Predators alter the scaling of diversity in prey metacommunities

Although predator effects on the number of locally coexisting species are well understood, there are few formal predictions of how these local predator effects influence patterns of prey diversity at larger spatial scales. Building on the theory of island biogeography, we develop a simple model that describes how predators can alter the scaling of diversity in prey metacommunities and compares the effects of generalist and specialist predators on regional prey diversity. Generalist predators, which consume prey randomly with respect to species identity, are predicted to reduce α‐diversity and increase β‐diversity thereby maintaining regional diversity (γ‐diversity). Alternatively, specialist predators, which filter out prey species intolerant of predators, are predicted to reduce bothα‐diversity andβ‐diversity by causing the same prey species to be extirpated in each locality, resulting in regional prey species extinctions and lower γ‐diversity. These distinct effects of generalist and specialist predators on prey diversity at different spatial scales are uniquely shaped by the extent of predation within those metacommunities. Overall, our model results make general predictions for how different types of predators can differentially affect prey diversity across spatial scales, allowing a more complete understanding of the possible implications of predator eradications or introductions for biodiversity.     

Evolution of dispersal in a predator-prey metacommunity

Dispersal is crucial to allowing species inhabiting patchy or spatially subdivided habitats to persist globally despite the possibility of frequent local extinctions. Theoretical studies have repeatedly demonstrated that species that exhibit a regional metapopulation structure and are subject to increasing rates of local patch extinctions should experience strong selective pressures to disperse more rapidly despite the costs such increased dispersal would entail in terms of decreased local fitness. We extend these studies to consider how extinctions arising from predator-prey interactions affect the evolution of dispersal for species inhabiting a metacommunity. Specifically, we investigate how increasing a strong extinction-prone interaction between a predator and prey within local patches affects the evolution of each species' dispersal. We found that for the predator, as expected, evolutionarily stable strategy (ESS) dispersal rates increased monotonically in response to increasing local extinctions induced by strong predator top-down effects. Unexpectedly for the prey, however, ESS dispersal rates displayed a nonmonotonic response to increasing predator-induced extinction rates-actually decreasing for a significant range of values. These counterintuitive results arise from how extinctions resulting from trophic interactions play out at different spatial scales: interactions that increase extinction rates of both species locally can, at the same time, decrease the frequency of interaction between the prey and predator at the metacommunity scale.     

Dispersal stochasticity mediates species richness in source–sink metacommunities

Although it is well‐known that dispersal of organisms within a metacommunity will influence patterns of coexistence and richness, theoretical and experimental studies generally assume that dispersal rates are constant through time. However, dispersal is often a highly variable process that can vary seasonally and/or when stochastic events (e.g. wind storms, droughts, floods) occur. Using a well‐known source–sink metacommunity model, we present novel predictions for local and regional species richness when stochasticity in dispersal is expressly considered. We demonstrate that dispersal stochasticity alters some of the predictions obtained with constant dispersal; the peak of the predicted hump‐shaped relationship between dispersal and local species richness is diminished and shifted towards higher values of dispersal. Dispersal stochasticity increases extinction probabilities of inferior competitor species particularly in metacommunities subjected to severe isolation events (i.e. decreases of dispersal) or homogenization events (i.e. sudden increases of dispersal). Our results emphasize how incorporating dispersal stochasticity into theoretical predictions will broaden our understanding of metacommunities dynamics and their responses to natural and human‐related disturbances.     

The effects of habitat fragmentation on persistence of source-sink metapopulations in systems with predators and prey or apparent competitors.

We consider systems with one predator and one prey, or a common predator and two prey species (apparent competitors) in source and sink habitats. In both models, the predator species is vulnerable to extinction, if productivity in the source is insufficient to rescue demographically deficient sink populations. Conversely, in the model with two prey species, if the source is too rich, one of the prey species may be driven extinct by apparent competition, since the predator can maintain a large population because of the alternative prey. Increasing the rate of predator movement from the source population has opposite effects on prey and predator persistence. High emigration rate exposes the predator population to danger of extinction, reducing the number of individuals that breed and produce offspring in the source habitat. This may promote coexistence of prey by relaxing predation pressure and apparent competition between the two prey species. The number of sinks and spatial arrangement of patches, or connectivity between patches, also influence persistence of the species. More sinks favor the prey and fewer sinks are advantageous to the predator. A linear pattern with the source at one end is profitable for the predator, and a centrifugal pattern in which the source is surrounded by sinks is advantageous to the prey. When the dispersal rate is low, effects of the spatial structure may exceed those of the number of sinks. In brief, productivity in patches and patterns of connectivity between patches differentially influence persistence of populations in different trophic levels.     

