Changing vulnerability to predation related to season and sex in an African ungulate assemblage

The consequences of predation for prey population dynamics depend on the extent to which this mortality is predisposed by malnutrition or senescence, or additive in the sense that animals that would otherwise not have died at that time were killed. In places lacking effective predators, few adult ungulates die during the summer or wet season months when food is plentifully available. Hence the seasonal distribution of predator kills as well as the age and sex classes of the prey indicates the extent to which malnutrition contributes to mortality as well as other influences on vulnerability. Using records of animal deaths assembled over 35 years in South Africa's Kruger National Park, these patterns were investigated for 12 ungulate species forming the prey of lions, and for three other large predators with respect to one prey species. Buffalo, kudu and giraffe were more strongly represented in kills made during the late dry season, while wildebeest and zebra made relatively greater contributions during the wet season. Impala, waterbuck, warthog and rarer antelope species became more prominent in kills during transitional periods between seasons. Five prey species showed an elevation in representation of males in lion kills during the mating season, as well as impala for all predator species. Females were more prominently represented in kills during the time of late gestation and parturition for three prey species. Hence reproductive activities as well as changing vegetation cover and food resources affected vulnerability to predation. Shifts in susceptibility to predation over the seasonal cycle corresponded with rainfall‐related variation in the annual representation of these ungulate species in lion kills. The availability of vulnerable prey species, age and sex classes at different stages of the seasonal cycle helps maintain a high abundance of lions. These factors contribute to the strong additive impact that predation has had on the abundance of some of these ungulate populations.     

A cross‐system meta‐analysis reveals coupled predation effects on prey biomass and diversity

Predator diversity and abundance are under strong human pressure in all types of ecosystems. Whereas predator potentially control standing biomass and species interactions in food webs, their effects on prey biomass and especially prey biodiversity have not yet been systematically quantified. Here, we test the effects of predation in a cross‐system meta‐analysis of prey diversity and biomass responses to local manipulation of predator presence. We found 291 predator removal experiments from 87 studies assessing both diversity and biomass responses. Across ecosystem types, predator presence significantly decreased both biomass and diversity of prey across ecosystems. Predation effects were highly similar between ecosystem types, whereas previous studies had shown that herbivory or decomposition effects differed fundamentally between terrestrial and aquatic systems based on different stoichiometry of plant material. Such stoichiometric differences between systems are unlikely for carnivorous predators, where effect sizes on species richness strongly correlated to effect sizes on biomass. However, the negative predation effect on prey biomass was ameliorated significantly with increasing prey richness and increasing species richness of the manipulated predator assemblage. Moreover, with increasing richness of the predator assemblage present, the overall negative effects of predation on prey richness switched to positive effects. Our meta‐analysis revealed strong general relationships between predator diversity, prey diversity and the interaction strength between trophic levels in terms of biomass. This study indicates that anthropogenic changes in predator abundance and diversity will potentially have strong effects on trophic interactions across ecosystems. Synthesis The past centuries we have experienced a dramatic loss of top–predator abundance and diversity in most types of ecosystems. To understand the direct consequences of predator loss on a global scale, we quantitatively summarized experiments testing predation effects on prey communities in a cross‐system meta‐analysis. Across ecosystem types, predator presence significantly decreased both biomass and diversity of prey, and predation effects were highly similar. However, with increasing predator richness, the overall negative effects of predation on prey richness switched to positive ones. Anthropogenic changes in predator communities will potentially have strong effects on prey diversity, biomass, and trophic interactions across ecosystems.     

