This post is a sequence to previous posts, Hunting by humans never constitutes compensatory mortality and Turnover in wolf packs – Voyageurs case studies (confirmed mortalities).
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 pup survival (possibility to recruit pups as helpers) influences the pack future fates.
Bluebird Lake Pack
In autumn 2020 a group of wolves had started travelling on the outskirts of Wiyapka Lake Pack.
They had 3 pups and this could have necessitated the group’s takeover of portions of WLP’s territory because the group was 9 wolves strong and such a large group might have been prompted and enabled to enlarge the territory (or to claim a new territory which I unfortunately do not know according to available data).
While pup survival success might have necessitated the redefining of the territory, I am not certain if it would have facilitated it (pups could have been to young and inexperienced to be counted) but the pack already had 4 older subordinates and the two additional, inexperienced pups were probably not a significant addition in terms of territorial challenges during their first winter.
1 wolf (or 2 wolves) apparently dispersed or died during winter 2020/21 but the pack still managed to hold the newly acquired range.
It is worth mentioning that the Wiyapka Lake Pack was only 3 wolves during the time of takeover of some of the portions of their territory and all of their pups had died in 2020.
The WLP failed to produce pups efficiently two years in a row (0 pups in 2020/21 and 1 surviving pup in winter of 2021/22) which is suggestive of lack of resources, harsh year or another factor.
The takeover was not aggressive and it was unlikely that a 3 wolf group would have stood against a 9 wolf group even if they were the residents (with local experience and higher stakes).
However, it does not appear that the WLP suffered due to these losses because the breeding female successfully produced 3 pups that survived to winter 2021/22.
Their only subordinate dispersed in April, 2022 and this dispersal resulted in his death outside of his natal range but it appeared to have been regular dispersal behaviour.
Thus, the chances were somewhat bettered for both packs although it is not clear what the benefits were to the WLP.
It is possible that there were no benefits to the WLP and they simply had survived through a difficult year (2020/21 was not an easy breeding season in many wolf packs).
But it seemed that the BLP improved their own fortunes without reducing chances for WLP and the takeover was fluent and did not result in dramatic disruptions.
However, it would appear that the BLP also intruded into Huron Pack’s territory and might have killed the Huron Pack’s subordinate (Huron Pack at the time consisted of 2 breeders, and possibly 2 subordinates (maybe some pups, up to 3) and was thereby smaller than the BLP).
But the BLP did not overtake the neighbouring Huron Pack’s territory, nor its portions although they had been seen on HP’s grounds (see below).
It is not clear whether the BLP actually killed Huron Pack’s subordinate (and later – breeding male) because the former incident occurred in the west part of HP’s territory while BLP intruded (and resided) in the east.
It is possible that these intrusions in the east part of HP’s territory had been mostly made by the BLP’s subordinates as is common with young wolves in large packs when they begin considering dispersal.
Of course, they could have roamed more widely, as well, at some point reaching HP’s west range.
Also, the BLP breeding female had likely been born on HP current range and the intrusions could have been exploratory attempts on her part to discover what has become of her natal range and her family (which had been displaced by the HP many years after the BLP breeding female had dispersed).
It is likelier because pre-dispersal forays would have involved individuals but the entire BLP group had been seen on HP range.
However, the situation has not been commented on extensively enough to know that happened.
In autumn, 2021 the pack was 8 wolves strong (5 adults and 3 pups).
As the last year pups would be considered adults (subadults) in autumn 2021, this would mean that the pack had gained 3 pups and lost 3 (sub)adults during spring – summer.
However, by winter 2021/22, two more members were lost (dispersal or mortality) and the pack was 6 wolves.
The pack remained stable despite the slight reduction in size and produced pups again in spring, 2022.
It appeared that the improved survival rate of pups (3 vs. 2) did not affect the pack’s social ambitions (they did not attempt to take over new portions of other packs’ territories) and the improved survival could have been the result of sufficient resources after the slight acquisition of range in 2020/21.
According to the new map posted by the Voyageurs Wolf Project, the Bluebird Lake Pack no longer exists.
I find it surprising because the success in pup survival and recruitment into helpers was not suggestive of any threats to the pack’s integrity.
