Has dispersal promoted individualism in wolves and has individuality, paradoxically, evolved due to pronounced sociality?

While watching the International Wolf Center livestream videos, I cannot help but wonder at the extraordinary ability by these individuals to keep track of highly dynamic (on physical and social level) interactions (and speedy, as well).

Also, these wolves seem to be capable of minding what everyone else is doing even if each individual acts in their own capacity and their behaviour is not synchronized in a directly cooperative manner although I suspect that it is coordinated even in situations where it coordination of intent and purpose do not come across as obvious, e.g., individuals who have ‘teamed up’ are not side by side, are not performing the same action, do not exchange conspicuous cues etc.).

At the same time, wolves are also keeping track of their social role and adjusting its expressions to the unfolding events.

Additionally, they seem to be following their individual purpose (in touch with their own physical and psychological responses).

This latter mode is often hard to distinguish from their social role because I believe that, frequently, when individuals appear to be acting out of their own will (e.g., they seem grumpy), it is, in truth, their social role they are enacting and if it were left to their own devices, they would be acting differently.

However, I am convinced that wolves are aware of themselves as individuals and as individualities (because they can overcome their individual inclinations to perform social behaviours instead which requests for being aware of themselves as having traits, needs, desires which are separate from their social functions).

This ability of separating social identity from personal identity manifests in their strategies where they imbue the ‘etiquette’ behaviours with their unique approach.

As a result, while several wolves in the group can act ‘in a subordinate manner’, mostly it is easy to distinguish which individual it is (from the manner in which they act rather than from their fur markings, colour etc.) despite the action being rather similar.

Wolves do not forget who they are even when they behaving as social entities, and it manifests.

Most long-time wolf watchers of specific packs will recognize their favourite wolves even if these individuals do not accomplish anything that other individuals would not accomplish – because they have their own signature mark to everything that they do which can allow to identify the individual during social interactions even if the external cues (colour, markings, size) are not clearly seen at the time.

Wolves can deny their individual will for the sake of their social role but they also make choices despite the prevailing social mood.

For example, while the majority is engaged in highly enthused play, some individuals (especially, the older dominants but not exclusively so) rest or eat.

I think that for a social species, it is difficult to restrain from going with the general mood because they are hormonally, behaviourally and psychologically set to interact with the group and to comply with the group’s will.

Still, these occasions (where the individual does not play but, e.g., rests) are not easy to classify as purely individualist vs. social decisions.

In species that forage cooperatively or engage in other cooperative activities upon which the group’s survival depends and which are physically demanding, individuality that is rooted in one’s ability to track one’s own physiological states and to respond to them accordingly – is not purely… individual.

In such species, it is a social obligation to keep fit which is especially true for group leaders (the breeding pair but also other group members who are key providers, defenders etc.).

Thereby, the ‘individual’ decision to rest rather than to ‘go with the group’s mood’ might not be as individual as it seems.

It might be a social decision based not on what the individual truly wants but on what the group expects of the individual.

Nevertheless, such decisions imply following one’s own internal states and they give rise to individuality.

Thus, when the older wolves in the IWC pack rest or otherwise refrain from participating in the gambols of the youth, they might be following ancient social duties and their personal choices (if they were based solely on their current and individual psychological and hormonal disposition) would be quite different.

The frolicking exuberance by the young wolves might not be as individually selected as one might surmise, either.

While they probably have more energy to spare and the group does not rely on them as vitally, perhaps it is their task to keep the group spirit and to give everyone a boost during the group’s resting periods (they are the post-game or pre-game cheerleaders).

I do not believe that they would prefer not to play but their decision to play might have twofold motivations – individual as well as social.

The requirement of establishing one’s own physical identity and maintaining it (with respect to social expectations) might be a feature of cooperative species.

As wolves are cooperatively to the point that they do almost everything together, they might have to mind their responsibility of being also individuals.

I suppose that in species which are social but which are not highly cooperative (e.g., ungulates) and where the loss of one member is tragic but it cannot impair the functioning of the whole group, individuality (in physical sense), paradoxically, might be less developed.

This could be the reason why, for example, many social deer species (e.g., elk or red deer) fail to recognize cues that their fitness is under threat while social carnivores (who hunt cooperatively) are better at maintaining balance with their environmental carrying capacity.

Non-cooperative social species might not be as aware of their individual physical condition and they might base their self-assessment (health- and success-wise) on the social group dynamics.

For example, wapiti might reckon they are doing well (under circumstances where they have severely depleted their resources, overpopulated their range and approaching a population crash) because – up to the point where they are ravaged by starvation or disease – there are no social cues that would inform them on the danger of their behaviour.

If they base their fitness evaluation not on how they feel physically but rather on social dominance, group size, group dynamics, it is possible that large herds are considered a success (apparently, there are enough resources to form large groups) and as the females start giving birth to smaller-sized offspring (due to poorer diet), the population might become more equal in terms of dominance (lack of clearly larger and clearly smaller individuals whereby the smaller individuals can no longer identify themselves as socially inferior and even poorly-faring individuals might be considered average failing to provide a cue that the group is not doing well).

Large groups could, coincidentally, awaken greater tolerance.

