Cool Headed Individuals Are Better Survivors: Non-Consumptive and Consumptive Effects of a Generalist Predator on a Sap Feeding Insect

Non-consumptive effects (NCEs) of predators are part of the complex interactions among insect natural enemies and prey. NCEs have been shown to significantly affect prey foraging and feeding. Leafhopper''s (Auchenorrhyncha) lengthy phloem feeding bouts may play a role in pathogen transmission in vector species and also exposes them to predation risk. However, NCEs on leafhoppers have been scarcely studied, and we lack basic information about how anti-predator behaviour influences foraging and feeding in these species. Here we report a study on non-consumptive and consumptive predator-prey interactions in a naturally co-occurring spider–leafhopper system. In mesocosm arenas we studied movement patterns during foraging and feeding of the leafhopper Psammotettix alienus in the presence of the spider predator Tibellus oblongus. Leafhoppers delayed feeding and fed much less often when the spider was present. Foraging movement pattern changed under predation risk: movements became more frequent and brief. There was considerable individual variation in foraging movement activity. Those individuals that increased movement activity in the presence of predators exposed themselves to higher predation risk. However, surviving individuals exhibited a ‘cool headed’ reaction to spider presence by moving less than leafhoppers in control trials. No leafhoppers were preyed upon while feeding. We consider delayed feeding as a “paradoxical” antipredator tactic, since it is not necessarily an optimal strategy against a sit-and-wait generalist predator.     

Consumptive and non-consumptive effects of an invasive marine predator on native coral-reef herbivores

Invasive predators typically have larger effects on native prey populations than native predators, yet the potential roles of their consumptive versus non-consumptive effects (CEs vs. NCEs) in structuring invaded systems remains unclear. Invasive lionfish (Pterois volitans) may have ecosystem-level effects by altering native fish grazing on benthic algae that could otherwise displace corals. Lionfish could reduce grazing by decreasing the abundance of herbivorous fishes (CEs), and/or the predation risk posed by lionfish could alter grazing behavior of fishes (NCEs). To test for these CEs, we manipulated lionfish densities on large reefs in The Bahamas and surveyed fish populations throughout June 2009–2011. In July 2011, NCEs of lionfish were measured by observing fish grazing behavior on algal-covered substrata placed in microhabitats varying in lionfish presence at different spatial scales, and quantifying any resulting algal loss. Lionfish reduced small herbivorous fish density by the end of the 2010 summer recruitment season. Grazing by small and large fishes was reduced on high-lionfish-density reefs, and small fish grazing further decreased when in the immediate presence of lionfish within-reefs. Lionfish had a negative indirect effect on algal loss, with 66–80 % less algae removed from substrata in high-lionfish-density reefs. Parrotfishes were likely driving the response of herbivorous fishes to both CEs and NCEs of lionfish. These results demonstrate the importance of considering NCEs in addition to CEs of invasive predators when assessing the effects of invasions.     

Mesopredator suppression by an apex predator alleviates the risk of predation perceived by small prey

Predators can impact their prey via consumptive effects that occur through direct killing, and via non-consumptive effects that arise when the behaviour and phenotypes of prey shift in response to the risk of predation. Although predators'' consumptive effects can have cascading population-level effects on species at lower trophic levels there is less evidence that predators'' non-consumptive effects propagate through ecosystems. Here we provide evidence that suppression of abundance and activity of a mesopredator (the feral cat) by an apex predator (the dingo) has positive effects on both abundance and foraging efficiency of a desert rodent. Then by manipulating predators'' access to food patches we further the idea that apex predators provide small prey with refuge from predation by showing that rodents increased their habitat breadth and use of ‘risky′ food patches where an apex predator was common but mesopredators rare. Our study suggests that apex predators'' suppressive effects on mesopredators extend to alleviate both mesopredators'' consumptive and non-consumptive effects on prey.     

