Hunting by humans never constitutes compensatory mortality

For a long time research studies dedicated to many wild species were largely aimed at determining ‘sustainable harvest quotas’, i.e., the amount of individuals that can be taken lethally from the population in order to maintain the population’s viability.

Through this perspective, terms ‘additive or compensatory’ mortality have been introduced where mortality by human hunting should be compensatory and not additive in that it substitutes natural mortality but does not complements it.

If hunting quotas exceed natural mortality levels, they are viewed as additive mortality factors, i.e., they add to the natural mortality.

However, the paradigm has been changing and population numbers alone are no longer regarded as an acceptable estimate of population health and sustainability.

Of course, ethics must also be considered and many studies lead to conclusion that lethal control is not necessary and not effective.

Many of these issues I have already addressed in the article, ‘Can You Tell the Difference? Opposing Lethal Wolf Management’.

This post will be dedicated to wolves, as well.

I will attempt to discuss how hunting by humans is never compensatory.

The article that I previously linked to provides insight into how hunting by humans can disrupt wolf social structures which results in reduced efficiency by wolves to perform their ecosystem services also in increased stress levels that, in the long term, can detrimentally affect population’s fitness (compromised immunity).

This means that even if the population is ‘successfully’ maintained as 200 wolves per area (e.g., per a country), these 200 wolves after (during) the years of harvest might not be as healthy as 200 wolves before the years of harvest (without harvest) and, hence, while 200 wolves still reside in the area, they constitute a deteriorating population which might collapse more easily due to some external influences (e.g., extreme weather events, prey density fluctuations, climate change etc.).

It is also possible that we have not yet understood some of the influences of legal harvesting of wolves.

For example, in my post, ‘Age structure effects in harvested wolf populations – indicator of additive impact?‘ I have looked at the age structuring in natural and hunted wolf populations where in natural populations there is a tendency for age groups to even out over time while hunted wolf populations exhibit age structures skewed toward younger individuals.

This can lead to consequences that might alter the population’s demographic performance in the long term.

In my view, it is highly possible that hunting is an additive mortality factor even if we perceive it as ‘population numbers’ alone and I will attempt to develop on several reasons.

One of such reasons could be the high stress levels mentioned before that are known to affect reproductive success in some species (likely, in wolves, as well).

While wolves appear to be able to ‘replenish’ their numbers through increased reproduction and immigration into population vacancies, I think that at some point this turnover would not be sustainable because the local population health will deteriorate and wolves will show reduced ability to produce enough pups to ensure adequate hunting and territory defense rates.

Thus, the local population will not be healthy and immigrants from other populations (especially, if those populations are hunted, as well) might not be able to revive it anymore.

If wolves are hunted in optimal habitats and the immigration occurs from suboptimal (former sink) habitats (not from other optimal habitats), the population will not be capable of producing enough individuals to populate the ‘sinks’ that later return dispersers to replace shot or trapped breeders.

On a side note, it might be useful to study whether populations suffering natural mortality and losing their breeders/pack members to natural mortality factors demonstrate equal stress levels to those suffering human-caused mortality.

For example, most of the natural mortality factors are not sudden and wolves (as a species that is famed for its selective potential to detect poor health status in others) might be able to prepare for the loss.

The loss would thus be more gradual – wolves would be aware of their situation and its risks.

Meanwhile, mortality caused by hunting is often sudden and unexpected affecting also healthy social groups and healthy individuals.

Sudden loss bears the additional shock effect which could raise the stress levels even further.

(It could be compared to losing one’s grandpa who had been suffering ill health for a longer while already to losing one’s strong, relatively young father who could have withstood disease and strife.)

There are other factors that might reduce the stress experienced upon natural losses and that will be touched upon later in the text.

For example, in wolf populations natural mortality is often accompanied with immediate breeding position or territory takeovers.

If the former breeder is displaced by a new breeder and suffers mortality due to a conflict or expulsion from the pack, the new breeder is often accepted by the old pack and the pack gets to remain together and to reduce its cohesion only slightly compared to what they would have experienced on occasion of complete dissolution.

