To begin with, I would like to issue a statement that this will be a highly speculative post but as a non-scientist I can probably afford some flights of fancy.
When I began reading about wolf sociality, I learned the basic facts – that wolves mostly form social units that are based on the breeding pair and their offspring of the current year and possibly also older offspring.
This family unit shares a large territory which is rather fixed although it can be reduced or extended (mainly through social mechanisms).
Their use of their entire range is limited during summer when the pack’s activities are restricted and concentrated around the den (and later rendevouz site) where the comparatively immobile pups are raised.
Once the pups are big enough to follow the adults, the pack resumes roaming on their entire range and the winter range is thereby much larger than the summer range and wolves do not use permanent homesites during winter but rather locate themselves respective to the prey distribution, last kills and territory boundary patrolling efforts.
There can be exceptions to this model but usually the pattern is rather consistent in temperate/boreal regions.
However, as I began reading about wolf populations in different biogeographic areas, I learned that wolves, for example, in arctic regions (but also boreal and boreal/temperate Canada) do not necessarily follow this social pattern.
Where prey is migratory, packs tend to follow the prey and their territories (or rather ranges) become vast extending over 1000 km2.
They can also overlap other packs’ ranges and some wolf populations (e.g., wolves in Algonquin Park) hold territories during summer which are then vacated during winter when the packs follow their prey and are variably territorial around the winter yards of their prey (many packs not being territorial at all but usually avoiding other packs and tolerating solitary wolves).
Caribou-focused wolves also follow the herds to the wintering and calving grounds although most frequently it appears that this spring/summer tracking is not as strict (wolves do not locate themselves in the closest possible proximity to calving grounds and rather stay, e.g., in the proximity of forests) because the den use tends to be traditional (wolves use the same dens over the years irrespective of how far or close to the den the caribou route lies in that particular summer).
It appears (from reports and publications that I have read) that in the more northerly regions (Manitoba, Alaska) the wolf population social structure is more flexible with overlapping and vast ranges, some packs that are sedentary (because their prey does not migrate, either, or because there are several prey species available some of which do not migrate) and some packs that are migratory as well as with pack territory boundary fluctuations that are not necessarily the result of social strife between packs.
The pack territories in the north appear to be in a greater state of flux relative to pack territories in the more southern regions.
For example, the Middle Fork Pack in the State of Oregon, USA had two territories in 2018 (winter and summer territory) which were separated by the territory of the South Snake Pack.
The Middle Fork Pack appeared to use higher elevation range during summer and lower elevation during winter which corresponds to typical prey species use of foraging grounds in the two seasons.
Apparently, in 2019 already this situation had changed and Middle Fork Pack had united its two ranges possibly due to South Snake Pack pressure (but the Middle Fork Pack also bred successfully over the years and could have been able to expand their range because their territory in 2019 seemed somewhat larger than in 2018).
I find it worth mentioning that the NE Oregon wolf population might have genetic lineages extending into Canada (through Idaho and Washington) where a greater number of migratory wolves might be found.
Also, I was curious to read about the events unfolding in the Voyageurs National Park (see the Twitter post by Voyageurs Wolf Project) which involve a very odd divorce by a long-time successful breeding pair and the subsequent ranging of the former breeding male on the neighbouring territory with a neighbouring subordinate.
In this case, two packs appear to be using the same territory.
I have written two posts about similar issue: Has prevalence of polygamy and promiscuity in wolves changed due to intense persecution by humans? and How did the Beringian wolves take over?
In this post, I would like to speculate (as the scientists sometimes say in a derisive tone) about the historical forms of wolf social life.
For many of the claims here I do not really have any evidence to support them.
But I will share my musings anyway.
I believe that wolves first evolved sociality (living in groups) and only then they evolved territoriality (defending set territories).
I think that territoriality probably arose from the necessity by wolves to use den sites during pup rearing season and the inability of pups to follow adults who potentially followed migratory prey.