I wonder if the pack might have split as sometimes happens in large packs.
Some of the new packs on BLP’s former range might be constituted of the BLP’s old members.
This might be especially likely if the pup survival in 2022 was also high and the pack ended up very large-sized (relative to the average pack sizes in Voyageurs) once again.
Half-Moon Pack
This pack is somewhat strange.
It has had very poor pup survival (none of supposedly 4 survived in 2020 and only 1 of supposedly 8 survived in 2021).
The Half-Moon Pack’s territory on map seems moderately large and they even appear to be frequenting neighbouring packs’ ranges slightly (or the neighbours are frequenting theirs but there is an overlap and not a takeover resulting in exclusion of one pack or the other from the overlapping zone).
There is a great pack turnover in this specific area and upon these conditions (unstable social groups, high pup mortality, difficulty to maintain an adequate range, possibly poor resources if pup mortality can be attributed to starvation etc.), one might expect that the HMP would suffer intrusions and attempts at takeovers of at least portions of their territory.
But it does not appear to be so.
I have no knowledge regarding the prey distribution or landscape features on HMP range (perhaps it is so large because it includes large or many unsuitable patches).
It simply seems that the poor pup survival and the dismissal by the neighbours of the opening opportunities (and the HMP had some decent-sized neigbouring packs at least in 2021/22 although not in 2020/21) indicates at the low quality of the pack’s home range.
Or else, the HMP has been simply lucky and the instability of the neighbouring groups has led to oversight by them of the HMP’s vulnerability (while preoccupied with their own ‘troubles’).
Half-Moon Pack appears to have persisted even into 2023 according to the new map and I am very interested to learn how they fared from 2022 to 2023.
Huron Pack
Huron Pack produced at least 3 pups in 2020 and apparently, during summer and autumn of 2020, it also accepted an unrelated subordinate which was a female from the pack (Sheep Ranch Pack) the territory of which they had taken over in the year before.
This female was also with the pack in February – March, 2021.
The breeding male disappeared sometime after November, 2020.
It is possible that he was killed by other wolves (see below).
They also had one related subordinate in 2020 and this subordinate was killed by intruding wolves that perhaps belonged to the Bluebird Lake Pack (possibly 9 wolves strong at the time with several subordinates conceivably inclined to explore and to claim home ranges.
It is not very clear whether any pups had been born in 2020 and survived until even fall because they were not observed in the spring, 2021 when the HP was only 3 wolves strong: the former breeding female, the unrelated subordinate and a new breeding male.
This example illustrates the danger of being a small wolf pack (if the pups did not survive until the events unfolded, the Huron Pack was merely 3 – 4 wolves strong) neighbouring a much larger pack (Bluebird Lake Pack, 9 wolves) seeking to expand its territory or producing potential dispersers who are exploring on neighbouring ranges and assessing the chances.
The fate of the 2020 pups is not really known (from the reports) and thus I am unwilling to comment on their mortality as a factor of influence regarding the pack’s fates and partial disruption (only the breeding female and the unrelated subordinate remained in the territory).
It is not know who the new breeding male was.
(It is possible that the former Sheep Ranch Pack subordinate lingered behind if she was hoping to be chosen for the breeding position by the new male.)
For example, 9 wolves were observed in the Bluebird Lake Pack until mid-January when the BLP lost 1 member to dispersal or mortality.
It is possible that this individual could have become the new breeder in the Huron Pack which had a vacant breeding male’s position.
The Voyageur’s Wolf Project team appears very skilled at identifying individuals from camera footage and I expect they would have noticed that the new breeding male was a former BBL member.
Also, it was noted that the new breeding male was large and therefore, he might not have been a subordinate (subadult age).
Perhaps the deaths and disappearances in the Huron Pack were attributable to either this new male (a lone roaming wolf) or to BLP, or to both, or to neither, and the large male outcompeted any potential BLP subordinates for the breeding vacancy in the neighbouring pack once the former breeding male had died (disappeared) already.
I find it somewhat strange to assume that he would have killed the related subordinate if she was female but the sex of the related subordinate has not been mentioned.
