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.     

An enemy of your enemy is your friend: Impact of predators on aggregation behavior of gammarids

Predation pressure may affect many aspects of prey behavior, including forming groups and changes in social interactions. We studied the aggregation behavior of competing gammarids Dikerogammarus villosus and Pontogammarus robustoides (Amphipoda, Crustacea) to check whether they modify their preferences for conspecifics or heterospecifics in response to predator (the racer goby Babka gymnotrachelus) kairomones in the presence or absence of stone shelters (alternative protection source). Both species exhibited preferences toward shelters occupied by conspecifics over empty shelters and conspecifics apart from shelters, suggesting that their aggregation depends not only on habitat heterogeneity, but also on their social interactions. Moreover, gammarids in the presence of shelters (safer conditions) preferred conspecifics over heterospecifics, but predator kairomones made them form aggregations irrespective of species. In the predator presence, P. robustoides increased its aggregation level only in the sheltered conditions, whereas D. villosus exhibited this response only in the absence of shelters, suggesting that this behavior can protect it against predators. Therefore, we tested the antipredator effectiveness of D. villosus aggregations by exposing them to fish predation. Gobies foraged most effectively on immobile single gammarids compared to moving and aggregated individuals. Fish also avoided aggregated prey, confirming the protective character of aggregations. We have demonstrated that the predator presence increases aggregation level of prey gammarids and affects their social behavior by reducing antagonistic interactions and avoidance between competing species. This is likely to affect their distribution and functioning in the wild, where predator pressure is a standard situation.     

The effects of habitat quality and female size on the productivity of the lesser spotted eagle Aquila pomarina in the light of the alternative prey hypothesis

Habitat quality is an important but insufficiently understood concept in ecology and conservation biology, due to geographic and temporal variation as well as interaction with individual quality. In 1994–2002, we studied the Estonian population of the lesser spotted eagle Aquila pomarina in order to (1) explore the relative contributions of habitat and female size in reproductive success; (2) check for a switch to alternative prey in vole‐poor years and the relevant variation in annual habitat quality as confirmed in the common buzzard Buteo buteo in the same area. We measured five landscape variables, the number of neighbouring conspecifics and the relative size of the female according to large moulted feathers in 77 nesting territories, and related this to the eagles’ productivity in vole‐rich and vole‐poor years. Nesting lesser spotted eagles benefited from heterogeneous landscapes and suffered from the neighbourhood of conspecifics. There was no evidence that different‐sized females used different habitats. In general, female size was positively related to productivity in vole‐poor but not vole‐rich years, but in the presence of competitors, large size appeared to be disadvantageous. The mean annual productivity of the eagle was well correlated with that of the buzzard, both having peaks after every three years. In contrast to the buzzard, the share of voles in the eagle's diet and its habitat quality did not differ significantly between good and poor years. We concluded that despite a superficial ecological similarity to the buzzard, the lesser spotted eagle did not behave as predicted by the alternative prey hypothesis, but the study confirmed that annual variation in prey utilization and relative habitat quality are parts of the same functional response. Non‐switching to alternative prey may be related to a historical foraging strategy, used by the eagles before they spread to agricultural landscapes, since the current effects of body size strongly suggested food shortage in vole‐poor years.     

Influence of prey availability and conspecifics on patch quality for a cannibalistic forager: laboratory experiments with the wolf spider Schizocosa

 Because cannibals are potentially both predator and prey, the presence of conspecifics and alternative prey may act together to influence the rate at which cannibals prey upon each other or emigrate from a habitat patch. Wolf spiders (Lycosidae) are cannibalistic-generalist predators that hunt for prey with a sit-and-wait strategy characterized by changes in foraging site. Little information is available on how both prey abundance and the presence of conspecifics influence patch quality for these cursorial, non-web-building spiders. To address this question, laboratory experiments were conducted with spiderlings and older juveniles of the lycosid genus Schizocosa. The presence of insect prey consistently reduced rates of spider emigration when spiders were housed either alone or in groups. Solitary juvenile Schizocosa that had been recently collected from the field exhibited a median giving-up time (GUT) of 10 h in the absence of prey (Collembola); providing Collembola increased the median GUT to 64 h. For solitary spiders, the absence of prey increased by about fourfold the rate of emigration during the first 24 h. In contrast, for spiders in patches with a high density of conspecifics, the absence of prey increased the 24-h emigration rate by only 1.6-fold. For successful cannibals in the no-prey patches, the presence of conspecifics improved patch quality by providing a source of food. Mortality by cannibalism was affected by both prey availability and openness of the patch to net emigration. In patches with no net emigration, the presence of prey reduced rates of cannibalism from 79% to 57%. Spiders in patches open to emigration but not immigration experienced a rate of cannibalism (16%) that was independent of prey availability. The results of these experiments indicate that for a cannibalistic forager such as the wolf spider Schizocosa, (1) the presence of conspecifics can improve average patch quality when prey are absent, and (2) cannibalism has the potential to be a significant mortality factor under natural field conditions because cannibalism persisted in prey patches that were open to emigration. Received: 12 April 1996 / Accepted: 14 August 1996     

