Two
SOCIAL STRUCTURE AND COMMUNICATION

THE SOCIAL STRUCTURE OF a wolf pack is all-important. Breeding, hunting, and feeding are tied to it, as is territorial maintenance and play behavior. What evidence we have of the wolf’s ability to teach its young to hunt (as well as their ability to learn) suggests that social structure plays a strong role here, too. Wolf pups raised without a pack structure adapt very poorly to life in the wild.

Generally speaking, there are three separate social structures in the pack: a hierarchy of males, a hierarchy of females, and a more seasonally related cross-sexual social structure. There is, typically, an “alpha,” or primary, male that dominates the other males and an alpha female that dominates the other females. This alpha pair is thought to be the breeding pair, but there are a good many cases in captivity and in the wild where a lower-ranking male has bred with the alpha female, the alpha male expressing apparent disinterest.

Females may head packs and they always strongly influence pack activities. We often think of animals like the wolf, who appear to have so many points in common with us, in human social terms. With respect to females, who have largely a subordinate standing in Western human societies, the analogy, I think, is poor. Female wolves may not only lead packs but outlast a succession of male alpha animals. It is females, moreover, who decide where to den and thus where the pack will have to hunt for five or six weeks. In the north, where wolves are following migrating caribou, a poor guess about where caribou are going to be can be disastrous, and the possibility of failure underlines the gravity of the female’s decision.

Young females, as mentioned, are thought to be slightly faster than young males and therefore better hunters under some circumstances. The male hunter-male leader image of the wolf pack is misleading but, unconsciously, I am sure, it is perpetuated by males, who dominate this field of study. For the same reason paramilitary descriptions of wolf behavior—where “lieutenant wolves” are “dispatched” to “patrol” the territory, and parents “instill discipline” in the pups—creep in and strongly color our impressions of the animal, often without our knowing it. I am certain this is part of the reason people believe that male wolves always do the killing, and why the males’ weights are so often exaggerated.

Social structure in a wolf pack has been observed in greatest detail among captive wolves, which makes extrapolating to wild wolves risky. Captive animals engage in no hunting activities, are penned in areas of incomprehensible size when compared with the one-hundred-plus square miles routinely used by a pack, often lack for suitable exercise, and are constantly interfered with by human beings seeking to establish or maintain various levels of socialization. Their social displays, because they can never get away from each other, are excessive. Fieldwork has substantiated that a strong social order does exist in the wild and that there is an alpha pair at the top. It is most evident during the mating season when the alpha female asserts herself and sometimes fights to keep other females in estrus away from the alpha male, and the alpha male fights, less vigorously it seems, to keep other males away from the alpha female. (These are ritualized squabbles with few physical injuries in most cases.) It is not known whether it is the alpha male that normally fathers the pups, but they are almost always whelped by the alpha female.

Lesson

A female wolf left four or five pups alone in a rendezvous area in the Brooks Range one morning and set off down a trail away from them. When she was well out of sight, she turned around and lay flat in the path, watching her back trail. After a few moments, a pup who had left the rendezvous area trotted briskly over a rise in the trail and came face to face with her. She gave a low bark. He stopped short, looked about as though preoccupied with something else, then, with a dissembling air, began to edge back the way he had come. His mother escorted him to the rendezvous site and departed again. This time she didn’t bother watching her back trail. Apparently the lesson had taken, for all the pups stayed put until she returned that evening.

But the term alpha—evolved to describe captive animals—is still misleading. Alpha animals do not always lead the hunt, break trail in snow, or eat before the others do. An alpha animal may be alpha only at certain times for a specific reason, and, it should be noted, is alpha at the deference of the other wolves in the pack.

The wolf is a social animal; it depends for its survival on cooperation, not strife. Human beings, particularly in recent years, have grown accustomed to speaking of “dominance hierarchies” in business corporations and elsewhere, and the tendency has been to want wolf packs (or troops of chimpanzees) to conform to similar molds. The social structure of a wolf pack is dynamic—subject to change, especially during the breeding season—and may be completely reversed during periods of play. It is important during breeding, feeding, travel, and territorial maintenance, and seems to serve a purpose when wolves gather to reassure each other of the positive aspects of their life-style as reflected in this social order, one that enhances survival by collective hunting and natural population control.

