Aaaah, you skim water off a ocean.
The gray wolf or grey wolf (Canis lupus), often known simply as the wolf, is the largest wild member of the Canidae family. It is an ice age survivor originating during the Late Pleistocene around 300,000 years ago.[3] DNA sequencing and genetic drift studies reaffirm that the gray wolf shares a common ancestry with the domestic dog (Canis lupus familiaris). Although certain aspects of this conclusion have been questioned, including recently,[4] the main body of evidence confirms it. A number of other gray wolf subspecies have been identified, though the actual number of subspecies is still open to discussion. Gray wolves are typically apex predators in the ecosystems they occupy. Though not as adaptable to human presence as more generalist canid species,[4] wolves have thrived in temperate forests, deserts, mountains, tundra, taiga, grasslands, and even urban areas.[5]
Though once abundant over much of Eurasia and North America, the gray wolf inhabits a very small portion of its former range because of widespread destruction of its territory, human encroachment, and the resulting human-wolf encounters that sparked broad extirpation. Even so, the gray wolf is regarded as being of least concern for extinction according to the International Union for Conservation of Nature, when the entire gray wolf population is considered as a whole. Today, wolves are protected in some areas, hunted for sport in others, or may be subject to extermination as perceived threats to livestock and pets.
Physiology
Physical characteristics
Gray wolf weight and size can vary greatly worldwide, tending to increase proportionally with latitude as predicted by Bergmann's Rule. In general, height varies from 0.6 to 0.95 meters (24 to 37 in) at the shoulder. Wolf weight varies geographically; on average, European wolves may weigh 38.5 kilograms (85 lb), North American wolves 36 kilograms (79 lb), and Indian and Arabian wolves 25 kilograms (55 lb).[6] Though rarely encountered, extreme specimens of more than 77 kilograms (170 lb) have been recorded in Alaska, Canada,[7] and the former Soviet Union.[8] The heaviest recorded gray wolf in the New World was killed on 70 Mile River in east-central Alaska on July 12, 1939 and weighed 79 kilograms (170 lb),[6] while the heaviest recorded wolf in the Old World was killed after World War II in the Kobelyakski Area of the Poltavskij Region in the Ukrainian SSR, and weighed 86 kilograms (190 lb).[9] Grey wolves are sexually dimorphic, with females in any given wolf population typically weighing 20% less than males.[10] Females also have narrower muzzles and foreheads; slightly shorter, smoother furred legs; and less massive shoulders.[6] Gray wolves can measure anywhere from 1.3 to 2 meters (4.3 to 6.6 ft) from nose to the tip of the tail, which itself accounts for approximately one quarter of overall body length.[11]
Gray wolves rely on their stamina rather than speed for hunting. Their narrow chests and powerful backs and legs facilitate efficient locomotion. They are capable of covering several miles trotting at about 10 kilometers per hour (6 mph), and have been known to reach speeds approaching 65 kilometers per hour (40 mph) during a chase.[12] One female gray wolf was recorded to have made 7-meter (23 ft) bounds when chasing prey.[9]
Gray wolf paws are able to tread easily on a wide variety of terrains, especially snow. There is a slight webbing between each toe, which allows them to move over snow more easily than comparatively hampered prey. Gray wolves are digitigrade, which, with the relative largeness of their feet, helps them to distribute their weight well on snowy surfaces. The front paws are larger than the hind paws, and have a fifth digit, the dewclaw, that is absent on hind paws.[13] Bristled hairs and blunt claws enhance grip on slippery surfaces, and special blood vessels keep paw pads from freezing.[14] Scent glands located between a wolf's toes leave trace chemical markers behind, helping the wolf to effectively navigate over large expanses while concurrently keeping others informed of its whereabouts.[14] Unlike dogs and western coyotes, gray wolves have a lower density of sweat glands on their paws. This trait is also present in Eastern Canadian Coyotes which have been shown to have recent wolf ancestry.[15] Wolves in Israel are unique due to the middle two toes of their paws being fused, a trait originally thought to be unique to the African Wild Dog.[16]
Genetic research has shown that black furred wolves owe their colouration to a mutation that first arose in domestic dogs
Gray wolves molt some of their coats in late spring or early summer.