Do anuran larvae respond behaviourally to chemical cues from an invasive crayfish predator? A community-wide study

Antipredator behaviour is an important fitness component in most animals. A co-evolutionary history between predator and prey is important for prey to respond adaptively to predation threats. When non-native predator species invade new areas, native prey may not recognise them or may lack effective antipredator defences. However, responses to novel predators can be facilitated by chemical cues from the predators’ diet. The red swamp crayfish Procambarus clarkii is a widespread invasive predator in the Southwest of the Iberian Peninsula, where it preys upon native anuran tadpoles. In a laboratory experiment we studied behavioural antipredator defences (alterations in activity level and spatial avoidance of predator) of nine anurans in response to P. clarkii chemical cues, and compared them with the defences towards a native predator, the larval dragonfly Aeshna sp. To investigate how chemical cues from consumed conspecifics shape the responses, we raised tadpoles with either a tadpole-fed or starved crayfish, or dragonfly larva, or in the absence of a predator. Five species significantly altered their behaviour in the presence of crayfish, and this was largely mediated by chemical cues from consumed conspecifics. In the presence of dragonflies, most species exhibited behavioural defences and often these did not require the presence of cues from predation events. Responding to cues from consumed conspecifics seems to be a critical factor in facilitating certain behavioural responses to novel exotic predators. This finding can be useful for predicting antipredator responses to invasive predators and help directing conservation efforts to the species at highest risk.     

Web-building management in an orb-weaving spider, Zygiella x-notata: influence of prey and conspecifics

According to optimal foraging theory, spiders should adapt their web building to environmental variations. Until now, there was no data on the influence of simultaneous information coming from different environmental factors on web building behaviour. Under laboratory conditions, we studied the behaviour of Zygiella x-notata in the presence of prey, conspecifics, or both simultaneously. There was a stimulating effect of prey, but web building was not affected by the presence of conspecifics. When spiders and prey were present simultaneously, the effect was similar to that of prey alone; it seemed that there was no interactive influence of both factors. We discussed about the use of environmental information by spiders in foraging behaviour.     

Landscape structure influences the use of social information in an insectivorous bat

In anthropogenic landscapes, aerial insectivores are often confronted with variable habitat complexity, which may influence the distribution of prey. Yet, high mobility may allow aerial insectivores to adjust their foraging strategy to different prey distributions. We investigated whether aerial-hunting common noctules Nyctalus noctula adjust their foraging strategy to landscapes with different habitat complexity and assumingly different prey distribution. We hypothesized that the movement behaviour of hunting common noctules and changes of movement behaviour in reaction towards conspecifics would depend on whether they hunt in a structurally poor cropland dominated landscape or a structurally rich forest dominated landscape. We tracked flight paths of common noctules in northeastern Germany using GPS loggers equipped with an ultrasonic microphone that recorded foraging events and presence of conspecifics. Above cropland, common noctules hunted mainly during bouts of highly tortuous and area restricted movements (ARM). Bats switched from straight flight to ARM after encountering conspecifics. In the forested landscape, common noctules hunted both during ARM and during straight flights. The onset of ARM did not correlate with the presence of conspecifics. Common noctules showed a lower feeding rate and encountered more conspecifics above the forested than above the cropland dominated landscape. We conjecture that prey distribution above cropland was patchy and unpredictable, thus making eavesdropping on hunting conspecifics crucial for bats during search for prey patches. In contrast, small scale structural diversity of the forested landscape possibly led to a more homogeneous prey distribution at the landscape scale, thus enabling bats to find sufficient food independent of conspecific presence. This suggests that predators depending on ephemeral prey can increase their foraging success in structurally poor landscapes by using social information provided by conspecifics. Hence, a minimum population density might be obligatory to enable successful foraging in simplified landscapes.     

