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.     

Behavioral responses to prey density by three acarine predator species with different degrees of polyphagy

Behavioral responses by three acarine predators, Phytoseiulus persimilis, Typhlodromus occidentalis, and Amblyseius andersoni (Acari: Phytoseiidae), to different egg and webbing densities of the spider mite Tetranychus urticae (Acari: Tetranychidae) on rose leaflets were studied in the laboratory. Prey patches were delineated by T. urticae webbing and associated kairomones, which elicit turning back responses in predators near the patch edge. Only the presence of webbing affected predator behavior; increased webbing density did not increase patch time. Patch time increased with increased T. urticae egg density in the oligophagous P. persimilis, but was density independent in the polyphagous species T. occidentalis and A. andersoni. Patch time in all three species was more strongly correlated with the number of prey encounters and attacks than with the actual prey number present in the patch. Patch time was determined by (a) the turning back response near the patch edge; this response decayed through time and eventually led to the abandonment of the patch, and (b) encounters with, and attacks upon, prey eggs; these prolonged patch time by both an increment of time spent in handling or rejecting prey and an increment of time spent searching between two successive prey encounters or attacks. Although searching efficiency was independent of prey density in all three species, the predation rate by P. persimilis decreased with prey density because its searching activity (i.e. proportion of total patch time spent in searching) decreased with prey density. Predation rates by T. occidentalis and A. andersoni decreased with prey density because their searching activity and success ratio both decreased with prey density. The data were tested against models of predator foraging responses to prey density. The effects of the degree of polyphagy on predator foraging behavior were also discussed.     

Effects of opportunistic predation on anti-predator behavioural responses in a guild of ground foragers

We studied factors that affect prey selection by a generalist predator that opportunistically attacks prey species, and the associated inter- and intra-specific responses of prey to this type of predation. Our model system was a guild of ground-foraging birds that are preyed upon by magpies (Pica pica) during the breeding season. We found that magpies attacked up to 12 species during three consecutive breeding seasons. The overall capture success was estimated to be 4.9%. Magpies tended to attack from the air, targeting solitary prey, either on the ground or flying. Inter-specific prey responses to the risk of magpie predation included a reduction in the mean number of species occupying a foraging patch when magpies were present and a decrease in the distance between heterospecific neighbours. Intra-specific responses to magpie predation varied between species that were subject to different attack rates. Preferentially attacked prey enhanced their risk responses (increase in scanning time and scanning rate in the presence of magpies) relative to those species attacked in proportion to their abundance (increase only in scanning rate with magpies). Species attacked infrequently, relative to their abundance, showed no antipredator response. The probability of being attacked, rather than mortality rate, appears to be the factor to which prey species respond.     

Behavioural and developmental responses of predatory coral reef fish to variation in the abundance of prey

Ecological theory suggests that the behaviour, growth and abundance of predators will be strongly influenced by the abundance of prey. Predators may in turn play an important role in structuring prey populations and communities. Responses of predators to variation in prey abundance have most commonly been demonstrated in low-diversity communities where food webs are relatively simple. How predators respond in highly diverse assemblages such as in coral reef habitats is largely unknown. This study describes an experiment that examined how the movement, diet and growth of the coral reef piscivore, Cephalopholis boenak (Serranidae) responded to variation in the abundance of its prey. Predator densities were standardised on small patch reefs made from the lagoonal reef-building coral, Porites cylindrica. These patch reefs exhibited natural variation in the abundance and community structure of multiple species of prey. However, our experiment generated a relatively simple predator–prey relationship, with C. boenak primarily responding to the most abundant species of prey. Three responses of predators were observed: aggregative, functional and developmental. Thirty-one per cent of individuals moved between patch reefs during the experiment, all from areas of relatively low to high prey density. Feeding rates were higher on patch reefs of high prey density, while growth rates of fish that remained on low prey density reefs throughout the experiment were lower. Growth rates of C. boenak on the experimental reefs were also much higher than for those living on natural patch reefs over the same time period, corresponding with overall differences in prey abundance. These results suggest that local abundance, feeding rate and growth of C. boenak were closely linked to the abundance of their main prey. This combination of predatory responses is a potential mechanism behind recent observations of density-dependent mortality and population regulation of prey in coral reef fish communities.     