Multi‐scale responses to warming in an experimental insect metacommunity

In metacommunities, diversity is the product of species interactions at the local scale and dispersal between habitat patches at the regional scale. Although warming can alter both species interactions and dispersal, the combined effects of warming on these two processes remains uncertain. To determine the independent and interactive effects of warming‐induced changes to local species interactions and dispersal, we constructed experimental metacommunities consisting of enclosed milkweed patches seeded with five herbivorous milkweed specialist insect species. We treated metacommunities with two levels of warming (unwarmed and warmed) and three levels of connectivity (isolated, low connectivity, high connectivity). Based on metabolic theory, we predicted that if plant resources were limited, warming would accelerate resource drawdown, causing local insect declines and increasing both insect dispersal and the importance of connectivity to neighboring patches for insect persistence. Conversely, given abundant resources, warming could have positive local effects on insects, and the risk of traversing a corridor to reach a neighboring patch could outweigh the benefits of additional resources. We found support for the latter scenario. Neither resource drawdown nor the weak insect‐insect associations in our system were affected by warming, and most insect species did better locally in warmed conditions and had dispersal responses that were unchanged or indirectly affected by warming. Dispersal across the matrix posed a species‐specific risk that led to declines in two species in connected metacommunities. Combined, this scaled up to cause an interactive effect of warming and connectivity on diversity, with unwarmed metacommunities with low connectivity incurring the most rapid declines in diversity. Overall, this study demonstrates the importance of integrating the complex outcomes of species interactions and spatial structure in understanding community response to climate change.     

Intra- and interspecific density-dependent dispersal in an aquatic prey-predator system

1. Dispersal intensity is a key process for the persistence of prey-predator metacommunities. Consequently, knowledge of the ecological mechanisms of dispersal is fundamental to understanding the dynamics of these communities. Dispersal is often considered to occur at a constant per capita rate; however, some experiments demonstrated that dispersal may be a function of local species density. 2. Here we use aquatic experimental microcosms under controlled conditions to explore intra- and interspecific density-dependent dispersal in two protists, a prey Tetrahymena pyriformis and its predator Dileptus sp. 3. We observed intraspecific density-dependent dispersal for the prey and interspecific density-dependent dispersal for both the prey and the predator. Decreased prey density lead to an increase in predator dispersal, while prey dispersal increased with predator density. 4. Additional experiments suggest that the prey is able to detect its predator through chemical cues and to modify its dispersal behaviour accordingly. 5. Density-dependent dispersal suggests that regional processes depend on local community dynamics. We discuss the potential consequences of density-dependent dispersal on metacommunity dynamics and stability.     

A cross‐system meta‐analysis reveals coupled predation effects on prey biomass and diversity

Predator diversity and abundance are under strong human pressure in all types of ecosystems. Whereas predator potentially control standing biomass and species interactions in food webs, their effects on prey biomass and especially prey biodiversity have not yet been systematically quantified. Here, we test the effects of predation in a cross‐system meta‐analysis of prey diversity and biomass responses to local manipulation of predator presence. We found 291 predator removal experiments from 87 studies assessing both diversity and biomass responses. Across ecosystem types, predator presence significantly decreased both biomass and diversity of prey across ecosystems. Predation effects were highly similar between ecosystem types, whereas previous studies had shown that herbivory or decomposition effects differed fundamentally between terrestrial and aquatic systems based on different stoichiometry of plant material. Such stoichiometric differences between systems are unlikely for carnivorous predators, where effect sizes on species richness strongly correlated to effect sizes on biomass. However, the negative predation effect on prey biomass was ameliorated significantly with increasing prey richness and increasing species richness of the manipulated predator assemblage. Moreover, with increasing richness of the predator assemblage present, the overall negative effects of predation on prey richness switched to positive effects. Our meta‐analysis revealed strong general relationships between predator diversity, prey diversity and the interaction strength between trophic levels in terms of biomass. This study indicates that anthropogenic changes in predator abundance and diversity will potentially have strong effects on trophic interactions across ecosystems. Synthesis The past centuries we have experienced a dramatic loss of top–predator abundance and diversity in most types of ecosystems. To understand the direct consequences of predator loss on a global scale, we quantitatively summarized experiments testing predation effects on prey communities in a cross‐system meta‐analysis. Across ecosystem types, predator presence significantly decreased both biomass and diversity of prey, and predation effects were highly similar. However, with increasing predator richness, the overall negative effects of predation on prey richness switched to positive ones. Anthropogenic changes in predator communities will potentially have strong effects on prey diversity, biomass, and trophic interactions across ecosystems.     

