Historical (red + green) and modern (green) range of wild
subspecies of C. lupus
The gray wolf (
Canis lupus),[a] also known as the timber wolf or
western wolf,[b] is a canine native to the wilderness and remote areas
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
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.
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
2 Taxonomy and evolution
2.2 Evolution and relationship with the dog
2.3 Population structure
2.4 Hybridization with other Canis
3 Physical description
3.1 Anatomy and dimensions
3.2 Skull and dentition
4.1 Social and territorial behaviors
4.2 Reproduction and development
4.3 Hunting and feeding behaviors
6.3 Enemies and competitors
7 Range and conservation
7.2.1 Historical range and decline
7.2.2 Modern range
7.3 North America
7.3.1 Historical range and decline
7.3.2 Modern range
8 Diseases and parasites
8.1 Viral and bacterial infections
8.2 Parasitic infections
9 Relationships with humans
9.1 In culture
9.1.1 In personal names
9.1.2 In folklore, religion and mythology
9.1.3 In fable and literature
9.1.4 In heraldry and symbolism
9.2.1 Wolf predation on livestock
9.2.2 Conflicts with dogs
9.2.3 Wolf predation on humans
Human predation on wolves
9.3 As pets and working animals
10 See also
11 Further reading
14 External links
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
Taxonomy and evolution
Canis lupus was first recorded by
Carl Linnaeus in his
Systema Naturae in 1758, with the Latin classification
translating into the English words "dog wolf". The thirty-seven
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.
Evolution and relationship with the dog
Main article: Evolution of the wolf
Artist's impression of a Beringian wolf
The evolution of the wolf occurred over a geologic time scale of
800,000 years, transforming the first
Middle Pleistocene wolf specimen
that is recognized as being morphologically similar to
into today's dog, dingo, and gray wolf. Ecological factors including
habitat type, climate, prey specialization and predatory competition
will greatly influence the wolf's genetic population structure and
cranio-dental plasticity. Wolves went through a population
bottleneck 20,000 years before present (YBP), which indicates that
many wolf populations had gone extinct at a time that coincided with
Last Glacial Maximum
Last Glacial Maximum 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.
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.
Italian wolf consisted of an isolated population with low genetic
diversity. Wolves from Croatia, Bulgaria, and Greece formed the
Balkans cluster. Wolves from Finland, Latvia, Belarus, Poland
Russia formed the north-central Europe cluster, with wolves from
Carpathians cluster coming from a mixture of wolves from the
north-central cluster and the Dinaric-
Balkans cluster. The wolves from
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
F1 wolf-dog hybrids from Wildlife Park Kadzidlowo, Poland. The first
is the product of a male wolf and a female spaniel, while the latter
comes from a female wolf and a male West Siberian Laika
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 USA, with 300,000 such animals being present
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
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
Anatomy and dimensions
Differences between gray wolf and coyote
Gray wolf skeleton
Wolf mandible diagram showing the names and positions of the teeth.
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
80 kg (176 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
Alaska on July 12, 1939 and weighed 79.4 kg
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
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 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).
Black and white-furred gray wolves
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
North America than in Eurasia, with about half the wolves in
Yellowstone National Park
Yellowstone National Park being black.
Social and territorial behaviors
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.[c] 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.
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
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
See also: Canine reproduction
Gray wolves mating
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
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 litter annually being the average.
Unlike the coyote, the gray wolf never reaches reproductive
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
American bison standing its ground, thereby increasing its chance of
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
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
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 moose in typical fashion: biting the hindquarters
The actual killing method varies according to prey species. With large
prey, mature wolves usually avoid attacking frontally, instead
focusing on the rear and sides of the animal. Large prey, such as
moose, is killed by biting large chunks of flesh from the soft
perineum area, causing massive blood loss. Such bites can cause wounds
10–15 cm (3.9–5.9 in) in length, with three such bites
to the perineum usually being sufficient to bring down a large deer in
optimum health. 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.
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 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.
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
Facial expressions (Konrad Lorenz, 1952). Bottom to top: increasing
fear (ears back); left to right: increasing aggression (snarl); top
right: maximum of both.
Expressive characteristics of visual features
used during social interactions in wolves
Closed to slits
Erect and forward
Flattened and turned down to side
Horizontal retraction ("submissive grin")
Extended ("lick intention")
Shortened (skin folded)
Lengthened (skin smoothed)
Contracted (bulging over eyes)
Tucked under body
Gray wolf howling
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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
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
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
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
Gray wolf carrying caribou hindquarter, Denali National Park
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.
