by Barry Lopez
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.
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.
Three
HUNTING AND TERRITORY
WOLVES DO NOT GET hungry in the way we normally understand hunger. Their feeding habits and digestive systems are adapted to a feast-or-famine existence and to procuring and processing massive amounts of food in a relatively short time. They are more or less always hungry. Wolves commonly go without food for three or four days and then gorge, eating as much as eighteen pounds of meat in one sitting. Then, “meat drunk,” they may lay out in the sun until digestion is completed (in two or three hours), and then start again. A Russian record reports a wolf going without food for seventeen days, and it is commonly recorded that wolves may eat up to one-fifth of their body weight at one time.
The wolf’s diet consists mostly of muscle meat and fatty tissue from various animals. Heart, lung, liver, and other internal organs are eaten. Bones are crushed to get at the marrow, and bone fragments are eaten as well; even hair and skin are sometimes consumed. The only part consistently ignored is the stomach and its contents. Some vegetable matter is taken separately, particularly berries, but Canis lupus does not seem to digest them very well. The red wolf commonly consumes a higher proportion of vegetable matter and subsists on smaller game, like swamp rabbit, and such things as fiddler crabs. All wolves eat grass, possibly to scour the digestive tract and remove worms. Consisting mostly of cellulose, the grass itself is never digested.
Wolves consume an average of five to ten pounds of meat a day and wash it down with large quantities of water to prevent uremic poisoning from the high production of urea associated with a meat diet. The wolf has a large liver and pancreas to aid digestion, and the feces provide an interesting example of efficiency in its large intestine. Droppings in the wild typically consist of chips and slivers of bone neatly packaged along with such items as the rubbery remains of deer hooves in a capsule of hair that moves very smoothly down the colon.
The major sources of meat in the wolf’s diet are deer, moose, elk, musk ox, Dall sheep, Rocky Mountain sheep, caribou, reindeer, or beaver, depending on the area, the season, and the year—a good one or a bad one, say, for moose. Wolves also prey on buffalo (in Wood Buffalo National Park, Canada), snowshoe hares (on Ellesmere Island), flightless ducks (in the James Bay region of Canada), marmots, mice, squirrels, grouse, geese, and rabbits. Wolves fish, too, wade-herding salmon, arctic grayling, or whitefish into shallow pools where they’re trapped. They also mouth-spear them in swift water from the bank with well-timed lunges. They eat carrion and occasionally insects, especially when they encounter them in epidemic populations. And they feed on domestic stock. They hunt by intent but are opportunists, too.
Many studies indicate that wolves prey largely on the aged, the diseased, and the very young. While this is the norm, it is not always true. Wolves do take animals in their prime. And they kill in excess (though excessive kills are often made at the time of denning to ensure a supply of meat while the pack is denbound). In reality, they will likely kill, or try to kill, any animal that presents itself at a disadvantage. The important thing to bear in mind is that generally wolves do not overrun the prey population and kill it out, although they have been known to do so, just as they have, extraordinary as it seems, practiced a kind of fallow-field farming by not killing deer in certain parts of their territories for four or five years, letting the prey population recover there.
Surplus Killing
Wolves sometimes kill all out of proportion to their need for food. That this is also an eerie human trait is suggested by the behavior of buffalo hunters in the nineteenth century. Under peculiar circumstances, having to do with wind, perhaps—no one really knows—a herd of buffalo would remain placid while a hunter dropped animal after animal in its midst. The sight of blood and the bellowing of the wounded did nothing to disturb them. They remained oblivious until something—a gust of wind, the sight of so many carcasses—finally stampeded them. A hunter would commonly shoot until his gun barrel overheated and threatened to explode. The men who made such stands later recalled being absolutely mesmerized by the apparent oblivion, that the moment seemed utterly suspended. When the buffalo finally reacted, the almost mindless impulse to shoot abated immediately and a feeling of remorse overcame them.
In cases where wolf populations are small, or where the prey population is outstripping its food supply, wolves may kill indiscriminately across all age classes.
Just as some people would like to believe that wolves never hurt each other, so some would like to believe that wolves only kill animals that are doomed anyway—the old, the sick, the injured. But the pruning of herds attributed to predators like the wol
f is, at best, crude. The question of which animals will die is also affected by severe winters, range deterioration, and human hunting, among other factors.
In a word, not enough is known.
The idea of wolves on the hunt powerfully engages the human imagination. The wolf spends perhaps one-third of his life in pursuit of food. It is a task for which he evolved and to which he is well suited. With powerful jaw muscles he will clamp down on a moose’s bulbous nose and hold on tenaciously while the moose swings him clear of the ground or stomps on him in a vain effort to throw him off. The wolf can course for miles behind fleeing game, and smell prey a couple of miles off. He has superb hearing and can read tracks as well.
Man admires the wolf’s prowess and indefatigable pursuit, but death itself—blood, gore, and the thought of a wounded animal bellowing in its death throes—makes human beings intensely uncomfortable. There have been people throughout history who would gladly have taken a stand for the preservation of the wolf if only they could have gotten over their own revulsion at the way wolves kill. They frequently could not abide the idea of a wolf killing a creature as “beautiful” as a deer. So the hunting down of prey species warrants careful attention here.
The wolf is a marvelous hunter. Whether we care for the act (or the smelling of anal glands or rolling in decaying meat) is moot—or should be. Wolves kill the largest ungulates by running alongside them, slashing at their hams, ripping at their flanks and abdomen, tearing at the nose and head, harassing the animal until it weakens enough through loss of blood and the severing of muscles to be thrown to the ground. At this point the wolves usually rip open the abdominal cavity and begin eating, sometimes before the animal is dead. If the chase has been a hard one, the wolves may rest before eating anything.
Stories of Teutonically organized raids, where some wolves act as decoys while others wait in ambush, create a sense of hypermilitary tactics, which is misleading. Wolves are intelligent hunters, not marauders. Stories of wolves shredding each other during an attack, like a cauldron of sharks in a bloody frenzy, are spurious. But the conjecture put forth to counteract a claim of stockmen, that wolves never kill beyond their needs, is also erroneous. Wolves do sometimes kill their natural prey in excess of their need for food.