Similarly, reproduction can continue with the new breeder taking care of the old breeder’s offspring or producing his/her own offspring that very year without skipping a season.

This can lead to continued tenancy by the pack on their territory because they have enough pack members to defend it and their social life is not threatened by the lack of reproductive effort which appears to be a significant ‘binding drive’ keeping packs together.

The ability to remain with the known pack members, to remain in their old home and to have a new litter of pups soon enough could compensate for the stress caused by the turnover and by the loss of their beloved pack member.

It would be useful to research these impacts of natural vs. human-caused mortality facts and to also assess whether packs remain together more often, produce pups in the following breeding season and are otherwise less affected in their traditions, social cohesion etc. upon losing pack members to natural mortality.

In this post, I have used the term ‘additive impact’ outside of the common paradigm where additive/compensatory are usually applied to refer to wolf numbers and not to other types of impact and processes.

Personally, I believe that the paradigm needs to change and we should regard the consequences of hunting as additive if they disrupt significant natural processes.

Therefore, hunting by humans can never be compensatory because humans are not integrated within the natural ecosystems and they cannot substitute natural mortality factors for reasons that I am going to discuss below.

There are three main deficiencies in human hunting activity that makes it a poor substitute for natural mortality:

1. Humans do not kill individuals that would have died naturally, nor they apply selectivity that is commonly applied in nature (I will not address this in great detail because it is known that hunters do not aspire to kill weak, sick, old etc. wolves but they are rather on the lookout for trophies, i.e., big wolves with nice fur);
2. Hunters are not an integral part of the ecosystem and thus hunting by humans does not fall into the greater ‘logic’ or pattern of natural processes that lead to mortality and disruption of packs (mortality caused by human hunters could be described at best as stochastic);
3. Hunting by humans does not lead to improved health of the wolf population through evolved adaptations to the natural environment.

Natural mortality ‘benefits’

While the benefits of natural mortality (vs. human-caused mortality) are real, I have used quotation marks because any mortality should be regarded as a sad event and the benefits are perhaps a blessing that gives a meaning to our loss.

Mortality can contribute to the health of populations and species as well as their evolution and local adaptations.

Wolves themselves take lives to improve lives (see ‘Research Studies for the Guiding of Wolf Policies’ section ‘Ecosystem roles’ and ‘Grey Wolf (Canis lupus) – a Keystone, Umbrella species‘).

Any mortality can create a brief opening for resources or breeding positions, and this also includes mortality caused by humans.

However, humans do not live in the natural ecosystems and their hunting efforts come from the outside.

Therefore, mortality by humans is more stochastic (resembling natural accidents) and less potentially beneficial than natural mortality.

Natural mortality can be beneficial because animals learn to adapt in order to avoid death and to improve survival.

But in order to do so the mortality factors have to represent some natural states or processes that are integrated in the wolf’s life and that the wolf can effectively respond to.

This is, in fact, where human-caused mortality differs from accidental mortality.

Unlike natural mortality, it does not allow for the species to adapt to its environment through attempting to improve foraging/defense strategies, through selection of genes and traits, through developing immunity etc. (because mortality is not caused by any factors inherent to the wolf’s environment).

However, unlike deaths by accident, human-caused mortality can be predictable (it occurs in specific time and in specific areas).

Wolves can and have to adapt to mortality variables introduced into their lives by humans – they learn to avoid certain places during certain times, they change their resource use patterns, they select habitats further from hunting threat etc.).

This does not truly benefit the wolves as a species – it does not help them to become stronger and healthier with respect to pathogens, climate, habitat selection, resource selection, foraging strategies, defense strategies, social cooperation etc.

Human-caused mortality is additive also in this respect – it demands for wolves to account for considerations that their survival depends on -but not their interests as a species and as a population.

It would be of interest for wolves, e.g., to learn hunting new prey species in order to better exploit the resources of their specific habitats or to develop immunity against certain pathogens, or to select partners that can help produce offspring with higher chances at survival (maintaining these important genes active in the population pool) etc.