Wolves have bulky coats consisting of two layers. The first layer is made up of tough guard hairs that repel water and dirt. The second is a dense, water-resistant undercoat that insulates. The undercoat is shed in the form of large tufts of fur in late spring or early summer (with yearly variations). A wolf will often rub against objects such as rocks and branches to encourage the loose fur to fall out. The undercoat is usually gray regardless of the outer coat's appearance. Wolves have distinct winter and summer pelages that alternate in spring and autumn. Females tend to keep their winter coats further into the spring than males.
At birth, wolf pups tend to have darker fur and blue irises that will change to a yellow-gold or orange color when the pups are between 8 and 16 weeks old.[19] Wolves' long, powerful muzzles help distinguish them from other canids, particularly coyotes and Golden Jackals, which have more narrow, pointed muzzles. In wolves, the anterior incisure of the nasal bones does not have a medial protrusion, unlike jackals. The cingulum on the external edge of the first upper molar is only slightly expressed, while it is broad and distinctly marked in jackals.[20]
Wolves and most larger dogs share identical dentition. The maxilla has six incisors, two canines, eight premolars, and four molars. The mandible has six incisors, two canines, eight premolars, and six molars.[22] The fourth upper premolars and first lower molars constitute the carnassial teeth, which are essential tools for shearing flesh. The long canine teeth are also important, in that they hold and subdue the prey. Capable of delivering up to 10,000 kilopascals (1,500 psi) of pressure, a wolf's teeth are its main weapons as well as its primary tools.[6] This is roughly twice the pressure that a domestic dog of similar size can deliver.[23] The dentition of grey wolves is better suited to bone crushing than those of other modern canids, though it is not as specialised as that found in hyenas.[24]
Wolf saliva has been shown to reduce bacterial infection in wounds and accelerate tissue regeneration.[25]
The pups, which weigh about 0.5-kilogram (1 lb) at birth, are born blind, deaf, and completely dependent on their mother.[26][31] The average litter size is 5–6 pups, though there are two Soviet records of litters consisting of 17 pups.[9] The pups reside in the den and stay there for two months. The den is usually on high ground near an open water source, and has an open chamber at the end of an underground or hillside tunnel that can be up to a few meters long.[14] During this time, the pups will become more independent, and will eventually begin to explore the area immediately outside the den before gradually roaming up to a mile away from it at around five weeks of age. Wolf growth rate is slower than that of coyotes and dholes.[32] They begin eating regurgitated foods after two weeks of feeding on milk, which in wolves has less fat and more protein and arginine than dog milk.[10] By this time, their milk teeth have emerged—and are fully weaned by 10 weeks. During the first weeks of development, the mother usually stays with her litter alone, but eventually most members of the pack will contribute to the rearing of the pups in some way.[26] After two months, the restless pups will be moved to a rendezvous site, where they can stay safely while most of the adults go out to hunt. One or two adults stay behind to ensure the safety of the pups. After a few more weeks, the pups are permitted to join the adults if they are able, and will receive priority on anything killed, their low ranks notwithstanding. Letting the pups fight for eating privileges results in a secondary ranking being formed among them, and allows them to practice the dominance/submission rituals that will be essential to their future survival in pack life.[26] During hunts, the pups remain ardent observers until they reach about eight months of age, by which time they are large enough to participate actively.
