Interspecific differences in antipredator strategies determine the strength of non‐consumptive predator effects on stream detritivores

Animal species differ considerably in their response to predation risks. Interspecific variability in prey behaviour and morphology can alter cascading effects of predators on ecosystem structure and functioning. We tested whether species‐specific morphological defenses may affect responses of leaf litter consuming invertebrate prey to sit‐and‐wait predators, the odonate Cordulegaster boltonii larvae, in aquatic food webs. Partly or completely blocking the predator mouthparts (mandibles and/or extensible labium), thus eliminating consumptive (i.e. lethal) predator effects, we created a gradient of predator‐prey interaction intensities (no predator < predator – no attack < predator – non‐lethal attacks < lethal predator). A field experiment was first used to assess both consumptive and non‐consumptive predator effects on leaf litter decomposition and prey abundances. Laboratory microcosms were then used to examine behavioural responses of armored and non‐armored prey to predation risk and their consequences on litter decomposition. Results show that armored and non‐armored prey responded to both acute (predator – non‐lethal attacks) and chronic (predator – no attack) predation risks. Acute predation risk had stronger effects on litter decomposition, prey feeding rate and prey habitat use than predator presence alone (chronic predation risk). Predator presence induced a reduction in feeding activity (i.e. resource consumption) of both prey types but a shift to predator‐free habitat patches in non‐armored detritivores only. Non‐consumptive predator effects on prey subsequently decreased litter decomposition rate. Species‐specific prey morphological defenses and behaviour should thus be considered when studying non‐consumptive predator effects on prey community structure and ecosystem functioning.     

Indirect interactions in a rice ecosystem: density dependence and the interplay between consumptive and non‐consumptive effects of predators

1. Density‐ and trait‐mediated indirect interactions (DMIIs and TMIIs, respectively) in food chains play crucial roles in community structure and processes. However, factors affecting the relative strength of these interactions are poorly understood, including in widespread and important freshwater rice ecosystems. 2. We studied the strength of DMIIs and TMIIs in a food chain involving a predator (the Reeve’s turtle Chinemys reevesii), its herbivorous prey (the apple snail Pomacea canaliculata) and a plant (rice Oryza sativa) in outdoor containers simulating rice fields. We also evaluated consumptive and non‐consumptive effects of the predator on the snail. We removed a fixed proportion of snails every 2 days to simulate prey consumption and introduced a caged turtle that was fed daily with snails to simulate non‐consumptive effects. 3. Direct consumptive effects increased growth of the remaining snails and their per capita feeding rate. Moreover, consumptive and non‐consumptive effects, and their interaction, affected the proportion of snails buried in the soil. This interaction was presumably because increasing food availability per snail induced their self‐burying behaviour. 4. Both DMIIs and TMIIs affected the number of rice plants remaining, whereas their interaction term was not significant. 5. In summary, density dependence and interactions between consumptive and non‐consumptive effects influenced snail growth and behaviour, respectively. However, no cascading effects of these complicated interactions on rice plants were detected.     

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.     

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.     

Non‐consumptive effects of a natural enemy on a non‐prey herbivore population

1. Predator–prey interactions have traditionally focused on the consumptive effects that predators have on prey. However, predators can also reduce the abundance of prey through behaviourally‐mediated non‐consumptive effects. For example, pea aphids (Acyrthosiphon pisum Harris) drop from their host plants in response to the risk of attack, reducing population sizes as a consequence of lost feeding opportunities. 2. The objective of the present study was to determine whether the non‐consumptive effects of predators could extend to non‐prey herbivore populations as a result of non‐lethal incidental interactions between herbivores and foraging natural enemies. 3. Polyculture habitats consisting of green peach aphids (Myzus persicae Sulzer) feeding on collards and pea aphids feeding on fava beans were established in greenhouse cages. Aphidius colemani Viereck, a generalist parasitoid that attacks green peach aphids but not pea aphids, was released into half of the cages and the abundance of the non‐host pea aphid was assessed. 4. Parasitoids reduced the population growth of the non‐host pea aphid by increasing the frequency of defensive drops; but this effect was dependent on the presence of green peach aphids. 5. Parasitoids probably elicited the pea aphid dropping behaviour through physical contact with pea aphids while foraging for green peach aphids. It is unlikely that pea aphids were responding to volatile alarm chemicals emitted by green peach aphids in the presence of the parasitoid. 6. In conclusion, the escape response of the pea aphid provided the opportunity for a parasitoid to have non‐target effects on an herbivore with which it did not engage in a trophic interaction. The implication is that natural enemies with narrow diet breadths have the potential to influence the abundance of a broad range of prey and non‐prey species via non‐consumptive effects.     

