Foraging and refuge use by a pond snail: Effects of physiological state,predators, and resources

The costs and benefits of anti-predator behavioral responses should be functions of the actual risk of predation, the availability of the prey's resources, and the physiological state of the prey. For example, a food-stressed individual risks starvation when hiding from predators, while a well-fed organism can better afford to hide (and pay the cost of not foraging). Similarly, the benefits of resource acquisition are probably highest for the prey in the poorest state, while there may be diminishing returns for prey nearing satiation. Empirical studies of state-dependent behavior are only beginning, however, and few studies have investigated interactions between all three potentially important factors. Here I present the results of a laboratory experiment where I manipulated the physiological state of pond snails (Physa gyrina), the abundance of algal resources, and predation cues (Belostoma flumineum waterbugs consuming snails) in a full factorial design to assess their direct effects on snail behavior and indirect effects on algal biomass. On average, snails foraged more when resources were abundant, and when predators were absent. Snails also foraged more when previously exposed to physiological stress. Snails spent more time at the water's surface (a refuging behavior) in the presence of predation cues on average, but predation, resource levels, and prey state had interactive effects on refuge use. There was a consistent positive trait-mediated indirect effect of predators on algal biomass, across all resource levels and prey states.     

The control of risk hypothesis: reactive vs. proactive antipredator responses and stress‐mediated vs. food‐mediated costs of response

Inducible defences against predators evolve because they reduce the rate of direct predation, but this benefit is offset by the cost (if any) of defence. If antipredator responses carry costs, the effect of predators on their prey is partitioned into two components, direct killing and risk effects. There is considerable uncertainty about the strength of risk effects, the factors that affect their strength, and the mechanisms that underlie them. In some cases, antipredator responses are associated with a glucocorticoid stress response, and in other cases they are associated with trade‐offs between food and safety, but there is no general theory to explain this variation. Here, I develop the control of risk (COR) hypothesis, predicting that proactive responses to predictable and controllable aspects of risk will generally have food‐mediated costs, while reactive responses to unpredictable or uncontrollable aspects of predation risk will generally have stress‐mediated costs. The hypothesis is grounded in laboratory studies of neuroendocrine stressors and field studies of food‐safety trade‐offs. Strong tests of the COR hypothesis will require more studies of responses to natural variation in predation risk and the physiological consequences of these responses, but its explanatory power can be illustrated with existing case studies.     

A review of the sentinel prey method as a way of quantifying invertebrate predation under field conditions

Sentinel prey can provide a direct, quantitative measure of predation under field conditions. Live sentinel prey provides more realistic data but rarely allows the partitioning of the total predation pressure; artificial prey is less natural but traces left by different predators are identifiable, making it suitable for comparative studies. We reviewed the available evidence of the use of both types of invertebrate sentinel prey. Fifty‐seven papers used real prey, usually measuring predation on a focal (often pest) species, with studies overwhelmingly from North America. The median predation was 25.8% d^?1. Artificial sentinel prey (45 papers) were used in both temperate and tropical areas, placed more above ground than at ground level. The most commonly used artificial prey imitated a caterpillar. Up to 14 predator groups were identified, registering a median of 8.8% d^?1 predation; half the studies reported only bird predation. Predation on real prey was higher than on artificial ones, but invertebrate predation was not higher than vertebrate predation. Invertertebrate but not vertebrate predation was negatively related to prey size. Predation near the Equator was not higher than at higher latitudes, nor in cultivated than noncultivated habitats. The use of sentinel prey is not yet standardised in terms of prey size, arrangement, exposure period or data reporting. Due to the simplicity and ease of use of the method, such standardisation may increase the usefulness of comparative studies, contributing to the understanding of the importance and level of predation in various habitats worldwide.     