The location of activity around den sites could have offered an insight into territorial living but it is possible that the early wolves were not territorial during the entire year and that, initially, they protected their pups and the restricted summer resources (which tend to be scarcer than winter resources, especially, if the prey species are located far from the traditional denning site) rather than protecting ‘a home’ in the area sense.
The traditional character of den sites might have arise during periods of harsh climate when permafrost reduced den site availability.
It is not entirely ludicrous to assume that traditional denning (reuse of den sites and denning areas) was also a later system which replaced denning close to calving grounds (in randomly chosen locations that were variable from year to year).
In order to understand this, it would be important to know whether wolves have always fixated on specific ungulate populations (e.g., herds) or if they used to roam over wide areas in order to meet any herd of ungulates or any solitary ungulate (if their original lifestyle was that of a pack in the state of constant ‘dispersal’ which is not restricted to traditional denning areas, nor it is restricted to a specific ungulate population).
On such occasion, wolves may have followed particular herds only during periods of high energy demands (reproduction) or during periods of low prey densities / high wolf densities (when a stricter and more predictable resource-keeping would have been more beneficial).
This might be elucidated if I knew more about Pleistocene faunal assemblages (the ungulate species available to wolves and their behaviour).
Be as it may, during denning wolves perhaps discovered benefits to restricted use of space and in areas where ungulates were not migratory but sedentary, wolves, too, could settle on a fixed all-year-round range.
However, I do not find it likely that is was some convenience derived from hunting efficiency that made wolves decide – this is the lifestyle for us (where possible).
Denning is always restrictive and it can bear fatal toll on the pack because the pack has to depend on limited resources and, in migratory prey systems, the fundamental resource is not always that readily accessible.
Where wolves den far from migratory routes and calving grounds, pup survival is greatly reduced (Frame, P.F. et al., 2008).
Nomadic lifestyle is more supportive of wolf fitness than territorial lifestyle which also leads to intraspecific strife, a leading cause of death in many non-hunted wolf populations.
Migratory wolves, as mentioned before, as less territorial at least during the winter which could promote survival due to lower aggression levels between social groups.
The way I see it – territorial lifestyle is not truly something that might be selected for in order to make life easier with regard to food availability and social tension on population level.
I believe that wolves began enjoying the social interactions around the den site and the playing with the pups and with one another.
Far-ranging wolves have to travel astounding distances on daily basis and it is possible that migration demands physical effort that does not leave much energy for social play and bonding.
I suppose there are other (hormonal and more practical) reasons involved in wolf inclination to reduce their ranges (where possible) and I hope to someday to be able to figure them out but currently I will simply state that I think wolves chose territoriality to promote social bonding.
I also believe that wolves attempted to resolve the constraints by food limitations by forming large kin groups that did not necessarily merge entirely.
In many wolf families it can still be observed that two females den together (or within one family unit) and I think this evolved not out of polygamous tendencies but rather out of ancient pack units of related family groups (e.g., a mother and a daughter and their respective mates) that denned separately (but not always) and later when the pups could leave the dens and gather in larger sites (rendevouz sites), the family units sometimes came together so that the pup defense and education could be carried out by some individuals of these united groups while others went on farther foraging forays.
Later in the winter, the groups might have travelled together initially forming the extremely large packs that have been observed in Alaska, US and Canada by wolf biologists (e.g., packs in the Wood Buffalo National Park etc.).
These groups are sometimes larger than necessary (according to the number of individual wolves needed to take down a prey species that is of a lesser size than a muskox or a bison).
This could have allowed for some individuals in the pack to sometimes stay back and to engage in social bonding and to enjoy a sense of a temporary homesite.
Where prey was more sedentary or there were several types of prey available, these large packs could have begun forming year-round territories with related family groups living side by side and not separating themselves in the strictest manner but also holding on each to their own den sites and core ranges.