To place a heavy emphasis on such supposed facets of behavior as “intimidation,” “pulling rank,” and games of psychological cruelty based on social structures, however, is simply to confuse the tools of human analysis with the actual behavior of wolves.

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Four weeks old.

Another factor to be considered is that wolf packs, like individual wolves, have personalities. Packs may contain autocrats, petulant individuals, or even cretins; and the personalities involved may make one pack more Prussian or austere in its organization than another.

With this to mind, the following can be taken as a “typical” pack structure: an alpha male and alpha female at the top, perhaps four or five years old; subordinate males and females, some sexually mature, in the middle, the dominant of which are called beta animals; and the pups. Deference is shown to alpha animals by subordinate animals of both sexes. Subordinates of the same sex establish their own order, usually with the yearlings at the bottom and a two- or three-year-old animal at the top. The pups, too, have their own social order, but theirs is without regard to sex. The alpha animals and the others gradually bring the pups into the social structure as the litter stabilizes toward the end of summer. Wolf discipline is firm but not ruthless. An errant pup is typically held by the muzzle and pinned momentarily to the ground.

After the pups are weaned, the other members of the pack play an increasingly greater role in their upbringing, providing both food and recreation. The pups eat partially digested food regurgitated for them by adults. Babysitting adults play rag doll to the pups, who now have needle-sharp milk teeth and are mobbing each other, wrestling, biting, and generally grabbing reclining adults vigorously by the ruff of the neck. This behavior will later be molded into efficient hunting technique.

Daily activities center around the mouth of the den until the pups are about eight weeks old, at which time the adults move them to the first of a series of rendezvous sites where they remain while the others hunt. By late fall the pups may weigh forty-five to fifty pounds and accompany hunting adults. The knack of taking large animals like deer, moose, elk, mountain sheep, and caribou is behavior that must be taught; trapping mice with a bilateral stab of the forelimbs or surprising and catching a snowshoe hare comes more naturally.

By the time the pups are a year old they are almost full grown. Some disperse and either spend time alone or attempt to join another pack, or follow their own pack at a distance; others stay on with the pack for another year, until they reach sexual maturity.

There is probably a significant difference from year to year in the way a particular pack of wolves gets along. The loss of a good hunter, a drop or rise in the prey population, a prolonged winter, excessive social tension within the pack—all these are part of the ebb and flow of life for the wolf. Although no formal studies have been made, field studies of more than a year’s duration suggest an annual cycle of individual identification with the pack that takes on unusual significance in the light of the suggestion that wolves seem to have both a sense of self and a sense of the pack and of preserving it.

Social tension increases in late winter with the evolution of a mating pair in the pack, and the level of anxiety is sometimes marked. Once two wolves have mated, however, emotions abate. The pack may split up to hunt more effectively. The pack’s identity at this time may be at its loosest, with members hunting in ones and twos. The pregnant female, perhaps accompanied by her mate, selects a den site a week or more before she whelps, and the pack drifts together again to provide for her and the pups. This is undoubtedly a most difficult time in the year for the wolf. Activities center on the den site, when the most efficient kind of hunting would be one without such a locus. The pups are vulnerable and a few inevitably succumb quickly to exposure or starvation. There must be among wolves some sense of the importance of the population, as defined more by the hunting and reproductive unit (the pack) than by the individual, that leads wolves to stay together at this time in an effort to raise the pups.