It is of little immediate interest for wolves to allocate cognitive and energy resources to track human activity because humans do not truly belong in the ecosystems that the wolf has to inhabit and to evolve in accordance with and to adapt to.

It is a strain that is very unlikely to give rise to wolves that are more efficient at their job in the nature.

The non-integral character of hunting by humans also leads to differential timing and consequences of the mortality that is induced.

Natural mortality follows a certain ‘logic’ that is universal but also very specific to the locality and to the pack that it affects as well as its immediate habitat.

‘Deaths by nature’, in my view, create incidents of turnover that are more contiguous, less disruptive and that, as discussed above, have a higher likelihood to maintain pack cohesion, pack traditions, pack knowledge of the local range and the continuous provision of the ecosystem services by wolves.

Natural mortality (intraspecific strife and starvation but also, to a lesser degree, disease) frequently is not a sudden occurrence but rather it has a lead-in and a follow-up weaving the sad event into the unfolding drama of the entire ecosystem.

This can be seen when one considers specific examples of how starvation or intraspecific strife (in more or less natural populations, not populations that are disrupted due to lack of wild prey or other unnatural factors) proceeds to impact the lives of individuals and their packs.

Natural occurs under specific conditions that are closely linked to the local environment and the particular species group and population demographics.

For example, intraspecific strife can take lives when one group has grown large and it needs to split or to expand its territory while the neighbouring group has declined in numbers, e.g., due to poor pup survival.

In this specific scenario, it is possible that one group has flourished because it has sufficient resources on its range while the other has dwindled because it does not.

Thus, after the clash, there will no longer be a group that cannot sustain itself.

There will be one big group that has plenty of optimal habitat and now also some suboptimal habitat.

It will retain its group cohesion and traditions and pack structure and knowledge of the area (unlike in situations where hunting results in pack disruption that leads to new groups settling in the area that yet have to build on cohesion, traditions, dominance hierarchies, local experience etc.).

In a different scenario, the larger pack might split into two groups but the group that claims the less optimal habitat might also retain parts of the optimal habitat and have better chances than the original small group.

In any case, this takeover could be the result of subtle changes in the environment (e.g., changes in prey distribution or ecosystem productivity, or the actual range available for wolves in their territory) leading to a redistribution of resources that, once more, create sustainable homes.

It is easily seen that the death event was not sudden, nor it was contingent to the causalities and conditions of the particular ecosystem and population.

Its consequences, similarly, while lamentable lead to redistribution of resources that might give better chances to all.

Moreover, the state of disorganization during this takeover is very brief and the local ranges are still managed by wolves that are coordinated as a group and that have acquired a certain balance with the local resources.

It is very unlike a situation where one group is disrupted suddenly through the effects of hunting by humans and the neighbouring groups may not have sufficient numbers and resources to take over because the situation itself has not arisen out of the local resources and population demographics and it cannot be evened out and ‘made better’ by what is available currently in the area.

It does not fall into the ‘local logic of events’.

Similarly, when old breeders are killed by new claimants, the new prospective breeder is already there when the old breeder dies, and the reproductive activities can proceed immediately. 

Wolves are very Christian that way – they forgive their ‘enemy’ and often the new breeder might be accepted by the old group which leads to even less disruption. 

Of course, the old breeder does not always have to die. They can take subordinate position or disperse and start a new family with their great experience.

This post is dedicated to mortalities which is why I am discussing a more fatal scenario.

When wolves are hunted by humans, their deaths do not follow the internal causalities of the local population and they place an additional strain on the survivors who have to fill the vacancies that have not arisen out of natural circumstances – by applying themselves also beyond natural capacities.

Hunting by humans may favour floaters (lone wolves awaiting their opportunity) as opposed to inheritance of the resources by the existent packs.

However, if not accepted within the former pack, floaters might also find it difficult to maintain ranges in the former pack’s territory because the need to reproduce is very pressing in order to have a pack that can hunt efficiently and defend their kills and territories efficiently.

They do not yet have the knowledge of the ‘local scene’ but they have to produce pups at once and this can lead to malnourished adults as well as higher pup mortality.

