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Gray wolf+ The gray wolf or grey wolf (Canis lupus), also known as the timber wolf or western wolf, is a canid native to the wilderness and remote areas of Eurasia and North America. The '''gray wolf''' or '''grey wolf''' (''Canis lupus'' is a canid+ native to the wilderness and remote areas of Eurasia+ and North America+. 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,Heptner, V.G. The gray wolf is one of the world's best known and well researched animals, with probably more books written about it than any other wildlife species.
Based on morphology from China, the Pliocene+ wolf ''Canis chihliensis'' may have been the ancestor for both ''Canis armbrusteri'' and ''Canis lupus'' before their migration into North America. In 2010, a study compared the mDNA+ haplotype+s of 947 modern gray wolves from across Europe with the published sequences of 24 Pleistocene+ wolves from western Europe dated between 1,200-44,000 years BP.
Dog+s and gray wolves do not voluntarily interbreed in the wild, though they can produce fertile wolf-dog+ offspring. The gray wolf has interbred extensively with the eastern wolf+ producing a hybrid population termed Great Lakes boreal wolves+. 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. The gray wolf is the largest extant member of the Canidae+, excepting certain large breeds of domestic dog. The gray wolf has very dense and fluffy winter fur, with short underfur and long, coarse guard hair+s. 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. The gray wolf is a social animal, whose basic social unit consists of a mated pair, accompanied by the pair's adult offspring. 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. Wolves defend their territories from other packs through a combination of scent marking+, direct attacks and howling (see Communication+). The gray wolf is generally monogamous+, with mated pairs usually remaining together for life, unless one of the pair dies. 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. 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 muskox+en unaided. Two gray wolves eating a white-tailed deer+Once prey is brought down, wolves begin to feed ex The breeding pair typically monopolizes food in order to continue producing pups. 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. 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. 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. Other vocalisations of wolves are usually divided into three categories: growls, barks and whines.
Olfaction+ is probably the wolf's most acute sense, and plays a fundamental role in communication. The gray wolf is a habitat generalist, and can occur in desert+s, grassland+s, forest+s and arctic tundra+s. Although wolves primarily feed on medium to large sized ungulate+s, they are not fussy eaters. Brown bear+s typically dominate wolf packs in disputes over carcasses, while wolf packs mostly prevail against bears when defending their den sites. Wolves may encounter striped hyena+s in Israel, Central Asia and India, usually in disputes over carcasses. The gray wolf was once one of the world's most widely distributed mammals, living throughout the northern hemisphere north of 15°N latitude in North America and 12°N in India. 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.Boitani, L. Old English literature+ contains several instances of Anglo-Saxon kings+ and warriors+ taking on ''wulf'' as a prefix+ or suffix+ in their names.
The study found that phylogenetically+ the haplotypes represented two haplogroups+ that were separated by 5 mutational steps and referred to these as haplogroup 1 and 2. In North America, black colored wolves+ acquired their coloration from wolf-dog hybridization, which occurred 10,000–15,000 years ago. Females tend to have narrower muzzles and foreheads, thinner necks, slightly shorter legs and less massive shoulders than males. 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 East+ern and South Asia+n cousins. Most of the underfur and some of the guard hairs are shed in the spring and grow back in the autumn period. Triggers for dispersal include the onset of sexual maturity+ and competition within the pack for food. Scent marking is used for territorial advertisement, and involves urination, defecation and ground scratching.
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. 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. 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 km. Wolves typically locate their prey by scent, though they must usually be directly downwind of it. 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.
This is the most critical stage of the hunt, as wolves may never catch up with prey running at top speed.
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. With large prey, mature wolves usually avoid attacking frontally, instead focusing on the rear and sides of the animal.
When food is scarce, this is done at the expense of other family members, especially non-pups. While less gregarious canids generally possess simple repertoires of visual signals, wolves have more varied signals that subtly inter grade in intensity.
Barking has a fundamental frequency between 320–904 Hz, and is usually emitted by startled wolves. The wolf has a large number of apocrine sweat gland+s on the face, lips, back, and between the toes.