Relative importance of consumptive and non-consumptive effects of predators on prey and plant damage: the influence of herbivore ontogeny

The non-consumptive (or trait-mediated) effects of predators on prey are known to contribute substantially to the negative impact of insect predators on herbivorous insects. Our goal now is to understand what factors alter the relative importance of the consumptive (or density-mediated) and non-consumptive components of the total predator impact. This is important both for understanding the effects of predators in natural systems as well as for successfully manipulating predators for biological control in agriculture. In this study, we tested whether herbivore ontogeny influenced the contribution of consumptive and non-consumptive effects of a predator on herbivore survivorship and plant damage by the herbivores. We addressed these questions using the native plant Solanum ptychanthum Dunal (Solanaceae), the predator Podisus maculiventris Say (Heteroptera: Pentatomidae), and first-, third-, and fourth-instar Manduca sexta L. (Lepidoptera: Sphingidae). In field cage experiments, we found that first- and third-instar M. sexta were more vulnerable to predators compared to fourth instars. In the presence of predators, M. sexta caterpillars spent less time on feeding compared to caterpillars in the absence of predators. The amount of damage the plants received was reduced in the presence of the predator and the consumptive and non-consumptive components contributed approximately equally to this reduction. Thus, the non-consumptive component of the predator is important for all of the herbivore stages vulnerable to predation in our study. We conclude with a discussion of possible implications of considering non-consumptive effects of predators in biological control of agricultural pests.     

Predator–prey naïveté, antipredator behavior,and the ecology of predator invasions

We present a framework for explaining variation in predator invasion success and predator impacts on native prey that integrates information about predator–prey naïveté, predator and prey behavioral responses to each other, consumptive and non‐consumptive effects of predators on prey, and interacting effects of multiple species interactions. We begin with the ‘naïve prey’ hypothesis that posits that naïve, native prey that lack evolutionary history with non‐native predators suffer heavy predation because they exhibit ineffective antipredator responses to novel predators. Not all naïve prey, however, show ineffective antipredator responses to novel predators. To explain variation in prey response to novel predators, we focus on the interaction between prey use of general versus specific cues and responses, and the functional similarity of non‐native and native predators. Effective antipredator responses reduce predation rates (reduce consumptive effects of predators, CEs), but often also carry costs that result in non‐consumptive effects (NCEs) of predators. We contrast expected CEs versus NCEs for non‐native versus native predators, and discuss how differences in the relative magnitudes of CEs and NCEs might influence invasion dynamics. Going beyond the effects of naïve prey, we discuss how the ‘naïve prey’, ‘enemy release’ and ‘evolution of increased competitive ability’ (EICA) hypotheses are inter‐related, and how the importance of all three might be mediated by prey and predator naïveté. These ideas hinge on the notion that non‐native predators enjoy a ‘novelty advantage’ associated with the naïveté of native prey and top predators. However, non‐native predators could instead suffer from a novelty disadvantage because they are also naïve to their new prey and potential predators. We hypothesize that patterns of community similarity and evolution might explain the variation in novelty advantage that can underlie variation in invasion outcomes. Finally, we discuss management implications of our framework, including suggestions for managing invasive predators, predator reintroductions and biological control.     

Ontogenetic differences of herbivory on woody and herbaceous plants: a meta-analysis demonstrating unique effects of herbivory on the young and the old, the slow and the fast

Defensive modifications in prey traits that reduce predation risk can also have negative effects on prey fitness. Such nonconsumptive effects (NCEs) of predators are common, often quite strong, and can even dominate the net effect of predators. We develop an intuitive graphical model to identify and explore the conditions promoting strong NCEs. The model illustrates two conditions necessary and sufficient for large NCEs: (1) trait change has a large cost, and (2) the benefit of reduced predation outweighs the costs, such as reduced growth rate. A corollary condition is that potential predation in the absence of trait change must be large. In fact, the sum total of the consumptive effects (CEs) and NCEs may be any value bounded by the magnitude of the predation rate in the absence of the trait change. The model further illustrates how, depending on the effect of increased trait change on resulting costs and benefits, any combination of strong and weak NCEs and CEs is possible. The model can also be used to examine how changes in environmental factors (e.g., refuge safety) or variation among predator–prey systems (e.g., different benefits of a prey trait change) affect NCEs. Results indicate that simple rules of thumb may not apply; factors that increase the cost of trait change or that increase the degree to which an animal changes a trait, can actually cause smaller (rather than larger) NCEs. We provide examples of how this graphical model can provide important insights for empirical studies from two natural systems. Implementation of this approach will improve our understanding of how and when NCEs are expected to dominate the total effect of predators. Further, application of the models will likely promote a better linkage between experimental and theoretical studies of NCEs, and foster synthesis across systems.     