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
predating on the vulnerable individuals of large prey, with pack of
timber wolves capable of bringing down a 500 kg (1,100 lb)
Digestion only takes a few hours, thus wolves can feed
several times in one day, making quick use of large quantities of
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
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 attacking brown bear with cubs
Gray wolves confronting coyotes over pronghorn carcass (1919), Louis
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
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
Main article: List of grey wolf populations by country
See also: History of wolves in Yellowstone, Wolves in Great Britain,
and Wolves in Ireland
The gray wolf was once one of the world's most widely distributed
mammals, living throughout the northern hemisphere north of 15°N
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,
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
Polychrome cave painting of a gray wolf, Font-de-Gaume, 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
mating seasons do not fully coincide, the likelihood of wild wolves
and dogs mating and producing surviving offspring is small.
The extermination of Northern Europe's wolves first became an
organized effort during the Middle Ages, and continued until the late
1800s. 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
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
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.
Last wolf to be killed in central Finland (Karstula, 1911)
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
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.
Present range of
Canis lupus subspecies in Eurasia
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
Mercantour National Park
Mercantour National Park in 1993, and at least fifty wolves
were discovered in the western
Alps in 2000. By 2013 the 250 wolves in
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
European wolf in Bavarian
Forest National Park, 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 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
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
Bosnia and Herzegovina since 1986, while the
species is fully protected in neighbouring
Croatia and Slovenia.
Historical range and decline
Monument to the last Japanese gray wolf, Nara Prefecture, Japan
During the 19th century, gray wolves were widespread in many parts of
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
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
Korean Peninsula and Japan, before it separated from
mainland Asia, 20,000 years ago during the Pleistocene. During the
Tsugaru Strait widened and isolated
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
Indian Wolf at Velavadar (Blackbuck National Park, Gujarat)
Gray wolf near Ardahan, Turkey. Although Turkish wolves have no legal
protection, they may number about 7000 individuals.
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
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)
Jammu and Kashmir
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
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.
Historical and present range of gray wolf subspecies in 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
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
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,: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 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
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
New Brunswick and
Nova Scotia between 1870 and 1921, and
in Newfoundland around 1911. It vanished from the southern regions of
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
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.
Captive Mexican gray wolf
The species' modern range in
North America is mostly confined to
Alaska and Canada, with populations also occurring in 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
British Columbia were
Yellowstone National Park
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
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
Endangered Species Act
Endangered Species Act (ESA) protection in Minnesota,
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
Diseases and parasites
Viral and bacterial infections
Footage of a gray wolf taken from Abruzzo Natural Park showing
advanced signs of canine distemper
Viral diseases carried by wolves include rabies, canine distemper,
canine parvovirus, infectious canine hepatitis, papillomatosis, and
canine coronavirus. Wolves are a major host for rabies in Russia,
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
Bacterial diseases carried by wolves include brucellosis, lyme
disease, leptospirosis, tularemia, bovine tuberculosis,
listeriosis, anthrax and foot and mouth disease. 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
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.
Wolves carry ectoparasites and endoparasites, with wolves in the
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
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
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"),
The Latin for "female prostitute" is lupa, and the most famous brothel
Pompeii was the Lupanar.
In folklore, religion and mythology
Main article: Wolves 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
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
Little Red Riding Hood
Little Red Riding Hood (1883), Gustave Doré
Aesop featured wolves in several of his fables, playing on the
concerns of Ancient Greece's settled, sheep-herding world. His most
famous is the fable of The Boy Who Cried Wolf, which is directed at
those who knowingly raise false alarms, and from which the idiomatic
phrase "to cry wolf" is derived. Some of his other fables concentrate
on maintaining the trust between shepherds and guard dogs in their
vigilance against wolves, as well as anxieties over the close
relationship between wolves and dogs. Although
Aesop used wolves to
warn, criticize and moralize about human behavior, his portrayals
added to the wolf's image as a deceitful and dangerous animal.
