NamingThe English 'wolf' stems from the Old English ''wulf'', which is itself thought to be derived from the Proto-Germanic ''*wulfaz''. The Latin ''lupus'' is a Sabine loanword. Both derive from the Proto-Indo-European root ''*wlqwos'' or ''*lukwos''.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 , Wolfhetan , Isangrim , Scrutolf , Wolfgang and Wolfdregil .
Ancient Greek literature is similar: Autolycus , Lycurgus .
The Latin for "female prostitute" is ''lupa'', and the most famous brothel in Pompeii was the Lupanar.
DistributionIn 2013, a genetic study found that the wolf population in Europe was divided along a north-south axis and formed five major clusters. Three clusters were identified occupying southern and central Europe in Italy, the Dinaric-Balkans, the Carpathians. Another two clusters were identified occupying north-central Europe and the Ukrainian steppe. The Italian wolf consisted of an isolated population with low genetic diversity. Wolves from Croatia, Bulgaria, and Greece formed the Dinaric-Balkans cluster. Wolves from Finland, Latvia, Belarus, Poland and Russia formed the north-central Europe cluster, with wolves from the Carpathians cluster coming from a mixture of wolves from the north-central cluster and the Dinaric-Balkans cluster. The wolves from the Carpathians were more similar to the wolves from the Pontic-Caspian Steppe than they were to wolves from north-central Europe. These clusters may have been the result of expansion from glacial refugia, an adaptation to local environments, and landscape fragmentation and the killing of wolves in some areas by humans.
In 2016, two genetic studies of North American gray wolves found that they formed six ecotypes – genetically and ecologically distinct populations separated from other populations by their different type of habitat. These six wolf ecotypes were named West Forest, Boreal Forest, Arctic, High Arctic, Baffin, and British Columbia. The studies found that precipitation and mean diurnal temperature range were the most influential variables. These findings were in accord with previous studies that precipitation influenced morphology and that vegetation and habitat type influenced wolf differences. The local adaptation of a wolf ecotype most likely reflects a wolf's preference to remain in the type of habitat that it was born into.
StatusThe gray wolf was once one of the world's most widely distributed mammals, living throughout the northern hemisphere north of 15°N latitude in North America and 12°N in India. However, deliberate human persecution has reduced the species' range to about one third, because of livestock predation and fear of attacks on humans. The species is now extinct in much of Western Europe, in Mexico and much of the United States. In modern times, the gray wolf occurs mostly in wilderness and remote areas, particularly in Canada, Alaska and northern USA, Europe, and Asia from about 75°N to 12°N. Wolf population declines have been arrested since the 1970s, and have fostered recolonization and reintroduction in parts of its former range, due to legal protection, changes in land-use and rural human population shifts to cities. Competition with humans for livestock and game species, concerns over the danger posed by wolves to people, and habitat fragmentation pose a continued threat to the species. Despite these threats, because of the gray wolf's relatively widespread range and stable population, it is classified as least concern by the IUCN.
BehaviorThe gray wolf is a social animal, whose basic social unit consists of a mated pair, accompanied by the pair's adult offspring.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 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. The average pack consists of a family of 5–11 animals , 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 , 390 km , and 670 km 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 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 , different wolf packs may temporarily join forces. Studies of wolves' cortisol levels show that they rise significantly when a pack member dies, indicating the presence of stress.
Wolves are highly territorial animals, and generally establish territories far larger than they require to survive in order to assure a steady supply of prey. Territory size depends largely on the amount of prey available and the age of the pack's pups, tending to increase in size in areas with low prey populations or when the pups reach the age of 6 months, thus having the same nutritional needs as adults. Wolf packs travel constantly in search of prey, covering roughly 9% of their territory per day ). The core of their territory is on average 35 km2 , 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 , while the largest was held by an Alaskan pack of ten wolves encompassing a 6,272 km2 area. Wolf packs are typically settled, and usually only leave their accustomed ranges during severe food shortages.
Wolves defend their territories from other packs through a combination of scent marking, direct attacks and howling . Scent marking is used for territorial advertisement, and involves urination, defecation and ground scratching. Scent marks are generally left every 240 m 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.
HabitatThe 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 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.