Predator biomass determines the magnitude of non-consumptive effects (NCEs) in both laboratory and field environments

Predator body size often indicates predation risk, but its significance in non-consumptive effects (NCEs) and predator risk assessment has been largely understudied. Although studies often recognize that predator body size can cause differing cascading effects, few directly examine prey foraging behavior in response to individual predator sizes or investigate how predator size is discerned. These mechanisms are important since perception of the risk imposed by predators dictates behavioral responses to predators and subsequent NCEs. Here, we evaluate the role of predator body size and biomass on risk assessment and the magnitude of NCEs by investigating mud crab foraging behavior and oyster survival in response to differing biomasses of blue crab predators using both laboratory and field methods. Cues from high predator biomass treatments including large blue crab predators and multiple small blue crab predators decreased mud crab foraging and increased oyster survival, whereas mud crab foraging in response to a single small blue crab did not differ from controls. Mud crabs also increased refuge use in the presence of large and multiple small, but not single small, blue crab predators. Thus, both predator biomass and aggregation patterns may affect the expression of NCEs. Understanding the impact of predator biomass may therefore be necessary to successfully predict the role of NCEs in shaping community dynamics. Further, the results of our laboratory experiments were consistent with observed NCEs in the field, suggesting that data from mesocosm environments can provide insight into field situations where flow and turbulence levels are moderate.     

Predator cue and prey density interactively influence indirect effects on Basal resources in intertidal oyster reefs

Predators can influence prey abundance and traits by direct consumption, as well as by non-consumptive effects of visual, olfactory, or tactile cues. The strength of these non-consumptive effects (NCEs) can be influenced by a variety of factors, including predator foraging mode, temporal variation in predator cues, and the density of competing prey. Testing the relative importance of these factors for determining NCEs is critical to our understanding of predator-prey interactions in a variety of settings. We addressed this knowledge gap by conducting two mesocosm experiments in a tri-trophic intertidal oyster reef food web. More specifically, we tested how a predatory fish (hardhead catfish, Ariopsis felis) directly influenced their prey (mud crabs, Panopeus spp.) and indirectly affected basal resources (juvenile oysters, Crassostrea virginica), as well as whether these direct and indirect effects changed across a density gradient of competing prey. Per capita crab foraging rates were inversely influenced by crab density, but they were not affected by water-borne predator cues. As a result, direct consumptive effects on prey foraging rates were stronger than non-consumptive effects. In contrast, predator cue and crab density interactively influenced indirect predator effects on oyster mortality in two experiments, with trait-mediated and density-mediated effects of similar magnitude operating to enhance oyster abundance. Consistent differences between a variable predator cue environment and other predator cue treatments (no cue and constant cue) suggests that an understanding of the natural risk environment experienced by prey is critical to testing and interpreting trait-mediated indirect interactions. Further, the prey response to the risk environment may be highly dependent on prey density, particularly in prey populations with strong intra-specific interactions.     

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.     

Rainfall and hydrological stability alter the impact of top predators on food web structure and function

Climate change will alter the distribution of rainfall, with potential consequences for the hydrological dynamics of aquatic habitats. Hydrological stability can be an important determinant of diversity in temporary aquatic habitats, affecting species persistence and the importance of predation on community dynamics. As such, prey are not only affected by drought‐induced mortality but also the risk of predation [a non‐consumptive effect (NCE)] and actual consumption by predators [a consumptive effect (CE)]. Climate‐induced changes in rainfall may directly, or via altered hydrological stability, affect predator–prey interactions and their cascading effects on the food web, but this has rarely been explored, especially in natural food webs. To address this question, we performed a field experiment using tank bromeliads and their aquatic food web, composed of predatory damselfly larvae, macroinvertebrate prey and bacteria. We manipulated the presence and consumption ability of damselfly larvae under three rainfall scenarios (ambient, few large rainfall events and several small rainfall events), recorded the hydrological dynamics within bromeliads and examined the effects on macroinvertebrate colonization, nutrient cycling and bacterial biomass and turnover. Despite our large perturbations of rainfall, rainfall scenario had no effect on the hydrological dynamics of bromeliads. As a result, macroinvertebrate colonization and nutrient cycling depended on the hydrological stability of bromeliads, with no direct effect of rainfall or predation. In contrast, rainfall scenario determined the direction of the indirect effects of predators on bacteria, driven by both predator CEs and NCEs. These results suggest that rainfall and the hydrological stability of bromeliads had indirect effects on the food web through changes in the CEs and NCEs of predators. We suggest that future studies should consider the importance of the variability in hydrological dynamics among habitats as well as the biological mechanisms underlying the ecological responses to climate change.     