Diseases recorded to be carried by wolves include brucella, deerfly fever, leptospirosis, foot-and-mouth disease, and anthrax. Wolves are major hosts for rabies in Russia, Iran, Afghanistan, Iraq, and India.[35] Though wolves are not reservoirs for the disease, they can catch it from other species. Wolves develop an exceptionally severe aggressive state when rabid and can bite numerous people in a single attack. Before a vaccine was developed, bites were almost always fatal. Today, rabid wolf bites can be treated, but the severity of rabid wolf attacks can sometimes result in outright death, or a bite near the head will make the disease act too fast for the treatment to take effect. Rabid attacks tend to cluster in winter and spring. With the reduction of rabies in Europe and North America, few rabid wolf attacks have been recorded, though some still occur annually in the Middle East.[36] Wolves also carry the Canine coronavirus, infections being most prevalent in winter months.[37]
Wolves are territorial animals. Studies have shown that the average size of a wolf pack's territory is close to 200 km2 (80 sq mi).[53] 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.[54] Prey density tends to be much higher in the territory's surrounding areas. Despite this higher abundance of prey, wolves tend to avoid hunting in the fringes of their territory unless desperate, due to the possibility of fatal encounters with neighboring packs.[55] Established wolf packs rarely accept strangers into their territories, with one study on wolf mortality in Minnesota and the Denali National Park and Preserve concluding that 14–65% of wolf deaths were due to predation by other wolves.[56] In fact, 91% of wolf fatalities occur within 3.2 km (2.0 mi) of the borders between neighboring territories.[57] The majority of killed wolves are dominant animals, due to their greater assertiveness in confronting other packs.[58] In rare cases in which a stranger is accepted into the pack, the animal itself is almost invariably a young specimen of 1–3 years of age, while the majority of killed wolves are adults.[59]
Communication between these boundaries is achieved in part through scent marking and howling. Howling is the principal means of spacing in wolf populations. It communicates the location of a core territory as well as enforcing a territory-independent buffer zone around the roaming wolf pack. This territory-independent buffer zone is a means of avoiding encounters with neighboring packs near territory borders.[60] Lone wolves, in contrast, rarely respond to howls, instead taking an "under the radar" approach. Howling communicates a core territory over time, as a wolf packs spends much of their time there.
Dispersion
Wolves, like other canines, use scent marking to lay claim to anything—from territory to fresh kills.[62] Breeding wolves scent mark the most often, with males doing so more than females. The most widely used scent marker is urine. Male and female breeding wolves urine-mark objects with a raised-leg stance (all other pack members squat) to enforce rank and territory. They also use marks to identify food caches and to claim kills on behalf of the pack. Defecation markers are used for the same purpose as urine marks, and serve as a more visual warning, as well.[62] Defecation markers are particularly useful for navigation, keeping the pack from traversing the same terrain too often and also allowing each wolf to be aware of the whereabouts of its pack members. Above all, though, scent marking is used to inform other wolves and packs that a certain territory is occupied, and that they should therefore tread cautiously.
Wolves have scent glands all over their bodies, including at the base of the tail, between toes, and in the eyes, genitalia, and skin.[62] Pheromones secreted by these glands identify each individual wolf. A dominant wolf will "rub" its body against subordinate wolves to mark such wolves as being members of a particular pack. Wolves may also "paw" dirt to release pheromones instead of urine marking.[63]
Diet
Wolves feed primarily on medium to large sized ungulates, though they are opportunistic feeders, and will generally eat any meat that is available,[64] including non-ungulate species,[65] carrion and garbage.[64] Cannibalism is not uncommon in wolves, and has been recorded to occur in times of food scarcity,[66] when a pack member dies,[67] and during territorial disputes.[58] Some wolf packs in Alaska and Western Canada have been observed to feed on salmon.[68][69] Humans are rarely, but occasionally preyed upon (see Attacks on humans).[36][70][71][72] Wolves will typically avoid a potential prey item which does not conform to what they experienced during their lives. Generally, the greater the discrepancy to what wolves are accustomed to, the greater their resistance to exploring it. This is only increased should the new prey act bold, assertive, and fearless. Nevertheless, even if there is no food shortage, wolves will explore alternative prey if they continually come into close contact with it and habituate themselves.[73]
An American Bison standing its ground, thereby increasing its chance for survival.
A bull elk running, thereby decreasing its chance for survival.
