Lessons from niche partitioning: rethinking human obligations in wolf coexistence
Coexistence among carnivores depends on niche partitioning—spatial, temporal, and trophic. Humans consistently refuse to apply the same logic to themselves, instead demanding that wolves alone adapt in a one-sided coexistence. Genuine coexistence requires humans to modify their behaviour, protect wolf sociality and culture, and create conditions that allow the wolves natural ecological and cultural processes to persist.
In writing about the ecology of feline and canine predators a few years ago, I attempted to navigate through some of the contentions that had arisen about the impact that wolves (Canis lupus) had after their reinstatement through human agency to Yellowstone National Park (1). A common disputation was whether the wolves were having any effect on the recovery of aspen and willow through reducing herbivory by elk (Cervus canadensis)(and see this current spat ((2,3)). As an example, I highlighted a study on predator avoidance that had asserted that wolves had negligible impact on elk movements in the Northern Range, that elk may simply ignore the risk of wolf predation while navigating the landscape in search of winter forage (4). A separate study by this same group suggested that the cougar (Puma concolor) and not wolf, was the main predator influencing the movement of elk across the winter range of northern Yellowstone National Park (5).
Some of the observations in the latter study made sense, that the probability of elk predation by wolves decreased with the ruggedness of territory, as they are cursorial predators that kill mainly in flat, open areas at morning and dusk, whereas predation by cougars increased with ruggedness as they are spot-and-stalk/ambush predators. The authors had concluded that elk avoided open areas during the day and forested areas during the night in a pattern that fitted avoidance of the predatory behaviour of the wolf and cougar by instead occupying the temporally vacant hunting domain i.e. when wolves and cougar were respectively at rest and not hunting. However, their bold claim that male cougars, not wolves, exerted the most pressure on elk habitat selection was worthless when it was an inference from modelling data tucked away in supplementary information that was not part of the paper. Moreover, they had skipped over any explanation of why the results of this latter study in terms of the spatiotemporal effects of wolves were at odds with their previous study.
Yellowstone as a case study in predator coexistence
I had to admit back then that I was not aware there were cougar in Yellowstone, nor had I seen any study of trophic interaction in Yellowstone involving cougars, let alone cougar and wolf. After a 50-year absence, cougar reestablished a viable, year-round population in northern Yellowstone in the 1980s (6). This natural reestablishment occurred during a period of high elk abundance and the absence of wolf. An estimated 15 to 22 cougars existed within Yellowstone National Park immediately before wolves were reintroduced, that number increasing to an estimated 26–42 cougar after wolf establishment (7) while their range expanded eastward further into the Park (see Fig 2c in (8)). The cougar is an elusive animal, making it difficult to accurately count its population. A current estimation is being made using a methodology where remote camera traps are deployed in a predefined grid pattern to monitor wildlife, and from which preliminary results indicate a stable population since 2017 of up to 45 cougars inhabiting northern Yellowstone (6,7). Meanwhile the wolf population after reinstatement in 1995, when 41 wild wolves from Canada and northwest Montana were released in Yellowstone, reached a peak around 2003 and then fluctuated between 83 and 123 wolves since 2009 (9). Given the potential for intraguild killing between predators to occur when they compete for resources, such as the killing of coyotes (Canis latrans) by wolves (10) and the killing of foxes (Vulpes vulpes) by lynx (Lynx lynx)(11) then clearly wolves and cougars have found some way to coexist within the Park without killing each other.
