Density-dependent prey behaviours and mutable predator foraging modes induce Allee effects and over-prediction of prey mortality rates

1. Predator–prey models are often used to represent consumptive interactions between species but, typically, are derived using simple experimental systems with little plasticity in prey or predator behaviours. However, many prey and predators exhibit a broad suite of behaviours. Here, we experimentally tested the effect of density-dependent prey and predator behaviours on per capita relative mortality rates using Florida bass (Micropterus floridanus) consuming juvenile Bluegill (Lepomis macrochirus).
2. Experimental ponds were stocked with a factorial design of low, medium, and high prey and predator densities. Prey mortality, prey–predator behaviours, and predator stomach contents were recorded over or after 7 days. We assumed the mortality dynamics followed foraging arena theory. This pathologically flexible predator–prey model separates prey into invulnerable and vulnerable pools where predators can consume prey in the latter. As this approach can represent classic Lotka–Volterra and ratio-dependent dynamics, we fit a foraging arena predator–prey model to the number of surviving prey.
3. We found that prey exhibited density-dependent prey behaviours, hiding at low densities, shoaling at medium densities, and using a provided refuge at high densities. Predators exhibited ratio-dependent behaviours, using an ambush foraging mode when one predator was present, hiding in the shadows at low prey–high predator densities, and shoaling at medium and high prey–high predator densities. The foraging arena model predicted the mortality rates well until the high prey–high predator treatment where group vigilance prey behaviours occurred and predators probably interfered with one another resulting in the model predicting higher mortality than observed.
4. This is concerning given the ubiquity of predator–prey models in ecology and natural resource management. Furthermore, as Allee effects engender instability in population regulation, it could lead to inaccurate predictions of conservation status, population rebuilding or harvest rates.

     

Piscivore efficiency and refuging prey: the importance of predator search mode

In predator-prey interactions, the efficiency of the predator is dependent on characteristics of both the predator and the prey, as well as the structure of the environment. In a field enclosure experiment, we tested the effects of a prey refuge on predator search mode, predator efficiency and prey behaviour. Replicated enclosures containing young of the year (0+) and 1-year-old (1+) perch were stocked with 3 differentially sized individuals of either of 2 piscivorous species, perch (Perca fluviatilis), pike (Esox lucius) or no piscivorous predators. Each enclosure contained an open predator area with three small vegetation patches, and a vegetated absolute refuge for the prey. We quantified the behaviour of the predators and the prey simultaneously, and at the end of the experiment the growth of the predators and the mortality and habitat use of the prey were estimated. The activity mode of both predator species was stationary. Perch stayed in pairs in the vegetation patches whereas pike remained solitary and occupied the corners of the enclosure. The largest pike individuals stayed closest to the prey refuge whereas the smallest individuals stayed farthest away from the prey refuge, indicating size-dependent interference among pike. Both size classes of prey showed stronger behavioural responses to pike than to perch with respect to refuge use, distance from refuge and distance to the nearest predator. Prey mortality was higher in the presence of pike than in the presence of perch. Predators decreased in body mass in all treatments, and perch showed a relatively stronger decrease in body mass than pike during the experiment. Growth differences of perch and pike, and mortality differences of prey caused by predation, can be explained by predator morphology, predator attack efficiency and social versus interference behaviour of the predators. These considerations suggest that pike are more efficient piscivores around prey refuges such as the littoral zones of lakes, whereas perch have previously been observed to be more efficient in open areas, such as in the pelagic zones of lakes.     

Consequences of size structure in the prey for predator-prey dynamics: the composite functional response

