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PubMed2005

Estimating the prevalence and strength of non-independent predator effects.

Understanding whether multiple predator species have independent effects on shared prey is critical for understanding community dynamics. We describe the prevalence and strength of non-independence between predators by quantifying the prey's risk of predation and the degree to which it deviates from the risk predicted from a null model of independent predator effects. Specifically, we document how frequently non-independent effects occur among ten different multiple predator combinations with mayfly larvae as prey. These predator combinations vary both predator density and predator species richness. Overall, the predator effects were non-independent and translated to an average of 27% fewer prey being consumed compared to independent predator effects. Non-independence of this magnitude is likely to have population level consequences for the prey and influence the distribution or prey preference of predators. Closer inspection shows that much of the risk reduction in this system is weak, to the point of being indistinguishable from independent predator effects, while few effects are strong. This pattern of many weak interactions and few strong ones parallels the pattern of interaction strengths documented previously in intertidal communities. Consequently, understanding strong interactors in multiple predator systems may help us understand the importance of a species.

Animals↗
PubMed2004

Spatial dynamics of specialist seed predators on synchronized and intermittent seed production of host plants.

Masting, the synchronized and intermittent seed production by plant populations, provides highly variable food resources for specialist seed predators. Such a reproductive mode helps minimize seed losses through predator satiation and extinction of seed predator populations. The seed predators can buffer the resource variation through dispersal or extended diapause. We developed a spatially explicit resource-consumer model to understand the effect of masting on specialist seed predators. The masting dynamics were assumed to follow a resource-based model for plant reproduction, and the population dynamics of the predator were represented by a spatially extended Nicholson-Bailey model. The resultant model demonstrated that when host plants reproduce intermittently, seed predator populations go locally extinct, but global persistence of the predator is facilitated by dispersal or extended diapause. Global extinction of the predator resulted when the intermittent reproduction is highly synchronized among plants. An approximate invasion criterion for the predators showed that negative lag-1 autocorrelation in seeding reduces invasibility, and positive lag-1 cross-correlation enhances invasibility. Spatial synchronization in seeding at local scale caused by pollen coupling (or climate forcing) further prevented invasion of the predators. If the predators employed extended diapause, extremely high temporal variability in reproduction was required for plants to evade the predators.

Animals↗
PubMed2004

A community-level evaluation of the impact of prey behavioural and ecological characteristics on predator diet composition.

Although predation avoidance is the most commonly invoked explanation for vertebrate social evolution, there is little evidence that individuals in larger groups experience lower predation rates than those in small groups. We compare the morphological and behavioural traits of mammal prey species in the Taï forest, Ivory Coast, with the diet preferences of three of their non-human predators: leopards, chimpanzees and African crowned eagles. Individual predators show marked differences in their predation rates on prey species of different body sizes, but clear patterns with prey behaviour were apparent only when differences in prey habitat use were incorporated into the analyses. Leopard predation rates are highest for terrestrial species living in smaller groups, whereas eagle predation rates are negatively correlated with group size only among arboreal prey. When prey predation rates are summed over all three predators, terrestrial species incur higher predation rates than arboreal species and, within both categories, predation rates decline with increasing prey group size and decreasing density of groups in the habitat. These results reveal that it is necessary to consider anti-predator strategies in the context of a dynamic behavioural interaction between predators and prey.

Animals↗
PubMed2006

Predator identity and additive effects in a treehole community.