Isengrim the wolf, a character first appearing in the 12th-century
Latin poem Ysengrimus, is a major character in the
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
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
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
In heraldry and symbolism
Main article: Wolves in heraldry
Coat of arms
Coat of arms of the Elvange family, featuring a wolf charge rampant
The wolf is a frequent charge in English armory. It is illustrated as
a supporter on the shields of Lord Welby, Rendel, and Viscount
Wolseley, and can be found on the coat of arms of Lovett and the vast
majority of the Wilsons and Lows. The demi-wolf is a common crest,
appearing in the arms and crests of members of many families,
including that of the Wolfes, whose crest depicts a demi-wolf holding
a crown in its paws, in reference to the assistance the family gave to
Charles II during the battle of Worcester. Wolf heads are common in
Scottish heraldry, particularly in the coats of Clan Robertson and
Skene. The wolf is the most common animal in Spanish heraldry, and is
often depicted as carrying a lamb in its mouth, or across its
back. The wolf is featured on the flags of the Confederated
Tribes of the Colville Reservation, the Oneida Nation of
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
Waiting for a Chinook (c. 1900), Charles Marion Russell
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 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
Conflicts with dogs
Being the most abundant carnivores, feral and 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
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
Wolf attacks on humans
Wolf attacks on humans and List of wolf attacks
Child snatched by a wolf (1914), Le Petit Journal
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,
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. 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
Human predation on wolves
Wolf hunting and
Wolf hunting with dogs
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, 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.
Human uses of hunted wolves
As pets and working animals
Main article: Wolves as pets and working animals
Gray wolves and coyotes used as draught animals in northern Ontario,
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.
OR-7, a gray wolf being electronically tracked in the northwest United
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^ The species
Canis lupus also includes the domestic dog (
familiaris) and the dingo (
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,
Canis lupus lupus.
^ The term "western wolf" is primarily used by American taxonomists in
Canis lupus from
Canis lycaon, the so-called "eastern
wolf" of North America.
^ 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 University of Basel,
who based his findings on researching the behavior of captive gray
wolves. This view on gray wolf pack dynamics was later popularized by
L. David Mech
L. David Mech 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.
^ a b Mech, L.D., Boitani, L. (
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IUCN Red List of
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^ Espuno, Nathalie; Lequette, Benoit; Poulle, Marie-Lazarine; Migot,
Pierre; Lebreton, Jean-Dominique (2004). "Heterogeneous response to
preventive sheep husbandry during wolf recolonization of the French
Alps". Wildlife Society Bulletin. 32 (4): 1195–1208.
^ Mertens, A., Schneider, H., 2005. What is wrong with Romanian
livestock guarding dogs? A discussion. Carnivore Damage Prevent. News
^ Mech & Boitani 2003, pp. 300–304
^ McNay, M. E. (2007). A Review of Evidence and Findings Related to
the Death of Kenton Carnegie on November 8, 2005 Near Points North,
Alaska Department of Fish and Game, Fairbanks, Alaska.
^ Butler, L.; Dale, B.; Beckmen, K. and Farley, S. (2011). "Findings
Related to the March 2010 Fatal Wolf Attack near Chignik Lake,
Special Publication, ADF&G/DWC/WSP-2011-2.
^ Roosevelt, T. (1909). Hunting the grisly and other sketches; an
account of the big game of the United States and its chase with horse,
hound, and rifle, New York, London, G. P. Putnam's sons, pp. 179-207
^ Harding, A. R. (1909). Wolf and coyote trapping; an up-to-date wolf
hunter's guide, giving the most successful methods of experienced
"wolfers" for hunting and trapping these animals, also gives their
habits in detail, Columbus, Ohio, A. R. Harding pub. co.
^ Lopez 1978, p. 108
^ a b c d e Graves 2007, pp. 121–40
^ Lopez 1978, pp. 159–60
^ Tucker, P. & Weide, B. (1998), "Can You Turn a Wolf into a Dog:
Commonly Asked Questions about Wolves and Hybrids in Captivity", Wild
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.
Mech, L. David (1981). The Wolf: The Ecology and Behaviour of an
Endangered Species. University of
Mech, L. David; Boitani, Luigi, eds. (2003). Wolves: Behaviour,
Ecology and Conservation. University of Chicago Press.
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.