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 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. Smaller sized animals that may supplement the diet of wolves include marmot+s, hare+s, badger+s, fox+es, weasel+s, ground squirrel+s, mice+, hamster+s, vole+s and other rodent+s, as well as insectivore+s. They willingly eat the berries+ of mountain ash+, lily of the valley+, bilberries+, blueberries+ and cowberry+. In North America, incidents of gray wolves killing coyote+s are common, particularly in winter, when coyotes feed on wolf kills.
Striped hyenas feed extensively on wolf-killed carcasses in areas where the two species interact.
Wolves encounter cougar+s along portions of the Rocky Mountains+ and adjacent mountain ranges. Wolverines usually wait until the wolves are done feeding, but have been known to drive away wolves from kills. 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+. However, deliberate human persecution has reduced the species' range to about one third, because of livestock predation and fear over attacks on humans. The glacier is situated in the Mission Mountains at an elevation of 8,000 feet (2,400 m) above sea level and is immediately northeast of Gray Wolf Peak. Examples include Wulfhere, Cynewulf, Ceonwulf, Wulfheard, Earnwulf, Wulfm?r, Wulfstan and ?thelwulf. The dog is a divergent subspecies of the gray wolf and was derived from a now-extinct population of Late Pleistocene wolves, and the dog and the dingo are not separate species. The 947 European wolves revealed 27 different haplotypes with haplogroup 1 forming a monophyletic+ clade+, and all other haplotypes forming haplogroup 2. 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.
Nevertheless, coyote genetic markers have been found in some wild isolated gray wolf populations in the southern United States. The gray wolf's head is large and heavy, with a wide forehead, strong jaws and a long, blunt muzzle. Scent marks are generally left every 240 metres throughout the territory on regular travelways and junctions.
Females are capable of producing pups every year, with one litter+ annually being the average. Because of this, it rarely manages to capture hidden hares or birds, though it can easily follow fresh tracks. When a breeze carrying the prey's scent is located, the wolves stand alert, and point their eyes, ears and nose towards their target. Should this occur, the wolves hold back, as they require the stimulus of a running animal to proceed with an attack. 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. Large prey, such as moose, is killed by biting large chunks of flesh from the soft perineum+ area, causing massive blood loss+.
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. 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.
The results suggested that that the facial color pattern of canid species is related to their gaze communication, and that especially gray wolves use the gaze signal in conspecific communication. Wolves do not bark as loudly or continuously as dogs do, but bark a few times and retreat from perceived danger. The odor produced by these glands varies according to the individual wolf's microflora+ and diet, giving each a distinct "odor fingerprint". In cold climates, the gray wolf can reduce the flow of blood near its skin to conserve body heat. Other prey species include reindeer+, argali+, mouflon+, wisent+, saiga+, ibex+, chamois+, wild goat+s, fallow deer+ and musk deer+. Wolves may attack coyote den sites, digging out and killing their pups, though rarely eating them.
One-to-one, hyenas dominate wolves, but wolf packs can drive off single or outnumbered hyenas. Wolves and cougars typically avoid encountering each other by hunting on different elevations. Wolves appear capable of escaping competitive exclusion+ from tigers only when human persecution decreases tiger numbers. The species is now extinct in much of Western Europe+, in Mexico and much of the United States. 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). However, one genetic study undertaken in the Caucasus Mountains+ showed that as many as 10% of dogs in the area, including livestock guardian dog+s, are first generation hybrids.
Especially long hairs are found on the shoulders, and almost form a crest on the upper part of the neck.
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.
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. Such markers can last for 2–3 weeks, and are typically placed near rocks, boulders, trees or the skeletons of large animals. 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. 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. In open areas, wolves may precede the hunt with group ceremonies involving standing nose-to-nose and wagging their tails. If the targeted animal stands its ground, the wolves either ignore it, or try to intimidate it into running. 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. Such bites can cause wounds 10–15 cm in length, with three such bites to the perineum usually being sufficient to bring down a large deer in optimum health. 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. Postural communication in wolves consists of a variety of facial expressions, tail positions and piloerection+. Growling has a fundamental frequency of 380–450 Hz, and is usually emitted during food challenges.