Predator-prey interactions in a nonequilibrium context: the metapopulation approach to modeling "hide-and-seek" dynamics in a spatially explicit tri-trophic system

Predators and prey are usually heterogeneously distributed in space so that the ability of the predators to respond to the distribution of their prey may have a profound influence on the stability and persistence of a predator‐prey system. A special type of dynamics is “hide‐and‐seek” characterized by a high turnover rate of local populations of prey and predators, because once the predators have found a patch of prey they quickly overexploit it, whereupon the starving predators either should move to better places or die. Continued persistence of prey and predators thus hinges on a long‐term balance between local extinctions and founding of new subpopulations. The colonization rate depends on the rate of emigration from occupied patches and the likelihood of successfully arriving at a suitable new patch, while extinction rate depends on the local population dynamics. Since extinctions and colonizations are both discrete probabilistic events, these phenomena are most adequately modeled by means of a stochastic model. In order to demonstrate the qualitative differences between a deterministic and stochastic approach to population dynamics, a spatially explicit tritrophic predator‐prey model is developed in a deterministic and a stochastic version. The model is parameterized using data for the two‐spotted spider mite (Tetranychus urticae) and the phytoseiid mite predator Phytoseiulus persimilis inhabiting greenhouse cucumbers.
Simulations show that the deterministic and stochastic approaches yield different results. The deterministic version predicts that the populations will exhibit violent fluctuations, implying that the system is fundamentally unstable. In contrast, the stochastic version predicts that the two species will be able to coexist in spite of frequent local extinctions of both species, provided the system consists of a sufficiently large number of local populations. This finding is in agreement with experimental results. It is therefore concluded that demographic stochasticity in combination with dispersal is capable of producing and maintaining sufficient asynchrony between local populations to ensure long‐term regional (metapopulation) persistence.     

Density-dependent dispersal and relative dispersal affect the stability of predator-prey metacommunities

Although density-dependent dispersal and relative dispersal (the difference in dispersal rates between species) have been documented in natural systems, their effects on the stability of metacommunities are poorly understood. Here we investigate the effects of intra- and interspecific density-dependent dispersal on the regional stability in a predator-prey metacommunity model. We show that, when the dynamics of the populations reach equilibrium, the stability of the metacommunity is not affected by density-dependent dispersal. However, the regional stability, measured as the regional variability or the persistence, can be modified by density-dependent dispersal when local populations fluctuate over time. Moreover these effects depend on the relative dispersal of the predator and the prey. Regional stability is modified through changes in spatial synchrony. Interspecific density-dependent dispersal always desynchronizses local dynamics, whereas intraspecific density-dependent dispersal may either synchronize or desynchronize it depending on dispersal rates. Moreover, intra- and interspecific density-dependent dispersal strengthen the top-down control of the prey by the predator at intermediate dispersal rates. As a consequence the regional stability of the metacommunity is increased at intermediate dispersal rates. Our results show that density-dependent dispersal and relative dispersal of species are keys to understanding the response of ecosystems to fragmentation.     