1. Current formulations of functional responses assume that the prey is homogeneous and independent of intraspecific processes. Most prey populations consist of different coexisting size classes that often engage in asymmetrical intraspecific interactions, including cannibalism, which can lead to nonlinear interaction effects. This may be important as the size structure with the prey could alter the overall density-dependent predation rates. 2. In a field experiment with damselfly and dragonfly larvae, 16 treatments manipulated the density of a small prey stage, the presence of large conspecific prey and the presence of heterospecific predators. 3. Size structure in the prey (i.e. when both prey stages were present) decreased the impact of the predator on overall prey mortality by 25-48% at mid and high prey densities, possibly due to density-dependent size-structured cannibalism in the prey. The predation rates on small prey stages were determined by the interaction of large prey and predators. Predation rates increased with prey density in the absence of large prey, but predation rates were constant across densities when large conspecifics were present. 4. The functional response for unstructured prey followed a Holling type III model, but the predation rate for size-structured prey was completely different and followed a complex pattern that could not be explained with any standard functional response. 5. Using additional laboratory experiments, a mortality model was developed and parameterized. It showed that the overall prey mortality of size-structured prey can be adequately predicted with a composite functional response model that modelled the individual functional responses of each prey stage separately and accounted for their cannibalistic interaction. 6. Thus, treating a prey population as a homogeneous entity will lead to erroneous predictions in most real-world food webs. However, if we account for the effects of size structure and the intraspecific interactions on functional responses by treating size classes as different functional groups, it is possible to reliably predict the dynamics of size-structured predator-prey systems.     

Predator impacts on stream benthic prey

The impact that predators have on benthic, macroinvertebrate prey density in streams is unclear. While some studies show a strong effect of predators on prey density, others show little or no effect. Two factors appear to influence the detection of predator impact on prey density in streams. First, many field studies have small sample sizes and thus might be unable to detect treatment effects. Second, streams contain two broad classes of predators, invertebrates and vertebrates, which might have different impacts on prey density for a variety of reasons, including availability of refuge for prey and prey emigration responses to the two types of predators. In addition, predatory vertebrates have more complex prey communities than predatory invertebrates; this complexity might reduce the impact that predatory vertebrates have on prey because of indirect effects. I conducted a meta-analysis on the results of field studies that manipulate predator density in enclosures to determine (1) if predators have a significant impact on benthic prey density in streams, (2) if the impacts that predatory invertebrates and vertebrates have differ, and (3) if predatory vertebrates have different impacts on predatory prey versus herbivorous prey. The results of the meta-analysis suggest that on average predators have a significant negative effect on prey density, predatory invertebrates have a significantly stronger impact than predatory vertebrates, and predatory vertebrates do not differ in their impact on predatory versus herbivorous invertebrate prey. Three methodological variables (mesh size of enclosures, size of enclosures, and experimental duration) were examined to determine if cross correlations exist that may explain the differences in impact between predatory invertebrates and vertebrates. No correlation exists between mesh size and predator impact. Over all predators, no correlation exists between experimental duration and predator impact; however, within predatory invertebrates a correlation does exist between these variables. Also, a correlation was found between enclosure size and predator impact. This correlation potentially explains the difference in impact between predatory invertebrates and predatory vertebrates. Results of the meta-analysis suggest two important areas for future research: (1) manipulate both types of predators within the same system, and (2) examine their impacts on the same spatial scale.     

Effects of size structure and habitat complexity on predator–prey interactions

1. A predator's ability to suppress its prey depends on the level of interference among predators. While interference typically decreases with increasing habitat complexity, it often increases with increasing size differences among individuals. However, little is known about how variation in intrinsic factors such as population size structure alters predator–prey interactions and how this intrinsic variation interacts with extrinsic variation. 2. By experimentally varying the level of vegetation cover and the size structure of the predatory damselfly Ischnura posita Hagen, we examined the individual and interactive effects of variation in habitat complexity and predator size structure on prey mortality. 3. Copepod prey survival linearly increased as the I. posita size ratio decreased and differed by up to 31% among different predator size structures. Size classes had an additive effect on prey survival, most likely because intraspecific aggression appeared size‐independent and size classes differed in microhabitat preference: large I. posita spent 14% more time foraging on the floor than small larvae and spent more time in the vegetation with increasing habitat complexity. Despite this difference in microhabitat use among size classes, habitat structure did not influence predation rates or interference among size classes. 4. In general, results suggest that seasonal and spatial variation in the size structure of populations could drive some of the discrepancies in predator‐mediated prey suppression observed in nature, and this variation could exceed the effects of variation in habitat structure.     