With the calving of local prey species and the increased mobility that comes with moving the pups from the den to a series of sites from which the adults hunt, a climb up from this annual low point begins. The pack is given to emotional greeting, general nuzzling, and increasing horseplay as the summer goes on. They eat well. The weather is good. Their emotional peak must come in the fall and early winter. The pups are able to travel with the adults. Game is fat. The healthiest wolves have a tone of well-being, a strength of bone and mind, that will take them safely through the winter and into spring. They look to each other more often now, knowing that soon it will be impossible to get by alone as one could have done in summer. The increase in howls, their coming together frequently and wagging their tails as they orient around the alpha animals, marks the onset of winter.

After many years of work in the wild studying wolves, Adolph Murie wrote that the strongest impression he was left with was of the wolves’ friendliness toward each other. Most systems of human description of animal behavior fall abysmally short in this area, which is unfortunate where wolves are concerned. Even as adults, wolves play tag with each other or romp with the pups, running about a clearing or on a snowbank with a rocking-horse gait. They scare each other by pouncing on sleeping wolves and by jumping in front of one another from hiding places. They bring things to each other, especially bits of food. They prance and parade about with sticks or bones in their mouths. I recall how one Alaska evening, the sun still bright at 11:30 P.M., we watched three wolves slip over the flanks of a hill in the Brooks Range like rafts dipping over riffles on a river. Sunlight shattered on a melt pond ahead of them. Spotting some pintail ducks there, the wolves quickly flattened out in the blueberries and heather. They squirmed slowly toward the water. At a distance of fifty feet they popped in the air like corks and charged the ducks. The pintails exploded skyward in a brilliant confusion of pounding wings, bounding wolves, and sheets of sunburst water. Breast feathers from their chests hung almost motionless in midair. They got away. The wolves cavorted in the pond, lapped some water, and were gone. It was all a game.

The social relationships of wolves are maintained through three systems of interpersonal and interpack communication: vocalization, postural signaling, and scent marking.

The wolf’s howl is the social signal perhaps most familiar to everyone. It typically consists of a single note, rising sharply at the beginning or breaking abruptly at the end as the animal strains for volume. It can contain as many as twelve related harmonics. When wolves howl together they harmonize, rather than chorus on the same note, creating an impression of more animals howling than there actually are. Wolves do not have to stand to howl. They can howl lying down or sitting on their haunches. I’ve even seen a wolf, with an air of not wanting to miss out, howl while defecating.

There has been more speculation about the nature and function of the wolf’s howl than the music, probably, of any other animal. It is a rich, captivating sound, a seductive echo that can moan on eerily and raise the hair on your head. Wolves apparently howl to assemble the pack, especially before and after the hunt; to pass on an alarm, especially at the den site; to locate each other in a storm or in unfamiliar territory; and to communicate across great distances. Some Eskimos, according to writer/naturalist Farley Mowat, claim to be able to understand what wolves are howling about and to take advantage of it when the howling reveals the approach of migrating caribou. The howl may carry six miles or more in still arctic air.

There is little evidence that wolves howl during a chase, but they may do so afterward, perhaps to celebrate a successful hunt (the presence of food), their prowess, or the fact that they are all together again, that no one has been injured. Adolph Murie, who had an eye for such things, reported a lone wolf howling while hunting mice.

There has never been any evidence that wolves howl at the moon, or howl more frequently during a full moon, though howling may be more frequent in the evening or early morning. Howling reaches a seasonal peak in the winter months, during the time of courtship and breeding; it is easy to see how the idea that wolves howl at the moon might have gained credence and played well on the imagination during these cold, clear nights when the sound carried far and a full moon lent an eerie aspect to a snowscape.

Howl

It was wild, untamed music and it

echoed from the hillsides

and filled the valleys. It sent

a queer shivering feeling along my

spine. It was not a feeling

of fear, you understand, but a sort of

tingling, as if there was hair on my back

and it was hackling.

—ALDA ORTON, Alaskan trapper

What emotions prompt a howl remain unknown, though field and laboratory researchers both suggest that solo howls and group howling alike are brought on by restlessness and anxiety. Loneliness is the emotion most often mentioned, but group howling has a quality of celebration and camaraderie about it, what wildlife biologist Durward Allen called “the jubilation of wolves.” Murie writes of four wolves assembled on a skyline, wagging their tails and frisking together. They began to howl, and while they did so a gray female ran up from the den a hundred yards away and joined them. She was greeted with energetic tail wagging and general good feeling, then they all threw back their heads and howled. The howling, wrote Murie, floated softly across the tundra. Then, abruptly, the assembly broke up. The mother returned to the den and the pups; the others departed on the evening hunt.

