Living and dying in a multi‐predator landscape of fear: roe deer are squeezed by contrasting pattern of predation risk imposed by lynx and humans

The theory of predation risk effects predicts behavioral responses in prey when risk of predation is not homogenous in space and time. Prey species are often faced with a tradeoff between food and safety in situations where food availability and predation risk peak in the same habitat type. Determining the optimal strategy becomes more complex if predators with different hunting mode create contrasting landscapes of risk, but this has rarely been documented in vertebrates. Roe deer in southeastern Norway face predation risk from lynx, as well as hunting by humans. These two predators differ greatly in their hunting methods. The predation risk from lynx, an efficient stalk‐and‐ambush predator is expected to be higher in areas with dense understory vegetation, while predation risk from human hunters is expected to be higher where visual sight lines are longer. Based on field observations and airborne LiDAR data from 71 lynx predation sites, 53 human hunting sites, 132 locations from 15 GPS‐marked roe deer, and 36 roe deer pellet locations from a regional survey, we investigated how predation risk was related to terrain attributes and vegetation classes/structure. As predicted, we found that increasing cover resulted in a contrasting lower predation risk from humans and higher predation risk from lynx. Greater terrain ruggedness increased the predation risk from both predators. Hence, multiple predators may create areas of contrasting risk as well as double risk in the same landscape. Our study highlights the complexity of predator–prey relationship in a multiple predator setting. Synthesis In this study of risk effects in a multi‐predator context, LiDAR data were used to quantify cover in the habitat and relate it to vulnerability to predation in a boreal forest. We found that lynx and human hunters superimpose generally contrasting landscapes of fear on a common prey species, but also identified double‐risk zones. Since the benefit of anti‐predator responses depends on the combined risk from all predators, it is necessary to consider complete predator assemblages to understand the potential for and occurrence of risk effects across study systems.     

Tiger and leopard co-occurrence: intraguild interactions in response to human and livestock disturbance

Intraguild interactions have important implications for carnivore demography and conservation. Differences in how predators respond to different forms of disturbance might alter their interaction patterns. We sought to understand how human and livestock disturbance impact co-occurrence of sympatric large carnivores such as tiger (Panthera tigris) and leopard (P. pardus) and thereby mediate the intraguild interaction pattern to enable coexistence of the species in a human-dominated landscape. We surveyed 361 locations in Chitwan National Park, Nepal, to examine how prey abundance and disturbance factors such as human and livestock presence might influence habitat use by tigers and leopards independently and when co-occurring. Single-species single-season models and two-species single-season models were developed to examine hypotheses on unconditional detection and occupancy and species interaction respectively. Pervasive human use of the park had negative impacts on tiger occupancy while the abundance of prey had a positive influence. Despite significant prey overlap between tigers and leopards, none of the native prey species predicted leopard habitat occupancy. However, habitats used extensively by livestock were also used by leopards. Further, we found strong evidence of intraguild competition. For instance tiger occupancy was higher in prey-rich areas and leopard occupancy was low in the sites where tigers were present. These findings, and a species interaction factor of < 1 clearly indicate that leopards avoid tigers, but their use of areas of disturbance enables them to persist in fringe habitats. We provide empirical evidence of how intraguild interaction may result in habitat segregation between competing carnivores, while also showing that human and livestock use of the landscape create disturbance patterns that facilitate co-occurrence of the predators. Thus, because large carnivores compete, some disturbance may mediate coexistence in small protected areas. Understanding such interactions can help address important conservation challenges associated with maintaining diverse carnivore communities in small or disturbed landscapes.     

