Gray wolf | Wikipedia audio article


The wolf (Canis lupus), also known as the
gray wolf, timber wolf, western wolf, and its other subspecies 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). Like the red wolf, 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..
The gray wolf is the second most specialised member of the genus Canis, after the Ethiopian
wolf, as demonstrated by its morphological 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 eastern wolf, coyote, and golden jackal,
to produce fertile hybrids. It is the only species of Canis to have a range encompassing
both the Old and New Worlds, 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 gray wolf is typically an apex predator throughout its range, with
only humans and tigers posing a serious threat to it. It feeds primarily on large 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
gray wolf population is estimated to be 300,000. 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 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. Both derive from
the Proto-Indo-European root *wlqwos or *lukwos.==Taxonomy and evolution=====Taxonomy===The species Canis lupus was first recorded
by Carl Linnaeus in his publication Systema Naturae 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 Mammal Species of the World third
edition that was published in 2005. The nominate subspecies is the Eurasian wolf (Canis lupus
lupus), also known as the common wolf. The subspecies includes the domestic dog, dingo,
eastern wolf and red wolf, but lists C. l. italicus and C. l. communis as synonyms of
C. l. lupus. However, the classification of several as either species or subspecies has
recently been challenged.===Origin===The evolution of the wolf occurred over a
geologic time scale of at least 300 thousand years.
The gray wolf Canis lupus 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 gray
wolves have identified distinct sub-populations that live in close proximity to each other.
This variation in sub-populations is closely linked to differences in habitat – precipitation,
temperature, vegetation, and prey specialization – which affect cranio-dental plasticity.The
archaeological and paleontological records show gray wolf continuous presence for at
least the last 300,000 years. This continuous presence contrasts with genomic analyses,
which suggest that all modern wolves and dogs descend from a common ancestral wolf population
that existed as recently as 20,000 years ago. These analyses indicate a population bottleneck,
followed by a rapid radiation from an ancestral population at a time during, or just after,
the Last Glacial Maximum. However, the geographic origin of this radiation is not known.===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. Three clusters were identified occupying southern
and central Europe in Italy, the Dinaric-Balkans, the Carpathians. 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 Pontic-Caspian
Steppe than they were to wolves from north-central Europe. These clusters may have been the result
of expansion from glacial refugia, 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 ecotypes – 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===It was once thought that dogs and gray wolves
did not voluntarily interbreed in the wild, though they can produce fertile wolf-dog offspring.
In 2010, a study of 74 Italian wolf 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,
black colored wolves 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
Caucasus Mountains showed that as many as 10% of dogs in the area, including livestock
guardian dogs, are first generation hybrids. The captive breeding of wolf-dog hybrids has
proliferated in the United States, where there is an estimated population of 300,000.The
gray wolf has interbred extensively with the eastern wolf producing a hybrid population
termed Great Lakes boreal wolves. 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 Y-chromosomes
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
artificial insemination, making this the first hybridization case between pure coyotes and
northwestern gray wolves. At six months of age, the hybrids were closely monitored and
were shown to display both physical and behavioral characteristics from both species. Although
hybridization between wolves and golden jackals 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
phenotypes, to the point of being mistaken for wolves by trained biologists.==Physical description=====
Anatomy and dimensions===The gray wolf is the largest extant member
of the Canidae, 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 Bergmann’s Rule, with the large wolves of Alaska and Canada sometimes weighing 3–6
times more than their Middle Eastern and South Asian 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 79.4 kg (175 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 Russia. 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 coyote and golden jackal), 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 ribcage, 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.Generally, wolves have a high heart weight of 0.93%-1.07%
total body mass compared to the average mammal at 0.59% total body mass. Wolves have a decreased
heart rate suggesting cardiac enlargement and hypertrophy. Tibetan gray wolves, which
occupy territories up to 3,000 above sea level, have evolved hearts that withstand the low
oxygen levels. Specifically, these wolves have a strong selection for RYR2, a gene that
initiates cardiac excitation.===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 hyenas. Its molars 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 German shepherd. 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 placental mammals, the bite force at the canines (in Newtons/kilogram
of body weight) was greatest in the extinct dire wolf (163), then followed among the extant
canids 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). A similar
trend was found with the carnassial tooth bite force, but with the extinct dire wolf
and gray wolf both measuring (141), then followed by the African hunting dog (136), the dhole
(114), and the dingo (113).===Fur===The gray wolf has very dense and fluffy winter
fur, with short underfur and long, coarse guard hairs. 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 elbows down to the calcaneal tendons. 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. Black-colored wolves in North America inherited the Kb allele responsible
for melanism 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.==Behavior=====
Social and territorial behaviors===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
pack 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 sexual maturity 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. Studies of wolves’ cortisol levels show that they
rise significantly when a pack member dies, indicating the presence of stress.Wolves are
highly territorial 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 km2 (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 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. 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 scent marking, direct attacks and howling (see Communication). 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 Denali National Park and Preserve were due
to predation by other wolves.===Reproduction and development===The gray wolf is generally monogamous, 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 “Casanova wolves” and, unlike males from established
packs, they do not form pair bonds with the females they mate with. Some gray wolf packs
may have multiple breeding females this way, as is the case in Yellowstone National Park.