At least many rodents who are also herbivores become less territorial during their population peaks (high population density) because it no longer makes sense to defeat resources and one can preserve energy by tolerating others.

Social ungulates are not strictly territorial but the large groups might bear the same impacts of reduced dominance – an altered state which is augmented by the decreasing quality of resources (why defend a specific foraging spot when it is no better than the rest?).

As a result, such species might not be able to properly assess their individual fitness because they base their fitness identity on social cues (which are not individualized) and the social cues are not effective.

Returning to the individuality of the wolves…

I do not think that the social requirement to keep oneself rested and healthy is the only mechanism through which wolves have become social-individualists.

I also believe that dispersal is a contributing factor because the ability to perform a successful dispersal journey is crucial in the wolf’s reproductive future but dispersal is frequently an undertaking one must assume one one’s own – making individual decisions, being self-reliant etc.

As wolves mostly establish new families after dispersal, the dispersal itself might act as a selective pressure, i.e., only those wolves survive and get to breed who have been capable of making individual choices during dispersal, providing for themselves while on their own, being self-sufficient.

My suspicion is that dispersal used to be longer, historically.

Nowadays, it is typically cut short by human-caused mortality (traffic, poaching, conflicts with livestock etc.).

In days when there was more of wilderness and less of civilization, individual wolves might have persisted in the state of dispersal for years before they have found a chance to establish a family and to reproduce (as they still do in some cases, e.g., the famous wolf OR7 and his brother OR3).

This ability to persist during prolonged dispersal would have been very important because wolf populations must have been denser, as well, and there would not have been many areas that were void of wolves but that were suitable for wolf residency.

Thus, in order to find a chance, wolves might have had to wait for longer periods until some vacancies opened in the local populations.

These ‘lonesome stages’ (during which the wolf, of course, can temporarily join packs or form alliances with other individuals) must have contributed to the evolution of wolf’s individualistic capabilities, and nature might have favoured wolves who could be individualistic in order to, once more, gain the opportunity to be social.

However, it is curious how wolves develop their self-reliance skills because I do not believe that wolf families specifically invest effort in isolating their packmates and forcing them to learn ‘their own ways’.

Partly, some individualism would be developed already in the phase of yearlings when, during the pup-raising period, wolf packs tend to split up in smaller hunting groups or they hunt individually.

Individual decision-making also shows up during cooperative activities because wolf hunting strategies are not based on some type of group behaviour akin to ‘synchronized swimming’.

Faster wolves have different tasks while subduing prey than larger, slower wolves (with greater jaw-strength) have.

Similarly, the group sometimes fans out exploring the area individually but at all times keeping awareness of the location of others to inform others if prey has been found.

Sometimes wolves split into task forces whereby some individuals flush the prey into a specific direction while others wait in ambush.

These strategies rely on individualized skills and individual decision making embedded within the social group’s objectives and ‘meta-activity’.

However, such activities do not necessarily prepare the individuals for a long (perhaps years-long) journeys on their own because, during these activities, the individual always has the chance of coming back and keeping in touch with one’s family through howling.

I believe that scent-marking is also a strong binding force and that individuals who travel on their own in territories marked by their families, do not develop the mindset of being truly alone.

During my efforts to reconstruct wolf pack recolonization histories in NE Oregon (they can be found under Articles > Wolf Pack Histories), I noticed that quite often wolf pack territory size increased (in this population where it was still possible because adjacent ranges were not inhabited by neighbouring packs) and sometimes also livestock conflict grew more pronounced during years when there was a large number of two-year-olds in the pack (livestock conflict risk is greater for lone wolves or wolves travelling individually than for packs/groups).

This made me arrive at a conclusion that a young wolf’s education was comprised of at least three initial stages:

  • learning basic motor skills, hunting skills, tracking skills, territory navigation skills etc. and learning to be a part of the group (pup);
  • learning basic care-taking skills and learning to be an efficient, actively participating, dependable part of the group beyond self-maintenance (yearling) while also acquiring some skills as individual hunters and decision-makers (during summer foraging which is not always done in groups);
  • developing a social and environmental interest beyond one’s group and one’s home; beginning to venture out on forays in order to explore areas and wolf societies outside of natal pack’s range.

The third phase is sometimes explained simply by the young individual’s growing interest in mating opportunities or perhaps social pressure in the pack.

However, my interpretation is that these older individuals who have been fortunate to acquire their fundamental education (sort of high school education) at home, now feel inclined to pursue the next stage in order to compare their experience at home with the environmental (resources, prey, terrain) and social qualities found outside of their home.

I believe that this is largely a social drive whereby the individual has grasped the essential dynamics of their own family group and they have grown curious about ‘those other wolves’ whom they might have already encountered as neighbours or trespassers or who must be out there (in recently recolonized populations where pack ranges do not overlap and immigration is relatively rare).

I believe that these forays are active studying experiences.

This is when the individual breaks touch with the group in order to pursue their interests independently and the individual also leaves the group territory.

Outside of the group’s range, there is little scent evidence of their own family (if any) and howling is dangerous.

