Comparative Patterns of Predation by Cougars and Recolonizing Wolves in Montana's Madison Range

Abstract: Numerous studies have documented how prey may use antipredator strategies to reduce the risk of predation from a single predator. However, when a recolonizing predator enters an already complex predator—prey system, specific antipredator behaviors may conflict and avoidance of one predator may enhance vulnerability to another. We studied the patterns of prey selection by recolonizing wolves (Canis lupus) and cougars (Puma concolor) in response to prey resource selection in the northern Madison Range, Montana, USA. Elk (Cervus elaphus) were the primary prey for wolves, and mule deer (Odocoileus hemionus) were the primary prey for cougars, but elk made up an increasingly greater proportion of cougar kills annually. Although both predators preyed disproportionately on male elk, wolves were most likely to prey on males in poor physical condition. Although we found that the predators partitioned hunting habitats, structural complexity at wolf kill sites increased over time, whereas complexity of cougar kill sites decreased. We concluded that shifts by prey to structurally complex refugia were attempts by formerly naïve prey to lessen predation risk from wolves; nevertheless, shifting to more structurally complex refugia might have made prey more vulnerable to cougars. After a change in predator exposure, use of refugia may represent a compromise to minimize overall risk. As agencies formulate management strategies relative to wolf recolonization, the potential for interactive predation effects (i.e., facilitation or antagonism) should be considered.     

The landscape of fear as an emergent property of heterogeneity: Contrasting patterns of predation risk in grassland ecosystems

The likelihood of encountering a predator influences prey behavior and spatial distribution such that non‐consumptive effects can outweigh the influence of direct predation. Prey species are thought to filter information on perceived predator encounter rates in physical landscapes into a landscape of fear defined by spatially explicit heterogeneity in predation risk. The presence of multiple predators using different hunting strategies further complicates navigation through a landscape of fear and potentially exposes prey to greater risk of predation. The juxtaposition of land cover types likely influences overlap in occurrence of different predators, suggesting that attributes of a landscape of fear result from complexity in the physical landscape. Woody encroachment in grasslands furnishes an example of increasing complexity with the potential to influence predator distributions. We examined the role of vegetation structure on the distribution of two avian predators, Red‐tailed Hawk (Buteo jamaicensis) and Northern Harrier (Circus cyaneus), and the vulnerability of a frequent prey species of those predators, Northern Bobwhite (Colinus virginianus). We mapped occurrences of the raptors and kill locations of Northern Bobwhite to examine spatial vulnerability patterns in relation to landscape complexity. We use an offset model to examine spatially explicit habitat use patterns of these predators in the Southern Great Plains of the United States, and monitored vulnerability patterns of their prey species based on kill locations collected during radio telemetry monitoring. Both predator density and predation‐specific mortality of Northern Bobwhite increased with vegetation complexity generated by fine‐scale interspersion of grassland and woodland. Predation pressure was lower in more homogeneous landscapes where overlap of the two predators was less frequent. Predator overlap created areas of high risk for Northern Bobwhite amounting to 32% of the land area where landscape complexity was high and 7% where complexity was lower. Our study emphasizes the need to evaluate the role of landscape structure on predation dynamics and reveals another threat from woody encroachment in grasslands.     

Spatial Partitioning of Predation Risk in a Multiple Predator-Multiple Prey System

ABSTRACT Minimizing risk of predation from multiple predators can be difficult, particularly when the risk effects of one predator species may influence vulnerability to a second predator species. We decomposed spatial risk of predation in a 2-predator, 2-prey system into relative risk of encounter and, given an encounter, conditional relative risk of being killed. Then, we generated spatially explicit functions of total risk of predation for each prey species (elk [Cervus elaphus] and mule deer [Odocoileus hemionus]) by combining risks of encounter and kill. For both mule deer and elk, topographic and vegetation type effects, along with resource selection by their primary predator (cougars [Puma concolor] and wolves [Canis lupus], respectively), strongly influenced risk of encounter. Following an encounter, topographic and vegetation type effects altered the risk of predation for both ungulates. For mule deer, risk of direct predation was largely a function of cougar resource selection. However, for elk, risk of direct predation was not only a function of wolf occurrence, but also of habitat attributes that increased elk vulnerability to predation following an encounter. Our analysis of stage-based (i.e., encounter and kill) predation indicates that the risk effect of elk shifting to structurally complex habitat may ameliorate risk of direct predation by wolves but exacerbate risk of direct predation by cougars. Information on spatiotemporal patterns of predation will be become increasingly important as state agencies in the western United States face pressure to integrate predator and prey management.     