I believe this is still reflected in what we deem ‘confusing’ wolf behaviour.
For example, I mentioned the curious Voyageurs wolf case where an established successful pair divorced suddenly.
I would explain it in a manner that would probably not be supported by wolf biologists.
I believe that in the early states of wolf sociality, related wolf families (clans, tribes) lived together and shared ‘problems’.
When someone had to leave temporarily, for some reason, there could be relatives who replaced this individual until the individual returned.
There were also social structures in place to prevent unwanted group living effects (such as breeding with the ‘wrong’ mate).
I think that what happened in the Voyageurs was that after the demise of the neighbouring Nashata Pack’s breeding male, the young Cranberry Bay wolves (subordinates) sought to pair up with the Nashata widowed female.
The original Cranberry pair might have favoured this because it would follow the ancient wolf tradition (that currently perhaps only exists in my mind) of having several related packs living adjacent one to another.
I observe that there is a tendency in many wolf packs to establish a territory and then to produce pups and then to retain subordinates and to expand territory.
I think that this tendency would be followed by growing the pack to a number which can then split in order to have two packs that, once again, begin the formation or larger and larger numbers.
Behind this mechanism, I see a strategy to form family groups not in the limited sense of a breeding pair and their offspring but of several related breeding pairs and their offspring (something between a nuclear family and a tribe).
I think that the breeding male of the Cranberry Bay Pack (the supposed divorcee) followed his offspring ‘to woo the Nashata female’ (however, the sex of the two Cranberry Bay subordinates who were seen with the Nashata breeding female and also later roamed with their father, has not been stated).
I believe that this was almost working out and the Nashata Pack began restructurizing (as suggested by the separation of a yearling Nashata female who was later found with the Cranberry Bay former breeding male and his subordinates).
But there came another prospective suitor who, for some reason, outcompeted the Cranberry Bay wolves.
Meanwhile, the Cranberry Bay Pack breeding female had stayed back with an unrelated adult male during the breeding season and, with her mate ‘managing suddenly very confusing business’ in the neighbouring territory and without anyone to prevent breeding on social basis, she probably bred with the adult male because I believe that wolves do not always follow their own feelings and preferences but rather they have to keep their genes going and to have pups not to lose the home that they have made for themselves.
The Cranberry Bay subordinates who had travelled with their father, dispersed and did not mate with the Nashata yearling possibly because they did not actually have an adequate territory and because the young female was not yet fully mature.
However, the young female remained with the Cranberry Bay former breeding male and I suppose he felt responsible over her and also he had lost his own home and mate, and therefore they stayed together roaming on the female’s mother range.
I believe it was one big misunderstanding and some ill timing.
But I also think that it is reflective of what would be natural for wolves and that is:
- Engaging in matters to establish neighbouring ranges for their offspring in order to create ‘family tribes”;
- Having other adults behind to ensure territory defense and hunting efficiency (normally, these adults would be related or there would be someone imparting breeding restriction);
- Territorial exclusivity can sometimes be overlooked and wolves even attest to some flexibility regarding territories in certain situations (especially, involving the breeding prospects of kin) while, at the same time, being territorial to death in other instances.
The disruption of the original strategy might have led to non-related neighbours, territories that are sometimes too small to support families through seasons, polygamy and divorces which would have, historically, originated in several females raising pups together while each having her own mate as well as in related adults from several family units temporarily performing roles of individuals who have had to take a leave in order to resolve ‘family matters’.
I am aware of the facts countering my theory.
For example, related neighbouring packs are not always tolerant of one another and they do not appear cooperative.
I think that something went wrong along the way.
Frequently, all the faults with nature going astray from its own decided path are attributed to human impacts but I think that multi-unit family groups would have been even more resilient to being hunted (and even to habitat fragmentation) than the current wolf social units.
Habitat fragmentation could have been an issue but the impact is dependent on how vast family areas wolves wished to attain (and several families would have found it easier to cover larger ranges).