Unlike lion prides, wolf packs numbering above 2 individuals show little strategic cooperation in hunting large prey.[64] Wolves typically attempt to conceal themselves as they approach their prey. Often, they will wait for the prey to graze, when it is distracted.[9] If the prey animal stands its ground or confronts the pack, the wolves will approach and threaten it. The wolves will eventually leave if their prey does not run, though the length of time can range from hours to days.[64] If their prey attempts to flee, the wolves will give chase. Wolves generally do not engage in long chases, and will usually stop a pursuit after a chase of 10–180 meters (10–200 yd), though there has been one documented case of a wolf chasing a moose for 36 kilometers (22 mi).[9] Female wolves tend to be better at chasing prey than males, while the latter are more adept at wrestling large prey to the ground once it is caught. Packs composed largely of female wolves thrive on fleet footed prey such as elk, while packs specialising in bison tend to have a greater number of males.[74] Though commonly portrayed as targeting solely sick or infirm animals,[33] there is little evidence that they actively limit themselves to such targets. Rather, the evidence shows that wolves will simply target the easiest options available, which as well as sick and infirm animals, can also include young animals and pregnant females.[75] Though wolves commonly hunt large prey in packs, there are cases in which single wolves have successfully killed large animals unaided. One wolf was recorded to have killed moose 11 times singlehandedly.[76]
Wolves will typically attempt to disable large prey by tearing at the haunches and perineum, causing massive bleeding and loss of coordination. A single bite can cause a wound up to 10–15 centimeters (4–6 in) in length. A large deer in optimum health generally succumbs to three bites at the perineum area after a chase of 150 meters (160 yd). Once their prey is sufficiently weakened, the wolves will grab it by the flanks and pull it down.[9] Sometimes, with medium sized prey such as dall sheep, wolves will bite the throat, severing the windpipe or jugular.[77] When attacking canid prey, such as dogs, coyotes or other wolves, wolves will kill by biting the back, neck or head.[58][78][79] With prey of equal or lesser weight to the wolf, such as lambs or small children, wolves will grab their quarry by the neck, chest, head or thigh and carry them off to a secluded spot.[9][72] Once the prey collapses, the wolves will tear open the abdominal cavity and commence feeding on the animal, sometimes before it has died.[9] On some occasions, wolves will not press an attack, and will wait for their prey to die from their wounds before feeding begins.[80] Wolves will occasionally attack pregnant ungulates to feed on the fetus(es), leaving the mother uneaten.[81] Usually, it is the dominant pair that works the hardest in killing the pack's target.[64] Wolves have on occasion been observed to engage in acts of surplus killing. This phenomenon is common when wolves target livestock.[82] In the wild, this usually occurs in late winter or spring when deep snow impedes their prey's escape.[82][83]
Pack status is reinforced during feeding. The breeding pair usually eats first, starting with the heart, liver, and lungs. Wolves of intermediate rank will prevent lower ranking pack members from feeding until the dominant pair finishes eating.[25] The stomach of prey is eaten, though the contents are left untouched if the killed animal is a herbivore. The leg muscles are eaten next, with the hide and bones being the last to be consumed.[64] If they are disturbed while feeding, they will instead focus their attention on their prey's fat deposits rather than internal organs.[84][85] A single wolf can eat up to 3.2–3.5 kilograms (7–8 lb) of food at a time, though they can eat as much as 13–15 kilograms (29–33 lb) when sufficiently hungry. A wolf's yearly requirement is 1,500 kilograms (3,307 lb).[9] Wolves can go without sustenance for long periods, with a Russian record showing how one specimen survived for 17 days without food.[6] Research has shown that 2 weeks without food will not weaken a wolf's muscle activity.[9] After eating, wolves will drink large quantities of water to prevent uremic poisoning.[6] A wolf's stomach can hold up to 7.5 liters (8 U.S. qt).[9] Wolves supplement their diet with vegetation. Scat analysis found 75% of samples found Yellowstone National Park wolves’ summer diet contained plants mostly grass (Graminae).[86] In some areas of the former Soviet Union wolves have been reported to cause serious damage to watermelon plantations.[75]
Studies on how wolves affect prey populations tend to vary considerably, with some results indicating that wolves dramatically reduce, sometimes locally extirpate some prey species, while others indicate that wolf predation simply takes over from other mortality factors present in wolf-free zones.[66][83] Wolves are not essential for the presence of many other species.[87]
Interspecific predatory relationships
Rolf Peterson investigating a coyote carcass killed by a wolf in Yellowstone National Park, January 1996
Wolves typically dominate other canid species in areas where they are sympatric. In North America, wolves are generally intolerant of coyotes in their territory; two years after their reintroduction to the Yellowstone National Park, the wolves were responsible for a near 50% drop in coyote populations through both competition and predation.[88] Wolves have been reported to dig coyote pups from their dens and kill them. Wolves typically do not consume the coyotes they kill. There are no records of coyotes killing wolves,[79] though they have been known to gang up on wolves if they outnumber them.[88] Wolves have been observed to allow coyotes to approach their kills, only to chase them down and kill them. Coyote specialist Robert Crabtree of the Yellowstone Ecological Research Center suggested that this behaviour could be linked to the intraspecific territoriality of wolves, even though coyote represent no danger: "Maybe you want to teach your pups tricks of the trade... Maybe wolves are killing coyotes to practice for conflicts with other wolves later in life."[89] Near identical interactions have been observed in Greece between wolves and Golden Jackals.[90] Wolves may kill foxes on kill sites, though not as frequently as they do with coyotes. Raccoon Dogs are also reportedly preyed upon.[79]