Landscapes of Coexistence for terrestrial carnivores: the ecological consequences of being downgraded from ultimate to penultimate predator by humans

Fear of predation can have major impacts on the behaviour of prey species. Recently the concept of the ecology of fear has been defined and formalised; yet there has been relatively little focus on how these ideas apply to large carnivore species which, although not prey sensu stricto, also experience fear as a result of threats from humans. Large carnivores are likely also subject to a Landscape of Fear similar to that described for prey species. We argue that although fear is generic, ‘human‐caused mortality’ represents a distinct and very important cause of fear for large carnivores, particularly terrestrial large carnivores as their activities overlap with those of humans to a greater degree. We introduce the idea of a ‘Landscape of Coexistence’ for large carnivores to denote a subset of the Landscape of Fear where sufficient areas of low human‐caused mortality risk are present in the landscape for long term coexistence of large carnivores and humans. We then explore aspects of terrestrial large carnivore behavioural ecology that may be best explained by risk of human‐caused mortality, and how the nature of a Landscape of Coexistence for these large carnivores is likely to be shaped by specific factors such as habitat structure, wild and domestic prey base, and human distribution and behaviour. The human characteristics of this Landscape of Coexistence may be as important in determining large carnivore distribution and behavioural ecology as the distribution of resources. Understanding the Landscape of Coexistence for terrestrial large carnivores is therefore important for their biology and conservation throughout large parts of their remaining ranges. Synthesis The Landscape of Fear concept describing the relationship between predator and prey also applies to the relationship between humans and top carnivores. We synthesise current research to introduce the Landscape of Coexistence concept, arguing that top predators respond to the risks of human‐caused mortality through spatiotemporal partitioning of activities to reduce contact with people. The character of the Landscape of Coexistence may be more important than the distribution of resources in determining large carnivore distribution and behavioural ecology in human dominated landscapes. Understanding their behavioural responses to human threats is crucial to successful conservation of large carnivores.     

Habitat selection by Eurasian lynx (Lynx lynx) is primarily driven by avoidance of human activity during day and prey availability during night

The greatest threat to the protected Eurasian lynx (Lynx lynx) in Central Europe is human‐induced mortality. As the availability of lynx prey often peaks in human‐modified areas, lynx have to balance successful prey hunting with the risk of encounters with humans. We hypothesized that lynx minimize this risk by adjusting habitat choices to the phases of the day and over seasons. We predicted that (1) due to avoidance of human‐dominated areas during daytime, lynx range use is higher at nighttime, that (2) prey availability drives lynx habitat selection at night, whereas high cover, terrain inaccessibility, and distance to human infrastructure drive habitat selection during the day, and that (3) habitat selection also differs between seasons, with altitude being a dominant factor in winter. To test these hypotheses, we analyzed telemetry data (GPS, VHF) of 10 lynx in the Bohemian Forest Ecosystem (Germany, Czech Republic) between 2005 and 2013 using generalized additive mixed models and considering various predictor variables. Night ranges exceeded day ranges by more than 10%. At night, lynx selected open habitats, such as meadows, which are associated with high ungulate abundance. By contrast, during the day, lynx selected habitats offering dense understorey cover and rugged terrain away from human infrastructure. In summer, land‐cover type greatly shaped lynx habitats, whereas in winter, lynx selected lower altitudes. We concluded that open habitats need to be considered for more realistic habitat models and contribute to future management and conservation (habitat suitability, carrying capacity) of Eurasian lynx in Central Europe.     

Temporal variation in habitat selection breaks the catch‐22 of spatially contrasting predation risk from multiple predators

Predator avoidance depends on prey being able to discern how risk varies in space and time, but this is made considerably more complicated if risk is simultaneously present from multiple predators. This is the situation for an increasing number of mammalian prey species, as large carnivores recover or are reintroduced in ecosystems on several continents. Roe deer Capreolus capreolus in southern Norway illustrate a case in which prey face two predators with contrasting patterns of predation risk. They face a catch‐22 situation: spatially avoiding the risk from one predator (lynx Lynx lynx in dense habitat) implies exposure to the other (hunters in open habitat). Using GPS‐data from 29 roe deer, we tested for daily and seasonal variation in roe deer selection for habitat with respect to the habitats’ year‐round average risk level. Generally, roe deer altered their habitat selection between night and day in a pattern consistent with being able to avoid predicted risk from the nocturnal lynx during night and predicted risk from human hunters during day. However, seasonal variation in habitat selection only partially corresponded with the predicted seasonal variation in risk. Whereas roe deer avoided areas with high risk from hunters more strongly during the hunting season than in other seasons, there was a lack of selection towards areas and time periods lowering the risk of lynx predation during winter. It seems likely that the risk of starvation and thermal stress constrain roe deer habitat selection during this energetically challenging season with cold temperatures, snow cover and limited natural forage. The habitat selection pattern of roe deer fits thus only partly with the two contrasting risk gradients they face. Adjusting risk‐avoidance behavior temporally can be an adaptive response in the case of several predators whose predation patterns differ in space and time.     