That this is so comes from a new study that used camera traps across the northern part of the park as well as fitting both wolves and cougars with GPS collars. By comparing data collected between 1998 and 2005 and again from 2016 to 2024 (covering nearly 4,000 kill sites) the team observed a striking shift in prey choice (12). Both predators hunted elk less over time, a prey species that had once dominated their diets. While wolves increasingly moved toward larger prey such as bison (Bison bison) cougars began targeting smaller deer species more frequently, such as mule deer (Odocoileus hemionus) and white-tailed deer (Odocoileus virginianus). Considering the growing overlap between cougar and wolf habitats, the study found that in terms of wolf–cougar interactions nearly half of all encounters occurred at sites where cougars had made a kill. Thus wolves, aided by the sociality of their hunting in packs, were actively attracted to areas where cougars had made a kill and then stole and consumed the elk killed by the cougar, a process called kleptoparasitism. In contrast, cougars are lone hunters that actively avoided wolf kills and never kleptoparasitized wolf kills. Indeed, there was only one encounter where a wolf had killed the prey first. Switching to smaller prey species meant that cougars spent less time consuming the kill, so reducing the chance that wolves would discover the kill. In addition, cougars tended to avoid areas where wolves had made kills and stayed close to spatial refugia, escape terrain that had climbable trees or rugged terrain that provided essential escape opportunities such as rocky outcroppings. Of the 12 documented cougar deaths, two were caused by wolves finding cougars on elk kills in areas lacking escape terrain. None of the 90 documented wolf deaths were caused by cougars, most being due to natural causes or human actions.
Niche partitioning beyond Yellowstone
The two observations, prey switching and increased selection for rugged terrain by cougars when wolves were close, seem to be the means by which intraguild mortality was avoided and which allowed the wolves and cougars to coexist. The authors suggested that coexistence among apex carnivores may thus depend less on overall prey abundance and more on prey diversity spanning body size classes that facilitates niche partitioning, both spatial and dietary, and mechanisms to reduce interference competition, a direct, antagonistic interaction between wolves and cougars. I’ve written before on niche partitioning as a primary mechanism regulating coexistence, and its breakdown amongst apex and meso-carnivores – wolves, coyotes, bobcats (Lynx rufus) red foxes, gray foxes (Urocyon cinereoargenteus) fishers (Pekania pennanti) and American martens (Martes americana) - across a gradient of human disturbance in a forested area of the Great Lakes region in eastern America (13). Individual carnivores consumed more human food subsidies in disturbed landscapes, resulting in significant increases in trophic niche expansion and trophic niche overlap among species. This is known to increase interspecific competition leading to intraguild predation – there is a decoupling of predator-prey dynamics as the dominant species started killing the others so that the state of coexistence no longer existed.
Niche partitioning is not something you may associate with the nature depleted ecology of Britain, and thus it’s no surprise that in a trawl of recent papers, I found just two, both of which looked at the terrestrial/marine divide in foraging ecology of different seagull species. Thus, in a colony in Scotland, Lesser Black-backed gulls (Larus fuscus) had significantly larger foraging ranges than Herring (Larus argentatus) and Great Black-backed gulls (Larus marinus) indicating spatial segregation (14). Herring and Great Black-backed gulls strongly selected for landfill and coastal habitats, while Lesser Black-backed gulls also selected for these habitats, but primarily used agricultural areas. The other study focussed on the movements of Lesser Black-backed and Herring gulls from their multiple breeding colonies around the UK (15). Again, Lesser Black-backed gulls had significantly larger ranges than Herring gulls. During active nesting periods, the marine foraging ranges of Lesser Black-backed gulls were greater than terrestrial ranges, whereas the opposite was true for Herring gulls.
Of more interest for niche partitioning was a recent study in Bulgaria that used camera traps to survey the spatiotemporal activity patterns of carnivore species of different body sizes, testing expectations based on mesopredator release theory (16). I have written about mesopredator release in Britain, the evidence coming from the interaction between badgers (Meles meles) foxes and hedgehogs (Erinaceus europaeus) revealed after mass culling of badgers (17) but this study in Bulgaria was of a more complete carnivore guild assembly: brown bear (Ursus arctos), wolf, golden jackal (Canis aureus) red fox, badger (Meles meles) European wildcat (Felis silvestris) and stone (Martes foina) and pine martens (Martes martes) (16). It was found that brown bear and wolf avoided each other, wolf overlapped with jackal, and red fox avoided both wolves and jackal. There was little segregation amongst the smaller carnivores (badger, wildcat, and martens) but these avoided the bear, wolf and jackal. The study also noted that carnivore activity varied by season. Thus the brown bear was crepuscular– nocturnal in colder months (where crepuscular means before sunrise or after sunset) but cathemeral (active at irregular or sporadic intervals during both day and night) in warmer months; wolf and red fox were nocturnal in colder months, but cathemeral in warmer months; golden jackal and wildcat were cathemeral year‑round; badger was nocturnal in colder months, but crepuscular–nocturnal in warmer months; and the martens were nocturnal all year round. These results highlighted complex mechanisms for niche partitioning and coexistence among these carnivores.