Changes in prey populations were likely favourable for territorial living because most ungulates nowadays are probably non-migratory offering a stable prey basis during all seasons on a restricted range (a year-round territory).
However, the reduction in prey size after Pleistocene could have complicated the maintenance of large family units (but the separate units could have decreased in size still retaining the tribal union structure).
I believe something happened that prevented the wolves from acquiring sufficient numbers within one family unit in order to split the pack and then split it again until several neighbouring kin territories were acquired.
Considering what factors could have attributed to promoting dispersal over permanent residency and gradual enlargement of family influence, I wonder about changes in climate after glaciations as well as perhaps genetic inbreeding issues.
The melting of glaciers shrank and finally sumberged the Beringian land bridge causing masses of species to disperse into both directions and this was how the native Eurasian wolves were largely replaced by the Beringian wolves.
I believe that Beringian wolves were partly migratory and smaller than native wolves and thereby they were very successful in taking over (they constituted larger dispersing groups rather than dispersing individuals and they also could have subside on the smaller prey after the megafaunal extinctions).
They were perhaps also better used to transient lifestyle because Beringia appeared to be a migratory corridor mediating the expansion of populations on either continent (Eurasia or North America).
Their genetics could have been better suited to a dynamic situation where they prey, as well, were migrating in and out of Beringia (making it more troublesome to follow a single herd or population).
Perhaps the waves of emigration and swift colonization during Late Pleistocene created a situation in which new groups and individuals pushing forth the line and travelling through any permanent ranges that the earlier colonizers attempted to establish.
The wolves had to begin defending their territories very fiercely if they did not want to be flushed out with the waves of new immigrants from the lands that were now covered with sea.
The prey movements could have become unpredictable, as well, because some mammals went extinct and any herding, traditional migratory behaviour or philopatry could have been disrupted due to the changes in climate, vegetation and faunal assemblages.
It could have become difficult to follow a particular herd or a particular migratory route and wolves might have had to establish territories in which they opportunistically consumed any ungulates that happened to be there at the time while fighting off the dispersants journeying through.
Under such conditions, it may have been impossible to maintain adequate and stable territories to retain subordinates or to ‘plan ahead’ with respect to prey basis on the selected range resulting in loss of ability to form large groups that could split and become neighbours to their parents.
The population density might have been very high, as well, due to the great number of floaters.
Instead subordinate offspring would have joined the waves of dispersants in the manner we observe today.
Hormonal and behavioural changes might have arisen due to the new level of territorial strife.
I believe that social living is more important in most wolves than some sort of conquest and fight and I think that the loss of some forms of social life became replaced by adrenaline-boosted social consolidation during high-stress situations.
Oxytocins which are the ‘social hormones’ are produced not only during play and affiliative interactions but also during hunting and territorial patrolling (Wirobski, G. et al., 2023) and wolves may have become increasingly territorial to compensate for the loss of opportunity to maintain large social groups over vast ranges (because the pleasant feelings associated with large family could be replaced with oxytocins produced during such defense-aimed territorial group activities.
As I mentioned before, some of the new conditions could have also affected the wolves’ ability to establish ‘multi-family settlements’ even where it was still possible.
One of the reasons could have been interbreeding avoidance.
In order to have these tribal groups, there should be certain mechanisms in place ensuring a supply of potential breeders who are available to pair up with the residents when vacancies open.
However, in ‘tribal settings’, residency would have been more permanent because tolerance toward neighbours would be higher and group turnover would be slower.
It would also have been difficult to ‘fight one’s way’ to the breeding status (if several families protected the current positions).
As a result the opening of breeding vacancies could have been more rare.
Thereby, the ‘breeding stock’ should have been comprised of individuals who were prepared to roam very far (travelling through the family residential ranges which could have been vast and passing on to another tribal area if no vacancies were open locally in the former one), who were prepared to roam for long (because opening of vacancies would have not occurred frequently and perhaps these individuals would have spent the entire life as a solitary, lone wolf).