The cost of migratory prey: seasonal changes in semi‐domestic reindeer distribution influences breeding success of Eurasian lynx in northern Norway

Migratory prey is a widespread phenomenon that has implications for predator–prey interactions. By creating large temporal variation in resource availability between seasons it becomes challenging for carnivores to secure a regular year‐round supply of food. Some predators may respond by following their migratory prey, however, most predators are sedentary and experience strong seasonal variation in resource availability. Increased predation on alternative prey may dampen such seasonal resource fluctuations, but reduced reproduction rates in predators is a predicted consequence of migratory primary prey behavior that has received little empirical attention. We used data from 23 GPS collared Eurasian lynx Lynx lynx monitored during 2007–2013 in northern Norway, to examine how spatio‐temporal variation in the migratory behavior of semi‐domestic reindeer Rangifer tarandus influences lynx spatial organization and reproductive success using estimates of seasonal home range overlap and breeding success. We found that lynx of both sexes maintained seasonally stable home ranges and exhibited site fidelity across years, independent of whether they had access to reindeer throughout the year or experienced a scarcity of reindeer in winter due to migration. However, lynx without access to reindeer in winter showed a decreased probability of reproducing and a tendency for lowered kitten survival into their first winter, when compared to female lynx with reindeer available year around. This supports the hypothesis that sedentary predators experience demographic costs in systems with migratory primary prey. Changes in the migratory behavior of ungulates, including disrupted migrations, is therefore likely to have bottom–up effects on the population dynamics of sedentary predators as well as the previously documented consequences for ungulate population dynamics.     

The effect of anthropogenic features on the habitat selection of a large carnivore is conditional on sex and circadian period,suggesting a landscape of coexistence

Improving coexistence between humans and large predators is one of the foremost issues for the survival of large carnivores, especially in the Neotropics, where conflicts for retaliation are still frequent. This problem was increased due to the expansion of agricultural areas, settlements, roads, and the loss of natural habitats. Therefore, a key component in the long-term conservation of carnivores is to reduce animal-human conflicts. We aimed to assess multi-scale habitat selection models, exploring specificities in the selection for each sex and variation for the circadian period. We found that jaguars live in real landscapes of fear with high human and livestock density, where the perception of risk related to humans governed the selection of their resources. However, depending on the sex of individuals and the circadian period, jaguars positively selected some anthropic structures, such as areas of crops and human settlements. This selection suggested an aptitude to use various human-dominated structures and indicated jaguars could locally perceive risks in different ways, depending of sex and day period. Unexpectedly, jaguars presented attraction to roads, sexual or circadian related, regardless of the natural environments. Our results demonstrate that male and female jaguars could use some anthropic features differently in the distinct circadian periods. In addition, we conclude that the knowledge of the habitat selection for jaguars is a crucial component to the structure of the landscape of coexistence of this species and can give us efficient guidance to better comprehend the behavior through different scales of selection and through different periods of the day. Finally, our results show fundamental observations on the movement plasticity of this species for the construction of conservation plans focusing on the coexistence in different landscapes of the Neotropics dominated by humans.     

