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Gray Wolf
Conservation status
Template:StatusConcern
Scientific classification
Kingdom:	Animalia
Phylum:	Chordata
Class:	Mammalia
Order:	Carnivora
Family:	Canidae
Genus:	Canis
Species:	C. lupus
Binomial name
Canis lupus Linnaeus, 1758

The Gray Wolf (Canis lupus; also spelled Grey Wolf, see spelling differences; other forms: Timber Wolf, Wolf). The Gray Wolf shares a common ancestry with the domestic dog (Canis lupus familiaris), and is known from DNA sequencing and genetic drift studies to be the progenitor of all dogs as they exist today. Gray wolves were once abundant and distributed over much of North America, Eurasia, and the Middle East. Today, for a variety of human-related reasons including widespread habitat destruction and excessive hunting, wolves inhabit only a very limited portion of their former range.

Gray wolves, being keystone predators, are integral components of the ecosystems to which they typically belong. The wide range of habitats in which wolves can thrive reflects their adaptability as a species, and includes temperate forests, mountains, tundra, taiga, and grasslands. In the contiguous U.S., with the exception of Minnesota and Wisconsin (where they have a threatened status), they are listed as endangered under the Endangered Species Act of 1973. They continue to be hunted in many areas of the world as perceived threats to livestock and human well-being, as well as for sport.

Carolus Linnaeus gave the wolf the scientific name Canis lupus in the 18th century.

Anatomy, physiology, and reproduction

Features and adaptations

Wolf weight and size can vary greatly worldwide, though both tend to increase proportionally with higher latitudes. Generally speaking, height varies from 0.6 to 0.9 meters (24 to 35 inches) at the shoulder, and weight can range anywhere from 25 to 65 kg (55-143 pounds), making wolves the largest among all wild canids. Although rarely encountered, extreme specimens reaching 80 kg (176 lb.) have been recorded in Alaska and Canada, though some people claim to have seen even larger anomalous individuals (90+ kg) roaming the Yukon, where some of the largest wolves in North America can be found. Customarily, however, wolves will be of a more typical physical capacity, with the females in a given population weighing about twenty percent less than their male counterparts. Wolves can measure anywhere between 1.3 and 2 meters (51 to 78 inches) from nose to tail tip, with the tail itself consisting of approximately one quarter of overall body length.

Wolves are built for stamina, possessing features tailored for long-distance travel. Narrow chests and powerful backs and legs contribute to the wolf's proficiency for efficient locomotion. They are capable of covering several miles trotting at about a 10 km/h (6 mph) pace, though they have been known to reach speeds approaching 65 km/h (40 mph) during a chase (wolves only run fast when testing potential prey). While sprinting thus, wolves can cover up to 5 m (16 ft) per bound.

Wolf paws are designed to traverse easily through a wide variety of terrains, especially snow. There is a slight webbing between each toe, which allows wolves to move over snow more easily than comparatively hampered prey. Wolves are digitigrade, so the relative largeness of their feet helps to better distribute their weight on snowy surfaces. The front paws are larger than the hind paws, and feature a fifth digit, a dewclaw, that is absent on hind paws. Bristled hairs and blunt claws enhance grip on slippery surfaces, and special blood vessels keep paw pads from freezing. Furthermore, scent glands located between a wolf's toes leave trace chemical markers behind, thereby helping the wolf to effectively navigate over large expanses while concurrently keeping others informed of its whereabouts.

A wolf sometimes seems more massive than it actually is due to its bulky coat, which is made of two layers. The first layer consists of tough guard hairs designed to repel water and dirt. The second is a dense, water-resistant undercoat that insulates. 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.

Coloration varies greatly, and runs from gray to gray-brown, all the way through the canine spectrum of white, red, brown, and black. These colors tend to mix in many populations to form predominantly blended individuals, though it is certainly not uncommon for an individual or an entire population to be entirely one color (usually all black or all white). A multicolor coat characteristically lacks any clear pattern other than it tends to be lighter on the animal's underside. Fur color sometimes corresponds with a given wolf population's environment; for example, all-white wolves are much more common in areas with perennial snow cover. Aging wolves acquire a grayish tint in their coats.

At birth, wolf pups tend to have darker fur and blue eyes that will change to a yellow-gold or orange color when the pups are 8-16 weeks old. Though extremely unusual, it is possible for an adult wolf to retain its blue-colored eyes.

Wolves have stout, blocky muzzles that help distinguish them from coyotes and dogs. Wolves also differ in certain skull dimensions, having a smaller orbital angle, for example, than dogs (>53 degrees for dogs compared to <45 degrees for wolves) while possessing a comparatively larger brain capacity. Larger paw size, yellow eyes, longer legs, and bigger teeth further distinguish adult wolves from other canids, particularly dogs. Also, precaudal glands at the base of the tail are present in wolves, whereas they are not in dogs.

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. 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 are designed to hold and subdue the prey. Powered by 1500 lb/sq. inch of pressure, a wolf's teeth are its main weapons as well as its primary tools. Therefore, any injury to the jaw line or teeth could devastate an individual, dooming it to starvation or incompetence.

Courtship and mating

Usually, the desire to pass on genetic material drives young wolves away from their birth packs, leading them to seek out mates and territories of their own. Dispersals occur at all times during the year, typically involving wolves who reached sexual maturity during the previous breeding season. It takes two such dispersals from two different packs for the process to take place (though it can occur, dispersing wolves from the same pack rarely mate). Once two dispersing wolves meet and begin traveling together, they immediately begin the process of seeking out territory, preferentially doing so in time for the next mating season. The bond that forms between such wolves lasts for the shorter of the two lifetimes, with few exceptions.

