Reduced Prey Density Research Articles

A healthy but depleted herd: Predators decrease prey disease and density.

The healthy herds hypothesis proposes that predators can reduce parasite prevalence and thereby increase the density of their prey. However, evidence for such predator-driven reductions in the prevalence of prey remains mixed. Furthermore, even less evidence supports increases in prey density during epidemics. Here, we used a planktonic predator-prey-parasite system to experimentally test the healthy herds hypothesis. We manipulated density of a predator (the phantom midge, Chaoborus punctipennis) and parasitism (the virulent fungus Metschnikowia bicuspidata) in experimental assemblages. Because we know natural populations of the prey (Daphnia dentifera) vary in susceptibility to both predator and parasite, we stocked experimental populations with nine genotypes spanning a broad range of susceptibility to both enemies. Predation significantly reduced infection prevalence, eliminating infection at the highest predation level. However, lower parasitism did not increase densities of prey; instead, prey density decreased substantially at the highest predation levels (a major density cost of healthy herds predation). This density result was predicted by a model parameterized for this system. The model specifies three conditions for predation to increase prey density during epidemics: (i) predators selectively feed on infected prey, (ii) consumed infected prey release fewer infectious propagules than unconsumed prey, and (iii) sufficiently low infection prevalence. While the system satisfied the first two conditions, prevalence remained too high to see an increase in prey density with predation. Low prey densities caused by high predation drove increases in algal resources of the prey, fueling greater reproduction, indicating that consumer-resource interactions can complicate predator-prey-parasite dynamics. Overall, in our experiment, predation reduced the prevalence of a virulent parasite but, at the highest levels, also reduced prey density. Hence, while healthy herds predation is possible under some conditions, our empirical results make it clear that the manipulation of predators to reduce parasite prevalence may harm prey density.

Local and large-scale spatial variation in a marine predator-prey interaction in the southwestern Atlantic.

Predator-prey interactions are a key ecological process which can be modified by environmental conditions over a range of spatial scales. Through two complementary short-term experiments, we assessed how local and large-scale environmental conditions affect a subtropical intertidal predator-prey interaction. At a local scale, we evaluated the effects of the degree of exposure to wave action and prey density on consumption rate and interaction strength using a whelk-barnacle system. Consumption rate decreased with wave exposure at experimentally reduced prey density but did not change at ambient density. Such an interactive effect occurred due to shifts in the whelk's feeding behaviour, likely linked to encounter rate and stress amelioration underpinned by prey density. Per capita interaction strength of the whelk on the barnacle weakened along the wave exposure gradient, but to a greater degree at reduced compared to ambient prey density. This confirms that environmental harshness can decrease the importance of predators, but the magnitude of change may be modified by density-dependent effects. A large-scale experiment did not reveal spatial patterns in the whelk-barnacle interaction, nor relationships to chlorophyll-a concentration or the minor change in sea temperature across the study area. Patterns in the size of consumed barnacles along the chlorophyll-a gradient suggest changes in food choice related to prey quality and size. We conclude that disentangling the effects of wave exposure and prey density revealed important potential mechanisms driving species locally. Large-scale variation in the whelk-barnacle interaction appeared to be linked to species' traits shaped by the environmental context.

Population responses of omnivorous arthropods to plant alternative resources suppress prey populations: A meta-analysis.

Omnivory is ubiquitous in ecological communities. Yet, we lack a consensus of how plant alternative resources impact the ability of omnivores to suppress prey populations. Previous work suggests that plant alternative resources can increase, decrease, or have no effect on the magnitude of omnivore-prey interactions. This discrepancy may arise from (1) the ability of omnivore populations to respond to plant alternative resources and (2) identity-specific effects of plant alternative resources. We used a meta-analysis to examine how omnivore population responses and the identity of plant alternative resources affect (1) omnivore predation rates (mainly reported as per capita predation rate) and (2) omnivore impacts on prey population density. Plant alternative resources reduced omnivore predation rate regardless of identity. The suppression of the predation rate by flowers and flowering plants was magnified when pollen alone was tested as the alternative resource. Surprisingly, plant alternative resource availability reduced prey density, suggesting that omnivore predation increased with plant alternative resources. This discrepancy (plant alternative resources not only decreased omnivore predation rates but also decreased prey density) resulted from experimental differences in the ability of omnivore populations to respond to plant alternative resources. In the presence of plant alternative resources, allowing omnivore population responses decreased prey density, while not allowing population responses increased prey density. Because omnivores commonly suppress prey density in the presence of plant alternative resources when population responses of omnivores are allowed, the effectiveness of biological control may depend upon the availability of such resources and the facilitation of population responses.