Beyond the encounter: Predicting multi‐predator risk to elk (Cervus canadensis) in summer using predator scats

1. There is growing evidence that prey perceive the risk of predation and alter their behavior in response, resulting in changes in spatial distribution and potential fitness consequences. Previous approaches to mapping predation risk across a landscape quantify predator space use to estimate potential predator‐prey encounters, yet this approach does not account for successful predator attack resulting in prey mortality. An exception is a prey kill site that reflects an encounter resulting in mortality, but obtaining information on kill sites is expensive and requires time to accumulate adequate sample sizes.
2. We illustrate an alternative approach using predator scat locations and their contents to quantify spatial predation risk for elk (Cervus canadensis) from multiple predators in the Rocky Mountains of Alberta, Canada. We surveyed over 1300 km to detect scats of bears (Ursus arctos/U. americanus), cougars (Puma concolor), coyotes (Canis latrans), and wolves (C. lupus). To derive spatial predation risk, we combined predictions of scat‐based resource selection functions (RSFs) weighted by predator abundance with predictions that a predator‐specific scat in a location contained elk. We evaluated the scat‐based predictions of predation risk by correlating them to predictions based on elk kill sites. We also compared scat‐based predation risk on summer ranges of elk following three migratory tactics for consistency with telemetry‐based metrics of predation risk and cause‐specific mortality of elk.
3. We found a strong correlation between the scat‐based approach presented here and predation risk predicted by kill sites and (r = .98, p < .001). Elk migrating east of the Ya Ha Tinda winter range were exposed to the highest predation risk from cougars, resident elk summering on the Ya Ha Tinda winter range were exposed to the highest predation risk from wolves and coyotes, and elk migrating west to summer in Banff National Park were exposed to highest risk of encountering bears, but it was less likely to find elk in bear scats than in other areas. These patterns were consistent with previous estimates of spatial risk based on telemetry of collared predators and recent cause‐specific mortality patterns in elk.
4. A scat‐based approach can provide a cost‐efficient alternative to kill sites of quantifying broad‐scale, spatial patterns in risk of predation for prey particularly in multiple predator species systems.

     

A spatial theory for characterizing predator–multiprey interactions in heterogeneous landscapes

Trophic interactions in multiprey systems can be largely determined by prey distributions. Yet, classic predator–prey models assume spatially homogeneous interactions between predators and prey. We developed a spatially informed theory that predicts how habitat heterogeneity alters the landscape-scale distribution of mortality risk of prey from predation, and hence the nature of predator interactions in multiprey systems. The theoretical model is a spatially explicit, multiprey functional response in which species-specific advection–diffusion models account for the response of individual prey to habitat edges. The model demonstrates that distinct responses of alternative prey species can alter the consequences of conspecific aggregation, from increasing safety to increasing predation risk. Observations of threatened boreal caribou, moose and grey wolf interacting over 378 181 km² of human-managed boreal forest support this principle. This empirically supported theory demonstrates how distinct responses of apparent competitors to landscape heterogeneity, including to human disturbances, can reverse density dependence in fitness correlates.     

Interspecific infanticide deters predators

It is well known that young, small predator stages are vulnerable to predation by conspecifics, intra-guild competitors or hyperpredators. It is less known that prey can also kill vulnerable predator stages that present no danger to the prey. Since adult predators are expected to avoid places where their offspring would run a high predation risk, this opens the way for potential prey to deter dangerous predator stages by killing vulnerable predator stages. We present an example of such a complex predator–prey interaction. We show that (1) the vulnerable stage of an omnivorous arthropod prey discriminates between eggs of a harmless predator species and eggs of a dangerous species, killing more eggs of the latter; (2) prey suffer a minor predation risk from newly hatched predators; (3) adult predators avoid ovipositing near killed predator eggs, and (4) vulnerable prey near killed predator eggs experience an almost fourfold reduction of predation. Hence, by attacking the vulnerable stage of their predator, prey deter adult predators and thus reduce their own predation risk. This provides a novel explanation for the killing of vulnerable stages of predators by prey and adds a new dimension to anti-predator behaviour.     