Multiple predator species can interact as well as strongly affect lower trophic levels, resulting in complex, nonadditive effects on prey populations and community structure. Studies of aquatic systems have shown that interactive effects of predators on prey are not necessarily predictable from the direct effects of each species alone. To test for complex interactions, the individual and combined effects of a top and intermediate predator on larvae of native and invasive mosquito prey were examined in artificial analogues of water-filled treeholes. The combined effects of the two predators were accurately predicted from single predator treatments by a multiplicative risk model, indicating additivity. Overall survivorship of both prey species decreased greatly in the presence of the top predator Toxorhynchites rutilus. By itself, the intermediate predator Corethrella appendiculata increased survivorship of the native prey species Ochlerotatus triseriatus and decreased survivorship of the invasive prey species Aedes albopictus relative to treatments without predators. Intraguild predation did not occur until alternative prey numbers had been reduced by approximately one-half. Owing to changes in size structure accompanying its growth, T. rutilus consumed more prey as time progressed, whereas C. appendiculata consumed less. The intermediate predator, C. appendiculata, changed species composition by preferentially consuming A. albopictus, while the top predator, T. rutilus, reduced prey density, regardless of species. Although species interactions were in most cases predicted from pairwise interactions, risk reduction from predator interference occurred when C. appendiculata densities were increased and when the predators were similarly sized.

Animals↗
PubMed2006

Survival trade-offs between two predator-induced phenotypes in Pacific treefrogs (Pseudacris regilla).

In many organisms, specific predator species induce defensive phenotypes that are qualitatively different from the phenotypes induced by other predator species. This differential induction implies that there is no optimal phenotype that works best against all predators. However, few studies have actually tested the hypothesis that each predator-induced phenotype provides the highest survival rate in encounters with the predator that induced that phenotype. In this experiment, I reared Pacific treefrog (Pseudacris regilla) larvae with chemical cues from two different predators (bluegill sunfish and predaceous diving-beetle larvae), and without predator cues. The Pacific treefrog larvae in the three treatments differed in their morphology and foraging behavior. I then exposed tadpoles from each treatment to free-foraging predaceous diving beetles and bluegill sunfish. Tadpoles survived best when exposed to the predator whose cues they were reared with, and worst when exposed to the other predator. In both predator environments, the tadpoles reared in the nonpredator control treatment had intermediate survival between the two predator-induced groups. Thus, there is no generalized "antipredator" response to these predators; rather, there was a clear trade-off in survival abilities between the predators.

Analysis of Variance↗
PubMed2002

Intraguild interference and biocontrol effects of generalist predators in a winter wheat field.

Arable land typically harbours communities of polyphagous invertebrate natural enemies, among them numerous soil-surface dwelling predators such as ground beetles (Carabidae) and spiders (Lycosidae, Linyphiidae). Numbers of these predators were experimentally manipulated in a winter wheat field in order to study the predation impact of a generalist predator assemblage on herbivorous insects, the possible interferences among the predators concerned, and subsequent effects on wheat plant parameters. Removing ground beetles doubled numbers of Lycosidae indicative of intraguild interference between these two predator groups. Aphid densities were highest in carabid removal plots implying a substantial predation impact of ground beetles on the pest population. The predation impact of ground beetles was strongest earlier and disappeared later in the season. In mid-season, at intermediate aphid densities, the combined impact of carabid beetles and spiders appeared to be responsible for the reduction in aphid abundance. This result was probably due to a biomass effect rather than to a synergistic effect of the predator community. Thysanoptera decreased when spiders were removed (perhaps because spiders were preying on a predator of thrips), while Cicadellidae and Delphacidae showed no effect at all. The rise of aphid numbers in carabid removal plots corresponded to an increase in protein content of the wheat grains, while other plant parameters such as plant numbers and grain mass were not affected. In conclusion, this study provided field evidence for intraguild interference among generalist ground predators in arable land. Despite this interference the polyphagous predator community was able to depress numbers of aphids in winter wheat, a result cascading down to plant quality parameters.

Animals↗
PubMed2005

Sequential predator effects across three life stages of the African tree frog, Hyperolius spinigularis.