Direct evidence of a prey depletion “halo” surrounding a pelagic predator colony

Colonially breeding birds and mammals form some of the largest gatherings of apex predators in the natural world and have provided model systems for studying mechanisms of population regulation in animals. According to one influential hypothesis, intense competition for food among large numbers of spatially constrained foragers should result in a zone of prey depletion surrounding such colonies, ultimately limiting their size. However, while indirect and theoretical support for this phenomenon, known as "Ashmole's halo," has steadily accumulated, direct evidence remains exceptionally scarce. Using a combination of vessel-based surveys and Global Positioning System tracking, we show that pelagic seabirds breeding at the tropical island that first inspired Ashmole's hypothesis do indeed deplete their primary prey species (flying fish; Exocoetidae spp.) over a considerable area, with reduced prey density detectable >150 km from the colony. The observed prey gradient was mirrored by an opposing trend in seabird foraging effort, could not be explained by confounding environmental variability, and can be approximated using a mechanistic consumption-dispersion model, incorporating realistic rates of seabird predation and random prey dispersal. Our results provide a rare view of the resource footprint of a pelagic seabird colony and reveal how aggregations of these central-place foraging, marine top predators profoundly influence the oceans that surround them.

Role of fear in predator–prey system with intraspecific competition

Predator–prey interaction is considered as a natural phenomenon in ecological system. An obvious question arises in ecosystem is that how predator affect the density of prey population. In most of the ecological models, predators reduce prey densities by direct killing which is a minor part in population dynamics. Experimental studies on vertebrate has shown that fear of predators can influence the behavioural pattern of prey population and reduce their reproduction. Mathematical models of predator–prey system with variety of predator functional responses with fear effect are studied recently. These studies ignore the fear effect on the death rate of prey species. Based on the above observation, we propose a mathematical model of predator–prey system with the cost of fear that affect reproduction and death rate of prey population. We also include a term modelling intraspecific competition within the predator population. This additional term limits the predator population growth for large predator to prey density ratios. Basic results on the positivity and boundedness of the solutions, feasibility of equilibria, local and global stability and uniform persistence (ensuring the long term survival of the species) and limit cycles and its nature are investigated. Numerical experiments are performed to test the theoretical results obtained for this model.

Study of fear mechanism in predator-prey system in the presence of competitor for the prey

Predator-prey relationships are one of the most commonly observed phenomena in ecosystem. In most of the ecological models, predators reduce prey densities by direct killing. A field study on vertebrate has shown that fear of predators can influence the behavioural pattern of prey population and reduce their reproduction. Most of the recent studies usually consider simplified systems in which predators only attack a single prey species. These studies ignore the impact of diversity of non-prey species. Based on the above observation, we propose a mathematical model of predator-prey system with cost of fear where prey population has a competitor (non-prey). Basic results on the positivity and boundedness of the solutions, feasibility of equilibria, local and global stability and uniform persistence (ensuring the long term survival of the species) and limit cycles and its nature are investigated. We also investigate the impact of fear effect on the model. We observe that increase amount of fear reduce the predator density at the coexistence equilibrium point. Numerical experiments are performed to test the theoretical results obtained for this model.

Consumptive effects and mismatch in predator-prey turnover rates cause inversion of biomass pyramids.

The mismatch between the turnover rates of predators and prey is one of the oldest explanations for the existence of inverted trophic pyramids. To date, the hypotheses regarding trophic pyramids have all been based on consumptive trophic links between predators and prey, and the relative contribution of non-consumptive effects is still unknown. In this study, we investigated if the inversion of pyramids in bromeliad ecosystems is driven by (i) a rapid colonization of organisms having short cohort interval production (CPI), and (ii) the prevalence of consumptive or non-consumptive effects of top predators. We used a manipulative experiment to investigate the patterns of prey colonization and to partition the net effects of the dominant predator (damselfly larvae) on biomass pyramids into consumptive (uncaged damselfly larvae) and non-consumptive effects (caged damselfly larvae). Consumptive effects of damselflies strengthened the inversion of trophic pyramids. Non-consumptive effects, however, did not affect the shape of biomass pyramids. Instead, the rapid colonization of organisms with predominantly short CPI sustained the large biomass of top predators found in natural bromeliad ecosystems. Prey colonized bromeliads rapidly, but this high production was never visible as standing stock because damselflies reduce prey densities by more than a magnitude through direct consumption. Our study adds to the growing evidence that there are a variety of possible ways that biomass can be trophically structured. Moreover, we suggest that the strength of biomass pyramids inversion may change with the time of ecological succession as prey communities become more equitable.