Gray wolves also practice alloparental care, 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 mount each other when the highest ranking female
in the pack comes into heat. 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 litter annually
being the average. Unlike the coyote, the gray wolf never reaches reproductive senescence.
Estrus 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 whelp in the den of their previous
litter, while younger females typically den near their birthplace. The gestation period
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.===Hunting and feeding behaviors===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. The gray 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, 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,
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. 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.
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. 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. 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) 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.
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 heart, liver, lungs
and stomach lining. The kidneys and spleen are eaten once they are exposed, followed
by the muscles. A single wolf can eat 15–19% of its body weight in a single feeding.==Communication=====
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 piloerection.
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. 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 dominant animal, and consists of the
submissive wolf lying partly on its back and allowing the dominant wolf to sniff its anogenital
area. 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.===Auditory===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 km2 (50 sq mi).
Wolf howls are generally indistinguishable from those of large dogs. Male wolves give
voice through an octave, passing to a deep bass with a stress on “O”, while females produce
a modulated nasal baritone 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 horned owl. 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 mutually intelligible, 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.===Olfactory===Olfaction is probably the wolf’s most acute
sense, and plays a fundamental role in communication. The wolf has a large number of apocrine sweat
glands on the face, lips, back, and between the toes. The odor produced by these glands
varies according to the individual wolf’s microflora and diet, giving each a distinct
“odor fingerprint”. A combination of apocrine and eccrine sweat glands on the feet allows
the wolf to deposit its scent whilst scratching the ground, which usually occurs after urine
marking and defecation during the breeding season. The follicles present on the guard
hairs from the wolf’s back have clusters of apocrine and sebaceous glands at their bases.
As the skin on the back is usually folded, this provides a microclimate 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 pre-caudal scent glands may play a role in
expressing aggression, as combative wolves raise the base of their tails whilst drooping
the tip, thus positioning the scent glands at the highest point. The wolf possesses a
pair of anal sacs 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 gender and reproductive state. The secretions
of the preputial glands 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
substances from the vagina, which communicate the females’ reproductive state, and can be
detected by males from long distances. Urine marking is the best-studied means of olfactory
communication 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 raised leg urination (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===The gray wolf is a habitat generalist, and
can occur in deserts, grasslands, forests and arctic tundras. 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 topography. 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 tissue-freezing 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 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.===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 ungulates, 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 preying
on the vulnerable individuals of large prey, with pack of timber wolves capable of bringing
down a 500 kg (1,100 lb) moose. Digestion 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 ungulates, they are not fussy eaters. Smaller sized animals
that may supplement the diet of wolves include marmots, hares, badgers, foxes, weasels, ground
squirrels, mice, hamsters, voles and other rodents, as well as insectivores. They frequently
eat waterfowl and their eggs. When such foods are insufficient, they prey on lizards, snakes,
frogs, rarely toads and large insects as available. In times of scarcity, wolves readily eat carrion,
visiting cattle burial grounds and slaughter houses. Cannibalism 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 Astrakhan hunt Caspian seals on the Caspian Sea coastline
and some wolf packs in Alaska and Western Canada have been observed to feed on salmon.
Humans are rarely, but occasionally preyed upon. Other primates occasionally taken by
wolves include grey langurs in Nepal and hamadryas baboons in Saudi Arabia.In Eurasia, many gray
wolf populations are forced to subsist largely on livestock and garbage in areas with dense
human activity, though wild ungulates such as moose, red deer, roe deer and wild boar
are still the most important food sources in Russia and the more mountainous regions
of Eastern Europe. Other prey species include reindeer, argali, mouflon, wisent, saiga,
ibex, chamois, wild goats, fallow deer and musk deer. 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, white-tailed deer, elk, mule deer, bighorn
sheep, Dall’s sheep, American bison, muskox and caribou.