While theoretical studies of the timing of key switch points in complex life cycles such as hatching and metamorphosis have stressed the importance of considering multiple stages, most empirical work has focused on a single life stage. However, the relationship between the fitness components of different life stages may be complex. Ontogenetic switch points such as hatching and metamorphosis do not represent new beginnings--carryover effects across stages can arise when environmental effects on the density and/or traits of early ontogenetic stages subsequently alter mortality or growth in later stages. In this study, I examine the effects of egg- and larval-stage predators on larval performance, size at metamorphosis, and post-metamorphic predation in the African tree frog Hyperolius spinigularis. I monitored the density and survival of arboreal H. spinigularis clutches in the field to estimate how much egg-stage predation reduced the input of tadpoles into the pond. I then conducted experiments to determine: (1) how reductions in initial larval density due to egg predators affect larval survival and mass and age at metamorphosis in the presence and absence of aquatic larval predators, dragonfly larvae, and (2) how differences in mass or age at metamorphosis arising from predation in the embryonic and larval environments affect encounters with post-metamorphic predators, fishing spiders. Reduction in larval densities due to egg predation tended to increase per capita larval survival, decrease larval duration and increase mass at metamorphosis. Larval predators decreased larval survival and had density-dependent effects on larval duration and mass at metamorphosis. The combined effects of embryonic and larval-stage predators increased mass at metamorphosis of survivors by 91%. Larger mass at metamorphosis may have immediate fitness benefits, as larger metamorphs had higher survival in encounters with fishing spiders. Thus, the effects of predators early in ontogeny can alter predation risk even two life stages later.

Age Factors↗
PubMed2005

How does the presence of predators influence the persistence of antipredator behavior?

We developed a virtual world to study the effect of predators on predator recognition. We trained a neural network to discriminate between the shapes of simulated aerial and terrestrial predators and non-predators. Then, the network's weighting values were fixed into the genomes of a set of autonomous agents. These animats were required to eat, avoid death due to starvation, and avoid predation, by fleeing from approaching predators. We systematically varied the predator's lethality, the mutation rate, the cost of fleeing a predator, and the presence or absence of aerial and terrestrial predators. We used ANOVA to analyse the average recognition ability (a measure of directional selection) and the standard deviation of recognition ability (a measure of relaxed selection) after 500 generations of selection. Mutation rate and the cost of flight had the greatest effect on both the average and standard deviation of recognition abilities. The loss of all predators relaxed selection on predator recognition abilities. The loss of specific predators had complex effects on recognition abilities. Persistence is largely influenced by escape costs.

Animals↗
PubMed2006

Productivity, dispersal and the coexistence of intraguild predators and prey.

A great deal is known about the influence of dispersal on species that interact via competition or predation, but very little is known about the influence of dispersal on species that interact via both competition and predation. Here, I investigate the influence of dispersal on the coexistence and abundance-productivity relationships of species that engage in intraguild predation (IGP: competing species that prey on each other). I report two key findings. First, dispersal enhances coexistence when a trade-off between resource competition and IGP is strong and/or when the Intraguild Prey has an overall advantage, and impedes coexistence when the trade-off is weak and/or when the Intraguild Predator has an overall advantage. Second, the Intraguild Prey's abundance-productivity relationship depends crucially on the dispersal rate of the Intraguild Predator, but the Intraguild Predator's abundance-productivity relationship is unaffected by its own dispersal rate or that of the Intraguild Prey. This difference arises because the two species engage in both a competitive interaction as well as an antagonistic (predator-prey) interaction. The Intraguild Prey, being the intermediate consumer, has to balance the conflicting demands of resource acquisition and predator avoidance, while the Intraguild Predator has to contend only with resource acquisition. Thus, the Intraguild Predator's abundance increases monotonically with resource productivity regardless of either species' dispersal rate, while the Intraguild Prey's abundance-productivity relationship can increase, decrease, or become hump-shaped with increasing productivity depending on the Intraguild Predator's dispersal rate. The important implication is that a species' trophic position determines the effectiveness of dispersal in sampling spatial environmental heterogeneity. The dispersal behavior of a top predator is likely to have a stronger effect on coexistence and spatial patterns of abundance than the dispersal behavior of an intermediate consumer.

Animals↗
PubMed2003

Evolution of prey behavior in response to changes in predation regime: damselflies in fish and dragonfly lakes.

