Predator dispersal determines the effect of connectivity on prey diversity

Linking local communities to a metacommunity can positively affect diversity by enabling immigration of dispersal-limited species and maintenance of sink populations. However, connectivity can also negatively affect diversity by allowing the spread of strong competitors or predators. In a microcosm experiment with five ciliate species as prey and a copepod as an efficient generalist predator, we analysed the effect of connectivity on prey species richness in metacommunities that were either unconnected, connected for the prey, or connected for both prey and predator. Presence and absence of predator dispersal was cross-classified with low and high connectivity. The effect of connectivity on local and regional richness strongly depended on whether corridors were open for the predator. Local richness was initially positively affected by connectivity through rescue of species from stochastic extinctions. With predator dispersal, however, this positive effect soon turned negative as the predator spread over the metacommunity. Regional richness was unaffected by connectivity when local communities were connected only for the prey, while predator dispersal resulted in a pronounced decrease of regional richness. The level of connectivity influenced the speed of richness decline, with regional species extinctions being delayed for one week in weakly connected metacommunities. While connectivity enabled rescue of prey species from stochastic extinctions, deterministic extinctions due to predation were not overcome through reimmigration from predator-free refuges. Prey reimmigrating into these sink habitats appeared to be directly converted into increased predator abundance. Connectivity thus had a positive effect on the predator, even when the predator was not dispersing itself. Our study illustrates that dispersal of a species with strong negative effects on other community members shapes the dispersal-diversity relationship. When connections enable the spread of a generalist predator, positive effects of connectivity on prey species richness are outweighed by regional extinctions through predation.     

Patterns, mechanisms and spatial scale of aggregation in generalist and specialist predatory mites (Acari: Phytoseiidae)

Prey species often distribute themselves patchily in their habitats. In response to this spatial variation in prey density, some predator species aggregate in patches of higher prey density. This paper reviews a series of laboratory experiments to demonstrate the patterns of responses by phytoseiid predators (Phytoseiulus persimilis, Typhlodromus occidentalis and Amblyseius andersoni) to spatial variation in the density of their spider mite prey (Tetranychus urticae) and reveal the behavioural mechanisms underlying the observed patterns. In addition, patterns of aggregation were examined at a variety of spatial scales on plants in greenhouses. The patterns, mechanisms and spatial scale of aggregation in three predatory species are discussed in relation to their varying degrees of polyphagy. The results show that a specialist predator species (1) aggregates more strongly than generalist predators, (2) does so not because it finds prey patches of high density more easily but because it remains in these patches longer than generalist predators and (3) tends to aggregate more often at lower levels of spatial scale than generalist predators. It is suggested that these conclusions, based mainly on laboratory studies of a small sample of species, should be tested in the future on a wider selection of specialist and generalist species at different scales in the field. This revised version was published online in November 2006 with corrections to the Cover Date.     

Numerical and dietary responses of a predator community in a temperate zone of Europe

The generalist predation hypothesis predicts that the functional responses of generalist predator species should be quicker than those of specialist predators and have a regulating effect on vole populations. New interpretations of their role in temperate ecosystems have, however, reactivated a debate suggesting generalist predators may have a destabilizing effect under certain conditions (e.g. landscape homogeneity, low prey diversity, temporary dominance of 1 prey species associated with a high degree of dietary specialization). We studied a rich predator community dominated by generalist carnivores ( Martes spp., Vulpes vulpes, Felis catus ) over a 6 yr period in farmland and woodland in France. The most frequent prey were small rodents (mostly Microtus arvalis , a grassland species, and Apodemus spp., a woodland species). Alternative prey were diverse and dominated by lagomorphs ( Oryctolagus cuniculus, Lepus europeus ). We detected a numerical response among specialist carnivores but not among generalist predators. The dietary responses of generalist predators were fairly complex and most often dependent on variation in density of at least 1 prey species. These results support the generalist predation hypothesis. We document a switch to alternative prey, an increase of diet diversity, and a decrease of diet overlap between small and medium-sized generalists during the low density phase of M. arvalis . In this ecosystem, the high density phases of small mammal species are synchronous and cause a temporary specializing of several generalist predator species. This rapid functional response may indicate the predominant role of generalists in low amplitude population cycles of voles observed in some temperate areas.     