Risk taking by Eurasian lynx (Lynx lynx) in a human-dominated landscape: effects of sex and reproductive status

This study aimed to test how the sex and reproductive status of Eurasian lynx influenced their use of 'attractive sinks'– habitats with high prey density and high mortality risks. Locations of 24 Eurasian lynx Lynx lynx were obtained by radio-telemetry in a mixed forest and agricultural habitat in south-eastern Norway. Roe deer, the major food source of lynx in the study area, occurred at higher densities closer to areas of human activity and infrastructure. Proximity of lynx locations to human activity and infrastructure was used as a risk index because the most common causes of death among Scandinavian lynx were of anthropogenic origin. This study shows that distances from lynx locations to human activity were significantly greater for females with newborn kittens than for males, but this decreased with kitten age. The data suggest that this response to human activity is influenced by the reproductive strategies of males and females, and might explain male-biased human-induced mortality in this study and in carnivores more generally.     

River otter and mink occupancy dynamics in riparian systems

Semi-aquatic mammals are dependent upon streams and riparian areas, which are a product of the landscapes they drain. Both local stream morphology and surrounding land use are likely to have important influences on current occupancy of semi-aquatic mammals and potentially affect future geographic distributions. We identified aspects of the riparian system and stream structure at multiple scales that relate to the presence of river otter (Lontra canadensis) and mink (Neovison vison) to better understand how changing landscapes affect occupancy dynamics of these semi-aquatic mammals and to facilitate future monitoring and management. We estimated multi-season occupancy using 103 sites sampled over 6 seasonal sampling periods in southern Illinois, USA (44,526 km²) during 2012–2014. We hypothesized river otter and mink occupancy were related to multiple aspects of landscape and local habitat attributes including land cover, water availability, human disturbance, and stream characteristics. Occupancy of river otter was predicted by large stream size, less developed area near the stream site, and proximity to areas with reintroduced or remnant populations of river otter. Mink were more likely to occupy sites with small streams and decreased water availability near the site. However, top models for both species had low weights and high uncertainty for multiple variables. Habitat-based models may not be the best predictors of occupancy for these carnivores because they are more likely to respond to prey diversity or availability, but landscape changes that decrease natural water availability and increase human disturbance to the stream at the local scale are likely to negatively affect river otter. © 2019 The Authors. The Journal of Wildlife Management published by Wiley Periodicals, Inc. on behalf of The Wildlife Society.     

Time‐to‐kill: measuring attack rates in a heterogenous landscape with multiple prey types

Although spatial heterogeneity of prey and landscapes are known to contribute to variation around predator‐prey functional response models, few studies have quantified these effects. We illustrate a new approach using data from winter movement paths of GPS‐collared wolves in the Rocky Mountains of Canada and time‐to‐event models with competing risks for measuring the effect of prey and landscape characteristics on the time‐to‐kill, which is the reciprocal of attack rate (aN) in a Holling's functional response. We evaluated 13 a priori models representing hypothesized mechanisms influencing attack rates in a heterogeneous landscape with two prey types. Models ranged from variants on Holling's disc equation, including search rate and prey density, to a full model including prey density and patchiness, search rates, satiation, and landscape features, which were measured along the wolf's movement path. Movement rates of wolves while searching explained more of the variation in time‐to‐kill than prey densities. Wolves did not compensate for low prey density by increasing movement rates and there was little evidence that spatial aggregation of prey influenced attack rates in this multi‐prey system. The top model for predicting time‐to‐kill included only search rate and landscape features. Wolves killed prey more quickly in flat terrain, likely due to increased vulnerability from accumulated snow, whereas attack rates were lower when wolves hunted near human‐made features presumably due to human disturbance. Understanding the sources of variation in attack rates provides refinements to functional response models that can lead to more effective predator–prey management in human‐dominated landscapes.     

