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Summary Escaping from a predator is a matter of life or death, and prey are expected to adaptively alter their physiology under chronic predation risk in ways that may affect escape. Theoretical models assume that escape performance is mass dependent, whereby scared prey strategically maintain an optimal body mass to enhance escape. Experiments testing the mass‐dependent predation risk hypothesis have demonstrated that prior experience of predation risk can affect body mass, and the behavioural decisions about evasive actions to take. Other studies on natural changes in body mass indicate that mass can affect escape. No single experiment has tested if all of these components are indeed linked, which is a critical necessary condition underpinning the mass‐dependent predation risk hypothesis. We tested all components of the mass‐dependent predation risk hypothesis in a repeated measures experiment by presenting predator and non‐predator cues to brown‐headed cowbirds housed in semi‐natural conditions. Exposure to predator cues affected body mass, fat, pectoral muscle thickness and evasive actions (take‐off angle and speed), but not the physiological capacity to escape, as measured by flying ability. Examining individual variation revealed that flying ability was unrelated to mass loss in either sex, unrelated to mass gain in males, and only females that gained a very large amount of mass flew poorly. We next conducted a body mass manipulation in the laboratory to rigorously test whether small to large perturbations in mass can ever affect flying ability. We induced either no change in mass (control), a moderate reduction of <10% or a more extreme reduction of >10% which the literature suggests should enhance flight. Flying ability was maintained regardless of treatment. Examining individual variation revealed the same precise patterns as in the first experiment. We conclude that prey may alter their mass and evasive actions in response to predation risk, but their escape ability remains robust and inelastic, presumably because disabling oneself is likely to lead to disastrous consequences. We suggest that animals may only face a mass‐dependent predation risk trade‐off in a narrow set of circumstances linked to life‐history stages that require large amounts of mass gain, for example, parturition and migration. A lay summary is available for this article. Lay Summary

The very presence of predators can strongly influence flexible prey traits such as behavior, morphology, life history, and physiology. In a rapidly growing body of literature representing diverse ecological systems, these trait (or “fear”) responses have been shown to influence prey fitness components and density, and to have indirect effects on other species. However, this broad and exciting literature is burdened with inconsistent terminology that is likely hindering the development of inclusive frameworks and general advances in ecology. We examine the diverse terminology used in the literature, and discuss pros and cons of the many terms used. Common problems include the same term being used for different processes, and many different terms being used for the same process. To mitigate terminological barriers, we developed a conceptual framework that explicitly distinguishes the multiple predation-risk effects studied. These multiple effects, along with suggested standardized terminology, are risk-induced trait responses (i.e., effects on prey traits), interaction modifications (i.e., effects on prey–other-species interactions), nonconsumptive effects (i.e., effects on the fitness and density of the prey), and trait-mediated indirect effects (i.e., the effects on the fitness and density of other species). We apply the framework to three well studied systems to highlight how it can illuminate commonalities and differences among study systems. By clarifying and elucidating conceptually similar processes, the framework and standardized terminology can facilitate communication of insights and methodologies across systems and foster cross-disciplinary perspectives

For foraging herbivores, both food quality and predation risk vary across the landscape. Animals should avoid low-quality food patches in favour of high-quality ones, and seek safe patches while avoiding risky ones. Herbivores often face the foraging dilemma, however, of choosing between high-quality food in risky places or low-quality food in safe places. Here, we explore how and why the interaction between food quality and predation risk affects foraging decisions of mammalian herbivores, focusing on browsers confronting plant toxins in a landscape of fear. We draw together themes of plant–herbivore and predator–prey interactions, and the roles of animal ecophysiology, behaviour and personality. The response of herbivores to the dual costs of food and fear depends on the interplay of physiology and behaviour. We discuss detoxification physiology in dealing with plant toxins, and stress physiology associated with perceived predation risk. We argue that behaviour is the interface enabling herbivores to stay or quit food patches in response to their physiological tolerance to these risks. We hypothesise that generalist and specialist herbivores perceive the relative costs of plant defence and predation risk differently and intra-specifically, individuals with different personalities and physiologies should do so too, creating individualised landscapes of food and fear. We explore the ecological significance and emergent impacts of these individual-based foraging outcomes on populations and communities, and offer predictions that can be clearly tested. In doing so, we provide an integrated platform advancing herbivore foraging theory with food quality and predation risk at its core.