During the mating season, breeding wolves become extremely affectionate with one another in anticipation for the female's ovulation cycle. Overall, pack tension rises, as each mature wolf begins to feel the urge to mate. In fact, during this time, the alpha male and alpha female may be forced to aggressively prevent other wolves from mating with each other. Under normal circumstances, a pack can only support one litter per year, so this type of dominance behavior is beneficial in the long run.

When the alpha female goes into estrus – a phenomenon that occurs once per year, and lasts between five and fourteen days – she and her mate will spend an increased amount of time in seclusion. Pheromones in the female's urine and the swelling of her vulva let the male know when his mate is in heat. She will be unreceptive for the first few days of estrus, during which time she sheds the lining of her uterus. Once the female begins to ovulate, mating occurs.

The male wolf will mount the female firmly from behind. After achieving coitus, the two form a copulatory tie once the bulb (bulbus glandis; located near the base of the canine penis) of the male's penis swells and the female's vaginal muscles tighten. Ejaculation is induced by the thrusting of the male's pelvis and the undulation of the female's cervix. The two become physically inseparable for anywhere between ten and thirty minutes, during which period the male will ejaculate multiple times. The male will occasionally lift one of his legs over the female such that they are standing end-to-end. This is believed to limit the leakage of semen; it may also be a defensive measure.

The mating ordeal is repeated many times throughout the female's brief ovulation period, which occurs once per year per female (unlike female dogs, with whom estrus usually occurs twice per year). It is believed that both males and females can continue to breed in this manner until at least ten years of age.

Breeding and life cycle

Normally, only the alpha pair of the pack breeds, which is a kind of organization not uncommon to other pack-hunting canids including the Dhole and the African Hunting Dog. Mating occurs between January and April, happening later in the year as latitude increases. A pack usually produces a single litter, though sometimes multiple litters will be born if the alpha male mates with one or more subordinate females. Under normal circumstances, the alpha female will try to prevent this by aggressively dominating other females and physically separating them from the alpha male during the mating season.

The gestation period lasts 60 to 63 days, and the pups are born blind, deaf, and completely dependent on their mother (i.e. altricial). There are 1–14 pups per litter, with the average litter size being about four to six. Pups reside in the den, where they are born, and stay there until they reach about 8 weeks of age (the den is usually on high ground near an open water source, and has an open "room" at the end of an underground/hillside tunnel that can be up to a few meters long). 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. They begin eating regurgitated foods at four weeks – by which time their milk teeth have emerged – and are weaned by six 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.

After two months, the restless pups will be moved to a rendezvous site, which gives them a safe place to reside while most of the adults go out to hunt. An adult or two will 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 (they tag along as observers until about eight months, by which time they are large enough to actively participate), and will receive first priority on anything killed, their low ranks notwithstanding. Letting the pups fight for the right to eat results in a secondary ranking being formed among them, and lets them practice the dominance/submission rituals that will be essential to their future survival in pack life.

Wolves typically reach sexual maturity after two or three years, at which point many of them will feel compelled to leave their birth packs and search out mates and territories of their own. Wolves that reach maturity generally live between 6 and 9 years in the wild, although in captivity they can live to be twice that age. High mortality rates result in a relatively low life expectancy for wolves on an overall basis. Pups die when food is scarce; they can also fall prey to other predators such as bears, or, less likely, coyotes, foxes, or other wolves. The most significant mortality factors for grown wolves are hunting and poaching by humans, car accidents, and wounds suffered while hunting prey. Wolves are susceptible to the same infections that affect domestic dogs, such as mange, heartworm, rabies and canine distemper, and such diseases can become epidemic, drastically reducing the wolf population in an area.

Behavior

Body language

Wolves can visually communicate an impressive variety of expressions and moods that range from subtler signals – such as a slight shift in weight – to the more obvious ones – like rolling on the back as a sign of complete submission.

• 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 vertical and curled toward the back. This display shows the wolf's rank to all others in the pack. A dominant lupine may stare penetratingly at a submissive one, pin it to the ground, "ride up" on its shoulders, or even stand on its hind legs.
• Submission (active) – In 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 partially 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 is often accompanied by whimpering.
• Anger – An angry lupine'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 snarl.
• Fear – A frightened wolf tries to make its body look small and therefore less conspicuous. The ears flatten down 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 lupine is suspicious. In addition, the wolf narrows its eyes. The tail of a wolf that senses danger points straight out, parallel to the ground.
• Relaxedness – A relaxed wolf's tail points straight down, and the wolf may rest sphinxlike or on its side. The wolf's tail may also wag. 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 lupine may wag its tail if it is in a joyful mood. The tongue may loll out of the mouth.
• Hunting – A wolf that is hunting is tensed, and therefore the tail is horizontal and straight.
• Playfulness – A playful lupine 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 is reminiscent of the playful behavior executed in domestic dogs.

Howling

Wolves howl for several reasons. Howling helps pack members keep in touch, allowing them to effectively communicate in thickly forested areas or over great distances. Furthermore, howling helps to summon pack members to a specific location. Howling can also serve as a declaration of territory, as portrayed by a dominant wolf's tendency to respond to a human imitation of a "rival" individual in an area that the wolf considers its own. This behavior is also 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. Thus, wolves tend to howl with great care.

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. During such choral sessions, wolves will howl at different tones and varying pitches, which tends to prevent a listener from accurately estimating the number of wolves involved. This concealment of numbers makes a listening rival pack wary of what action to take. For example, confrontation could mean bad news if the rival pack gravely underestimates the howling pack's numbers.