Predator identity and time of day interact to shape the risk–reward trade-off for herbivorous coral reef fishes

Non-consumptive effects (NCEs) of predators occur as prey alters their habitat use and foraging decisions to avoid predation. Although NCEs are recognized as being important across disparate ecosystems, the factors influencing their strength and importance remain poorly understood. Ecological context, such as time of day, predator identity, and prey condition, may modify how prey species perceive and respond to risk, thereby altering NCEs. To investigate how predator identity affects foraging of herbivorous coral reef fishes, we simulated predation risk using fiberglass models of two predator species (grouper Mycteroperca bonaci and barracuda Sphyraena barracuda) with different hunting modes. We quantified how predation risk alters herbivory rates across space (distance from predator) and time (dawn, mid-day, and dusk) to examine how prey reconciles the conflicting demands of avoiding predation vs. foraging. When we averaged the effect of both predators across space and time, they suppressed herbivory similarly. Yet, they altered feeding differently depending on time of day and distance from the model. Although feeding increased strongly with increasing distance from the predators particularly during dawn, we found that the barracuda model suppressed herbivory more strongly than the grouper model during mid-day. We suggest that prey hunger level and differences in predator hunting modes could influence these patterns. Understanding how context mediates NCEs provides insight into the emergent effects of predator–prey interactions on food webs. These insights have broad implications for understanding how anthropogenic alterations to predator abundances can affect the spatial and temporal dynamics of important ecosystem processes.     

The Effect of Simulated African Wild Dog Presence on Anti‐predator Behaviour of Kudu and Impala

In this study, we examined the behavioural, temporal and spatial effects of simulated African wild dog (Lycaon pictus) presence on its two main prey species: kudu (Tragelaphus strepsiceros) and impala (Aepyceros melampus). We spread African wild dog faeces around waterholes and played African wild dog sounds at different intervals to mimic immediate and non‐immediate predation pressure. We looked at anti‐predator behaviour at both a herd and individual level and distinguished between high‐quality (detracts from all other activities), high‐cost vigilance and low‐quality (used to monitor the surrounding in spare time), low‐cost vigilance to determine costs involved. We found that simulated African wild dog presence had little effect on anti‐predator behaviour of their free‐ranging prey. Only when immediate predation risk was mimicked did kudu invest in (additional) high‐quality vigilance, whereas impala showed no response. Regardless of direct cues of African wild dog presence, behavioural adjustments to reduce predation risk were primarily based on environmental factors such as time of the day and broad‐scale habitat structure. Predators have been shown to utilize waterholes to hunt, and prey species are therefore likely to maximize anti‐predator behaviour in this high‐risk environment based on environmental variables affecting predation risk, the main predator within the system, and water requirements, leaving little flexibility to respond to (simulated) African wild dog presence.     

Inside‐container effects drive mosquito community structure in Brazilian Atlantic forest

Metacommunity theory is a convenient framework in which to investigate how local communities linked by dispersal influence patterns of species distribution and abundance across large spatial scales. For organisms with complex life cycles, such as mosquitoes, different pressures are expected to act on communities due to behavioral and ecological partitioning of life stages. Adult females select habitats for oviposition, and resulting offspring are confined to that habitat until reaching adult stages capable of flight; outside‐container effects (OCE) (i.e., spatial factors) are thus expected to act more strongly on species distributions as a function of adult dispersal capability, which should be limited by geographic distances between sites. However, larval community dynamics within a habitat are influenced by inside‐container effects (ICE), mainly interactions with conspecifics and heterospecifics (e.g., through effects of competition and predation). We used a field experiment in a mainland‐island scenario to assess whether environmental, spatial, and temporal factors influence mosquito prey and predator distributions and abundances across spatial scales: within‐site, between‐site, and mainland‐island. We also evaluated whether predator abundances inside containers play a stronger role in shaping mosquito prey community structure than do OCE (e.g., spatial and environmental factors). Temporal influence was more important for predators than for prey mosquito community structure, and the changes in prey mosquito species composition over time appear to be driven by changes in predator abundances. There was a negligible effect of spatial and environmental factors on mosquito community structure, and temporal effects on mosquito abundances and distributions appear to be driven by changes in abundance of the dominant predator, perhaps because ICE are stronger than OCE due to larval habitat restriction, or because adult dispersal is not limited at the chosen spatial scales.     