The influence of temporally variable predation risk on indirect interactions in an aquatic food chain

We know little about how temporally variable predation risk influences prey behavior. The risk allocation hypothesis predicts that prey facing more frequent risk should show weak anti-predator responses, and should be particularly active foragers during rare periods of safety, compared to prey facing infrequent risk. Several studies offer support for the risk allocation hypothesis, but how these responses might propagate through the larger ecological community remains largely unknown. We experimentally investigated the relative strength of trait- and density-mediated indirect effects of a predator on its prey’s resource across predation treatments that varied the lethality (caged or free-swimming predators) and temporal variability (always, often, or sometimes present) of predation. We performed this experiment in pond mesocosms using a giant water bug predator (Belostoma lutarium), an herbivorous pond snail (Physa gyrina), and algae as the basal resource. Snails greatly reduced the abundance of their algal resource when in the absence of predation. Lethal predation at low and medium intensities had significant positive indirect effects on the abundance of algae, mostly by reducing snail density. Snails responded behaviorally to high levels of deadly predation by foraging more and hiding less than in other situations, as predicted by the risk allocation hypothesis, and thus ameliorated the density-mediated indirect effects of predators on algae. Behavioral responses to caged predators, and the subsequent trait-mediated indirect effects, were negligible regardless of predation intensity. Our previous work has demonstrated that trait-mediated indirect effects are weak when resources are abundant, as they were in this experiment. This work demonstrates that temporal variation in predation intensity plays a key role in determining the relative strength of TMIIs and DMIIs in an aquatic food chain.     

Predation risk perception and food scarcity induce alterations of life-cycle traits of the mosquito Culex pipiens

Abstract.  1. Predators may affect prey populations by direct consumption, and by inducing defensive reactions of prey to the predation risk. Food scarcity frequently has effects on the inducible defences of prey, but no consistent pattern of food–predation risk interaction is known.
2. In this study the combined effect of food shortage and predation-risk perception in larvae of the mosquito Culex pipiens was investigated. Water exposed to the aquatic predator bug Notonecta glauca was used as a source of predation intimidation. Mosquito larvae were reared in three different media containing either no predator cues or the cues of N. glauca that had been fed on either C. pipiens larvae or on Daphnia magna . Food was provided in favourable or limited amount for these set-ups.
3. The results showed that chemical cues from the predators fed with prey's conspecifics caused a decreased survival, delayed pre-imaginal development, and reduction in body size of emerged mosquitoes, whereas chemical cues from predators fed with D. magna caused only delayed development. Food scarcity significantly exacerbates the negative effect of the predator cues on pre-imaginal development of C. pipiens . Effects of the cues on larval development and body size of imagoes are significantly stronger for females than for males.
4. The present study suggests that when food is limited, predators can affect population dynamics of prey not only by direct predation, but also by inducing lethal and sublethal effects due to perception of risk imposed by chemical cues. To understand the effects of predators on mosquito population dynamics, environmental parameters such as food deficiency should be considered.     

Under pressure: Short- and long-term response to predation varies in two populations of a live-bearing fish

The non-lethal effects of predation can significantly influence animal behavior and population composition. Research has often centered around prey response to predator exposure in the short term, but fewer studies have highlighted the effects of long-term predator exposures. In addition, studies of responses to predation risk are not always calibrated against the ecological history of predation risk in specific populations. We address these gaps by examining the effects of both long- and short-term predator exposure on the behavior of individuals from populations that have different ecological histories of predation risk. We exposed individuals from high-predation and low-predation populations of the live-bearing freshwater poeciliid, Heterandria formosa, to predators to assess changes in male reproductive behavior toward females. We also assessed longer-term reproductive responses by exposing male and female H. formosa to predators at a random time of day, every day, for 30 days. In the presence of a predator, in the short term, males changed the frequency of their behaviors and females varied in their concentration of cortisol, demonstrating immediate responses to the perceived risk. The magnitude of these changes was larger in the population without a long history of predator exposure. However, we found that males and females did not change their reproductive output when exposed to predators over longer periods of time, suggesting that individuals acclimatize to the level of predation risk they experience. These results also suggest that short-term variation in behavior or stress hormone responses should not be used as proxies for long-term responses or fitness effects. Future work should assess both short-term behavior and long-term responses while simultaneously considering the ecological history of populations.     

