The advantage of alternative tactics of prey and predators depends on the spatial pattern of prey and social interactions among predators

Individual variation in behavioral strategies is ubiquitous in nature. Yet, explaining how this variation is being maintained remains a challenging task. We use a spatially-explicit individual-based simulation model to evaluate the extent to which the efficiency of an alternative spacing tactic of prey and an alternative search tactic of predators are influenced by the spatial pattern of prey, social interactions among predators (i.e., interference and information sharing) and predator density. In response to predation risk, prey individuals can either spread out or aggregate. We demonstrate that if prey is extremely clumped, spreading out may help when predators share information regarding prey locations and when predators shift to area-restricted search following an encounter with prey. However, dispersion is counter-selected when predators interact by interference, especially under high predator density. When predators search for more randomly distributed prey, interference and information sharing similarly affect the relative advantage of spreading out. Under a clumped prey spatial pattern, predators benefit from shifting their search tactic to an area-restricted search following an encounter with prey. This advantage is moderated as predator density increases and when predators interact either by interference or information sharing. Under a more random prey pattern, information sharing may deteriorate the inferior search tactic even more, compared to interference or no interaction among predators. Our simulation clarifies how interactions among searching predators may affect aggregation behavior of prey, the relative success of alternative search tactics and their potential to invade established populations using some other search or spacing tactics.     

Temperature dependence of predation depends on the relative performance of predators and prey

The temperature dependence of predation rates is a key issue for understanding and predicting the responses of ecosystems to climate change. Using a simple mechanistic model, we demonstrate that differences in the relative performances of predator and prey can cause strong threshold effects in the temperature dependence of attack rates. Empirical data on the attack rate of northern pike (Esox lucius) feeding on brown trout (Salmo trutta) confirm this result. Attack rates fell sharply below a threshold temperature of +11°C, which corresponded to a shift in relative performance of pike and brown trout with respect to maximum attack and escape swimming speeds. The average attack speed of pike was an order of magnitude lower than the escape speed of brown trout at 5°C, but approximately equal at temperatures above 11°C. Thresholds in the temperature dependence of ecological rates can create tipping points in the responses of ecosystems to increasing temperatures. Thus, identifying thresholds is crucial when predicting future effects of climate warming.     

Spatial processes can determine the relationship between prey encounter rate and prey density

Theoretical models frequently assume that the rate at which a searching predator encounters prey increases linearly with prey density. In a recent experiment using great tits searching for winter moth caterpillars, the time to find the first prey item did not decline as quickly with density as the standard theory assumes. Using a spatial simulation model, we show that prey aggregation and/or spatially correlated searching behaviour by the predator can generate a range of relationships, including results that are qualitatively similar to those found in the great tit experiment. We suggest that further experiments are required to determine whether the explanation proposed here is correct, and that theoretical work is needed to determine how this behaviour is likely to influence the ecological and evolutionary dynamics of predator-prey communities.     

Non-lethal effects of predators on prey growth rates depend on prey density and nutrient additions

A number of studies show that predators can depress prey growth rates by inducing reductions in foraging activity, but the size of this non-lethal effect is quite variable. Here I investigate how prey density and resource productivity may alter the extent to which predators depress the growth rates of their prey. Theory predicts that when resources are overgrazed, an increase in predation risk will have little net effect on individual food intake because the decline in foraging effort will be offset by an increase in resource level. Thus, the non-lethal effects of predators on prey growth rates should depend upon prey density and resource productivity in a predictable manner, with the growth penalty imposed by predators being strongest when resources are undergrazed and weakest when resources are overgrazed. I tested this hypothesis by manipulating predation risk, prey density, and nutrient additions in a mesocosm experiment with the pulmonate snail Helisoma trivolvis . Refuge use by snails was 45% higher in the presence of caged crayfish than in their absence. Snail growth rates were reduced, on average, by 24% in the presence of caged crayfish. However, the magnitude of the growth penalty exacted by crayfish depended on snail density and nutrient additions. When snails were stocked at high density and nutrient additions were low, growth suppression was just 2.6%. At the other extreme, when snails were at low density and nutrient additions were high, growth suppression was 44.6%. Thus, the non-lethal effects of predators on prey growth depend on environmental context, illustrating an important link between individual traits and system-level properties.     

Adaptation of prey and predators between patches

Mathematical models are proposed to simulate migrations of prey and predators between patches. In the absence of predators, it is shown that the adaptation of prey leads to an ideal spatial distribution in the sense that the maximal capacity of each patch is achieved. With the introduction of co-adaptation of predators, it is proved that both prey and predators achieve ideal spatial distributions when the adaptations are weak. Further, it is shown that the adaptation of prey and predators increases the survival probability of predators from the extinction in both patches to the persistence in one patch. It is also demonstrated that there exists a pattern that prey and predators cooperate well through adaptations such that predators are permanent in every patch in the case that predators become extinct in each patch in the absence of adaptations. For strong adaptations, it is proved that the model admits periodic cycles and multiple stability transitions.     

