Turnover in wolf packs – Voyageurs case studies (dispersal)

This post is a sequence to previous posts, Hunting by humans never constitutes compensatory mortality, Turnover in wolf packs – Voyageurs case studies (confirmed mortalities), Turnover in wolf packs – Voyageurs case studies (pup survival).

Reasons for composing the text can accordingly be found in the former materials.

In the post I am going to analyze data from the Greater Voyageurs Ecosystem, Wolf Pack and Population Size Reports 2020/21 (Gable, T. et al., 2021) and 2021/22 (Gable, T. et al., 2022) in order to determine how dispersal (mainly by subordinates, e.g., yearlings) influences the pack future fates.

It is often difficult to disentangle dispersal vs. mortality because non-radiocollared individuals mostly just ‘disappear’ (unless they establish in the neighbouring (study area) packs and are genetically confirmed at a later time as new resident pack members or breeders).

However, I would like to mostly focus on what happens when a family loses its non-breeding, non-pup members (as compared to when it does not).

Most of such losses would be natural concerning individuals that are not dispersing due to environmental constraint, social disruption etc. but that are dispersing in accordance with what is usual in their age group in the study area.

Bluebird Lake Pack

Bluebird Lake Pack (or rather its home territory) was likely founded in fall/winter 2020 by a range-expanding large wolf pack led by the breeding female V052 who was a Sheep Ranch Pack offspring (born in 2015).

The BLP pack named in 2020 and it is not known when exactly it had been formed.

It is possible that V052 dispersed in the autumn-winter of 2016/17 (because she was collared in spring, 2016 while still as a Sheep Ranch Pack wolf) and thus the Bluebird Lake Pack might have formed in 2017 already or during some time between 2017 and 2020.

They were spotted in October 2020 using parts of Wiyapka Lake Pack range but this does not mean they had not been around prior to that (in a more elusive capacity).

The breeding female apparently had not dispersed that far from her natal territory and it is possible that she was even ‘making it back’ because if I am not mistaken (no official maps including the Sheep Ranch Pack exist), Bluebird Lake Pack took over portions of Wiyapka Lake Pack that lie adjacent to where the SRP used to be found although at the time its range was already occupied by the Huron Pack).

(But due to ambiguous phrasing, it is also possible that the BLP had already existed as an un-named neighbour to both Huron Pack and Wiyapka Lake Pack and in 2020, they simply expanded their range, not just showed up in the area for the first time.)

But if there had been a gap between the former Sheep Ranch Pack and the previous BLP location, it is interesting because V052 was probably already the breeding female in her new pack and she was not trying to ‘go back home’ (which had been lost at some point during her absence) in the sense of resuming a subordinate position.

Yet she was compelled to settle closer to her natal range.

This could be related to, e.g., familiarity with resources.

The year 2020 appeared to be rather tough on the wolves in the Voyageurs Ecosystem and the hardships as well as the large pack size that the BLP had attained might have prompted to attain greater security.

Maybe she also was not aware of the pack turnover and relied on tolerance by her former pack members toward the needs of her family.

However, this cannot be known because it is unclear where V052 used to live before 2020 and what her breeding status was.

It appears that she had been the breeding female for at least 2 years prior to appearing on BLP new range because in October, 2020 (first official spotting on portions of Wiyapka Pack range), she was travelling with 8 other wolves and 2 of those wolves were pups born in 2020.

This means V052 had denned in an unknown location in the spring of 2020 but most likely also at least in the spring of 2019 which would make 2 breeders + 2 pups + 5 subordinates (5 young wolves born in 2019 or maybe in 2018 and 2019).

5 pup survival through 1.5 years without any lost to mortality would be wondrous in the area and thereby it is possible the 5 subordinates were offspring of 2018 and 2019.

The Bluebird Lake Pack lost 1 member in the winter of 2020/21, most likely to dispersal because it had 5 subordinates and some of these individuals should have been inclined to disperse.

Such a big pack did not suffer great consequences due to this dispersal.

However, it is worth noting that the retention of the subordinates is a likely contributing factor to the pack’s success at taking over some of the Wiyapka Lake Pack’s territory thus selecting their range.

The move slightly back closer to V052 natal range might have been deliberate and this indicates that large packs in which not all yearlings disperse can make more active decisions regarding where they want to reside.

In fact, the Bluebird Lake Pack even intruded in the Huron Pack’s territory and they may have wished to claim V052’s natal range but this claim was unsuccessful.

Or the challenge was not attempted at all due to unknown reasons because the BLP was two or three times stronger than the HP.

Huron Pack had highly experienced breeders who had even managed to take over a far larger pack’s territory and this could have been one of the reasons why the BLP (with its own experienced breeders) did not attempt a strategic move (experienced pack members can considerably determine pack success in conflicts and perhaps the BLP did not wish to risk it with the old, wise Huron wolves; see Cassidy, K.A. et al., 2015).

It might have resulted in death of 1 or 2 Huron Pack’s members – it is not conclusive because the confirmed death of Huron’s subordinate occurred on the western part of HP’s range while the BLP had settled in the east.

Thus, it is possible that the Bluebird Lake Pack did not have any actual territorial claims over HP’s territory but simply intruded by necessity (lack of resources for the large group) or out of curiosity (the BLP breeding female V052 would have wished to know what had happened to her family and its home).