However, if the other subordinate had been a male, the new breeder might have killed him to prevent competition over the breeding status because the subordinate was related to his mother (the former breeding female) but not to the other subordinate (the former member of the Sheep Ranch Pack) and he could have attempted to breed the unrelated female thus taking over the pack’s breeding male position.
It is not determinable without knowing the sex of the deceased subordinate.
This could be true due to the reason that BLP resided and intruded to the east of HP while the HP subordinate’s death occurred in the west and might have been another incident entirely.
BLP’s intrusion could have been not the cause of changes in the HP but rather a consequence (sensed vulnerability followed by inspection of new opportunities).
The Huron Pack’s breeding female produced pups in spring 2021.
The Huron Pack apparently dissolved in 2021 and their territory was taken over by the Paradise Pack.
The HP dissolved in summer and thereby it is likely they lost their pups.
In this case, the loss of pups led to a complete pack disruption as can happen in such small groups (larger groups are less likely to dissolve).
Another factor could have been the relatively recent pair bond by the breeding male and breeding female of 2021.
Thus, if the new breeding male had, indeed, killed the related subordinate, it could have been a mistake because retention of another member would have increased the chances of HP to stay together (especially, if the female was related to the subordinate increasing the motivation to keep the group intact).
In such cases, nevertheless, subordinates tend to disperse.
However, if the other subordinate had been a male, the new breeder might have killed him to prevent competition over the breeding status because the subordinate was related to his mother (the former breeding female) but not to the other subordinate (the former member of the Sheep Ranch Pack) and he could have attempted to bred the unrelated female.
It is not determinable, however, without knowing the sex of the deceased subordinate.
The Paradise Pack did not seem to have taken over through means of territorial skirmish (they were also only 2 adult wolves strong at the time + pups who were too young to participate), but they likely expanded their range as the neighbouring pack disappeared.
Which was very beneficial for them because the PP appears to suffer from resource deficiency.
It is easily seen how the turnovers did not create dramatic disruptions and all the new alliances and residencies were fundamentally those of experienced locals shifting their bounds of influence.
Huron Pack’s story also prompts me to wonder about wolves’ motivation to defend and to keep their territory.
Of course, wolves are territorial animals and sometimes even a pair of wolves or a single wolf would maintain their chosen home and protect it against other wolves (e.g., before mating up and producing pups for the first time or during years when reproduction has not been successful for some reason).
However, such actions are not dependent only on ‘calculations’ and aware decisions.
The drive to keep one’s home might be supplemented and/or enhanced by hormones that are produced with respect to having pups (planned or around) and with respect to sociality.
Perhaps wolves have a harder time maintaining territories or groups if these hormones have not been produced (or the production has ceased, e.g., due to pup loss) and the adversity is great (or they have suffered a loss that has traumatized them briefly or permanently).
Pup defense might spur territorial behaviour for self-evident reasons.
But I wonder if pack size does not influence this motivation through the contrast in production of oxytocins (and other hormones) during pack cooperation and positive social interactions vs. the threat that is correspondingly perceived if the territory / group cohesion has been compromised.
If the group is tight, it has pups and subordinates and it works as a unit with high hopes for the following season, the pack members might experience better everyday ‘feeling’ than members in small packs without pups and with less certain prospects.
Thus, the physiological perception of what could be lost (derived as a comparison between the pleasant hormonal states and the danger of not having them anymore) might alter the wolves’ motivation to persist on a territory or as a group.
This could be one of the mechanisms behind lower chances at pack break-ups and reproductive disruptions in larger packs as well as the overall success in larger packs during territorial conflicts (which is also attributed, of course, to mere force, not merely motivation).
Lightfoot Pack
The Lightfoot Pack is an example of amazing persistence where the breeding pair had remained together on the territory they took over in 2017 until January, 2022 when the breeding male was killed and the pack structure changed but the pack itself apparently persisted at least to the winter 2022/23.
As claimed by the Voyageurs Wolf Project team, they have had very low pup survival rates (all 7 pups died in 2020).
It is possible that they lived on a poor quality range which might have led also to lower interest in their resources by neighbours.