Competition during early ontogeny: Effects of native and invasive planktivores on the growth, survival, and habitat use of bluegill

Abstract Early life stages of fishes are sensitive to ecological and environmental disturbances and experience very high mortality rates. During early ontogeny, the growth and survival of larval fish often depends on food availability. Because habitat and diet shifts are strongly tied to individual body size, factors that influence the growth rates of individuals (e.g. resource limitation, competition) also affect other aspects of ontogeny including the timing of habitat or diet shifts. In the context of biological invasions, non‐native species can potentially disrupt the interaction of larval fish with their food via competition for shared prey, reducing growth and survival during a vulnerable period of an organism's life history. We hypothesised that invasive planktivores negatively affect native species through the vulnerable larval life stage via competition for zooplankton resources. To test this hypothesis, we conducted a series of experiments to assess and contrast the effects of invasive (bighead carp, Hypophthalmichthys nobilis) and native (golden shiners, Notemigonus crysoleucas) planktivores on zooplankton densities, and their effects on the growth, survival, abundance, and habitat use of larval bluegill (Lepomis macrochirus). Overall, the effects of the invasive planktivore were consistently greater than the native planktivore in terms of reduced prey densities, reduced bluegill growth rates, and delays to the timing of ontogenetic habitat shifts. Growth rates of bluegill larvae were reduced by 58–87% in the presence of bighead carp and 37% in the presence of golden shiners (relative to controls), but such reductions did not consistently lead to reduced survival (in mesocosm experiment) or relative abundance (in pond experiment). However, we estimated that bighead carp and golden shiners delayed ontogenetic habitat shifts in bluegill by 9–24 and 1–3 days, respectively. Although we did not detect an effect of planktivores on the numbers of larval bluegill, our findings suggest that bighead carp may still affect bluegill ontogeny and freshwater food webs by disrupting the timing of ontogenetic habitat shifts. By affecting the coupling of habitats via organism movements during early ontogeny, bighead carp may indirectly disrupt predator–prey interactions of native taxa.

Prey density affects strengths of density- and trait-mediated indirect interactions of predators on an algal community

In trophic cascades, predators can indirectly affect resources by reducing prey density (density-mediated indirect interactions; DMIIs) or by changing prey traits, such as their behavior, morphology, physiology, or life history (trait-mediated indirect interactions; TMIIs). Although the importance of predator (e.g., foraging strategy) and resource characteristics (e.g., quantity) in these indirect interactions is well recognized, little attention has been paid to prey characteristics (e.g., density), especially in field studies. We focused on a marine trophic cascade involving the carnivorous snail Thais clavigera, its limpet prey Siphonaria sirius, and the algae Lithoderma sp. and Ulva sp. Using intertidal rocks as natural replicates, we experimentally evaluated the in situ effects of the DMIIs and TMIIs on the algal community under two density ranges of prey. The strengths of consumptive effects (CEs) and non-consumptive effects (NCEs) of the predator on the prey limpet were also monitored to elucidate the mechanisms of the indirect interactions. At high densities, CEs decreased the percentage of individual limpets that disappeared (that likely died); however, neither DMIIs nor TMIIs were detected. At low densities, both CEs and NCEs reduced the per capita feeding rate and CEs reduced the growth rate of limpets. Moreover, Lithoderma sp. was replaced by Ulva sp. through both DMIIs and TMIIs. These results suggest that prey density is a key determinant of the strengths of CEs and NCEs, and notably, of DMIIs and TMIIs.