In the absence of a “landscape of fear”: How lions,hyenas, and cheetahs coexist

Aggression by top predators can create a “landscape of fear” in which subordinate predators restrict their activity to low‐risk areas or times of day. At large spatial or temporal scales, this can result in the costly loss of access to resources. However, fine‐scale reactive avoidance may minimize the risk of aggressive encounters for subordinate predators while maintaining access to resources, thereby providing a mechanism for coexistence. We investigated fine‐scale spatiotemporal avoidance in a guild of African predators characterized by intense interference competition. Vulnerable to food stealing and direct killing, cheetahs are expected to avoid both larger predators; hyenas are expected to avoid lions. We deployed a grid of 225 camera traps across 1,125 km² in Serengeti National Park, Tanzania, to evaluate concurrent patterns of habitat use by lions, hyenas, cheetahs, and their primary prey. We used hurdle models to evaluate whether smaller species avoided areas preferred by larger species, and we used time‐to‐event models to evaluate fine‐scale temporal avoidance in the hours immediately surrounding top predator activity. We found no evidence of long‐term displacement of subordinate species, even at fine spatial scales. Instead, hyenas and cheetahs were positively associated with lions except in areas with exceptionally high lion use. Hyenas and lions appeared to actively track each, while cheetahs appear to maintain long‐term access to sites with high lion use by actively avoiding those areas just in the hours immediately following lion activity. Our results suggest that cheetahs are able to use patches of preferred habitat by avoiding lions on a moment‐to‐moment basis. Such fine‐scale temporal avoidance is likely to be less costly than long‐term avoidance of preferred areas: This may help explain why cheetahs are able to coexist with lions despite high rates of lion‐inflicted mortality, and highlights reactive avoidance as a general mechanism for predator coexistence.     

What shapes Eurasian lynx distribution in human dominated landscapes: selecting prey or avoiding people?

In the multi-use landscape of southern Norway, the distribution of lynx is likely to be determined both by the abundance of their favoured prey – the roe deer – and the risk associated with the presence of humans because most lynx mortalities are caused by humans (recreational harvest, poaching, vehicle collisions). We described the distribution of the reproductive portion of the lynx population based on snow-track observations of females with dependent kittens collected over 10  yr (1997–2006) in southern Norway. We used the ecological-niche factor analysis to examine how lynx distribution was influenced by roe deer, human activity, habitat type, environmental productivity and elevation. Our first prediction that lynx should be found in areas of relatively high roe deer abundance was supported. However, our second prediction that lynx should avoid human activity was rejected, and lynx instead occupied areas more disturbed in average than those available (with the exception of the most densely occupied areas). Lynx, however, avoided the most disturbed areas and our third prediction of a trade-off between abundance of prey and avoidance of human activity was supported. On the one hand, roe deer in the most disturbed areas benefit to a large extent from current human land use practices, potentially allowing them to escape predation from lynx. On the other hand, the situation is not so favourable for the predators who are restricted in competition refuges with medium to low prey densities. The consequence is that lynx conservation will have to be achieved in a human modifed environment where the potential for a range of conflicts and high human-caused mortality will remain a constant threat.     

The importance of fine-scale breeding site selection patterns under a landscape-sharing approach for wolf conservation

For large carnivores persisting in human-dominated landscapes, conservation planning is often hindered by the large spatial requirements of these species, availability of protected areas, and human land uses. Protected areas are usually too small to support viable populations, and scattered across a human land-use matrix. Therefore, large carnivore conservation should be planned at large spatial scales under a land-sharing approach (allowing the coexistence between large carnivores and people in the same landscape), which means increasing the focus on the human-dominated matrix. Most of the critical factors determining large carnivore persistence (i.e., those related to food availability and vulnerability to humans) interact synergistically in space and time during the breeding season. Here, using as a case study a wolf population in NW Iberia, we studied fine-scale breeding site selection patterns (1 and 9 km²) in relation to human pressure, and the availability of food and refuge. The selection of wolf breeding sites in this human-dominated landscape was not determined by potential availability of prey biomass in the immediate vicinity (1 km²). However, wolves selected breeding sites with high availability of refuge (concealing vegetation), and low human accessibility and activity levels. Paved roads showed the highest proportion of independent contribution to explaining breeding site selection patterns (negative influence), being followed by refuge availability (positive influence) and the remoteness of breeding sites in relation to the surrounding spatial context (positive influence). Refuge availability, even at very small spatial scales taking into account the spatial requirements of wolves, may compensate for moderate levels of human activities in the vicinity of breeding sites. The strength of breeding selection patterns varied along a hierarchical process at different spatial scales. Under a landscape-sharing approach, integrating key processes observed in the human-dominated matrix, such as breeding site selection patterns, into landscape planning is of paramount importance for carnivore conservation. By temporally restricting human use on breeding sites and small portions of surrounding lands (~ 1 km²), and by maintaining several small refuge areas interspersed within the human-dominated matrix, we could favor wolf persistence without reducing land availability for other uses, improving the conditions for coexistence between wolves and humans.     