Estimating the prevalence and strength of non-independent predator effects

Understanding whether multiple predator species have independent effects on shared prey is critical for understanding community dynamics. We describe the prevalence and strength of non-independence between predators by quantifying the prey’s risk of predation and the degree to which it deviates from the risk predicted from a null model of independent predator effects. Specifically, we document how frequently non-independent effects occur among ten different multiple predator combinations with mayfly larvae as prey. These predator combinations vary both predator density and predator species richness. Overall, the predator effects were non-independent and translated to an average of 27% fewer prey being consumed compared to independent predator effects. Non-independence of this magnitude is likely to have population level consequences for the prey and influence the distribution or prey preference of predators. Closer inspection shows that much of the risk reduction in this system is weak, to the point of being indistinguishable from independent predator effects, while few effects are strong. This pattern of many weak interactions and few strong ones parallels the pattern of interaction strengths documented previously in intertidal communities. Consequently, understanding strong interactors in multiple predator systems may help us understand the importance of a species.     

Statistical Methods for Identifying Wolf Kill Sites Using Global Positioning System Locations

Abstract: Accurate estimates of kill rates remain a key limitation to addressing many predator—prey questions. Past approaches for identifying kill sites of large predators, such as wolves (Canis lupus), have been limited primarily to areas with abundant winter snowfall and have required intensive ground-tracking or aerial monitoring. More recently, attempts have been made to identify clusters of locations obtained using Global Positioning System (GPS) collars on predators to identify kill sites. However, because decision rules used in determining clusters have not been consistent across studies, results are not necessarily comparable. We illustrate a space—time clustering approach to statistically define clusters of wolf GPS locations that might be wolf kill sites, and we then use binary and multinomial logistic regression to model the probability of a cluster being a non—kill site, kill site of small-bodied prey species, or kill site of a large-bodied prey species. We evaluated our approach using field visits of kills and assessed the accuracy of the models using an independent dataset. The cluster-scan approach identified 42–100% of wolf-killed prey, and top logistic regression models correctly classified 100% of kills of large-bodied prey species, but 40% of small-bodied prey species were classified as nonkills. Although knowledge of prey distribution and vulnerability may help refine this approach, identifying small-bodied prey species will likely remain problematic without intensive field efforts. We recommend that our approach be utilized with the understanding that variation in prey body size and handling time by wolves will likely have implications for the success of both the cluster scan and logistic regression components of the technique. (JOURNAL OF WILDLIFE MANAGEMENT 72(3):798–807; 2008)     

Intraguild predation reduces redundancy of predator species in multiple predator assemblage

1. Interference between predator species frequently decreases predation rates, lowering the risk of predation for shared prey. However, such interference can also occur between conspecific predators. 2. Therefore, to understand the importance of predator biodiversity and the degree that predator species can be considered functionally interchangeable, we determined the degree of additivity and redundancy of predators in multiple- and single-species combinations. 3. We show that interference between two invasive species of predatory crabs, Carcinus maenas and Hemigrapsus sanguineus, reduced the risk of predation for shared amphipod prey, and had redundant per capita effects in most multiple- and single-species predator combinations. 4. However, when predator combinations with the potential for intraguild predation were examined, predator interference increased and predator redundancy decreased. 5. Our study indicates that trophic structure is important in determining how the effects of predator species combine and demonstrates the utility of determining the redundancy, as well as the additivity, of multiple predator species.     

Seasonal patterns of predation for gray wolves in the multi-prey system of Yellowstone National Park

1.?For large predators living in seasonal environments, patterns of predation are likely to vary among seasons because of related changes in prey vulnerability. Variation in prey vulnerability underlies the influence of predators on prey populations and the response of predators to seasonal variation in rates of biomass acquisition. Despite its importance, seasonal variation in predation is poorly understood. 2.?We assessed seasonal variation in prey composition and kill rate for wolves Canis lupus living on the Northern Range (NR) of Yellowstone National Park. Our assessment was based on data collected over 14 winters (1995-2009) and five spring-summers between 2004 and 2009. 3.?The species composition of wolf-killed prey and the age and sex composition of wolf-killed elk Cervus elaphus (the primary prey for NR wolves) varied among seasons. 4.?One's understanding of predation depends critically on the metric used to quantify kill rate. For example, kill rate was greatest in summer when quantified as the number of ungulates acquired per wolf per day, and least during summer when kill rate was quantified as the biomass acquired per wolf per day. This finding contradicts previous research that suggests that rates of biomass acquisition for large terrestrial carnivores tend not to vary among seasons. 5.?Kill rates were not well correlated among seasons. For example, knowing that early-winter kill rate is higher than average (compared with other early winters) provides little basis for anticipating whether kill rates a few months later during late winter will be higher or lower than average (compared with other late winters). This observation indicates how observing, for example, higher-than-average kill rates throughout any particular season is an unreliable basis for inferring that the year-round average kill rate would be higher than average. 6.?Our work shows how a large carnivore living in a seasonal environment displays marked seasonal variation in predation because of changes in prey vulnerability. Patterns of wolf predation were influenced by the nutritional condition of adult elk and the availability of smaller prey (i.e. elk calves, deer). We discuss how these patterns affect our overall understanding of predator and prey population dynamics.     