Predator diet breadth influences the relative importance of bottom-up and top-down control of prey biomass and diversity

We investigated the effects of predator diet breadth on the relative importance of bottom-up and top-down control of prey assemblages, using microbial food webs containing bacteria, bacterivorous protists and rotifers, and two different top predators. The experiment used a factorial design that independently manipulated productivity and the presence or absence of two top predators with different diet breadths. Predators included a "specialist" predatory ciliate Euplotes aediculatus, which was restricted to feeding on small prey, and a "generalist" predatory ciliate Stentor coeruleus, which could feed on the entire range of prey sizes. Both total prey biomass and prey diversity increased with productivity in the predator-free control and specialist predator treatments, a pattern consistent with bottom-up control, but both remained unchanged by productivity in the generalist predator treatment, a pattern consistent with top-down control. Linear food chain models adequately described responses in the generalist predator treatment, whereas food web models incorporating edible and inedible prey (which can coexist in the absence of predators) adequately described responses in the specialist predator treatment. These results suggest that predator diet breadth can play an important role in modulating the relative strength of bottom-up and top-down forces in ecological communities.     

Quantifying predator dependence in the functional response of generalist predators

A long‐standing debate concerns how functional responses are best described. Theory suggests that ratio dependence is consistent with many food web patterns left unexplained by the simplest prey‐dependent models. However, for logistical reasons, ratio dependence and predator dependence more generally have seen infrequent empirical evaluation and then only so in specialist predators, which are rare in nature. Here we develop an approach to simultaneously estimate the prey‐specific attack rates and predator‐specific interference (facilitation) rates of predators interacting with arbitrary numbers of prey and predator species in the field. We apply the approach to surveys and experiments involving two intertidal whelks and their full suite of potential prey. Our study provides strong evidence for predator dependence that is poorly described by the ratio dependent model over manipulated and natural ranges of species abundances. It also indicates how, for generalist predators, even the qualitative nature of predator dependence can be prey‐specific.     

Effects of competition, predation, and dispersal on species richness at local and regional scales

This study explores the consequences of predator-mediated coexistence among competitors for patterns of incidence and diversity at local and regional scales. We develop a model that draws on elements of metapopulation models of competitors and food chains by allowing competitors to coexist locally in the presence of predators but not in their absence. The model predicts that predators promote regional coexistence by greatly expanding the range of conditions under which two competitors persist at equilibrium. Predators could have positive or negative effects on mean local diversity within the region depending on their dispersal rates, those of the prey, and their effects on prey extinction rates. The presence of predators increased the abundance of inferior competitors, thereby expanding the conditions for positive relationships between local and regional diversity. The model also predicted positive correlations between local diversity of predators and prey. These predictions were supported by patterns of phytoplankton, zooplankton, and fish species richness among lakes. The model may help to resolve the apparent contrast between linear patterns of local and regional richness and experimental evidence for strong invasion resistance and rapid dispersal in zooplankton.     

Predators temper the relative importance of stochastic processes in the assembly of prey metacommunities

Communities assemble through a combination of stochastic processes, which can make environmentally similar communities divergent (high β-diversity), and deterministic processes, which can make environmentally similar communities convergent (low β-diversity). Top predators can influence both stochasticity (e.g. colonization and extinction events) and determinism (e.g. size of the realized species pool), in community assembly, and thus their net effect is unknown. We investigated how predatory fish influenced the scaling of prey diversity in ponds at local and regional spatial scales. While fish reduced both local and regional richness, their effects were markedly more intense at the regional scale. Underlying this result was that the presence of fish made localities within metacommunities more similar in their community composition (lower β-diversity), suggesting that fish enhance the deterministic, relative to the stochastic, components of community assembly. Thus, the presence of predators can alter fundamental mechanisms of community assembly and the scaling of diversity within metacommunities.     

Interactive effects of productivity and predation on zooplankton diversity

Recent studies suggest the necessity of understanding the interactive effects of predation and productivity on species coexistence and prey diversity. Models predict that coexistence of prey species with different competitive abilities can be achieved if inferior resource competitors are less susceptible to predation and if productivity and/or predation pressure are at intermediate levels. Hence, predator effects on prey diversity are predicted to be highly context dependent: enhancing diversity from low to intermediate levels of productivity or predation and reducing diversity of prey at high levels of productivity or predation. While several studies have examined the interactive effects of herbivory and productivity on primary producer diversity, experimental studies of such effects in predator‐prey systems are rare. We tested these predictions using an aquatic field mesocosm experiment in which initial density of the zooplankton predator Notonecta undulata and productivity were manipulated to test their interactive effects on diversity of seven zooplankton, cladoceran species that were common in surrounding ponds. Two productivity levels were imposed via phosphorus enrichment at levels comparable to low and intermediate levels found within neighboring natural ponds. We used open systems to allow for natural dispersal and behaviorally‐mediated numerical responses by the flight‐capable predator. Effects of predators on zooplankton diversity depended on productivity level. At low and high productivity, prey species richness declined while at high productivity it showed a unimodal relationship with increasing the predator density. Effects of treatments were weaker when using Pielou's evenness index or the inverse Simpson index as measures of prey diversity. Our findings are generally consistent with model predictions in which predators can facilitate prey coexistence and diversity at intermediate levels of productivity and predation intensity. Our work also shows that the functional form of the relationship between prey diversity and predation intensity can be complex and highly dependent on environmental context.     

