Insect herbivory and vertebrate grazing impact food limitation and grasshopper populations during a severe outbreak

1. Competition between herbivores often plays an important role in population ecology and appears strongest when densities are high or plant production is low. Phytophagous insects are often highly abundant, but relatively few experiments have examined competition between vertebrates and phytophagous insects. 2. In grassland systems worldwide, grasshoppers are often the dominant phytophagous insect, and livestock grazing is a dominant land use. For this study, a novel experiment was conducted examining competition between vertebrates and invertebrates, where both grasshopper densities and sheep grazing were manipulated inside 10‐m² caged mesocosms during a grasshopper outbreak. We examined how grasshopper densities and the timing of vertebrate herbivory affected grasshopper densities, if the effects of vertebrates on survival and reproduction changed with grasshopper density, and how a naturally occurring grasshopper outbreak affected grasshopper populations in the following year. 3. Densities of grasshoppers at the site peaked at 130 m^–2. Food‐limited competition was stronger in treatments with higher grasshopper densities and repeated or late livestock herbivory, leading to reduced survival, femur length, and functional ovarioles, a measure of future reproduction. Strong food‐limited density‐dependent reproduction and survival led to reduced hatching densities in 2001. 4. As competition was typically stronger with high grasshopper densities than with livestock grazing, competition from vertebrates could be relatively less important for phytophagous insect population dynamics during outbreaks. The experiment provides insights into how competition between insect and vertebrate herbivores influences insect population dynamics, and indicates that severe outbreaks can rapidly subside with strong competition from vertebrate and insect herbivores.     

Predation- and competition-mediated brain plasticity in Rana temporaria tadpoles

An increasing number of studies have demonstrated phenotypic plasticity in brain size and architecture in response to environmental variation. However, our knowledge on how brain architecture is affected by commonplace ecological interactions is rudimentary. For example, while intraspecific competition and risk of predation are known to induce adaptive plastic modifications in morphology and behaviour in a wide variety of organisms, their effects on brain development have not been studied. We studied experimentally the influence of density and predation risk on brain development in common frog (Rana temporaria) tadpoles. Tadpoles grown at low density and under predation risk developed smaller brains than tadpoles at the other treatment combinations. Further, at high densities, tadpoles developed larger optic tecta and smaller medulla oblongata than those grown at low densities. These results demonstrate that ecological interactions - like intraspecific competition and predation risk - can have strong effects on brain development in lower vertebrates.     

Non-lethal effects of predation in birds

Predators can affect individual fitness and population and community processes through lethal effects (direct consumption or ‘density’ effects), where prey is consumed, or through non‐lethal effects (trait‐mediated effects or interactions), where behavioural compensation to predation risk occurs, such as animals avoiding areas of high predation risk. Studies of invertebrates, fish and amphibians have shown that non‐lethal effects may be larger than lethal effects in determining the behaviour, condition, density and distribution of animals over a range of trophic levels. Although non‐lethal effects have been well described in the behavioural ecology of birds (and also mammals) within the context of anti‐predation behaviour, their role relative to lethal effects is probably underestimated. Birds show many behavioural and physiological changes to reduce direct mortality from predation and these are likely to have negative effects on other aspects of their fitness and population dynamics, as well as affecting the ecology of their own prey and their predators. As a consequence, the effects of predation in birds are best measured by trade‐offs between maximizing instantaneous survival in the presence of predators and acquiring or maintaining resources for long‐term survival or reproduction. Because avoiding predation imposes foraging costs, and foraging behaviour is relatively easy to measure in birds, the foraging–predation risk trade‐off is probably an effective framework for understanding the importance of non‐lethal effects, and so the population and community effects of predation risk in birds and other animals. Using a trade‐off approach allows us to predict better how changes in predator density will impact on population and community dynamics, and how animals perceive and respond to predation risk, when non‐lethal effects decouple the relationship between predator density and direct mortality rate. The trade‐off approach also allows us to identify where predation risk is structuring communities because of avoidance of predators, even when this results in no observable direct mortality rate.     