Fleeing towards death – leech‐induced behavioural defences increase freshwater snail susceptibility to predatory fish

Prey species are often exposed to multiple predators, which presents several difficulties to prey species. This is especially true when the response to one predator influences the prey’s susceptibility to other predators. Predator‐induced defences have evolved in a wide range of prey species, and experiments involving predators with different hunting strategies allow researchers to evaluate how prey respond to multiple threats. Freshwater snails are known to respond to a variety of predators with both morphological and behavioural defences. Here we studied how freshwater snails Radix balthica responded behaviourally to fish and leech predators, both separately and together. Our aim was to explore whether conflicting predator‐induced responses existed and, if so, what effect they had on snail survival when both predatory fish and leeches were present. We found that although R. balthica increased refuge use when exposed to predatory fish, they decreased refuge use when exposed to predatory leeches. When both predators were present, snails showed a stronger response towards leech than fish and responded by leaving the refuge. This response made the snails more susceptible to fish predation, which increased snail mortality when exposed to both fish and leech compared to fish only. We show that predators that have a relatively low predation rate can substantially increase mortality rates by indirect effects. By forcing snails out of refuges such as rock and macrophyte habitats, leeches can indirectly increase predation from molluscivorous fish and may thus affect snail densities.     

Spatial refuge from intraguild predation: implications for prey suppression and trophic cascades

The ability of predators to elicit a trophic cascade with positive impacts on primary productivity may depend on the complexity of the habitat where the players interact. In structurally-simple habitats, trophic interactions among predators, such as intraguild predation, can diminish the cascading effects of a predator community on herbivore suppression and plant biomass. However, complex habitats may provide a spatial refuge for predators from intraguild predation, enhance the collective ability of multiple predator species to limit herbivore populations, and thus increase the overall strength of a trophic cascade on plant productivity. Using the community of terrestrial arthropods inhabiting Atlantic coastal salt marshes, this study examined the impact of predation by an assemblage of predators containing Pardosa wolf spiders, Grammonota web-building spiders, and Tytthus mirid bugs on herbivore populations (Prokelisia planthoppers) and on the biomass of Spartina cordgrass in simple (thatch-free) and complex (thatch-rich) vegetation. We found that complex-structured habitats enhanced planthopper suppression by the predator assemblage because habitats with thatch provided a refuge for predators from intraguild predation including cannibalism. The ultimate result of reduced antagonistic interactions among predator species and increased prey suppression was enhanced conductance of predator effects through the food web to positively impact primary producers. Behavioral observations in the laboratory confirmed that intraguild predation occurred in the simple, thatch-free habitat, and that the encounter and capture rates of intraguild prey by intraguild predators was diminished in the presence of thatch. On the other hand, there was no effect of thatch on the encounter and capture rates of herbivores by predators. The differential impact of thatch on the susceptibility of intraguild and herbivorous prey resulted in enhanced top-down effects in the thatch-rich habitat. Therefore, changes in habitat complexity can enhance trophic cascades by predator communities and positively impact productivity by moderating negative interactions among predators.     

Influence of intra‐ and interspecific variation in predator–prey body size ratios on trophic interaction strengths

1. Predation is a pervasive force that structures food webs and directly influences ecosystem functioning. The relative body sizes of predators and prey may be an important determinant of interaction strengths. However, studies quantifying the combined influence of intra‐ and interspecific variation in predator–prey body size ratios are lacking.
2. We use a comparative functional response approach to examine interaction strengths between three size classes of invasive bluegill and largemouth bass toward three scaled size classes of their tilapia prey. We then quantify the influence of intra‐ and interspecific predator–prey body mass ratios on the scaling of attack rates and handling times.
3. Type II functional responses were displayed by both predators across all predator and prey size classes. Largemouth bass consumed more than bluegill at small and intermediate predator size classes, while large predators of both species were more similar. Small prey were most vulnerable overall; however, differential attack rates among prey were emergent across predator sizes. For both bluegill and largemouth bass, small predators exhibited higher attack rates toward small and intermediate prey sizes, while larger predators exhibited greater attack rates toward large prey. Conversely, handling times increased with prey size, with small bluegill exhibiting particularly low feeding rates toward medium–large prey types. Attack rates for both predators peaked unimodally at intermediate predator–prey body mass ratios, while handling times generally shortened across increasing body mass ratios.
4. We thus demonstrate effects of body size ratios on predator–prey interaction strengths between key fish species, with attack rates and handling times dependent on the relative sizes of predator–prey participants.
5. Considerations for intra‐ and interspecific body size ratio effects are critical for predicting the strengths of interactions within ecosystems and may drive differential ecological impacts among invasive species as size ratios shift.