Wolves, jackals, and expanding ranges in Europe
Unlike other European countries, the wolf was never entirely extirpated from Bulgaria (18). Similarly, the golden jackal has had a long presence in Bulgaria (19) and so they and wolves had plenty of time to work out how to avoid each other. In terms of size, the jackal is larger than a fox, but smaller than a wolf (20). Into the 20th century, though, the golden jackal started to break out of the boundaries of its long-term distribution in the Balkans and Dalmatia at the SE edge of Europe, and expanded towards northern and western Europe (see Fig 5 in (19)). It is now found in 29 European countries, having increased its total range by 46% since 2016 for an estimated population around 150,000 (21). One possible driver that enabled that expansion was the historical extirpation of wolves across Europe (22). This inference was derived from the observation that there were inverse patterns of relative abundance and distribution for both canid species at various spatial scales in seven out of eight cases of wolf re-colonization of jackal territories across Europe. Jackals disappeared or were displaced out or to the periphery of the newly established wolf home-range.
This seems supported by a study in NE Italy, which was one of the first areas in Western Europe to experience golden jackal colonization from SE Europe, followed by an internal wolf recolonization in a west-to-east direction into the Eastern Italian Alps (23). Using systematic and opportunistic data collected over 11 years (2013–2023) to assess the presence of both species, the data showed that jackals preferred less rugged and more fragmented areas typical of the lowland - which fits with their diet of mostly small mammal (20) - whereas wolves preferred rugged terrain with extensive forested patches in the Alpine range. At a local scale, golden jackal presence was negatively related to wolf presence, suggesting a potential top-down effect of the wolf on golden jackal colonization dynamics, but the authors recognised that wolf influence may vary in intensity depending on environmental context, with a weaker effect in areas of higher human pressure, such as the lowlands. A literature search on studies that investigated the spatial, temporal or trophic niche partitioning between pairs of carnivore species found that the most frequently studied niche dimension was the trophic (use of different food resources) followed by the spatial and temporal dimensions (24). Few of those studies reported an effect of human disturbance used as an experimental design, but from those that did, the authors of the literature search found evidence that human disturbance can affect all three niche dimensions, impeding and unbalancing niche partitioning and which increased intraguild competition.
I find intriguing the widespread dispersal of jackal into new territories in Europe while at the same time wolves were recolonising their former range (25). It would and will be a challenging time for many extant species, both prey and predators, to come to terms with the newly arrived presence of these two carnivores. New behaviours will develop around the partitioning that prey will adopt to avoid predation. Other members of the predator guild will seek to avoid direct competition for the same limited resources, since the superior competitor may eventually drive the other to local extinction or force it to adapt to a new niche, as the wolf has with the cougar in Yellowstone (see above) and as the wolf does to the jackal. It is niche partitioning by trophic, temporal, or spatial means that ensures coexistence. While there is ample evidence of the spatiotemporal strategies that wolves have to avoid humans so that they adapt to human presence by acting as ghosts in the landscape (see references in (26-28) and (29)). I despair that the human species is so blind to its own obligation to keep out of the way of wolves. I'm finding it difficult to persist in wading through the arguments of why humans should have a monopolistic opinion on the very existence of wild nature when we are such latecomers in the 3.5 billion years of the existence of life (30).
Lessons for human–wolf coexistence
It has made me consider what lessons niche partitioning has for the human species, considering how the relationship with wolves becomes increasingly more fractious as political will outpaces ecological understanding (31-33). As I have documented, the alleged wolf expert, Luigi Boitani, takes an entirely anthropocentric view of killing for coexistence, a lethal control of wolves to limit their populations, and with a complete disregard that the human species make any allowances, change its behaviour, cede spatially, but expects the wolf to bear the burden of death in a one-sided aim for coexistence (34,35). Equally as obstructive are those that believe their special insight on coexistence lays in giving prominence to victimhood and grievances in the decision-making process, but which is described as a pragmatic/dynamic coexistence discourse giving social legitimacy for a socially sustainable coexistence, and which elevates rural people in Europe almost to the level of indigeneity – as if there are any people now in Europe living a truly autochthonous existence (36) - while claiming that wolf protectionism promotes a “narrow interpretation of what nature and coexistence should be” (37-39).