Wolves supplement their diet with fruit and vegetable matter. They willingly eat the berries
of mountain ash, lily of the valley, bilberries, blueberries and cowberry. Other fruits include
nightshade, apples and pears. They readily visit melon fields during the summer months.
A well-fed wolf stores fat under the skin, around the heart, intestines, kidneys, and
bone marrow, particularly during the autumn and winter.===Enemies and competitors===Gray wolves typically dominate other canid
species in areas where they both occur. In North America, incidents of gray 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. Near identical interactions have been observed in Eurasia between gray wolves and
golden jackals, with the latter’s numbers being comparatively small in areas with high
wolf densities. Gray wolves are the most important predator of raccoon dogs, killing large numbers
of them in the spring and summer periods. Wolves also kill red, arctic and corsac foxes,
usually in disputes over carcasses, sometimes eating them. In Asia, they may compete with
dholes, though there is at least one record of a lone wolf associating with a pair of
dholes in Debrigarh Wildlife Sanctuary.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. Wolf interactions with American black bears 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, polar bears are rarely encountered by wolves,
though there are two records of wolf packs killing polar bear cubs. Wolves also kill
the cubs of Asian black bears.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-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
felines. 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 kills. They have been reported killing mothers and their kittens.
Wolves hunt steppe cats, and may scavenge from snow leopard kills. Wolves may also reduce
Eurasian lynx 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 bobcats
have been documented.Leftovers of wolf kills are sometimes scavenged by wolverines. 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, tigers appear to be the only serious predators of wolves. 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.==Range and conservation==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 of attacks
on humans. The species is now extinct in much of Western Europe, in Mexico and much of the
United States. In modern times, the gray wolf occurs mostly in wilderness and remote areas,
particularly in Canada, Alaska and northern USA, Europe, and Asia 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
least concern by the IUCN.===Europe=======
Decline====In Europe, the oldest gray wolf remains were
found in France and date to 400,000-350,000 years ago. Wolf populations strongly declined
across Europe during the 18th and 19th centuries largely due to human persecution, and by the
end of the Second World War they had been eradicated from all of Central Europe and
almost all of Northern Europe.The extermination of Northern Europe’s wolves first became an
organized effort during the Middle Ages, and continued until the late 19th century. In
England, wolf persecution was enforced by legislation, and the last wolf was killed
in the early sixteenth century during the reign of Henry VII. Wolves lasted longer in
Scotland, where they sheltered in vast tracts of forest, which were subsequently burned
down. Wolves managed to survive in the forests of Braemar and Sutherland until 1684. The
extirpation of wolves in Ireland followed a similar course, with the last wolf believed
to have been killed in 1786. A wolf bounty was introduced in Sweden in 1647, after the
extermination of moose and reindeer forced wolves to feed on livestock. The Sami 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 World War II.In Central Europe, wolves were
dramatically reduced in number during the early nineteenth century, because of organized
hunts and reductions in ungulate populations. In Bavaria, the last wolf was killed in 1847,
and had disappeared from the Rhine regions by 1899. In Switzerland, wolves were extinct
in the twentieth century; they are naturally coming back from Italy since the 1990s. In
1934, Nazi Germany 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 Germany before 1945 was the
so-called “Tiger of Sabrodt”, which was shot near Hoyerswerda, Lusatia (then Lower Silesia)
in 1904. Today, wolves have returned to the area. Wolf hunting in France was first institutionalized
by Charlemagne between 800–813, when he established the louveterie, a special corps
of wolf hunters. The louveterie was abolished after the French Revolution in 1789, but was
re-established in 1814. In 1883, up to 1,386 wolves were killed, with many more by poison. In Eastern Europe, 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 Slovakia 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 Balkans benefitted from the region’s contiguity with the former
Soviet Union 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 Carpathian Basin. 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 Peloponnese in 1930. Despite periods of intense hunting
during the eighteenth century, wolves never disappeared in the western Balkans, from Albania
to the former Yugoslavia. Organized persecution of wolves began in Yugoslavia in 1923, with
the setting up of the Wolf Extermination Committee (WEC) in Kocevje, Slovenia. The WEC was successful
in reducing wolf numbers in the Dinaric Alps.In Southern Europe, 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 Iberian Peninsula 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 Alps by 1800, and numbered