If BLP had lived side by side with the former Sheep Ranch Pack previously, these intrusions could have been a regular occurrence (tolerated by the kin) or they could have begun after the relatively successful reproductive season (for BLP) as the non-related pack was no longer entitled to peaceful coexistence side by side (as it would have been with V052’s natal pack).

However, when Huron Pack eventually dissolved (perhaps due to pup mortality in 2021), it was not BLP but the northerly Paradise Pack which overtook HP territory.

It is also interesting that V052 younger sister V082 (whom she probably had never met because this female wolf was born in 2018 or 2019) did not join sister’s pack but rather joined the Huron Pack which had taken over her natal territory (Sheep Ranch Pack’s territory).

The Huron Pack took over in January, 2020 but the Bluebird Lake Pack was noticed in October, 2020 (if the BLP had not been a neighbour to the Sheep Ranch Pack prior to 2020).

Thus, it is not surprising that this SRP female V082 initially (summer, fall) stayed with the Huron Pack (if she, for some reason, was reluctant to disperse).

This could attest to these two packs (BLP and SRP) not having resided side by side which could have created familiarity between the related individuals in the two packs and which could have induced V082’s joining her sister’s pack.

(However, the BLP was large and it might not have been willing to accept a new member while the SRP could use ‘extra paws’.)

If the Huron Pack had run-ins with the Bluebird Lake Pack, it is possible that V082 met her sister V052 in an agonistic situation.

Interestingly, no observations exist of V082 with the Huron Pack during winter when the intrusions by BLP occurred.

But later V082 was, once again, with the Huron Pack (February, March, 2021) which means that she either had not recognized her sister (whom she had never met but whom she might have identified through other kin cues, e.g., scent which was left as markings near the common territory boundary between HP and BLP), could not join her sister’s already large pack (if she was too reluctant to disperse in the first place, she might have been too timid to approach 8 – 9 wolf pack) or briefly investigated and decided to return to the HP which had accepted her once already and which had been diminished in numbers (tolerant toward non-related subordinates).

Back to Bluebird Lake Pack’s story…

After claiming the territory, they produced pups in 2021 and appeared to have lost interest in expanding their dominance further.

Their subordinates began dispersing (3 dispersed by the end of summer and 2 more between summer and fall).

Still, the Bluebird Lake Pack was one of the largest packs in the Voyageurs National Park in the winter of 2021/22 and the dispersal of subordinates probably did not change much after they had established themselves or they did not attempt to retain their yearlings or older siblings.

However, in the new map (2022/23) posted by the Voyageurs Wolf Project team the Bluebird Lake Pack no longer exists.

My prediction is that the pack dissolved after the death of their breeding female who was old for the average wolf life span in Voyageurs (7 yrs in 2022).

Her death could have resulted in loss of pups, as well. If many subordinates dispersed during 2022, this could have left the pack very small (minus at least one breeder, no pups).

At least one of the new packs on the BLP former range might have been founded by former BLP wolves.

If the number of retained subordinates did not drop dramatically during 2022/23, the pack could have divided in smaller groups rather than dissolved because dissolution is less common in larger packs (which will be known only when the VWP team publishes the report for 2022/23).

Splitting of packs is not unusual but 3 packs are now seen where BLP used to reside (splitting usually results in two packs).

It is known that the Vermilion River Pack is led by the breeding male that was born into the neighbouring Bug Creek Pack but this male could have paired up with a former BLP female.

The age of the BLP old breeding male VP0C has not been given and he could have remained as a breeder in one of the groups.

I wonder that the VWP team would not have changed the name of his particular pack if that were the case (unless his pack moved a considerable distance from the Blue Lake) but that is not what they consistently do (e.g., they renamed Moonshadow >> Blood Moon but in this case, they were not 100% sure that it was the same breeding female).

We will hopefully see what happened.

Cranberry Bay Pack

Cranberry Bay Pack was 6 wolves strong in early December, 2020, but by January, 2021, only 4 wolves were observed and 2 wolves either died or dispersed.

I lack data in order to determine the likelihood of either scenario (it has not been mentioned if the pack had any subordinates and how many pups had survived).

1.5 year old wolves (or older subordinates) are likelier to disperse while pups might be likelier to have died.

In 2021/22 Cranberry Bay pup survival rate was very high (3 or 4 of the produced 4 pups survived at least until 2022) and it is possible (and more delightful) to assume that the individuals who were no longer seen in later winter 2020/21 were dispersing older siblings.

Later note – upon reviewing some of the VWP videos, I noticed a comment in this video that at least one of the 2021/22 4-year-wolf combo was a pup.

Regardless of the status, it did not appear to bear much impact on the pack’s affairs.

Intruders had been noted on Cranberry Bay Pack’s territory in February, 2021, when the pack was more vulnerable (having lost two members) but apparently 4 wolves were still formidable enough of a force (the intruders also were of the count of 4).

It is interesting to observe such intrusions of large groups that probably constitute entire packs or subdivisions of packs.

It could have been the newly formed neighbouring Nashata Pack which was 4 wolves strong and whose territory was rather cramped.

However, it would have been mentioned by the Voyageurs Wolf Project team because one of the Nashata Pack’s wolves was collared and recognizable.