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. The pups themselves begin howling towards the end of July, and can be provoked into howling sessions relatively easily over the following two months. Such indiscriminate howling usually has a communicative intent, and has no adverse consequences so early in a wolf's life. Howling becomes less indiscriminate as wolves learn to distinguish howling pack members from rival wolves.

Other vocalizations

Growling, used in tandem with bared teeth, is the most visual and effective warning wolves use. Wolf growls have a distinct, deep, bass-like quality, and are used much of the time as a threat, though they are not always necessarily used for defense. Wolves will also growl at other wolves while being aggressively dominant.

Wolves can also bark, which they do when nervous or to warn other wolves of danger. Wolves bark very discreetly, and will not generally bark loudly or repeatedly as dogs do; rather, they use a low-key, breathy "whuf" sound to get attention immediately from other wolves. Wolves will also "bark-howl" by adding a brief howl to the end of a bark. Wolves bark-howl for the same reasons they normally bark. Actually, pups bark and bark-howl much more frequently than adults, using such vocalizations as cries for attention, care, or food.

Wolves can also whimper, which they usually do only while submitting to other wolves. Wolf pups will whimper when they need a reassurance of security from their parents or other wolves.

Scent marking

Wolves, like other canines, use scent marking to lay claim to anything from territory to fresh kills. Alpha wolves scent mark the most often, with males doing so more than females. The most widely used scent marker is urine. Male alpha wolves urine-mark objects using a raised-leg stance (all females squat) so as to enforce rank and territory. They will also use such marks to identify food caches and to claim kills on behalf of the whole pack. Defecation markers are used for the same purposes as urine marks, and serve as a more visual warning, as well. These types of scent markings are particularly useful for navigational purposes, keeping the pack from traversing the same terrain too often while also allowing each individual to be aware of the whereabouts of its pack members. Above all, though, scent marking is used to notify other wolves and packs that a given 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. Pheromones secreted by these glands identify each individual wolf. A dominant wolf will "rub" his or her body against subordinate wolves to mark such individuals as being members of a particular pack. Wolves may also "paw" dirt to release pheromones in lieu of urine marking.

That wolves rely so heavily on odoriferous signals testifies greatly to their olfactory capabilities. Wolves can pick up any scent, including marks, from great distances, and can distinguish among them just as well or better than humans can distinguish other humans visually.

Social structure and hunting

The pack

Wolves function as social predators and hunt in packs organized according to strict, rank-oriented social hierarchies. It was originally thought that this comparatively high level of social organization had more to do with hunting success, and while this still may be true to a certain extent, emerging theories suggest that the pack has less to do with hunting and more to do with reproductive success.

The pack is led by the two individuals that sit atop the social hierarchy — the alpha male and the alpha female. The alpha pair (of whom only one may be the "top" alpha) has the greatest amount of social freedom compared to the rest of the pack, but they are not "leaders" in the human sense of the term. The alphas do not give the other wolves orders; rather, they simply have the most freedom in choosing where to go, what to do, and when to do it. Possessing strong instincts for fellowship, the rest of the pack usually follows.

While most alpha pairs are monogamous with each other, there are exceptions. An alpha animal may preferentially mate with a lower-ranking animal, especially if the other alpha is closely related (a brother or sister, for example). The death of one alpha does not affect the status of the other alpha, who will quickly take another mate.

Usually, only the alpha pair is able to successfully rear a litter of pups (other wolves in a pack may breed, but will usually lack the resources required to raise the pups to maturity). All the wolves in the pack assist in raising wolf pups. Some mature individuals, usually females, may choose to stay in the original pack so as to reinforce it and help rear more pups. Most, males particularly, will disperse, however.

The size of the pack may change over time and is controlled by several factors, including habitat, personalities of individual wolves within a pack, and food supply. Packs can contain between two and 20 wolves, though an average pack consists of six or seven. New packs are formed when a wolf leaves its birth pack and claims a territory. Lone wolves searching for other individuals can travel very long distances seeking out suitable territories. Dispersing individuals must avoid the territories of other wolves because intruders on occupied territories are chased away or killed, a behavior that may explain wolf "predation" of dogs. Most dogs, except perhaps large, specially bred attack dogs, do not stand much of a chance against a pack of wolves protecting its territory from an unwanted intrusion.

Hierarchy

The hierarchy – led by the alpha male and female – affects all activity in the pack to some extent. In most larger packs, there are two separate hierarchies in addition to an overbearing one: the first consists of the males, led by the alpha male, and the other consists of the females, led by the alpha female. In this situation, the alpha male usually assumes the "top" alpha position, though alpha females have been known to take control over entire packs in some cases. The male and female hierarchies are interdependent, and are maintained constantly by aggressive and elaborate displays of dominance and submission.

After the alpha pair, there may also, especially in larger packs, be a beta wolf or wolves – a "second-in-command" to the alphas. Betas typically assume a more prominent role in assisting with the upbringing of the alpha pair's litter, often serving as surrogate mothers or fathers while the alpha pair is away. Beta wolves are the most likely to challenge their superiors for the role of the alpha, though some betas seem content with being second, and will sometimes even let lower ranking wolves leapfrog them for the position of alpha should circumstances necessitate such a happening (death of the alpha, etc.). More ambitious beta wolves, however, will only wait so long before challenging for the top spot; unless, of course, they choose to disperse and create their own pack instead.

Loss of rank can happen gradually or suddenly. An older wolf may simply choose to give way when a motivated challenger presents itself, yielding its position without bloodshed. On the other hand, the challenged individual may choose to fight back, with varying degrees of intensity. While the majority of wolf aggression is non-damaging and ritualized, a high-stakes fight can easily result in injury for either or both parties. The loser of such a confrontation is frequently chased away from the pack or, rarely, may be killed as other aggressive wolves contribute to the insurgency. This kind of dominance encounter is more common during the mating season.