I believe it is rather rare that failure to produce pups would not interfere with the pair’s determination to stay on their territory and to keep trying.
The count of possibly surviving pups has not been given, however, for other years prior to 2020 and ‘low pup survival’ might not necessarily translate into ‘zero pup survival’.
If the pack even managed to take over another pack’s territory (which could have happened due to the former pack’s own dissolution and not through conflict; not mentioned in the reports), they should have had some better success at least in 2016 or 2017.
Year 2021 appeared to have been better for this pack (at least up to January, 2022 when the breeding male was killed by other wolves) because in winter 2021/22 there were observations of 5 or 6 wolves (the breeding pair including the late breeding male + possibly 3 or 4 surviving pups).
The pup survival was likely the only factor securing the persistence of the pack on the territory because after the death of the breeding male, the breeding female did not find a mate (according to VWP estimates) and the pack did not reproduce.
The breeding female was not observed with the pack, either, which could have left the 3 subadults alone.
However, the new map still pins the Lightfoot Pack on its former territory and this is thereby a case where pup survival was the most important factor retaining the genes of a specific group on its range it had occupied since 2017.
Moonshadow Pack
The Moonshadow Pack did not have pups in 2020 and this could be a reason (or one of the reasons) why the pair split up (another important consequence of reproductive failures).
The former breeding female dispersed while the breeding male remained on the Moonshadow Pack’s territory and found a new mate.
It appears that perhaps mate incompatibility (and not territory quality) could have been an issue which was resolved through mate change because the MP produced pups in 2021 of which 2 might have survived.
However, the breeding male died of natural causes in January, 2022 and the breeding male’s position was taken over by a new male (a former floater).
It is possible that the pack retained its structure (the same breeding female and two of her 2021 pups), and it is now known as the Blood Moon Pack.
As far as I understand, the pack did not reproduce in 2022 but they did reproduce in 2023.
On this occasion, it is difficult to say whether pup survival influenced the outcome of these events (if those were, indeed, the former Moonshadow Pack’s pups).
Namely, the roaming male appeared to have been around for a while and he might have found the recently widowed female relatively quickly (before she dispersed due to loss of mate) even if there had been no pups holding her to the territory.
The pup presence unlikely influenced the new male’s decision to stay because an available mate and an available home should have been factors sufficient to sway his intent.
However, the area is known for a high pack turnover some of which is also through strife and aggressive shifting of territory boundaries (usurpation of portions of territories or even whole ranges) and the pups would have possibly contributed to defending the existing territory for the purposes of reproducing in future (since 2023).
It is not known (to me) how many of the 2021 pups remained with the group because they could have as well dispersed in 2022 contributing very little to the breeding pair’s interests.
Paradise Pack
Paradise Pack is one of those cases where the VWP team surmised they would not endure due to high pup mortality yet they have.
This is highly interesting because pup mortality is one of the top factors causing pack dissolution especially when the pack only consists of the breeding pair.
Paradise Pack’s story is somewhat similar to Lightfoot Pack’s story (see above) – both packs have claimed territories that might not be optimal (if pup survival rates are any indicator) and their reproductive success is low but they endure and the pair does not break up.
It prompts the question – could there be any additional considerations that wolves use to make the final decision (to remain together or not; to stay in the suboptimal territory or not?).
Can they anticipate the future?
Namely, can they estimate their actual compatibility or the potential of their chosen home despite the yearly variables that can be stochastic or non-permanent?
This could indicate that wolves do not react merely to the immediate circumstance and they do not make ‘rush’ decisions (i.e., decisions based on one year’s success or failure, decisions based on hormones, emotions etc.).
Moreover, it would be understandable that wolves could evaluate probabilities based on past experience, e.g., in situations where the breeding pair has been together for a long time and one or two years of ill-fortunes does not cancel out their common experience and their knowledge of one another’s qualities.
Lightfoot Pack had such prior experience (they had been together since before 2017) but Paradise Pack was newly formed.
Paradise Pack was established by the breeding pair in the winter of 2020 (during the breeding season) and they produced pups in the spring of 2020.
Unfortunately, none of the pups survived and they even appeared to have deceased by July.
