Editing Wolf (section)

== Behaviour ==

{{anchor|Behaviour}}
{{See also|Dog behaviour}}

=== Social structure ===

{{See also|Pack (canine)#Pack behavior in grey wolves}}
[[File:Canis lupus pallipes Mysore Zoo 2.jpg|thumb|alt=Photograph of three wolves running and biting each other|Indian wolves at the [[Mysore Zoo]]]]

The wolf is a [[Sociality|social animal]].{{sfn|Heptner|Naumov|1998|pp=164–270}} Its populations consist of packs and lone wolves, most lone wolves being temporarily alone while they disperse from packs to form their own or join another one.{{sfn|Mech|Boitani|2003|p=164}} The wolf's basic social unit is the [[nuclear family]] consisting of a [[mated pair]] accompanied by their offspring.{{sfn|Heptner|Naumov|1998|pp=164–270}} The average pack size in North America is eight wolves and 5.5 in Europe.<ref name=Miklosi2015/> The average pack across Eurasia consists of a family of eight wolves (two adults, juveniles, and yearlings),{{sfn|Heptner|Naumov|1998|pp=164–270}} or sometimes two or three such families,<ref name=Paquet2003/> with examples of exceptionally large packs consisting of up to 42 wolves being known.{{sfn|Mech|Boitani|2003|pp=2–3, 28}} [[Cortisol]] levels in wolves rise significantly when a pack member dies, indicating the presence of stress.<ref name=Molnar2015/> During times of prey abundance caused by calving or migration, different wolf packs may join together temporarily.{{sfn|Heptner|Naumov|1998|pp=164–270}}

Offspring typically stay in the pack for 10–54 months before dispersing.{{sfn|Mech|Boitani|2003|pp=1–2}} Triggers for dispersal include the onset of [[sexual maturity]] and competition within the pack for food.{{sfn|Mech|Boitani|2003|pp=12–13}} 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 upwards of {{convert|206|km|mi|abbr=on}}, {{convert|390|km|mi|abbr=on}}, and {{convert|670|km|mi|abbr=on}} from their natal (birth) packs.<ref name=Nowak1983/> 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.{{sfn|Mech|Boitani|2003|p=38}} 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 of one to three years old, and unlikely to compete for breeding rights with the mated pair. This usually occurs between the months of February and May. Adopted males may mate with an available pack female and then form their own pack. In some cases, a lone wolf is adopted into a pack to replace a deceased breeder.{{sfn|Mech|Boitani|2003|pp=2–3, 28}}

Wolves are [[Territory (animal)|territorial]] and generally establish territories far larger than they require to survive assuring a steady supply of prey. Territory size depends largely on the amount of prey available and the age of the pack's pups. They tend to increase in size in areas with low prey populations,<ref name=Jedrzejewski2007/> or when the pups reach the age of six months when they have the same nutritional needs as adults.{{sfn|Mech|Boitani|2003|pp=19–26}} Wolf packs travel constantly in search of prey, covering roughly 9% of their territory per day, on average {{cvt|25|km/d}}. The core of their territory is on average {{cvt|35|km2}} where they spend 50% of their time.<ref name=Jedrzejewski2007/> Prey density tends to be much higher on the territory's periphery. Wolves tend to avoid hunting on the fringes of their range to avoid fatal confrontations with neighbouring packs.<ref name=Mech1977/> The smallest territory on record was held by a pack of six wolves in northeastern Minnesota, which occupied an estimated {{cvt|33|km2}}, while the largest was held by an Alaskan pack of ten wolves encompassing {{cvt|6,272|km2}}.{{sfn|Mech|Boitani|2003|pp=19–26}} Wolf packs are typically settled, and usually leave their accustomed ranges only during severe food shortages.{{sfn|Heptner|Naumov|1998|pp=164–270}} Territorial fights are among the principal causes of wolf mortality, one study concluding that 14–65% of wolf deaths in Minnesota and the [[Denali National Park and Preserve]] were due to other wolves.<ref name=Mech2003/>