Diet quality in a wild grazer declines under the threat of an ambush predator

Predators influence prey populations not only through predation itself, but also indirectly through prompting changes in prey behaviour. The behavioural adjustments of prey to predation risk may carry nutritional costs, but this has seldom been studied in the wild in large mammals. Here, we studied the effects of an ambush predator, the African lion (Panthera leo), on the diet quality of plains zebras (Equus quagga) in Hwange National Park, Zimbabwe. We combined information on movements of both prey and predators, using GPS data, and measurements of faecal crude protein, an index of diet quality in the prey. Zebras which had been in close proximity to lions had a lower quality diet, showing that adjustments in behaviour when lions are within short distance carry nutritional costs. The ultimate fitness cost will depend on the frequency of predator–prey encounters and on whether bottom-up or top-down forces are more important in the prey population. Our finding is the first attempt to our knowledge to assess nutritionally mediated risk effects in a large mammalian prey species under the threat of an ambush predator, and brings support to the hypothesis that the behavioural effects of predation induce important risk effects on prey populations.     

A ‘dynamic’ landscape of fear: prey responses to spatiotemporal variations in predation risk across the lunar cycle

Ambiguous empirical support for ‘landscapes of fear’ in natural systems may stem from failure to consider dynamic temporal changes in predation risk. The lunar cycle dramatically alters night‐time visibility, with low luminosity increasing hunting success of African lions. We used camera‐trap data from Serengeti National Park to examine nocturnal anti‐predator behaviours of four herbivore species. Interactions between predictable fluctuations in night‐time luminosity and the underlying risk‐resource landscape shaped herbivore distribution, herding propensity and the incidence of ‘relaxed’ behaviours. Buffalo responded least to temporal risk cues and minimised risk primarily through spatial redistribution. Gazelle and zebra made decisions based on current light levels and lunar phase, and wildebeest responded to lunar phase alone. These three species avoided areas where likelihood of encountering lions was high and changed their behaviours in risky areas to minimise predation threat. These patterns support the hypothesis that fear landscapes vary heterogeneously in both space and time.     

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.     

Combined climate‐ and prey‐mediated range expansion of Humboldt squid (Dosidicus gigas), a large marine predator in the California Current System

Climate‐driven range shifts are ongoing in pelagic marine environments, and ecosystems must respond to combined effects of altered species distributions and environmental drivers. Hypoxic oxygen minimum zones (OMZs) in midwater environments are shoaling globally; this can affect distributions of species both geographically and vertically along with predator–prey dynamics. Humboldt (jumbo) squid (Dosidicus gigas) are highly migratory predators adapted to hypoxic conditions that may be deleterious to their competitors and predators. Consequently, OMZ shoaling may preferentially facilitate foraging opportunities for Humboldt squid. With two separate modeling approaches using unique, long‐term data based on in situ observations of predator, prey, and environmental variables, our analyses suggest that Humboldt squid are indirectly affected by OMZ shoaling through effects on a primary food source, myctophid fishes. Our results suggest that this indirect linkage between hypoxia and foraging is an important driver of the ongoing range expansion of Humboldt squid in the northeastern Pacific Ocean.     

Proximate stimuli: An overlooked driving force for risk-induced trait responses affecting interactions in aquatic ecosystems

Inducible responses in prey to predation risk can influence species interaction strength, with significant ecological consequences. Much of the past research on interactions in aquatic ecosystems has focused on remote stimuli (e.g., diffusible chemicals emitted from predators and injured conspecifics, which easily propagate through environmental water), as cues triggering trait responses in prey, and has overlooked the importance of proximate stimuli (e.g., physical disturbance and less-diffusible chemicals), which occur in attack or direct contact to prey by predators. Proximate stimuli from predators as well as remote stimuli may induce significant responses in prey functional traits such as behavior, morphology, and life history and, therefore, act as an important mechanism of top-down effects in aquatic ecosystems. In this opinion paper, we argue that studying the effects of proximate stimuli is essential to better understanding of individual adaptation to predation risk in nature and ecological consequences of predator–prey interactions. Here, we propose research directions to examine the role of proximate stimuli for phenotypic plasticity and interaction systems.     