Additive multiple predator effects can reduce mosquito populations

1. Multiple predator interactions may profoundly alter ecological community dynamics and can complicate predictions of simpler pairwise predator–prey interaction strengths. In particular, multiple predator effects may lessen or enhance prey risk, with implications for community-level stability. Such emergent effects may modulate natural enemy efficacy towards target organisms. 2. In the present study, a functional response approach was used to quantify emergent multiple predator effects among natural enemies towards the disease vector mosquito complex, Culex pipiens. Conspecific multiple predator–predator interactions of the cyclopoid copepod Macrocyclops albidus (intermediate predator) were quantified by comparing multiple predator consumption simulations, based on individual consumption rates, with multiple predator consumption rates that were experimentally observed. Further, the study examined the influence of the presence of a predator at a higher trophic level, Chaoborus flavicans, on copepod group predation. 3. Both predators displayed type II functional responses, with C. flavicans consuming significantly more prey than M. albidus individually. Overall consumption levels of mosquitoes increased with greater predator density and richness. Antagonistic or synergistic emergent multiple predator effects between conspecifics of M. albidus were not detected, and the higher-level predator did not reduce effects of the intermediate predator. Accordingly, evidence for additive multiple predator interactions was found. 4. The lack of predator–predator interference between cyclopoid copepods and larval chaoborid midges provides strong support for their combined application in mosquito biocontrol. It is proposed that there should be increased examination of multiple predator effects in assessments of natural enemy efficacies to better understand overall predatory effects within communities and utilities in vector control.     

Influence of visual conditions on foraging and growth of juvenile fishes with dissimilar sensory physiology

The influence of turbidity on foraging and growth of young-of-the-year (YOY) of two percid species with relatively dissimilar sensory physiology, perch Perca fluviatilis and pikeperch Sander lucioperca was tested. A littoral mysid, Neomysis integer , was used as prey. Functional response was investigated indoors at two levels of turbidity, 3 NTU (clear) and 25 NTU (turbid). Growth and consumption rate were studied both during day and night in outdoor experiments at similar turbidity levels. Pikeperch were not affected by turbid conditions in either of the experiments. Foraging of perch, on the other hand, was significantly negatively affected by higher turbidity in the functional response experiments and during the night-time in the outdoor experiments. As opposed to pikeperch, consumption rates of perch decreased markedly during nights. Perch also grew more slowly in the treatments with turbid water. The dissimilar reactions of the two species indicate that sensory physiological adaptations and foraging behaviour are important factors that partly can explain disparate reactions of YOY fishes to the level of turbidity.     

Effect of prey size on attack components of the functional response by Notonecta undulata

The number of encounters per prey, the proportion of encounters resulting in attacks, and the proportion of attacks that were successful were observed while fourth-instar Notonecta undulata nymphs preyed on smaller N. undulata nymphs. While encounters per prey and proportion of encounters resulting in attacks increased with prey size, the proportion of attacks that were successful decreased. The increase in encounter rate per prey was due in part to an increase in the predator's reactive distance to prey as prey size increased. While none of the attack parameters varied significantly with prey density, logarithmic regression of the number of encounters per unit search time on prey density suggested that prey density tends to have a positive effect on encounters per first-instar prey but a negative effect on encounters per second-instar prey. A functional response model is presented that incorporates components of the predator's attack rate as exponential functions of prey density and allows for effects of the time the predator may spend evaluating prey encountered but not attacked and time spent attacking prey not captured. Estimates of the attack parameters derived from the experimental data are used in the model to generate functional response curves for fourth-instar N. undulata preying on first- or second-instar conspecifics. The predicted curve for second-instar prey is typical type II but the curve for firstinstar prey is slightly positively density dependent at low prey densities, i.e., type III.     

