Eat or be eaten: Implications of potential exploitative competition between wolves and humans across predator-savvy and predator-naive deer populations.

Recolonization of predators to their former ranges is becoming increasingly prevalent. Such recolonization places predators among their prey once again; the latter having lived without predation (from such predators) for a considerable time. This renewed coexistence creates opportunities to explore predation ecology at both fundamental and applied levels. We used a paired experimental design to investigate white-tailed deer risk allocation in the Upper and Lower Peninsulas (UP and LP) in Michigan, USA. Wolves are functionally absent in the LP, while deer in the UP coexist with a re-established wolf population. We treated 15 sites each in UP and LP with wolf olfactory cues and observed deer vigilance, activity, and visitation rates at the interface of habitat covariates using remote cameras. Such a paired design across wolf versus no-wolf areas allowed us to examine indirect predation effects while accounting for confounding parameters such as the presence of other predators and human activity. While wolf urine had no effect across most metrics in both UP and LP, we observed differences in deer activity in areas with versus without wolves. Sites treated with wolf urine in the UP showed a reduction in crepuscular deer activity, compared to control/novel-scent treated sites. Furthermore, we observed a strong positive effect of vegetation cover on deer vigilance in these sites. This indicates that simulated predator cues likely affect deer vigilance more acutely in denser habitats, which presumably facilitates predation success. Such responses were however absent among deer in the LP that are presumably naïve toward wolf predation. Where human and non-human predators hunt shared prey, such as in Michigan, predators may constrain human hunting success by increasing deer vigilance. Hunters may avoid such exploitative competition by choosing hunting/bait sites located in open areas. Our results pertaining to fundamental predation ecology have strong applied implications that can promote human-predator coexistence.

Wolves alter the trajectory of forests by shaping the central place foraging behaviour of an ecosystem engineer.

Predators can directly and indirectly alter the foraging behaviour of prey through direct predation and the risk of predation, and in doing so, initiate indirect effects that influence myriad species and ecological processes. We describe how wolves indirectly alter the trajectory of forests by constraining the distance that beavers, a central place forager and prolific ecosystem engineer, forage from water. Specifically, we demonstrate that wolves wait in ambush and kill beavers on longer feeding trails than would be expected based on the spatio-temporal availability of beavers. This pattern is driven by temporal dynamics of beaver foraging: beavers make more foraging trips and spend more time on land per trip on longer feeding trails that extend farther from water. As a result, beavers are more vulnerable on longer feeding trails than shorter ones. Wolf predation appears to be a selective evolutionary pressure propelled by consumptive and non-consumptive mechanisms that constrain the distance from water beavers forage, which in turn limits the area of forest around wetlands, lakes and rivers beavers alter through foraging. Thus, wolves appear intricately linked to boreal forest dynamics by shaping beaver foraging behaviour, a form of natural disturbance that alters the successional and ecological states of forests.

Differential provisioning roles, prey size, and prey abundance shape the dynamic feeding behavior of gray wolves.

The demands of raising dependent young can influence the feeding behaviors of social carnivores, especially for individuals that are primarily responsible for provisioning young. We investigated how the feeding and provisioning behavior of a social carnivore, gray wolves (Canis lupus), are connected and shaped by extrinsic and intrinsic factors, and whether and how these patterns changed throughout the pup-rearing season (April-August). We found breeding wolves had shorter handling times of prey, lower probability of returning to kills, and greater probability of returning to homesites after kills compared to subordinate individuals. However, the feeding and provisioning behaviors of breeding individuals changed considerably over the pup-rearing season. Wolves had longer handling times and returned to provision pups directly after kills less frequently as annual prey abundance decreased. These patterns indicate that adult wolves prioritize meeting their own energetic demands over those of their pups when prey abundance decreases. We suggest that differential provisioning of offspring based on prey abundance is a behavioral mechanism by which group size adjusts to available resources via changes in neonate survival.

Logging, linear features, and human infrastructure shape the spatial dynamics of wolf predation on an ungulate neonate.