Rank order within a pack is established and maintained through a series of ritualized fights and posturing best described as ritual bluffing. Wolves prefer psychological warfare to physical confrontations, meaning that high-ranking status is based more on personality or attitude than on size or physical strength. Rank, who holds it, and how it is enforced varies widely between packs and between individual animals. In large packs full of easygoing wolves, or in a group of juvenile wolves, rank order may shift almost constantly, or even be circular (e.g., animal A dominates animal B, who dominates animal C, who dominates animal A).

In a more typical pack, however, only one wolf will assume the role of the omega – the lowest-ranking member of a pack. These individuals absorb the greatest amount of aggression from the rest of the pack, and may be subjected to different forms of truculence at any given point – anything from constant dominance from other pack members to inimical, physical harassment. Though this arrangement may seem objectionable after cursory analysis, the nature of pack dynamics demands that one wolf be at the bottom of the ranking order, and such individuals are perhaps better suited for constant displays of active and passive submission than they are for living alone. For wolves, camaraderie – no matter what the form – is preferable to solitude, and, indeed, submissive wolves tend to choose low rank over potential starvation.

Cooperative hunting and diet

Packs of wolves cooperatively hunt any large herbivores in their range, whereas lone wolves are limited to consuming smaller animals due to their relative inability to catch anything larger. Pack hunting methods range from surprise attacks to long-lasting chases, though they strongly favor the latter. Through meticulous cooperation, a pack of wolves is able to pursue large prey for several hours before relenting, though the success rate of such chases is rather low. Solitary wolves depend on small animals, capturing them by pouncing and pinning them to the ground with their front paws – a common technique among canids such as foxes and coyotes. Wolves' diets include, but are not limited to, elk, caribou, moose, deer, and other large ungulates. They also prey on rodents and small animals in a limited manner, as a typical wolf requires between 1.3 to 4.5 kg (3 and 10 lb) of meat per day for sustenance; however, this certainly doesn't mean that a wolf will get the chance to eat everyday. In fact, wolves rarely eat on a daily basis, and so they make up for this by eating up to 9 kg (20 lb) of meat when they get the chance.

When pursuing large prey, wolves generally attack from all angles, targeting the necks and sides of such animals. Wolf packs test large populations of prey species by inducing a chase, thereby picking and ganging up on an individual that they perceive to be less fit. Less fit animals typically include the elderly, diseased, and young, and though these individuals are among the most likely to fall victim to predation, healthy animals may also succumb through circumstance or by chance. Hence, while wolves are certainly capable of culling the least fit from the communities of animals upon which they prey, the process certainly doesn't target the feeble or the ill-suited to an outright degree. Healthy, fit individuals will stand their ground against wolves, and are simply better able to effectively defend themselves, increasing the possibility of injury for the wolves involved. Realizing this, wolves are not likely to spend much time testing, chasing, or harassing such individuals. Probability dictates that these tactics are much more useful against lame, young, or old prey animals, and so it is these individuals that are most likely to fall to wolf predation. Even so, pack hunting efforts are usually fruitless. In one study, less than 1 out of 10 chases of moose resulted in a successful kill. Therefore, wolves must hunt almost constantly to sustain themselves.

Wolves, like many other keystone predators, are somewhat sensitive to fluctuations in prey abundance, making them likely to experience minor changes within their own populations as the abundance of their primary prey species gradually rises and drops over long periods of time. This balance between wolves and their prey prevents the mass starvation of all species involved.

Historical perceptions

Summary

The relationship between humans and wolves has had a very long and turbulent history. Traditionally, humans have viewed wolves negatively, perceiving them to be dangerous or as nuisances to be destroyed. European folklore exacerbated this negative image, which was brought over to North America as it was settled. In brief, the gray wolf, which, at one point, could be found in any ecosystem on every continent in the Northern Hemisphere, was persistently one of the first species to go once a significant population of humans settled in a given area. As technology made the killing of wolves and other predators easier, simple control became something more like complete annihilation.

Historically, baseless fear of the wolf has been responsible for most of the trouble the species has received, including why it was nearly hunted out of existence in the U.S. and Europe prior to the twentieth century. However, ecological research conducted during the 20th century shed new light on wolves and other predators, specifically with regard to the critical role they play in maintaining the ecosystems to which they belong. As a result of this and other important factors, wolves have come to be viewed in a much more realistic, equitable way.

A general environmental awareness began to take root sometime in the middle of the twentieth century that forced people to re-think former notions, including those regarding predators. In North America, people realized that in over one hundred years of documentation, there had been no verified human fatality caused by an attack from a healthy wolf. Eventually, they found out that wolves, being naturally cautious and rightfully wary of humans, will almost always flee, perhaps only carefully approaching a person out of curiosity. In fact, it is likely that any documented wolf attack (of any severity) to this date was, actually, from a feral dog, a rabid individual, or the result of some sort of human provocation.

While progress has been made, traditional opinions prove difficult to change in some cases. Today and in the past, improved understanding and education have been the main allies in altering perceptions. As research improves and as people figure out better ways to educate people about the true nature of this planet's ecosystems, public acceptance of the wolf as an integral member of the earth's biological society will likely prevail.