Similar actions among Cape hunting dogs have been called mood-synchronizing activities by one researcher.

The wolf’s other vocalizations have received less attention, though wolves seem to use these other sounds more often, to communicate more information. They are commonly divided into three categories: growls, barks, and whines or squeaks. Howls have been recorded and studied in the wild. Growls, squeaks, and barks have only rarely been heard in the field, so we must proceed here solely on the basis of information from captive animals.

Wolves only infrequently bark, and then it is a quiet “woof” more often than a dog-type bark. They do not bark continuously like dogs but woof a few times and then retreat, as for example when a stranger approaches the pen. Barks reported from the field are associated with a pack’s being surprised at its den and an animal, usually the female, rising to bark a warning.

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The wolf in the middle in the photograph on the left begins to howl in response to the others. In chorus like this, each wolf chooses a different pitch. The production of harmonics (see chart, page 42) may create the impression of fifteen or twenty wolves where there are in fact only three or four.

Growling is heard during food challenges and, like the bark, is associated with threat behavior or an assertion of rights in some social context. To the human ear this is perhaps the most doglike wolf sound in terms of its association with other behaviors, such as a squabble over a bone. Growling is common among pups when they’re playing. Pups also growl when they jerk at the ruff of a reclining adult and comically will even try to growl adults off a piece of food. Another type of growl is a higher-pitched one that begins to sound like a whine and often precedes a snapping lunge at another wolf.

Perhaps the most interesting sounds are the whines and high-pitched social squeaks associated with greeting, feeding the pups, play, pen pacing, and other situations of anxiety, curiosity, and inquiry. They are the sounds of intimacy.

Over a period of weeks one spring I observed a captive pack that included four pups. With the aid of underground microphones in the den, I was able to listen to sounds that otherwise would have been inaudible. The pups frequently wrestled down there, growling and yipping, but they would stop immediately when the mother came to the den entrance and squeaked—usually, but not always, a call for food. Sometimes it was a call to play. It was the custom with this pack for the alpha animals to take meat (chicken) from the hand of their caretaker and to then call the others—the pups and yearlings—with squeaks. They would all trot over. Then adults and young alike would engage in a twisting, supple dance —everyone squeaking, the young jabbing at the adults’ muzzles—until the adults regurgitated the meat.

The mother would squeak on occasion when the pups were playing too roughly; the father would occasionally call the pups over to him with a squeak, and they would all just stand there and nuzzle. Reassurance, perhaps.

GROWL

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BARK

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HOWL

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WHINE

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Some squeaks were repeated often enough to be recognized; these were associated with certain specific behaviors, leading one to think of them as bits of true communication.

Wolves hear well up to a frequency of 26kH. (beyond the range of human hearing, in the range where bats and porpoises produce sound), but what wolves do with high-frequency information remains a mystery. The ability to detect high-pitched sounds—wolves can distinguish between sounds a single tone apart in the range of 10-15kH.—may help them locate rodents under a snowpack. Many Russian scientists believe that wolves hunt even large game more by sound than smell and that the wolf’s range of hearing and the fineness of its auditory discrimination make this its keenest sense.

Postural communication is composed of a variety of facial expressions and tail positionings, as well as such things as piloerection (raising the hackles). Lunging, chasing, body slamming and fighting, and more subtle gestures may also be considered in this category.

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Rudolph Schenkel in 1947 was the first to document the great range of facial expression in the wolf and to relate it (and tail positions) to moods or feelings in the animal. These simple classifications for gestures—“suspicion,” “threat,” “anxiety,” “submission”—later came to be understood in a more complicated fashion. Today the tendency is to treat postural gestures dynamically, as part of a complex of behaviors, augmented by vocalizations, flavored by personal idiosyncrasies.