     

Human activity helps prey win the predator-prey space race

Predator-prey interactions, including between large mammalian wildlife species, can be represented as a "space race", where prey try to minimize and predators maximize spatial overlap. Human activity can also influence the distribution of wildlife species. In particular, high-human disturbance can displace large carnivore predators, a trait-mediated direct effect. Predator displacement by humans could then indirectly benefit prey species by reducing predation risk, a trait-mediated indirect effect of humans that spatially decouples predators from prey. The purpose of this research was to test the hypothesis that high-human activity was displacing predators and thus indirectly creating spatial refuge for prey species, helping prey win the "space race". We measured the occurrence of eleven large mammal species (including humans and cattle) at 43 camera traps deployed on roads and trails in southwest Alberta, Canada. We tested species co-occurrence at camera sites using hierarchical cluster and nonmetric multidimensional scaling (NMS) analyses; and tested whether human activity, food and/or habitat influenced predator and prey species counts at camera sites using regression tree analysis. Cluster and NMS analysis indicated that at camera sites humans co-occurred with prey species more than predator species and predator species had relatively low co-occurrence with prey species. Regression tree analysis indicated that prey species were three times more abundant on roads and trails with >32 humans/day. However, predators were less abundant on roads and trails that exceeded 18 humans/day. Our results support the hypothesis that high-human activity displaced predators but not prey species, creating spatial refuge from predation. High-human activity on roads and trails (i.e., >18 humans/day) has the potential to interfere with predator-prey interactions via trait-mediated direct and indirect effects. We urge scientist and managers to carefully consider and quantify the trait-mediated indirect effects of humans, in addition to direct effects, when assessing human impacts on wildlife and ecosystems.     

Implications of a fluctuating fish predator guild on behavior, distribution, and abundance of a shared prey species: the grass shrimp Palaemonetes pugio

In some systems, the identity of a prey species' dominant predator(s) may not be constant over time. In cases in which a prey species exhibits different responses to various predator species, such changes in predator identity may have population-wide consequences. Our goals were to determine (1) whether mortality of and refuge use by the grass shrimp, Palaemonetes pugio, were predator-specific, and (2) how effects of prey size and habitat interacted with predator type. Striped bass (Morone saxatilis) exerted twice as much predation pressure as mummichog (Fundulus heteroclitus), although not equally as great on large (female) and small (male) shrimp. Mummichog, which fed preferentially on large shrimp, forced a partitioning of habitat between the two shrimp size classes. In contrast, large and small shrimp occupied similar habitats when subjected to striped bass, which fed on both size classes equally. Refuge use of grass shrimp depended on predator type. In the presence of mummichog, which occupied shallower depths in the water column than striped bass, shrimp stayed deep and close to structural habitat. Striped bass, which were deeper, caused shrimp to move high in the water column away from structural habitat. When both predators were present, shrimp distribution was similar to that when only striped bass were present, striped bass predation rate was enhanced, and overall mortality was higher than with either predator alone. Results suggest that at times when mummichogs are the dominant predators, large (female) shrimp experience higher predation than small (male) shrimp and are physically separated from their potential mates. When striped bass are more abundant, male and female shrimp may share a similar, shallow, less structure-oriented distribution and be subjected to higher mortality. When both predators are present, mortality rates may be higher still. This predator-, size-, and habitat-specificity of grass shrimp behavior suggests significant population and distribution consequences of fluctuating predator guilds and fluctuating cover of structural habitats in the field.     