Suspended sediment alters predator–prey interactions between two coral reef fishes

Sediment derived from agriculture and development increases water turbidity and threatens the health of inshore coral reefs. In this study, we examined whether suspended sediment could change predation patterns through a reduction in visual cues. We measured survivorship of newly settled Chromis atripectoralis exposed to Pseudochromis fuscus, a common predator of juvenile damselfishes, in aquaria with one of four turbidity levels. Increased turbidity led to a nonlinear response in predation patterns. Predator-induced mortality was ~50 % in the control and low turbidity level, but exhibited a substantial increase in the medium level. In the highest turbidity level, predation rates declined to the level seen in the control. These results suggest an imbalance in how the predator and prey cope with turbidity. A turbidity-induced change to the outcome of predator–prey interactions represents a major change to the fundamental processes that regulate fish assemblages.     

Discrimination of Variation in a Male Signaling Trait Affects Detection Time in Visual Predators

In many species, expression of elaborate male characteristics likely represents a balance between sexual selection and natural selection via predation, as male traits selected to elicit rapid detection or response on the part of females also increase detection by predators. Predation costs are frequently inferred, but the underlying mechanisms associated with specific traits have rarely been directly explored. Males of the wolf spider Schizocosa ocreata (Araneae: Lycosidae) exhibit a sexually selected signaling trait (dark tufts of bristles) on their forelegs, and are sympatric with a number of visually hunting generalist predators, including cannibalistic conspecifics, that may impact spider survival. Here we use latency of orientation response of the American toad, Bufo americanus (Anura: Bufonidae), to video, 'virtual' courting S. ocreata male stimuli as an index of predator detection, and latency of orientation response of female S. ocreata to the same stimuli as an index of conspecific detection. When compared with stimuli representing the population average, elimination of the signal trait had no significant effect on predator detection but did increase latency to orient in conspecifics. Increasing the size of the signal trait had no effect on conspecific detection but did significantly reduce latency to orient for predatory toads. Results clearly indicate that for a courting male spider of a given size and vigor level, variation in the expression of a secondary sexual characteristic alone can incur differential direct costs and benefits by influencing latency of orientation to visual signals by predators and conspecifics.     

The feeding success of cattle egrets in flocks

The feeding rates of grouped (<1.5 m from conspecifics) and solo (>5 m from conspecifics) cattle egrets (Bubulcus ibis) in loose flocks away from cows were compared, to test the hypothesis that grouped cattle egrets benefit from feeding on prey flushed inadvertently by nearby conspecifics. The flock feeding rates were also compared to those of grouped and solo egrets near cows, to determine the effects of flock membership on feeding rates. Birds in flocks captured prey faster than those with cows, and tended to capture larger prey, but field observations and captive experiments failed to show that the feeding success of flock members was enhanced by the hypothesized ‘beater’ effect. Increases in prey density, however, always resulted in higher feeding rates, so some cattle egret groups may form in response to local concentrations of prey. Prey size may also play a role in group formation, because birds in the field tended to feed at greater distances from their neighbours when larger prey were captured, regardless of prey density. When small groups did form among cattle egrets feeding on relatively large prey, group members occasionally captured prey items that had been discovered by nearby conspecifics. This behaviour was not observed among birds in dense aggregations, which fed on small, highly abundant prey. These data indicate that there is a potential cost associated with feeding too near others unless the prey are relatively small and abundant.     

The Effect of Refuge on Dermestes ater (Coleoptera: Dermestidae) Predation on Musca domestica (Diptera: Muscidae): Refuge for Prey or the Predator?

It is believed that habitat heterogeneity can change the extent of predator-prey interactions. Therefore, in this study we examined the effect of habitat heterogeneity (characterized here as an addition of refuge) on D. ater predation on M. domestica. Predation of D. ater on M. domestica larvae was carried out in experimental habitats with and without refuge, and examined at different prey densities. The number of prey eaten by beetles over 24 h of predator-prey interaction was recorded, and we investigated the strength of interaction between prey and predator in both experimental habitats by determining predator functional response. The mean number of prey eaten by beetles in the presence of refuge was significantly higher than in the absence of refuge. Females had greater weight gains than males. Logistic regression analyses revealed the type II functional response for both experimental habitats, even though data did not fit well into the random predator model. Results suggest that the addition of refuge in fact enhanced predation, as prey consumption increased in the presence of refuge. Predators kept in the presence of refuge also consumed more prey at high prey densities. Thus, we concluded that the addition of refuge was an important component mediating D. ater-M. domestica population interactions. Refuge actually acted as a refuge for predators from prey, since prey behaviors detrimental to predators were reduced in this case.     