When an alpha male encounters a low-ranking male in the same pack, he may stand erect and still, his tail horizontal in line with his spine, and stare at the other animal. The subordinate animal generally lowers his body, holds his tail down, turns slightly away from the alpha male, and lays his ears back. In a more serious encounter he might retract the corners of his mouth to reveal his teeth in what is called a submissive grin and twist his head so that he is looking up at the alpha animal. This is called passive submission. The subordinate’s attempt to lick the alpha animal’s muzzle would be active submission.

To a casual observer such a display appears to clearly reveal a dominant/submissive relationship; however, an observer who was with the animals regularly and was familiar with their personalities and the history of the pack—fights, matings, alliances—could tell much more. Many encounters are simple, but it would be misleading to establish simplicity of gesture as the rule. An obviously “submissive” wolf may be expressing submission, fear, and defensive aggression simultaneously. Even trained ethologists have mistaken who the submissive animal in an encounter was, thinking that an inhibited bite (a submissive gesture) was an attack (a display of dominance).

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An intense moment between a dominant ten-year-old male on the left and a yearling female on the right. The split second captured here reveals, in the body language of wolves, annoyance on the part of the older animal with a partial erection of his tail (1), the raising of his hackles (2), the forward movement of his ears (3), and the vertical retraction of his lips (4). The subordinate animal indicates acquiescence with a flattening of her ears (5), a flashing of the white of her eyes (6), an appeasement gesture called “licking intention” (7), and a general lowering of her body (8). Her raised paw (9) may be the tail end of the gesture that elicited the adult’s response, or part of another appeasement gesture.

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Subtle gestures of postural and facial communication are accentuated by a set of dark lines marking the ears, eyes, muzzle, and shoulders.

Markings on the fur already mentioned emphasize detail in postural expression, just as lipstick emphasizes the lips or eye shadow the eyes in human beings. The dark tip of the tail creates simple contrast for increased visibility and also marks the area of the tail that twitches, a sign of excitement. A wolf’s black lips are set off against white hairs on the muzzle and lower jaw to emphasize this important area of the face, and the face itself is subtly marked, especially around the eyes. The ears are lined with light hairs and rimmed with dark.

The face is the focus of silent gesture in the wolf and Schenkel and others have identified a number of facial gestures, especially in connection with movements of the eyes, ears, and lips. The terms used convey something of the range and complexity: “licking intention,” “agonistic pucker,” “tooth snapping,” “intimidating stare.” In the wild, observers are rarely close enough to see any of this. Schenkel was particularly attentive to the angle of the ears and the wrinkling of the forehead as clues to the wolf’s intentions. One senses, I think, the excitement the man must have felt, sitting outside the wolf cage at the Basel Zoo, as he contemplated the slow emergence of a new kind of language, a priceless key to understanding behavior.

Nose pushing, jaw wrestling, cheek rubbing, and facial licking are common when wolves are together. One wolf mouthing another’s muzzle is a friendly gesture; clamping another’s muzzle between bared teeth is less friendly but not as ferocious as it appears to the human eye. “Standing over” and “riding up” are distinguished as body gestures; an animal may emphasize his dominance by standing over, or straddling, a reclining subordinate. The subordinate may respond by rolling over on his back and, in an extreme case, urinating a few drops on himself. When one wolf approaches another from the side, he may deliver a “hip slam,” and after more pushing and shoving ride up on the other animal by placing his forepaws on or over the other’s shoulders. This, too, is a gesture of dominance and is common among pups. Another gesture commonly seen is the stiff-legged approach of an alpha animal, which is at variance with the looser, bicycling motion of the other pack members.