Adaptive theory predicts that the fundamental trade-off between starvation and predation risk shapes diurnal patterns in foraging activity and mass gain in wintering passerine birds. Foragers mitigating both types of risk should exhibit a bimodal distribution (increased foraging and mass gain early and late in the day), whereas both foraging and mass gains early (versus late) during the day are expected when the risk of starvation (versus predation) is greatest. Finally, relatively constant rates of foraging and mass gain should occur when the starvation–predation risk trade-off is independent of body mass. Using automated feeders with integrated digital balances, we estimated diurnal patterns in foraging and body mass gain to test which ecological scenario was best supported in wintering great tits Parus major. Based on data of 40 consecutive winter days recording over 12 000 body masses of 28 individuals, we concluded that birds foraged and gained mass early during the day, as predicted by theory when the starvation–predation risk trade-off is mass-dependent and starvation risk outweighs predation risk. Slower explorers visited the feeders more often, and decreased their activity along the day more strongly, compared with faster explorers, thereby explaining a major portion of the individual differences in diurnal patterning of foraging activity detected using random regression analyses. Birds did not differ in body mass gain trajectories, implying both that individuals differed in the usage of feeders, and that unbiased conclusions regarding how birds resolve starvation–predation risk trade-off require the simultaneous recording of foraging activity and body mass gain trajectories. Our study thereby provides the first unambiguous demonstration that individual birds are capable of adjusting their diurnal foraging and mass gain trajectories in response to ecological predictors of starvation risk as predicted by starvation–predation risk trade-off theory.

Predation is a strong ecological force that shapes animal communities through natural selection. Recent studies have shown the cascading effects of predation risk on ecosystems through changes in prey behavior. Minimizing predation risk may explain why multiple prey species associate together in space and time. For example, mixed-species flocks that have been widely documented from forest systems, often include birds that eavesdrop on sentinel species (alarm calling heterospecifics). Sentinel species may be pivotal in (1) allowing flocking species to forage in open areas within forests that otherwise incur high predation risk, and (2) influencing flock occurrence (the amount of time species spend with a flock). To test this, we conducted a short-term removal experiment in an Amazonian lowland rainforest to test whether flock habitat use and flock occurrence was influenced by sentinel presence. Antshrikes (genus Thamnomanes) act as sentinels in Amazonian mixed-species flocks by providing alarm calls widely used by other flock members. The alarm calls provide threat information about ambush predators such as hawks and falcons which attack in flight. We quantified home range behavior, the forest vegetation profile used by flocks, and the proportion occurrence of other flocking species, both before and after removal of antshrikes from flocks. We found that when sentinel species were removed, (1) flock members shifted habitat use to lower risk habitats with greater vegetation cover, and (2) species flock occurrence decreased. We conclude that eavesdropping on sentinel species may allow other species to expand their realized niche by allowing them to safely forage in high-risk habitats within the forest. In allowing species to use extended parts of the forest, sentinel species may influence overall biodiversity across a diverse landscape.

Fear of predation produces large effects on prey population dynamics through indirect risk effects that can cause even greater impacts than direct predation mortality. As yet, there is no general theoretical framework for predicting when and how these population risk effects will arise in specific prey populations, meaning that there is often little consideration given to the key role predator risk effects can play in understanding conservation and wildlife management challenges. Here, we propose that population predator risk effects can be predicted through an extension of individual risk trade‐off theory and show for the first time that this is the case in a wild vertebrate system. Specifically, we demonstrate that the timing (in specific months of the year), occurrence (at low food availability), cause (reduction in individual energy reserves), and type (starvation mortality) of a population‐level predator risk effect can be successfully predicted from individual responses using a widely applicable theoretical framework (individual‐based risk trade‐off theory). Our results suggest that individual‐based risk trade‐off frameworks could allow a wide range of population‐level predator risk effects to be predicted from existing ecological theory, which would enable risk effects to be more routinely integrated into consideration of population processes and in applied situations such as conservation.