Nonconsumptive effects in a multiple predator system reduce the foraging efficiency of a keystone predator

Many studies have demonstrated that the nonconsumptive effect (NCE) of predators on prey traits can alter prey demographics in ways that are just as strong as the consumptive effect (CE) of predators. Less well studied, however, is how the CE and NCE of multiple predator species can interact to influence the combined effect of multiple predators on prey mortality. We examined the extent to which the NCE of one predator altered the CE of another predator on a shared prey and evaluated whether we can better predict the combined impact of multiple predators on prey when accounting for this influence. We conducted a set of experiments with larval dragonflies, adult newts (a known keystone predator), and their tadpole prey. We quantified the CE and NCE of each predator, the extent to which NCEs from one predator alters the CE of the second predator, and the combined effect of both predators on prey mortality. We then compared the combined effect of both predators on prey mortality to four predictive models. Dragonflies caused more tadpoles to hide under leaf litter (a NCE), where newts spend less time foraging, which reduced the foraging success (CE) of newts. Newts altered tadpole behavior but not in a way that altered the foraging success of dragonflies. Our study suggests that we can better predict the combined effect of multiple predators on prey when we incorporate the influence of interactions between the CE and NCE of multiple predators into a predictive model. In our case, the threat of predation to prey by one predator reduced the foraging efficiency of a keystone predator. Consequently, the ability of a predator to fill a keystone role could be compromised by the presence of other predators.     

Tradeoffs involved in site selection and foraging in a wolf spider: effects of substrate structure and predation risk

Understanding how animals weigh habitat features, exposure to predators and access to resources is important to determining their life history and distribution across the landscape. For example, when predators accumulate in structurally complex habitats, they face an environment with different competitive interactions, foraging opportunities and predatory risks. The wolf spider Pardosa milvina inhabits the soil surface of highly disturbed habitats such as agricultural fields throughout eastern North America. Pardosa displays effective antipredator behavior in the presence of chemical cues produced by a larger coexisting wolf spider, Hogna helluo . We used those cues to simulate predation risk in laboratory and field experiments designed to test the effects of habitat substrate and predation risk on site selection and prey consumption of Pardosa . In general, Pardosa preferred more complex substrates over bare dirt but those preferences were eliminated or reversed when cues from Hogna were present. Feeding trials revealed that substrate alone had few effects on Pardosa prey consumption, which we measured by documenting the change in the abdomen width. Although the presence of Hogna cues reduced prey consumption overall in field feeding trials, the negative effect of predation risk on prey consumption was only observed in grass and bare dirt substrates in the laboratory. We also found that prey capture was negatively affected by habitat complexity for both spider species but that same complexity offered Pardosa protection from predation by Hogna. This study provides insight into how two predator species interact to balance site selection and feeding in order to avoid predation. Shifts in foraging and distributional patterns of predators can have profound implications for their role in the food web.     

Evaluating the effects of large marine predators on mobile prey behavior across subtropical reef ecosystems

The indirect effect of predators on prey behavior, recruitment, and spatial relationships continues to attract considerable attention. However, top predators like sharks or large, mobile teleosts, which can have substantial top–down effects in ecosystems, are often difficult to study due to their large size and mobility. This has created a knowledge gap in understanding how they affect their prey through nonconsumptive effects. Here, we investigated how different functional groups of predators affected potential prey fish populations across various habitats within Biscayne Bay, FL. Using baited remote underwater videos (BRUVs), we quantified predator abundance and activity as a rough proxy for predation risk and analyzed key prey behaviors across coral reef, sea fan, seagrass, and sandy habitats. Both predator abundance and prey arrival times to the bait were strongly influenced by habitat type, with open homogenous habitats receiving faster arrival times by prey. Other prey behaviors, such as residency and risk‐associated behaviors, were potentially driven by predator interaction. Our data suggest that small predators across functional groups do not have large controlling effects on prey behavior or stress responses over short temporal scales; however, habitats where predators are more unpredictable in their occurrence (i.e., open areas) may trigger risk‐associated behaviors such as avoidance and vigilance. Our data shed new light on the importance of habitat and context for understanding how marine predators may influence prey behaviors in marine ecosystems.     