Humans are increasingly recognized as important players in predator-prey dynamics by modifying landscapes. This trend has been well-documented for large mammal communities in North American boreal forests: logging creates early seral forests that benefit ungulates such as white-tailed deer (Odocoileus virginianus), while the combination of infrastructure development and resource extraction practices generate linear features that allow predators such as wolves (Canis lupus) to travel and forage more efficiently throughout the landscape. Disturbances from recreational activities and residential development are other major sources of human activity in boreal ecosystems that may further alter wolf-ungulate dynamics. Here, we evaluate the influence that several major types of anthropogenic landscape modifications (timber harvest, linear features, and residential infrastructure) have on where and how wolves hunt ungulate neonates in a southern boreal forest ecosystem in Minnesota, USA. We demonstrate that each major anthropogenic disturbance significantly influences wolf predation of white-tailed deer fawns (n = 427 kill sites). In contrast with the "human shield hypothesis" that posits prey use human-modified areas as refuge, wolves killed fawns closer to residential buildings than expected based on spatial availability. Fawns were also killed within recently-logged areas more than expected. Concealment cover was higher at kill sites than random sites, suggesting wolves use senses other than vision, probably olfaction, to detect hidden fawns. Wolves showed strong selection for hunting along linear features, and kill sites were also closer to linear features than expected. We hypothesize that linear features facilitated wolf predation on fawns by allowing wolves to travel efficiently among high-quality prey patches (recently logged areas, near buildings), and also increase encounter rates with olfactory cues that allow them to detect hidden fawns. These findings provide novel insight into the strategies predators use to hunt ungulate neonates and the many ways human activity alters wolf-ungulate neonate predator-prey dynamics, which have remained elusive due to the challenges of locating sites where predators kill small prey. Our research has important management and conservation implications for wolf-ungulate systems subjected to anthropogenic pressures, particularly as the range of overlap between wolves and deer expands and appears to be altering food web dynamics in boreal ecosystems.

The ethology of wolves foraging on freshwater fish in a boreal ecosystem.

Through global positioning system (GPS) collar locations, remote cameras, field observations and the first wild wolf to be GPS-collared with a camera collar, we describe when, where and how wolves fish in a freshwater ecosystem. From 2017 to 2021, we recorded more than 10 wolves (Canis lupus) hunting fish during the spring spawning season in northern Minnesota, USA. Wolves ambushed fish in creeks at night when spawning fish were abundant, available and vulnerable in shallow waters. We observed wolves specifically targeting sections of rivers below beaver (Castor canadensis) dams, suggesting that beavers may indirectly facilitate wolf fishing behaviour. Wolves also cached fish on shorelines. We documented these findings across five different social groups at four distinct waterways, suggesting that wolf fishing behaviour may be widespread in similar ecosystems but has probably remained difficult to study given its annual brevity. Spawning fish may serve as a valuable pulsed resource for packs because the spring spawning season coincides with low primary prey (deer Odocoileus virginianus) availability and abundance, and when packs have higher energetic demands owing to newly born pups. We demonstrate the flexibility and adaptability of wolf hunting and foraging behaviour, and provide insight into how wolves can survive in a myriad of ecosystems.

Influence of infrastructure, ecology, and underpass-dimensions on multi-year use of Standard Gauge Railway underpasses by mammals in Tsavo, Kenya.

Rail and road infrastructure is essential for economic growth and development but can cause a gradual loss in biodiversity and degradation of ecosystem function and services. We assessed the influence of underpass dimensions, fencing, proximity to water and roads, Normalized Difference Vegetation Index (NDVI), presence of other species and livestock on underpass use by large and medium-sized mammals. Results revealed hyenas and leopards used the underpasses more than expected whereas giraffes and antelopes used the underpasses less than expected. Generalized linear mixed-effects models revealed that underpass height influenced use by wildlife, with several species preferring to use taller underpasses. Electric fencing increased underpass use by funneling species towards underpasses, except for elephants and black-backed jackal for which it reduced underpass passage. We also found that the use of underpasses by livestock reduced the probability of use by nearly 50% for wildlife species. Carnivore species were more likely to cross underpasses used by their prey. Buffalo, livestock, and hyenas used underpasses characterized by vegetation with higher NDVI and near water sources while baboons, dik-diks and antelope avoided underpasses with high NDVI. Our findings suggest a need for diverse and comprehensive approaches for mitigating the negative impacts of rail on African wildlife.