The effects of resource enrichment, dispersal, and predation on local and metacommunity structure

Community structure is the observable outcome of numerous processes. We conducted a laboratory experiment using a microbial model system to disentangle effects of nutrient enrichment, dispersal, and predation on prey species richness and predator abundance at local and metacommunity scales. Prey species included: Chilomonas sp., Colpidium striatum, Colpoda cucullus, Paramecium tetraurelia, P. caudatum, Philodina sp., Spirostomum sp., Tetrahymena thermophila, and Uronema sp., and Stentor coeruleus was the predator used. We hypothesized that: (1) increased basal resources should maintain greater species richness and higher predator abundance; (2) dispersal should maintain greater species richness; and (3) predation should reduce species richness, especially in the high resource treatments relative to no-predator treatments. Our results support all three hypotheses. Further, we show that dispersal affects richness at the local community scale but not at the metacommunity scale. However, predation seems to have major effects at both the local and metacommunity scale. Overall, our results show that effects of resource enrichment, dispersal, and predation were mostly additive rather than interactive, indicating that it may be sometimes easier to understand their effects than generally thought due to complex interactive effects.     

Dispersal rates affect species composition in metacommunities of Sarracenia purpurea inquilines

Dispersal among local communities can have a variety of effects on species composition and diversity at local and regional scales. Local conditions (e.g., resource and predator densities) can have independent effects, as well as interact with dispersal, to alter these patterns. Based on metacommunity models, we predicted that local diversity would show a unimodal relationship with dispersal frequency. We manipulated dispersal frequencies, resource levels, and the presence of predators (mosquito larvae) among communities found in the water-filled leaves of the pitcher plant Sarracenia purpurea. Diversity and abundance of species of the middle trophic level, protozoa and rotifers, were measured. Increased dispersal frequencies significantly increased regional species richness and protozoan abundance while decreasing the variance among local communities. Dispersal frequency interacted with predation at the local community scale to produce patterns of diversity consistent with the model. When predators were absent, we found a unimodal relationship between dispersal frequency and diversity, and when predators were present, there was a flat relationship. Intermediate dispersal frequencies maintained some species in the inquiline communities by offsetting extinction rates. Local community composition and the degree of connectivity between communities are both important for understanding species diversity patterns at local and regional scales.     

The role of habitat connectivity and landscape geometry in experimental zooplankton metacommunities

Theory predicts that inter-patch dispersal rates and patterns of patch heterogeneity both have the potential to alter patterns of local and regional species diversity. To test this, we manipulated both rates of habitat connectivity and the geometric arrangement of habitat heterogeneity within regions of experimental zooplankton communities. We found no effects of habitat geometry on any metric of species diversity or composition. Additionally, we found no effect of habitat connectivity rate on local species diversity. We did, however, find that increasing connectivity led to a decrease in regional diversity, as well as an increase in the percent similarity of local communities within regions. Of all of the species in these communities, the relatively large cladoceran Ceriodaphnia reticulata significantly responded to the treatments, and had a higher probability of achieving high densities when connectance was high. As such, we suggest that this species played a large role in driving the increased local community similarity and decreased regional species richness as connectivity increased. These findings are in opposition to previous experimental studies of metacommunities, but support the notion that increased connectance among local patches may decrease regional diversity when patches are heterogeneous.     

Consequences of varying regional heterogeneity in source–sink metacommunities

Although the influence of dispersal on coexistence mechanisms in metacommunities has received great emphasis, few studies have addressed how such influence is affected varying regional heterogeneity. We present a mechanistic model of resource competition in a metacommunity based on classical models of plant competition for limiting resources. We defined regional heterogeneity as the differences in resource supply rates (or resource availabilities) across local communities. As suggested by previous work, the highest diversify occurred at intermediate levels of dispersal among local communities. However our model shows how the effects of dispersal depend on the amount of heterogeneity among local communities and vice versa. Both regional and local species richness were the highest when heterogeneity was intermediate. We suggest that empirical studies that found no evidence for source–sink or mass effects at the community level may have examined communities with limited ranges of dispersal and regional heterogeneity. This model of species coexistence contributes to a broader understanding of patterns in real communities.     