Also, there is always the possibility that the individual will not be able to return (either due to intraspecific strife or because they might find a mate, accidentally) although some individuals return to their natal packs after prolonged periods of absence (even years) which might suggest – all through this time they have retained the natal group’s location in their memory for a reason, and this reason is the possibility to go back which must be ingrained in the wolf’s make-up suggesting that perhaps, earlier in wolf evolution, individuals did not necessarily disperse ‘irrevocably’ and they often came back (or else, wolves simply have far too excellent spatial memory to lose this knowledge).

Thereby, these must be the first ventures that are driven by individual motivations and that bargain with one’s social identity.

As I have been watching the interactions among the captive wolves at ICW, I refuse to believe that these young wolves embark on forays because they have grown bored of their family and its home.

It seems that wolves can invent millions and billions of ways how to perk up their social dynamics and they love their home.

Then what is it that propels these wolves out of their homes and into the unknown (or vaguely apprehended)?

As I mentioned before, I think that it is social curiosity.

Wolves, for the large part, seem transfixed on other wolves.

In fact, at least in Oregon, new packs failed to establish for years while the population was still in its sparse, initial phases of development (despite the fact that migrating wolves were spotted frequently and they must have had opportunities to pair up).

Meanwhile, once the population was denser, almost all at once, there was a boom of new packs establishing beside the founders even if they did not have the most optimal ranges.

Should a wolf not be drawn to a more abundant range while it is still available?

Instead these wolves seemed drawn to other wolves (other wolf packs) even if it meant the ranges were not of highest quality.

Many dispersing wolves did not leave the natal population and tried to hang out near its edges seeking for a chance to start a family.

Meanwhile, it would have been more profitable to strike out on their own and to leave the natal population because its edge habitats were already less than suboptimal.

I believe that besides the terms ‘natal pack’ and ‘natal territory’, two other terms should be introduced – ‘natal population’ and ‘natal region’ which are perhaps considered by wolves as extensions to their parent-based society and home.

Perhaps there is some desire felt by the individual to understand who they are and who their family is based on comparing themselves to those who are not kin, who lead different lives, who possess different traits.

Maybe the young wolves who depart on these forays want to come back and to look at their family ‘with new eyes’ applying their comparison-based insight creatively.

Perhaps this is achieved because the new cohort of yearlings have already taken the role of performing essential chores and the older subordinates feel they must carve out a more… individualistic role for themselves if they want to remain in the family as needed, efficient, productive members.

They have to offer what yearlings cannot offer not because they must compete with their younger siblings but rather because the yearlings have to be given their chance to study what befits their developmental stage (forming ‘yearling niche’ which the two-year-olds must separate from in order not to interfere with the education by their younger siblings).

I also wonder if wolves remember that they once had older siblings who left the pack.

Thus, they would know that somewhere there are related individuals, related packs which might be perceived as their own family and their own home.

What if these two-year-olds venture out to find the family and the home that lies outside of their immediate family and home but which is not a new family and a new home in the sense of having been formed with an unrelated mate?

This claim might be supported by a pattern I observed in Oregon packs where some wolves tended to disperse in the same direction as their older siblings (who had dispersed before them).

Dispersal direction can be affected by many factors, e.g., social factors, topographic factors even perhaps genetic factors.

Also, even if these individuals are following in the literal footsteps by their older siblings not because they have been ‘coerced to’ but because they want to track their siblings’ path, the reason might be that these wolves are cautious on their own and they find comfort in knowing that they sibling had trodden that way (and perhaps made wise choices by doing so).

However, there was another supportive tendency that I noticed in Oregon’s packs, namely, that wolves in newer packs where no older siblings had yet dispersed, tended to stay for longer (dispersing as two-year-olds or even as three-year-olds).

What if these individuals were not as prompted to leave because they yet had no siblings to find?

Perhaps this sense of kinship has co-evolved with hormonal changes because the inclination to find a mate entitles the search for an unrelated individual which implies differentiation between kin and non-kin (which could be a new revelation sparking interest also in the nearby non-related packs which are, for the first time, consciously apprehended as different in a recently highly relevant manner that is closely connected to one’s identity).

Reproductively immature wolves might not have this curiosity about kin vs. non-kin and they might not yet have come by a physiological and psychological awareness that some wolves are relatives and some are not (and that this affects them very urgently and directly) as well as by an understanding that their genetic family is, in fact, extended outside of the immediate group.

Upon maturation, not only a desire to find an unrelated mate might emerge but also a more conscious longing for the related individuals who are no longer in the group could appear (because both of these types of social relations are discovered outside of the group and the awareness of there being an unrelated individual who must be found might arise alongside a realization that there are related individuals who could be found).

Interestingly, the apprehension that unrelated individuals can turn into related individuals (and, thereby, into members of one’s social group and one’s social identity) could transform the wolf’s outlook toward neighbours and immigrants offering a gradient that surpasses ‘my group / not my group’ bimodality differentiating the array of possible encounters into more subtle categories of social potential.

I find it entertaining and paradoxical (from human viewpoint) that wolves might have evolved individuality in order to be better, more useful, more deeply ingrained members of their social group and that individuality in wolves is the result of extreme sociality.

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