In a large behavioral experiment we reconstructed the evolution of behavioral responses to predators to explore how interactions with predators have shaped the evolution of their prey's behavior. All Enallagma damselfly species reduced both movement and feeding in the presence of coexisting predators. Some Enallagma species inhabit water bodies with both fish and dragonflies, and these species responded to the presence of both predators, whereas other Enallagma species inhabit water bodies that have only large dragonflies as predators, and these species only responded to the presence of dragonflies. Lineages that shifted to live with large dragonflies showed no evolution in behaviors expressed in the presence of dragonflies, but they evolved greater movement in the absence of predators and greater movement and feeding in the presence of fish. These results suggest that Enallagma species have evolutionarily lost the ability to recognize fish as a predator. Because species coexisting with only dragonfly predators have also evolved the ability to escape attacking dragonfly predators by swimming, the decreased predation risk associated with foraging appears to have shifted the balance of the foraging/predation risk trade-off to allow increased activity in the absence of mortality threats to evolve in these lineages. Our results suggest that evolution in response to changes in predation regime may have greater consequences for characters expressed in the absence of mortality threats because of how the balance between the conflicting demands of growth and predation risk are altered.

Animals↗
PubMed2006

Species identity dominates the relationship between predator biodiversity and herbivore suppression.

Agricultural pest suppression is an important ecosystem service that may be threatened by the loss of predator diversity. This has stimulated interest in the relationship between predator biodiversity and biological control. Multiple-predator studies have shown that predators may complement or interfere with one another, but few experiments have determined if the resulting effects on prey are caused by changes in predator abundance, identity, species richness, or some combination of these factors. We experimentally isolated the effect of predator species richness on the biological control of an important agricultural pest, the green peach aphid. We found no evidence that increasing predator species richness affects aphid biological control; overall there was no strong complementarity or interference among predator species that altered the strength of aphid suppression. Instead, our experiments revealed strong effects of predator species identity, because predators varied dramatically in their per capita consumption rates. Our results are consistent with other multiple-predator studies finding strong species-identity effects and suggest that, for the biological control of aphids, conservation strategies that directly target key species will be more effective than those targeting predator biodiversity more broadly.

Animals↗
PubMed2005

Behavior of lead in a model microbial predator-prey system.

Predation at the microbial level can affect the fate of toxic trace metals. Metals associated with bacterial prey can be released into the dissolved phase following digestion by a predator, and/or metals can remain in the predator and be transferred potentially to the next level of the food chain. Toxic metal ions in the aqueous phase also are expected to modify the growth and predation rate of a microbial predator. A model predator-prey system was developed to test the effects of Pb on cells and to help elucidate the fate of Pb in this type of interaction. Established methods that have been shown to be suitable for distinguishing dissolved, prey-bound, predator-bound, and ingested Pb were used to establish the pathway of Pb over time. Growth parameters were measured using batch reactors for the protozoan predator Tetrahymena thermophila and the bacterial prey Pseudomonas putida without Pb and at several concentrations of Pb. The effect of prey density on predation and Pb phase distribution also was investigated. Results demonstrate that some kinetic parameters related to prey consumption and growth of T. thermophila are altered by Pb. Upon addition of predator to prey cells in equilibrium with dissolved Pb, dissolved and prey-bound Pb become associated with the predator through ingestion and adsorption. Ingested Pb is excreted later as a bound metal associated with T. thermophila waste matter. A preliminary mathematical model was developed to describe predator-prey dynamics and their influence on the behavior and fate of Pb. Growth data were used to obtain model parameters, and model simulations for Pb fractionation are compared to experimental observations.

Animals↗
PubMed2000

The Interaction between Competition and Predation: A Meta-analysis of Field Experiments.