Wolves in folklore

In many ancient myths, the wolf is portrayed as being brave, honorable, or intelligent. The best examples of such portrayals are demonstrated by some of the stories of the Native Americans. Elsewhere, the wolf was revered as the totem animal of Ancient Rome (see Romulus and Remus and Lupercalia). In Proto-Indo-European society, the wolf was most likely associated with the warrior class, and the word itself was subject to taboo deformation (the Latin lupus being an example of a mutated form of the original Proto-Indo-European *wlkwos).

The wolf has typically assumed a more negative image in legend, however, including in most of modern western folklore. Specifically, the wolf has often been portrayed as a creature to be feared. Two iconic examples of this image are the Big Bad Wolf and the werewolf (a human that transforms into a wolf by means of magic or a curse, and who is shunned and reviled by the rest of society). Norse mythology prominently includes three malevolent wolves, in particular: the giant Fenrisulfr, eldest child of Loki and Angrboda who was feared and hated by the Æsir, and Fenrisulfr's children, Skoll and Hati, who, according to legend, will devour the sun and moon at Ragnarok.

Despite their often negative image, wolves have periodically been credited in mythology, fiction, and reality with adopting, nursing, and raising human feral children, with the most famous examples being Romulus and Remus and Mowgli of The Jungle Book, respectively. In Mongolian mythology, the Mongols believe that they are descended from a male Gray Wolf and a white doe. In fact, the Mongols' greatest hero Genghis Khan called his people 'Clan of the Gray Wolf'.

Early perceptions of wolves in North America

From colonial times right up through World War II, the wilderness was perceived as something to be conquered, settled, and cultivated, as through Manifest Destiny or by man's inherent worldly right. Wolves, for a time, were partially valued for their fur, but in the majority of circumstances (and in some instances today), they were viewed as wholly worthless and decidedly despicable creatures. Humans destroyed the vast majority of their habitat and food source in North America, replacing forests and prairies with farms and wild ungulates with cattle. When the wolves preyed on what little of their natural prey was left, hunters complained; when they then began preying on the cattle, ranchers, of course, complained, too.

Eventually, American society's perception of the wolf was one defined by indifference, misapprehension, or outright hatred. With few vouching for them, wolves and other predators were destroyed en masse, resulting in a so-called "hunters' paradise" free from competing predators. To accomplish this, there was no limit to the extent hunters and trappers were willing to go in order to kill predators in large numbers. Besides traps, snares, and other mechanical methods, hunters would line carcasses with poison (usually strychnine), which would then kill the animals that preyed upon it. The corpses of affected animals would then themselves become poisonous, which tended to result in a rather long chain reaction of death. Essentially, wolves, like many predators during that time, managed to garner an incredible amount of hatred for a great variety of reasons, many of which were unfounded or embellished so as to fuel public cynicism and expedite the wolves' removal; and, indeed, they were removed from the contiguous 48 states almost completely over the subsequent decades (a small population in northern Minnesota was the only exception to the otherwise successful extirpation campaign).

Changing attitudes

In the early twentieth century, some significant research performed in the blossoming young field of ecology led to an important realization about the relationship between predators and the land. Aldo Leopold, a botanist, forester, and naturalist of the early twentieth century (who is now world-renowned), was at the forefront of this new science. Early in his career during the first quarter of the twentieth century, he encountered a pack of wolves in the mountains of the southwest. He described his encounter thus:

"We saw what we thought was a doe fording the torrent, her breast awash in white water. When she climbed the bank toward us and shook out her tail, we realized our error: it was a wolf. A half-dozen others, evidently grown pups, sprang from the willows and all joined in a welcoming melee of wagging tails and playful maulings. What was literally a pile of wolves writhed and tumbled in the center of an open flat at the foot of our rimrock.

"In those days we had never heard of passing up a chance to kill a wolf. In a second we were pumping lead into the pack, but with more excitement than accuracy; how to aim a steep downhill shot is always confusing. When our rifles were empty, the old wolf was down, and a pup was dragging a leg into impassable side-rocks.

"We reached the old wolf in time to watch a fierce green fire dying in her eyes. I realized then, and have known ever since, that there was something known only to her and to the mountain. I was young then, and full of trigger-itch; I thought that because fewer wolves meant more deer, that no wolves would mean hunters' paradise. But after seeing the green fire die, I sensed that neither the wolf nor the mountain agreed with such a view."

—an excerpt from "Thinking Like a Mountain" (A Sand County Almanac)

Encounters similar to Leopold's were being repeated throughout North America. Early ecologists were the first to recognize the fundamentally flawed rationale behind the creating of the "hunters' paradise." It soon became apparent that overzealous deer, lacking predators, could cause unimaginable devastation to a landscape by browsing every edible bush, tree, and shrub, thereby preventing new growth from ever reaching prominence and ultimately reducing entire ranges to a state of utter desolation. In other words, "...just as a deer herd lives in mortal fear of its wolves, so does a mountain live in mortal fear of its deer. (Leopold)" Eventually, the deer herd itself would starve of its own inordinateness.

After Leopold's death, the groundbreaking research of wolf scientists such as Farley Mowat, L. David Mech, and Adolph Murie helped to shift the wolf's image to that of an intelligent and affectionate creature essential to the proper functioning of a conventional North American ecosystem. This awareness widely exposed the beneficial nature of wolves, and helped lead to their eventual endangered classification and subsequent reintroduction efforts. Accordingly, while the stereotype of wolves as malicious, wanton killers and vile, worthless beasts still has influence in certain circles, a significant portion of the public has developed a more positive opinion of wolves as interesting, valuable, and noble animals. Society as a whole has begun to realize the morality in attempting to make up for centuries of undue persecution, and knows of the justification behind trying to bring some ecological integrity back to the American landscape.