Intraguild interactions among generalist predator functional groups drive impact on herbivore and decomposer prey

Different functional groups of generalist predators may complement each other in controlling prey populations; but intraguild interactions, common among generalist predators, may also reduce the strength of top–down control. In natural communities greater alterations to ecosystem function are expected if a whole functional group declines in abundance or is lost. Therefore studying functional group diversity is important for predicting effects of predator loss. We studied the top–down impact of web‐building spiders, hunting spiders and ants, which are highly abundant generalist predators in most terrestrial ecosystems, on prey from the herbivore and decomposer system of a grassland food web. The density of the three predator groups was manipulated by continuous removal in a three‐factorial designed field experiment, which was carried out for two years. We found no positive effect of increasing predator functional group richness on prey control. However there was evidence for strong composition effects between the functional groups. The presence of ants in predator assemblages reduced the prey suppression through mostly trait‐mediated intraguild interactions, while hunting and web‐building spiders contributed additively to prey suppression and reduced the density of herbivore and decomposer prey by 50–60%. A trophic cascade on plant biomass triggered by web‐builders and hunting spiders was diminished at levels of higher predator group diversity. In conclusion, our experiments showed that intraguild interactions strongly influence the strength of top–down control by generalist predators. Among spiders there was evidence for a positive relation between functional group richness and prey suppression but the overall outcome strongly depended on the occurrence of interference, driven by trait‐mediated indirect interactions.     

Context-dependent effects of predator removal from experimental microcosm communities

The loss of a predator from an ecological community can cause large changes in community structure and ecosystem processes, or have very little consequence for the remaining species and ecosystem. Understanding when and why the loss of a predator causes large changes in community structure and ecosystem processes is critical for understanding the functional consequences of biodiversity loss. We used experimental microbial communities to investigate how the removal of a large generalist predator affected the extinction frequency, population abundance and total biomass of its prey. We removed this predator in the presence or absence of an alternative, more specialist, predator in order to determine whether the specialist predator affected the outcome of the initial species removal. Removal of the large generalist predator altered some species' populations but many were unaffected and no secondary extinctions were observed. The specialist predator, though rare, altered the response of the prey community to the removal of the large generalist predator. In the absence of the specialist predator, the effects of the removal were only measurable at the level of individual species. However, when the specialist predator was present, the removal of the large generalist predator affected the total biomass of prey species. The results demonstrate that the effect of species loss from high trophic levels may be very context-dependent, as rare species can have disproportionately large effects in food webs.     

The influence of generalist predators in spatially extended predator–prey systems

The presence of generalist predators is known to have important ecological impacts in several fields. They have wide applicability in the field of biological control. However, their role in the spatial distribution of predator and prey populations is still not clear. In this paper, the spatial dynamics of a predator–prey system is investigated by considering two different types of generalist predators. In one case, it is considered that the predator population has an additional food source and can survive in the absence of the prey population. In the other case, the predator population is involved in intraguild predation, i.e., the source of the additional food of the predator coincides with the food source of the prey population and thus both prey and predator populations compete for the same resource. The conditions for linear stability and Turing instability are analyzed for both the cases. In the presence of generalist predators, the system shows different pattern formations and spatiotemporal chaos which has important implications for ecosystem functioning not only in terms of their predictability, but also in influencing species persistence and ecosystem stability in response to abrupt environmental changes. This study establishes the importance of the consideration of spatial dynamics while determining optimal strategies for biological control through generalist predators.     