Contrasting levels of social distancing between the sexes in lions.

Understanding sexual segregation is crucial to comprehend sociality. A comparative analysis of long-term lion data from Serengeti and Ngorongoro in Tanzania, and Gir in India, reveals that male-female associations are contingent upon male and female group size, prey-size and availability, and the number of prides that each male coalition currently resides. Males maintain proximity with females, whereas females are responsible for segregation except at large kills. Lions feed on the largest prey in Ngorongoro and the smallest in Gir, and females spend the most time with males in Ngorongoro and the least in Gir. Females roar less often in prey-scarce circumstances in Serengeti and throughout the year in Gir possibly to prevent being tracked by males that parasitize on female kills. However, females readily associate with males when available prey is large and abundant. Contrasting availability of resources between Gir and Serengeti/Ngorongoro helps explain the varying degrees of sexual segregation and appears to drive differences in mating systems between these lion populations.

Outsized effect of predation: Wolves alter wetland creation and recolonization by killing ecosystem engineers.

Gray wolves are a premier example of how predators can transform ecosystems through trophic cascades. However, whether wolves change ecosystems as drastically as previously suggested has been increasingly questioned. We demonstrate how wolves alter wetland creation and recolonization by killing dispersing beavers. Beavers are ecosystem engineers that generate most wetland creation throughout boreal ecosystems. By studying beaver pond creation and recolonization patterns coupled with wolf predation on beavers, we determined that 84% of newly created and recolonized beaver ponds remained occupied until the fall, whereas 0% of newly created and recolonized ponds remained active after a wolf killed the dispersing beaver that colonized that pond. By affecting where and when beavers engineer ecosystems, wolves alter all of the ecological processes (e.g., water storage, nutrient cycling, and forest succession) that occur due to beaver-created impoundments. Our study demonstrates how predators have an outsized effect on ecosystems when they kill ecosystem engineers.

The role of kinship and demography in shaping cooperation amongst male lions.

The influence of kinship on animal cooperation is often unclear. Cooperating Asiatic lion coalitions are linearly hierarchical; male partners appropriate resources disproportionately. To investigate how kinship affect coalitionary dynamics, we combined microsatellite based genetic inferences with long-term genealogical records to measure relatedness between coalition partners of free-ranging lions in Gir, India. Large coalitions had higher likelihood of having sibling partners, while pairs were primarily unrelated. Fitness computations incorporating genetic relatedness revealed that low-ranking males in large coalitions were typically related to the dominant males and had fitness indices higher than single males, contrary to the previous understanding of this system based on indices derived from behavioural metrics alone. This demonstrates the indirect benefits to (related) males in large coalitions. Dominant males were found to 'lose less' if they lost mating opportunities to related partners versus unrelated males. From observations on territorial conflicts we show that while unrelated males cooperate, kin-selected benefits are ultimately essential for the maintenance of large coalitions. Although large coalitions maximised fitness as a group, demographic parameters limited their prevalence by restricting kin availability. Such demographic and behavioural constraints condition two-male coalitions to be the most attainable compromise for Gir lions.

Territoriality drives preemptive habitat selection in recovering wolves: Implications for carnivore conservation.

