positive
portrayal of wolves in popular literature, and is largely considered to be the
most popular book on wolves, having been adapted into a 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. In heraldry and symbolism.
Of the Elvange family, featuring a wolf rampant The wolf is a frequent in
English armory. It is illustrated as a on the of,, and, and can be found on the
of Lovett and the vast majority of the and Lows. The demi-wolf is a common,
appearing in the arms and crests of members of many families, including that of
the, whose crest depicts a demi-wolf holding a crown in its paws, in reference
to the assistance the family gave to during the. Wolf heads are common in,
particularly in the coats of and.

The wolf is the most common animal in, and is often depicted as carrying a lamb
in its mouth, or across its back. The wolf is featured on the of the, the and
the.

In modern times, the wolf is widely used as an emblem for military and
paramilitary groups. It is the unofficial symbol of the, and serves as the logo
of the Turkish. During the, several units adopted the wolf as their symbol,
including the White Wolves and the. Waiting for a Chinook (c.

1900), 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. Some
nations help offset economic losses to wolves through compensation programmes or
state insurance. Domesticated animals are easy prey for wolves, as they have
evolved under constant human protection, and are thus unable to defend
themselves very well. 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 incidences are rare in North America, where
healthy populations of wild prey have been largely restored. The majority of
losses occur during the summer grazing period, with untended livestock in remote
pastures being the most vulnerable to wolf predation.

The most frequently targeted livestock species are (Europe), (northern
Scandinavia), (India), (Mongolia), and (North America). The number of animals
killed in single attacks varies according to species: most attacks on cattle and
horses result in one death, while turkeys, sheep and domestic reindeer may be
killed in. Wolves mainly attack livestock when the animals are grazing, though
they occasionally break into fenced enclosures. 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, weight loss
and a decrease in meat quality. Conflicts with dogs Being the most abundant
carnivores, feral and have the greatest potential to compete with wolves. A
review of the studies in the competitive effects of dogs on carnivores did not
mention any research on competition between dogs and wolves. 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.

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. 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. The most frequently killed hunting breeds in Scandinavia are, 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 are more likely to survive wolf attacks because of their better
ability to defend themselves. 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. Dogs that are employed to guard sheep
help to mitigate human–wolf conflicts, and are often proposed as one of the
non-lethal tools in the conservation of wolves.

Shepherd dogs are not particularly aggressive, but they can disrupt potential
wolf predation by displaying what is to the wolf ambiguous behaviors, such as
barking, social greeting, invitation to play or aggression. The historical use
of shepherd dogs across Eurasia has been effective against wolf predation,
especially when confining sheep in the presence of several livestock-guarding
dogs. However, shepherd dogs are sometimes killed by wolves. Wolf predation on
humans. Child snatched by a wolf (1914), The fear of wolves has been pervasive
in many societies, though humans are not part of the wolf's natural prey. How
wolves react to humans depends largely on their prior experience with people:
wolves lacking any negative experience of humans, or which are food-conditioned,
may show little fear of people.

Although wolves may react aggressively under provocation, such attacks are
mostly limited to quick bites on extremities, and the attacks are not pressed.
Predatory attacks (attacks by wolves treating humans as food) may be preceded by
a long period of, 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 or
berry and mushroom picking) increase, though cases of non-rabid wolf attacks in
winter have been recorded in, and oblasts, and. Also, wolves with pups
experience greater food stresses during this period.

The majority of victims of predatory wolf attacks are children under the age of
18 and, in the rare cases where adults are killed, the victims are almost always
women. Cases of 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, and. 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,
with bites from rabid wolves being 15 times more dangerous than those of rabid
dogs. Rabid wolves usually act alone, travelling large distances and often
biting large numbers of people and domestic animals.

Most rabid wolf attacks occur in the spring and autumn periods. Unlike with
predatory attacks, the victims of rabid wolves are not eaten, and the attacks
generally only occur on a single day. Also, the victims are chosen at random,
though the majority of cases involve adult men. In the half-century up to 2002,
there were eight fatal attacks in Europe and Russia, and more than 200 in south
Asia.