Disentangling interference competition from exploitative competition in a crab–bivalve system using a novel experimental approach

In predator–prey relationships such as those between crabs and their bivalve prey, interference competition is a topic of intense investigation as it can have profound consequences on the dynamics of both predator and prey populations. However in laboratory experiments – also those on crab–bivalve systems – workers never adequately disentangled interference competition from exploitative competition, as prey depletion was never compensated. Hitherto, experimental studies on crab–bivalve systems lack direct behavioural observations and have provided only indirect and thus inconclusive evidence of interference competition. We studied interference competition in adult male shore crabs Carcinus maenas that foraged on blue mussels Mytilus edulis. We developed a novel type of experimental tank to replenish each consumed mussel, and thus to keep prey levels constant. We conducted two experiments in which we varied number of crabs (1, 2, 4) and number of mussels (first experiment: 4, 8, 16, 32; second experiment: 8, 32, 128) and directly observed the foraging behaviour of crabs (foraging area=0.25 m²). In the first experiment, feeding rates decreased with increasing crab density only at mussel density 16 because both search time and time spent in agonistic interactions increased. At other mussel densities, variation in crab density did not affect feeding rates, possibly because of low statistical power and the narrow range of mussel densities offered. In the second experiment feeding rates decreased with increasing crab density because crabs spent more time in agonistic interactions and handling their prey. Feeding rates increased with increasing mussel density. Overall, crabs spent on average 14–18% of their foraging time in agonistic behaviours, while on three out of 64 occasions feeding rates decreased because mussels were stolen (kleptoparasitism). Concluding, we have shown that interference competition occurs in absence of prey depletion, while conducting direct behavioural observations aid to identify the behavioural processes that underlie interference competition.     

Partitioning the sources of demographic variation reveals density‐dependent nest predation in an island bird population

Ecological factors often shape demography through multiple mechanisms, making it difficult to identify the sources of demographic variation. In particular, conspecific density can influence both the strength of competition and the predation rate, but density‐dependent competition has received more attention, particularly among terrestrial vertebrates and in island populations. A better understanding of how both competition and predation contribute to density‐dependent variation in fecundity can be gained by partitioning the effects of density on offspring number from its effects on reproductive failure, while also evaluating how biotic and abiotic factors jointly shape demography. We examined the effects of population density and precipitation on fecundity, nest survival, and adult survival in an insular population of orange‐crowned warblers (Oreothlypis celata) that breeds at high densities and exhibits a suite of traits suggesting strong intraspecific competition. Breeding density had a negative influence on fecundity, but it acted by increasing the probability of reproductive failure through nest predation, rather than through competition, which was predicted to reduce the number of offspring produced by successful individuals. Our results demonstrate that density‐dependent nest predation can underlie the relationship between population density and fecundity even in a high‐density, insular population where intraspecific competition should be strong.     

Contrasting effects of habitat fragmentation,population density,and prey availability on body condition of two orb‐weaving spiders

1. Habitat fragmentation is a major threat to biodiversity because it disrupts movement between habitat patches. In addition, arthropod fitness may be reduced in fragmented habitats, e.g. due to reduced prey availability. 2. We studied the relationship of spider body condition with habitat fragmentation, population density, and prey availability. We expected that prey availability and population density of spiders would be affected by landscape composition and patch isolation. Body condition should be enhanced by high prey availability, but negatively affected by population density due to competition. 3. We sampled spiders on 30 groups of cherry trees that varied independently in the level of isolation from other woody habitats and in the percentage of woody habitat within 500 m radius. As a measure of body condition, we used residuals of the relationship between individual body mass/opisthosoma width and prosoma width of the two most common orb‐weaving spider species, Nuctenea umbratica Clerck and Araniella opisthographa Kulczynski. 4. Body condition of A. opisthographa was positively correlated with the abundance of flies, which increased with the percentage of forest in the landscape. In contrast, body condition of N. umbratica was reduced at high population densities, presumably due to intraspecific competition. In addition, body condition and population density of A. opisthographa was lower at isolated sites. 5. Our study suggests that effects of landscape fragmentation on body condition vary strongly between spider species, depending on the relative role of food limitation and intraspecific competition.     