Using Predator-Prey Theory to Predict Outcomes of Broadscale Experiments to Reduce Apparent Competition

Apparent competition is an important process influencing many ecological communities. We used predator-prey theory to predict outcomes of ecosystem experiments aimed at mitigating apparent competition by reducing primary prey. Simulations predicted declines in secondary prey following reductions in primary prey because predators consumed more secondary prey until predator numbers responded to reduced prey densities. Losses were exacerbated by a higher carrying capacity of primary prey and a longer lag time of the predator's numerical response, but a gradual reduction in primary prey was less detrimental to the secondary prey. We compared predictions against two field experiments where endangered woodland caribou (Rangifer tarandus caribou) were victims of apparent competition. First, when deer (Odocoileus sp.) declined suddenly following a severe winter, cougar (Puma concolor) declined with a 1-2-year lag, yet in the interim more caribou were killed by cougars, and caribou populations declined by 40%. Second, when moose (Alces alces) were gradually reduced using a management experiment, wolf (Canis lupus) populations declined but did not shift consumption to caribou, and the largest caribou subpopulation stabilized. The observed contrasting outcomes of sudden versus gradual declines in primary prey supported theoretical predictions. Combining theory with field studies clarified how to manage communities to mitigate endangerment caused by apparent competition that affects many taxa.

Habitat selection and consumption across a landscape of multiple predators.

Predator community composition can alter habitat quality for prey by changing the strength and direction of consumptive effects. Whether predator community composition also alters prey density via nonconsumptive effects during habitat selection is not well known, but is important for understanding how changes to predator communities will alter prey populations. We tested the hypothesis that predator community composition (presence of caged trout, caged dragonflies, or caged trout + dragonflies) alters colonization of aquatic mesocosms by ovipositing aquatic insects. In a previous experiment in this system, we found a spatial contagion effect, in which insects avoided pools with predators, but only when predator-free pools were isolated (∼5 m away from predator pools). Here, we removed the isolated predator-free pools, allowing us to test whether insects would make fine-scale (∼1 m) oviposition decisions in the absence of preferred isolated pools. We also estimated consumptive effects by allowing predators to feed on colonists for 5 days following colonization. All insects collected after 21 days were dipterans, dominated by Chironomidae. Total colonization, measured as the number of developing larvae after 21 days, was not affected by either predator presence or composition. Consumption was significant in the trout only treatment, reducing larval insect density by 46 ± 37% (mean ± SE). No other predator treatment significantly reduced prey density, although the proportion of chironomid larvae in protective cases increased in response to direct predation from dragonflies, indicating an antipredatory behavioral response. Taken together, these results reveal that predator community composition altered larval survival and behavior, but colonizing females either did not or could not assess these risks across small scales during oviposition.

Attracted to the enemy: Aedes aegypti prefers oviposition sites with predator-killed conspecifics

Oviposition habitat choices of species with aquatic larvae are expected to be influenced by both offspring risk of mortality due to predation, and offspring growth potential. Aquatic predators may indirectly influence growth potential for prey by reducing prey density and, for filter-feeding prey, by increasing bacterial food for prey via added organic matter (feces, partially eaten victims), creating the potential for interactive effects on oviposition choices. We tested the hypothesis that the mosquito Aedes aegypti preferentially oviposits in habitats with predatory Toxorhynchites larvae because of indirect effects of predation on chemical cues indicating bacterial abundance. We predicted that A. aegypti would avoid oviposition in sites with Toxorhynchites, but prefer to oviposit where bacterial food for larvae is abundant, and that predation by Toxorhynchites would increase bacterial abundances. Gravid A. aegypti were offered paired oviposition sites representing choices among: predator presence; the act of predation; conspecific density; dead conspecific larvae; and bacterial activity. A. aegypti preferentially oviposited in sites with Toxorhynchites theobaldi predation, and with killed conspecific larvae, but failed to detect preferences for other treatments. The antibiotic tetracycline eliminated the strongest oviposition preference. Both predation by Toxorhynchites and killed larvae increased bacterial abundances, suggesting that oviposition attraction is cued by bacteria. Our results show the potential for indirect effects, like trophic cascades, to influence oviposition choices and community composition in aquatic systems. Our results suggest that predators like Toxorhynchites may be doubly beneficial as biocontrol agents because of the attraction of ovipositing mosquitoes to bacterial by-products of Toxorhynchites feeding.