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.     

Effects of predation risk and plant resistance on Manduca sexta caterpillar feeding behaviour and physiology

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Predator presence dramatically reduces copepod abundance through condition‐mediated non‐consumptive effects

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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.     

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.     

Independent and combined effects of multiple predators across ontogeny of a dominant grazer

Ecosystems host multiple coexisting predator species whose interactions may strengthen or weaken top–down control of grazers. Grazer populations often exhibit size‐structure, but the nature of multiple predator effects on suppression of size‐structured prey has seldom been explicitly considered. In a southeastern US salt‐marsh, we used both field (additive design) and mesocosm (additive‐substitutive design) experiments to test the independent and combined effects of two species of predatory crab on the survival and predator‐avoidance behavior (i.e. a non‐consumptive effect) of both juveniles and adults of a dominant grazing snail. Results showed: 1) juvenile snails were more vulnerable to predation; 2) consumptive impacts of predators were hierarchically nested, i.e. the larger predator consumed both juvenile and adult snails, while the smaller‐bodied predator consumed only juvenile snails; 3) there were no emergent multiple predator effects on snail consumption; and 4) non‐consumptive effects differed from consumptive effects, with only the large predator inducing predator‐avoidance behavior of individuals within either snail ontogenetic class. The smaller predator therefore played a functionally redundant trophic role across the prey classes considered, augmenting and potentially stabilizing trophic regulation of juvenile snails. Meanwhile, the larger predator played a complementary and functionally unique role by both expanding the size‐spectrum of prey trophic regulation and non‐consumptively altering prey behavior. While our study suggests that nestedness of consumptive interactions determined by predator and prey body sizes may allow prediction of the functional redundancy of particular predator species, it also shows that traits beyond predator body size (e.g. habitat domain) may be required to predict potentially cascading non‐consumptive effects. Future studies of multiple predators (and predator biodiversity) should continue to strive towards greater realism by incorporating not only size‐structured prey, but also other aspects of resource and environmental heterogeneity typical of natural ecosystems.     

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.     

Non‐consumptive effects of a top‐predator decrease the strength of the trophic cascade in a four‐level terrestrial food web

The fear of predators can strongly impact food web dynamics and ecosystem functioning through effects on herbivores morphology, physiology or behaviour. While non‐consumptive predator effects have been mostly studied in three‐level food chains, we lack evidence for the propagation of non‐consumptive indirect effects of apex predators in four level food‐webs, notably in terrestrial ecosystems. In experimental mesocosms, we manipulated a four‐level food chain including top‐predator cues (snakes), mesopredators (lizards), herbivores (crickets), and primary producers (plants). The strength of the trophic cascade induced by mesopredators through the consumption of herbivores decreased in the presence of top‐predator cues. Specifically, primary production was higher in mesocosms where mesopredators were present relative to mesocosms with herbivores only, and this difference was reduced in presence of top‐predator cues, probably through a trait‐mediated effect on lizard foraging. Our study demonstrates that non‐consumptive effects of predation risk can cascade down to affect both herbivores and plants in a four‐level terrestrial food chain and emphasises the need to quantify the importance of such indirect effects in natural communities.     

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.     