I wonder who these wolves were and whether they lived nearby (but outside of the Voyageurs National Park study area) and performed an excursion out of their own range or if they were a ‘dispersing pack’ (a displaced pack or a pack disrupted due to loss of breeders etc.).

‘Dispersing pack/group’ situations are rarely described in the literature but they have been known to happen (e.g., the famous story of the Yellowstone male wolf 302M Casanova who dispersed from the Druid Pack with 5 other males).

This could have been a new group, as well.

February is the mating season and a breeding pair could have not only met one another but also ‘ganged up’ with some other dispersers from their natal families (which was the case, for example, in the founding event of the Nashata Pack, see below).

However, data has not been provided to make any assumptions in this case.

This particular event might be indicative of the ‘power of residency’ where of two packs of equal size (4 and 4; size being determinant in intraspecific strife outcomes, see Cassidy, K.A. et al., 2015) the residential pack prevails (if there were any territorial claim at all regarding the intent of the intruding group).

The Cranberry Bay Pack bred successfully in 2021 and 3 or 4 pups survived until the winter of 2021/22.

Once again, during late fall and early December, the Cranberry Bay Pack was larger (7 wolves) and their number had decreased by January, 2022.

I find it interesting that, for the second year, the dispersal occurred during late December / January.

Unless there are severe food constraints specifically on this pack’s range during this period (resulting in death or social tension and dispersal), there might be other reasons determining temporal patterns of dispersal.

For example, the Cranberry Bay Pack lives on Kabetogama Peninsula which is rather surrounded with water and which has many wetlands.

Perhaps midwinter is the most suitable time for dispersal when the ice cover allows for movement in different directions (without the need to intrude into the neighbouring packs’ territories which can be dangerous).

The observation of intruding wolves during this period could confirm increased ease of access.

If I understand correctly, the year of 2020/21 was harder on the VNP wolves than the subsequent year.

It might be useful to consider the number of wolves that in every particular year disperse from a territory respective to the pup survival and to interannual resource availability as well as mortality (quantified as some climate variables).

It might point at the carrying capacity of the particular wolf pack’s range, also allowing to track whether the demographic changes over time are normal or abnormal relative to average rates or extreme but natural events (abnormal changes could indicate at issues of species conservation interest).

E.g., if the wolves do not expand their range and their territory does not contract, either, there could be a limit to how many wolves can stay in the family in an average year (not too harsh, not too abundant).

For example, the Cranberry Bay Pack had 4 wolves in winter 2020/21 (the harsh winter) and 6 wolves in winter 2021/22 (the ‘easier on wolves’ winter).

Thus, the dispersal of the two wolves in 2020/21 could have been the result of lower carrying capacity due to the climate of the particular year while 6 wolves could be the actual carrying capacity for the CBP (unless it changes its range).

On the other hand, retention of subordinates in 2021/22 could have been an adaptive reaction to the pressure of the newly established neighbouring Nashata Pack which did not truly keep to its own and frequently overlapped its activity with the CBP range.

Such studies of individual territory carrying capacity might be difficult to conduct because long-term stable packs are probably scarce in the wolf distribution area (discovered mainly in relatively undisturbed populations and conditions).

I also find it curious why some wolf packs appear to rather try to retain their subordinates (seeking to expand their territory if the range becomes too small to sustain the large family) while in others dispersal regulates the balance between the home resources and its family.

For example, the above story of the Bluebird Lake Pack could suggest that the BLP had ‘accumulated’ subordinates in order to take over new areas.

But it could have also resulted from their wish to have a larger family and the respective need to expand their territory rather than allow dispersal.

The desirable pack size could be dependent on prey size or scavenger pressure.

But it could also be, for example, explained by genetic factors where some wolf groups are more sedentary and wish to just be with their families.

While others are more exploratory and the young are more eager to leave and to discover their own fortunes (also, reproductive ambition could be of importance).

Strangely, sometimes family-oriented groups might appear even more excursory (in a localized way) if they cannot satisfy the needs of the large family without intruding on other packs’ territories.

I believe that the Cranberry Bay Pack will need to retain an adequate number of subordinates because they appear to a have territorial overlap with the Nashata Pack (with at least one conflict possibly observed in March, 2022).

It remains to be seen if their number ever falls below 6 after 2021/22 (unless there is no other option due to very poor pup survival) while the Nashata Pack is still actively around.

Half-Moon Pack

In 2020, Half-Moon Pack produced pups but all 4 died until winter.

Meanwhile, the Half-Moon subordinate stayed with the pack until spring 2021 when the subordinate dispersed.

The Voyageurs Wolf Project team described this behaviour as usual for 2 year olds in their study area.

And it is common, indeed, from the perspective of wolves everywhere.

However, it made me wonder if poor pup survival leads to older dispersal age in subordinates (e.g., 2 years vs 1 year).

This could be another mechanism how poor pup survival or reproductive failure (e.g., due to hunting) may retain subordinates but can also contribute to pack turnover in the area because packs with subordinates appear to be more likely to expand their range or claim new ranges.

This would be different from takeovers that result from wolf abundance (packs that have pups + subordinates).

In such cases, the packs challenge other packs while still themselves vulnerable and with their fates unknown (dependent on retained older offspring that eventually disperses and on hopes to produce more pups in the following season).