Phantoms of the forest: legacy risk effects of a regionally extinct large carnivore

The increased abundance of large carnivores in Europe is a conservation success, but the impact on the behavior and population dynamics of prey species is generally unknown. In Europe, the recolonization of large carnivores often occurs in areas where humans have greatly modified the landscape through forestry or agriculture. Currently, we poorly understand the effects of recolonizing large carnivores on extant prey species in anthropogenic landscapes. Here, we investigated if ungulate prey species showed innate responses to the scent of a regionally exterminated but native large carnivore, and whether the responses were affected by human‐induced habitat openness. We experimentally introduced brown bear Ursus arctos scent to artificial feeding sites and used camera traps to document the responses of three sympatric ungulate species. In addition to controls without scent, reindeer scent Rangifer tarandus was used as a noncarnivore, novel control scent. Fallow deer Dama dama strongly avoided areas with bear scent. In the presence of bear scent, all ungulate species generally used open sites more than closed sites, whereas the opposite was observed at sites with reindeer scent or without scent. The opening of forest habitat by human practices, such as forestry and agriculture, creates a larger gradient in habitat openness than available in relatively unaffected closed forest systems, which may create opportunities for prey to alter their habitat selection and reduce predation risk in human‐modified systems that do not exist in more natural forest systems. Increased knowledge about antipredator responses in areas subjected to anthropogenic change is important because these responses may affect prey population dynamics, lower trophic levels, and attitudes toward large carnivores. These aspects may be of particular relevance in the light of the increasing wildlife populations across much of Europe.     

Insights into wolf presence in human‐dominated landscapes: the relative role of food availability,humans and landscape attributes

Aim Understanding which human or environmental factors interact to enable or to limit the occurrence and persistence of large carnivores in human‐dominated landscapes is an important issue for their effective conservation, especially under the current scenario of global change where most of their former habitat is being transformed by humans. Location NW Iberian Peninsula. Methods We combine data on the distribution of Iberian wolves (Canis lupus signatus) living in a human‐dominated landscape in NW Spain and variation and partitioning methods to investigate the relative importance of three groups of predictors: food availability, humans and landscape attributes – each group expected to have unequal effects on wolf reproduction and survival – and their interactions on the occurrence of this species. Results We found that the group of predictors related with landscape attributes (altitude, roughness and refuge) strongly determined wolf occurrence, followed by humans and food availability. Variance partitioning analysis revealed that the three most important components determining wolf occurrence were related with landscape attributes: (1) the joint effects of the three predictor groups, (2) the joint effect of humans and landscape attributes and (3) the pure effect of landscape attributes. Altitude had the main independent contribution to explain the probability of wolf occurrence. Main conclusions In human‐dominated landscapes, the occurrence of wolves is the result of a complex interaction among several environmental and human factors. Our results suggest that the characteristics of the landscape (spatial context) – factors associated with the security of wolves facilitating that animals go unnoticed by humans, wolf movements, dispersal events and short‐time colonization – become more important in human‐dominated landscapes and may have played a key role in the occurrence and persistence of this species throughout decades modulating the relationship between humans and wolf distribution.     