Match and mismatch: Integrating consumptive effects of predators,prey traits,and habitat selection in colonizing aquatic insects

Predators are a particularly critical component of habitat quality, as they affect survival, morphology, behavior, population size, and community structure through both consumptive and non‐consumptive effects. Non‐consumptive effects can often exceed consumptive effects, but their relative importance is undetermined in many systems. Our objective was to determine the consumptive and non‐consumptive effects of a predaceous aquatic insect, Notonecta irrorata, on colonizing aquatic beetles. We tested how N. irrorata affected survival and habitat selection of colonizing aquatic beetles, how beetle traits contributed to their vulnerability to predation by N. irrorata, and how combined consumptive and non‐consumptive effects affected populations and community structure. Predation vulnerabilities ranged from 0% to 95% mortality, with size, swimming, and exoskeleton traits generating species‐specific vulnerabilities. Habitat selection ranged from predator avoidance to preferentially colonizing predator patches. Attraction of Dytiscidae to N. irrorata may be a natural ecological trap given similar cues produced by these taxa. Hence, species‐specific habitat selection by prey can be either predator‐avoidance responses that reduce consumptive effects, or responses that magnify predator effects. Notonecta irrorata had both strong consumptive and non‐consumptive effects on populations and communities, while combined effects predicted even more distinct communities and populations across patches with or without predators. Our results illustrate that an aquatic invertebrate predator can have functionally unique consumptive effects on prey, attracting and repelling prey, while prey have functionally unique responses to predators. Determining species‐specific consumptive and non‐consumptive effects is important to understand patterns of species diversity across landscapes.     

Multi-predator effects across life-history stages: non-additivity of egg- and larval-stage predation in an African treefrog

The effects of multiple predators on their prey are frequently non‐additive because of interactions among predators. When prey shift habitats through ontogeny, many of their predators cannot interact directly. However, predators that occur in different habitats or feed on different prey stages may still interact through indirect effects mediated by prey traits and density. We conducted an experiment to evaluate the combined effects of arboreal egg‐stage and aquatic larval‐stage predators of the African treefrog, Hyperolius spinigularis. Egg and larval predator effects were non‐additive – more Hyperolius survived both predators than predicted from their independent effects. Egg‐stage predator effects on aquatic larval density and size and age at hatching reduced the effectiveness of larval‐stage predators by 70%. Our results indicate that density‐ and trait‐mediated indirect interactions can act across life‐stages and habitats, resulting in non‐additive multi‐predator effects.     

Effects of prey availability, facultative plant feeding, and plant defenses on a generalist insect predator

This study examined the effects of feeding interval, access to host plants (thus, a source of sap), and plant defenses on the predatory insect, Podisus maculiventris Say (Hemiptera: Pentatomidae). The experiment consisted of a 2 × 2 design with two feeding intervals (1 day or 5 days) and predators living on either tomato plants or plastic plants. Females fed every day had greater body weights and egg hatch rates than females fed every five days. Females on tomato plants lived longer than females on plastic plants. However, access to plants did not alleviate the effects of low prey level on predator weight or reproductive output. In a second experiment, third instar nymphs were placed on either tomato plants or plastic plants for four days to examine the effects of tomato trichome defenses on these predators. Nymphs on tomato plants experienced 50% mortality compared to 15% mortality for nymphs on plastic plants. Some nymphs living on tomato plants were trapped by the hairy trichomes of the plant; others had gummed up legs from the exudates of the plants’ glandular trichomes, which inhibited their movement and ability to feed on prey. Although predators appeared to benefit from feeding on tomato plants, their ability to live on the plants was negatively affected by the defensive features of the plants. The potential effects of trichome defenses on predator survival and population dynamics must be considered when evaluating the benefits of plants on insect predator life histories and efficacy as biological control agents.     

Encounters between aphids and their predators: the relative frequencies of disturbance and consumption

Ecologists may wish to evaluate the potential for predators to suppress prey populations through the costs of induced defensive behaviors as well as through consumption. In this paper, we measure the ratio of non‐consumptive, defense‐inducing encounters relative to consumptive encounters (henceforth the ‘disturbed : consumed ratio’) for two species of aphids and propose that these disturbed : consumed ratios can help evaluate the potential for behaviorally mediated prey suppression. For the pea aphid, Acyrthosiphon pisum (Harris) (Homoptera: Aphididae), the ratio of induced disturbances to consumption events was high, 30 : 1. For the cotton aphid, Aphis gossypii (Glover) (Homoptera: Aphididae), the ratio of induced disturbances to consumption events was low, approximately 1 : 14. These results indicate that the potential for predators to suppress pea aphid populations through induction of defensive behaviors is high, whereas the potential for predators to suppress cotton aphid populations through induced behaviors is low. In measuring the disturbed : consumed ratios of two prey species, this paper makes two novel points: it highlights the variability of the disturbed : consumed ratio, and it offers a simple statistic to help ecologists draw connections between predator–prey behaviors and predator–prey population dynamics.     