===Communication===
{{main article|Wolf communication}}
{{listen
| filename  = Wolf howls.ogg
| title = Wolves howling
| format = [[Ogg]]
| filename2 = rallying.ogg
| title2 = Rallying cry
| format2 = [[Ogg]]
}}
Wolves communicate using vocalizations, body postures, scent, touch, and taste.{{sfn|Mech|Boitani|2003|pp=66–103}} The phases of the moon have no effect on wolf vocalization, and despite popular belief, wolves do not howl at the Moon.{{sfn|Busch|2007|p=59}} Wolves [[howling|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, while crossing unfamiliar territory, and to communicate across great distances.{{sfn|Lopez|1978|p=38}} Wolf howls can under certain conditions be heard over areas of up to {{cvt|130|km2}}.<ref name=Paquet2003/> Other vocalizations include [[growling|growls]], [[bark (sound)|barks]] and whines. Wolves do not bark as loudly or continuously as dogs do in confrontations, rather barking a few times and then retreating from a perceived danger.{{sfn|Lopez|1978|pp=39–41}} Aggressive or self-assertive wolves are characterized by their slow and deliberate movements, high body [[Posture (psychology)|posture]] and raised [[hackles]], while submissive ones carry their bodies low, flatten their fur, and lower their ears and tail.{{sfn|Mech|Boitani|2003|p=90}}

Scent marking involves urine, feces, and [[preputial gland|preputial]] and anal gland scents. This is more effective at advertising territory than howling and is often used in combination with scratch marks. Wolves increase their rate of scent marking when they encounter the marks of wolves from other packs. Lone wolves will rarely mark, but newly bonded pairs will scent mark the most.<ref name=Paquet2003/> These marks are generally left every {{cvt|240|m}} throughout the territory on regular travelways and junctions. Such markers can last for two to three weeks,{{sfn|Mech|Boitani|2003|pp=19–26}} and are typically placed near rocks, boulders, trees, or the skeletons of large animals.{{sfn|Heptner|Naumov|1998|pp=164–270}} Raised leg [[urination]] 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.<ref name=Peters1975/>

=== Reproduction{{anchor|Reproduction_and_development}} ===

{{See also|Canine reproduction}}
[[File:Korean wolves mating (cropped).jpg|left|thumb|alt=Photograph of a pair of mating wolves|Korean wolves mating in the [[Tama Zoological Park]], Japan]]
Wolves are [[Monogamy in animals|monogamous]], mated pairs usually remaining together for life. Should one of the pair die, another mate is found quickly.{{sfn|Heptner|Naumov|1998|p=248}} With wolves in the wild, inbreeding does not occur where outbreeding is possible.{{sfn|Mech|Boitani|2003|p=5}} Wolves become mature at the age of two years and sexually mature from the age of three years.{{sfn|Heptner|Naumov|1998|p=248}} The age of first breeding in wolves depends largely on environmental factors: when food is plentiful, or when wolf populations are heavily managed, wolves can rear pups at younger ages to better exploit abundant resources. Females are capable of producing pups every year, one [[Litter (zoology)|litter]] annually being the average.{{sfn|Mech|Boitani|2003|p=175}} [[Oestrus]] and [[Rut (mammalian reproduction)|rut]] begin in the second half of winter and lasts for two weeks.{{sfn|Heptner|Naumov|1998|p=248}}
[[File:Iberian Wolf 01.jpg|thumb|alt=Photograph of wolf pups stimulating their mother to regurgitate some food|[[Iberian wolf]] pups stimulating their mother to regurgitate some food]]

Dens are usually constructed for pups during the summer period. When building dens, females make use of natural shelters like 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 one to three openings. The den is usually constructed not more than {{cvt|500|m}} away from a water source. It typically faces southwards where it can be better warmed by sunlight exposure, and the snow can thaw more quickly. 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 like [[magpie]]s and [[raven]]s. Though they mostly avoid areas within human sight, wolves have been known to nest near homes, paved [[road]]s and [[railway]]s.{{sfn|Heptner|Naumov|1998|pp=234–238}} During pregnancy, female wolves remain in a den located away from the peripheral zone of their territories, where violent encounters with other packs are less likely to occur.{{sfn|Mech|Boitani|2003|pp=42–46}}