According to the ideal-free distribution (IFD), individuals within a population are free to select habitats that maximize their chances of success. Assuming knowledge of habitat quality, the IFD predicts that average fitness will be approximately equal among individuals and between habitats, while density varies, implying that habitat selection will be density dependent. Populations are often assumed to follow an IFD, although this assumption is rarely tested with empirical data, and may be incorrect when territoriality indicates habitat selection tactics that deviate from the IFD (e.g. ideal-despotic distribution or ideal-preemptive distribution). When territoriality influences habitat selection, species' density will not directly reflect components of fitness such as reproductive success or survival. In such cases, assuming an IFD can lead to false conclusions about habitat quality. We tested theoretical models of density-dependent habitat selection on a species known to exhibit territorial behaviour in order to determine whether commonly applied habitat models are appropriate under these circumstances. We combined long-term radiotelemetry and census data from grey wolves Canis lupus in the Upper Peninsula of Michigan, USA to relate spatiotemporal variability in wolf density to underlying classifications of habitat within a hierarchical state-space modelling framework. We then iteratively applied isodar analysis to evaluate which distribution of habitat selection best described this recolonizing wolf population. The wolf population in our study expanded by >1,000% during our study (~50 to >600 individuals), and density-dependent habitat selection was most consistent with the ideal-preemptive distribution, as opposed to the ideal-free or ideal-despotic alternatives. Population density of terrestrial carnivores may not be positively correlated with the fitness value of their habitats, and density-dependent habitat selection patterns may help to explain complex predator-prey dynamics and cascading indirect effects. Source-sink population dynamics appear likely when species exhibit rapid growth and occupy interspersed habitats of contrasting quality. These conditions are likely and have implications for large carnivores in many systems, such as areas in North America and Europe where large predator species are currently recolonizing their former ranges.

Comparing survey and multiple recruitment-mortality models to assess growth rates and population projections.

Estimation of population trends and demographic parameters is important to our understanding of fundamental ecology and species management, yet these data are often difficult to obtain without the use of data from population surveys or marking animals. The northeastern Minnesota moose (Alces alces Linnaeus, 1758) population declined 58% during 2006-2017, yet aerial surveys indicated stability during 2012-2017. In response to the decline, the Minnesota Department of Natural Resources (MNDNR) initiated studies of adult and calf survival to better understand cause-specific mortality, calf recruitment, and factors influencing the population trajectory. We estimated population growth rate (λ) using adult survival and calf recruitment data from demographic studies and the recruitment-mortality (R-M) Equation and compared these estimates to those calculated using data from aerial surveys. We then projected population dynamics 50 years using each resulting λ and used a stochastic model to project population dynamics 30 years using data from the MNDNR's studies. Calculations of λ derived from 2012 to 2017 survey data, and the R-M Equation indicated growth (1.02 ± 0.16 [SE] and 1.01 ± 0.04, respectively). However, the stochastic model indicated a decline in the population over 30 years (λ = 0.91 ± 0.004; 2014-2044). The R-M Equation has utility for estimating λ, and the supporting information from demographic collaring studies also helps to better address management questions. Furthermore, estimates of λ calculated using collaring data were more certain and reflective of current conditions. Long-term monitoring using collars would better inform population performance predictions and demographic responses to environmental variability.

Territorial landscapes: incorporating density-dependence into wolf habitat selection studies.

Habitat selection is a process that spans space, time and individual life histories. Ecological analyses of animal distributions and preferences are most accurate when they account for inherent dynamics of the habitat selection process. Strong territoriality can constrain perception of habitat availability by individual animals or groups attempting to colonize or establish new territory. Because habitat selection is a function of habitat availability, broad-scale changes in habitat availability or occupancy can drive density-dependent habitat functional responses. We investigated density-dependent habitat selection over a 19-year period of grey wolf (Canis lupus) recovery in Michigan, USA, using a generalized linear mixed model framework to develop a resource selection probability function (RSPF) with habitat coefficients conditioned on random effects for wolf packs and random year intercepts. In addition, we allowed habitat coefficients to vary as interactions with increasing wolf density over space and time. Results indicated that pack presence was driven by factors representing topography, human development, winter prey availability, forest structure, roads, streams and snow. Importantly, responses to many of these predictors were density-dependent. Spatio-temporal dynamics and population changes can cause considerable variation in wildlife-habitat relationships, possibly confounding interpretation of conventional habitat selection models. By incorporating territoriality into an RSPF analysis, we determined that wolves' habitat use in Michigan shifted over time, for example, exhibiting declining responses to winter prey indices and switching from positive to negative responses with respect to stream densities. We consider this an important example of a habitat functional response in wolves, driven by colonization, density-dependence and changes in occupancy during a time period of range expansion and population increase.