Predation risk tradeoffs in prey: effects on energy and behaviour

The complexity of behavioural interactions in predator-prey systems has recently begun to capture trait-effects, or non-lethal effects, of predators on prey via induced behavioural changes. Non-lethal predation effects play crucial roles in shaping population and community dynamics, particularly by inducing changes to foraging, movement and reproductive behaviours of prey. Prey exhibit trade-offs in behaviours while minimizing predation risk. We use a novel evolutionary ecosystem simulation EcoSim to study such behavioural interactions and their effects on prey populations, thereby addressing the need for integrating multiple layers of complexity in behavioural ecology. EcoSim allows complex intra- and inter-specific interactions between behaviourally and genetically unique individuals called predators and prey, as well as complex predator-prey dynamics and coevolution in a tri-trophic and spatially heterogeneous world. We investigated the effects of predation risk on prey energy budgets and fitness. Results revealed that energy budgets, life history traits, allocation of energy to movements and fitness-related actions differed greatly between prey subjected to low-predation risk and high-predation risk. High-predation risk suppressed prey foraging activity, increased total movement and decreased reproduction relative to low-risk. We show that predation risk alone induces behavioural changes in prey which drastically affect population and community dynamics, and when interpreted within the evolutionary context of our simulation indicate that genetic changes accompanying coevolution have long-term effects on prey adaptability to the absence of predators.     

Is arthropod predation exclusively satiation‐driven?

Functional response models differ in which factors limit predation (e.g. searching efficiency, prey handling time, digestion) and whether predation behaviour is governed by an internal physiological state (e.g. satiation). There is now much evidence that satiation is a key factor in understanding changes in foraging behaviour, and that many predators are effectively digestion limited. Here, we ask if predation in a predatory arthropod can be explained from satiation-driven behaviour alone, or if behaviour is also influenced by the density of prey other than via the effect of prey ingestion on satiation. To address this question a satiation-based predation model is formulated, for which parameters are estimated on the basis of observations on digestion rate, satiation-related prey searching rate and prey capture behaviour, basically under high prey density conditions. The model predictions are subsequently tested against longer term predation experiments carried out at high and low prey densities. Since satiation can easily be linked with egg production, these tests are carried out both for predation and oviposition.
The predator–prey systems under study consist of females of two predatory mite species ( Neoseiulus barkeri and N. cucumeris ) and the larvae of two thrips species ( Thrips tabaci and Frankliniella occidentalis ) as their prey. For N. barkeri foraging on T. tabaci , the model gives good predictions at both high (4 larvae cm⁻¹) and low (0.1–1 larvae cm⁻²) prey densities. For N. cucumeris foraging on F. occidentalis , the predictions hold at the high prey density, but are too low at low prey densities. Thus our analysis indicates that we cannot fully explain density-dependent predation rates from satiation-driven behaviour alone. Different mechanisms are suggested on how prey density may affect foraging efficiency other than via satiation.     

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

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

Multiple predators induce risk reduction in coexisting vole species

Large predators may affect the hunting efficiency of smaller ones directly by decreasing their numbers, or indirectly by altering their behaviour. Either way this may have positive effects on the density of shared prey. Using large outdoor enclosures, we experimentally studied whether the presence of the Tengmalm's owl Aegolius funereus affects the hunting efficiency of the smallest member of the vole-eating predator guild, the least weasel Mustela nivalis, as measured by population responses of coexisting prey species, the field vole Microtus agrestis and the sibling vole M. levis . We compared the density and survival probability of vole populations exposed to no predation, weasel predation or combined predation by a weasel and an owl. The combined predation of both owl and weasel did not result in obvious changes in the density of sibling and field vole populations compared to the control populations without predators, while predation by least weasel alone decreased the densities of sibling voles and induced a similar trend in field vole densities. Survival of field voles was not affected by predator treatment while sibling vole survival was lower in predator treated populations than in control populations. Our results suggest that weasels are intimidated by avian predators, but without changing the effects of predators on competitive situations between the two vole species. Non-lethal effects of intraguild predation therefore will not necessarily change competitive interactions between shared prey species.     

