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Authors: John Bradshaw

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Kin selection explains that, when the younger members of the pack appear to deliberately put their own breeding rights to one side, they are actually acting in their own interest—but the advantages of this behavior are not purely familial. In addition to the advantage they gain from kin selection, it is also safer for the wolves themselves not to leave their pack while they are still young. Their lack of experience means that their chances of forming their own pack are actually rather slim. This accounts for the rare occasions when unrelated wolves have been recorded as joining existing packs; it appears that in such cases they are being recruited as a replacement when one of the most experienced members of the pack, maybe one of the original founders, leaves or dies.

Packs that form naturally, in the wild, are usually harmonious entities, with aggression being the exception rather than the norm. As in any family, there are occasional conflicts of interest within wolf packs,
but in general the parents have to do very little to keep their grown-up young in order. The young are essentially volunteers—they could leave the family and set up their own, but they choose not to, preferring to stay safely within the family unit until they are older and more experienced, and hence more likely to survive the risks of finding a mate and a new place to live. They regularly reinforce their bond with their parents and, at the same time, reassure them that they are helpers, not rivals, by performing a special ritual. The youngster crouches slightly as it approaches the parent, ears back and close to its head, and tail held low and wagging. It then nuzzles the side of the parent's face, an imitation of the food-soliciting behavior that it used when it was a cub. (This is very similar to the greeting ritual of the African wild dog, and so possibly a very ancient canid behavior, predating the evolution of both wolf and wild dog.)

The image of a harmonious pack is not the picture of wolf society that you will find in most books on dog behavior. Wolf biologists originally based most of their ideas on captive packs, which were easy to observe. Some of these packs were random assemblies of unrelated individuals, while others were fragments of packs, usually with one or both of the parents missing—basically composed of whatever individuals were available for the zoo to make an exhibit. What almost all of these packs had in common was that their structure had been irrevocably disrupted by captivity, so that the wolves were thrown into a state of confusion and conflict. Moreover, unless their human captors decided to separate them, none had the opportunity to leave. As a result, the relationships that emerged were based not on long-established trust but on rivalry and aggression.

The true picture of wolf society emerged only as wolves became protected, allowing packs to form and thrive over several years, without becoming fragmented by continual persecution. At roughly the same time, better technology for tracking and observing wolves in the wild became available: GPS, miniature radio transmitters with batteries robust enough to allow tracking over a whole season, and so on. Within a decade, descriptions of wolf society had changed from the image of the hierarchical pack run by two tyrants, one male, one female, to that of the harmonious family group, where, barring accidents, the younger
adults in the family voluntarily assisted their parents in raising their younger brothers and sisters. Coercion was replaced by cooperation as the underlying principle.

This radical change in our conception of pack behavior has required that we also reappraise the social signals that wolves use. Under zoo conditions, signals that wolf parents would normally use to remind their offspring to cooperate instead became the precursors of out-and-out fighting and were labeled “dominance indicators.” Similarly, the cohesive behaviors that adult young wolves would normally use to bond with their parents were now being used in desperate attempts to avoid conflict and so came to be labeled as “submission.”

Contrary to long-standing theories of wolf behavior, it is now believed that “submissive” behavior may be nothing of the sort. An effective “submissive” display should, by definition, indicate to an attacker that the attack is not worth pursuing—and, indeed, when wolves from two different packs happen to meet, the smaller one will try to avoid being attacked by performing such a display. This rarely works, however, and if the smaller wolf fails to run away, it will be attacked and often killed by the larger. Wolves from different packs have no common interests; they compete for food and are probably only very distantly related, if at all. Nevertheless, if the “submissive” display was truly an indication of submission, it ought to work in these circumstances, since the attacking wolf is putting itself at risk of injury, even if it wins. The fact that this display doesn't work under these circumstances indicates that it isn't a “submissive” display at all. Moreover, when it's performed between members of the same family, for the most part it is
not
preceded by any form of threat from the recipient. Rather, it usually appears spontaneously, reinforcing the bond between the members of the pack. Only in artificially constituted “packs,” kept in zoos, do “submissive” displays come to be a standard response to threat. Presumably the younger, weaker wolves learn by trial and error that such displays (sometimes) work under these unnatural circumstances, where pack loyalties have been totally disrupted and there is nowhere for them to escape to.

Wolves perform two signals that used to be labeled “submissive”: “active” and “passive.” Domestic dogs perform very similar signals, and these, too, are referred to as “active submission” and “passive submission.” One
might expect that any reinterpretation of these signals in the wolf would quickly have been followed by a reappraisal of what they mean when performed by dogs, but this has been slow to happen.