Clearly, in this socio-political interpretation of what “nature and coexistence should be”, it is the wolves that are expected to adapt, and not the human species. It would seem that wolves can adapt to human perturbations, as is shown by a recent experimental study in a region of Italy where wolf colonization of human-dominated landscapes began earliest and has progressed furthest (40). Wolves were confronted across a wide urbanization gradient with novel objects (plastic children's toys of varying shapes and colours) either attached to trees or on top of rods, as well as separately human voices or bird song, their reactions being monitored by motion-sensitive video camera traps. Overall, the likelihood of a fearful response of wolves to the toys was low, and did not change for wilder wolves when presented with a second toy. However, wolves from more urbanized areas showed heightened caution when the object changed. This is interesting, as it suggests wolves fear of interacting with artificial objects/new things (neophobia) is a learned behaviour derived from direct experiences.
By contrast, playbacks of human voices elicited strong fear responses, more so than bird sounds, like running away or avoidance/change of direction, regardless of urbanization. Wolves in groups reacted less fearfully than wolves alone, suggesting that the sociality of wolf packs buffered fear. In both tests, toys and human voices, wolves showed a reduced likelihood of exhibiting a fearful response over repeated exposures, the authors noting that a generalized reduction in fearfulness could be maladaptive for the existence of the wolf. While this last point is important, it should be considered that these fear-inducing events were contrived and did not result in any penalty to the wolves, thus the apparent re-adjustment in their behavioural responses was explicable.
A missing piece: animal culture
There have been attempts to define the niche of the human species, its manifestation as a superpredator (41,42) how it shapes wildlife behavioural responses (43) and how the ecological concept of niche is not solely a matter of biological traits when in human-influenced ecosystems a species' niche is purportedly shaped by a variety of social factors, including its historical, legal, economic, political and cultural status, as well as the diverse claims of knowledge and justice that these social factors inform – it is a socio-ecological niche (44). I still don’t see in all this any concessions on the part of the human species, just ultimate imperiousness in determining how other species get to exist. That this is entirely dangerous to the future of non-human species past any limit on their population that we tolerate, is in the recognition that at stake is animal culture and maintenance of the diversity of that culture.
Culture was previously considered to be unique to humans, but knowledge of sociality, socially learnt behaviour and information in many animal species that is shared within a group also indicates the prevalence of a culture, that learnt behaviour and information is being transmitted from parent to offspring; from older to younger, often unrelated, individuals; and horizontal transmission between peers of the same generation (45). Social learning can be a cause of ecological niche separation, as the activities where social learning and culture operate include development of foraging behaviour in many species; migration in group-living species; and communication by vocalization, facial expression, and body stance. This goes beyond just considering habitat suitability and maintaining population density at basic effective population size in supporting the long-term persistence of viable, natural populations of wild species. It requires us to ensure environments and circumstances that maintain cultural diversity through supporting the capacity for cultural transmission to emerge in these populations, where animals have strong social networks and be able to communicate with one another, and which allow culture to be transmitted (46). This adds another dimension to the intrinsic value of wild species, not in our valuation, but for the animals themselves.
A recent paper arguing for an inclusion of sociality as a criterion in grey wolf recovery plans and policies in America, described wolves as having a fast-paced life history (early sexual maturation, high fecundity, short lifespan); of being territorial, which increases survival of offspring; and possessing rich social lives with distinct cultures that are vertically transmitted (parent to offspring) through social learning (47). Packs can be composed of a mixture of maturing offspring and unrelated wolves, each with specific roles in their social group to support pack activities as well as provide learning experience for cubs as they develop (see Box 1 in (47)). The authors asserted that recovery is more than population viability and should include sociality, which they say is the foundation for eco-evolutionary potential and ecological function. They indicated the need in policy formulation to collect evidence for social behaviour and animal culture, and the transgenerational transmission which they equated to cultural inheritance. In addition, there needed to be an assessment of the degree to which behaviour and culture were associated with rates of survival, reproduction, and dispersal, as well as identifying variation in social behaviour and culture between social groups so that it can be protected as part of ensuring cultural diversity.