Do the antipredator strategies of shared prey mediate intraguild predation and mesopredator suppression?

Understanding the conditions that facilitate top predator effects upon mesopredators and prey is critical for predicting where these effects will be significant. Intraguild predation (IGP) and the ecology of fear are hypotheses used to describe the effects of top predators upon mesopredators and prey species, but make different assumptions about organismal space use. The IGP hypothesis predicts that mesopredator resource acquisition and risk are positively correlated, creating a fitness deficit. But if shared prey also avoid a top predator, then mesopredators may not have to choose between risk and reward. Prey life history may be a critical predictor of how shared prey respond to predation and may mediate mesopredator suppression. We used hierarchical models of species distribution and abundance to test expectations of IGP using two separate triangular relationships between a large carnivore, smaller intraguild carnivore, and shared mammalian prey with different life histories. Following IGP, we expected that a larger carnivore would suppress a smaller carnivore if the shared prey species did not spatially avoid the large carnivore at broad scales. If prey were fearful over broad scales, we expected less evidence of mesopredator suppression. We tested these theoretical hypotheses using remote camera detections across a large spatial extent. Lagomorphs did not appear to avoid coyotes, and fox detection probability was lower as coyote abundance increased. In contrast, white‐tailed deer appeared to avoid areas of increased wolf use, and coyote detection probability was not reduced at sites where wolves occurred. These findings suggest that mesopredator suppression by larger carnivores may depend upon the behavior of shared prey, specifically the spatial scale at which they perceive risk. We further discuss how extrinsic environmental factors may contribute to mesopredator suppression.     

Elephants facilitate impact of large predators on small ungulate prey species

Predators and megaherbivores have profound impacts on ecosystem structure and functioning. Following the reintroduction of apex predators (lion and spotted hyaena) into the Main Camp Section (Main Camp) of the Addo Elephant National Park (Addo – Eastern Cape, South Africa) populations of small (5–50 kg) prey species declined. Following the recent reintroduction of apex predators into the neighbouring Colchester Section, a similar decline in small prey species might have occurred. However, we predict that the dense nature of Thicket vegetation in Colchester has provided the small prey species a refuge from predation from the reintroduced apex predators. Using camera trap data collected over three years following the apex predator reintroduction into Colchester, we show that declines in small ungulate prey species have not taken place. The primary difference between these two sections at the time of the apex predator reintroduction was the state of the Thicket (dense vegetation type characteristic of both sections). Main Camp is characterized by fragmented Thicket that has been altered as a result of high elephant densities, whereas Colchester has intact Thicket following a long history of elephant absence. The fragmented Thicket in Main Camp allowed access to the Thicket habitats (as indicated by GPS collar data on lions), in which these small prey species reside, potentially increasing the predation on these species in Main Camp. The intact Thicket in Colchester, however, may provide a refuge away from the apex predators (and possibly meso-predators) for the small prey species. We suggest that the impact of predators on this prey community is conditional on the long history of ecosystem transformation by ecosystem engineers such as elephants.     