Such pack distribution and tenancy could be less stable although, as evident from this very same Half-Moon Pack, it is stable enough.

Thus, in years of low pup survival and reproductive failure pack turnover could become more probable (just like in years with above average pup production) – in areas where subordinates do not disperse too young (e.g., low population density areas).

But it seems that this often results in territorial shifts and less often in complete displacements.

It is interesting that by autumn the Half-Moon pack consisted of the aforedescribed 3 wolves (2 breeders + one 2-year old) + 2 pups (that died by mid-December, 2020) + 1 other subordinate which was apparently a 2.5 year old subordinate from Bowman Bay Pack (this wolf dispersed or died also by mid-December, 2020 leading the pack 3 wolves strong).

During 2020, Bowman Bay Pack had taken over Sheep Ranch Pack’s territory to the south and they had been renamed the Huron Pack.

One of their subordinates V093 apparently stayed with the BBP / HP but this other subordinate (born in 2019 perhaps a sibling to V093) ended up with the Half-Moon Pack.

According to this video (description) by Voyageurs Wolf Project, the Half-Moon Pack had caused the Bowman Bay Pack to move south (forced out of their territory).

The Bowman Bay Pack appeared to have suffered social destabilization during which one of the subordinates remained with the parents who somehow managed to take over the larger Sheep Ranch Pack’s territory but the other subordinate, for some reason, stayed with the invading Half-Moon Pack.

Meanwhile, a Sheep Ranch Pack’s subordinate joined the Bowman Bay Pack turned Huron Pack.

I find this behaviour rather strange.

Both wolves were probably old enough to disperse if they did not wish to join the struggles of their parents and thought that the misfortunes of their natal pack indicated they had to take matters into their own hands.

It appeared as if these particular individuals felt they needed more time not in their family but in their natal territory.

Usually it is assumed that subordinates stay behind if population density is too high and/or they can gain important experience that lead to established skills as breeders, providers and defenders.

These are normally skills thought to become derived from the social group and not from the territory itself.

It is also unclear whether the BBP subordinate helped its parents to take over the Sheep Pack territory and then dispersed back to what was now Half-Moon Pack’s territory actually travelling with the HMP (and not just roaming on their range as a lone wolf).

VWP mentions 4 wolves forming the Huron Pack but the SRP subordinate obviously joined the HP later when her own family had broken up.

It is possible that there were additional subordinates (apart from the one that ended up in Half-Moon Pack) in the BBP (I lack such data).

However, it is interesting that these wolves would rather stay behind (and at a great age, too – 2.5 years) when their family was gone and they probably could assume independence.

It almost appears as if they wished to learn more from their natal home.

It is also possible they were hoping to attain a breeding position.

This could be the case in the Huron Pack because at leas the breeding male (and possibly the breeding female, as well) was already rather old.

The Half-Moon pair, however, did not show much vulnerability.

Half-Moon Pack’s pup survival in 2021 was very poor (1 out of 8 survived past August, 2021).

But Half-Moon Pack has endured and they still show up on the map of 2022/23.

Moonshadow Pack

Moonshadow Pack is one of the examples where the breeding pair split up (perhaps due to incompatibility, it has not been stated in reports).

It appears that on such occasions, it is the male who inherits the territory while the female disperses more often than the other way around.

While, under the scenario of death of the breeding male, the breeding female remains on the territory and does not have great trouble finding a mate.

It also seems that in the latter case, breeding occurs with less interruption (without a ‘gap year’) although it is not always so.

Perhaps the individuals of the original pair are not as eager to mate again because they have suffered loss.

If the newcomer is a male, they might be more persuasive because males generally appear to be the initiators of copulation in the species.

If the newcomer is a female and the male is not as motivated due to emotional upset or bad experience, breeding might be postponed more often.

For example, in Blood Moon Pack (which will come into the story soon enough because it is the transformed Moonshadow Pack in 2021) the breeding male died in January, 2020 and the new breeding male appeared to have replaced him within the same month but pup production did not occur in the same year.

Perhaps it also depends on the age of the widowed individual or the reasons behind the pair’s break-up (motivation and caution).

This would be interesting to study (also from the perspective of the origin of the individual who remains on the territory in the case of a pair splitting and whether the individual is more likely to stay than to disperse if they have been born nearby vs. immigrated into the area).

Dispersal is also the process which ensures such quick replacement of mates.

Indeed, it appears that it is important to have some lone wolves roaming the general area and looking for opportunities.

They seem to fill the vacancies even within weeks after the vacancy has emerged.

The timing of dispersal could be thereby related to winter mortality because winter could be a time when it is profitable to be hanging around resident packs.

If the reproductive activity does not occur in the same year, it might still be beneficial to accept a new prospective mate because this ensures the ability to keep the territory (to defend it against claims by other wolves).

This was what happened in 2021 when the lone wolf Y1T who had been a floater in the Voyageurs National Park almost immediately took over when the former Moonshadow’s breeding male died.

Y1T was very invested in bonding with the former breeding female of the Moonshadow Pack and her pups (see this VWP video) and because he was a large, young and healthy wolf, I find it interesting that he did not select other options available to him.