Modelling the Effects of Prey Size and Distribution on Prey Capture Rates of Two Sympatric Marine Predators

Understanding how prey capture rates are influenced by feeding ecology and environmental conditions is fundamental to assessing anthropogenic impacts on marine higher predators. We compared how prey capture rates varied in relation to prey size, prey patch distribution and prey density for two species of alcid, common guillemot (Uria aalge) and razorbill (Alca torda) during the chick-rearing period. We developed a Monte Carlo approach parameterised with foraging behaviour from bird-borne data loggers, observations of prey fed to chicks, and adult diet from water-offloading, to construct a bio-energetics model. Our primary goal was to estimate prey capture rates, and a secondary aim was to test responses to a set of biologically plausible environmental scenarios. Estimated prey capture rates were 1.5±0.8 items per dive (0.8±0.4 and 1.1±0.6 items per minute foraging and underwater, respectively) for guillemots and 3.7±2.4 items per dive (4.9±3.1 and 7.3±4.0 items per minute foraging and underwater, respectively) for razorbills. Based on species' ecology, diet and flight costs, we predicted that razorbills would be more sensitive to decreases in 0-group sandeel (Ammodytes marinus) length (prediction 1), but guillemots would be more sensitive to prey patches that were more widely spaced (prediction 2), and lower in prey density (prediction 3). Estimated prey capture rates increased non-linearly as 0-group sandeel length declined, with the slope being steeper in razorbills, supporting prediction 1. When prey patches were more dispersed, estimated daily energy expenditure increased by a factor of 3.0 for guillemots and 2.3 for razorbills, suggesting guillemots were more sensitive to patchier prey, supporting prediction 2. However, both species responded similarly to reduced prey density (guillemot expenditure increased by 1.7; razorbill by 1.6), thus not supporting prediction 3. This bio-energetics approach complements other foraging models in predicting likely impacts of environmental change on marine higher predators dependent on species-specific foraging ecologies.

Disease in group-defending prey can benefit predators

Infectious diseases have the capacity not only to influence the host population but also to interacting species like predators. In particular, they can reduce host densities, which can have knock-on effects on predators. Here, we consider how an infectious disease in the prey affects the predator–prey relationship where the prey exhibit some kind of group defence against the predator (using a Holling type IV functional response). We find that the disease can reduce prey densities to levels where the group defence is weaker. This weakened group defence allows predators to survive in many situations where they could not without the disease.

Growth and survival of larval and early juvenile Lesser Sandeel in patchy prey field in the North Sea: An examination using individual-based modeling

Disentangling physical–biological interaction processes during early life-stages of fish is crucial for the understanding of fish stock recruitment. Among many individual and environmental aspects affecting mortality during the early life-stages of fish, encountering food at greater than average concentrations is regarded important for survival. Intense aggregations of zooplankton in near-surface waters provide these conditions for larval fish. Simulation studies by individual-based modeling can help understanding of the mechanisms for survival during early life-stages. In this study, we examined how growth and survival of larvae and early juveniles of Lesser Sandeel (Ammodytes marinus) in the North Sea are influenced by availability and patchiness of the planktonic prey by adapting and applying a generic bioenergetic individual-based model for larval fish. Input food conditions were generated by modeling copepod size spectra dynamics and patchiness based on particle count transects and Continuous Plankton Recorder time series data. The study analyzes the effects of larval hatching time, presence of zooplankton patchiness and within patch abundance on growth and survival of sandeel early life-stages in the North Sea. Simulations of patchiness related starvation mortality are able to explain observed patterns of variation in sandeel growth. Reduced prey densities within patches decrease growth and survival rate of larvae and match–mismatch affect growth and survival of larvae with different hatch time due to plankton seasonality. Of general scientific and environmental management interest, the results indicate a steep threshold concentration critical for survival at around 0.04–0.05 no. zooplankton/mL.

The advantage of no defense: predation enhances cohort survival in a desert amphibian