The interplay between multiple predators and prey colour divergence

Evolutionary divergence in the coloration of toxic prey is expected when geographic variation in predator composition and behavior favours shifts in prey conspicuousness. A fundamental prediction of predator‐driven colour divergence is that the local coloration should experience lower predation risk than novel prey phenotypes. The dorsal coloration of the granular poison frog varies gradually from populations of conspicuous bright red frogs to populations of dull green and relatively cryptic frogs. We conducted experiments with clay models in four populations to examine the geographic patterns of taxon‐specific predation. Birds avoided the local phenotype while lizards consistently selected for decreased conspicuousness and crab predation did not depend on frog coloration. Importantly, birds and lizards favoured low conspicuousness in populations where relatively cryptic green morphs have evolved. This study provides evidence for the interplay among distinct selective pressures, from multiple‐predator taxa, acting on the divergence in protective coloration of prey species. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 113 , 580–589.     

Facilitation of fisheries by natural predators depends on life history of shared prey

Predators commonly share prey with human exploiters, intuitively suggesting that there is an inherent human–predator conflict through competition for prey. Here we studied the effects of fishing and predation mortality on biomass distributions and yields of shared prey using a size‐structured model of competing populations, describing the life histories of Baltic Sea sprat and herring. Whereas both species responded in a similar fashion to increased fishing mortality, with decreasing juvenile and adult biomasses, we found that responses to predation mortality differed between species. Sprat only display weak compensatory responses with increasing predation mortality, while over a substantial range of mortalities there was a strong increase in adult (and total) herring biomass, i.e. overcompensation. The observed biomass overcompensation results from relaxed intraspecific competition as predation mortality increased, allowing for faster individual growth rates that in turn lead to a change in population composition (juvenile:adult biomass ratio). Our results suggest that the potential for biomass overcompensation is higher for species exhibiting substantial growth after maturation. Differences in size‐selectivity of predators and fishing mortality resulted in a positive effect of predation mortality on fisheries yields, which can be explained by an overcompensatory response in adult herring biomass. Thus, somewhat counter intuitive, our results suggest that fishermen, depending on prey life history, may actually benefit from allowing for a higher abundance of predators, despite competing for shared prey.     

The effects of hydrological disturbance on the densities of macroinvertebrate predators and their prey in a coastal stream

1. Environmental Stress Models (ESMs) predict that abiotic disturbance or harshness will differentially affect predators and prey. Consumer Stress Models (CSMs) predict that consumers will be relatively more inhibited by disturbance than prey, and therefore predator impacts will be reduced. Conversely, Prey Stress Models (PSMs) predict that prey will be more adversely affected and consequently predator impacts will increase in disturbed habitats. This study compared the relative tolerances of lotic invertebrate predators and their prey to hydrological disturbance in an Australian coastal stream to test the initial predictions of ESMs.
2. Macroinvertebrates were sampled with a suction sampler at monthly intervals and immediately following four high flow events at five sites on the Cumberland River, in south-west Victoria, Australia. Various statistical procedures were used to compare the relative resistance and resilience of predatory and prey taxa to each high flow event.
3. The relative resistances of seven predator and nine prey taxa to four floods over a 12-month period were highly variable between floods and between runs within the same flood. Prey taxa appeared to be more resilient than predators to the largest flood event, but there were no differences in the resilience of predators and prey following smaller floods. If disturbance tolerance is determined by resistance and resilience, then there was no consistent pattern of differential tolerance to floods among invertebrate predators and prey in this system.
4. The variability in the relative tolerances of taxa to different disturbance events makes general predictions about the effects of disturbance on the community-wide impact of predation extremely difficult.     

Behavior of fish predators and their prey: habitat choice between open water and dense vegetation

Synopsis Behavior of largemouth bass, Micropterus salmoides, and northern pike, Esox lucius, foraging on fathead minnows, Pimephales promelas, or bluegills, Lepomis macrochirus, was quantified in pools with 50% cover (half the pool had artificial stems at a density of 1000 stems m⁻²). Both predators spent most of their time in the vegetation. Largemouth bass searched for bluegills and ambushed minnows, whereas the relatively immobile northern pike ambushed all prey. Minnows were closer to predators and were captured more frequently than bluegills. Even when minnows dispersed, they moved continually and eventually wandered within striking distance of a predator. Bluegills dispersed in the cover with predators. Bass captured the few bluegills that strayed into the open and pike captured those that approached too closely in the cover. The ability of predators to capture prey while residing in habitats containing patches of dense cover may explain their residence in areas often considered to be poor ones for foraging. The unit is sponsored jointly by the United States Fish and Wildlife Service, Ohio Department of NaturalResources, The Ohio State University, and the Wildlife Management Institute     