Do prey select for vacant hunting domains to minimize a multi‐predator threat?

Many ecosystems contain sympatric predator species that hunt in different places and times. We tested whether this provides vacant hunting domains, places and times where and when predators are least active, that prey use to minimize threats from multiple predators simultaneously. We measured how northern Yellowstone elk (Cervus elaphus) responded to wolves (Canis lupus) and cougars (Puma concolor), and found that elk selected for areas outside the high‐risk domains of both predators consistent with the vacant domain hypothesis. This enabled elk to avoid one predator without necessarily increasing its exposure to the other. Our results demonstrate how the diel cycle can serve as a key axis of the predator hunting domain that prey exploit to manage predation risk from multiple sources. We argue that a multi‐predator, spatiotemporal framework is vital to understand the causes and consequences of prey spatial response to predation risk in environments with more than one predator.     

Wolves,white‐tailed deer,and beaver: implications of seasonal prey switching for woodland caribou declines

Population increases of primary prey can negatively impact alternate prey populations via demographic and behavioural responses of a shared predator through apparent competition. Seasonal variation in prey selection patterns by predators also can affect secondary and incidental prey by reducing spatial separation. Global warming and landscape changes in Alberta's bitumen sands have resulted in prey enrichment, which is changing the large mammal predator–prey system and causing declines in woodland caribou Rangifer tarandus caribou populations. We assessed seasonal patterns of prey use and spatial selection by wolves Canis lupus in two woodland caribou ranges in northeastern Alberta, Canada, that have undergone prey enrichment following recent white‐tailed deer Odocoileus virginianus invasion. We determined whether risk of predation for caribou (incidental prey) and the proportion of wolf‐caused‐caribou mortalities varied with season. We found that wolves showed seasonal variation in primary prey use, with deer and beaver Castor canadensis being the most common prey items in wolf diet in winter and summer, respectively. These seasonal dietary patterns were reflected in seasonal wolf spatial resource selection and resulted in contrasting spatial relationships between wolves and caribou. During winter, wolf selection for areas used by deer maintained strong spatial separation between wolves and caribou, whereas wolf selection for areas used by beaver in summer increased the overlap with caribou. Changing patterns in wolf resource selection were reflected by caribou mortality patterns, with 76.2% of 42 adult female caribou mortalities occurring in summer. Understanding seasonal patterns of predation following prey enrichment in a multiprey system is essential when assessing the effect of predation on an incidental prey species. Our results support the conclusion that wolves are proximately responsible for woodland caribou population declines throughout much of their range.     

Stochastic predation exposes prey to predator pits and local extinction

Understanding how predators affect prey populations is a fundamental goal for ecologists and wildlife managers. A well-known example of regulation by predators is the predator pit, where two alternative stable states exist and prey can be held at a low density equilibrium by predation if they are unable to pass the threshold needed to attain a high density equilibrium. While empirical evidence for predator pits exists, deterministic models of predator–prey dynamics with realistic parameters suggest they should not occur in these systems. Because stochasticity can fundamentally change the dynamics of deterministic models, we investigated if incorporating stochasticity in predation rates would change the dynamics of deterministic models and allow predator pits to emerge. Based on realistic parameters from an elk–wolf system, we found predator pits were predicted only when stochasticity was included in the model. Predator pits emerged in systems with highly stochastic predation and high carrying capacities, but as carrying capacity decreased, low density equilibria with a high likelihood of extinction became more prevalent. We found that incorporating stochasticity is essential to fully understand alternative stable states in ecological systems, and due to the interaction between top–down and bottom–up effects on prey populations, habitat management and predator control could help prey to be resilient to predation stochasticity.     

Herbivore release through cascading risk effects

Predators influence prey through consumption, and through trait-mediated effects such as emigration in response to predation risk (risk effects). We studied top-down effects of (sub-) adult wolf spiders (Lycosidae) on arthropods in a meadow. We compared risk effects with the overall top-down effect (including consumption) by gluing the chelicers of wolf spiders to prevent them from killing the prey. In a field experiment, we created three treatments that included either: (i) intact (‘predation’) wolf spiders; (ii) wolf spiders with glued chelicers (‘risk spiders’); or (iii) no (sub-) adult wolf spiders. Young wolf spiders were reduced by their (sub-) adult congeners. Densities of sheetweb spiders (Linyphiidae), a known intraguild prey of wolf spiders, were equally reduced by the presence of risk and predation wolf spiders. Plant- and leafhoppers (Auchenorrhyncha) showed the inverse pattern of higher densities in the presence of both risk and predation wolf spiders. We conclude that (sub-) adult wolf spiders acted as top predators, which reduced densities of intermediate predators and thereby enhanced herbivores. Complementary to earlier studies that found trait-mediated herbivore suppression, our results demonstrate that herbivores can be enhanced through cascading risk effects by top predators.     