ReproductionThe gray wolf is generally monogamous, with mated pairs usually remaining together for life. Upon the death of one mated wolf, pairs are quickly re-established. Since males often predominate in any given wolf population, unpaired females are a rarity. If a dispersing male gray wolf is unable to establish a territory or find a mate, he mates with the daughters of already established breeding pairs from other packs. Such gray wolves are termed "Casanova wolves" and, unlike males from established packs, they do not form pair bonds with the females they mate with. Some gray wolf packs may have multiple breeding females this way, as is the case in Yellowstone National Park. Gray wolves also practice alloparental care, in which a wolf pair may adopt the pup or pups of another. This might take place if the original parents die or are for some reason separated from them. In addition to heterosexual behavior, homosexual behavior has been observed in gray wolves. Male gray wolves often mount each other when the highest ranking female in the pack comes into heat.
The age of first breeding in gray wolves depends largely on environmental factors: when food is plentiful, or when wolf populations are heavily managed, wolves can rear pups at younger ages in order to better exploit abundant resources. This is further demonstrated by the fact that captive wolves have been known to breed as soon as they reach 9–10 months, while the youngest recorded breeding wolves in the wild were 2 years old. Females are capable of producing pups every year, with one litter annually being the average. Unlike the coyote, the gray wolf never reaches reproductive senescence. Estrus typically occurs in late winter, with older, multiparous females entering estrus 2–3 weeks earlier than younger females. During pregnancy, female wolves remain in a den located away from the peripheral zone of their territories, where violent encounters with other packs are more likely. Old females usually whelp in the den of their previous litter, while younger females typically den near their birthplace. The gestation period lasts 62–75 days, with pups usually being born in the summer period.
Wolves bear relatively large pups in small litters compared to other canid species. The average litter consists of 5–6 pups, with litter sizes tending to increase in areas where prey is abundant, though exceptionally large litters of 14–17 pups occur only 1% of the time. Pups are usually born in spring, coinciding with a corresponding increase in prey populations. Pups are born blind and deaf, and are covered in short soft grayish-brown fur. They weigh 300–500 g 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.
FoodGlobally, gray wolf diet is predominantly composed of large ) and medium-sized ) 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 moose. Digestion only takes a few hours, thus wolves can feed several times in one day, making quick use of large quantities of meat.
Although wolves primarily feed on medium to large sized ungulates, they are not fussy eaters. Smaller sized animals that may supplement the diet of wolves include marmots, hares, badgers, foxes, weasels, ground squirrels, mice, hamsters, voles and other rodents, as well as insectivores. They frequently eat waterfowl and their eggs. When such foods are insufficient, they prey on lizards, snakes, frogs, rarely toads and large insects as available. In times of scarcity, wolves readily eat carrion, visiting cattle burial grounds and slaughter houses. Cannibalism is not uncommon in wolves: during harsh winters, packs often attack weak or injured wolves, and may eat the bodies of dead pack members. Wolf packs in Astrakhan hunt Caspian seals on the Caspian Sea coastline and some wolf packs in Alaska and Western Canada have been observed to feed on salmon. Humans are rarely, but occasionally preyed upon. Other primates occasionally taken by wolves include grey langurs in Nepal and hamadryas baboons in Saudi Arabia.
In Eurasia, many gray wolf populations are forced to subsist largely on livestock and garbage in areas with dense human activity, though wild ungulates such as moose, red deer, roe deer and wild boar are still the most important food sources in Russia and the more mountainous regions of Eastern Europe. Other prey species include reindeer, argali, mouflon, wisent, saiga, ibex, chamois, wild goats, fallow deer and musk deer. The prey animals of North American wolves have largely continued to occupy suitable habitats with low human density, and cases of wolves subsisting largely on garbage or livestock are exceptional. Animals preferred as prey by North American wolves include moose, white-tailed deer, elk, mule deer, bighorn sheep, Dall's sheep, American bison, muskox and caribou.
Wolves supplement their diet with fruit and vegetable matter. They willingly eat the berries of mountain ash, lily of the valley, bilberries, blueberries and cowberry. Other fruits include nightshade, apples and pears. They readily visit melon fields during the summer months. A well-fed wolf stores fat under the skin, around the heart, intestines, kidneys, and bone marrow, particularly during the autumn and winter.
UsesWild 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 , so they can exercise.
CulturalThe wolf is a common motif in the foundational mythologies and cosmologies of peoples throughout Eurasia and North America . The obvious attribute of the wolf is its nature of a predator, and correspondingly it is strongly associated with danger and destruction, making it the symbol of the warrior on one hand, and that of the devil on the other. The modern trope of the Big Bad Wolf is a development of this. The wolf holds great importance in the cultures and religions of the nomadic peoples, both of the Eurasian steppe and of the North American Plains. In many cultures, the identification of the warrior with the wolf gave rise to the notion of Lycanthropy, the mythical or ritual identification of man and wolf.
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