only 100 by 1973, inhabiting only 3–5% of their former Italian range.====Recovery====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 Mercantour National Park in 1993,
and at least fifty wolves were discovered in the western Alps in 2000. By 2013 the 250
wolves in the Western Alps imposed a significant burden on traditional sheep and goat husbandry
with a loss of over 5,000 animals in 2012. There are approximately 2,000 wolves inhabiting
the Iberian Peninsula, of which 150 reside in northeastern Portugal. In Spain, the species
occurs in Galicia, Leon, and Asturias. Although hundreds of Iberian wolves are illegally killed
annually, the population has expanded south across the river Duero and east to the Asturias
and Pyrenees Mountains. 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 breeding pairs. 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 Republic of Karelia, 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 Central European countries. In the east, its range overlaps with populations
in Lithuania, Belarus, Ukraine, and Slovakia. 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 Berlin; 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 Croatia 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 Albania and Macedonia is largely unknown, despite the importance the two countries have
in linking wolf populations from Greece to those of Bosnia and Herzegovina 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 Slovenia.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 mating seasons do not fully
coincide, the likelihood of wild wolves and dogs mating and producing surviving offspring
is small.===Asia=======Historical range and decline====During the 19th century, gray wolves were
widespread in many parts of the Holy Land east and west of the Jordan River. 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 Saudi Arabia
during the 19th century, with most early publications involving animals either from southwestern
Asir, northern rocky areas bordering Jordan, or areas surrounding Riyadh.The gray wolf’s
range in the Soviet Union encompassed nearly the entire territory of the country, being
absent only on the Solovetsky Islands, Franz-Josef Land, Severnaya Zemlya, and the Karagin, Commander
and Shantar Islands. The species was exterminated twice in Crimea, once after the Russian Civil
War, and again after World War II. 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 European Russia. The population increased
again by 1980 to about 75,000, with 32,000 being killed in 1979. Wolf populations in
northern inner Mongolia declined during the 1940s, primarily because of poaching of gazelles,
the wolf’s main prey. In British-ruled India, wolves were heavily persecuted because of
their attacks on sheep, goats and children. In 1876, 2,825 wolves were bountied in the
North-Western Provinces (NWP) and Bihar. By the 1920s, wolf extermination remained a priority
in the NWP and Awadh. Overall, over 100,000 wolves were killed for bounties in British
India between 1871 and 1916.Wolves in Japan became extinct during the Meiji restoration
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 Korean Peninsula and Japan, before it separated from mainland
Asia, 20,000 years ago during the Pleistocene. During the Holocene, the Tsugaru Strait widened
and isolated Honshu from Hokkaidō, thus causing climatic changes leading to the extinction
of most large bodied ungulates inhabiting the archipelago. Japanese wolves likely underwent
a process of island dwarfism 7,000–13,000 years ago in response to these climatological
and ecological pressures. C. l. hattai (formerly native to Hokkaidō) was significantly larger
than its southern cousin C. l. hodophilax, 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====There is little reliable data on the status
of wolves in the Middle East, save for those in Israel and Saudi Arabia, 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 Syria. A small wolf population occurs in the Golan Heights, and is well protected by
the military activities there. Wolves living in the southern Negev desert are contiguous
with populations living in the Egyptian Sinai and Jordan. Throughout the Middle East, the
species is only protected in Israel. Elsewhere, it can be hunted year-round by Bedouins.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 Afghanistan and Pakistan
are important strongholds for the wolf. It has been estimated that there are about 300
wolves in approximately 60,000 km2 (23,000 sq mi) of Jammu and Kashmir in northern India,
and 50 more in Himachal Pradesh. 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 Nepal and Bhutan, there is no information of wolves
occurring there.Wolf populations throughout Northern and Central Asia are largely unknown,
but are estimated in the hundreds of thousands based on annual harvests. Since the fall of
the Soviet Union, continent-wide extermination of wolves has ceased, and wolf populations
have increased to about 25,000–30,000 animals throughout the former Soviet Union. In China
and Mongolia, 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 Xinjiang and Tibet hold
about 10,000 and 2,000 respectively. 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 Western world that
the wolf had never existed in southern China.===North America=======
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, California west of the Sierra Nevada, and the tropical and subtropical areas
of Mexico. Large continental islands occupied by wolves included Newfoundland, Vancouver
Island, southeastern Alaskan islands, and throughout the Arctic Archipelago and Greenland.