Resource-mediated impact of spider predation risk on performance in the grasshopper Ageneotettix deorum (Orthoptera: Acrididae)

In response to increased exposure to predators when searching for food, many prey increase the frequency of antipredator behaviors, potentially reducing foraging rate and food intake. Such direct, nonlethal interactions between predators and prey resulting in reduced food intake can indirectly influence lifecycle development through effects on growth, developmental rate, and survival. We investigated the general hypothesis that individual performance of a herbivorous insect can be negatively affected when exposed to nonlethal predation risk, and that the response can be mediated by food quality. This hypothesis was tested using the common rangeland grasshopper Ageneotettix deorum with and without exposure to common wolf spider predators (Lycosidae, Schizocosa spp.) on both untreated natural and fertilized vegetation. All spiders were rendered temporarily incapable of direct feeding by restricting function of the chelicerae with beeswax. Detectable responses by grasshoppers to spiders indicate indirect consequences for lifecycle development. Grasshopper performance was measured as hind femur growth, duration of nymphal lifecycle stages, and survivorship in a caged field experiment conducted over 2 years. Grasshoppers developed faster and grew 3–5% larger when allowed to forage on fertilized vegetation in the absence of risk from a spider predator. Failure-time analysis illustrated enhanced survival probability in response to elevated food quality and the negative effects of grasshopper susceptibility to nonlethal predation risk. Performance on food of relatively low, ambient quality with no predation risk equaled that of grasshoppers caged with high quality vegetation in the presence of a modified spider. Increased resource quality can clearly moderate the negative life history responses caused by the behavioral modification of grasshoppers when exposed to spider predation risk, a compensatory response.     

Interactive effects of predation risk and conspecific density on the nutrient stoichiometry of prey

The mere presence of predators (i.e., predation risk) can alter consumer physiology by restricting food intake and inducing stress, which can ultimately affect prey‐mediated ecosystem processes such as nutrient cycling. However, many environmental factors, including conspecific density, can mediate the perception of risk by prey. Prey conspecific density has been defined as a fundamental feature that modulates perceived risk. In this study, we tested the effects of predation risk on prey nutrient stoichiometry (body and excretion). Using a constant predation risk, we also tested the effects of varying conspecific densities on prey responses to predation risk. To answer these questions, we conducted a mesocosm experiment using caged predators (Belostoma sp.), and small bullfrog tadpoles (Lithobates catesbeianus) as prey. We found that L. catesbeianus tadpoles adjust their body nutrient stoichiometry in response to predation risk, which is affected by conspecific density. We also found that the prey exhibited strong morphological responses to predation risk (i.e., an increase in tail muscle mass), which were positively correlated to body nitrogen content. Thus, we pose the notion that in risky situations, adaptive phenotypic responses rather than behavioral ones might partially explain why prey might have a higher nitrogen content under predation risk. In addition, the interactive roles of conspecific density and predation risk, which might result in reduced perceived risk and physiological restrictions in prey, also affected how prey stoichiometry responded to the fear of predation.     

Diagnosing predation risk effects on demography: can measuring physiology provide the means?

Predators kill prey thereby affecting prey survival and, in the traditional top-down view of predator limitation, that is their sole effect. Bottom-up food limitation alters the physiological condition of individuals affecting both fecundity and survival. Predators of course also scare prey inducing anti-predator defences that may carry physiological costs powerful enough to reduce prey fecundity and survival. Here, we consider whether measuring physiology can be used as a tool to unambiguously diagnose predation risk effects. We begin by providing a review of recent papers reporting physiological effects of predation risk. We then present a conceptual framework describing the pathways by which predators and food can affect prey populations and give an overview of predation risk effects on demography in various taxa. Because scared prey typically eat less the principal challenge we see will be to identify measures that permit us to avoid mistaking predator-induced reductions in food intake for absolute food shortage. To construct an effective diagnostic toolkit we advocate collecting multiple physiological measures and utilizing multivariate statistical procedures. We recommend conducting two-factor predation risk × food manipulations to identify those physiological effects least likely to be mistaken for responses to bottom-up food limitation. We suggest there is a critical need to develop a diagnostic tool that can be used when it is infeasible to experimentally test for predation risk effects on demography, as may often be the case in wildlife conservation, since failing to consider predation risk effects may cause the total impact of predators to be dramatically underestimated.     