Emergent impacts of cannibalism and size refuges in prey on intraguild predation systems

Many organisms undergo ontogenetic niche shifts due to considerable changes in size during their development. These ontogenetic shifts can alter the trophic position of individuals, the type and strength of ecological interactions across species, and allow for cannibalism within species. In this study we ask if and how the interaction of a size refuge and cannibalism in the prey alters the dynamics of intraguild predation (IGP) systems. By manipulating the composition of large cannibalistic (Aeshna umbrosa) and predatory (Anax junius) dragonfly larvae in mesocosms we show that the interaction of cannibals and predators was non-linear and increased the survival of prey. The structure of the final resource community shared by prey and predator differed between small and large dragonfly treatments but not within size classes across species. In general, the small prey stage showed similar shifts in microhabitat use and refuge use when exposed to either conspecific cannibals or predators, while large cannibals showed no clear anti-predator response. However, further behavioral experiments revealed that specific behavioral components, such as distances between individuals or number of movements, differed when individuals were exposed to either cannibals or predators. This indicates that individuals discriminated between conspecific or heterospecific predators. Furthermore, in similar experiments large cannibals and predators showed different behaviors when exposed to conspecifics rather than to each other. These changes in behavior are consistent with the observed increase in prey survival. In general, the results indicate that cannibalism and ontogenetic niche shifts can result in behavior-mediated indirect interactions that reduce the impact of the predator on the mortality of its prey and alter the interactions of IGP systems. However, they also indicate that size is not the sole determinant and that we also need to account for the species identity when predicting the dynamics of communities.     

Indirect effects between shared prey: Predictions for

Suppression of a target prey by a predator can depend on its surrounding community, including the presence of nontarget, alternative prey. Basic theoretical models of two prey species that interact only via a shared predator predict that adding an alternative prey should increase predator numbers and ultimately lower target pest densities as compared to when the target pest is the only prey. While this is an alluring prediction, it does not explain the numerous responses empirically observed. To better understand and predict the indirect interactions produced by shared predation, we explore how additional prey species affect three broad ecological mechanisms, the predator's reproductive, movement, and functional responses. Specifically, we review current theoretical models of shared predation by focusing on these mechanisms, and make testable predictions about the effects of shared predation. We find that target predation is likely to be higher in the two prey system because of predator reproduction, especially when: predators are prey limited, alternative or total prey density is high, or alternative prey are available over time. Target predation may also be greater because of predator movement, but only under certain movement rules and spatial distributions. Predator foraging behavior is most likely to cause lower target predation in the two-prey system, when per capita predation is limited by something other than prey availability. It is clear from this review that no single theoretical generalization will accurately predict community-level effects for every system. However, we can provide testable hypotheses for future empirical and theoretical investigations of indirect interactions and help enhance their potential use in biological control.     

Female blue tits with brighter yellow chests transfer more carotenoids to their eggs after an immune challenge

Predicting the consequences of predator biodiversity loss on prey requires an understanding of multiple predator interactions. Predators are often assumed to have independent and additive effects on shared prey survival; however, multiple predator effects can be non-additive if predators foraging together reduce prey survival (risk enhancement) or increase prey survival through interference (risk reduction). In marine communities, juvenile reef fish experience very high mortality from two predator guilds with very different hunting modes and foraging domains—benthic and pelagic predator guilds. The few previous predator manipulation studies have found or assumed that mortality is independent and additive. We tested whether interacting predator guilds result in non-additive prey mortality and whether the detection of such effects change over time as prey are depleted. To do so, we examined the roles of benthic and pelagic predators on the survival of a juvenile shoaling zooplanktivorous temperate reef fish, Trachinops caudimaculatus, on artificial patch reefs over 2 months in Port Phillip Bay, Australia. We observed risk enhancement in the first 7 days, as shoaling behaviour placed prey between predator foraging domains with no effective refuge. At day 14 we observed additive mortality, and risk enhancement was no longer detectable. By days 28 and 62, pelagic predators were no longer significant sources of mortality and additivity was trivial. We hypothesize that declines in prey density led to reduced shoaling behaviour that brought prey more often into the domain of benthic predators, resulting in limited mortality from pelagic predators. Furthermore, pelagic predators may have spent less time patrolling reefs in response to declines in prey numbers. Our observation of the changing interaction between predators and prey has important implications for assessing the role of predation in regulating populations in complex communities.     