Does a top predator reduce the predatory impact of an invasive mesopredator on an endangered rodent?

The mesopredator release hypothesis (MRH) predicts that reduced abundance of top‐order predators results in an increase in the abundance of smaller predators (mesopredators) due to a reduction in intra‐guild predation and competition. The irruption of mesopredators that follows the removal of top‐order predators can have detrimental impacts on the prey of the mesopredators. Here we investigated the mechanisms via which the presence of a top‐order predator can benefit prey species. We tested predictions made according to the MRH and foraging theory by contrasting the abundances of an invasive mesopredator (red fox Vulpes vulpes) and an endangered prey species (dusky hopping mouse Notomys fuscus), predator diets, and N. fuscus foraging behaviour in the presence and absence of a top‐predator (dingo Canis lupus dingo). As predicted by the MRH, foxes were more abundant where dingoes were absent. Dietary overlap between sympatric dingoes and foxes was extensive, and fox was recorded in 1 dingo scat possibly indicating intra‐guild predation. Notomys fuscus were more likely to occur in fox scats than dingo scats and as predicted by the MRH N. fuscus were less abundant in the absence of dingoes. The population increase of N. fuscus following rainfall was dampened in the absence of dingoes suggesting that mesopredator release can attenuate bottom‐up effects, although it remains conceivable that differences in grazing regimes associated with dingo exclusion could have also influenced N. fuscus abundance. Notomys fuscus exhibited lower giving‐up densities in the presence of dingoes, consistent with the prediction that their perceived risk of predation would be lower and foraging efficiency greater in the presence of a top‐predator. Our results suggest that mesopredator suppression by a top predator can create a safer environment for prey species where the frequency of fatal encounters between predators and prey is reduced and the non‐consumptive effects of predators are lower.     

Constitutive exposure to the volatile methyl salicylate reduces per‐capita foraging efficiency of a generalist predator to learned prey associations

Understanding the factors that influence the ability of predators to find and kill herbivores is central to enhancing their impact on prey populations, but few studies have tested the impact of these factors on predation rates in realistic foraging environments. Using the tri‐trophic system consisting of tomato, Solanum lycopersicum L. (Solanaceae), hornworm caterpillars, Manduca sexta L. (Lepidoptera: Sphingidae), and the predaceous stink bug Podisus maculiventris (Say) (Hemiptera: Pentatomidae), we measured the effects of associative learning and plant volatile camouflage on predator behavior and foraging efficiency in field enclosures. To do so, we compared experienced vs. naive individuals under varying environmental contexts. Experienced predators were those with prior exposure to induced volatiles from the tomato–caterpillar association, whereas naive predators had not experienced tomato, only prey (caterpillars). We varied their environmental foraging matrix using either (1) tomato surrounded by basil (Ocimum basilicum L.; Lamiaceae) or (2) tomato exposed to the synthetic volatile, methyl salicylate (MeSA). We found that (1) experienced predators were more efficient than naive predators, capturing 28% more prey; (2) the tomato–basil combination did not affect predator–prey interactions; and (3) constitutive emission of synthetic MeSA caused a 22% reduction in P. maculiventris predation rate. These differences corresponded with distinct shifts in predator foraging; for example, experienced individuals were less stationary and exhibited unique behaviors such as stylet extension. Taken together, these results suggest that it is possible to improve the function of generalist predators in suppressing prey by coupling odors with food. However, constitutive emission of volatiles to attract natural enemies may ultimately camouflage neighboring plants, reducing the benefits of orientation to learned stimuli such as induced volatiles.