Such early pup mortality often leads to pack/pair dissolution (see above, e.g., Huron’s Pack case) and I suppose the pair/pack is less likely to dissolve if the pups have survived at least until the ‘officially harsh’ period between the exhaustion of ungulate fawn prey and the winter-induced weakening of potential kills.
Late summer, fall, early winter can be a time when it is, generally, hard to survive and perhaps pup mortality during this period is less suggestive (to the wolves) of their own qualities, abilities, compatibility, territory suitability etc.
While mortality prior to this ‘official’ period might be taken under different type of considerations by the wolves.
As indicated above, the Paradise Pack did not dissolve and they remained together and bred once more in 2021.
In 2021, 2 of their produced 5 pups survived which is not amazing but adequate.
Perhaps this higher survival was attributable to the good fortunes (at the expense of Huron Pack’s bad fortunes) – the Paradise Pack managed to acquire more territory when the Huron Pack dissolved in summer (possibly due to pup loss, see above).
It could have been of assistance to this nifty pack which also learned to feed on blueberries and did so extensively.
Their territory of 2020 had been very small and carved out between 4 four other packs (with high overlap).
Perhaps they did not give up in 2020 because they sensed vulnerability in some of the neighbours (the territorial situation was not very stable in their area with frequent trespassings and fatal conflicts, see above, e.g., Huron Pack).
However, the overlap with other packs is still high according to the map of 2022/23.
While the takeover of Huron Pack’s range was not an aggressive one, the survival of the PP’s pups would have aided it because there were other ready claimants around and one of these potential candidates was the far larger Bluebird Lake Pack (see above).
Two extra pack members could have facilitated the incorporation of the territory into PP’s range.
Tamarack Pack
Tamarack Pack resided in an area of frequent pack turnovers and their fates were not altogether very fortunate.
It is not known (due to lack of trail camera positions on their range) if they produced pups in 2020 but in the winter of 2020/21 the pack consisted of a breeding pair.
They persisted into 2021 when they had produced at least 1 pup but the pup did not survive to winter.
The pup’s death was followed by a takeover of the former Tamarack Pack’s territory by the Stub-Tail Pack.
The pair still remained together (note the resilience and determination in these packs south of Kabetogama Peninsula) and they occupied a range north of their former home.
However, in March, 2022, the breeding female was shot illegally and the breeding male became a lone wolf abandoning the territory.
The experiences of the Paradise Pack and also of the Lightfoot Pack (see above) shows that the poor pup survival and even displacement could not have necessarily resulted in the disappearance of this pack.
But there is a limit to wolves’ resilience.
Still, it is also interesting to consider why the breeding male did not remain on the territory.
Some of these stories have the surviving breeder stay and wait for new breeding opportunities that are not scarce with local dispersers and roamers from outside of the VNP.
Perhaps the new territory that he had been forced to inhabit after the takeover by the Stub-Tail Pack was not adequate to hold on to.
Or else, a factor could have been the loss of his mate after the breeding season and after the most intense dispersal period.
He might not have felt motivated enough or hopeful enough to expect a new partner showing up.
Mourning should not be dismissed.
For example, it is possible that if the mate is lost during the breeding season or shortly before it, the survivor experiences grief and anxiety in a different manner than these experiences affect the wolf prior or after the hormonal rush.
The hormonal state might contribute to overcoming depression and making different decisions while losing a mate during the period when the breeding is over and when perhaps other hormones are at play (parental ones) adding to the perception of loss (loss of the mate and loss of potential pups that could have had if the mate had not died), the individual is more prone to ‘give up’ or to evaluate the circumstance as less satisfactory (not being content with the current territory etc.).
It would be curious to analyze whether such decisions differ statistically during periods of particular hormonal concentrations.
Whiskey Point Pack
It is possible that the very existence of the Whiskey Point Pack is owed to the poor pup survival of the former Sand Point Pack in 2019.
The Sand Point Pack had held its territory since perhaps 2015 (but definitely since at least 2018).
Their breeding history is not known until 2019 when the SPP pair was observed alone (no pups) on the trail cameras.
It is likely they would have had pups before otherwise it would have been exceedingly difficult for a pair to persist for 4 years.