Shifts in breeding habitat selection behaviour in response to population density

We tested whether mountain pine beetles Dendroctonus ponderosae, an insect herbivore that exhibits outbreak population dynamics, modifies its habitat selection behaviour in response to density‐dependent environmental shifts. Using an individual‐based habitat selection model, we formulated predictions of how beetle population density will influence breeding habitat selectivity. Our model predicted that beetles should be more selective at intermediate densities than at low or high densities. The mechanisms influencing optimal selectivity differed between low and high density populations. In low density populations, breeding site availability was the primary factor affecting selectivity, whereas intraspecific competition and the reliability of habitat quality cues were important in high density populations. We tested our model predictions in natural populations that encompassed a range of beetle population densities. Our empirical findings supported the two key predictions from our model. First, habitat quality was more variable in high density populations. Second, individuals in high density populations were less selective compared to beetles from intermediate density populations. Our results demonstrate that beetles alter their habitat selection behaviour in response to density‐dependent shifts. We propose that the behavioural changes we identified may influence the rate at which beetle populations transition between density states.     

A sublethal effect on native Anthocoris nemoralis through competitive interactions with invasive Harmonia axyridis

1. The invasive ladybird Harmonia axyridis Pallas (Coleoptera: Coccinellidae) is implicated in declines of native ladybirds with intraguild predation identified as an underlying mechanism, however, less studied are the effects on non‐coccinellid predators. Intraguild predation between Anthocoris nemoralis Fabricius (Heteroptera: Anthocoridae) and H. axyridis and the relative effects of inter‐ and intraspecific competition on predators on potted Tilia cordata Mill. was investigated, at two aphid densities. 2. Intraguild predation was unidirectional in favour of H. axyridis in Petri dishes although a low level of ladybird egg predation by A. nemoralis occurred. However, on trees intraguild predation was rare. 3. A positive effect of aphid density on H. axyridis weight gain, and increased survival when reared with A. nemoralis on potted trees indicated stronger effects of intraspecific competition on H. axyridis. Consistent suppression of aphids by ladybirds revealed their superior competition for aphids. 4. Anthocoris nemoralis weight gain was reduced when reared with H. axyridis on potted trees, indicating a potential for a sublethal effect through interspecific competition. Survivorship of A. nemoralis did not differ between con‐ and heterospecific treatments, but high aphid density enhanced A. nemoralis survival. Overall, high mortality among nymphs across treatments suggested effects related to both inter‐ and intraspecific competition for resources. 5. Results provide evidence of a sublethal effect by invasive H. axyridis through resource competition, suggesting that where these predators co‐occur, competitive interactions for shared prey are more likely to affect A. nemoralis populations than intraguild predation by H. axyridis.     

Influence of a large late summer precipitation event on food limitation and grasshopper population dynamics in a northern Great Plains grassland

The complex interplay between grasshoppers, weather conditions, and plants that cause fluctuations in grasshopper populations remains poorly understood, and little is known about the ecological processes that generate grasshopper outbreaks. Grasshopper populations respond to interacting extrinsic and intrinsic factors, with yearly and decadal weather patterns and the timing of precipitation all potentially important. The effects of initial and increasing grasshopper densities on grasshopper survival and reproductive correlates were examined at a northern mixed-grass prairie site through manipulations of grasshopper densities inside 10-m2 cages. High-quality grass growth occurred after a 9.1-cm mid-August rain. Reduced proportional survival was apparent in the two higher density treatments before the rain, indicative of food-limited density-dependent mortality. However, the large late summer rainfall event mediated the effects of exploitative competition on demographic characteristics because of the high-quality vegetation growth. This led to weaker effects of food limitation on survival and reproduction at the end of the experiment. The results indicate a direct link between weather variation, resource quality and grasshopper population dynamics led to a severe grasshopper outbreak and show that infrequent large precipitation events can have significant effects on population dynamics. Additional research is needed to examine the importance of infrequent large precipitation events on grasshopper population dynamics in grassland ecosystems.     