Wikimedia Commons has media related to
Wikispecies has information related to
Quotations related to Wolf at Wikiquote
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
African palm civet
African palm civet (N. binotata)
Marsh mongoose (A. paludinosus)
Bushy-tailed mongoose (B. crassicauda)
Jackson's mongoose (B. jacksoni)
Black-footed mongoose (B. nigripes)
Alexander's kusimanse (C. alexandri)
Angolan kusimanse (C. ansorgei)
Common kusimanse (C. obscurus)
Flat-headed kusimanse (C. platycephalus)
Yellow mongoose (C. penicillata)
Pousargues's mongoose (D. dybowskii)
Angolan slender mongoose
Angolan slender mongoose (G. flavescens)
Black mongoose (G. nigrata)
Somalian slender mongoose
Somalian slender mongoose (G. ochracea)
Cape gray mongoose
Cape gray mongoose (G. pulverulenta)
Slender mongoose (G. sanguinea)
Ethiopian dwarf mongoose
Ethiopian dwarf mongoose (H. hirtula)
Common dwarf mongoose
Common dwarf mongoose (H. parvula)
Short-tailed mongoose (H. brachyurus)
Indian gray mongoose
Indian gray mongoose (H. edwardsii)
Indian brown mongoose
Indian brown mongoose (H. fuscus)
Egyptian mongoose (H. ichneumon)
Small Asian mongoose
Small Asian mongoose (H. javanicus)
Long-nosed mongoose (H. naso)
Collared mongoose (H. semitorquatus)
Ruddy mongoose (H. smithii)
Crab-eating mongoose (H. urva)
Stripe-necked mongoose (H. vitticollis)
White-tailed mongoose (I. albicauda)
Liberian mongoose (L. kuhni)
Gambian mongoose (M. gambianus)
Banded mongoose (M. mungo)
Selous' mongoose (P. selousi)
Meller's mongoose (R. melleri)
Meerkat (S. suricatta)
Spotted hyena (C. crocuta)
Brown hyena (H. brunnea)
Striped hyena (H. hyaena)
Aardwolf (P. cristatus)
Large family listed below
Large family listed below
Small family listed below
Cheetah (A. jubatus)
Caracal (C. caracal)
African golden cat
African golden cat (C. aurata)
Bay cat (C. badia)
Asian golden cat
Asian golden cat (C. temminckii)
European wildcat (F. silvestris)
African wildcat (F. lybica)
Jungle cat (F. chaus)
Black-footed cat (F. nigripes)
Sand cat (F. margarita)
Chinese mountain cat
Chinese mountain cat (F. bieti)
Domestic cat (F. catus)
Ocelot (L. pardalis)
Margay (L. wiedii)
Pampas cat (L. colocola)
Geoffroy's cat (L. geoffroyi)
Kodkod (L. guigna)
Andean mountain cat
Andean mountain cat (L. jacobita)
Oncilla (L. tigrinus)
Southern tigrina (L. guttulus)
Serval (L. serval)
Canadian lynx (L. canadensis)
Eurasian lynx (L. lynx)
Iberian lynx (L. pardinus)
Bobcat (L. rufus)
Pallas's cat (O. manul)
Marbled cat (P. marmorata)
Fishing cat (P. viverrinus)
Leopard cat (P. bengalensis)
Sundaland leopard cat (P. javanensis)
Flat-headed cat (P. planiceps)
Rusty-spotted cat (P. rubiginosus)
Cougar (P. concolor)
Jaguarundi (H. yagouaroundi)
Lion (P. leo)
Jaguar (P. onca)
Leopard (P. pardus)
Tiger (P. tigris)
Snow leopard (P. uncia)
Clouded leopard (N. nebulosa)
Sunda clouded leopard
Sunda clouded leopard (N. diardi)
Viverridae (includes Civets)
Binturong (A. binturong)
Small-toothed palm civet
Small-toothed palm civet (A. trivirgata)
Sulawesi palm civet
Sulawesi palm civet (M. musschenbroekii)
Masked palm civet
Masked palm civet (P. larvata)
Golden wet-zone palm civet (P. aureus)
Asian palm civet
Asian palm civet (P. hermaphroditus)
Jerdon's palm civet (P. jerdoni)
Golden palm civet
Golden palm civet (P. zeylonensis)
Owston's palm civet
Owston's palm civet (C. owstoni)
Otter civet (C. bennettii)
Hose's palm civet
Hose's palm civet (D. hosei)
Banded palm civet
Banded palm civet (H. derbyanus)
Banded linsang (P. linsang)
Spotted linsang (P. pardicolor)
African civet (C. civetta)
Abyssinian genet (G. abyssinica)
Angolan genet (G. angolensis)
Bourlon's genet (G. bourloni)
Crested servaline genet
Crested servaline genet (G. cristata)