Ecologists working with a range of organisms and environments have carried out manipulative field experiments that enable us to ask questions about the interaction between competition and predation (including herbivory) and about the relative strength of competition and predation in the field. Evaluated together, such a collection of studies can offer insight into the importance and function of these factors in nature. Using a new factorial meta-analysis technique, we combined the results of 20 articles reporting on 39 published field experiments to ask whether the presence of predators affects the intensity of competitive effects and to compare the average effects of competition and predation. Across all studies, the effects of competition in the presence of predators were less than in the absence of predators, and the interaction between competition and predation for most response variables was statistically significant. Removal of competitors had much more positive effects on organisms' growth and mass than did exclusion of predators. Predator exclusion had much more beneficial effects on organisms' survival than did competition. The mean effects of competition and predation on density did not differ from one another. The results differed among trophic levels. Further understanding would benefit greatly from more field experiments that manipulate both competition and predation, that focus on a wider range of organisms and environments, that focus on population-level parameters such as density, and that report results more completely, including data such as sample sizes and variances.

competition↗
PubMed2000

Sex-specific predation on a monogamous rat, Hypogeomys antimena (Muridae: Nesomyinae).

Sex-specific predation on adult individuals is often predicted by different behaviour in males and females resulting from different reproductive strategies and social systems. High predation pressure and the need for biparental care are considered a possible ecological basis for the evolution of monogamy, the most puzzling social system in mammals. In species where adults and offspring are vulnerable to the same predators, males and females may protect their offspring to different extents because of conflicting demands of investment in current and future offspring. I present the first empirical data on age- and sex-specific predation pressure by top predators on a monogamous rodent and the sex-specific behavioural responses of the prey species to different rates of predation. During an annual predation peak, only offspring and adult males were killed but no females. Whereas males and females travelled similar distances at night before the period of high predation on offspring, males moved further during this period. At the same time, males and females increased their distance from their offspring but males stayed closer to them than females. As a consequence, the distance between the members of the pair increased during the predation peak. The males' behaviour could lead to their encountering predators more frequently which would reduce survival prospects. The different behaviour of males and females provides empirical evidence that males invest in the welfare of current offspring at the cost of higher predation risk whereas females protect their residual reproductive value. Copyright 2000 The Association for the Study of Animal Behaviour.

Journal Article↗
PubMed1999

Predator Inspection: Closer Approach as a Way to Improve Assessment of Potential Threats.

When detecting a predator, some prey animals respond in a counterintuitive fashion by approaching, rather than fleeing, that potential threat of extinction. This seemingly paradoxical behaviour, known as predator inspection, has been reported for a wide variety of taxa-and therefore can be assumed to be adaptive. However, the view of predator inspection as a paradoxical behaviour rests on two implicit assumptions: (a) initial predator detecting is unambiguous, with no uncertainty in discriminating between hunting and non hunting members of predator species, or members of predator species and unrelated phenomena; (b) the costs of flight are negligible relative to the risk of predation. Upon reflection assumption (a) is not really tenable. Whereas assumption (b) is not consistent with experimental evidence [Godin & Crossman (1994) Behav. Ecol. Sociobiol.34, 359-366]. Given that predator detection is ambiguous and the costs of flight are not negligible, a prey individual may benefit by a closer approach to the source of the alarming signals, thus improving its assessment of the situation-despite the increased risk of predation. In this paper, the above statement is given rigor by reformulating the problem in game theoretical terms. The results indicate that a prey will minimize its costs by performing predator inspection whenever its degree of certainty regarding predator identification and/or assessment of its intentions is less than a threshold, which is determined by the model's parameters. Copyright 1999 Academic Press.

Journal Article↗
PubMed2006

The effects of different predator species on antipredator behavior in the Trinidadian guppy, Poecilia reticulata.

Different types of predators often elicit different antipredator responses in a common type of prey. Alternatively, some prey species may adopt a general response, which provides limited protection from many different types of predator. The Trinidadian guppy, Poecilia reticulata, is faced with a wide range of different predators throughout its range and is known to display varying levels of antipredator behavior depending on the predator assemblage. Pike cichlids, Crenicichla frenata, are regarded as the primary aquatic guppy predator in streams in the northern mountain range in Trinidad. As such, they are seen to be responsible for many of the differences in morphology, life history traits, and behavior between guppy populations from areas with few predators and those from areas with many pike cichlids. In this study we investigated how guppies responded when faced with different predator species using three common aquatic predators. We exposed shoals of ten guppies to one out of four treatments: no predator (control), pike cichlid, acara cichlid (Aequidens pulcher), and wolf fish (Hoplias malabaricus); and we made behavioral observations on both focal individuals and the shoal as a whole. Guppies showed significantly greater levels of predator inspection and shoaling behavior, foraged less, spent more time in the surface water, and stayed in significantly larger shoals when faced with pike cichlids than in other treatments. We discuss these results in the context of multiple predator effects.