Many packs in the Voyageurs NP were vulnerable in the winter of 2020/21 and consisted only of breeding pairs (6 packs) or was a ‘disputed area’ (Whiskey Point).

It would not have been difficult for Y1T to pair up with another wolf (or even in a solitary capacity) and to displace a resident pair (just like the Stub-Tail Pack did with respect to the Tamarack Pack and like the Whiskey Point Pack did with respect to the Sand Point Pack territory).

There were probably other opportunities nearby enough (regarding packs and areas that are not included in the VWP study area).

I wonder if there are wolves who prefer one strategy over another and if Y1T was rather set on gaining acceptance by an existing pack than challenging the residents (or other roamers).

I am quite certain that Y1T was informed of other options because he had had been around since before January and he had roamed over several pack ranges.

I have developed a theory that young, healthy wolves simply are not predisposed to aggression.

Y1T was fairly large and he could have been able to provide for himself as a lone wolf maintaining adequate condition without assistance (a potential reason why, unlike other wolves, he had not paired up while a floater).

Perhaps such healthy wolves find it difficult to muster aggression in order to launch an attack and to force other wolves out of their ranges.

Perhaps a greater degree of despair is needed or a pair bond (e.g., motivation to secure a breeding site for the newfound partner and anticipation to breed when exposed to the proximity of a potential mate).

Maybe it is not as easy to displace other wolves and two wolves are likelier to achieve it (e.g, we only observe the results of the unfolding events and we cannot know what Y1T had attempted prior to joining the Blood Moon Pack which did not result in attaining a breeding position and a territory)

We do not know, either, what happened to the Moonshadow breeding male.

The WVP team concluded his death due to starvation and not due to strife.

But that does not mean Y1T had not actively displaced him (starvation being the consequence of inability to feed without his pack’s support) in a manner that did not cause injury.

It could have been sufficient for Y1T to linger in the area and demonstrate his intent relatively passively.

His size and youth could have induced a retreat by the older wolf.

For example, Nashata Pack’s breeding male claimed his position amicably (possibly through being the larger male) over another male who had greater ‘rights’ to the specific territory (having been born in the neighbouring Shoepack Lake Pack and having thereby elicited tolerance by the SLP that allotted some of their own range to the newformed Nashata Pack).

Large wolves might enjoy lower conflict rates because many conflicts are resolved through comparison and not through actual interaction.

Nashata Pack

Nashata Pack has a very curious story of origin.

It was formed after the Shoepack Lake Pack wolf V074 dispersed and began roaming between the SLP and Cranberry Bay Pack territories.

There he apparently met other dispersers which is also interesting because I believe it is more frequent for wolves to pair up and then to settle or for a lone wolf to find an unoccupied range and then to wait out a mate than it is for wolves to form groups.

Especially, in this case where V074 was probably not related to the 3 other wolves he met.

Two of those wolves had formed a breeding pair and they could have been a displaced family group (breeders + their subordinate offspring) but they could have also met and set up a pack without prior acquaintance.

This group benefited from V074’s relatedness to the Shoepack Lake Pack as they could take over some of the SLP’s range without territorial conflicts (tolerated by the SLP probably due to V074’s status).

Partly, it is also possible that SLP was unwilling to provoke the new pack although at the time they had superior numbers (5 adult wolves + 2 pups).

It might be a strategy for dispersing wolves to ‘gang up’ in order to hunt together and to secure their territorial claims.

However, it is difficult to say how this strategy benefits wolves that do not become the breeders because they mostly disperse after pups have been produced (or a bit later).

Perhaps their profit is related to food provisioning and safety that comes with a defended territory (vs. the ‘sneaking around’ on other packs’ ranges where any larger kill is likely nonviable because it cannot be guarded against the residents and the lone wolves thus have to rely on ‘one-time meals’).

During spring-summer V074 dispersed from the Nashata Pack, as well.

Once again, there is the tendency for a subadult/adult wolf to stay nearby natal range (but not with their actual family).

This could be the result of high enough population density.

The wolf might disperse and then realize that all territories have been taken and it makes more sense to rejoin the natal pack or to briefly join a non-related pack especially if there might be a breeding opportunity at some point.

As mentioned before, 2020/21 was not an easy year and dispersing individuals could have learned that life was too difficult on their own.

This could have prompted the acceptance of other breeders on what should have been V074’s range (partly consisting of his parents’ territory).

The other Nashata subordinate also dispersed in winter 2020/21.

Apparently the range had been secured and their dispersal no longer threatened Nashata Pack’s territorial status.

Nashata Pack will be interesting to observe because they are cramped between two other packs with a slight range overlap and I wonder if this would prompt them to attempt to retain subordinates (because other packs on the Kabetogama Pensinsula tend to be relatively large) or if they simply cannot afford it because their range is comparatively small.

If they stick to their own territory (not using the overlap), they might starve.

But in order to be able to sneak around at all times, they might need a larger pack size which, once again, puts a strain on their resources.

Thus, dispersal could involve a trade-off between the opportunity to frequent dangerous grounds and per capita food availability.

Shoepack Lake Pack

One dispersal case in Shoepack Lake Pack has been described under Nashata Pack entry (a SLP male participated in founding the Nashata Pack).