The impacts that predators have on prey behavior, growth, survival, and ultimately the composition of many ecological communities are mediated by prey defenses and the susceptibility of prey to predators. We hypothesized that prey populations inhabiting short-lived, species-poor, aquatic environments should lack significant morphological, developmental, and behavioral responses to predators and are therefore highly susceptible to predation. Furthermore, we predicted that the resultant decrease in prey density and increase in per capita resources due to high susceptibility to predators should enhance overall cohort survival because of enhanced growth of surviving prey. To test these ideas, we performed laboratory and outdoor mesocosm experiments to disentangle multiple effects of predators on an anuran (Scaphiopus couchii); a species highly adapted to breeding in ephemeral habitats and that has one of the shortest larval periods of all anurans. Chemical (presence of predator) and lethal predator cues (predator plus consumed conspecific) elicited no response in behavior, development, or morphology, indicating a lack of defensive mechanisms. Survivorship was significantly reduced in treatments where tadpoles were exposed to predators. However, this reduction in prey density led to accelerated time to metamorphosis, conferring an advantage to survivors who must metamorphose before ephemeral ponds dry. Our experiments demonstrated that in short-lived environments, prey may exhibit little or no response to the presence of predators presumably because selection for anti-predator defenses is countered by selection for rapid metamorphosis. However, predation actually resulted in an increase in overall cohort survival. Although predators are relatively rare in highly ephemeral aquatic environments, they may play an important role in facilitating the long-term persistence of their prey by reducing prey density.

Experimental test of preference by a predatory fish for prey at different densities

Preference for a particular prey implies that a behavioral choice is made by the predator, requiring an experimental approach to separate process from observed patterns in nature. For example, pink shrimp ( Farfantepenaeus duorarum) and pinfish ( Lagodon rhomboides) are dominant prey observed in the diet of juvenile gag ( Mycteroperca microlepis) collected from natural habitats, but it is unclear whether their approximate equal representation is due to a lack of preference between the two. Furthermore, both prey are captured in high quantities as targeted (shrimp) and non-targeted (pinfish) components of a trawl fishery operating in the same seagrass habitats where juvenile gag are found, thus requiring examination as to whether reductions in prey densities affect consumption rates and preference. In the current study, I used laboratory feeding experiments and applied new analytical techniques derived in a series of recent papers published in this journal to examine preference by juvenile gag on these two morphologically and behaviorally different prey. Consumption rates were higher for shrimp than pinfish and were not related to the initial density of prey presented to gag. However, gag did not exhibit a feeding preference for either prey across 10 controlled comparisons. Experimental results in the laboratory therefore reflected patterns observed in the field for the diet of this predatory fish. Moreover, by examining the effects of reduced prey densities on food web processes, the approach used in this study may be applied to inform ecosystem-based management on indirect effects of fishing.

Bird predation alters infestation of desert lizards by parasitic mites

Theory predicts that predators can reduce parasite abundance on prey by reducing prey density and through disproportionate predation on heavily infested individuals. We experimentally tested this prediction by examining the effects of bird predation on parasitic mite infestation of the prey lizard Acanthodactylus beershebensis. We manipulated predation by adding perches to arid scrubland, allowing avian predators to hunt for lizards in a habitat the birds would not normally use. Host density influenced parasite abundance in hatchlings, but not in older aged individuals and parasite abundance did not affect lizard host survival. Contrary to expectation mite abundance on adult lizards increased under low predation intensities. We explain these results by suggesting a novel hypothesis based on the assumption that the two components of predation, i.e. actual removal of prey and risk, exert contradictory effects on macroparasite abundance.

Disentangling multiple predator effects in biodiversity and ecosystem functioning research

Disentangling multiple predator effects in biodiversity and ecosystem functioning research

The Many Faces of Fear: Comparing the Pathways and Impacts of Nonconsumptive Predator Effects on Prey Populations

BackgroundMost ecological models assume that predator and prey populations interact solely through consumption: predators reduce prey densities by killing and consuming individual prey. However, predators can also reduce prey densities by forcing prey to adopt costly defensive strategies.Methodology/Principal FindingsWe build on a simple Lotka-Volterra predator-prey model to provide a heuristic tool for distinguishing between the demographic effects of consumption (consumptive effects) and of anti-predator defenses (nonconsumptive effects), and for distinguishing among the multiple mechanisms by which anti-predator defenses might reduce prey population growth rates. We illustrate these alternative pathways for nonconsumptive effects with selected empirical examples, and use a meta-analysis of published literature to estimate the mean effect size of each pathway. Overall, predation risk tends to have a much larger impact on prey foraging behavior than measures of growth, survivorship, or fecundity.Conclusions/SignificanceWhile our model provides a concise framework for understanding the many potential NCE pathways and their relationships to each other, our results confirm empirical research showing that prey are able to partially compensate for changes in energy income, mitigating the fitness effects of defensive changes in time budgets. Distinguishing the many facets of nonconsumptive effects raises some novel questions, and will help guide both empirical and theoretical studies of how predation risk affects prey dynamics.