Seasonal shifts in predator body size diversity and trophic interactions in size-structured predator-prey systems

1.?Theory suggests that the relationship between predator diversity and prey suppression should depend on variation in predator traits such as body size, which strongly influences the type and strength of species interactions. Prey species often face a range of different sized predators, and the composition of body sizes of predators can vary between communities and within communities across seasons. 2.?Here, I test how variation in size structure of predator communities influences prey survival using seasonal changes in the size structure of a cannibalistic population as a model system. Laboratory and field experiments showed that although the per-capita consumption rates increased at higher predator-prey size ratios, mortality rates did not consistently increase with average size of cannibalistic predators. Instead, prey mortality peaked at the highest level of predator body size diversity. 3.?Furthermore, observed prey mortality was significantly higher than predictions from the null model that assumed no indirect interactions between predator size classes, indicating that different sized predators were not substitutable but had more than additive effects. Higher predator body size diversity therefore increased prey mortality, despite the increased potential for behavioural interference and predation among predators demonstrated in additional laboratory experiments. 4.?Thus, seasonal changes in the distribution of predator body sizes altered the strength of prey suppression not only through changes in mean predator size but also through changes in the size distribution of predators. In general, this indicates that variation (i.e. diversity) within a single trait, body size, can influence the strength of trophic interactions and emphasizes the importance of seasonal shifts in size structure of natural food webs for community dynamics.     

Lack of prey switching and strong preference for mosquito prey by a temporary pond specialist predator

1. The strengths of trophic interactions within ecosystems can be mediated by complex mechanisms that require elucidation if researchers are to understand and predict population- and community-level stabilities. Where multiple prey types co-occur, prey switching (i.e. frequency-dependent predation) by predators may facilitate low-density prey refuge effects which promote coexistence. On the other hand, lack of switching and strong preferences by predators can strongly suppress prey populations, which is especially important considering vector species such as mosquitoes. 2. The present study quantifies prey switching and preference patterns of the temporary pond specialist copepod Lovenula raynerae towards larvae of the medically important Culex pipiens mosquito complex in the presence of different proportions of alternative Daphnia pulex prey. Further, it examines whether prey switching and preferences are contingent on the sex of the predator. 3. Lovenula raynerae exhibited a lack of prey switching and strong preference for larval mosquito prey overall, irrespective of predator sex. Also, when larval mosquitoes were available in higher proportions over daphniids, the strength of this positive selectivity increased. There was very little low-density refuge for mosquitoes where they were rare. 4. Lack of prey switching and strong preferences towards mosquitoes by predatory paradiaptomid copepods may enhance population-level regulation of disease vector mosquitoes that exploit temporary pond-style habitats. Accordingly, the conservation and promotion of these predators might enable better management of medically important species across landscapes.     

18S rRNA metabarcoding diet analysis of a predatory fish community across seasonal changes in prey availability

Predator–prey relationships are important ecological interactions, affecting biotic community composition and energy flow through a system, and are of interest to ecologists and managers. Morphological diet analysis has been the primary method used to quantify the diets of predators, but emerging molecular techniques using genetic data can provide more accurate estimates of relative diet composition. This study used sequences from the 18S V9 rRNA barcoding region to identify prey items in the gastrointestinal (GI) tracts of predatory fishes. Predator GI samples were taken from the Black River, Cheboygan Co., MI, USA (n = 367 samples, 12 predator species) during periods of high prey availability, including the larval stage of regionally threatened lake sturgeon (Acipenser fulvescens Rafinesque 1817) in late May/early June of 2015 and of relatively lower prey availability in early July of 2015. DNA was extracted and sequenced from 355 samples (96.7%), and prey DNA was identified in 286 of the 355 samples (80.6%). Prey were grouped into 33 ecologically significant taxonomic groups based on the lowest taxonomic level sequences that could be identified using sequences available on GenBank. Changes in the makeup of diet composition, dietary overlap, and predator preference were analyzed comparing the periods of high and low prey abundance. Some predator species exhibited significant seasonal changes in diet composition. Dietary overlap was slightly but significantly higher during the period of high prey abundance; however, there was little change in predator preference. This suggests that change in prey availability was the driving factor in changing predator diet composition and dietary overlap. This study demonstrates the utility of molecular diet analysis and how temporal variability in community composition adds complexity to predator–prey interactions.     