Towards Quantifying Carrion Biomass in Ecosystems.

The decomposition of animal biomass (carrion) contributes to the recycling of energy and nutrients through ecosystems. Whereas the role of plant decomposition in ecosystems is broadly recognised, the significance of carrion to ecosystem functioning remains poorly understood. Quantitative data on carrion biomass are lacking and there is no clear pathway towards improved knowledge in this area. Here, we present a framework to show how quantities derived from individual carcasses can be scaled up using population metrics, allowing for comparisons among ecosystems and other forms of biomass. Our framework facilitates the generation of new data that is critical to building a quantitative understanding of the contribution of carrion to trophic processes and ecosystem stocks and flows.

Does persistent snowpack inhibit degradation of fecal stress indicators?

Physiological stress in wildlife can be a useful indicator of a population's response to environmental factors. By using non-invasive endocrinological techniques, such as fecal sampling, potential confounding factors associated with the stress of capture can be avoided. A potential drawback of fecal sampling, however, is degradation of samples which may produce aberrant measurements of fecal glucocorticoid metabolites. In vertebrates, glucocorticoids, such as corticosterone, become elevated in response to stress. We sought to gauge the reliability of measurement of fecal glucocorticoid metabolites from white-tailed deer (Odocoileus virginianus) fecal samples exposed to a temperate winter with substantial snow cover and cold temperatures for up to 90 days, by repeatedly subsampling fecal samples every 10 days and performing a corticosterone enzyme-linked immunosorbent assay (ELISA). Measurements of fecal glucocorticoid metabolites at 10 days were consistent with initial measurements, after which (20 days) they became aberrant following a period of thawing. Consequently, glucocorticoid metabolite levels in feces appear to remain stable under ambient conditions if temperatures remain below freezing at least for 10 days. While it's possible that samples may remain useful beyond this time frame based on previous laboratory studies of samples stored in a freezer, further work is needed to determine how samples weather in situ under extreme cold (e.g. Arctic) or periods of partial thawing.

Finding wolf homesites: improving the efficacy of howl surveys to study wolves.

Locating wolf (Canis lupus) homesites is valuable for understanding the foraging behavior, population dynamics, and reproductive ecology of wolves during summer. During this period wolf pack members (adults and pups) readily respond to simulated wolf howls (i.e., howl surveys), which allows researchers to estimate the location of the homesite via triangulation. Confirming the actual locations of homesites via ground truthing is labor intensive because of the error surrounding estimated locations. Our objectives were (1) to quantify observer error during howl surveys and compare amongst experience levels, (2) provide a simple method for locating homesites in the field by incorporating observer error, and (3) further document the value of this method for monitoring wolf packs throughout the summer. We located 17 homesites by howl surveys during 2015-2017 in the Greater Voyageurs Ecosystem, Minnesota, USA. Of 62 bearings taken by observers during howl surveys, bearings erred by an average of 7.6° ± 6.3° (SD). There was no difference in observer error between novice and experienced observers. A simple way to increase efficiency when searching for homesites is to search concentric areas (bands) based on estimated observer error, specifically by: (1) adding ±10° error bands around howl survey bearings when ≥3 bearings can be obtained, (2) ±10° and ±20° error bands when 2 bearings are obtained, and (3) ±10° and ±26° error bands when 1 bearing is obtained. By incorporating observer error and understanding how frequently and how far wolves move homesites, it is possible to monitor wolf packs and confirm most, if not all, homesites used by a pack from at least June until August without having a collared individual in a pack.

Fertilizing riparian forests: nutrient repletion across ecotones with trophic rewilding.