Fear in the dark? Community‐level effects of non‐lethal predators change with light regime

The total effect of predators on prey is a combination of direct consumption, and non‐consumptive effects (NCEs), such as predator‐induced changes to prey morphology, behaviour and life history. Past research into NCEs has tended to focus on pair‐wise interactions between predators and prey, while in natural ecosystems, species exist in complex communities with several trophic levels made up of multiple autotrophic and heterotropic species. To address how predator NCEs alter the photosynthetic and heterotrophic components of communities, we exposed microbial microcosms to one of three predator treatments: live predators (full predator effect), freeze‐killed predators (NCEs only) or no predators (control), and incubated them under either 12 h:12 h light:dark conditions or continual darkness. Under 12 h:12 h light:dark conditions, NCEs‐only communities never differed from predator‐free communities, but differed from live predator communities. Under conditions of continual darkness, the structure of NCEs‐only communities differed from predator‐free controls, but not from live predator communities, suggesting NCEs can be strong enough to structure communities. Predation threat may cause certain prey to induce defences, such as reductions in movement, which make them less competitive in a community setting. This reduction in competitive ability could lead to these species being driven to extinction through interspecific competition, resulting in similar communities to those in which live predators are present. Heterotrophic species whose rates of resource acquisition depend on movement rates may be affected to a greater extent than autotrophs by predator‐induced reductions in movement, accounting for our observed differences in predator NCEs in ‘dark’ and ‘light’ communities. Our results suggest that the community‐level consequences of fear are greater in the dark. Synthesis Predators affect prey through consumptive and non‐consumptive effects (NCEs) such as alterations to prey behaviour, morphology, and life history. However, predators and prey do not exist in isolated pairs, but in complex communities where they interact with many other species. Using a long term study (>10 predator generations), we show that predator NCEs alone can alter community structure under conditions of darkness, but not in a 12h:12h light:dark cycle. Our results demonstrate for the first time that although the community‐level consequences of predator NCEs may be dramatic, they depend upon the abiotic conditions of the ecosystem.     

The context dependence of non‐consumptive predator effects

Non‐consumptive predator effects (NCEs) are now widely recognised for their capacity to shape ecosystem structure and function. Yet, forecasting the propagation of these predator‐induced trait changes through particular communities remains a challenge. Accordingly, focusing on plasticity in prey anti‐predator behaviours, we conceptualise the multi‐stage process by which predators trigger direct and indirect NCEs, review and distil potential drivers of contingencies into three key categories (properties of the prey, predator and setting), and then provide a general framework for predicting both the nature and strength of direct NCEs. Our review underscores the myriad factors that can generate NCE contingencies while guiding how research might better anticipate and account for them. Moreover, our synthesis highlights the value of mapping both habitat domains and prey‐specific patterns of evasion success (‘evasion landscapes’) as the basis for predicting how direct NCEs are likely to manifest in any particular community. Looking ahead, we highlight two key knowledge gaps that continue to impede a comprehensive understanding of non‐consumptive predator–prey interactions and their ecosystem consequences; namely, insufficient empirical exploration of (1) context‐dependent indirect NCEs and (2) the ways in which direct and indirect NCEs are shaped interactively by multiple drivers of context dependence.     

The effect of predation pressure and predator adaptive foraging on the relative importance of consumptive and non‐consumptive predator net effects in a freshwater model system

An important challenge in community ecology is identifying the functional characteristics capable of predicting the nature and strength of predator effects on food webs. We developed an individual‐based model, based on a shallow lake model system, to evaluate the total, consumptive, and non‐consumptive indirect effect that predators have on basal resources when the predators differ in their foraging types (active adaptive foraging or sedentary foraging). Overall, both predator types caused similar total indirect effects on lower trophic levels. However, the nature net effects of predators diverged between predator foraging types. Active predators caused larger non‐consumptive effects, relative to the total indirect effect, irrespective of predation pressure levels. On the other hand, sedentary predators caused larger non‐consumptive effects for lower predation pressure levels, but consumptive effects became more important as predation pressure increased. Our simulations showed that the reliance on a particular mechanism driving consumer–resource interactions is altered by predator foraging behavior and highlight the importance of both prey and predator foraging behaviors to predict the causes and consequences of cascading effects observed in food webs.     