Predator size and phenology shape prey survival in temporary ponds

Theoretical efforts suggest that the relative sizes of predators and their prey can shape community dynamics, the structure of food webs, and the evolution of life histories. However, much of this work has assumed static predator and prey body sizes. The timing of recruitment and the growth patterns of both predator and prey have the potential to modify the strength of predator–prey interactions. In this study, I examined how predator size dynamics in 40 temporary ponds over a 3-year period affected the survival of spotted salamander (Ambystoma maculatum) larvae. Across communities, gape-limited predator richness, but not size, was correlated with habitat duration (pond permanence). Within communities, mean gape-limited predator size diminished as the growing season progressed. This size reduction occurred because prey individuals grew into a body size refuge and because the largest of the predators left ponds by mid-season. Elevated gape-limited predation risk across time and space was predicted by the occurrence of two large predatory salamanders: marbled salamander larvae (Ambystoma opacum) and red-spotted newt adults (Notophthalmus viridescens). The presence of the largest gape-limited predator, A. opacum, predicted A. maculatum larval survival in the field. The distribution of large predatory salamanders among ponds and across time is expected to lead to differing community dynamics and to generate divergent natural selection on early growth and body size in A. maculatum. In general, a dynamic perspective on predator size often will be necessary to understand the ecology and evolution of species interactions. This will be especially true in frequently disturbed or seasonal habitats where phenology and ontogeny interact to determine body size asymmetries. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

The influence of predator regime on the behaviour and mortality of a freshwater amphipod, <Emphasis Type="Italic">Gammarus pulex</Emphasis>

In species with restricted dispersal, traits may become genetically fixed leading to local adaptations. Therefore, predator avoidance in a prey species may differ between populations experiencing different predator regimes, but also between sexes within a population due to different vulnerability to predators. In this study we used male and female Gammarus pulex from two different predator regimes: fishless ponds, where invertebrates are the dominant predators and ponds with predatory fish. In the laboratory we examined refuge use, mortality, leaf decomposition rate and pair-formation in G. pulex when exposed to predator cues from either invertebrate predators or fish. Individuals from fish ponds spent more time in refuge and had a higher mortality than those from fishless ponds independent of predator cues. There was no effect of pond predator regime or predator cues on leaf decomposition rates. Further, fewer individuals formed pairs in G. pulex from fish ponds than from fishless ponds. Male G. pulex had a higher mortality and a higher decomposition rate than females independent of predator cues. However, there was no difference in refuge use between sexes. Our study shows that there are general differences in behaviour traits, both between predator regimes and sexes in G. pulex.     

The non-lethal effects of predators and the influence of food availability on life history of adult Chironomus tentans (Diptera: Chironomidae)

1. We used a laboratory experiment to determine effects of a predator (other than straightforward prey consumption) and food availability, during the larval stage, on adult size, age at emergence and fecundity of Chironomus tentans. 2. Predator presence and decreased food availability resulted in reduced adult emergence. 3. Predator presence and food availability resulted in smaller size and greater age at emergence of male and female chironomids. There were no significant interactions between effects of predator presence and food availability. 4. Predator presence had no significant effect on fecundity. 5. Smaller size and greater age at emergence can have important implications for adult survival, size and age at first reproduction and, thus, intrinsic rate of population growth.     

食饵庇护所对斑块环境下Leslie-Gower捕食系统的影响

建立了一个显式含有空间庇护所的两斑块Leslie-Gower捕食者-食饵系统。假设只有食饵种群在斑块间以常数迁移率迁移,且在每个斑块上食饵间的迁移比局部捕食者-食饵相互作用发生的时间尺度要快。利用两个时间尺度,可以构建用来描述所有斑块总的食饵和捕食者密度的综合系统。数学分析表明,在一定条件下,存在唯一的正平衡点,并且此平衡点全局稳定。进一步,捕食者的数量随着食饵庇护所数量增加而降低;在一定条件下,食饵的数量随着食饵庇护所数量增加先增加后降低,在足够强的庇护所强度下,两物种出现灭绝。对比以往研究,利用显式含有和隐含空间庇护所的数学模型所得结论不一致,这意味着在研究庇护所对捕食系统种群动态影响时,空间结构可能起着重要作用。