It should be borne in mind that there are misunderstandings among wolves, just as there are in human relationships, and that observers occasionally glimpse amazement on the face of a wolf who catches an intimidating look meant for someone else. The idea of “submission” in a wolf pack, made visible to the human observer in ways human beings consider debasing, like rolling over on one’s back, can be taken, I think, too seriously. The submissiveness is not neurotic; it is essential to the maintenance of group harmony. Submissive gestures have quite logical and very pleasant antecedents. Puppies first roll over on their backs so their parents can lick them clean. They first nuzzle adults to trigger regurgitation. Much of what is seen in a wolf pen might more profitably be viewed as “reassurance displays” aimed at group harmony rather than submissive whimpering at the feet of an ogre.

Our attempts to understand wolf language are crude and based not a little on the belief that the animal is simple-minded and therefore speaks a simple language. There are sounds we can’t hear and there are signals we don’t see. As we begin to put vocalizations together with a more fluid and careful description of body movement, we are bound to discover considerably more about the wolf and his language of gesture.

The third kind of communication, about which we unfortunately know very little, is the complex of olfactory cues associated with scent marking and glandular secretions.

In the wild, a traveling alpha animal either scent-marks or inspects a scent mark on the average of once every two minutes—an indication of the singular importance of the activity. It is commonly believed that the primary function of scent marking is to mark out a pack’s territory and so warn off intruders; but American behavioral psychologist Roger Peters, one of the first to study the phenomenon, believes that territorial marking is a secondary function. The primary one is to mark territory on a regular basis for the benefit of the resident pack. The marks are an aid in establishing cognitive maps for the younger wolves, mental pictures of the home range, so that they know where they are with respect to certain creeks or recent kills and how to get where they wish to go. Secondly, scent marks help the pack to communicate when it is broken up. By reading scent marks one wolf can determine whether an area has been hunted recently, if a certain member of the pack is in the general area, or who has traveled through recently with whom. This, writes Peters, would ensure the efficient use of all parts of a pack’s territory. An analogy would be forest dwellers leaving messages for each other, stuck in a hole in a designated oak.

Peters divides scent marks into four categories: raised-leg urination, squat urination, defecation, and ground scratching or dirt kicking. The first category is the most important. The alpha animals (primarily the males) make this kind of scent mark. They are made against objects above ground level to ensure a large evaporative surface (for a stronger odor) and to keep the mark clear of rain or snow. Squat urination and defecation probably have scent-marking as well as eliminative functions; defecation may trigger anal glands, imparting a personal scent to a scat. Dirt scratching probably functions as a visual display of dominance for the benefit of other pack members, but it may serve still another purpose if glands between the toes are stimulated and another olfactory message is left behind.

Wolves carry on a regular pattern of scent marking, visiting each section of their territory on the average of once every three weeks as they travel established routes. The area is, in effect, studded with olfactory hotspots. To ensure that a scent mark will be detected by others in a minimum of time, the marks are concentrated around trail junctions. But wolves even scent-mark when they are pursuing game cross-country, so presumably wolves can always tell whether they are in their own territory or not.

Scent marks certainly warn off intruders, but their larger role seems to be in the maintenance of a sense of spacial organization for the resident pack. They may also help wolves to find open territories, and loners of the opposite sex—the nucleus of a new pack—to find each other.

Glandular secretions form another olfactory stimulus that plays an important role in intrapack communication, but we know little about them. Anal (gland) inspection is common among male wolves, with the dominant males readily presenting their anal parts for inspection and subordinate animals withdrawing theirs or presenting them only reluctantly. Females rarely engage in anal inspection except during the breeding season when, like the males, they may be attracted to the presence of vulvar blood.

Other glands include the supracaudal gland on top of the tail, usually marked off by a dark patch of slightly stiffer hair, the glands between the toes, and glands around the cheeks. The habit dogs have of rolling in putrid substances is also found in wolves. It seems possible that odors picked up in this way and carried to other pack members have some communicative function.

Roger Peters told me once that wolves in the Superior National Forest defecate sometimes on beer cans. Like any scent mark, these scats give off both visual and olfactory signals. We should see more here than what the wolf might be telling us about our littering habits. The animals may be marking things they consider dangerous to other wolves, especially pups, for wolves also mark traps and poisoned baits by defecating on them. If Peters is correct in thinking that the olfactory information in a scat is intended for other pack members, the idea makes even better sense.