Common genet (G. genetta)
Johnston's genet (G. johnstoni)
Rusty-spotted genet (G. maculata)
Pardine genet (G. pardina)
Aquatic genet (G. piscivora)
King genet (G. poensis)
Servaline genet (G. servalina)
Haussa genet (G. thierryi)
Cape genet (G. tigrina)
Giant forest genet
Giant forest genet (G. victoriae)
African linsang (P. richardsonii)
Leighton's linsang (P. leightoni)
Malabar large-spotted civet
Malabar large-spotted civet (V. civettina)
Large-spotted civet (V. megaspila)
Malayan civet (V. tangalunga)
Large Indian civet
Large Indian civet (V. zibetha)
Small Indian civet
Small Indian civet (V. indica)
Fossa (C. ferox)
Eastern falanouc (E. goudotii)
Western falanouc (E. major)
Malagasy civet (F. fossana)
Ring-tailed mongoose (G. elegans)
Broad-striped Malagasy mongoose
Broad-striped Malagasy mongoose (G. fasciata)
Grandidier's mongoose (G. grandidieri)
Narrow-striped mongoose (M. decemlineata)
Brown-tailed mongoose (S. concolor)
Durrell's vontsira (S. durrelli)
Caniformia (cont. below)
Giant panda (A. melanoleuca)
Sun bear (H. malayanus)
Sloth bear (M. ursinus)
Spectacled bear (T. ornatus)
American black bear
American black bear (U. americanus)
Brown bear (U. arctos)
Polar bear (U. maritimus)
Asian black bear
Asian black bear (U. thibetanus)
Molina's hog-nosed skunk
Molina's hog-nosed skunk (C. chinga)
Humboldt's hog-nosed skunk
Humboldt's hog-nosed skunk (C. humboldtii)
American hog-nosed skunk
American hog-nosed skunk (C. leuconotus)
Striped hog-nosed skunk
Striped hog-nosed skunk (C. semistriatus)
Hooded skunk (M. macroura)
Striped skunk (M. mephitis)
Sunda stink badger
Sunda stink badger (M. javanensis)
Palawan stink badger
Palawan stink badger (M. marchei)
Southern spotted skunk
Southern spotted skunk (S. angustifrons)
Western spotted skunk
Western spotted skunk (S. gracilis)
Eastern spotted skunk
Eastern spotted skunk (S. putorius)
Pygmy spotted skunk
Pygmy spotted skunk (S. pygmaea)
Eastern lowland olingo
Eastern lowland olingo (B. alleni)
Northern olingo (B. gabbii)
Western lowland olingo
Western lowland olingo (B. medius)
Olinguito (B. neblina)
Ring-tailed cat (B. astutus)
Cacomistle (B. sumichrasti)
White-nosed coati (N. narica)
South American coati
South American coati (N. nasua)
Western mountain coati (N. olivacea)
Eastern mountain coati (N. meridensis)
Kinkajou (P. flavus)
Crab-eating raccoon (P. cancrivorus)
Raccoon (P. lotor)
Cozumel raccoon (P. pygmaeus)
Red panda (A. fulgens)
Caniformia (cont. above)
(includes fur seals
and sea lions)
South American fur seal
South American fur seal (A. australis)
Australasian fur seal (A. forsteri)
Galápagos fur seal
Galápagos fur seal (A. galapagoensis)
Antarctic fur seal
Antarctic fur seal (A. gazella)
Juan Fernández fur seal
Juan Fernández fur seal (A. philippii)
Brown fur seal
Brown fur seal (A. pusillus)
Guadalupe fur seal
Guadalupe fur seal (A. townsendi)
Subantarctic fur seal
Subantarctic fur seal (A. tropicalis)
Northern fur seal
Northern fur seal (C. ursinus)
Steller sea lion
Steller sea lion (E. jubatus)
Australian sea lion
Australian sea lion (N. cinerea)
South American sea lion
South American sea lion (O. flavescens)
New Zealand sea lion
New Zealand sea lion (P. hookeri)
California sea lion (Z. californianus)
Galápagos sea lion
Galápagos sea lion (Z. wollebaeki)
Walrus (O. rosmarus)
Hooded seal (C. cristata)
Bearded seal (E. barbatus)
Gray seal (H. grypus)
Ribbon seal (H. fasciata)
Leopard seal (H. leptonyx)
Weddell seal (L. weddellii)
Crabeater seal (L. carcinophagus)
Northern elephant seal
Northern elephant seal (M. angustirostris)
Southern elephant seal
Southern elephant seal (M. leonina)
Mediterranean monk seal (M. monachus)
Hawaiian monk seal
Hawaiian monk seal (M. schauinslandi)