On the role of sinigrin (mustard oil) in a tritrophic context: plant–aphid–aphidophagous hoverfly

1. Plant secondary metabolites can govern prey–predator interactions by altering the diet breadth of predators and sometimes provide an ecological refuge to prey. Brassicaceae plants and their specialist pests can be used as a model system for understanding the role of chemically mediated effects restricting the diet breadth of natural enemies, and consequently the occurrence of enemy‐free space for the specialist pest. 2. The objective of the present study was to test the performance of the generalist predator Episyrphus balteatus De Geer (Diptera: Syrphidae) fed on the specialist herbivore Brevicoryne brassicae L.(Homoptera: Aphididae), reared on two different brassica species: black mustard (Brassica nigra), a wild species with high levels of sinigrin; and canola (Brassica napus), a cultivated species without sinigrin. 3. The preference and performance of the predator and the performance of the prey were measured. Sinigrin was quantified by high‐performance liquid chromatography in both leaf samples and aphids reared on the two host plants. 4. The cabbage aphid performed better on canola than on black mustard. The performance of the predator on this aphid when reared on canola was clearly better than when reared on black mustard. Females had a higher overall preference for cabbage aphids reared on canola than on black mustard. 5. The ability of aphids reared on plants with high glucosinolate content to reduce the performance of their generalist predators indicates that the presence of B. nigra may provide enemy‐free space for the cabbage aphid from its predator, a concept that has useful application in the context of biological control for agricultural systems.     

Influence of extraguild prey density on intraguild predation by heteropteran predators: A review of the evidence and a case study

Heteropteran predators constitute an important component of predatory guilds in terrestrial and aquatic ecosystems. Most heteropteran species have generalist diets, and intraguild predation has been documented in most heteropteran families. Zoophytophagous species also frequently engage in intraguild interactions. An increase in extraguild prey density is often predicted to reduce intraguild predation between guild members by providing abundant alternate prey. However, an increase of extraguild prey density may also be associated with an increase in the density of intraguild predators, which could instead strengthen intraguild predation. Evaluating the combined effect of these potentially opposing influences on intraguild predation is difficult. Most studies have been carried out in the laboratory, using artificially simplified communities of predators and prey and employing spatial and temporal scales that may not reflect field conditions. We review experimental studies examining how extraguild prey density influences the intensity of intraguild predation and then report an observational case study examining the influence of extraguild prey density on the intensity of intraguild predation at larger spatial and temporal scales in unmanipulated cotton fields. Fields with more abundant extraguild prey (aphids, mites) were not associated with elevated densities of intraguild predators, and were strongly associated with increased survival of intraguild prey (lacewing larvae). In this system, the ability of extraguild prey to relax the intensity of intraguild predation, as previously documented in small-scale field experiments, also extends to the larger spatial and temporal scales of commercial agriculture.     

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.     

Are Infaunal Predators Important in Structuring Marine Soft-Bottom Communities?

The role of infaunal predators in structuring marine soft-bottomcommunities was evaluated according to these predators': 1)effects on prey density based on manipulative field experiments,2) feeding rates, 3) effects on prey distribution, 4) effectson species diversity, and 5) interactions with their prey. Estimatesof feeding rates indicate that many predatory taxa have thepotential to reduce the size of prey populations and suggestthat nemerteans are likely to have a larger impact on infaunalabundances than polychaetes. Infaunal predators have been demonstratedto have a significant effect on infaunal densities and to affectthe spatial and temporal distribution of their prey. The effectsof these predators on species diversity apparently depend onthe predator and the diversity of the system. These conclusionsmay not be applicable to all soft-bottom habitats or all groupsof infaunal predators because they are based on studies of veryfew taxa conducted almost exclusively in intertidal, unvegetated,mud habitats. Additional studies are needed on the effects ofpredation by infauna on infaunal population dynamics and onthe mechanisms of interactions between predator and prey. Furtherinvestigation will probably reveal that different groups ofinfaunal predators play different roles in structuring soft-bottomcommunities.     

Testing the Threat‐Sensitive Hypothesis with Predator Familiarity and Dietary Specificity

In a system with multiple predators, the threat‐sensitive predator avoidance hypothesis predicts that prey respond differently to predators relative to the risks each poses (e.g., degree of dietary specialization). Aquatic animals often rely heavily on detecting predators via chemical cues (kairomones) and respond with a suite of behaviors including detection and avoidance. However, little is known about how animals respond to kairomones of specialist versus generalist predators. In laboratory experiments, we compared behavioral responses of a poorly studied aquatic salamander, the greater siren (Siren lacertina), to cues from specialist and generalist predator snakes to evaluate threat‐sensitive responses. Sirens exhibited a novel behavior (gill‐flushing) most often in the presence of specialist predator cues. Avoidance behavior (reversing direction following cue detection) was higher in response to specialist predator and novel animal control cues and lowest in response to generalist predator cues. An intermediate response to the animal control, an unfamiliar amphibian predator, indicated that sirens respond cautiously to a novel cue. The gradient of observed responses to different snake cues indicates that sirens may be evaluating predation potential of animals based on their foraging specificity and familiarity.     