Metacommunity size influences aquatic community composition in a natural mesocosm landscape

Ecosystems are often arranged in naturally patchy landscapes with habitat patches linked by dispersal of species in a metacommunity. The size of a metacommunity, or number of patches, is predicted to influence community dynamics and therefore the structure and function of local communities. However, such predictions have yet to be experimentally tested using full food webs in natural metacommunities. We used the natural mesocosm system of aquatic macroinvertebrates in bromeliad phytotelmata to test the effect of the number of patches in a metacommunity on species richness, abundance, and community composition. We created metacommunities of varying size using fine mesh cages to enclose a gradient from a single bromeliad up to the full forest. We found that species richness, abundance, and biomass increased from enclosed metacommunities to the full forest size and that diversity and evenness also increased in larger enclosures. Community composition was affected by metacommunity size across the full gradient, with a more even detritivore community in larger metacommunities, and taxonomic groups such as mosquitoes going locally extinct in smaller metacommunities. We were able to divide the effects of metacommunity size into aquatic and terrestrial habitat components and found that the importance of each varied by species; those with simple life cycles were only affected by local aquatic habitat whereas insects with complex life cycles were also affected by the amount of terrestrial matrix. This differential survival of obligate and non‐obligate dispersers allowed us to partition the beta‐diversity between metacommunities among functional groups. Our study is one of the first tests of metacommunity size in a natural metacommunity landscape and shows that both diversity and community composition are significantly affected by metacommunity size. Synthesis Natural food webs are sensitive to meta‐community size, i.e. the number of patches connected through dispersal. We provide an empirical test using the aquatic foodweb associated within bromeliads as a model system. When we reduced the number of bromeliad patches connect through dispersal, we found a clear change of the foodweb in terms of population sizes, beta diversity, community composition and predator‐prey ratios. The response of individual taxa was predictable based on species traits including dispersal modes, life cycle, and adult resource requirements. Our study demonstrates that community structure is strongly influenced by the interplay of species traits and landscape properties.     

The effect of predation on the prevalence and aggregation of pathogens in prey

Although pathogens and predators have been widely used as bio-control agents against problematic prey species, little has been done to examine the prevalence and aggregation of pathogens in spatially structured eco-epidemiological systems. Here, we present a spatial model of a predator-prey/host-parasite system based on pair approximation and spatially stochastic simulations, with the predation pressure indicated by predator abundance and predation rates. Susceptible prey can not only be infected by contacting adjacent infected individuals but also by the global transmission of pathogens. The disease prevalence was found to follow a hump-shaped function in response to predation pressure. Moreover, predation pressure was not always negatively correlated with pathogen aggregation as proposed from empirical studies, but depending on the level of predation pressure. Highly connected site network facilitated the parasites infection, especially under high predation pressure. However, the connectivity of site network had no effect on the prevalence and aggregation of pathogens that can infect health prey through global transmission. It is thus possible to better design biological control strategies for target species by manipulating predation pressure and the range of pathogen transmission.     

Transitory versus Persistent Effects of Connectivity in Environmentally Homogeneous Metacommunities

While the effect of habitat connectivity on local and regional diversity has been analysed in a number of studies, time-dependent dynamics in metacommunities have received comparatively little consideration. When local patches of a metacommunity are identical in environmental conditions but differ in initial community composition, dispersal among patches may result in homogenization of local communities. In a microcosm experiment with benthic ciliates, we tested the hypothesis that the effect of connectivity on diversity is time-dependent and only transitory, with the degree of connectivity affecting the time to homogenization but not the final outcome. Six microcosms were connected to a metacommunity with one of three levels of connectivity. The six patches differed in initial community composition but were identical in environmental conditions. We found a time-dependent and transitory effect of connectivity on local and regional richness and on local Shannon diversity, while Bray-Curtis dissimilarity and regional Shannon diversity were persistently affected by connectivity. Local richness increased and regional richness decreased with connectivity during the initial phase of the experiment but soon converged to similar values in all three connectivity treatments. Local Shannon diversity was unimodally related to time, with maximum diversity reached sooner with high than with medium or low connectivity. Eventually, however, local diversity converged to similar values irrespective of connectivity. At the regional scale, Shannon diversity was persistently lower with high than with low connectivity. While initial differences in community composition vanished with medium and high connectivity, they were maintained with low connectivity resulting in persistently high beta and regional diversity. The effect of connectivity on ciliate community composition translated down to the algal resource, as stronger dominance of the superior competitor with high and medium connectivity resulted in stronger depletion of the resource.     