Presence of conspecifics triggers host searching behavior in an egg parasitoid wasp

Parasitoids and predators compete for host or prey species. The efficiency of obtaining prey or host items is reduced by intraspecific competition. As the optimal search behavior depends on the intensity of competition, it is important for the parasitoid or predator to obtain information on this intensity. Previous studies have shown that parasitoids can obtain information regarding competition from encounters with already parasitized hosts. They then change their host searching behavior accordingly. However, whether parasitoids obtain the information directly from observing the presence of conspecifics remains unclear. We used Tiphodytes gerriphagus (Marchal) (Hymenoptera: Scelionidae), the solitary egg parasitoid of water striders, for testing the effect of density of conspecifics on host searching behavior. Females of T. gerriphagus dive into the water to search for hosts and sometimes they dive without hosts present. Thus, we investigated whether T. gerriphagus changed underwater activities in response to the density of conspecifics in the absence of hosts. Four densities (1, 2, 4, and 8 female T. gerriphagus) were investigated. Females in competitive situations (groups of 2, 4, and 8 females) displayed host searching behavior, but the solitary females did not. This indicates that the presence of conspecifics triggers host searching behavior and that T. gerriphagus females obtain information on competition directly from conspecifics and use it for modifying their behavior.     

Antipredator behaviour of 0+ year Perca fluviatilis: effect of vegetation density and turbidity

In this study, the combined influence of vegetation density and water turbidity on habitat utilization of a prey fish, 0+ year perch Perca fluviatilis , under predation risk (pike, Esox lucius ) was investigated. The vegetated habitat was overall preferred over the open habitat in the presence of a predator. The level of turbidity, and to a lesser extent vegetation density, however, influenced the response of 0+ year perch. The use of the vegetated habitat was lower in very turbid than in clear and turbid conditions, suggesting reduced antipredator behaviour in very turbid water. The effect of vegetation density on antipredator behaviour was only present in clear water, where the use of a structural refuge decreased with increasing vegetation density. No such effect was observed in turbid and very turbid water. The results showed that the structuring role of vegetation or habitat complexity may diminish with increased turbidity. The observed masking effect of turbidity suggests that predator‐prey interactions in vegetated habitats are more complex than what has generally been thought.     

Predation risk cues associated with killed conspecifics affect the behavior and reproduction of prey animals

Organisms often perceive risk to be attacked by natural enemies through cues that accompany or persist after attacks on conspecifics. Under the risk of attack, preys may prioritize anti-enemy behaviours, sacrificing their feeding activities. These enemy-induced changes can affect the prey population growth. The presence of killed congeners is a past attack events cue but, in terrestrial prey–enemy systems, its effect on prey population biology is an open question. This was addressed here by studying both the space use and reproductive effort of aphids exposed to congeners killed by parasitoids. To test whether preys adjust their responses with the threat intensity, aphids' responses in presence of killed conspecifics were compared to those measured when directly exposed to non-consumptive parasitoids. The results showed that killed congeners are perceived by aphids and led to both changes in their space use and a strong decline in their population size. This experiment is the first to demonstrate that the presence of congeners killed by an enemy can contribute to the suppressive effects of natural enemies on prey populations.     

Consequences of refuge for the functional response of Dermestes ater (Coleoptera: Dermestidae) to Musca domestica (Diptera: Muscidae)

It is well known that a predator has the potential to regulate a prey population only if the predator responds to increases in prey density and inflicts greater mortality rates. Predators may cause such density-dependent mortality depending on the nature of the functional and numerical responses. Yet, few studies have examined the relationship between the addition of refuges and the characteristic of functional response fits. We investigated whether addition of a refuge changed the type of functional response exhibited by Dermestes ater on Musca domestica, comparing the inherent ability of D. ater to kill houseflies in the absence and in the presence of refuge. An additional laboratory experiment was also carried out to assess handling and searching times exhibited by D. ater. Logistic regression analyses revealed a type III functional response for predator–prey interaction without refuge, and results were described by the random predator equation. The mean number of prey killed did not differ between experimental habitats, indicating that the addition of refuge did not inhibit predation. However, predators that interacted with prey without refuge spent less time searching for prey at higher densities, increasing predatory interaction. We concluded that this interaction may be weak, because data from experiments with refuge fitted poorly to models. However, the high variability and the nonsignificance of the data from the experiment with refuge show the importance of refuge for promoting spatial heterogeneity, which may prevent prey extinction.     