It is probable that, under such systems, lone wolves were rare and they were tolerated better (perhaps accepted in the large, established packs as temporary guests because they hardly posed threat to the extended families and if resources allowed, they could be abided because they held the potential to provide an important demographic rescue).
There would have been essentially two types of wolves:
- Residentially-inclined, philopatric individuals who are very prepared to assume subordinate position in a pack without breeding opportunities for possibly the entire lifetime (however, dispersal might occur through settling in direct proximity of parent group);
- Exploratory, ambitious, bold individuals who left their packs to have adventures, to discover new ranges, to briefly join packs, to provide for themselves while roaming alone and to sometimes die without ever having attained their own family.
If the environmental conditions (as well as population density) changed and packs suddenly became more cramped on ranges with limited resources and without ability to migrate with the herds of ungulates or to split up temporarily in order to hunt outside their territories or to use the resources of adjoining territories communally in times of need, these packs could not tolerate lone wolves who were ‘extra mouths to feed’.
It might have become harder to survive on one’s own, as well, and if humans played a role at all, it might have become dangerous to spend substantial amounts of time on no-wolf land where humans had founded their settlements.
Dispersal mortality could have risen and habitat fragmentation could have restricted safe travels between populations.
Rather than having megafaunal species, the world was now inhabited by a greater number of smaller-sized animals which reduced the size of prey (even lone wolves can take down muskoxen or moose and this would have sustained a solitary individual for a long time while on their own) as well as proliferated the number of mesocarnivores (competitors to the lone wolf, scavengers on his kills).
The old system would have collapsed with the solitary individual mortality on the rise, with movement between populations restricted and with the necessity to provide dispersing individuals on a more frequent basis and more locally.
I believe that the echos of the old times are still reflected in wolves who disperse from their natal packs and who wander very far and long at times apparently challenging themselves ‘for the sake of the challenge’ (e.g., wolves crossing Alps, swimming across the Danube even though other routes were accessible and mates could be found more easily).
The ability by some wolves to ‘track down’ individuals who had dispersed earlier from their natal areas, could be related to initial visitations of distant relatives by the wolves who assumed long-term solitary roaming lifestyle.
Long-term migration is nowadays mostly cut short by mortality (often human-caused) or the overall instability of persecuted or otherwise vulnerable wolf groups who are in constant need of new breeders.
However, some long-term and even lifetime dispersers are still known in contemporary wolf social history.
For example, the two brothers from Oregon OR-3 and OR-7 might represent the genotype of roaming wolves who seem content and self-sufficient while on their own.
OR-7 is the famous wolf who was the first to reach California in about 90 years.
His brother OR-3 (also from Imnaha Pack, NE Oregon) dispersed a few months before his brother in 2011 (possibly at 3 years old, born in 2008).
OR-3 travelled in W and Central Oregon and likely located himself (albeit not permanently) nearby his brother’s pack (when OR-7 had settled down in 2014).
However, OR-3 never returned to an area which had a denser wolf population (NE Oregon) and which he knew well (having been born there).
Moreover, his brother’s pack was the only other pack in the area at the time and therefore, it was unlikely OR-3 would have found an unrelated mate unless another individual arrived from the NE.
Yet OR-3 did not appear to prioritize breeding and he only started a family (with the 3-year-old female OR28 who was 5 years younger than OR-3 and who had dispersed from Mt. Emily Pack which was relatively close to OR-3 natal range) in 2016 (the origins of the Silver Lake Pack).
Unfortunately, it did not work out.
The pair produced pup(s) in 2016 but the breeding female died in October, 2016 (poaching) and OR-3 became a lone wolf again possibly staying in the area for 2 more years and perhaps raising one of the 2016 pups to adulthood (fate of the pup unknown).