Anurans as prey: an exploratory analysis and size relationships between predators and their prey

The vertebrate predators of post-metamorphic anurans were quantified and the predator–prey relationship was investigated by analysing the relative size of invertebrate predators and anurans. More than 100 vertebrate predators were identified (in more than 200 reports) and classified as opportunistic, convenience, temporary specialized and specialized predators. Invertebrate predators were classified as solitary non-venomous, venomous and social foragers according to 333 reviewed reports. Each of these categories of invertebrate predators was compared with the relative size of the anurans, showing an increase in the relative size of the prey when predators used special predatory tactics. The number of species and the number of families of anurans that were preyed upon did not vary with the size of the predator, suggesting that prey selection was not arbitrary and that energetic constraints must be involved in this choice. The relatively low predation pressure upon brachycephalids was related to the presence of some defensive strategies of its species. This compounding review can be used as the foundation for future advances in vertebrate predator–prey interactions.     

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.     

Effects of Structural Refuge and Density on Foraging Behaviour and Mortality of Hungry Tadpoles Subject to Predation Risk

Theoretical models of prey behaviour predict that food‐limited prey engage in risk‐prone foraging and thereby succumb to increased mortality from predation. However, predation risk also may be influenced by factors including prey density and structural cover, such that the presumed role of prey hunger on predation risk may be obfuscated in many complex predator–prey systems. Using a tadpole (prey) – dragonfly larva (predator) system, we determined relative risk posed to hungry vs. sated prey when both density and structural cover were varied experimentally. Overall, prey response to perceived predation risk was primarily restricted to increased cover use, and hungry prey did not exhibit risk‐prone foraging. Surprisingly, hungry prey showed lower activity than sated prey when exposed to predation risk, perhaps indicating increased effort in search of refuge or spatial avoidance of predator cues among sated animals. An interaction between hunger level and predation risk treatments indicated that prey state affected sensitivity to perceived risk. We also examined the lethal implications of prey hunger by allowing predators to select directly between hungry and sated prey. Although predators qualitatively favoured hungry prey when density was elevated and structural cover was sparse, the overall low observed variation in mortality risk between hunger treatments suggests that preferential selection of hungry prey was weak. This implies that hunger effects on prey mortality risk may not be readily observed in complex landscapes with additional factors influencing risk. Thus, current starvation‐predation trade‐off theory may need to be broadened to account for other mechanisms through which undernourished prey may cope with predation risk.     

Development, growth, and egg production of Ageneotettix deorum (Orthoptera: Acrididae) in response to spider predation risk and elevated resource quality

Abstract.  1. Predation risk to insects is often size- or stage-selective and usually decreases as prey grow. Any factor, such as food quality, that accelerates developmental and growth rates is likely to reduce the period over which prey are susceptible to size-dependent predation.
2. Using field experiments, several hypotheses that assess growth, development, and egg production rates of the rangeland grasshopper Ageneotettix deorum (Scudder) were tested in response to combinations of food quality and predation risk from wolf spiders to investigate performance variation manifested through a behaviourally mediated path affecting food ingestion rates.
3. Grasshoppers with nutritionally superior food completed development ≈ 8–18% faster and grew 15–45% larger in the absence of spiders, in comparison with those subjected to low quality food exposed to spider predators. Growth and development did not differ for grasshoppers feeding on high quality food when predators were present in comparison with lower quality food unimpeded by predators. Responses indicated a compensatory relationship between resource quality and predation risk.
4. Surviving grasshoppers produced fewer eggs compared with individuals not exposed to spiders. Because no differences were found in daily egg production rate regardless of predation treatment, lower egg production was attributed to delayed age of first reproduction. Results compare favourably with responses observed in natural populations.
5. Risk of predation from spiders greatly reduced growth, development, and ultimately egg production. Increased food quality counteracts the impact of predation risk on grasshoppers through compensatory responses, suggesting that bottom-up factors mediate effects of spiders.     