Since the late 1960s, wolves have been officially protected in the U.S. in some form or another by a national endangered species bill, the latest of which is the Endangered Species Act of 1973. Additionally, in a somewhat novel development, they have been allowed to naturally propagate in the upper Midwest, and have been reintroduced to areas in Wyoming, Idaho, and Arizona. In Alaska, where they are not protected by the Act, their populations continue to be controlled (usually by aerial hunting) in an effort to increase yields for hunters.

Today, organizations such as the International Wolf Center, Defenders of Wildlife, and Mission: Wolf continue to attempt to educate people about the true nature of wolves, and such action has proven helpful to past reintroduction efforts, especially in places like Yellowstone National Park.

The large amount of research done on wolves in the last half century has also helped to educate people in a way that helps them to realize how sociologically similar humans are to wolves, and how people have little to fear from these naturally cautious, complex animals. Biologists L. David Mech, Rolf Peterson and Luigi Boitani have arguably been the three leaders in contemporary wolf research.

This onslaught of pro-wolf publicity, including that which has been procured from nature documentaries and books, has undoubtedly played a role in changing attitudes for the better. Such mediums tend to emphasize the wolf as an affectionate, devoted parent and fraternal animal that is deserving of our respect and protection as integral members of our global biodiversity.

Reintroduction

Yellowstone National Park and Idaho

Wolf packs were reintroduced to Yellowstone National Park and Idaho starting in 1995. These wolves were considered as “experimental, non-essential” populations per article 10(j) of the Endangered Species Act. Such classification gave government officials greater leeway in dealing with problem wolves, which was considered one of a series of compromises wolf reintroduction proponents made with concerned local ranchers.

Indeed, local industry and environmental groups battled for years over the Yellowstone and Idaho wolf reintroduction effort. The idea of wolf reintroduction was first brought to Congress in 1966 by biologists who were concerned with the critically high elk populations in Yellowstone. Officially, 1926 was the year the last wolves were killed within Yellowstone’s boundaries, and over the succeeding decades, populations of elk and other large prey animals had soared, and new growth vegetation suffered as a result. Moreover, coyotes tried to fill in the niche left by wolves, but were unable to control the large ungulate populations. Booming coyote numbers, furthermore, also had a negative effect on other species, particularly the red fox. Ranchers, though, remained steadfastly opposed to reintroducing a species of animal that they considered to be analogous to a plague, citing the hardships that would ensue with the potential loss of stock caused by wolves.

The government, which was charged with creating, implementing, and enforcing a compromise, struggled for over two decades to find middle ground. A wolf recovery team was appointed in 1974, and the first official recovery plan was released for public comment in 1982. General public apprehension regarding wolf recovery forced the U.S. Fish and Wildlife Service to revise their plan to implement more control for local and state governments, so a second recovery plan was released for public comment in 1985. That same year, a poll conducted at Yellowstone National Park showed that 74% of visitors thought wolves would improve the park, while 60% favored reintroducing them. The preparation of an environmental impact statement, the last critical step before reintroduction could be green-lighted, was halted when Congress insisted that further research be done before an EIS was to be funded.

In 1987, in an effort to shift the burden of financial responsibility from ranchers to the proponents of wolf reintroduction, Defenders of Wildlife set up a “wolf compensation fund” that would use donations to pay ranchers market value for any stock that was lost to wolf depredation. That same year, a final recovery plan was released. Following a long period of research, public education, and public commenting, a draft EIS was released for public review in 1993 and it received over 150,000 comments from interested parties. It was finalized in May 1994, and included a clause that specified that all wolves reintroduced to the recovery zones would be classified under the “experimental, non-essential” provision of the ESA. Though the original plan called for three recovery zones – one in Idaho, another in Montana, and a final one in the Greater Yellowstone Area – the Montana recovery zone was eliminated from the final EIS after it had been proven that a small, but breeding population had already established itself in the northwestern part of the state.

A pair of lawsuits filed in late 1994 put the whole recovery plan in jeopardy. Interestingly, while one of the lawsuits was filed by the Wyoming Farm Bureau, the other was filed by a coalition of concerned environmental groups. The latter pointed to unofficial wolf sightings as proof that wolves had already migrated down to Yellowstone from the north, which, they argued, made the plan to reintroduce an experimental population in the same area unlawful. According to their argument, if wolves were already present in Yellowstone, they should rightfully be afforded full protection under the ESA, which, they reasoned, was preferable to the limited “experimental” classification that would be given to any reintroduced wolves.

Nevertheless, both cases were thrown out on January 3, 1995. Adolescent members from packs of Mackenzie Valley wolves in Alberta, Canada were tranquilized and carted down to the recovery zones later that week, but a last minute court order delayed the planned releases. The stay came from an appellate court in Denver and was instigated by the Wyoming Farm Bureau. After spending an additional 36 hours in transport cages inside the recovery zones, the wolves were finally released following official judicial sanction. Yellowstone’s wolves stayed in acclimation pens for two more months before being released into the wild. Idaho’s wolves, conversely, were given a hard release. A total of 66 wolves were released to the two areas in this manner in January 1995 and January 1996.

2005 estimates of wolf populations in the two recovery zones reflect the success the species has had in both areas:

• Greater Yellowstone Area: 325
• Central Idaho: 565

These numbers, added with the estimated number of wolves in northwestern Montana (130), puts the total number of wolves in the Northern Rocky Mountains recovery area at over 1000 individuals. This includes approximately 134 packs (two or more wolves traveling together) and 71 breeding pairs (male and female that successfully rear a litter of at least two until Dec. 31). The recovery goal for the area was 30 breeding pairs total, and this number has been surpassed for some time.