The “active” display is the more common one among wolves and, rather than being a sign of submission, is in fact a bonding signal that scientists now refer to—much more appropriately—as the affiliation display. In the affiliation display, the wolf approaches with a low posture, holding its tail low; its ears are pulled slightly back, and its tail and hindquarters wag enthusiastically. This display forms part of what's called the “group ceremony,” which occurs when the pack reassembles or as a precursor to a hunting trip. Under these circumstances it can be performed by the parents (the “alphas”) as well as by their offspring, confirming its role as a mechanism whereby affectionate bonds are reinforced. It's difficult to figure out how this display was ever labeled as “submissive” behavior. A wolf performing the affiliation display is actually in a rather good position to attack its recipient—a swift twist of the head and it could sink its teeth into the other's throat. Therefore, accepting the performance of the affiliation display is, if anything, more an expression of trust on the recipient's part than on the performer's. It's undeniable that the younger members of the family perform the affiliation display toward their parents much more frequently than vice versa, but this behavior is typical of all parent-offspring relationships and does not mean that the offspring are allowing themselves to be “dominated” by their parents. Indeed, it simply reflects the asymmetry of the relationship between parents and offspring. The parents are the only parents that each of the young wolves is likely ever to have, and therefore their attachment is total. The parents may—indeed, probably do—have other offspring, so their attachment to each cub must unavoidably be a shared one.

Also in urgent need of reinterpretation is the wolf's other, less common submissive display, “passive submission.” Unlike the affiliation display, this may be an actual sign of submission, one that is derived from an infantile behavior in which cubs roll over to allow their mother to groom their belly and stimulate urination, which the cubs cannot yet do on their own. This display, which seems to have been adopted by adults as a way of deflecting possible attack, involves one wolf lying down, rolling onto its back, and exposing its abdomen for inspection by another. Some wolf biologists now refer to it as the “belly-up display,” which is more descriptive and presumes nothing about its function. Nevertheless, this display has precisely the characteristics one would expect of submission, since the wolf that performs it is placing itself in a position where it is at the mercy of the other.

A wolf (left) performing the affiliation display

The belly-up display

In wolves, the belly up-display is much rarer than the affiliation display and is more commonly seen in captive wolves than in the wild. When observed in zoos, it is most likely to be performed by wolves that are on the fringes of the captive “packs,” are often involved in fights, and rarely participate in group-howls. These are the very wolves that would almost certainly have gone off on their own if the fence surrounding their enclosure hadn't been there.
5
Continually stressed by being forced to remain in close quarters with other wolves that threaten to attack them at every turn, these outcasts will try any tactic that might deflect aggression. In cases where the affiliation display fails, behaving like a helpless cub evidently works, so they learn to use it when they are in desperation. Thus in wolves this display may be an artifact of captivity—not a normal part of adult behavior at all but, rather, a signal artificially carried over from infancy into adulthood, under unnatural circumstances.

Studies of wild wolf packs have made clear that the traditional interpretations of wolf submission—both aggression within the pack and the “submissive” behavior that is an attempt to defuse or deflect this aggression—reflect artifacts of captivity and therefore cannot be applied to wolves as a species. Wolf packs that have not been manipulated by man and are allowed to manage their own affairs, so to speak, are generally peaceful. This is not to say that all of what has been written about aggression between wolves is wrong; for instance, it is an undeniable fact that wolves—captive or not—can be very forceful and aggressive when they want to be. In the wild, even though relationships within packs are usually congenial, aggression toward outsiders, though infrequent, is unrestrained and potentially fatal. In captivity, however, pack “identity” is either nonexistent or severely disrupted, resulting in the expression of behaviors that would normally be seen only in skirmishes between members of different packs.

Observations of captive wolf packs have led not only to mistaken assessments of wolf behavior but also to fundamental misunderstandings about the structure of wolf families themselves—misunderstandings that have warped the popular conception of dogs as well. In captive wolf packs, the breeding pair are conventionally referred to as the “alpha male” and the “alpha female.” Many dog trainers, borrowing from this
conception, insist that owners must impress their own “alpha” status on their dogs, who would otherwise be driven to seek “alpha” status for themselves. Thanks to our new understanding of the way that wild wolves construct their packs, however, it has become clear that “alpha” status comes automatically with being a parent. The term “alpha,” as applied to a parent wolf in a normal pack, thus doesn't describe much about the wolf's status beyond its role as a parent.
6
It is meaningful only when used to describe the eventual victor of the warfare that is endemic to captive groups of wolves, which lack the family ties that would ensure peace in a natural pack. Which of these two models points to the most appropriate way to understand pet dogs and their relationships with their owners? Is it the “alpha” model, based on the unnatural captive pack, or the “family” model, based on the behavior of wolves that have been allowed to make their own choices as to who to live with and who not to? The family model is the product of millions of years of evolution, allowing the formation and refinement of an elaborate set of signals that serve to keep the peace. The alpha model emerges only in artificial social groupings that evolution has never had the opportunity to act on, and in which individual wolves have to draw upon every ounce of their intelligence and adaptability just in order to survive the relentless social tensions inherent in such groupings.

BOOK: Dog Sense
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