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The gray wolf usually carries its head at the same level as the back, raising it only when alert. Generally, wolves have a high heart weight of 0. Tibetan gray wolves, which occupy territories up to 3, above sea level , have evolved hearts that withstand the low oxygen levels. The gray wolf's head is large and heavy, with a wide forehead, strong jaws and a long, blunt muzzle.
This force is sufficient to break open most bones. A similar trend was found with the carnassial tooth bite force, but with the extinct dire wolf and gray wolf both measuring , then followed by the African hunting dog , the dhole , and the dingo The gray wolf has very dense and fluffy winter fur, with short underfur and long, coarse guard hairs. 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.
Wolf fur provides better insulation than dog fur, and does not collect ice when warm breath is condensed against it. 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. 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. 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.
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.
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.
Such markers can last for 2—3 weeks,  and are typically placed near rocks, boulders, trees, or the skeletons of large animals. 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. 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.
This might take place if the original parents die or are for some reason separated from them. The age of first breeding in gray wolves depends largely on environmental factors: This is further demonstrated by the fact that captive wolves have been known to breed as soon as they reach 9—10 months, while the youngest recorded breeding wolves in the wild were 2 years old.
Females are capable of producing pups every year, with one litter annually being the average. Unlike the coyote, the gray wolf never reaches reproductive senescence. 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 milk canines erupt after one month. Pups first leave the den after 3 weeks. 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: 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. 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.
Because of this, it rarely manages to capture hidden hares or birds, though it can easily follow fresh tracks.
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. With small, mouse -like prey, wolves leap in a high arc and immobilize it with their forepaws. 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.
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.
When food is scarce, this is done at the expense of other family members, especially non-pups. 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.
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. 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. 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 , a study compared the facial color pattern across 25 canid species.
The results suggested that the facial color pattern of canid species is related to their gaze communication, and that especially gray wolves use the gaze signal in conspecific communication. 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.
Pups almost never howl, while yearling wolves produce howls ending in a series of dog-like yelps. The pitch usually remains constant or varies smoothly, and may change direction as many as four or five times.
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. The two are however mutually intelligible , as North American wolves have been recorded to respond to European-style howls made by biologists. Other vocalisations of wolves are usually divided into three categories: Wolves do not bark as loudly or continuously as dogs do, but bark a few times and retreat from perceived danger.
Pups commonly growl when playing. One variation of the howl is accompanied by a high pitched whine, which precedes a lunging attack. Olfaction is probably the wolf's most acute sense, and plays a fundamental role in communication. The wolf has a large number of apocrine sweat glands on the face, lips, back, and between the toes. The odor produced by these glands varies according to the individual wolf's microflora and diet, giving each a distinct "odor fingerprint".
A combination of apocrine and eccrine sweat glands on the feet allows the wolf to deposit its scent whilst scratching the ground, which usually occurs after urine marking and defecation during the breeding season. The follicles present on the guard hairs from the wolf's back have clusters of apocrine and sebaceous glands at their bases.
As the skin on the back is usually folded, this provides a microclimate for bacterial propagation around the glands. During piloerection, the guard hairs on the back are raised and the skin folds spread, thus releasing scent. The pre-caudal scent glands may play a role in expressing aggression, as combative wolves raise the base of their tails whilst drooping the tip, thus positioning the scent glands at the highest point.
The 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. 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. 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.
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 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.
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. 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. 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. 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.
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. 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.
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. 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 may kill lynxes by running them down, or killing them before they can escape into trees. 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.
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.
However, deliberate human persecution has reduced the species' range to about one third, because of livestock predation and fear of attacks on humans. Wolf population declines have been arrested since the s, and have fostered recolonization and reintroduction in parts of its former range, due to legal protection, changes in land-use and rural human population shifts to cities.
Competition with humans for livestock and game species, concerns over the danger posed by wolves to people, and habitat fragmentation pose a continued threat to the species. Despite these threats, because of the gray wolf's relatively widespread range and stable population, it is classified as least concern by the IUCN. 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 19th century. In England , wolf persecution was enforced by legislation , and the last wolf was killed in the early sixteenth century during the reign of Henry VII.
Wolves lasted longer in Scotland , where they sheltered in vast tracts of forest, which were subsequently burned down. Wolves managed to survive in the forests of Braemar and Sutherland until The extirpation of wolves in Ireland followed a similar course, with the last wolf believed to have been killed in The Sami extirpated wolves in northern Sweden in organized drives.
By , few wolves remained in Sweden, because of the use of snowmobiles in hunting them, with the last specimen being killed in The gray wolf was exterminated in Denmark in and Norway's last wolf was killed in 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 , 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 , and had disappeared from the Rhine regions by Today, wolves have returned to the area. The louveterie was abolished after the French Revolution in , but was re-established in In , up to 1, wolves were killed, with many more by poison.
In Eastern Europe , wolves were never fully exterminated, because of the area's contiguity with Asia and its large forested areas. However, Eastern European wolf populations were reduced to very low numbers by the late nineteenth century.
Wolves were extirpated in Slovakia during the first decade of the twentieth century and, by the mid-twentieth century, could only be found in a few forested areas in eastern Poland.
Wolves in the eastern Balkans benefitted from the region's contiguity with the former Soviet Union and large areas of plains, mountains and farmlands. Wolves in Hungary occurred in only half the country around the start of the 20th century, and were largely restricted to the Carpathian Basin. Wolf populations in Romania remained largely substantial, with an average of 2, wolves being killed annually out of a population of 4, from — An all-time low was reached in , when the population was reduced to 1, animals.
The extermination of wolves in Bulgaria was relatively recent, as a previous population of about 1, animals in was reduced to about — in In Greece, the species disappeared from the southern Peloponnese in Despite periods of intense hunting during the eighteenth century, wolves never disappeared in the western Balkans, from Albania to the former Yugoslavia.
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 Wolf bounties were regularly paid in Italy as late as The recovery of European wolf populations began after the s, when traditional pastoral and rural economies declined and thus removed the need to heavily persecute wolves.
By the s, 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 , and now holds a population of over 1,, By the wolves in the Western Alps imposed a significant burden on traditional sheep and goat husbandry with a loss of over 5, animals in In Spain , the species occurs in Galicia , Leon , and Asturias.
Although hundreds of Iberian wolves are illegally killed annually, the population has expanded south across the river Duero and east to the Asturias and Pyrenees Mountains. In , wolves began recolonising central Sweden after a twelve-year absence, and have since expanded into southern Norway. As of , 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. Furthermore, the decline in the moose populations has reduced the wolf's food supply.