Often several individuals from different packs are killed by humans which altogether changes the distribution of the packs and some vacancies might open that are too small or too large to be filled by a new pair.

The pack territories might even become ‘illogical’ with respect to the resource distribution.

For example, if several individuals have been killed from two neighbouring packs and one of the packs dissolves while the other remains together but has dwindled in numbers, two new pairs might move in pushing the surviving pack to the brink of their former territory.

Thus, three packs now exist where two packs once lived and the third pack might suffer even more if they experience territorial clashes with their other neighbours where previously they had a ‘bufferzone’ between them.

This bufferzone has to be utilized by the dwindling pack because they have been pushed out from their former range.

Three packs live where two packs once resided and while initially it might be good enough, as the packs grow to average sizes, the limitations in their ranges can push them to starve, to fight or to become infected with disease.

In natural populations, the first wolves who colonize an area likely settle in accordance with the distribution of prey and denning sites.

These territories can be rather stable regardless of who inherits them – the actual offspring or neighbours.

It is very important to maintain (at least approximately) the original ranges because they can provide everything that a wolf family needs.

If this natural inheritance process is disrupted and wolves that ‘move in’, do not have knowledge of the resource distribution, the new territories might not present enough prey during all seasons or proper, safe den sites.

Thus, while the population manages to recover its numbers (and perhaps even grows in density), it is possible that the wolves now live on ranges that are not optimal, that are not formed in accordance with the logics of the local ecosystem and the logics of the wolf, and that, consequently, hinder pup survival and compromise the health (as well as the reproductive success) by adults.

In order to rearrange, wolves have to come in conflict with other packs but with continued hunting pressure, packs might not be able to rearrange because none of them reaches a size that can claim a proper territory pushing out the extra individuals.

It is sometimes assumed that there are always floaters available to fill in the vacancies.

But if the local population has to provide individuals who replace lost breeders and form new packs constantly, the population cannot stabilize because packs cannot reach their average pack size that is necessary to maintain a sustainable range and to ensure that the breeders do not exert all energy in pup rearing activities.

This might even cause genetic bottlenecks on a larger scope.

If many local individuals have to remain nearby in order to replace lost breeders and to repopulate the area, dispersal might be reduced out of the population and into other populations limiting gene flow (and if not reduced due to dispersing members of disrupted packs, then altered in a non-beneficial manner).

If the hunted population is considered a source population, lack of available emigrants from the source population can lead to extinction of the sink population, as well.

At some point the populations might not even be able to rely on immigration from other populations and some vacancies might not become filled after the loss of local individuals to any type of mortality.

I find it noteworthy that many studies discussing the effects of population turnovers mention dispersal of the disrupted packs that have been pushed out by the new claimants or that being roaming because they have lost pack cohesion and they cannot sustain themselves as a cooperative unit.

It is strange because in stable wolf populations vacancies in neighbouring territories are often filled in by locals and many local wolves (e.g., subadults) do not disperse far but they wait around for an opening in breeding position or territory availability.

Therefore, I would have predicted that dispersal is actually reduced in hunted populations because a greater number of individuals should stay behind in order to repopulate the ranges.

However, it does not appear necessarily so and it seems that instead the residents emigrate and newcomers settle in (which is not beneficial because the knowledge of local resources and adaptations to them that result in a more harmonious balance with the ecosystem, are lost).

To me, this points at a crucial difference between hunting by humans and other mortality factors, including hostile mortality incidents such as intraspecific strife.

I hope to delve deeper in case studies, but it seems to me that wolves do not perceive even intraspecific strife as stressful and disruptive as hunting by humans perhaps because intraspecific strife, as mentioned before, can be predicted and avoided and, to some extent, regulated by wolves themselves while hunting by humans is a persistent factor that drives wolves out from the area (seeking more peaceful ranges?) rather than intrudes temporarily later allowing for wolves’ lives to stabilize and even out the impact.

It could be that wolves react differently to situations when other wolves have killed pack members and to situations when pack members have been killed by humans.