No one knows what a wolf can smell, but the guess is that he is not so highly sensitive to faint odors—the human nose is even good at that—but that he is able to distinguish among many similar odors. What to the human nose just smells like “the woods” may for the wolf be hundreds of discrete bits of information. This power to discriminate, together with an olfactory memory (analogous to the auditory memory some people develop for hundreds of bird songs, i.e., which bird, which of its songs, at what time of day, etc.), becomes another way for the wolf to fathom his universe.

It is sometimes suggested that the wolf’s long nose is the result of selection for a keen sense of smell, but it is more probable that it evolved or was maintained as part of the need for large, powerful jaws.

There is another, far less obvious, kind of communication wolves employ which is perhaps extrasensory, or at least beyond our range of perception. I have noticed that captive animals at rest seem to pick up cues from each other even though there is no audible sound and they are out of visual contact. Their backs may be turned to each other or one may be off in some trees in a corner of the pen. When one animal stares intently at something, for example, it apparently creates some kind of tension. Other animals respond by lifting their heads and turning without hesitation to look at the area where the first animal is staring. In my experience it was most often the subordinate animals that responded first and the alpha animals last. Perhaps further research will establish a firmer foundation for this. It hints, of course, at much.

This may be the place to mention something that receives little attention, the fact that wolves kill each other. Recent efforts to change the bad image many people have of the wolf have led to the suggestion that while wolves may fight, the encounters aren’t fatal. This is not true. Wolves do kill each other, especially in captivity. In the wild, deaths are related most often to territorial trespass, especially when pups are threatened. Strangely behaving wolves—epileptic pups, wolves caught in steel traps and thrashing about, wolves crippled by a moose or gunshot—have been killed by their pack members. Wolves in the same pack rarely square off and fight to the death, flight being the rule for the loser, and they demonstrate an ability to work out disputes ritually, but disputes over an alpha position do sometimes come down to a bloody, eerily silent fight to the death.

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Scent rolling, a practice that enables the wolf to transport odors wherever he goes.

It is fairly common to observe a scapegoat animal in a captive pack, typically one that has fought for or once held the alpha or beta position. If he was once dominant and abused other animals from that position, he will likely be abused in turn. If he was benevolent as an alpha animal, he will be treated kindly. Interestingly, this “omega” animal is often relegated to the area in the wolf enclosure which is directly adjacent to the area outside the pen with the greatest human traffic. In the wild an omega animal might trail the pack at a distance, feeding on leftovers, occasionally trying to join them. He might even be permitted to do so briefly—to repel an outsider from another pack, for example—and eventually he might be reintegrated.

In one captive situation I observed, the outcast, a former beta male, still enjoyed the alpha male’s protection. One day the alpha male broke up a silent, serious fight between the outcast and the rest of the pack—nine animals—that left the outcast bloody and stiff with wounds for two days. One of the curious elements in this case was that the attacks were triggered by the outcast’s attempts to defecate outside its assigned area. The animal seemed to be suffering from constipation and nipped at its hindquarters as though bothered by worms. It occurred to me that the ousting could have been related to this intestinal problem. It would be interesting to know whether wild wolves banished by packs in the wild suffer from infections or diseases that threaten the health of the pack.

The reader may have been tempted by now to look at wolf behavior in terms of the sorts of things domestic dogs do. It should be stressed here that this is not wise. Dogs suffer from a wide variety of emotional disorders, many of them brought on by the sort of selective breeding that destroys or radically alters their systems of communication. Tail docking (boxers), excessive facial hair (sheep dogs), ear cropping (Doberman pinschers), pendulous ears (bloodhounds), and uniform coloration (Weimaraners) all have forced dogs to seek other means of communication. Often the behavior that we see in them—scent-marking a fire hydrant, for example—is an example of frustrated communication.