Fear and lethality in snowshoe hares: the deadly effects of non‐consumptive predation risk

Predators play a critical, top–down role in shaping ecosystems, driving prey population and community dynamics. Traditionally, studies of predator‐prey interactions have focused on direct effects of predators, namely the killing of prey. More recently, the non‐consumptive effects of predation risk are being appreciated; e.g. the ‘ecology of fear’. Prey responses to predation risk can be morphological, behavioural, and physiological, and are assumed to come at a cost to prey fitness. However, few studies have examined the relationship between predation risk and survival in wild animals. We tested the hypothesis that predation risk itself could reduce survival in wild‐caught snowshoe hares. We exposed female snowshoe hares to a simulated predator (a trained dog) during gestation only, and measured adult survival and, in surviving females, their ability to successfully wean offspring. We show for the first time in a wild mammal that the risk of predation can itself be lethal. Predation risk reduced adult female survival by 30%, and had trans‐generational effects, reducing offspring survival to weaning by over 85% – even though the period of risk ended at birth. As a consequence of these effects the predator‐exposed group experienced a decrease in number, while the control group substantially increased. Challenges remain in determining the importance of risk‐induced mortality in natural field settings; however, our findings show that non‐lethal predator encounters can influence survival of both adults and offspring. Future work is needed to test these effects in free‐living animals.     

Non-lethal effects of predation in birds

Predators can affect individual fitness and population and community processes through lethal effects (direct consumption or ‘density’ effects), where prey is consumed, or through non‐lethal effects (trait‐mediated effects or interactions), where behavioural compensation to predation risk occurs, such as animals avoiding areas of high predation risk. Studies of invertebrates, fish and amphibians have shown that non‐lethal effects may be larger than lethal effects in determining the behaviour, condition, density and distribution of animals over a range of trophic levels. Although non‐lethal effects have been well described in the behavioural ecology of birds (and also mammals) within the context of anti‐predation behaviour, their role relative to lethal effects is probably underestimated. Birds show many behavioural and physiological changes to reduce direct mortality from predation and these are likely to have negative effects on other aspects of their fitness and population dynamics, as well as affecting the ecology of their own prey and their predators. As a consequence, the effects of predation in birds are best measured by trade‐offs between maximizing instantaneous survival in the presence of predators and acquiring or maintaining resources for long‐term survival or reproduction. Because avoiding predation imposes foraging costs, and foraging behaviour is relatively easy to measure in birds, the foraging–predation risk trade‐off is probably an effective framework for understanding the importance of non‐lethal effects, and so the population and community effects of predation risk in birds and other animals. Using a trade‐off approach allows us to predict better how changes in predator density will impact on population and community dynamics, and how animals perceive and respond to predation risk, when non‐lethal effects decouple the relationship between predator density and direct mortality rate. The trade‐off approach also allows us to identify where predation risk is structuring communities because of avoidance of predators, even when this results in no observable direct mortality rate.     

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.     

Predation on primates: Ecological patterns and evolutionary consequences

It has long been thought that predation has had important ecological and evolutionary effects on primates as prey. Predation has been theorized to have been a major selective force in the evolution of hominids.¹ In modern primates, behaviors such as active defense, concealment, vigilance, flight, and alarm calls have been attributed to the selective pressures of predation, as has group living itself. It is clear that primates, like other animals, have evolved ways to minimize their risk of predation. However, the extent to which they have been able to do so, given other constraints of living such as their own need to acquire food, has not yet been resolved. Perhaps most hotly debated is whether predation has been the primary selective force favoring the evolution of group living in primates. Part of the difficulty in resolving the debate lies in a paucity of direct evidence of predation. This is regrettable yet understandable since primatologists, by definition, focus on the study of primates, not predators of primates (unless these are also primates). Systematic direct evidence of the effects of predation can best be obtained by studying predators that are as habituated to observers as are their primate prey. Until this is done, we must continue to rely on opportunistic accounts of predation and predation attempts, and on systematically obtained indirect evidence. Such data reveal several interesting patterns: (1) although smaller primates may have greater predation rates than larger primates, even the largest primates are not invulnerable to predation; (2) the use by primates of unfamiliar areas can result in higher predation rates, which might be one pressure favoring philopatry, or site fidelity; (3) arboreal primates are at greater risk of predation when they are more exposed (at forest edges and tops of canopies) than in more concealed locations; (4) predation by mammalian carnivores may often be episodic; and (5) terrestrial primates may not experience greater predation than arboreal primates.     