Wolf following a brown bear
Brown Bears are encountered in both Eurasia and North America. The majority of interactions between wolves and Brown Bears usually amount to nothing more than mutual avoidance. Serious confrontations depend on the circumstances of the interaction, though the most common factor is defence of food and young. Brown Bears will use their superior size to intimidate wolves from their kills and, when sufficiently hungry, will raid wolf dens. Brown Bears usually dominate wolves on kills, though they rarely prevail against wolves defending den sites. Wolves in turn have been observed killing bear cubs, to the extent of even driving off the defending mother bears. Deaths in wolf/bear skirmishes are considered very rare occurrences, the individual power of the brown bear and the collective strength of the wolf pack usually being sufficient deterrents to both sides.[79] Encounters with American Black Bears occur solely in the Americas; their interactions with wolves are much rarer than those of Brown Bears, due to differences in habitat preferences. 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 to kill Black Bears on numerous occasions without eating them. Unlike Brown Bears, Black Bears frequently lose against wolves in disputes over kills.[79] While encounters with brown and black bears appear to be common, polar bears are rarely encountered by wolves, though there are two records of wolf packs killing polar bear cubs.[91]
Large wolf populations limit the numbers of small to medium sized felines. Wolf predation is recorded to reduce lynx populations wherever the two species are sympatric. Lynx populations in Slovakia plummeted during World War II, when large numbers of wolves entered the cat's range. Similarly, in Russia, lynx populations drop in areas with high wolf densities.[92] In the Rocky Mountains and adjacent mountain areas of North America, wolves are usually hostile toward cougars and will kill cubs if given the opportunity. A pack will on occasion appropriate the kills of adult cougars, which respond by increasing their kill rate. Both species have been recorded to kill each other.[79] National Park Service cougar specialist Kerry Murphy stated that the cougar usually is at an advantage on a one to one basis, considering it can effectively use its claws, as well as its teeth, unlike the wolf which relies solely on its teeth.[89] Yellowstone officials have reported that attacks between cougars and wolves are not uncommon. Multiple incidents of cougars taking wolves and vice versa have been recorded in Yellowstone National Park. Researchers in Montana have found that wolves regularly kill cougars in the area.[93] Similarly, large numbers of wolves have been reported to reduce leopard populations in Tibet.[94] However, the reverse is true for larger cats such as tigers. In areas where wolves and tigers share ranges, such as the Russian Far East, the two species typically display a great deal of dietary overlap, resulting in intense competition. Wolf and tiger interactions are well documented in Sikhote-Alin, which until the beginning of the 20th century, held very few wolves. It is thought by certain experts that wolf numbers increased in the region after tigers were largely eliminated during the Russian colonization in the late 1800s and early 1900s. This is corroborated by native inhabitants of the region claiming that they had no memory of wolves inhabiting Sikohte-Alin until the 1930s, when tiger numbers decreased.[95] 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. Wolves appear capable of escaping competitive exclusion from tigers only when human persecution decreases the latter's numbers.[96] Today wolves are considered scarce in tiger inhabited areas, being found in scattered pockets, and usually seen traveling as loners or in small groups. First hand accounts on interactions between the two species indicate that tigers occasionally chase wolves from their kills, while wolves will scavenge from tiger kills. Tigers are not known to prey on wolves, though there are four records of tigers killing wolves without consuming them.[95] This competitive exclusion of wolves by tigers has been used by Russian conservationists to convince hunters in the Far East to tolerate the big cats, as they limit ungulate populations less than wolves, and are effective in controlling the latter's numbers.[97]
Wolves may occasionally encounter Striped Hyenas in the Middle East and Central and South Asia, mostly in disputes over carcasses. Though hyenas usually dominate wolves on a one to one basis, wolf packs have been reported to displace lone hyenas from carcasses.[98] Wolf remains have been found in Cave Hyena den sites, though it is unknown if the wolves were killed or scavenged upon.[99] Unlike cave hyenas, which preferentially preyed on lowland animals such as horses, wolves relied more on slope-dwelling ibex and Roe Deer, thus minimising competition. Wolves and Cave Hyenas seem to display negative abundance relations over time, with wolf populations expanding their ranges as hyenas disappeared.[100]
Communication
Body language
See also: Dog communication
The posture and facial expression of this Arabian wolf is defensive and gives warning to other wolves to be cautious.