However, pup mortality (zero pups in winter) of 2019/20 resulted in vulnerability that was exploited by a roaming pair in early 2020.
On this occasion, it is likely that poor pup survival not only during the prior season but potentially before (lack of subordinates retained in the pack) contributed to the loss of their territory to the new pair of wolves which then formed the Whiskey Point Pack.
This takeover was, nevertheless, very gradual because both pairs had been observed intermittently on the same range and the last observation of the SPP was even dated in December, 2020.
This means that they could have had a chance to reproduce in 2020 which they apparently either did not, or their pups did not survive, once again.
No further comments have been given.
From the winter 2020/21 (Dec – Jan), only the new pair was observed and the old pair disappeared (unknown fates).
It is perhaps worth noting that when the pairs fail to reproduce, the takeovers of their territories appear rather peaceful as if they are themselves relatively ready to give up.
This makes me wonder (especially on occasions when both individuals of the pair survive and when they disappear together indicating that they might not have necessarily split up but that they have abandoned their former range) whether there is such thing as compatibility of the particular breeders (or pack) with the specific home range.
In many species (e.g., white storks) the pair can either look for a new partner if they have trouble reproducing, or they can look for a new home/nest (if they consider the territory/range to be the problematic factor reducing their reproductive success).
These assessments are probably partly genetic because there can be statistical trends derived from these decisions (to stay or to go) (e.g., see Białas, J.T. et al., 2023).
These statistical trends are likely the result of evolutionary adaptations where the species has over millennia adjusted its behaviour to improve reproductive performance.
But the strange nature of these pack stories where some pairs/packs tend to break up and/or leave following (repeated) reproductive failure but others do not (even though as unsuccessful), and the fact that not all of the packs/pairs that remain, do so on optimal ranges, makes me wonder what determines these decisions in wolves.
I have already touched upon the issue of hormonal cycles and timing.
However, there might be some other aspects that make the wolves feel like ‘this is my home’ in some places but not others.
These factors of compatibility could be based on prior experience (natal range, experience during dispersal), or they could be based on the memories that the wolves have had since their arrival (traumatic – e.g., intraspecific strife; or pleasant – encountering one’s mate, having successful first kills, other types of enjoyable or empowering situations).
Other determinants might affect the pair’s (pack’s) perception of whether it is worth staying.
It could be, as mentioned, emotional attachment or perhaps some physical qualities that make the habitat more delightful, comfortable, easily used by the individual.
For example, larger wolves might have trouble settling in an area which is very densely overgrown with shrubs etc.
Smaller wolves could benefit from improved thermal regulation (e.g., shelter from winds).
Similarly, the colour of the wolves could be of interest (camouflage but also thermoregulation).
In any case, I will proceed with the Whiskey Point Pack’s story as they permanently established on the Sand Point Pack’s former range in the year of 2021.
The new pair was successful and they produced at least 4 pups (pup count observed in autumn, 2021).
It seems that all the 4 pups survived until winter.
I believe this is curious.
If the new pair was successful at breeding and pup survival was high, it denounces assumptions that the lack of success in the former Sand Point Pack was due to poor territory quality.
However, if the VWP team is correct in their ‘guesstimates’ that the SPP had been around since 2015, it is unlikely that the pair itself was incompatible.
They also appeared to attempt to stay together despite the pup mortality in 2019 and lack of reproduction or pup mortality in 2020.
This is suggestive of other factors having been at play and such factors could be incompatibility with the particular territory.
The SSP had lived on their range for many years but it could have changed or it could not have been supportive of older wolves (if they were about 6 or more years old).
Reproductive senescence has been observed in wolves (starting from age 4 – 5) and predatory senescence can start at the age of 3 already.
Both factors could have affected the SPP breeding pair’s success.
Predatory senescence would result in complications using the territory.
It would be interesting to see if other wolf families also have a harder time on this particular range as they are getting older.
There could be habitats (terrains, prey species communities) that are not very senior-friendly resulting in decreasing fitness.
However, pack size should be accounted for because senescent individual lower performance could be compensated by peak age individuals.