Prey Responses to Predator's Sounds: A Review and Empirical Study

Many animals assess their risk of predation by listening to and evaluating predators' vocalizations. We reviewed the literature to draw generalizations about predator discrimination abilities, the retention of these abilities over evolutionary time, and the potential underlying proximate mechanisms responsible for discrimination. Broadly, we found that some prey possess an ability to respond to a predator after having been evolutionarily isolated from a specific predator (i.e., predators are allopatric) and that some prey are predisposed to respond to certain types of predators that they coevolved with but without having ecological experience. However, these types of studies are lacking, and relatively, few studies have examined predator discrimination abilities in ungulates. To begin addressing these knowledge gaps, we performed field experiments on Mule deer (Odocoileus hemionus) in which we investigated the ability of deer to discriminate among familiar predators [coyotes (Canis latrans) and mountain lions (Puma concolor)] and an evolutionary relevant predator with which deer have had no recent exposure [locally extinct wolves (Canis lupus)]. We found that Mule deer respond to and discriminate among predators based on predator vocalizations and have retained an ability to respond to wolves that have been extinct from the study area since the early 20th century. Previous playback studies have shown that responses vary among human‐habituated and non‐habituated populations and differ according to human proximity. Deer greater than 0.5 km from human residences allocated more time to heightened responses both before and after stimulus playback. Our findings may help predict how prey–predator interactions may change as a result of the recovering wolf population with a basis in ecological and evolutionary experience in predator discrimination and desensitization.     

Linking anti‐predator behaviour to prey demography reveals limited risk effects of an actively hunting large carnivore

Ecological theory predicts that the diffuse risk cues generated by wide‐ranging, active predators should induce prey behavioural responses but not major, population‐ or community‐level consequences. We evaluated the non‐consumptive effects (NCEs) of an active predator, the grey wolf (Canis lupus), by simultaneously tracking wolves and the behaviour, body fat, and pregnancy of elk (Cervus elaphus), their primary prey in the Greater Yellowstone Ecosystem. When wolves approached within 1 km, elk increased their rates of movement, displacement and vigilance. Even in high‐risk areas, however, these encounters occurred only once every 9 days. Ultimately, despite 20‐fold variation in the frequency of encounters between wolves and individual elk, the risk of predation was not associated with elk body fat or pregnancy. Our findings suggest that the ecological consequences of actively hunting large carnivores, such as the wolf, are more likely transmitted by consumptive effects on prey survival than NCEs on prey behaviour.     

Spatial decomposition of predation risk using resource selection functions: an example in a wolf–elk predator–prey system

Predation is a fundamental ecological and evolutionary process that varies in space, and the avoidance of predation risk is of central importance in foraging theory. While there has been a recent growth of approaches to spatially model predation risk, these approaches lack an adequate mechanistic framework that can be applied to real landscapes. In this paper we show how predation risk can be decomposed into encounter and attack stages, and modeled spatially using resource selection functions (RSF) and resource selection probability functions (RSPF). We use this approach to compare the effects of landscape attributes on the relative probability of encounter, the conditional probability of death given encounter, and overall wolf and elk resource selection to test whether predation risk is simply equivalent to location of the predator. We then combine the probability of encounter and conditional probability of death into a spatially explicit function of predation risk following Lima and Dill's reformulation of Holling's functional response. We illustrate this approach in a wolf–elk system in and adjacent to Banff National Park, Alberta, Canada. In this system we found that the odds of elk being encountered by wolves was 1.3 times higher in pine forest and 4.1 times less in grasslands than other habitats. The relative odds of being killed in pine forests, given an encounter, increased by 1.2. Other habitats, such as grasslands, afforded elk reduced odds (4.1 times less) of being encountered and subsequently killed (1.4 times less) by wolves. Our approach illustrates that predation risk is not necessarily equivalent to just where predators are found. We show that landscape attributes can render prey more or less susceptible to predation and effects of landscape features can differ between the encounter and attack stages of predation. We conclude by suggesting applications of our approach to model predator–prey dynamics using spatial predation risk functions in theoretical and applied settings.