In 2019, he would have been 10 years old which is a considerable age for a wolf (although not an entirely extraordinary age for wolves in Oregon at the time and for wolves of his natal family) and he might have died due to natural causes (or he might have not died at all by then because nobody knows what happened to him).
I believe that his behaviour demonstrates rather clearly that this wolf was not as intent on starting a family and that there are wolves (OR-3, OR-7 and others) who might be genetically predisposed to long-term wanderings and who do just fine on their own although they might seek the proximity of relatives at least at times (perhaps to compensate for the lack of opportunity to temporarily integrate within a family unit in a visitor’s capacity).
Historically, such wolves might have even periodically returned to their natal packs (as sometimes occurs nowadays, as well, after months or years of absence).
The tendency to ‘visit the natal pack’ might also be evidenced by a slight inclination (observed in some cases) of an established breeder to expand their territory toward the natal pack (although this could also be suggestive of the strategy to reunite packs and to form adjoined kin units).
I think that OR-3 is not the perfect illustration of what I meant under my claims of gene-dispersing forever-roaming lone types because it seems to me that he preferred the area close to his brother’s pack and that he rather wished to live there.
Perhaps in the old times, he would have been a wolf who finds an area and then woos a subordinate wolf out of some resident pack to follow him to that home he has chosen.
He might demonstrate the inclination to form territories close to kin as well as the self-sufficient genotype who can spend a lifetime roaming and dreaming of possibilities.
Nothing is probably as clear-cut anymore because times have changed and evolution has made characters complex and ambitions combined of even contradictory tendencies.
The friendly roaming type who seeks opportunities might have been represented also by famous Casanova from Yellowstone National Park.
His story is one of promiscuity but perhaps he carries the genes of wolves who joined packs briefly, made friends, roamed on until their chance arrived (if it ever did).
Currently, with the altered social rules, Casanova did not act in such a morally ‘pure’ manner but it seemed that he wished to, first and foremost, make friends and to fill any vacancies in that opened up in the social structures (without assuming breeding position).
As would have been observed under the tribal settlement conditions where the social guard might have been higher preventing breeding events between the guest and the resident females (it is hard to look after the youth when there are fewer adults around and all individuals are constantly busy trying to provide for the pack and to monitor territories – a task which would have been almost non-existent in character in tribal joint-territories).
Under such scenarios, the roaming wolves who act as temporal guests, might have contributed some of the functions that are currently mainly ensured by genetic exchange, more precisely, a novel approach in perceiving the local resources and opportunities.
Thereby, the tribal packs would develop their local traditions and they would adapt to the local environment.
Immigrant breeders, meanwhile, would prevent inbreeding and improve the genetic diversity that is important for population resilience.
However, genetic diversity, in some of its functional aspects, can be replaced by learning from a new individual who has arrived from a different place and who has probably encountered many diverse situations during the migratory process.
Such individuals-educators/innovators (with their long-term travelling and short-term social mingling with several packs) would accumulate knowledge that they can impart with the local wolf societies through education (for example, through participating in the hunt and applying strategies that are novel to the pack traditions).
These are my musings over the wolf social systems of ancient which wolves might have liked to keep.
If I am correct, our current wolf management methods are rather in clash with the objectives of wolves themselves.
We aim to maintain genetic diversity in areas that the wolves might have preferred to occupy with genetically distinct groups.
Dispersal is considered a temporary stage in a wolf’s life and while we encourage dispersal (corridors, connected habitats), we do not really aim to encourage long-term survival and self-provisioning of dispersing individuals.
The dispersal corridors are not established to become optimal habitats.
Also, wolf lethal management often removes many residential individuals leading to a constant need of replacement of breeders (reducing the time that a dispersing individual can spend roaming).
If they do not end up in a new pack soon, they frequently due to poaching, legal hunting, traffic accidents, starvation etc.
Thus, if I were correct in my assumptions, our vision of wolf management should be altered profoundly.
Notes by Ieva 