The authors acknowledged that the space required for natural behavioural ecology to persist for a social species like the wolf is larger than protected spaces, and thus gave guidance on how plans for the wider landscape should incorporate the social group as the most fundamental unit of recovery. This included limiting anthropogenically driven mortalities that extended past intentional wolf mortality (i.e., hunting or harvest or poaching) to include also vehicular collisions through signage, speed limit restrictions, or green bridges (wildlife crossings); implementing a buffer zone design on the basis of the landscape used by wolves when they traverse political boundaries into neighbouring jurisdictions; banning killing during the breeding season and no killing of dispersers; maintaining full protection of wolves regardless of where wolves occur, and with increased legal consequences for wolf poaching; and the study of the consequences of wolf removal as a function of pack size to identify whether it was possible to reduce the population without impeding social function.
The actions above in ensuring persistence and diversity of wolf culture can be viewed in the context of partitioning, separating the influence of humans from wolves, in a form of niche partitioning that accepts that we have to make the efforts to coexist, rather than the wolf alone bearing the burden, a one-sided coexistence. Another example of our modifying our actions comes from the recommendations in a paper that looked at the reasons why aggressive human-carnivore encounters occurred in two case studies of young wolves in Poland. The wolves were 13–14 months old during attacks and 8–9 months old when they appeared in the vicinity of households (48). Positive food conditioning was a factor in both cases, leading the authors to argue for proper management of waste from households, food markets, restaurants, camp grounds, and livestock breeding farms, as well as a ban on baiting for hunting or photographic purposes. The recommendations, however, went wider, arguing that detailed rigorous analyses of factors that led to every conflict situation would help to develop good practices for various human activities conducted in areas inhabited by wolves, such as forestry, farming, tourism, recreation, hunting, wildlife watching and photography. In the longer term, they said there should be education of the wider public on rules of coexistence with large carnivores, including science-based knowledge on wolf ecology, explaining the differences between natural and unnatural behaviour, the proper reaction in cases of unintentional or deliberate actions leading to wolf habituation, as well as guidance on how to behave in close encounters with wolves.
Toward a human niche that allows coexistence
I was struck by an article in the December issue of Argus, the magazine of De Faunabescherming, a Dutch fauna protection foundation (49). The article was by Wolves in Utrecht, and it prompted some of my first thoughts about applying niche partitioning to human-wolf coexistence. Wolves in Utrecht have consistently questioned the picture that has been painted in the Netherlands of life-threatening wolves that attack women, children, and dogs, out of nowhere and without provocation, having doubts about the incidents that have been alleged, and seeking to better understand what works in terms of how to eliminate their possibility (50). The first proposal amongst the seven in the article was to create genuine quiet areas by closing off paths in core wolf areas, especially during the cub season, and giving animals the chance to retreat (51). I see this proposal having universality in an approach of partitioning for coexistence, since I have noted before that Denmark saw an imperative in protecting established breeding pairs and their young offspring, when a Danish municipality introduced a ban on public access from April to October to the wolf breeding area in the Klelund Plantation near Hovborg in South Jutland, and the Danish Environmental Protection Agency advised the public against actively searching for the wolf cubs on the days when a military training area in West Jutland was open (52).
The turmoil in the perception of the return of wolves to the Netherlands is especially febrile (34,35). It will need more space to do justice to all the proposals that Wolves in Utrecht put forward in that article and elsewhere, as it does to what sensible scientific approaches that have been advanced to understand in the Netherlands how to separate humans and wolves in space and/or time (53,54).
Mark Fisher 19 March 2026
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url:www.self-willed-land.org.uk/articles/niche.htm
www.self-willed-land.org.uk mark.fisher@self-willed-land.org.uk