In the former case, if I am not mistaken, wolves feel more confident that they can once again take charge of the situation and they do not feel as if their home has been lost forever.

They more often remain behind and they reorganize and carry on.

In the latter case, wolves apparently are affected by greater stress that causes them to feel that ‘this is the end’ and to try and begin elsewhere – anew.

There is a slight possibility that the physiological stress response is not natural (does not resemble natural stress levels following natural mortality) and the wolves ‘overreact’ feeling less capable of persisting and re-establishing themselves.

This might not even be an accurate self-assessment on behalf of the wolves but rather a reaction resulting from abnormal hormonal charge.

I believe it is easily seen how natural mortality creates ‘a story’ which is continuous, logical, contiguous, which allows for fluent inheritance of resources, knowledge and traditions, which reduces stress experienced by the packs and which promotes cohesion and cooperation.

Packs do not become too large, nor they become too small.

Territories are of a size appropriate for a long-term family and the territories can provide all the necessary resources through all seasons (e.g., deer do not move out to winter yards leaving the wolves starved and den sites are central and not peripheral etc.).

Individuals can be retained in packs to ensure cooperative lifestyle essential to wolves or they can participate in the local dynamics, or they can disperse to ensure gene flow between different populations as well as the maintenance of populations dwelling in suboptimal habitats.

Hunting by humans is not a part of these stories, and the link between the past of the local population within its local ecosystem becomes severed from its future.

I believe that ‘compensatory harvest’ does not exist unless humans agree to leave their houses and reside in the forests alongside wolves and other species.

I also believe that wolf researchers should study the following matters:

1. Does natural mortality cause fewer pack disruptions and gaps in reproductive years than human-caused mortality (do packs stay together and reproduce in the following season at a greater rate)?
2. Does natural mortality cause lower stress responses in wild-living wolves than human-caused mortality does?
3. Is it likelier for natural mortality to result in takeovers that allow for the territories to retain their sustainable resource distribution and that allow for a greater number of locally-experienced individuals to become the new territory-holders (even if some pack members are replaced by newcomers)?
4. Are territories and pack sizes more stable in non-hunted populations and does this lead to lower overall mortality as well as a reduced turnover rate?
5. Are far-ranging dispersal rates lower in hunted populations? (This factor might be confounding because disrupted packs can disperse, as well, and perhaps it would be more efficient to study successful dispersal which results in recruitment into a new pack with its territory.)

Other questions might be asked and I think that it would be most interesting to research case studies of specific packs and their histories and to see how natural mortality events arise out of local conditions and whether they lead into new conditions that give a better chance for everyone.

***

Dispersal is a confounding factor in this that I yet find difficult to clearly integrate into the picture.

It seems that in cases where human-caused mortality is high or when it occurs outside of typical dispersal seasons, there might not be a sufficient number of floaters around to fill in the vacancies.

If too many vacancies emerge, the sheer number of floaters might not be sufficient to fill them and to provide some roaming individuals that are available upon ’emergencies’.

I think these effects are combined.

The packs try to produce pups and to recruit them in order to ensure territory maintenance, expansion and proper hunting groups (and perhaps also to satisfy wolf’s social demands that might not be based on any real and actual needs but they might be based on the evolutionary requirement to have one’s group).

If reproduction has failed and no subordinates exist (because the pack is new itself), the individual might be less likely to remain on the old territory and more likely to disperse.

However, this dispersal might be different from that of dispersal after having reached independence.

For example, wolves with former breeding experience might find it more difficult to approach other packs, to roam in their territories or to initially take on more subordinate roles.

Perhaps it might even be hard for such a wolf to start a family with a pack that has already existed under a different leadership (compared to starting a pack with a new mate who has not bred before).

While the young wolves (dispersing pups) might not be able to start a new pack or to take on a breeder’s role in an existing pack due to their immaturity.

If dispersal is shifted toward, for example, subadult (not sexually mature) individuals and former breeders, dispersal itself might not be able to secure the maintenance of the resident packs (by replacing lost breeders).