While Lohr and Ballard postulated that the gray wolf had never been present on Prince
Edward Island, 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 November 6, 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 Northumberland Strait.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 Appalachians and the northwestern
Great Lakes Region. In Canada, the gray wolf was extirpated in New Brunswick and Nova Scotia
between 1870 and 1921, and in Newfoundland around 1911. It vanished from the southern
regions of Quebec and Ontario between 1850 and 1900. The gray wolf’s decline in the prairies
began with the extermination of the American bison 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 Minnesota. 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====The species’ modern range in North America
is mostly confined to Alaska and Canada, with populations also occurring in northern Minnesota,
northern Wisconsin and Michigan’s Upper Peninsula, and small portions of Washington, Idaho, northern
Oregon, and Montana. 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 Glacier National Park 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 Alberta and British Columbia were relocated
to Yellowstone National Park and Idaho. In addition, the Mexican wolf (Canis lupus baileyi)
was reintroduced to Arizona and New Mexico 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 province and territory. First Nations residents may hunt wolves without restriction, and some
provinces require licenses for residents to hunt wolves while others do not. In Alberta,
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 woodland caribou.In
Alaska, the gray wolf population is estimated at 6,000–7,000, and can be legally harvested
during hunting and trapping seasons, with bag limits 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 Endangered Species Act (ESA) protection in Minnesota, Wisconsin, and Michigan
in 1974, and was re-classed 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. A female wolf shot in 2013 in Hart County, Kentucky 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.==Diseases and parasites=====Viral and bacterial infections===Viral diseases carried by wolves include rabies,
canine distemper, canine parvovirus, infectious canine hepatitis, papillomatosis, canine coronavirus,
and foot and mouth disease. Wolves are a major host for rabies in Russia, Iran, Afghanistan,
Iraq and India. In wolves, the incubation period is 8–21 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 doesn’t 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.Bacterial diseases carried by wolves include brucellosis,
lyme disease, leptospirosis, tularemia, bovine tuberculosis, listeriosis and anthrax. 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.===Parasitic infections===
Wolves carry ectoparasites and endoparasites, with wolves in the former Soviet Union having
been recorded to carry at least 50 species. Most of these parasites infect wolves without
adverse effects, though the effects may become more serious in sick or malnourished specimens.
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.Wolves are often infested with a variety of arthropod
exoparasites, including fleas, ticks, lice, and mites. The most harmful to wolves, particularly
pups, is Sarcoptes scabiei (or mange mite), though they rarely develop full blown mange,
unlike foxes. 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. The tick Dermacentor pictus also infests wolves. Other ectoparasites include biting
lice, sucking lice and the fleas Pulex irritans and Ctenocephalides canis.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. Wolves may carry Neospora caninum,
which is of particular concern to farmers, as the disease can be spread to livestock,
with infected animals being 3–13 times more likely to miscarry than those not infected.
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 isn’t 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.==Relationships with humans==
Human presence appears to stress wolves, as seen by increased cortisol levels in instances
such as snowmobiling near their territory.===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).Ancient Greek literature is similar: Autolycus (“the
wolf itself”), Lycurgus (“wolf-work”). The Latin for “female prostitute” is lupa,
and the most famous brothel in Pompeii was the Lupanar.====In folklore, religion and mythology====The wolf is a common motif in the foundational
mythologies and cosmologies of peoples throughout Eurasia and North America (corresponding to
the historical extent of the habitat of the gray wolf). The obvious attribute of the wolf
is its nature of a predator, and correspondingly it is strongly associated with danger and
destruction, making it the symbol of the warrior on one hand, and that of the devil on the
other. The modern trope of the Big Bad Wolf is a development of this. The wolf holds great
importance in the cultures and religions of the nomadic peoples, both of the Eurasian
steppe and of the North American Plains. In many cultures, the identification of the warrior
with the wolf (totemism) gave rise to the notion of Lycanthropy, the mythical or ritual
identification of man and wolf.====In fable and literature====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. 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.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. 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. Farley Mowat’s largely fictional 1963 memoir Never Cry Wolf 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 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.====In heraldry and symbolism====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. The wolf is featured on
the flags of the Confederated Tribes of the Colville Reservation, the Oneida Nation of
Wisconsin and the Pawnee. 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.===Conflicts=======
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 incidents
are rare in North America, where healthy populations of wild prey have been largely restored. The
majority of losses occur during the summer grazing period, with untended livestock in
remote pastures being the most vulnerable to wolf predation. The most frequently targeted
livestock species are sheep (Europe), domestic reindeer (northern Scandinavia), goats (India),
horses (Mongolia), cattle and turkeys (North America). The number of animals killed in
single attacks varies according to species: most attacks on cattle and horses result in
one death, while turkeys, sheep and domestic reindeer may be killed in surplus. Wolves
mainly attack livestock when the animals are grazing, though they occasionally break into
fenced enclosures. 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.====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. 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 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.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====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 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, though cases of non-rabid wolf attacks in winter have been recorded
in Belarus, Kirov and Irkutsk oblasts, Karelia and Ukraine. Also, wolves with pups experience
greater food stresses during this period.The majority of victims of predatory wolf attacks
are children under the age of 18 and, in the rare cases where adults are killed, the victims
are almost always women. 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, 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. 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 south Asia. Between 2005–2010, two people
were killed in North America.====Human predation on wolves====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, coursing
with dogs (usually combinations of sighthounds, bloodhounds and fox terriers), poisoning with
strychnine, and trapping. A popular method of wolf hunting in Russia involves trapping
a pack within a small area by encircling it with fladry poles carrying a human scent.