A combination of apocrine and eccrine sweat gland+s 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 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.
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. When such foods are insufficient, they prey on lizard+s, snake+s, frog+s, rarely toad+s and large insect+s as available.
There are no records of coyotes killing wolves, though coyotes may chase wolves if they outnumber them. Wolf interactions with American black bear+s are much rarer than with brown bears, because of differences in habitat preferences. In winter, however, when snow accumulation forces their prey into valleys, interactions between the two species become more likely. 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. In modern times, the gray wolf occurs mostly in wilderness and remote areas, particularly in Canada, Alaska+ and northern USA, Europe+, and Asia+ from about 75°N to 12°N. Arcing northeast from the Mount Deception - Needles - Mount Clark - Mount Walkinshaw massif, Gray Wolf Ridge is covered by alpine and subalpine forests up to the timberline, leaving the upper elevations of the ridge's six rocky peaks vegetated only by fragile alpine meadows.
The captive breeding+ of wolf-dog+ hybrids has proliferated in the USA, with 300,000 such animals being present there. In tests performed on a putative chupacabra+ carcass, 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. 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 hyena+s. The smallest territory on record was held by a pack of six wolves in northeastern Minnesota, which occupied an estimated , while the largest was held by an Alaskan pack of ten wolves encompassing a area. 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. Such gray wolves are termed "Casanova+ wolves" and, unlike males from established packs, they do not form pair bond+s with the females they mate with. Estrus+ typically occurs in late winter, with older, multiparous females entering estrus 2–3 weeks earlier than younger females.
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. Wolves typically commence feeding by consuming the larger internal organs of their prey, such as the heart+, liver+, lungs+ and stomach+ lining.
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. The follicle+s present on the guard hairs from the wolf's back have clusters of apocrine and sebaceous gland+s at their bases. 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. 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.
In times of scarcity, wolves readily eat carrion+, visiting cattle burial grounds and slaughter house+s. A well-fed wolf stores fat+ under the skin, around the heart, intestines, kidneys, and bone marrow, particularly during the autumn and winter. In Asia, they may compete with dhole+s, though there is at least one record of a lone wolf associating with a pair of dholes in Debrigarh Wildlife Sanctuary+.
The majority of black bear encounters with wolves occur in the species' northern range, with no interactions being recorded in Mexico.
Two cases are known however from southern Israel, where wolves and striped hyenas associated closely with each other in an apparently amicable way.
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. In 2013, a captive breeding experiment in Utah between gray wolves and western coyotes produced six hybrids through artificial insemination+, making this the very first hybridization case between pure coyotes and northwestern gray wolves+. 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. Wolf packs are typically settled, and usually only leave their accustomed ranges during severe food shortages. Some gray wolf packs may have multiple breeding females this way, as is the case in Yellowstone National Park+. 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.
With small, mouse+-like prey, wolves leap in a high arc and immobilize it with their forepaws. 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. 749-788 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''". One variation of the howl is accompanied by a high pitched whine, which precedes a lunging attack.
As the skin on the back is usually folded, this provides a microclimate+ for bacterial propagation around the glands. During the autumn-spring period, when wolves are more active, they willingly lie out in the open, whatever their location. Digestion+ only takes a few hours, thus wolves can feed several times in one day, making quick use of large quantities of meat. Wolves have been recorded on numerous occasions to actively seek out black bears in their dens and kill them without eating them. 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.
It is nonetheless closely related enough to smaller ''Canis'' species, such as the eastern wolf+, coyote+, posing a serious threat to it. At six months of age, the hybrids were closely monitored and were shown to display both physical and behavioral characteristics from both species.
Short, elastic and closely adjacent hairs are present on the limbs from the elbow+s down to the calcaneal tendon+s. Gray wolves also practice alloparental care+, in which a wolf pair may adopt the pup or pups of another.
Old females usually whelp+ in the den of their previous litter, while younger females typically den near their birthplace. Pups almost never howl, while yearling wolves produce howls ending in a series of dog-like yelps. Whining is associated with situations of anxiety, curiosity, inquiry and intimacy such as greeting, feeding pups and playing.