Between 2005–2010, two people were killed in North America. Human predation on
wolves. Carcasses of hunted wolves in Russia. Wolves are difficult to hunt
because of their elusiveness, sharp senses, high endurance, and ability to
quickly incapacitate and kill hunting dogs. Historic methods include killing of
spring-born litters in their dens, with dogs (usually combinations of, and ),
poisoning with, and. A popular method of wolf hunting in Russia involves
trapping a pack within a small area by encircling it with 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 and, wolves
are traditionally hunted with eagles and falcons, though this practise 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, but is controversial. Several types of dog, including like the,, and,
have been specifically bred for wolf hunting. Gray wolves and coyotes used as
draught animals in northern Ontario, 1923 Wild wolves are sometimes kept as and,
in some rarer occasions, as.

Although closely related to domesticated, 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. See also • •, a gray wolf being electronically tracked in the
northwest United States • Further reading • Apollonio, Marco; Mattioli, Luca
(2006). Il Lupo in Provincia di Arezzo (in Italian). Editrice Le Balze..

• Bibikov, D. 'Volk: Proiskhozhdenie, sistematika, morfologia, ekologia [The
Wolf: History, Systematics, Morphology and Ecology]' (in Russian).

Nauka, Moscow, USSR.. • Busch, Robert H.

Wolf Almanac. The Lyons Press.. • Coleman, Jon T. Vicious: Wolves and Men in
America. Yale University Press..

• Dutcher, Jim; Dutcher, Jamie (2003). Wolves at Our Door: The Extraordinary
Story of the Couple Who Lived with Wolves.

William Andrew.. • Fischer, Hank (1995). • Fuller, Todd K. 'Wolves of the
World'. Voyageur Press.. • Grooms, Steve (1999).

'Return of the Wolf'. Northword Press..

• Hampton, Bruce (1997). The Great American Wolf. Holt Paperbacks.. •
Harrington, Fred H.; Paquet, Paul C. Wolves of the world: perspectives of
behavior, ecology, and conservation. Simon & Schuster.. • McIntyre, Rick (1996).

A Society of Wolves: National Parks and the Battle over the Wolf. Voyageur
Press..

• McNamee, Thomas (1998). The Return of the Wolf to Yellowstone. Holt
Paperbacks..

David (1966).. Department of the Interior, Park Service. David (1993). 'The Way
of the Wolf'.

Voyageur Press.. • Murie, Adolph (1944).. Department of the Interior, Park
Service. • Musiani, Marco; Boitani, Luigi; Paquet, Paul C. The World of Wolves:
New Perspectives on Ecology, Behaviour, and Management.

University of Calgary Press.. • Nie, Martin (2003). Beyond Wolves: The Politics
of Wolf Recovery and Management. University of Minnesota Press.. • Peterson,
Rolf Olin (1977).. National Park Service Scientific Monograph Series.

• Weaver, John (1978).. Department of the Interior, Park Service. • The species
Canis lupus also includes the ( Canis lupus familiaris) and the ( Canis lupus
dingo), both of which are regarded as man-made variants. However, neither dogs
nor dingoes are referred to as gray wolves. Throughout this article, the term
'gray wolf' will be used to refer collectively to naturally occurring
subspecies, especially the nominate subspecies,. • The term 'western wolf' is
primarily used by in distinguishing Canis lupus from, the so-called 'eastern
wolf'. • In the past, the prevailing view on gray wolf packs was that they
consisted of individuals vying with each other for dominance, with dominant gray
wolves being referred to as the 'alpha' male and female, and the subordinates as
'beta' and 'omega' wolves.

This terminology was first used in 1947 by Rudolf Schenkel of the, who based his
findings on researching the behavior of captive gray wolves. This view on gray
wolf pack dynamics was later popularized by in his 1970 book The Wolf. He
formally disavowed this terminology in 1999, explaining that it was heavily
based on the behavior of captive packs consisting of unrelated individuals, an
error reflecting the once prevailing view that wild pack formation occurred in
winter among independent gray wolves. Later research on wild gray wolves
revealed that the pack is usually a family consisting of a breeding pair and its
offspring of the previous 1–3 years.

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