However, the pack decreased from 7 wolves in summer, 2020, (including 2 pups) to 6 wolves by mid-December, 2020, to 5 wolves in late December, 2020, to 4 wolves in January, 2020.

2 of the wolves constituted the breeding pair and it is not known who the other 2 wolves were but they could have been either the pups (less likely to disperse) or pup + subordinate combination (if one of the pups died and two subordinates dispersed).

The packs on Kabetogama Pensinsula are rather successful breeders which is why I find it likelier that at least 1 pup if not both survived (and I find it improbable that both pups died while 2 subordinates stayed because subordinates would have dispersed first if there was food shortage or other such circumstance).

It is known that at least one of the dispersants was a subordinate.

These fluctuations could have contributed to permanent establishment of the Nashata Pack (overlapping the Shoepack Lake Pack range) although the SLP wolves disappeared in winter when Nashata Pack was already an influential unit.

The two subordinates were apparently retained because until December, 2021, the SLP was 8 wolves strong and even with very successful pup production, they would not have attained such size without subordinates staying behind (the 2020 pups, presumably).

Once again, during the course of winter 2021/22, the pack became smaller from 8 wolves to 6 wolves in April.

If the pack was 8 wolves in December (assuming that both 2020 pups had stayed behind as subordinates by then), 4 pups had survived in 2021.

Then the two subadults (or to be more precisely, 1.5 year old wolves) would have dispersed (or, once again, some combination between subadult dispersal and pup mortality/dispersal).

This pack appeared to follow a very regular and ‘normal’ pattern of reproduction and dispersal, almost a text-book case.

I would have been very confident that the Shoepack Lake Pack continues with their ‘model family’ strategy but, surprisingly, the SLP does not show up on the new map of 2022/23.

Considering their large size and subordinate retention, it is hopeful at least (without knowing what happened) that the territory could have been inherited by one of the original breeders or 2021 offspring forming a new unit.

The Shoepack Lake Pack had existed since before 2020 and it is possible that one of the breeding wolves was old and might have died causing pack restructuring or break-up.

Tamarack Pack

Tamarack Pack represents one of the occasions when a pair disperses together after having been displaced by another pack (the Stub-Tail Pack or, at the time, pair).

They only moved slightly north from the territory which they had established and settled adjacent to the Stub-Tail Pack.

Perhaps the proximity of the opportunity to resettle determines whether packs and pairs remain together in such situations.

It would be interesting to study if displaced pairs / packs have a greater probability of remaining together with decreasing distance from available patches (or vulnerable packs nearby).

All in all, not much information is given on what happens after the pack or pair experiences a disturbance that is great enough to potentially prompt a break-up.

For example, there might be some temporal gradient according to which complete dissolution becomes likelier and likelier.

E.g., the former pack mates (or the pair) might persist and roam together for a certain period after which they determine that they have better chances if they try their fortunes individually (or they cannot remain together because it is impossible to provide for a group on other groups’ ranges).

Obviously, these chances are affected by resource availability (prey availability even if vacant territories are not available), perhaps pair bond length (e.g. long-term partners less willing to split up), pack member age (yearlings and subadults might be more prone to disassociate from the roaming group than pups and established breeders).

In Tamarack Pack’s case, the dispersal did not result in pack persistence although not due to unwillingness of the pair to overcome the death of their only pup in 2021 + displacement from their range after the pup’s death – but rather due to the death of B1T, the breeding female, who was shot illegally in March 2022, after which event her former mate dispersed.

I believe that the mortality risk could have been higher because apparently the Tamarack Pack was forced slightly out of the official Voyageurs National Park range in which wolves might be better protected.

It has been mentioned that the Tamarack Pack was forced out to reside between the Stub-Tail Pack and Highway 53 and major roads provide access for poachers (hunting and poaching rates are higher close to roads).

I suppose that her mate did not attempt to stay on the territory because it was suboptimal.

Otherwise, he could have deployed the strategy of waiting for a new mate.

Also, it was past the mating season and his motivation to retain breeding status and an established territory might have been lower (as compared to prospects during winter).

The Tamarack Pack story is, indeed, very tragic.

Whiskey Point Pack

Whiskey Point Pack was formed after a pair of roaming wolves (possibly two paired-up young dispersers) displaced the Sand Point Lake Pack which had been a breeding pair for possibly 5 years until then.

It has not been mentioned what happened to the SPLP pair after diplacement but none of their 2019 pups survived and beginning from the winter of 2020 the pack’s territory started being frequented by another pair of wolves.

The SPLP maintained residency until at least December 2020 but there were no pups with them.

The other pair (the Whiskey Point Pack to be) was also spotted on the same range throughout 2020 but apparently they force the takeover (possibly due to the experience of the older wolves or maybe some other factors were involved).

I find this story interesting because it potentially juxtaposes two types of dispersing pairs (if the SPP pair dispersed together after the displacement which cannot be fully confirmed).

One pair probably consisted of two 1.5-year-old or 2.5-yeard old dispersers who had paired up after leaving their natal pack.

The other pair consisted of established breeders who had suffered poor pup survival and eventual displacement (which could have occurred peacefully if the SPLP began doubting their reproductive success as a pair or on the particular range but it also could have been pressured through the constant presence of the strange pair on SPLP territory reminding the SPLP pair that there were other claimants and causing higher stress levels).