Climate change effects on behavioral and physiological ecology of predator–prey interactions: Implications for conservation biological control

Habitat management under the auspices of conservation biological control is a widely used approach to foster conditions that ensure a diversity of predator species can persist spatially and temporally within agricultural landscapes in order to control their prey (pest) species. However, an emerging new factor, global climate change, has the potential to disrupt existing conservation biological control programs. Climate change may alter abiotic conditions such as temperature, precipitation, humidity and wind that in turn could alter the life-cycle timing of predator and prey species and the behavioral nature and strength of their interactions. Anticipating how climate change will affect predator and prey communities represents an important research challenge. We present a conceptual framework—the habitat domain concept—that is useful for understanding contingencies in the nature of predator diversity effects on prey based on predator and prey spatial movement in their habitat. We illustrate how this framework can be used to forecast whether biological control by predators will become more effective or become disrupted due to changing climate. We discuss how changes in predator–prey interactions are contingent on the tolerances of predators and prey species to changing abiotic conditions as determined by the degree of local adaptation and phenotypic plasticity exhibited by species populations. We conclude by discussing research approaches that are needed to help adjust conservation biological control management to deal with a climate future.     

Functionally different predators break down antipredator defenses of spider mites

Prey that lives with functionally different predators may experience enhanced mortality risk, because of conflicts between the specific defenses against their predators. Because natural communities usually contain combinations of prey and functionally different predators, examining risk enhancement with multiple predators may help to understand prey population dynamics. It is also important in an applied context: risk enhancement with multiple biological control agents could lead to successful suppression of pests. We examined whether risk enhancement occurs in the spider mite Tetranychus kanzawai Kishida (Acari: Tetranychidae) when exposed to two predator species: a generalist ant, Pristomyrmex punctatus Mayr (Hymenoptera: Formicidae), and a specialist predatory mite, Neoseiulus womersleyi Schicha (Acari: Phytoseiidae). We replicated microcosms that consisted of spider mites, ants, and predatory mites. Spider mites avoided generalist ants by staying inside their webs on leaf surfaces. In contrast, spider mites avoided specialist predatory mites that intruded into their webs by exiting the web, which obviously conflicts with the defense against ants. In the presence of both predators, enhanced mortality of spider mites was observed. A conflict occurred between the spider mites’ defenses: they seemed to move out of their webs and be preyed upon by ants. This is the first study to suggest that risk enhancement occurs in web‐spinning spider mites that are exposed to both generalist and specialist predator species, and to provide evidence that ants can have remarkable synergistic effects on the biological control of spider mites using specialist predatory mites.     

Predation across spatial scales in heterogeneous environments

A hierarchy of scales is introduced to the spatially heterogeneous Lotka-Volterra predator-prey diffusion model, and its effects on the model's spatial and temporal behavior are studied. When predators move on a large scale relative to prey, local coupling of the predator-prey interaction is replaced by global coupling. Prey with low dispersal ability become narrowly confined to the most productive habitats, strongly amplifying the underlying spatial pattern of the environment. As prey diffusion rate increases, the prey distribution spreads out and predator abundance declines. The model retains neutrally stable Lotka-Volterra temporal dynamics: different scales of predator and prey dispersal do not stabilize the interaction. The model predicts that, for prey populations that are limited by widely ranging predators, species with low dispersal ability should be restricted to discrete high density patches, and those with greater mobility should be more uniformly distributed at lower density.     

Bistability and limit cycles in generalist predator–prey dynamics

Since generalist predators feed on a variety of prey species they tend to persist in an ecosystem even if one particular prey species is absent. Predation by generalist predators is typically characterized by a sigmoidal functional response, so that predation pressure for a given prey species is small when the density of that prey is low. Many mathematical models have included a sigmoidal functional response into predator–prey equations and found the dynamics to be more stable than for a Holling type II functional response. However, almost none of these models considers alternative food sources for the generalist predator. In particular, in these models, the generalist predator goes extinct in the absence of the one focal prey. We model the dynamics of a generalist predator with a sigmoidal functional response on one dynamic prey and fixed alternative food source. We find that the system can exhibit up to six steady states, bistability, limit cycles and several global bifurcations.