Non-additivity among multiple cues of predation risk: a behaviorally-driven trophic cascade between owls and songbirds

Non-lethal effects of predators on prey are initiated in the form of responses to direct and indirect cues of predation risk. Like their lethal equivalents, non-lethal effects may affect species further down the food web initiating a behaviorally-driven trophic cascade. I presented a direct cue of predation risk, owl vocalizations, to white-footed mice ( Peromyscus leucopus ) during either a new or full moon (indirect cue). Mice reduced their activity in space by nearly two-thirds in response to playbacks of owl vocalizations during a full moon. However, neither moonlight (full vs new) nor the presence/absence of owl calls had an effect on space use when each cue varied singly. Previous studies have demonstrated a tight correlation between spatial activity in mice as used in the current experiment and nest predation rates on ground-nesting birds. Because moonlight is a ubiquitous deterrent of activity in nocturnal rodents I used of long-term nesting records the veery ( Catharus fuscescens ) to test whether nest predation rates were correlated negatively with moonlight. For half the lunar cycle (∼full moon to new moon) predation rates decreased with moonlight as predicted. During the second half of the lunar cycle predation and moonlight did not correlate as expected, but this was likely due to the depletion of vulnerable nests after a period of in which predation rates were at their maximum near the full moon. These studies suggest that the non-lethal effects of predatory risk on mice (i.e. changes in space use) cascade to affect their prey. Through the mechanism of reduced space use by rodents, perceived predation risk has the potential to significantly and indirectly affect songbird nest predation rates.     

Assessing the risk effects of native predators on the exotic American bullfrog (Lithobates catesbeianus) and their indirect consequences to ecosystem function

Understanding the factors and mechanisms that affect the impacts of invasive species in invaded environments has been widely debated among researchers. However, few studies about invasive species have explored the effects of predation risks by native predators on exotic prey. Herein, the traditional invasive predator-native prey framework was reversed. We tested if tadpoles, of the worldwide invasive American Bullfrog Lithobates catesbeianus, were affected by the predation risk imposed by native predators. We used two different species of belostomatid predators and tested whether and how predation-induced phenotypic plasticity on L. catesbeianus reverberated in morphological, physiological, and ecosystem-level processes. Individuals of L. catesbeianus modified their morphological (tail muscle width), behavioral (activity and foraging), and physiological (growth and growth efficiency) traits in the presence of predation risk. Based on the observed morphological changes, our results suggest that prey may recognize predator-specific cues. In addition, we observed that L. catesbeianus' responses to predation risk can affect ecosystem-level properties, by inducing trophic cascades and reducing animal-mediated nutrient recycling rates. In summary, our study supports that exotic prey species who are subjected to native predators may display anti-predator responses, with implications for their development, as well as possible ecosystem-level effects.     

Too risky to settle: avian community structure changes in response to perceived predation risk on adults and offspring

Predation risk is widely hypothesized as an important force structuring communities, but this potential force is rarely tested experimentally, particularly in terrestrial vertebrate communities. How animals respond to predation risk is generally considered predictable from species life-history and natural-history traits, but rigorous tests of these predictions remain scarce. We report on a large-scale playback experiment with a forest bird community that addresses two questions: (i) does perceived predation risk shape the richness and composition of a breeding bird community? And (ii) can species life-history and natural-history traits predict prey community responses to different types of predation risk? On 9 ha plots, we manipulated cues of three avian predators that preferentially prey on either adult birds or offspring, or both, throughout the breeding season. We found that increased perception of predation risk led to generally negative responses in the abundance, occurrence and/or detection probability of most prey species, which in turn reduced the species richness and shifted the composition of the breeding bird community. Species-level responses were largely predicted from the key natural-history trait of body size, but we did not find support for the life-history theory prediction of the relationship between species'' slow/fast life-history strategy and their response to predation risk.