Trophic rewilding maintains that large mammals are functionally important to resource subsidies and nutrient repletion, yet this prediction is understudied. Here, I report on the potential magnitude and variability of nitrogen that moose populations move from aquatic to terrestrial ecosystems. My aim is to provide justified approximations of the role of moose in the flux of a limiting nutrient across ecotones and to illustrate how this role is linked to wolf predation and climate warming. Using Isle Royale and northeastern Minnesota, USA as contrasting focal systems, I found that the long-term annual N gain for riparian forests likely ranges from 1 to 10 kg N ha-1 yr-1, depending on the heterogeneity of moose movements. For these systems, this range is equivalent to approximately 4-30% of net annual N mineralization, approximately 62-625% of annual N runoff, approximately 28-333% of annual atmospheric N deposition and approximately 31-312% of the N sequestered by trees. The N flux approximation is most sensitive to moose population levels and, as such, is influenced by wolves, climate warming and disease. The potential for other terrestrial ungulates that feed on aquatic plants to provide significant nutrient repletion across ecotones is unknown but important to examine in the context of trophic rewilding. The extent to which predators influence ungulate abundance indirectly impacts this nutrient repletion.This article is part of the theme issue 'Trophic rewilding: consequences for ecosystems under global change'.

Spatially varying density dependence drives a shifting mosaic of survival in a recovering apex predator (Canis lupus).

Understanding landscape patterns in mortality risk is crucial for promoting recovery of threatened and endangered species. Humans affect mortality risk in large carnivores such as wolves (Canis lupus), but spatiotemporally varying density dependence can significantly influence the landscape of survival. This potentially occurs when density varies spatially and risk is unevenly distributed. We quantified spatiotemporal sources of variation in survival rates of gray wolves (C. lupus) during a 21-year period of population recovery in the Upper Peninsula of Michigan, USA. We focused on mapping risk across time using Cox Proportional Hazards (CPH) models with time-dependent covariates, thus exploring a shifting mosaic of survival. Extended CPH models and time-dependent covariates revealed influences of seasonality, density dependence and experience, as well as individual-level factors and landscape predictors of risk. We used results to predict the shifting landscape of risk at the beginning, middle, and end of the wolf recovery time series. Survival rates varied spatially and declined over time. Long-term change was density-dependent, with landscape predictors such as agricultural land cover and edge densities contributing negatively to survival. Survival also varied seasonally and depended on individual experience, sex, and resident versus transient status. The shifting landscape of survival suggested that increasing density contributed to greater potential for human conflict and wolf mortality risk. Long-term spatial variation in key population vital rates is largely unquantified in many threatened, endangered, and recovering species. Variation in risk may indicate potential for source-sink population dynamics, especially where individuals preemptively occupy suitable territories, which forces new individuals into riskier habitat types as density increases. We encourage managers to explore relationships between adult survival and localized changes in population density. Density-dependent risk maps can identify increasing conflict areas or potential habitat sinks which may persist due to high recruitment in adjacent habitats.

Habitat capacity for cougar recolonization in the Upper Great Lakes region.

BACKGROUND: Recent findings indicate that cougars (Puma concolor) are expanding their range into the midwestern United States. Confirmed reports of cougar in Michigan, Minnesota, and Wisconsin have increased dramatically in frequency during the last five years, leading to speculation that cougars may re-establish in the Upper Great Lakes (UGL) region, USA. Recent work showed favorable cougar habitat in northeastern Minnesota, suggesting that the northern forested regions of Michigan and Wisconsin may have similar potential. Recolonization of cougars in the UGL states would have important ecological, social, and political impacts that will require effective management. METHODOLOGY/PRINCIPAL FINDINGS: Using Geographic Information Systems (GIS), we extended a cougar habitat model to Michigan and Wisconsin and incorporated primary prey densities to estimate the capacity of the region to support cougars. Results suggest that approximately 39% (>58,000 km2) of the study area could support cougars, and that there is potential for a population of approximately 500 or more animals. An exploratory validation of this habitat model revealed strong association with 58 verified cougar locations occurring in the study area between 2008 and 2013. CONCLUSIONS/SIGNIFICANCE: Spatially explicit information derived from this study could potentially lead to estimation of a viable population, delineation of possible cougar-human conflict areas, and the targeting of site locations for current monitoring. Understanding predator-prey interactions, interspecific competition, and human-wildlife relationships is becoming increasingly critical as top carnivores continue to recolonize the UGL region.