Since wolves have been recovered, there have been hundreds of confirmed incidents of livestock depredation, though such predation represents a minute proportion of a wolf’s diet on a per wolf basis. While the majority of wolves ignore livestock entirely, a few rogue wolves or wolf packs will become chronic livestock depredators, and will either be relocated or killed depending on the area and number of incidents. Since the year Defenders of Wildlife implemented their compensation fund, they have allocated over $500,000 to private owners for proven and probable livestock depredation by wolves.

Arizona

The five last known wild Mexican grey wolves were captured in 1980 in accordance with an agreement between the United States and Mexico intended to save the critically endangered species. Since then, a comprehensive captive breeding program has brought the subspecies back from the brink. Currently, there are 300 captive Mexican wolves taking part in the program.

The ultimate goal for these wolves, however, is to reintroduce them to areas of their former range. In March 1998, this reintroduction campaign began with the releasing of three packs into the Apache-Sitgreaves National Forest in Arizona. Today, there may be up to 50 wild Mexican wolves in Arizona and New Mexico. The final goal for Mexican wolf recovery is a wild, self-sustaining population of at least 100 individuals.

Scandinavia

In Sweden and Norway, there has been a long and ongoing conflict between some groups whose belief it is that wolves no longer have a place in nature and those who wish the wolf to be allowed to expand out into more of the area’s vast boreal forests. The former mostly consists of members of the rural working class who fear competition for certain large ungulate species (Roe Deer, moose, etc.), and who consider the wolf to be a foreign element. They argue that modern Scandinavian wolves are actually recent migrants from Russia and not the remnants of old native wolf packs, which, they reason, is why they don’t belong in Sweden and Norway. Recent DNA research seems to support this claim.

Scandinavian wolves had been nearly completely eliminated from the range due to extirpation campaigns in the nineteenth and twentieth centuries, and were considered to be gone from the area by the 1960s. In the early 1980s, however, a single breeding pack was discovered in southern Sweden, over 1000 km away from the nearest known population in Russia or Eastern Finland. The pack was small – about ten animals – and it stayed that way for many years until its population began to noticeably increase starting in 1991. Prior to 1991, the small population lacked ideal genetic diversity, and inbreeding had been occurring to a potentially dangerous degree. Furthermore, low birth rates suggest that the wolves were apprehensive to mate with each other, which was most likely due to their close relation. Genetic data suggests that, in 1991, a lone immigrant wolf from Russia migrated to the area and single-handedly restored genetic diversity to the population. A particular study showed that of the 72 wolves born between 1993 and 2001, 68 of them could trace their genetic heritage to this lone migrant wolf. Today, there are over one hundred individuals that range across this southern area of Scandinavia. The population remains genetically isolated, however, which is a cause of concern for some. On the other hand, there is reason to believe that as the number of wolves living in this area increases, the boundaries of the population's range will creep towards the ranges of other, separate populations in Finland, thus promoting dispersal. Direct reintroduction remains an intriguing option to foster genetic diversity in the Scandinavian population in the meantime.

There has been much speculation as to how the original population came to be in the early 80s. Some believe that they might be a native species – remnants of a population that somehow survived persecution. Much genetic research has been performed on this population, however, and this particular theory isn’t supported by the findings. Genetic analysis seems to support the idea that the wolves were immigrants that had traveled over 1000 km from Russia to southern Scandinavia along one of several possible dispersal routes. Certain conspiracy theorists believe that they were artificially reintroduced per some secret agenda by the Swedish government.

Since the wolves have reestablished themselves, Norwegian and Swedish farmers have complained of sheep and dog depredation. Indeed, many ranchers in Norway were forced to give up their practice once local wolves discovered sheep as potential prey. This exemplifies the general trend that the people who are usually the most skeptical about wolf recovery, though they typically represent the minority, are also the ones most directly affected by it. Most of the proponents of wolf reintroduction in Norway and Sweden can be found in urban populations, which is a pattern that can be seen wherever wolf reintroduction is a hot button issue. As a result, some are calling for the legalization of hunting wolves in this area. E.U. regulation doesn't make this an option in Sweden. However, government action could be taken to cull wolf populations if either of the two countries involved should sanction such action.

Wolf hunting

See also: Wolf hunting

Livestock predation

As long as there are enough prey, wolves seem to avoid taking livestock. However, some wolves or packs can specialize in hunting livestock once the behavior is learned despite relative food supply. Sheep are frequently the most vulnerable, while horses and cattle are also at risk. Wolf-secure fences, relocation (in some cases), or local extermination of wolves are today the only known methods to effectively stop livestock predation.

Over several centuries in some countries, shepherds and dog breeders have used selective breeding to "create" large livestock-guarding dogs that can stand up to wolves preying on flocks. In the United States, as the timber wolf and other large predators been reintroduced, the USDA has been looking into the use of breeds such as the Akbash from Turkey, the Maremma from Italy, the Great Pyrenees from France and the Kuvasz from Hungary, among others.

Some ranchers in the United States hunt wolves from helicopters or light planes, some of them calling it an effective method of controlling wolf numbers, others calling it a sport. This practice is seen as highly controversial. Poisons have been used to kill wolves during the extermination campaigns in Europe and America. Today, most of the hunting is done on the ground or from helicopters. Other, non- or less-lethal methods of protecting livestock from wolves have been under development for the past decade. Such methods include rubber ammunition, and use of guard animals.