Local sex ratio affects the cost of reproduction

1.?Costs and benefits of reproduction are central to life-history theory, and the outcome of reproductive trade-offs may depend greatly on the ecological conditions in which they are estimated. In this study, we propose that costs and benefits of reproduction are modulated by social effects, and consequently that selection on reproductive rates depends on the social environment. 2.?We tested this hypothesis in a great tit Parus major population. Over 3 years, we altered parental reproductive effort via brood size manipulations (small, intermediate, large) and manipulated the local social environment via changes in the local fledgling density (decreased, increased) and the local sex ratio (female-biased, control, male-biased). 3.?We found that male-biased treatment consistently increased the subsequent local breeding densities over the 3-year study period. We also found that parents rearing small broods in these male-biased plots had increased survival rates compared with the other experimental groups. 4.?We conclude that reproductive costs are the product of an interaction between parental phenotypic quality after reproduction and the social environment: raising a small brood had long-lasting effects on some phenotypic traits of the parents and that this increased their survival chances in male-biased environment where habitat quality may have deteriorated (via increased disease/predation risk or intraspecific competition). 5.?Our results provide the first experimental evidence that local sex ratio can affect reproductive costs and thus optimal clutch size.     

Predation and searching efficiency of a ladybird beetle, Coccinella septempunctata Linnaeus in laboratory environment

The predation and searching efficiency of fourth instar of predatory C. septempunctata at various densities of mustard aphid, Lipaphis erysimi (Kaltenbach) and predator was investigated under laboratory conditions. The feeding rate of predatory stage decreased at increased prey- and predator densities. Highest percent (92.80%) prey consumption was observed at initial prey density and lowest percent (40.86%) prey consumption at highest prey density by the fourth instar, though the total prey consumption increased with increase in either prey- or predator densities. Similarly, the individual prey consumption was also highest at initial predator density and lowest at highest predator density owing to the mutual interference between the predators at higher densities. The area of discovery (searching efficiency) also decreased with increase in prey- and predator densities. Handling time of predator was highest at lower prey densities, which decreased with increased prey densities. The highest percentage of prey consumption at the prey density of 50 revealed that 1:50 predator-prey ratio was the best to reduce the pest population.     

Predator-induced changes in population structure and individual quality of Microtus voles: a large-scale field experiment

In small mammal populations with multiannual oscillations in density, observational data have revealed cyclic changes in population structure, reproduction, and individual quality, but mechanisms inducing these changes have remained an open question. We analysed data collected during a 3-year predator reduction experiment to find out the effects of predators on population structure, reproductive parameters, and individual quality of Microtus voles (the field vole M. agrestis and the sibling vole M. rossiaemeridionalis ) in western Finland. Voles were collected by snap trapping in April, June, August, and October during 1997–1999. The yearly reduction of predators from April to October had a clear positive effect on the abundance of sibling voles but did not significantly affect the densities of field voles. Predator reduction apparently also affected the age ratio and mean body size in late summer, as well as pancreatic weights of voles. However, all observed differences between predator reduction and control areas, except those in abundance, were small and may mainly reflect a generally higher survival leading to higher densities of voles in predator reduction areas. Our results also indicated a relative lack of high quality food at population peaks but not because of reduced foraging activity in the presence of predators. We conclude that the indirect effects of vole-eating predators on the population growth of main prey are small compared to the detrimental direct effects on prey survival. In the case of less preferred prey, indirect effects of predation through reduced interspecific competition may play a role at high densities.     

Consequences of an amphibian malformity for development and fitness in complex environments

1. Environmental stressors have both lethal and sublethal effects, such as altered developmental rates and the induction of malformations. Ecological interactions, including predation and competition, often amplify such effects, for instance by inducing behavioural changes that increase susceptibility to the stress. 2. Using experimental mesocosms, we asked whether the density of conspecific competitors and predation risk from larval water beetles (Dytiscus spp.) affect the development of malformations in tadpoles of the wood frog (Rana sylvatica). We also examined whether such malformities increase the susceptibility of tadpoles to predation. 3. The risk of predation decreased the frequency of malformities in both low‐ and high‐density treatments, although this effect was greater at low density. Behavioural observations suggested that reductions in activity by amphibian larvae induced by predators mediated these responses by decreasing cumulative exposure to ultraviolet‐B radiation, the putative stressor causing the observed malformity. These results suggest that predators can reduce negative impacts of stressors by inducing behavioural changes in prey organisms. 4. Malformed individuals were twice as vulnerable to predators as non‐malformed individuals, suggesting that sublethal effects can ultimately cause increased mortality.     