This facial expression shows fear.
Wolves can communicate visually through a variety of expressions and moods ranging from subtle signals, such as a slight shift in weight, to more obvious ones, such as rolling on their backs to indicate complete submission.[101]
* Dominance – A dominant wolf stands stiff legged and tall. The ears are erect and forward, and the hackles bristle slightly. Often the tail is held vertically and curled toward the back. This display asserts the wolf's rank to others in the pack. A dominant wolf may stare at a submissive one, pin it to the ground, "ride up" on its shoulders, or even stand on its hind legs.
* Submission (active) – During active submission, the entire body is lowered, and the lips and ears are drawn back. Sometimes active submission is accompanied by muzzle licking, or the rapid thrusting out of the tongue and lowering of the hindquarters. The tail is placed down, or halfway or fully between the legs, and the muzzle often points up to the more dominant animal. The back may be partly arched as the submissive wolf humbles itself to its superior; a more arched back and more tucked tail indicate a greater level of submission.
* Submission (passive) – Passive submission is more intense than active submission. The wolf rolls on its back and exposes its vulnerable throat and underside. The paws are drawn into the body. This posture is often accompanied by whimpering.
* Anger – An angry wolf's ears are erect, and its fur bristles. The lips may curl up or pull back, and the incisors are displayed. The wolf may also arch its back, lash out, or snarl.
* Fear – A frightened wolf attempts to make itself look small and less conspicuous; the ears flatten against the head, and the tail may be tucked between the legs, as with a submissive wolf. There may also be whimpering or barks of fear, and the wolf may arch its back.
* Defensive – A defensive wolf flattens its ears against its head.
* Aggression – An aggressive wolf snarls and its fur bristles. The wolf may crouch, ready to attack if necessary.
* Suspicion – Pulling back of the ears shows a wolf is suspicious. The wolf also narrows its eyes. The tail of a wolf that senses danger points straight out, parallel to the ground.
* Relaxation – A relaxed wolf's tail points straight down, and the wolf may rest sphinx-like or on its side. The wolf may also wag its tail. The further down the tail droops, the more relaxed the wolf is.
* Tension – An aroused wolf's tail points straight out, and the wolf may crouch as if ready to spring.
* Happiness – As dogs do, a wolf may wag its tail if in a joyful mood. The tongue may roll out of the mouth.
* Hunting – A wolf that is hunting is tensed, and therefore the tail is horizontal and straight.
* Playfulness – A playful wolf holds its tail high and wags it. The wolf may frolic and dance around, or bow by placing the front of its body down to the ground, while holding the rear high, sometimes wagged. This resembles the playful behavior of domestic dogs.
Howling and other vocalisations
Howling adult wolf at the UK Wolf Conservation Trust.