Response of predators to prey abundance: separating the effects of prey density and patch size

We tested the relative and combined effects of prey density and patch size on the functional response (number of attacks per unit time and duration of attacks) of a predatory reef fish (Cheilodactylus nigripes (Richardson)) to their invertebrate prey. Fish attacked prey at a greater rate and for longer time in large than small patches of prey, but large patches had naturally greater densities of prey. We isolated the effects of patch size and prey density by reducing the density of prey in larger patches to equal that of small patches; thereby controlling for prey density. We found that the intensity at which fish attacked prey (combination of attack rate and duration) was primarily a response to prey density rather than the size of patch they occupied. However, there was evidence that fish spent more time foraging in larger than smaller patches independent of prey density; presumably because of the greater total number of prey available. These experimental observations suggest that fish can distinguish between different notions of prey abundance in ways that enhance their rate of consumption. Although fish may feed in a density dependent manner, a critical issue is whether their rate of consumption outstrips the rate of increase in prey abundance to cause density dependent mortality of prey.     

Understanding predator densities for successful co‐existence of alien predators and threatened prey

The high failure rate of threatened species translocations has prompted many managers to fence areas to protect wildlife from introduced predators. However, conservation fencing is expensive, restrictive and exacerbates prey naïveté reducing the chance of future co‐existence between native prey and introduced predators. Here, we ask whether two globally threatened mammal species protected in fenced reserves, with a history of predation‐driven decline and reintroduction failure, could co‐exist with introduced predators. We defined co‐existence as population persistence for at least 3 years and successful recruitment. We manipulated the density of feral cats within a large fenced paddock and measured the impact on abundance and reproduction of 353 reintroduced burrowing bettongs and 47 greater bilbies over 3 years. We increased cat densities from 0.038 to 0.46 per square km and both threatened species survived, reproduced and increased their population size. However, a previous reintroduction trial of 66 bettongs into the same paddock found one red fox (Vulpes vulpes), at a density of 0.027 per square km, drove the bettong population extinct within 12 months. Our results show that different predator species vary in their impact and that despite a history of reintroduction failure, threatened mammal species can co‐exist with low densities of feral cats. There may be a threshold density below which it is possible to maintain unfenced populations of reintroduced marsupials. Understanding the numerical relationships between population densities of introduced predators and threatened species is urgently needed if these species are to be re‐established at landscape scales. Such knowledge will enable a priori assessment of the risk of reintroduction failure thereby increasing the likelihood of reintroduction success and reducing the financial and ethical cost of failed translocations.     

Habitat-choice interactions between pike predators and perch prey depend on water transparency

A mesocosm experiment indicated that water transparency influenced antipredator behaviour in young-of-the-year perch Perca fluviatilis , which partly contradicts another study by showing that high transparency decreases rather than increases perch antipredatory use of vegetated habitats when predators are also free to choose habitat. The present study emphasizes the importance of simultaneously considering both prey and predator habitat-choice behaviours when evaluating predator–prey interactions in relation to water visibility     

The effect of habitat structure on prey mortality depends on predator and prey microhabitat use

Structurally complex habitats provide cover and may hinder the movement of animals. In predator–prey relationships, habitat structure can decrease predation risk when it provides refuges for prey or hinders foraging activity of predators. However, it may also provide shelter, supporting structures and perches for sit-and-wait predators and hence increase their predation rates. We tested the effect of habitat structure on prey mortality in aquatic invertebrates in short-term laboratory predation trials that differed in the presence or absence of artificial vegetation. The effect of habitat structure on prey mortality was context dependent as it changed with predator and prey microhabitat use. Specifically, we observed an ‘anti-refuge’ effect of added vegetation: phytophilous predators that perched on the plants imposed higher predation pressure on planktonic prey, while mortality of benthic prey decreased. Predation by benthic and planktonic predators on either type of prey remained unaffected by the presence of vegetation. Our results show that the effects of habitat structure on predator–prey interactions are more complex than simply providing prey refuges or cover for predators. Such context-specific effects of habitat complexity may alter the coupling of different parts of the ecosystem, such as pelagic and benthic habitats, and ultimately affect food web stability through cascading effects on individual life histories and trophic link strengths.     