In this case, SPP did not have any subordinates that could have further complicated their effort to stay at their old home.
Windsong Pack
Windsong Pack’s story is probably illustrative of why it is important to have bred successfully and to have also retained some of the older offspring in their natal range (not dispersed during winter).
Windsong Pack lost its former breeding male to lethal management (depredation retaliations) in the summer of 2020.
His son took over and the pack was renamed, as well (Fawn Crick Pack -> Windsong Pack).
The son and his new mate also adopted at least one of the pups who survived to winter 2020/21.
It has not been stated how many pups there had been and what happened with the rest of the former pack members.
The change in breeders has been termed a turnover and the entire process has been referred to as pack dissolution (probably because none of the former breeders maintained their positions contrary to some situations where the surviving breeder, in this case, the female, would have found a new mate and continued the pack’s tradition).
If this occurred during summer, pups might have been too young to follow their dispersing mother and it is possible that the pup observed with the new breeding male and his mate was the only survivor.
It is also possible that not even this pup would have survived without his older brother.
The pup helped its brother through the new breeding season and two pups survived in 2021 leading to a pack size of 5 wolves in the winter of 2021/22.
Both pups were removed due to livestock depredation in May, 2022.
The Windsong Pack persisted, however, at least to 2022/23 (according to VWP Twitter post map).
This pack is thereby interesting with its subordinates not dispersing too early which even led to the inheritance of the original FCP’s territory by FCP breeding pair’s son.
I wonder what determined the direction of the turnover because, as I have mentioned before, the breeding female could have retained the breeding position and she could have found a mate by the following breeding season.
It could have been the chance of having met a mate already in summer that determined her son would be the future breeder in the pack.
Their territory size was not that much larger than in other packs and I wonder if there could be a genetic predisposition in some packs for its older sons and daughters to stay behind.
This is, of course, also determined by other factors such as food availability, population density etc.
If the two 2021 spring pups also got into trouble due to depredation, this could signal either difficulties acquiring wild prey or pre-dispersal type of behaviour where the subadults roam more widely and encounter novel situations.
As far as I understand, however, there is a ranch nearby which is essentially found in the ‘middle of the forest’ fragmenting the wolf families’ range.
Their father was removed due to depredation in summer.
It has not been stated if he actually killed livestock in summer or spring, or if the retaliation occurred in summer.
But if he did kill livestock in summer, this would indicate the ranch being within their range and not marginal because I would expect that he was keeping closer to the den site and that their den site was not on territorial boundaries.
Thus, the fact that the two pups of 2021 depredated does not necessarily mean that they were ranging far outside of their territory and they, too, could have been of the ‘late-dispersing’ type if there is such in wolves.
Their territory cannot have been overabundant because this would have allowed for the mother (the former FCP breeding female) to stay behind and to raise her pups to maturity as a subordinate to her son and his mate.
Wiyapka Lake Pack
Wiyapka Lake Pack had very poor pup survival for 2 years (2019/20 and 2020/21) and in the autumn of 2020 this likely resulted in loss of portions of their territories to the Bluebird Lake Pack.
The Bluebird Lake Pack was 9 wolves strong at the time while the Wiyapka Lake Pack was only 3 wolves strong (the breeding pair and a subordinate born in 2019).
Their story evidences how important pup survival can be also with respect to retaining subordinates with a larger wolf pack nearby.
It is difficult to say whether successful pup production in 2020 would have helped them retain their whole range (although it might have motivated them to a greater extent to stand up to the challenge) but 2019 pups could have been experienced and strong aids in defending their territory.
In 2021, Wiyapka Lake Pack was more successful which is somewhat counterintuitive because they had lost parts of their territory yet for the first year after two poorer years they did well (3 surviving pups that survived until at least winter 2021/22).
Perhaps they did not lose the most significant parts of their range (important prey concentrations, den sites).
Perhaps the reduced need for patrolling the formerly relatively large range for a small pack saved them some energy which could be allocated to reproduction.
Wiyapka Lake Pack still existed in 2022/23 (according to the new map) and they apparently survived their two poor year crisis with losses that did not put their pack integrity to hazard.
Notes by Ieva 