Trapping and breeding for fur

Wolves are frequently trapped, in the areas where it is legal, using snares or leg-hold traps. The economic value of wolf pelts is limited, so it is mainly a recreation activity. Wolf trapping has come under heavy fire from animal rights groups, who also use it to attack other forms of trapping and hunting. It is alleged that trapping, using the right tools and equipment, can be considered as humane as hunting; however, unskilled trappers can create unnecessary suffering in animals.

Wolves are bred for their fur in very few locations, as they are considered as a rather problematic animal to breed, and combined with the low value of the pelt, it has driven most of the fur farms to change to utilizing other animals, such as the fox.

Darting and box traps are important tools used by wildlife biologists who use the techniques to catch wolves, fit them with collars holding radio transmitters, and check their health before releasing the wolves. Use of such radio collars enable biologists in airplanes to track the wolves and count pack sizes. The radio telemetry is an important component in the suite of tools used in Wisconsin to calculate the overall wolf population.

Taxonomy

Classification and relation to the dog

Much debate has occurred over the relationship between the wolf and the domestic dog. Most authorities see the wolf as the dog's direct ancestor, but others have postulated descent from the Golden Jackal. Because the canids have evolved recently and different canids interbreed readily, untangling the true relationships has presented difficulties. However, molecular systematics now indicate very strongly that domestic dogs and wolves are more closely related than either is to any other canid, and the domestic dog is now normally classified as a subspecies of the wolf: Canis lupus familiaris.

The classification of wolves and closely allied creatures offers many challenges. Although taxonomists have proposed many species over the years, most types clearly do not comprise true species. Indeed, only a single wolf species may exist. Scientists have proposed a host of subspecies. Many of these seem unlikely to stand. Further taxonomic clarification may well take decades.

Subspecies of the wolf

The subspecies for the Gray Wolf has been a very controversial issue among taxonomists. It was once believed there were as many as 50 separate subspecies. However, the last decade has seen a new and widely accepted list that has been condensed to 13 living subspecies, and 2 recently extinct subspecies. This takes into account the anatomy, distribution, and migration of various wolf colonies.

Media

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See also

• Maned Wolf
• Werewolf
• Wolfdog
• Wild Canid Survival and Research Center
• Wolves in fiction
• Wiktionary:Wolf

Three other extant canid species also known as wolves:

• Red Wolf
• Eastern Canadian Wolf
• Ethiopian Wolf
• Mexican Wolf

Extinct species of wolf:

• Dire Wolf

References

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2. Wolves. What is a wolf? URL accessed on August 18, 2005.
3. "Eyes on Wildlife". Natural History of the Gray Wolf. Retrieved August 21. {{cite web}}: Check date values in: |accessdate= (help); Unknown parameter |accessyear= ignored (|access-date= suggested) (help)
4. "World of the Wolf". The skull of Canis lupus. Retrieved August 21. {{cite web}}: Check date values in: |accessdate= (help); Unknown parameter |accessyear= ignored (|access-date= suggested) (help)
5. "Animal Diversity Web". Canis lupus. Retrieved August 18. {{cite web}}: Check date values in: |accessdate= (help); Unknown parameter |accessyear= ignored (|access-date= suggested) (help)
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7. Fred H. Harrington. "NOVA Online." What's in a Howl? Accessed on August 21, 2005.
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10. "Wolf Web". Wolf Pack. Retrieved August 21. {{cite web}}: Check date values in: |accessdate= (help); Unknown parameter |accessyear= ignored (|access-date= suggested) (help)
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12. "Smithsonian National Zoological Park". Gray Wolf Facts. Retrieved August 15. {{cite web}}: Check date values in: |accessdate= (help); Unknown parameter |accessyear= ignored (|access-date= suggested) (help)
13. Mary Taylor Young. "Colorado's Wildlife Company". CWC Summer 2004: A Wolf Primer. Retrieved August 15. {{cite web}}: Check date values in: |accessdate= (help); Unknown parameter |accessyear= ignored (|access-date= suggested) (help)
14. "Natures Wolves". Wolf Attacks On Humans. Retrieved October 31. {{cite web}}: Check date values in: |accessdate= (help); Unknown parameter |accessyear= ignored (|access-date= suggested) (help)
15. "Defenders of Wildlife". A Yellowstone Chronology. Retrieved May 3. {{cite web}}: Check date values in: |accessdate= (help); Unknown parameter |accessyear= ignored (|access-date= suggested) (help)
16. "USFWS" (PDF). Rocky Mountain Wolf Recovery 2005 Interagency Annual Report. Retrieved May 3. {{cite web}}: Check date values in: |accessdate= (help); Unknown parameter |accessyear= ignored (|access-date= suggested) (help)
17. "USFWS" (PDF). Mexican Grey Wolf Fact Sheet. Retrieved May 4. {{cite web}}: Check date values in: |accessdate= (help); Unknown parameter |accessyear= ignored (|access-date= suggested) (help)
18. "Nature". Conservation biology: Lone wolf to the rescue. Retrieved May 4. {{cite web}}: Check date values in: |accessdate= (help); Unknown parameter |accessyear= ignored (|access-date= suggested) (help)
19. "Wildlife Biology" (PDF). The origin of the southern Scandinavian wolf ‘’Canis Lupus’’ population. Retrieved May 4. {{cite web}}: Check date values in: |accessdate= (help); Unknown parameter |accessyear= ignored (|access-date= suggested) (help)

External links

• Sinapu
• International Wolf Center
• Mission: Wolf
• Canids.org
• Wolf Park
• The Iberian Wolf
• Species Overview at Enature.com
• The National Wildlife Federation
• Click for Wolves
• Defenders of Wildlife

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• Wolves
• Mammals of India
• Arctic land animals
• Fauna of Canada
• Extinct animals of the United Kingdom