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Social and territorial behaviorsEdit

Although some wolves are solitary, most are highly gregarious animals. The basic social unit of a wolf pack is the mated pair, accompanied by the pair's adult offspring. In ideal conditions, the mated pair produces pups every year, with such offspring typically staying in the pack for 10–54 months before dispersing. 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 42 wolves being known.
WolfPack01

Triggers for dispersal include the onset of sexual maturity and competition within the pack for food. A new pack is usually founded by an unrelated 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 or run them out of the territory. 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 or 15 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 territory unless desperate, due to the possibility of fatal encounters with neighboring packs. The smallest territory on record was held by a pack of six wolves in northeastern Minnesota, which occupied an estimated 33 km2 (13 sq mi), While the largest was held by an Alaskan pack of ten wolves encompassing a 6,272 km2 (2,422 sq mi) area. Wolf packs are typically settled, and usually only leave their accustomed ranges during severe food shortages. Wolves defend their territories from other packs through a combination of scent marking, direct attacks and howling (see Communication).

Scent marking is used for territorial advertisement, and involves urination, defecation and ground scratching. Scent marks are generally left every 240 metres 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. When scent marking and howling fail to deter strange wolf packs from entering another's territory, violent interactions can ensue. Territorial fights are among the principal causes of wolf mortality: one study on wolf mortality in Minnesota and the Denali National Park and Preserve concluded that 14–65% of wolf deaths were due to predation by other wolves.

Hunting and Feeding BehaviorEdit

WolfHunt01
Although social animals, single wolves or mated pairs typically have higher success rates in hunting than do large packs. Single wolves having occasionally been observed to kill large prey such as moose, bison and muskoxen unaided. A wolf hunt can be divided into five stages which are the following: 
  • Locating Prey
  • Stalking
  • Encounter
  • Rush
  • Chasing


Locating prey: Where the wolves travel in search of prey through their power of scent, chance encounter, and tracking. Wolves typically locate their prey by scent, though they must usually be directly downwind of it. When a breeze carrying the prey's scent is located, the wolves stand alert, and point their eyes, ears and nose towards their target. In open areas, wolves may precede the hunt with group ceremonies involving standing nose-to-nose and wagging their tails. Once concluded, the wolves head towards their prey.


The stalk: The wolves attempt to conceal themselves as they approach. As the gap between the wolves and their prey closes, the wolves quicken their pace, wag their tails, and peer intently, getting as close to their quarry as possible without making it flee.


The encounter: Once the prey detects the wolves, it can either approach the wolves, stand its ground, or flee. Large prey, such as moose, elk, and muskoxen, usually stand their ground. Should this occur, the wolves hold back, as they require the stimulus of a running animal to proceed with an attack. If the targeted animal stands its ground, the wolves either ignore it, or try to intimidate it into running.


The rush: If the prey attempts to flee, the wolves immediately pursue it. This is the most critical stage of the hunt, as wolves may never catch up with prey running at top speed. If their prey is travelling in a group, the wolves either attempt to break up the herd, or isolate one or two animals from it.


The chase: A continuation of the rush, the wolves attempt to catch up with their prey and kill it. When chasing small prey, wolves attempt to catch up with their prey as soon as possible, while with larger animals, the chase is prolonged, in order to wear the selected prey out. Wolves usually give up chases after 1–2 km (0.62–1.3 mi), though one wolf was recorded to chase a deer for 21 km (13 mi). Both Russian and North American wolves have been observed to drive prey onto crusted ice, precipices, ravines, slopes and steep banks to slow them down.


Illustration (1909) of wolves killing a caribou 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 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 imobilize 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. Surplus killing may also occur when adult wolves are teaching their young to hunt. 


WolfFeasting01
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.

Denning and Sheltering BehaviorEdit

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.Wolves do not line their denning places, a likely precaution against parasites. The den is usually constructed not more than 500 meters 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 odor 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.

After the puppies are too large to inhabit the den, they are moved to a rendezvous site, where they stay while the pack is away hunting. Now considered sub-adults, the wolves will never use a den again in their life, unless one of them later becomes a mother herself. Dens are only used for birthing puppies, not as a sleeping area or to escape the weather as you may think. In captivity, wolves will occasionally use dens to escape dominance from other members, but this is not a natural behavior, rather an ingenious way to use the available environment to cope with the negative aspects of captivity.

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