Predictable changes in predation mortality as a consequence of changes in food availability and predation risk

Theory predicts that animals will have lower activity levels when either the risk of predation is high or the availability of resources in the environment is high. If encounter rates with predators are proportional to activity level, then we might expect predation mortality to be affected by resource availability and predator density independent of the number of effective predators. In a factorial experiment, we tested whether predation mortality of larval wood frogs, Rana sylvatica, caused by a single larval dragonfly, Anax junius, was affected by the presence of additional caged predators and elevated resource levels. Observations were consistent with predictions. The survival rate of the tadpoles increased when additional caged predators were present and when additional resources were provided. There was no significant interaction term between predator density and food concentration. Lower predation rates at higher predator density is a form of interference competition. Reduced activity of prey at higher predator density is a potential general mechanism for this widespread phenomenon. Higher predation rates at low food levels provides an indirect mechanism for density-dependent predation. When resources are depressed by elevated consumer densities, then the higher activity levels associated with low resource levels can lead to a positive association between consumer density and consumer mortality due to predation. These linkages between variation in behaviour and density-dependent processes argue that variation in behaviour may contribute to the dynamics of the populations. Because the capture rate of predators depends on the resources available to prey, the results also argue that models of food-web dynamics will have to incorporate adaptive variation in behaviour to make accurate predictions.     

Interactions among intraspecific competition, emergence patterns, and host selection behaviour in Ips pini (Coleoptera: Scolytinae)

Abstract.  1. Variations in developmental conditions of bark beetles, particularly intraspecific competition, can induce morphological and physiological modifications in the offspring. It is hypothesised that intraspecific competition could also affect host selection behaviour. Such behavioural changes might be manifested in response to host (alpha-pinene) or beetle (verbenone) compounds.
2.  Ips pini were bred at different densities. The offspring were measured for size, weight, and lipid concentration, and then subjected to tunnelling bioassays in agar media amended with varying amounts of alpha-pinene or verbenone.
3. High parental colonisation densities reduced emergence time of parents and offspring. Increasing colonisation density and emergence time had a negative influence on offspring vigour, resulting in a reduction of the distance tunnelled by the beetles.
4. Both alpha-pinene and verbenone were repellent. Surprisingly, verbenone was also toxic at high concentrations, with its effect being greater on beetles arising from high densities. The repellent effect of these compounds did not vary according to colonisation density, but for both compounds, at a 1.5 mg g^–1 concentration, it increased with emergence time.
5. The consequences of varying vigour and behaviour in relation to developmental conditions and emergence patterns on population dynamics of I. pini are discussed.     

Behaviourally mediated indirect effects: interference competition increases predation mortality in foraging redshanks

1. The effect of competition for a limiting resource on the population dynamics of competitors is usually assumed to operate directly through starvation, yet may also affect survival indirectly through behaviourally mediated effects that affect risk of predation. Thus, competition can affect more than two trophic levels, and we aim here to provide an example of this. 2. We show that the foraging success of redshanks Tringa totanus (L.) foraging on active prey was highest in the front of flocks, whereas this was not the case for redshanks foraging on inactive prey. Also, when foraging on active prey, foraging success in a flock decreased as more birds passed through a patch, while overall foraging success was not lower on subsequent visits to the same patch. Thus, redshanks foraging on active prey suffered from interference competition, whereas this was not the case for redshanks foraging on inactive prey. 3. This interference competition led to differences in activity: redshanks attaining a lower foraging success had a higher walking rate. Greater activity was associated with wider flock spacing and shorter distances to cover, which has previously been shown to increase predation risk and mortality from sparrowhawks Accipiter nisus (L.). 4. We conclude that behavioural adaptations of prey species can lead to interference competition in foraging redshanks, and thus can affect their predation risk and mortality through increased activity. This study is one of the first to show how interference competition can be a mechanism for behaviourally mediated indirect effects, and provides further evidence for the suggestion that a single species occupying an intermediate trophic level may be simultaneously top-down controlled by a predator and bottom-up controlled by a behavioural response of its prey.