Wolf howls, which can last from 0.5–11 seconds, typically have a frequency of 150-780 Hz.[17] Howling helps pack members keep in touch, allowing them to communicate effectively in thickly forested areas or over great distances. Howling also helps to call pack members to a specific location. Howling can also serve as a declaration of territory, as shown in a dominant wolf's tendency to respond to a human imitation of a "rival" wolf in an area the wolf considers its own. This behavior is stimulated when a pack has something to protect, such as a fresh kill. As a rule of thumb, large packs will more readily draw attention to themselves than will smaller packs. Adjacent packs may respond to each others' howls, which can mean trouble for the smaller of the two. Wolves therefore tend to howl with great care.[102] Wolves will also howl for communal reasons. Some scientists speculate that such group sessions strengthen the wolves' social bonds and camaraderie—similar to community singing among humans.[102] During such choral sessions, wolves will howl at different tones and varying pitches, making it difficult to estimate the number of wolves involved. This confusion of numbers makes a listening rival pack wary of what action to take. For example, confrontation could be disastrous if the rival pack gravely underestimates the howling pack's numbers. A wolf's howl may be heard from up to 16 kilometers (10 mi) away, depending on weather conditions. Observations of wolf packs suggest that howling occurs most often during the twilight hours, preceding the adults' departure to the hunt and following their return. Studies also show that wolves howl more frequently during the breeding season and subsequent rearing process.[103] The pups themselves begin howling soon after emerging from their dens and can be provoked into howling sessions easily over the following two months. Such indiscriminate howling usually is intended for communication, and does not harm the wolf so early in its life.[102] Howling becomes less indiscriminate as wolves learn to distinguish howling pack members from rival wolves. Wolves from the Middle East and Southern Asia are unusual as they are not known to howl, and have more doglike vocalisations comprised of short, sharp barks.[6][51]
Growling, while teeth are bared, is the most visual warning wolves use. Wolf growls have a distinct, deep, bass-like quality which can range from 250 to 1,500 Hz.[17] It is often used to threaten rivals, though not necessarily to defend themselves. Wolves also growl at other wolves while being aggressively dominant. Wolves bark when nervous or when alerting other wolves of danger but do so very discreetly and will not generally bark loudly or repeatedly as dogs do. Instead they use a low-key, breathy "whuf" sound which can measure from 320–904 Hz[17] to immediately get attention from other wolves. Wolves also "bark-howl" by adding a brief howl to the end of a bark. Wolves bark-howl for the same reasons they normally bark. Generally, pups bark and bark-howl much more frequently than adults, using these vocalizations to cry for attention, care, or food. A lesser known sound is the rally. Wolves will gather as a group and, amidst much tail-wagging and muzzle licking, emit a high-pitched wailing noise interspersed with something similar to (but not the same as) a bark. Rallying is often a display of submission to an alpha by the other wolves.[104] Wolves also whimper, a sound with a maximum range of 3,500 Hz,[17] usually when submitting to other wolves. Wolf pups whimper when they need a reassurance of security from their parents or other wolves.
Taxonomy
The gray wolf is a member of the genus Canis, which comprises between 7 and 10 species. It is one of six species termed 'wolf', the others being the Red Wolf (Canis rufus), the Indian Wolf (Canis indica), the Himalayan Wolf (Canis himalayaensis), the Eastern Wolf (Canis lycaon), and the Ethiopian Wolf (Canis simensis), although there is still some uncertainty as to whether some of these should be considered subspecies of Canis lupus or species in their own right. Recent genetic research suggests that Indian Wolf populations in the Indian subcontinent may represent a distinct species. Similar results were obtained for the Himalayan Wolf, which is traditionally placed into the Tibetan Wolf (Canis lupus laniger).[105]
With respect to common names, spelling differences result in the alternative spelling grey wolf. As the first-named and most widespread of species termed "wolf", gray wolves are often simply referred to as wolves. It was one of the many species originally described by Carl Linnaeus in his eighteenth-century work, Systema Naturae, and it still bears its original classification, Canis lupus.[106] The binomial name is derived from the Latin Canis, meaning "dog", and lupus, "wolf".[107]
Timeline of canids including Canis lupus in red. (Tedford & Xiaoming Wang)
Desert dwelling grey wolf subspecies, such as this Arabian wolf, tend to be smaller than their more northern cousins.
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