Antipredator Behavior in Desmognathus ochrophaeus: Threat‐Specific Responses to Chemical Stimuli in a Foraging Context

Prey species may reduce the likelihood of injury or death by engaging in defensive behavior but often incur costs related to decreased foraging success or efficiency. To lessen these costs, prey may adjust the intensity or type of antipredator behavior according to the nature of the perceived threat. We evaluated the potential for threat‐sensitive responses by Allegheny Mountain dusky salamanders (Desmognathus ochrophaeus) exposed to chemical stimuli associated with predation by asking three questions: (1) Do individual D. ochrophaeus respond to chemical cues in a threat‐sensitive manner? (2) Do salamanders exhibit the same pattern of behavioral response while foraging? and (3) Is foraging efficiency reduced when focal individuals are exposed to stimuli from predators or predation events? In our first experiment, we evaluated salamander chemosensory movements (nose‐taps), locomotor activity (steps), and edge behavior in response to chemical stimuli from disturbed and injured conspecifics as well as predatory Gyrinophilus porphyriticus and found that individual D. ochrophaeus show a significant graded increase in nose‐taps when exposed to cues from conspecifics and a reduction in activity when exposed to the predator. In our second experiment, we again observed salamander responses to the same chemical stimuli but in this instance added five Drosophila prey to the test dishes. We found that salamanders exhibited a similar pattern of response to the chemical stimuli in the presence of prey, showing a graded increase in nose‐taps to cues from conspecifics and a reduction in activity when exposed to the predator. However, foraging efficiency (i.e. the proportion of successful strikes) did not vary significantly among treatments. Our data show that individual D. ochrophaeus detect and differentially respond to chemical stimuli associated with predation, but do not significantly reduce foraging efficiency. Overall, the type and relative intensity of these responses is largely unaffected by the presence of potential prey.     

A laboratory study of predator-prey interactions of stoneflies and mayflies

SUMMARY. 1. Reactions of individual predatory stoneflies to contact by individual mayfly prey, and vice versa, were studied in the laboratory using two sets of species, one from a stream in southern Ontario. Canada, and the other from a stream in North Wales. 2. Based largely on information received via their antennae, the stoneflies were capable of distinguishing between conspecifics and mayfly prey and between prey species. A clear preference for certain prey species was evident but less preferred species were made more attractive when damaged or when the stoneflies were very hungry. Detection of conspecifics by stoneflies was confused in the presence of prey or the local smell of prey body fluids and this led to interference during hunting and feeding at high predator densities. 3. Some mayfly species were capable of distinguishing between conspecifics. other mayflies and predators. The cerci were important in intra- and interspecific contact encounters between mayflies and were used in a fencing fashion to establish a minimum distance between nymphs. However, the cerci did not appear to play an anti-predation role as stoneflies captured and ate mayflies with and without cerci in equal numbers. 4. Reactions to conspecifics by the Palaearetie mayflies Rhithrogena semicolorata (Curt.) and Baetis rhodani (Pictet) were similar to those of the ecologically-equivalent Nearctic species Stenonema vicarium (Walker) and Baetis vagans McDunnough. 5. In encounters between mayflies, intra- and interspecific responses were similar. However, the responses of some mayfly species to conspeeifics were modified in the physical presence of a predator or the water from its holding tank. Different mayfly species made specific responses to predaeeous stoneflies. Responses to a herbivorous stonefly were either similar to conspeeific encounters or were intermediate between them and encounters with predators. 6. Experiments to reveal the exact nature of the stonefly stimulus detected by the mayfly nymphs strongly suggested that there were two components to the stimulus. One was physical contact by the predator's antennae, the other was chemical in nature and could be detected only over short distances (up to a few mm). Independently each could elicit a response, but combined their effect appeared to be synergistic. 7. Different mayfly species responded differently to the stonefly Phasganophora capitata (Pictet) and this sequence of sensitivity was mirrored by the predator's choice of prey species—the most sensitive to the predator being the favoured prey and the less sensitive the least favoured. The significance of these findings to optimal foraging theory, as applied to tactile predators, is discussed.