The influence of prey abundance on the feeding ecology of two piscivorous species of coral reef fish

Despite the potential importance of predation as a process structuring coral reef fish communities, few studies have examined how the diet of piscivorous fish responds to fluctuations in the abundance of their prey. This study focused on two species of rock-cod, Cephalopholis cyanostigma (Valenciennes, 1828) and Cephalopholis boenak (Bloch, 1790) (Serranidae), and monitored their diet in two different habitats (patch and contiguous reef) at Lizard Island on the northern Great Barrier Reef, Australia, over a 2-year period. The abundance of the rock-cods and the abundance and family composition of their prey were monitored at the same time. Dietary information was largely collected from regurgitated samples, which represented approximately 60% of the prey consumed and were unbiased in composition. A laboratory experiment showed that fish were digested approximately four times faster than crustaceans, leading to gross overestimation of the importance of crustaceans in the diet. When this was taken into account fish were found to make up over 90% of the diet of both species. Prey fish of the family Apogonidae, followed by Pomacentridae and Clupeidae, dominated the diet of both species of rock-cod. The interacting effect of fluctuations in prey abundance and patterns of prey selection caused dietary composition to vary both temporally and spatially. Mid-water schooling prey belonging to the families Clupeidae and to a lesser extent Caesionidae were selected for over other families. In the absence of these types of prey, apogonids were selected for over the more reef-associated pomacentrids. A laboratory experiment supported the hypothesis that such patterns were mainly due to prey behaviour. Feeding rates of both species of rock-cod were much higher in summer than in winter, and in summer they concentrated on small recruit sized fish. However, there was little variation in feeding rates between habitats, despite apparent differences in prey abundance. In summary, our observations of how the feeding ecology of predatory fish responded to variation in prey abundance provide potential mechanisms for how predation may affect the community structure of coral reef fishes.     

Prey preference in stoneflies: a comparative analysis of prey vulnerability

Summary Laboratory feeding trials were conducted with the predaceous stonefly Hesperoperla pacifica and a number of mayfly and dipteran prey species to investigate the effects of predator size, and prey size and morphology, on the predator's success. Observations under dim red light permitted estimation of encounter rate (E/min), attack propensity (A/E), capture success (C/A) and handling time (HT). For prey of a particular species and size, HT decreased log-linearly with increasing predator size. Across all prey categories, HT increased log-linearly with increasing values of the ratio prey dry wt/predator dry wt, and differences among species appeared to be small. Overall, capture success was low, but C/A was higher for dipterans than for mayflies, especially with large H. pacifica. Predator size affected C/A when prey fell within a certain size range, but was not a detectable influence with very small or very large prey. Values of A/E of near 10% typified many predatorprey combinations; however, ephemerellid mayflies suffered markedly fewer attacks, and values of A/E up to 30% were obtained with some species-size combinations. We estimated benefit to the predator first as prey wt ingested per unit time (dry wt/HT), and second by mutliplying the former term by capture success. Values increased with increasing size of the predator, and inclusion of the C/A term indicated that predators would obtain greater reward from small relative to large prey, and from dipterans relative to mayflies. Howerver, there was little evidence that attacks were biased toward more profitable prey. We compare the relative contributions of E/min, A/E and C/A to prey choice, and discuss their applicability to predation events in nature.     

Prey choice by marten during a decline in prey abundance

Summary We examined variation in diet choice by marten (Martes americana) among seasons and between sexes and ages from 1980–1985. During this period prey populations crashed simultaneously, except for ruffed grouse (Bonasa umbellus) which was common at the beginning and end of the study, and masked shrews (Sorex cinereus) which were abundant in 1983. Marten were catholic in selection of prey and made use of most available mammalian prey, ruffed grouse, passerine birds, berries, and insects. Diet niche was widest during the latter three years when prey was scare, particularly in late winter. Diet niche breadth was negatively correlated with abundance of all common prey species. Proportion of small prey species in the diet was correlated with absolute abundance of those species, but proportion of some large prey was related to their relative abundance. Diet choice varied among years and among seasons. Berries and insects were common in summer diets while large prey, particularly varying hare (Lepus americanus), were more frequent in winter diet than in summer diet. We found little evidence that any small mammal species was a preferred prey. Sexual size dimorphism between the sexes did not affect prey choice, nor did age. Reduced foraging effort in winter resulted in a wider diet niche only when prey was scarce. The only prediction of optimal foraging models fully supported by our data was a wider diet niche with reduced prey abundance. However, among the three most profitable prey species choice was dependent on the absolute abundance of the most profitable type (varying hare). We suggest that marten primarily forage for large prey but employ a strategy which results in encounters with small prey as well. These small prey are eaten as they provide energy at minimal cost, between captures of large prey.     