Living on the edge: Multiscale habitat selection by cheetahs in a human‐wildlife landscape

Animals select habitats that will ultimately optimize their fitness through access to favorable resources, such as food, mates, and breeding sites. However, access to these resources may be limited by bottom‐up effects, such as availability, and top‐down effects, such as risk avoidance and competition, including that with humans. Competition between wildlife and people over resources, specifically over space, has played a significant role in the worldwide decrease in large carnivores. The goal of this study was to determine the habitat selection of cheetahs (Acinonyx jubatus) in a human‐wildlife landscape at multiple spatial scales. Cheetahs are a wide‐ranging, large carnivore, whose significant decline is largely attributed to habitat loss and fragmentation. It is believed that 77% of the global cheetah population ranges outside protected areas, yet little is known about cheetahs’ resource use in areas where they co‐occur with people. The selection, or avoidance, of three anthropogenic variables (human footprint density, distance to main roads and wildlife areas) and five environmental variables (open habitat, semiclosed habitat, edge density, patch density and slope), at multiple spatial scales, was determined by analyzing collar data from six cheetahs. Cheetahs selected variables at different scales; anthropogenic variables were selected at broader scales (720–1440 m) than environmental variables (90–180 m), suggesting that anthropogenic pressures affect habitat selection at a home‐range level, whilst environmental variables influence site‐level habitat selection. Cheetah presence was best explained by human presence, wildlife areas, semiclosed habitat, edge density and slope. Cheetahs showed avoidance for humans and steep slopes and selected for wildlife areas and areas with high proportions of semiclosed habitat and edge density. Understanding a species’ resource requirements, and how these might be affected by humans, is crucial for conservation. Using a multiscale approach, we provide new insights into the habitat selection of a large carnivore living in a human‐wildlife landscape.     

Vulnerability of Domestic Sheep to Lynx Depredation in Relation to Roe Deer Density

Abstract The Eurasian lynx (Lynx lynx) causes large loss of free-ranging domestic sheep in Norway. We tested whether the observed higher kill rates by male lynx than female lynx were related to an association between the availability of the main natural prey, as measured by roe deer (Capreolus capreolus) habitat suitability, presence of sheep, and habitat selection of male and female lynx. We found that lynx selected areas with high roe deer suitability during summer and winter. Moreover, during summer, compared to male lynx, females had greater selection for roe deer areas and a stronger avoidance for sheep grazing areas, which suggests that previously observed differences in kill rates between male and female lynx can be attributed to sex-specific habitat use during summer. The connection between lynx habitat use and roe deer also was reflected in a positive relationship between the roe deer suitability of a sheep grazing area and the total loss of lambs, which suggests that livestock, rather than being actively selected, are mainly killed by lynx incidentally when encountered during other lynx activities (e.g., searching for natural prey species). Therefore, any management practice that separates lynx and sheep, such as concentrating livestock into small patches or less preferred habitats, may reduce depredation.     

Canada lynx use of burned areas: Conservation implications of changing fire regimes

A fundamental problem in ecology is forecasting how species will react to major disturbances. As the climate warms, large, frequent, and severe fires are restructuring forested landscapes at large spatial scales, with unknown impacts on imperilled predators. We use the United States federally Threatened Canada lynx as a case study to examine how predators navigate recent large burns, with particular focus on habitat features and the spatial configuration (e.g., distance to edge) that enabled lynx use of these transformed landscapes. We coupled GPS location data of lynx in Washington in an area with several recent large fires and a number of GIS layers of habitat data to develop models of lynx habitat selection in recent burns. Random Forest habitat models showed lynx‐selected islands of forest skipped by large fires, residual vegetation, and areas where some trees survived to use newly burned areas. Lynx used burned areas as early as 1 year postfire, which is much earlier than the 2–4 decades postfire previously thought for this predator. These findings are encouraging for predator persistence in the face of fires, but increasingly severe fires or management that reduces postfire residual trees or slow regeneration will likely jeopardize lynx and other predators. Fire management should change to ensure heterogeneity is retained within the footprint of large fires to enable viable predator populations as fire regimes worsen with climate change.     