Linking the green and brown worlds through nonconsumptive predator effects

Predators can indirectly affect lower trophic levels by either consuming their prey (consumptive effect, CE) or by changing the physiology or behavior of their prey (nonconsumptive effect, NCE). Cascading effects of predators on primary producers are common, and can be propagated by CEs, NCEs, or a combination of both mechanisms. Predator impacts in detrital food webs (the ‘brown world’) have received considerably less attention than their effects on systems with primary producers at the base (the ‘green world’), and only recently have we begun to appreciate the importance of above‐ground predators indirectly impacting below‐ground processes. Numerous studies reveal the total impact (CEs and NCEs) of predators in brown food webs, but our understanding of the role of isolated NCEs is limited. Many habitats and major taxa have not been studied, and patterns are difficult to distinguish due to frequent reporting of mixed effects. Predators play an important role as connectors between brown and green worlds when they feed from both food webs (multichannel feeding). We are only beginning to understand how NCEs influence detrital food webs, and it is unknown whether multichannel fear is an essential component of predator–prey ecology that regulates ecosystem function. Synthesis Predators have been shown to impact ecosystems through both consumptive and nonconsumptive effects on their prey Historically, herbivory‐based ‘green’ systems have been the venue for documenting these predator effects, while detritus‐based ‘brown’ systems received considerably less attention. However, similar mechanisms exist in green and brown worlds, suggesting strong parallels. We review and synthesize predator effects in detrital systems, highlighting important shortcomings in current understanding. Furthermore, we build upon the idea of multichannel feeding (i.e. consumption of prey from both green and brown food webs) to propose the existence of ‘multichannel fear’. We provide a framework for documenting multichannel fear to facilitate continued exploration of how predators link seemingly disparate systems.     

Remembering the good and the bad: memory-based mediation of the food–safety trade-off in dynamic landscapes

Predator–prey interactions are central to fitness as animals simultaneously avoid death and consume resources to ensure growth and reproduction. Along with direct effects, predators can also exert strong non-consumptive effects. For example, prey shift habitat use in the presence of predators, a potentially learned behavior. The impact of cognition on movement and predator interactions is largely unexplored despite evidence of learned responses to predation threat. We explore how learning and spatial memory influence predator–prey dynamics by introducing predators into a memory-driven movement modeling framework. To model various aspects of risk, we vary predator behavior: their persistence and spatial correlation with the prey’s resources. Memory outperforms simpler movement processes most in patchy environments with more predictable predators that are more easily avoided once learned. In these cases, memory aids foragers in managing the food–safety trade-off. For example, particular parameterizations of the predation memory reduce encounters while maintaining consumption. We found that non-consumptive effects are highest in landscapes of concentrated, patchy resources. These effects are intensified when predators are highly correlated with the forager’s resources. Smooth landscapes provide more opportunities for foragers to simultaneously consume resources and avoid predators. Predators are able to effectively guard all resources in very patchy landscapes. These non-consumptive effects are also seen with the shift away from the best quality habitat compared to foraging in a predator-free environment.     

Litter identity mediates predator impacts on the functioning of an aquatic detritus-based food web

During past decades, several mechanisms such as resource quality and habitat complexity have been proposed to explain variations in the strength of trophic cascades across ecosystems. In detritus-based headwater streams, litter accumulations constitute both a habitat and a resource for detritivorous macroinvertebrates. Because litter edibility (which promotes trophic cascades) is usually inversely correlated with its structural complexity (which weakens trophic cascades), there is a great scope for stronger trophic cascades in litter accumulations that are dominated by easily degradable litter species. However, it remains unclear how mixing contrasting litter species (conferring both habitat complexity and high quality resource) may influence top–down controls on communities and processes. In enclosures exposed in a second-order stream, we manipulated litter species composition by using two contrasting litter (alder and oak), and the presence–absence of a macroinvertebrate predator (Cordulegaster boltonii larvae), enabling it to effectively exert predation pressure, or not, on detritivores (consumptive versus non-consumptive predation effects). Leaf mass loss, detritivore biomass and community structure were mostly controlled independently by litter identity and mixing and by predator consumption. However, the strength of predator control was mediated by litter quality (stronger on alder), and to a lesser extent by litter mixing (weaker on mixed litter). Refractory litter such as oak leaves may contribute to the structural complexity of the habitat for stream macroinvertebrates, allowing the maintenance of detritivore communities even when strong predation pressure occurs. We suggest that considering the interaction between top–down and bottom–up factors is important when investigating their influence on natural communities and ecosystem processes in detritus-based ecosystems.     