Animals↗
PubMed2006

The effects of prey patchiness, predator aggregation, and mutual interference on the functional response of Phytoseiulus persimilis feeding on Tetranychus urticae (Acari: Phytoseiidae, Tetranychidae).

The spatial distributions of two-spotted spider mites Tetranychus urticae and their natural enemy, the phytoseiid predator Phytoseiulus persimilis, were studied on six full-grown cucumber plants. Both mite species were very patchily distributed and P. persimilis tended to aggregate on leaves with abundant prey. The effects of non-homogenous distributions and degree of spatial overlap between prey and predators on the per capita predation rate were studied by means of a stage-specific predation model that averages the predation rates over all the local populations inhabiting the individual leaves. The empirical predation rates were compared with predictions assuming random predator search and/or an even distribution of prey. The analysis clearly shows that the ability of the predators to search non-randomly increases their predation rate. On the other hand, the prey may gain if it adopts a more even distribution when its density is low and a more patchy distribution when density increases. Mutual interference between searching predators reduces the predation rate, but the effect is negligible. The stage-specific functional response model was compared with two simpler models without explicit stage structure. Both unstructured models yielded predictions that were quite similar to those of the stage-structured model.

Animals↗
PubMed2005

Role of NMDA receptors in the lateralized potentiation of amygdala afferent and efferent neural transmission produced by predator stress.

The present study investigated the role of NMDA receptors in behavioral and neuroplastic changes in amygdala efferent (central amygdala to periaqueductal gray-ACE-PAG) and amygdala afferent (ventral angular bundle to basolateral amygdala-VAB-BLA) pathways in response to predator stress. Effects on brain and behavioral response to predator stress of competitive block of NMDA receptors with a dose of 10 mg/kg of CPP (3-(2-carboxypiperazin4-yl)propyl-l-phosphonic acid) were studied. Behavioral response to stress was tested with hole board, elevated plus maze, light/dark box, social interaction and acoustic startle tests. CPP was administered i.p. 30 min prior to predator stress and blocked the effects of predator on some but not all behaviors measured 8-9 days later. Effects of predator stress and CPP on potentials evoked in the PAG by single pulse stimulation of the ACE and in the BLA by single pulse stimulation of VAB were assessed 10-11 days after predator stress. Predator stress potentiated ACE-PAG evoked potentials in the right but not the left hemisphere, replicating previous work. Predator stress potentiated VAB-BLA transmission in both hemispheres 10-11 days after predator stress. Right hemisphere VAB-BLA potentiation replicated and extended past studies showing right hemisphere potentiation at 1 and 9 days after stress. Left VAB-BLA potentiation effects differed from the long term depression seen in VAB-BLA at 1 and 9 days after stress in previous studies. CPP blocked predator stress-induced potentiation of ACE-PAG and VAB-BLA evoked potentials in the right hemisphere. CPP did not block left VAB-BLA potentiation, rather CPP amplified it. Left hemisphere effects of CPP were interpreted as reflecting block of NMDA dependent long term depression, which unmasked a non-NMDA dependent potentiation. Taken together, the findings add to a body of evidence suggesting that a syndrome of behavioral changes follows predator stress. Components of this syndrome likely depend on changes in separable neural substrates. Potentiation of ACE-PAG and VAB-BLA evoked potentials in the right hemisphere likely mediates a subset of changes in behavior. Moreover, a medial ACE-PAG pathway is implicated in mediating stress-induced changes in startle amplitude. In contrast, a lateral ACE-PAG pathway is implicated in mediating changes in startle habituation. Finally, consistent with cat and human studies, the right hemisphere appears particularly important in long term response to stress.

Amygdala↗