The Lion King and the Hyaena Queen: large carnivore interactions and coexistence

Interactions among species, which range from competition to facilitation, have profound effects on ecosystem functioning. Large carnivores are of particular importance in shaping community structure since they are at the top of the food chain, and many efforts are made to conserve such keystone species. Despite this, the mechanisms of carnivore interactions are far from understood, yet they are key to enabling or hindering their coexistence and hence are highly relevant for their conservation. The goal of this review is thus to provide detailed information on the extents of competition and facilitation between large carnivores and their impact in shaping their life histories. Here, we use the example of spotted hyaenas (Crocuta crocuta) and lions (Panthera leo) and provide a comprehensive knowledge of their interactions based on meta‐analyses from available literature (148 publications). Despite their strong potential for both exploitation and interference competition (range and diet overlap, intraguild predation and kleptoparasitism), we underline some mechanisms facilitating their coexistence (different prey‐age selection and scavenging opportunities). We stress the fact that prey abundance is key to their coexistence and that hyaenas forming very large groups in rich ecosystems could have a negative impact on lions. We show that the coexistence of spotted hyaenas and lions is a complex balance between competition and facilitation, and that prey availability within the ecosystem determines which predator is dominant. However, there are still many gaps in our knowledge such as the spatio‐temporal dynamics of their interactions. As both species' survival becomes increasingly dependent on protected areas, where their densities can be high, it is critical to understand their interactions to inform both reintroduction programs and protected area management.     

Mechanisms of coexistence in diverse herbivore–carnivore assemblages: demographic,temporal and spatial heterogeneities affecting prey vulnerability

Simple models coupling the dynamics of single predators to single prey populations tend to generate oscillatory dynamics of both predator and prey, or extirpation of the prey followed by that of the predator. In reality, such oscillatory dynamics may be counteracted by prey refugia or by opportunities for prey switching by the predator in multi‐prey assemblages. How these mechanisms operate depends on relative prey vulnerability, a factor ignored in simple interactive models. I outline how compositional, temporal, demographic and spatial heterogeneities help explain the contrasting effects of top predators on large herbivore abundance and population dynamics in species‐rich African savanna ecosystems compared with less species‐diverse northern temperate or subarctic ecosystems. Demographically, mortality inflicted by predation depends on the relative size and life history stage of the prey. Because all animals eventually die and are consumed by various carnivores, the additive component of the mortality inflicted is somewhat less than the predation rate. Prey vulnerability varies annually and seasonally, and between day and night. Spatial variation in the risk of predation depends on vegetation cover as well as on the availability of food resources. During times of food shortage, herbivores become prompted to occupy more risky habitats retaining more food. Predator concentrations dependent on the abundance of primary prey species may restrict the occurrence of other potential prey species less resistant to predation. The presence of multiple herbivore species of similar size in African savannas allows the top predator, the lion, to shift its prey selection flexibly dependent on changing prey vulnerability. Hence top–down and bottom–up influences on herbivore populations are intrinsically entangled. Models coupling the population dynamics of predators and prey need to accommodate the changing influences of prey demography, temporal variation in environmental conditions, and spatial variation in the relative vulnerability of alternative prey species to predation. Synthesis While re‐established predators have had major impacts on prey populations in northern temperate regions, multiple large herbivore species typically coexist along with diverse carnivores in African savanna ecosystems. In order to explain these contrasting outcomes, certain functional heterogeneities must be recognised, including relative vulnerability of alternative prey, temporal variation in the risk of predation, demographic differences in susceptibility to predation, and spatial contrasts in exposure to predation. Food shortfalls prompt herbivores to exploit more risky habitats, meaning that top–down and bottom–up influences on prey populations are intrinsically entangled. Models coupling the interactive dynamics of predator and prey populations need to incorporate these varying influences on relative prey vulnerability.     

Predators can influence the host‐parasite dynamics of their prey via nonconsumptive effects

Ecological communities are partly structured by indirect interactions, where one species can indirectly affect another by altering its interactions with a third species. In the absence of direct predation, nonconsumptive effects of predators on prey have important implications for subsequent community interactions. To better understand these interactions, we used a Daphnia‐parasite‐predator cue system to evaluate if predation risk affects Daphnia responses to a parasite. We investigated the effects of predator cues on two aspects of host–parasite interactions (susceptibility to infection and infection intensity), and whether or not these effects differed between sexes. Our results show that changes in response to predator cues caused an increase in the prevalence and intensity of parasite infections in female predator‐exposed Daphnia. Importantly, the magnitude of infection risk depended on how long Daphnia were exposed to the cues. Additionally, heavily infected Daphnia that were constantly exposed to cues produced relatively more offspring. While males were ~5× less likely to become infected compared to females, we were unable to detect effects of predator cues on male Daphnia–parasite interactions. In sum, predators, prey, and their parasites can form complex subnetworks in food webs, necessitating a nuanced understanding of how nonconsumptive effects may mediate these interactions.     