The ‘regular’ floaters (dispersing yearlings) might not be sufficient in numbers to both provide mating resources for the resident packs with vacancies and for the dispersing former pack members.

The lack of subadult age group might also lower the number of dispersing individuals.

***

Regarding the strange dispersal aspects and the reasons why packs appear less capable of retaining their territories after human-induced mortality (vs. natural mortality), some of the findings of this study could be of relevance (Oliynyk, R.T., 2023).

Even if we disregard the overall mortality increase (by 20% after the initiation of hunting), the study mentions that overall mortality more than doubled in females and almost tripled in yearlings in the hunted population.

Moreover, human-caused mortality more than quadrupled for females and juveniles while their natural mortality decreased greatly initially.

This means that natural mortality became replaced by human-caused mortality (and acquired higher rates than pre-hunting mortality).

This is significant because human-caused mortality may have different timing, targets and consequences than natural mortality.

Male mortality less than doubled and male natural mortality increased possibly due to territorial conflicts.

While this is a case study (Minnesota wolf population), the results could be attributable to other wolf populations.

It is important to understand how these trends alter wolf social structures, dispersal patterns and other aspects of wolf biology and ecology.

For example, in many situations the death of one breeder does not necessarily translate into the dissolution of the pack or loss of their territory if there is someone else to take over.

These ‘someones’ are frequently the other breeder or an older subordinate (yearling or 2-yeard-old son/daughter who has the fortune to find a mate).

There are at least two conditions under which the pack does not break up and the territory is still held by the former group:

• there is an individual of the former pack that is capable of breeding (sexually mature individual);
• there is an unrelated disperser (floater) that can replace the lost breeder or that can form a new partnership with the sexually mature subordinate.

If juvenile mortality triples, this limits the number of older subordinates that could take over the breeding position or that could assist in defending the territory against other claimants on behalf of the surviving parent until the parent finds a mate and produces a new litter of pups.

It can also limit the number of dispersers that are roaming and available as mate substitutes.

This is especially true if all of the nearby wolf populations are hunted (because dispersal often occurs as immigration) and such adjacent populations could be at greater risk.

If female mortality doubles, this could lead to lack of access to mates for males.

If the breeding male survives, he might be forced to roam to actually find a female.

It is interesting to determine who actually dispersers with such high mortality of juveniles.

Because territorial conflicts (resulting in increased male mortality) attest to dispersing individuals or pairs seeking to settle.

However, if there are fewer juveniles (due to tripled mortality), who are these roamers?

Are they immigrants from other wolf populations (but are those populations then not hunted)?

It appears to me that the lack of yearlings and older subordinates results in a situation where the potential breeders themselves are forced to disperse because otherwise they cannot encounter a mate.

Thus, the natural model of dispersing subadults and subordinate adults (who are, in some respects, ‘dispensable’ as the breeding pair keeps producing new litters of pups) is replaced by dispersing breeders or subordinates that would have been otherwise able to stay on their natal range and to inherit the territory and breeding status from their parents.

This causes a turnover in packs and their territories that is not characterized by retention of some of the ‘original settlers’ on their familiar range that they have adapted to (pack traditions).

The pack dissolution and abandonment of territories could be enhanced by increased pup mortality.

While pups are lost to hunting during winter (and not to other factors such as incompatibility and otherwise poor performance by the breeders, senescence or deterioration of the resources on the territory), the mechanisms responsible over making decisions to stay together and to stay at home could be instinctual and not always as conscious.

If the wolves are incapable of actively preventing these losses, the pup mortality might signal to them (on a physiological and psychological level – through ancient mechanisms) that there is something wrong with their pair bond or with their home range.

They might thereby seek to improve their fortunes by making the only active decisions they have been evolutionary adapted to make, i.e., to replace mate or territory.

As far as I gather, mate replacement is not as frequent as territory replacement (especially, under circumstance of potential scarcity of mates), and territory replacement could be the choice that the wolves make in order to attempt to resolve the ‘mystery’ of the high mortality of their pups.

This results in territory abandonment and migration of former breeders or even breeding pairs.