This method relies heavily on the wolf’s fear of human scents, though it can lose its effectiveness
when wolves become accustomed to the 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 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 Borzoi, Irish wolfhound,
and Kyrgyz Tajgan, have been specifically bred for wolf hunting.===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.==See also==
Ethiopian wolf OR-7, a gray wolf being electronically tracked
in the northwest United States Custer wolf==Further reading==
Apollonio, Marco; Mattioli, Luca (2006). Il Lupo in Provincia di Arezzo (in Italian).
Editrice Le Balze. ISBN 88-7539-123-8. Bibikov, D. I. (1985). “Volk: Proiskhozhdenie,
sistematika, morfologia, ekologia [The Wolf: History, Systematics, Morphology and Ecology]”
(in Russian). Nauka, Moscow, USSR. ASIN B001A1TKK4. Busch, Robert H. (2009). Wolf Almanac. The
Lyons Press. ISBN 1-59921-069-X. Coleman, Jon T. (2006). Vicious: Wolves and
Men in America. Yale University Press. ISBN 0-300-11972-0.
Dutcher, Jim; Dutcher, Jamie (2003). Wolves at Our Door: The Extraordinary Story of the
Couple Who Lived with Wolves. William Andrew. ISBN 0-7434-0049-6.
Fischer, Hank (1995). Wolf Wars. Falcon. ISBN 1-56044-352-9.
Fuller, Todd K. (2004). “Wolves of the World”. Voyageur Press. ISBN 0-89658-640-5.
Grooms, Steve (1999). “Return of the Wolf”. Northword Press. ISBN 1-55971-717-3.
Hampton, Bruce (1997). The Great American Wolf. Holt Paperbacks. ISBN 0-8050-5528-2.
Harrington, Fred H.; Paquet, Paul C. (1982). Wolves of the world: perspectives of behavior,
ecology, and conservation. Simon & Schuster. ISBN 0-8155-0905-7.
McIntyre, Rick (1996). A Society of Wolves: National Parks and the Battle over the Wolf.
Voyageur Press. ISBN 0-89658-325-2. McNamee, Thomas (1998). The Return of the
Wolf to Yellowstone. Holt Paperbacks. ISBN 0-8050-5792-7.
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 0-89658-179-9.
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 1-55238-269-9.
Nie, Martin (2003). Beyond Wolves: The Politics of Wolf Recovery and Management. University
of Minnesota Press. ISBN 0-300-11972-0. 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.==Notes====References=====Bibliography===
Graves, Will (2007). Wolves in Russia: Anxiety throughout the ages. Detselig Enterprises.
ISBN 1-55059-332-3. Lopez, Barry H. (1978). Of Wolves and Men.
J. M. Dent and Sons Limited. ISBN 0-7432-4936-4. 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 0-8166-1026-6.
Mech, L. David; Boitani, Luigi, eds. (2003). Wolves: Behaviour, Ecology and Conservation.
University of Chicago Press. ISBN 0-226-51696-2. Van Nuys, Frank (2015). Varmints and Victims:
Predator Control in the American West. Lawrence, KS: University Press of Kansas.
Walker, Brett L. (2005). The Lost Wolves Of Japan. University of Washington Press. ISBN
0-295-98492-9. 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 0-285-62411-3.==External links==
Quotations related to Wolf at Wikiquote Dog (mammal) at Encyclopædia Britannica
Gray wolf (mammal) at Encyclopædia Britannica Wolf (mammal) at 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

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