It is worth noting that the Whiskey Point pair stayed together and persisted in their search for territory for several months.

It has not been mentioned when precisely the future WPP pair first appeared in the SPLP territory but from the context it seems that this pair had been roaming around and evaluating their chances for over half a year.

I come by a subjective impression that there are differences in persistence between pairs that have been displaced or that have split up (after they have become roamers) to stay together vs. pairs that have bonded recently and are on the lookout for their first breeding range.

In the publication by Fritts, S.H & Mech, L.D., 1981 such newly-bonded pairs roamed and settled and roamed again for at least a year in some cases (although mate change or mate abandonment also occurred during this period in some pairs prior to actually reproducing).

The WPP pair also roamed together patiently and for a longer while.

At the same time, former breeders give an impression that they split up more easily once the pack has been disrupted/displaced.

If this is true at all (and it could be quantified), it might be interesting to study the reasons behind such differences.

Perhaps I am wrong because a long-term pair bond should encourage endurance.

However, it could be precisely the bond, the memories, the attachments, the experience that could increase the amount of stress due to the losses suffered.

Such pairs could feel more doomed than young pairs ‘with nothing to lose and all to gain’ because they could become depressed due to losing their home and having their bonds threatened.

Their experience and potentially greater age might interfere with the ‘let us try our luck’ approach possibly entertained by younger dispersers.

These old pairs might regard novel situations with a more critical attitude because they have been used to assuming active leadership rather than relying on fate and chance.

Pairs with former territorial experience could also have created specific foraging routines that are no longer as adaptive and flexible as would be necessary to survive in a highly dynamic circumstance.

All in all, it would make for an interesting research.

Windsong Pack

Windsong Pack’s dispersal case was that of a dispersing breeding female after her mate (of the former Fawn Crick Pack) had been lethally removed due to depredation in summer, 2020.

It is not known precisely how the events unfolded after that because the breeding female had had pups.

It would appear that the FCP female dispersed but one of her pups survived and joined his brother (a former FCP subordinate) who took over and found a mate.

It is difficult to imagine why and at which point exactly the female dispersed.

She was related to the new pack leader (her son V087) and she had pups to take care of.

Did she disperse right after her mate’s death or did she wait until the pups (one pup that survived) were old enough to survive without her supervision?

Why did she not become a subordinate in her son’s pack.

Overall, packs are more open to accept male members as subordinates likely due to resource acquisition and defense efficiency reasons.

Rick McIntyre has claimed in some of his interviews that while the breeding pair has rather equal role division (regarding to dominance), the female is the true matriarch of the group and she dominates the male at least during the breeding season.

From what I have read, subjectively, it feels that in these cases where the former breeder has lost their mate and/or status, it is easier for males to integrate into the new pack structure as subordinates than it is is for females (unless the female becomes the new breeder).

As mentioned before, this could be the result of greater tolerance toward males as pack members.

But it could also indicate at social behaviour where the female finds it harder to ‘step down the ladder’.

It is not impossible, however. For example, see the famous Ellesmere pack where Whitey (daughter) took over the breeding position from her mother who stayed in the pack for 2 years helping to raise the pups (Mech, L.D., 1995).

Perhaps the degree of relatedness and the sex of the replaced breeder (non-kin) are of importance because in Windsong Pack’s case the new non-related breeder was of the same sex as the surviving former breeder while on Ellesmere Whitey was Mom’s own daughter.

It has not been stated in the report how old the Fawn Crick Pack breeding female had been (perhaps she was young enough to make it harder to accept she would not be producing more pups while on Ellesmere Mom had been 7 – 9 years old and reproductive ability in wolves decreases after about age 4 – 5 (Stahler, D.R. et al., 2012) and 11 years is already of the final age groups for reproduction if the individual even survives that long).

I believe the reasons behind these changes in dominance positions could be hormonal.

Female breeders also produce testosterone (at higher levels than non-breeders).

Perhaps while in male wolves testosterone production is more natural and can be invested in other pack-related activities, long-term testosterone production in females results in other types of reactions disallowing to physically adjust her behaviour as necessary to accept the subordinate position.

In older wolves, this could no longer be as marked.

Other reproductive hormones could be of importance, as well, altering female body in ways that prevent ‘resetting’ after a certain count of years have passed.

Perhaps these hormones cannot be ‘sublimated’ into other types of pack activities (unlike in the male).

I also wonder about the possible pup abandonment by the FCP female after she lost her mate.

It is not known if it happened at all (apparently one pup was healthy enough and nurtured enough to survive and 1 pup survival is not uncommon in Voyageurs wolves also under more regular circumstances).

But if it did, I wonder if the psychological impact suffered by the female upon losing her mate during the reproductively most active period might affect her hormones also in a manner that disrupts caregiving behaviours.

Wolves might be less adapted to process grief over loss of a mate during summer season because wolf pack ranges shrink and intraspecific strife can lessen (involving attacks on pups rather than on adults).

Intraspecific strife and mortality due to starvation etc. could be more frequent in winter when hormonal status is different in the female.

Maybe the combination of mate loss in summer + hormonal production in summer is highly non-adaptive which would caution against any lethal activity against wolves during this period in order not to endanger the litter of the year through disrupting maternal (parental) caregiving.