The [[gestation period]] lasts 62–75 days with pups usually being born in the spring months or early summer in very cold places such as on the tundra. Young females give birth to four to five young, and older females from six to eight young and up to 14. Their mortality rate is 60–80%.{{sfn|Heptner|Naumov|1998|pp=249–254}} Newborn wolf pups look similar to [[German Shepherd|German Shepherd Dog]] pups.{{sfn|Mech|Boitani|2003|p=47}} They are born blind and deaf and are covered in short soft greyish-brown fur. They weigh {{cvt|300|–|500|g}} at birth and begin to see after nine to 12 days. The milk canines erupt after one month. Pups first leave the den after three weeks. At one-and-a-half 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 three to four weeks. They have a fast growth rate during their first four months of life: during this period, a pup's weight can increase nearly 30 times.{{sfn|Heptner|Naumov|1998|pp=249–254}}{{sfn|Mech|Boitani|2003|pp=46–49}} Wolf pups begin play-fighting at the age of three weeks, though unlike young coyotes and foxes, their bites are gentle and controlled. Actual fights to establish hierarchy usually occur at five to eight weeks of age. This is in contrast to young coyotes and foxes, which may begin fighting even before the onset of play behaviour.<ref name=Fox1978/> By autumn, the pups are mature enough to accompany the adults on hunts for large prey.{{sfn|Mech|Boitani|2003|pp=42–46}}

===Hunting and feeding===
{{main article|Hunting behavior of gray wolves}}
[[File:Wolves and elk (cropped).jpg|thumb|right|alt=Aerial photograph a bull elk in winter being pursued by four wolves |Wolves pursuing a bull elk]]
Single wolves or mated pairs typically have higher success rates in hunting than do large packs; single wolves have occasionally been observed to kill large prey such as moose, [[bison]] and [[muskox]]en unaided.{{sfn|Mech|Boitani|2003|pp=119–121}}<ref name=Thurber1993/> The size of a wolf hunting pack is related to the number of pups that survived the previous winter, adult survival, and the rate of dispersing wolves leaving the pack. The optimal pack size for hunting elk is four wolves, and for bison a large pack size is more successful.{{sfn|Mech|Smith|MacNulty|2015|p=4}} Wolves move around their territory when hunting, using the same trails for extended periods.{{sfn|Heptner|Naumov|1998|p=233}} Wolves are nocturnal predators. During the winter, a pack will commence hunting in the twilight of early evening and will hunt all night, traveling tens of kilometres. Sometimes hunting large prey occurs during the day. During the summer, wolves generally tend to hunt individually, ambushing their prey and rarely giving pursuit.{{sfn|Heptner|Naumov|1998|p=240}} 

When hunting large gregarious prey, wolves will try to isolate an individual from its group.<ref name=MacNulty2007/> If successful, a wolf pack can bring down game that will feed it for days, but one error in judgement can lead to serious injury or death. Most large prey have developed defensive adaptations and behaviours. Wolves have been killed while attempting to bring down bison, elk, moose, muskoxen, and even by one of their smallest hoofed prey, the white-tailed deer. With smaller prey like [[beaver]], geese, and hares, there is no risk to the wolf.{{sfn|Mech|Smith|MacNulty|2015|pp=1–3}} Although people often believe wolves can easily overcome any of their prey, their success rate in hunting hoofed prey is usually low.{{sfn|Mech|Smith|MacNulty|2015|p=7}} 
[[File:Wolves eating deer.jpeg|thumb|right|alt=Photograph of two wolves eating a deer carcass at night|Two wolves feeding on a [[white-tailed deer]]]]
The wolf must give chase and gain on its fleeing prey, slow it down by biting through thick hair and hide, and then disable it enough to begin feeding.{{sfn|Mech|Smith|MacNulty|2015|pp=1–3}} Wolves may wound large prey and then lie around resting for hours before killing it when it is weaker due to blood loss, thereby lessening the risk of injury to themselves.{{sfn|Mech|Smith|MacNulty|2015|p=82–89}} With medium-sized prey, such as roe deer or [[sheep]], wolves kill by [[Throat clamp|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.<ref name=Zimen/>

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.{{sfn|Mech|1981|p=185}} The breeding pair typically monopolizes food to continue producing pups. When food is scarce, this is done at the expense of other family members, especially non-pups.{{sfn|Mech|Boitani|2003|p=58}} Wolves typically commence feeding by gorging on the larger internal organs, like the [[heart]], [[liver]], [[lung]]s, and [[stomach]] lining. The [[kidney]]s and [[spleen]] are eaten once they are exposed, followed by the muscles.{{sfn|Mech|Boitani|2003|pp=122–125}} A wolf can eat 15–19% of its body weight in one sitting.{{sfn|Mech|Boitani|2003|p=201}}