During piloerection, the guard hairs on the back are raised and the skin folds spread, thus releasing scent.
Subspecies of Canis lupus+ Canis lupus has 40 subspecies currently described, including the dingo, Canis lupus dingo, and the domestic dog, Canis lupus familiaris, and many subspecies of wolf throughout the Northern Hemisphere.
It feeds primarily on large ungulate+s, though it also eats smaller animals, livestock, carrion, and garbage.Mech, L. Although hybridization between wolves and golden jackal+s has never been observed, evidence of such occurrences was discovered through mtDNA analysis on jackals in Bulgaria.
The gray wolf usually carries its head at the same level as the back, raising it only when alert.
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. During times of ungulate abundance (migration, calving etc.), different wolf packs may temporarily join forces.
This might take place if the original parents die or are for some reason separated from them. 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+.
Howling consists of a fundamental frequency that may lie between 150 and 780 Hz, and consists of up to 12 harmonically related overtones.
When building dens, females make use of natural shelters such as fissures in rocks, cliffs overhanging riverbanks and holes thickly covered by vegetation. While encounters with brown and black bears appear to be common, polar bear+s are rarely encountered by wolves, though there are two records of wolf packs killing polar bear cubs. 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 phenotype+s, to the point of being mistaken for wolves by trained biologists.
Wolf fur provides better insulation than dog fur, and does not collect ice when warm breath is condensed against it. In addition to heterosexual behavior, homosexual behavior has been observed in gray wolves. 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.
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. The pitch usually remains constant or varies smoothly, and may change direction as many as four or five times.
Sometimes, the den is the appropriated burrow of smaller animals such as foxes, badgers or marmots. 3-12, Edmonton: Canadian Circumpolar Institute 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+. Male gray wolves often mount each other+ when the highest ranking female in the pack comes into heat. 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. When wolves are together, they commonly indulge in behaviors such as nose pushing, jaw wrestling, cheek rubbing and facial licking. Howls used for calling pack mates to a kill are long, smooth sounds similar to the beginning of the cry of a horned owl+. Female wolves tend to have smoother furred limbs than males, and generally develop the smoothest overall coats as they age. The mouthing of each other's muzzles is a friendly gesture, while clamping on the muzzle with bared teeth is a dominance display. On rare occasions, female wolves dig burrows themselves, which are usually small and short with 1–3 openings. Older wolves generally have more white hairs in the tip of the tail, along the nose and on the forehead. Its exact function is debated, though most researchers agree that its primary purpose is to establish boundaries. The den is usually constructed not more than 500 metres away from a water source, and typically faces southwards, thus ensuring enough sunlight exposure, keeping the denning area relatively snow free.
The winter fur is retained longest in lactating females, though with some hair loss around their nipples. 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. Wolves urine mark more frequently and vigorously in unfamiliar areas, or areas of intrusion, where the scent of other wolves or canids is present. Resting places, play areas for the pups and food remains are commonly found around wolf dens. Wolves may kill lynxes by running them down, or killing them before they can escape into trees. 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.
The odour of urine and rotting food emanating from the denning area often attracts scavenging birds such as magpie+s and raven+s. 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).
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. Only dominant wolves typically use RLU, with subordinate males continuing to use the juvenile standing posture throughout adulthood. As there are few convenient places for burrows, wolf dens are usually occupied by animals of the same family. The two are however mutually intelligible+, as North American wolves have been recorded to respond to European-style howls made by biologists.
RLU is considered to be one of the most important forms of scent communication in the wolf, making up 60–80% of all scent marks observed. Though they mostly avoid areas within human sight, wolves have been known to nest near domiciles+, paved road+s and railway+s. Wolf packs in Astrakhan+ hunt Caspian seal+s on the Caspian Sea+ coastline and some wolf packs in Alaska and Western Canada have been observed to feed on salmon+. Other primate+s occasionally taken by wolves include grey langur+s in Nepal and hamadryas baboon+s in Saudi Arabia.

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