But there is nothing truly wrong with the territory itself (resource-wise) and it might be impossible for the wolves to resolve the problem if the nearby habitats are included in the hunting range, as well.

This depends on how the wolves perceive reproductive success (because in winter pups die as weaned subadults having reached a relative independence and because some of these lost pack members might not be known to have died – assumed dispersed).

These are just a few of the scenarios that might unfold under conditions of increased juvenile and female mortality.

Increased natural mortality in males, on the other hand, probably also occurs in winter coinciding with increased human-caused mortality in females.

Therefore, it becomes more likely that both breeders die during the same season and neither one of them can attempt to retain the breeding position in their former pack.

And if those are pairs rather than individuals who form the new social group of ‘floaters’, it might reduce the opportunity for the survivor to find a new mate even further.

***

When mortality or other unwanted phenomena strike natural populations, individuals attempt to adapt because they are used to regarding any events as integrated within cause-consequence network in which they are a part and therefore in which they can take certain actions to change the course of causality (to improve the chances).

For example, if birds fail to raise their brood, they can consider altering nest site or divorcing the former partner and pairing up with a new partner.

If there is resource scarcity, animals might attempt to broaden their niche or to expand their territories even if it involved territorial strife.

Animals are proactive and they likely perceive failures (especially, repeated failures) in a responsible manner (what can I do to prevent this from happening again?) even if sometimes it is beyond them to improve the situation.

I believe that hunted wolf populations might suffer despair because there is nothing that they can do in order to prevent the disaster from striking and the losses are ‘not logical’ – they do not follow from any of the patterns that wolves are familiar with (climate, prey availability, pup care, den site selection, home range selection, partner selection etc.).

Helplessness is one of the leading causes of depression in humans, and, for example, animals who are kept in laboratories or tiny cages, often exhibit frantic behaviour (attempts to change the situation without understanding how to achieve it) or suicidal behaviour.

Hunting does not remove individuals ‘who had it coming’ (I am not speaking in the moral sense here but in the natural sense, i.e., individuals who were old, diseased, who were not successful, who had suffered some adversity beyond their scope of changing the situation but not beyond their scope of understanding the reasons beyond their bad fortunes – e.g., weather events).

It is possible that wolves exhibit maladaptive behaviours in attempts to prevent future misfortunes.

One of the reasons human-caused mortality might be more disruptive could be that wolves attempt to adapt to it in a manner that is familiar to them – by changing their territories (dispersing) or by changing partner.

But it is not the territory, nor it is the partner that is at fault, and nothing becomes resolved unless the wolves, by chance, end up in a protected area.

I have mentioned some of this before but I wanted to add that through hunting wolves, we might even cause them to become maladaptive.

By ‘maladaptive’ I mean adapting to this huge mortality factor (human-caused mortality) in a manner that contradicts what is adaptive in the natural setting (and thereby what is adaptive for the species on a genetic, ecological and behavioural level).

We sometimes actually ‘punish’ wolves for being too successful (having produced too many pups, having colonized new ranges, having explored the lands that belong to them also, having followed their curiosity to discover new skills, new objects etc.).

If wolves have to alter their social life and reproductive activity due to human disturbance in situations when they have actually been doing well and their efforts have been fruitful, wolves do not get to enjoy the fruits of their success, nor they learn to manage the consequences of success (e.g., higher population density).

Where wolves should have expanded their territories and stabilized their social groups (thus, reaching lower densities and reducing pack turnover), having suffered this undeserved mortality, they react in a manner which is not consequential to their natural reality, i.e., they divorce, they emigrate, they break up packs, they fail to reproduce with the former partner and in the following breeding season etc.

Thus, wolves do not learn to adapt to the natural consequences of having done well and having grown in numbers (which is important in order to attain self-regulation, balance with the local ecosystem and knowledge of its resources and limitations).

Instead, they react by resuming ‘default’ (starting from the beginning).

Essentially, wolves cease to adapt to natural course of events and instead they adapt to human-imposed course of events, and these two courses are rarely compatible.