Spatial averaging and disturbance lead to high productivity in aquatic metacommunities

Dispersal in heterogeneous ecosystems, such as coastal metacommunities, is a major driver of diversity and productivity. According to theory, both species richness and spatial averaging shape a unimodal relationship of productivity with dispersal. We experimentally tested the hypothesis that disturbances acting on local patches would buffer the loss of productivity at high dispersal by preventing synchronized species oscillations. To simulate these disturbances, our experimental assemblages involved species that self‐organized in isolation under three inflow pulsing frequencies, where hydraulic displacement and nutrient loading affected assemblage diversity and composition. At steady‐state, the emerging isolated assemblages were connected at three levels of dispersal creating three metacommunities of different connectivity. Consistent with theory, as dispersal increased, species richness in the metacommunity declined; productivity however remained high. This occurred because the most productive species in our study (which dominated the isolated patch of intermediate inflow pulsing frequency) dominated all three patches (low, intermediate and high inflow pulsing frequencies) after dispersal commenced in our metacommunities. This experimental result provides empirical support for the mechanism of spatial averaging. Furthermore, disturbances, in the form of localized pulsed inflows, prevented population oscillation synchrony caused by homogenization. Overall, our observations suggest that localized environmental fluctuations and the identity of species seem to be more influential than dispersal in shaping the diversity and composition of phytoplankton assemblages and stabilizing productivity.     

Positive indirect effect of aquatic insects on terrestrial prey is not offset by increased predator density

1. Changes in one prey species' density can indirectly affect the abundance of another prey species if a shared predator eats both species. Sometimes, indirect effects occur when prey straddle habitats, including when riparian predator populations grow in response to emergent aquatic insects and increase predation on terrestrial prey. However, predators may largely switch to aquatic insects or become satiated, reducing predation on terrestrial prey. 2. To determine the net indirect effect of aquatic insects on terrestrial arthropods via generalist spider predators, a field experiment was conducted mimicking midge influx and a wolf spider numerical response inside enclosures near an Icelandic lake. Lab mesocosms were also used to assess per capita rates of spider predation u nder differing levels of midge abundance. 3. Midges always decreased sentinel prey predation, but this effect increased with predator density. When midges were absent, predation increased 30% at a high spider density, but predation was equal between spider treatments when midges were present. In situ arthropods showed no effect of midge or spider treatments, although non‐significant abundance patterns were observed congruent with sentinel prey results. 4. In lab mesocosms, prey survivorship increased ≥50% where midges were present and rapidly saturated; the addition of 5, 20, 50, and 100 midges equivalently reduced spider predation, supporting predator distraction rather than satiation as the root cause. 5. The present results demonstrate a strong positive indirect effect of midges and broadly support the concept that predator responses to alternative prey are a major influence on the magnitude and direction of predator‐mediated indirect effects.     

The importance of blue and green landscape connectivity for biodiversity in urban ponds

The negative impact of urbanization on biodiversity can be buffered by blue (e.g., rivers, ponds) and green (e.g., parks, forests) spaces. However, to prevent biodiversity loss and reduce the risk of local extinctions, blue and green spaces need to be connected by corridors, so that organisms may disperse between sites. Landscape connectivity affects local community composition and metacommunity dynamics by facilitating dispersal. The goal of this study was to test the relative roles of pond environmental properties, spatial structure, and functional landscape connectivity on differentiation of invertebrate metacommunities in urban ponds in the city of Stockholm, Sweden. We characterized functional connectivity as blue connectivity (distance to water bodies), green connectivity (land use), and combined blue-green connectivity. We estimated functional connectivity by using electrical circuit theory to identify dispersal corridors. Interestingly, while circuit theory is often used in single-taxon studies, this method has rarely been applied to multiple taxa forming a metacommunity, as we have done in this study. Indeed, our study contributes toward an increased focus on the role of dispersal at the metacommunity level. We determined that functional connectivity was the most important factor in explaining community differentiation, with the local environment contributing comparatively little, and spatial structure the least. Combined blue-green functional connectivity had a major influence on structuring urban pond communities, explaining 7.8% of the variance in community composition across ponds. Furthermore, we found that increased functional connectivity was associated with an increase in the number of species. In summary, our results suggest that to preserve biodiversity in urban ponds, it is important to enhance functional connectivity, and that open green spaces could augment blue corridors in maintaining functional connectivity in urban pond metacommunities. To generalize these findings, future urban biodiversity studies should compare how functional connectivity affects metacommunities across multiple major cities.     

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.