This was specifically what occurred in Windsong Pack’s case – the FCP breeding male was lethally removed due to depredation and so were Windsong Pack’s two pups of 2020.

While I generally disapprove of lethal removals, during summer they could bear an even more detrimental impact on wolf’s (female’s) psychology.

Windsong Pack is also an important example demonstrating the ‘salvaging factor’ of subordinate retention (at an age which allows to assume the breeding position if the breeding parent has been lost).

V087 took over and probably invested greatly in the survival of his younger sibling.

The pack was also rescued by the dispersing female who paired up with V087 in a manner that did not disrupt pack’s productivity.

Thereby floaters are of great importance in wolf populations.

Perhaps this is why they are somewhat tolerated.

On one hand, they represent competition and threat.

On the other hand, they are there to rescue disrupted packs when calamity strikes which can be unpredictable.

Wiyapka Lake Pack

Wiyapka Lake Pack was one of those packs that retained a subordinate throughout 2020 which possibly was beneficial for the pack’s persistence because the pack suffered some hard times.

They lost all pups in 2020 and some of their territory was taken over by the Bluebird Lake Pack (see above).

Their subordinate could not have helped much against the BLP (which was 9 wolves strong) but there were several territorial changes taking place in 2020 all around the Wiyapka Lake Pack (e.g., Fawn Crick Pack > Windsong, intrusion by BLP into Huron’s territory).

Their neighbouring Lightfoot Pack already had experience in taking over other packs’ ranges (in 2017) and their pup survival had been rather poor (perhaps due to insufficient resources and too small territory size) which could have prompted them to expand their territory (but they were 2 wolves while the Wiyapka Lake Pack was 3 wolves strong).

It is always difficult to make claims regarding what could have been or what could not have been but I find it rather probable that the subordinate facilitated pack’s persistence until he dispersed in April, 2021.

It was just as well because the Wiyapka Lake Pack had successful pup production on 2021 and appears to have held its own until at least 2022/23.

Stub-Tail Pack

Stub-Tail Pack’s story is closely linked to Tamarack Pack’s story (see above) because in 2021 they took over the latter’s territory.

Before that they were yet another nomadic pair of wolves seeking opportunities to settle.

The takeover does not appear to have been aggressive (but perhaps stressful due to the presence of strangers just like in Whiskey Point Pack’s case).

This takeover occurred in mid-autumn 2021 and it is difficult to say for how long the Stub-Tail pair had been together and how often they had trespassed prior to the final takeover.

I do not know where the name for the Stub-Tail Pack came from but the Shoepack Lake Pack has tail genetic tail deformities (see this Voyageurs Wolf Project video).

If the Stub-Tail Pack’s name refers to tail deformities in one of the partners, this wolf could have dispersed from the SLP.

Possibly three wolves dispersed from the SLP in winter 2020/21 and one of these wolves could have found a pair.

The Shoepack Lake Pack is rather far from where the Stub-Tail Pack was eventually formed (also accounting for the fact that the SLP resided on a peninsula and the population density appears to be high which makes it more difficult to move through the many territories of other wolves).

But it is possible.

When a video is posted of the Stub-Tail Pack wolves, we might see whether any of the individuals have tail deformities and whether it is a genetically short (but intact) tail (like in the SLP case) or a tail deformed due to injury lacking its tip altogether (which better corresponds to a ‘stub’).

Be as it may, I would be very interested to know what happens to dispersers once they form a pair and if their movements thence are different from lone wolf movements (due to differential nutritional needs, possibly lower stealth potential etc.).

I find it curious to understand when precisely the wolf pair decides it has bonded somewhat ‘irreversibly’.

As mentioned before, there are causes where wolves pair up briefly during dispersal, then separate and pair up with other wolves.

The determinant could be pup production (successful mating) but many of these pairs appear to exhibit loyalty also prior to having mated.

It could be territory acquisition and commonly performed scent-marking which is an important bonding activity in wolf pair.

See, for example, this Austrian + Polish wolf pair which is scent marking together with such devotion that it fails to notice and be concerned with a roe deer in the background – in a video by Association for Nature ‘Wolf’.

The video appears to have been recorded outside of the breeding season.

If the failure to notice the roe deer is indicative of immersion in another activity, it is not a mating activity which overrides hunger but it is the scent-marking activity with the potential mate (demonstrating its significance).

If scent-marking is essential to pair bonding, it might be constrained to when the wolf pair has actually found a territory.

However, if attachment (e.g., emotional attachment) may precede territorial scent-marking and the need to ritualize the bond is overpowering, the nomadic wolf pairs might engage in the scent-marking activity before they have decided on a territory or before a territory is even available.

Thus, the bond itself might determine where the pair would live as soon as it becomes strong enough to demand for common rituals (which are then executed in the respective place where the pair happens to be).

This could also result in conflicts with residential wolves if the pair begins scent-marking on their territory because they ‘cannot help it’ and they have a strong need to assure their bond.

Some takeovers could not have been as deliberate as they are the effect of pair-bonding activity which could be dependent on hormones.

For example, male and female hormones might change as they spend a lot of time together and they have close physical contact.

Also, hormone production during peak reproductive periods might be more pronounced in pairs (exposed to the potential mate) than in lone wolves.

This would be interesting to study, as well.