Nature of Predation Risk Cues in Container Systems: Mosquito Responses to Solid Residues From Predation

In aquatic systems, prey animals associate predation risk with cues that originate either from the predator or from injured conspecifics. Sources and benefits of these cues have received considerable attention in river, lake, and pond ecosystems but are less well understood in small container ecosystems that can hold less than a liter of water. Mosquitoes Aedes triseriatus (Say) and Aedes albopictus (Skuse) encounter predatory Corethrella appendiculata (Grabham) and Toxorhynchites rutilus (Coquillett) in small containers and show antipredatory behavioral responses. We investigated the sources of the predation cues to which these prey larvae respond. We tested whether Ae. albopictus larvae show behavioral responses to cues emanating from the predator or from damage to prey caused by the act of predation. We also tested whether Ae. triseriatus respond to cues present in fluid or solid residues from predator activity. Ae. albopictus showed behavioral modifications only in response to waterborne cues from a feeding predator and not to cues from a starving predator, indicating that Ae. albopictus respond to cues created by the act of predation, which could include substances derived from damaged prey or substances in predator feces. Ae. triseriatus showed behavioral responses to solid residues from predation but not to fluid without those solids, indicating that the cues to which they respond originate in predator feces or uneaten prey body parts. Our results suggest that cues in this system may be primarily chemicals that are detected upon contact with solid residues that are products of the feeding processes of these predators.     

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.     

Dragonfly larvae and tadpole frog space use games in varied light conditions

Predators and prey often engage in a game where predators attemptto be in areas with higher prey densities and prey attempt tobe in areas with lower predator densities. A few models havepredicted the resulting distributions of predators and prey,but little empirical data exist to test these predictions andto examine how abiotic and biotic factors shape the distributions.Thus, we observed how Anax dragonfly nymphs and Pacific treefrog tadpoles (Pseudacris regilla) either together or separatelydistributed themselves in an arena with a high- and a low-preyresource patch. Trials were conducted in high- and low-lightconditions to manipulate predation risk and to view the effectsof this abiotic factor. Counter to the model predictions, wefound that predators were not more abundant in high-resource(HR) patches, and they thus did not force prey toward beinguniformly distributed. Using a model selection approach to assesswhat factors affected predator and prey patch-switching movement,we found that prey more often left patches that had more predatorspresent, but predators surprisingly more often left patcheswith more prey present. Light levels did not affect predationrisk; however, in the dark with the associated reduction invisual information predators preferred HR patches. This causeda lower coincidence of prey and predators in patches. Predatorsalso switched patches less often when they occupied the samepatch as the other predator. This suggests that predator distributions,and indirectly prey distributions, are affected by the riskof intraguild predation.     

Anthropogenic influences on primate antipredator behavior and implications for research and conservation

Predation risk affects prey species' behavior, even in the absence of a direct threat, but human-induced environmental change may disturb ecologically significant predator–prey interactions. Here, we propose various ways in which knowledge of antipredator tactics, behavioral risk effects, and primate–predator interactions could assist in identifying human-caused disruption to natural systems. Using behavior to evaluate primate responses to the ongoing environmental change should be a potentially effective way to make species conservation more predictive by identifying issues before a more dramatic population declines. A key challenge here is that studies of predation on primates often use data collected via direct observations of habituated animals and human presence can deter carnivores and influence subjects' perception of risk. Hence, we also review various indirect data collection methods to evaluate their effectiveness in identifying where environmental change threatens wild species, while also minimizing observer bias.     

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.