Prey perception of predation risk: volatile chemical cues mediate non-consumptive effects of a predator on a herbivorous insect

Predators can affect prey in two ways—by reducing their density (consumptive effects) or by changing their behavior, physiology or other phenotypic traits (non-consumptive effects). Understanding the cues and sensory modalities prey use to detect predators is critical for predicting the strength of non-consumptive effects and the outcome of predator–prey encounters. While predator-associated cues have been well studied in aquatic systems, less is known about how terrestrial prey, particularly insect larvae, detect their predators. We evaluated how Colorado potato beetle, Leptinotarsa decemlineata, larvae perceive predation risk by isolating cues from its stink bug predator, the spined soldier bug, Podisus maculiventris. When exposed to male “risk” predators that were surgically manipulated so they could hunt but not kill, beetles reduced feeding 29 % compared to controls. Exposure to risk females caused an intermediate response. Beetles ate 24 % less on leaves pre-exposed to predators compared to leaves never exposed to predators, indicating that tactile and visual cues are not required for the prey’s response. Volatile odor cues from predators reduced beetle feeding by 10 % overall, although male predators caused a stronger reduction than females. Finally, visual cues from the predator had a weak effect on beetle feeding. Because multiple cues appear to be involved in prey perception of risk, and because male and female predators have differential effects, beetle larvae likely experience tremendous variation in the information about risk from their local environment.     

Resource dynamics influence the strength of non-consumptive predator effects on prey

Predators influence prey populations both by consuming individual prey, and by inducing changes in prey behaviour that limit reproduction and survival. Because prey trade-off predation risk for forageing gains, the magnitude of predators' non-consumptive effects should depend on resource availability. Studies of non-consumptive effects generally adopt either of two strategies: (i) maintaining a static ration of the prey's resources; and (ii) using resource populations that vary dynamically in response to prey behaviour. Contrasting these experimental designs using meta-analysis, we evaluated whether resource dynamics influence the magnitude of non-consumptive effects on prey growth, survival, fecundity, population density, forageing rate and habitat use. Predators had a more negative effect on prey demography in dynamic- vs. static-resource experiments. Our results highlight the importance of resource dynamics in mediating the magnitude of non-consumptive effects of predators on prey, and illustrate the often-unintended impacts of experimental design on estimates of effect size in ecological interactions.     

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.     

Within-Plant Bottom-Up Effects Mediate Non-Consumptive Impacts of Top-Down Control of Soybean Aphids

There is increasing evidence that top-down controls have strong non-consumptive effects on herbivore populations. However, little is known about how these non-consumptive effects relate to bottom-up influences. Using a series of field trials, we tested how changes in top-down and bottom-up controls at the within-plant scale interact to increase herbivore suppression. In the first experiment, we manipulated access of natural populations of predators (primarily lady beetles) to controlled numbers of A. glycines on upper (i.e. vigorous-growing) versus lower (i.e. slow-growing) soybean nodes and under contrasting plant ages. In a second experiment, we measured aphid dispersion in response to predation. Bottom-up and top-down controls had additive effects on A. glycines population growth. Plant age and within-plant quality had significant bottom-up effects on aphid size and population growth. However, top-down control was the dominant force suppressing aphid population growth, and completely counteracted bottom-up effects at the plant and within-plant scales. The intensity of predation was higher on upper than lower soybean nodes, and resulted in a non-consumptive reduction in aphid population growth because most of the surviving aphids were located on lower plant nodes, where rates of increase were reduced. No effects of predation on aphid dispersal among plants were detected, suggesting an absence of predator avoidance behavior by A. glycines. Our results revealed significant non-consumptive predator impacts on aphids due to the asymmetric intensity of predation at the within-plant scale, suggesting that low numbers of predators are highly effective at suppressing aphid populations.