Topcats and underdogs: intraguild interactions among three apex carnivores across Asia's forestscapes

Intraguild interactions among carnivores have long held the fascination of ecologists. Ranging from competition to facilitation and coexistence, these interactions and their complex interplay influence everything from species persistence to ecosystem functioning. Yet, the patterns and pathways of such interactions are far from understood in tropical forest systems, particularly across countries in the Global South. Here, we examined the determinants and consequences of competitive interactions between dholes Cuon alpinus and the two large felids (leopards Panthera pardus and tigers Panthera tigris) with which they most commonly co-occur across Asia. Using a combination of traditional and novel data sources (N = 118), we integrate information from spatial, temporal, and dietary niche dimensions. These three species have faced catastrophic declines in their extent of co-occurrence over the past century; most of their source populations are now confined to Protected Areas. Analysis of dyadic interactions between species pairs showed a clear social hierarchy. Tigers were dominant over dholes, although pack strength in dholes helped ameliorate some of these effects; leopards were subordinate to dholes. Population-level spatio-temporal interactions assessed at 25 locations across Asia did not show a clear pattern of overlap or avoidance between species pairs. Diet-profile assessments indicated that wild ungulate biomass consumption by tigers was highest, while leopards consumed more primate and livestock prey as compared to their co-predators. In terms of prey offtake (ratio of wild prey biomass consumed to biomass available), the three species together harvested 0.4–30.2% of available prey, with the highest offtake recorded from the location where the carnivores reach very high densities. When re-examined in the context of prey availability and offtake, locations with low wild prey availability showed spatial avoidance and temporal overlap among the carnivore pairs, and locations with high wild prey availability showed spatial overlap and temporal segregation. Based on these observations, we make predictions for 40 Protected Areas in India where temporally synchronous estimates of predator and prey densities are available. We expect that low prey availability will lead to higher competition, and in extreme cases, to the complete exclusion of one or more species. In Protected Areas with high prey availability, we expect intraguild coexistence and conspecific competition among carnivores, with spill-over to forest-edge habitats and subsequent prey-switching to livestock. We stress that dhole–leopard–tiger co-occurrence across their range is facilitated through an intricate yet fragile balance between prey availability, and intraguild and conspecific competition. Data gaps and limitations notwithstanding, our study shows how insights from fundamental ecology can be of immense utility for applied aspects like large predator conservation and management of human–carnivore interactions. Our findings also highlight potential avenues for future research on tropical carnivores that can broaden current understanding of intraguild competition in forest systems of Asia and beyond.     

Habitat-related heterogeneity in breeding in a metapopulation of the Iberian lynx

Identifying attributes associated with good breeding habitat is critical for understanding animal population dynamics. However, the association between environmental heterogeneity and breeding probability has been often overlooked in habitat analyses. We evaluated habitat quality in a metapopulation of the endangered Iberian lynx Lynx pardinus by analyzing spatiotemporal patterns in breeding records. Data summarizing successful production of litters after emergence from dens over four years within 13 lynx territories were examined. We designed a set of generalized linear mixed models representing different hypotheses regarding how patterns in breeding records relate to environmental heterogeneity. Environmental heterogeneity was described by two characteristics: 1) a landscape index measured in lynx territories indicative of time‐averaged prey availability and 2) yearly variability in prey abundance not captured with this index. By including the random effect of the lynx territory we also accounted for other territory‐specific effects on reproduction. We found significant differences in yearly prey density dynamics among lynx territories. However, temporal variation in prey density contributed poorly to explaining lynx breeding. The most parsimonious model included the landscape structure as the only effect explaining breeding patterns. A multinomial‐model‐representation of the landscape hypothesis explained nearly 50% of variability in breeding records. Results pointed to the existence of a habitat quality gradient associated with particular landscape structures influencing lynx habitat selection and breeding performance. Underlying this gradient was time‐averaged prey availability. Probably as a result of long‐term fitness strategies in long‐lived territorial species, the short‐term fluctuations in prey availability had a minor influence. Our results illustrate how habitat inferences can be enhanced by incorporating the link between spatiotemporal patterns in reproduction and environmental heterogeneity.