Predation may indirectly influence prey's fitness and population dynamics through behavioural adjustments in response to perceived predation risk. These non‐consumptive effects of predation can also arise from hunting by humans, but they remain less documented. Advances in biologging allow detailed assessments of the activity budgets of elusive wildlife, increasing the potential to uncover the non‐consumptive effects of human activities on animals. We used tri‐axial accelerometry to record the daily activity of 24 Scandinavian brown bears (20 females and 4 males) from a heavily hunted population in Sweden, for a total of 29 bear‐years (2015–2022). We used a random forest algorithm trained with observations of captive brown bears to classify the accelerometry data into four behaviours, running, walking, feeding and resting, with an overall precision of 95%. We then used these classifications to evaluate changes in bear activity budgets before and during the hunting season. Bears exhibited a bimodal daily activity pattern, being most active at dusk and dawn and resting around midday and midnight. However, during the hunting season, males became more nocturnal compared to before the hunting season, suggesting a proactive behavioural adjustment to reduce encounters with hunters. Females showed the opposite pattern and had a higher probability of being active during the day, potentially to increase nutritional gains before denning. Additionally, daily number of running bouts did not vary between the pre‐hunting and hunting seasons in both sexes, but females' proportion of running bouts occurring during legal hunting hours was higher during the hunting season than prior to it, which suggests a reactive behavioural adjustment to encounters with hunters. Detailed assessments of wild animal behaviours, allowed through recording of movement data at high frequencies, have the potential to improve our understanding of the impacts of human activity on wildlife. We used accelerometry data to evaluate behavioural adjustments to temporal variation in hunting risk in Scandinavian brown bears. We find that male bears become more nocturnal after the onset of the hunting season, while female bears do the opposite by becoming more diurnal. These results suggest sex‐differences in brown bear responses to hunting risk.

An increase in predation risk triggers a trait response of prey, which alters the interactions between the prey and other species, ultimately affecting other species in the ecosystem. Such predator‐driven trait‐mediated indirect effects (TMIEs) may have been shaped by long‐term evolutionary processes involving the organisms involved, but learning may also be important, especially in contemporary ecosystems experiencing repeated biological invasions. The apple snail Pomacea canaliculata is an important introduced pest of rice, Oryza sativa. Recently, the carrion crow Corvus corone has been found to prey on this species only in some areas, suggesting that learning is involved in this predation. In addition, apple snails can learn to escape from predators and exhibit predator‐specific responses. Thus, the “chain of learning” by the crow and the snail may shape novel TMIEs in the rice ecosystem. We conducted field and mesocosm experiments to test this hypothesis. In the field experiment, we simulated predation by crows in rice fields and investigated the behavior of apple snails. The snails exhibited escape behaviors in response to the simulated predation, and both the proportion of individuals showing the escape response and the degree of escape response were greater in fields with predation by crows than those without predation. In the mesocosm experiment, apple snails from fields with and without predation by crows were separately introduced into mesocosms simulating rice fields, and the behaviors of the snails and the number of remaining rice plants were recorded for 16 days at three levels of predation risk (daily, every 4 days, or no predation). Both the presence/absence of predation in the collection fields and simulated predation affected the escape responses of the snails. Moreover, damage to rice was more severe in mesocosms containing snails from fields without predation than those containing snails from fields with predation. These results suggest that the “chain of learning” causes TMIEs in agroecosystems.

Prey often retreat into the safety of refuges for protection from predators. This shift into refuge can reduce foraging opportunities, escalating the costs of risk and the strength of nonconsumptive effects. Such costs, however, may be shaped by the variation in resources that refuges harbor for prey foraging (i.e., refuge quality), and change dynamically via impacts on prey state. Despite its potential importance, we lack an explicit understanding of how refuge quality impacts prey performance under risk. Using a rocky intertidal food chain, we examined the interaction between predation risk and the amount of resources available for prey in refuge. We found that refuges with more resources greatly reduce the costs of refuge use, and that nonconsumptive effects are thereby weakened by as much as one-half, with especially strong impacts on prey growth and growth efficiency. These results suggest that failure to consider refuge quality could result in overestimation of the negative effects associated with prey refuge use.

The ecological impacts of predation risk are influenced by how prey allocate foraging effort across periods of safety and danger. Foraging decisions depend on current danger, but also on the larger temporal, spatial or energetic context in which prey manage their risks of predation and starvation. Using a rocky intertidal food chain, we examined the responses of starved and fed prey (Nucella lapillus dogwhelks) to different temporal patterns of risk from predatory crabs (Carcinus maenas). Prey foraging activity declined during periods of danger, but as dangerous periods became longer, prey state altered the magnitude of risk effects on prey foraging and growth, with likely consequences for community structure (trait-mediated indirect effects on basal resources, Mytilus edulis mussels), prey fitness and trophic energy transfer. Because risk is inherently variable over time and space, our results suggest that non-consumptive predator effects may be most pronounced in productive systems where prey can build energy reserves during periods of safety and then burn these reserves as ‘trophic heat’ during extended periods of danger. Understanding the interaction between behavioural (energy gain) and physiological (energy use) responses to risk may illuminate the context dependency of trait-mediated trophic cascades and help explain variation in food chain length.