Temperate trees and shrubs as global invaders: the relationship between invasiveness and native distribution depends on biological traits

Many woody plants have been recently recognized as major invasive species with serious impacts on species diversity and functioning of invaded ecosystems. We evaluated the global invasion success of temperate trees and shrubs with native ranges in central Europe and explored the role of their native distribution and that of biological traits in determining whether they have become invasive in other parts of the world. Of the 94 species forming the source-area species pool, 27 % are known to be invasive in at least one region of the world. Generalized linear models on phylogenetically non-informed analyses revealed that tall woody plants flowering early in the season, and occupying many grid cells in the native range are significantly more likely to become successful invaders than species not possessing these traits. However, other traits can partly reduce the disadvantages resulting from low regional frequencies in the native range and consequent lower probability of them being introduced elsewhere. Species that do not depend for reproduction on another individual, those pollinated by wind and dispersed by animals are likely to become invasive even if they do not have extensive native distributions. However, of these traits only the regional frequency in the native range remained significant when phylogenetic relationships were taken into account. This indicates that the effect of the regional frequency is generic, valid across all woody species, and fine-tuned by advantageous biological traits inherited from common ancestors, shared by groups of phylogenetically-related species. Traits that only appeared significant in the phylogenetically non-informed analyses should be considered as specific for shrubs and trees of central Europe. Mode of reproduction was only significant in the phylogenetically-informed analysis, indicating that reproduction exclusively by seed favours invasiveness of woody species. From the management perspective, however, the predictive value of some traits is not diminished by them being phylogenetically constrained since we are not interested whether the behaviour of particular species is a result of evolutionary history but how we can treat specific cases of invasion.     

The non-consumptive effects of predators and personality on prey growth and mortality

Individual organisms vary in personality, and the ecological consequences of that variation can affect the strength of predator–prey interactions. Prey with bolder tendencies can mitigate the strength of species interactions by altering growth and initiating ontogenetic niche shifts (ONS). While the link between personality and growth has been established, recent research has highlighted the important interplay between ONS and predator cues in community ecology. The objective of this study was to evaluate the effects of prey personality and predator cues on prey growth and ONS. We predicted growth–mortality trade-offs among personalities with higher survival, larger size, and accelerated ONS for bold individuals in comparison with shy individuals. To evaluate this objective, we conducted behavioral assays and a mesocosm experiment to test how southern leopard frog (Rana sphenocephala) tadpole personality and predatory fish (bluegill, Lepomis macrochirus) cues affects tadpole growth and metamorphosis. On average, bold tadpoles had higher mortality across all treatments in comparison with shy tadpoles. The effects of fish cues were dependent on tadpole personality with shy tadpoles metamorphosing significantly later than bold tadpoles. Bold tadpoles were larger than shy tadpoles at metamorphosis; however, that pattern reversed with fish cues as shy individuals metamorphosed larger than bold individuals. Our results suggest personality may be useful for predicting growth and life history for some prey species with predators. Specifically, the threat of predation can interact with personality to incur a benefit (earlier ONS) while also incurring a cost (size at metamorphosis). Hence by incorporating predator cues with personality, ecologists will be able to elucidate growth–mortality trade-offs mediated by personality.     

Linear scaling between microbial predator and prey densities in the global ocean

It has been proposed that microbial predator and prey densities are related through sublinear power laws. We revisited previously published biomass and abundance data and fitted Power-law Biomass Scaling Relationships (PBSRs) between marine microzooplankton predators (Z) and phytoplankton prey (P), and marine viral predators (V) and bacterial prey (B). We analysed them assuming an error structure given by Type II regression models which, in contrast to the conventional Type I regression model, accounts for errors in both the independent and the dependent variables. We found that the data support linear relationships, in contrast to the sublinear relationships reported by previous authors. The scaling exponent yields an expected value of 1 with some spread in different datasets that was well-described with a Gaussian distribution. Our results suggest that the ratios Z/P, and V/B are on average invariant, in contrast to the hypothesis that they systematically decrease with increasing P and B, respectively, as previously thought.     

Control of aphids on wheat by generalist predators: effects of predator density and the presence of alternative prey

There is evidence for both positive and negative effects of generalist predators on pest populations and the various reasons for these contrasting observations are under debate. We studied the influence of a generalist predator, Pardosa lugubris (Walckenaer) (Araneae: Lycosidae), on an aphid pest species, Rhopalosiphum padi (L.) (Hemiptera: Aphididae; low food quality for the spider), and its host plant wheat, Triticum spec. (Poaceae). We focused on the role of spider density and the availability of alternative prey, Drosophila melanogaster Meigen (Diptera: Drosophilidae; high food quality). The presence of spiders significantly affected plant performance and aphid biomass. Alternative prey and spider density strongly interacted in affecting aphids and plants. High spider density significantly improved plant performance but also at low spider density plants benefited from spiders especially in the presence of alternative prey. The results suggest that generalist arthropod predators may successfully reduce plant damage by herbivores. However, their ability to control prey populations varies with predator nutrition, the control of low-quality prey being enhanced if alternative higher-quality prey is available.