Ross seal (O. rossi)
Harp seal (P. groenlandicus)
Spotted seal (P. largha)
Harbor seal (P. vitulina)
Caspian seal (P. caspica)
Ringed seal (P. hispida)
Baikal seal (P. sibirica)
Large family listed below
Large family listed below
Canidae (includes dogs)
Short-eared dog (A. microtis)
Side-striped jackal (C. adustus)
African golden wolf
African golden wolf (C. anthus)
Golden jackal (C. aureus)
Coyote (C. latrans)
Gray wolf (C. lupus)
Black-backed jackal (C. mesomelas)
Red wolf (C. rufus)
Ethiopian wolf (C. simensis)
Crab-eating fox (C. thous)
Maned wolf (C. brachyurus)
Dhole (C. alpinus)
Culpeo (L. culpaeus)
Darwin's fox (L. fulvipes)
South American gray fox
South American gray fox (L. griseus)
Pampas fox (L. gymnocercus)
Sechuran fox (L. sechurae)
Hoary fox (L. vetulus)
African wild dog
African wild dog (L. pictus)
Raccoon dog (N. procyonoides)
Bat-eared fox (O. megalotis)
Bush dog (S. venaticus)
Gray fox (U. cinereoargenteus)
Island fox (U. littoralis)
Bengal fox (V. bengalensis)
Blanford's fox (V. cana)
Cape fox (V. chama)
Corsac fox (V. corsac)
Tibetan sand fox
Tibetan sand fox (V. ferrilata)
Arctic fox (V. lagopus)
Kit fox (V. macrotis)
Pale fox (V. pallida)
Rüppell's fox (V. rueppelli)
Swift fox (V. velox)
Red fox (V. vulpes)
Fennec fox (V. zerda)
African clawless otter
African clawless otter (A. capensis)
Oriental small-clawed otter
Oriental small-clawed otter (A. cinerea)
Sea otter (E. lutris)
Spotted-necked otter (H. maculicollis)
North American river otter
North American river otter (L. canadensis)
Marine otter (L. felina)
Neotropical otter (L. longicaudis)
Southern river otter
Southern river otter (L. provocax)
Eurasian otter (L. lutra)
Hairy-nosed otter (L. sumatrana)
Smooth-coated otter (L. perspicillata)
Giant otter (P. brasiliensis)
Hog badger (A. collaris)
Tayra (E. barbara)
Lesser grison (G. cuja)
Greater grison (G. vittata)
Wolverine (G. gulo)
Saharan striped polecat
Saharan striped polecat (I. libyca)
Striped polecat (I. striatus)
Patagonian weasel (L. patagonicus)
American marten (M. americana)
Yellow-throated marten (M. flavigula)
Beech marten (M. foina)
Nilgiri marten (M. gwatkinsii)
European pine marten
European pine marten (M. martes)
Japanese marten (M. melampus)
Sable (M. zibellina)
Fisher (P. pennanti)
Japanese badger (M. anakuma)
Asian badger (M. leucurus)
European badger (M. meles)
Honey badger (M. capensis)
Bornean ferret-badger (M. everetti)
Chinese ferret-badger (M. moschata)
Javan ferret-badger (M. orientalis)
Burmese ferret-badger (M. personata)
(Weasels and Ferrets)
Amazon weasel (M. africana)
Mountain weasel (M. altaica)
Stoat (M. erminea)
Steppe polecat (M. eversmannii)
Colombian weasel (M. felipei)
Long-tailed weasel (M. frenata)
Japanese weasel (M. itatsi)
Yellow-bellied weasel (M. kathiah)
European mink (M. lutreola)
Indonesian mountain weasel
Indonesian mountain weasel (M. lutreolina)
Black-footed ferret (M. nigripes)
Least weasel (M. nivalis)
Malayan weasel (M. nudipes)
European polecat (M. putorius)
Siberian weasel (M. sibirica)
Back-striped weasel (M. strigidorsa)
Egyptian weasel (M. subpalmata)
American mink (N. vison)
African striped weasel
African striped weasel (P. albinucha)
American badger (T. taxus)
Marbled polecat (V. peregusna)
Law of heraldic arms
Grant of arms
Officers of arms
King of Arms, Herald, Pursuivant
Private Officer of Arms
Conventional elements of coats of arms
Slogan (battle cry)
coats of arms
National coat of arms
Augmentation of honour
Sanguine (blood red)
Crowns and coronets
Mantle and pavilion
List of oldest heraldry
Heraldic flag (Banner of arms)
Women in heraldry
1 Non-traditional, rarely used traditions in italic (typically
regional or modern, considered unheraldic by some)
Fauna Europaea: 305289