Prey susceptibilities,prey utilization and variable attack efficiencies of Ural owls

Summary To investigate the factors that influence prey utilization among predators with active prey, three series of experiments were performed in which Ural owls (Strix uralensis) searched for and attacked three prey species of wild mice, Microtus montebelli, Apodemus speciosus, and A. argenteus, in a large flight cage. Over the whole study, owls attacked mice about ten times a night. The number of attacks on each prey species did not differ from that predicted by a random attack model. M. montebelli was taken more than either Apodemus species. Prey utilization appears to be influenced by prey susceptibilities only and it is unlikely that prey selection by the owls affected prey utilization patterns. Under the experimental conditions, random attack is predicted by optimal foraging theory. However, random attack may be explained just as well by the inability of the owl to discriminate prey type. The owls, energy gain was adjusted not by alteration in the number of attacks on a prey species but rather by alteration in the capture success between experiments. Capture success increased in poor food conditions for the same prey species. This flexibility in capture success has not been considered in the assumptions of optimal foraging theory. In conventional optimal foraging theory, the probability of capture success is implicitly assumed as constant and unity. We suggest that this assumption is inadequate to understand the foraging behavior of owls.     

Dynamic changes in prey choice by stickleback during simultaneous encounter with large prey

When a three-spined stickleback Gasterosteus aculeatus encountered prey simultaneously the probability of hanging and the median pursuit time were greater than when prey were encountered sequentially. During simultaneous prey encounter fish did not choose to attack the more profitable prey but instead the nearer prey was handled first except when the difference between the two prey sizes was large. No difference was found in the level of total energetic intake by the fish regardless of prey size pairing. Fish that handled and ate the first prey of a pair in <5 s attacked the second prey with a high probability of success, demonstrating an opportunistic feeding strategy. Importantly however, the fish did not choose to maximize long term energy intake rate by eating both prey, but rather short-term considerations over the course of feeding took precedence. With an empty stomach, the probability of a fish eating ( P _eat) the first prey handled was high regardless of prey size. As stomach fullness increased, the P _eat the first prey handled decreased if it was the larger prey. Hence, the fish were unselective when the stomach was empty but thereafter there was a shift in preference towards the smaller prey. The decision of which prey to attack and eat appeared to be based on short-term energy considerations and the level of stomach fullness. This study demonstrates that feeding on a short-term scale is a crucial factor to take account of when analysing fish feeding during simultaneous prey encounter.     

Optimal Bayesian foraging policies and prey population dynamics-some comments on Rodriguez-Girones and Vasquez

In this paper we show the density-dependent harvest rates of optimal Bayesian foragers exploiting prey occurring with clumped spatial distribution. Rodríguez-Gironés and Vásquez (1997) recently treated the issue, but they used a patch-leaving rule (current value assessment rule) that is not optimal for the case described here. An optimal Bayesian forager exploiting prey whose distribution follows the negative binomial distribution should leave a patch when the potential (and not instantaneous) gain rate in that patch equals the best long-term gain rate in the environment (potential value assessment rule). It follows that the instantaneous gain rate at which the patches are abandoned is an increasing function of the time spent searching in the patch. It also follows that the proportion of prey harvested in a patch is an increasing sigmoidal function of the number of prey initially present. In this paper we vary several parameters of the model to evaluate the effects on the forager's intake rate, the proportion of prey harvested per patch, and the prey's average mortality rate in the environment. In each case, we study an intake rate maximizing forager's optimal response to the parameter changes. For the potential value assessment rule we find that at a higher average prey density in the environment, a lower proportion of the prey is taken in a patch with a given initial prey density. The proportion of prey taken in a patch of a given prey density also decreases when the variance of the prey density distribution is increased and if the travel time between patches is reduced. We also evaluate the effect of using predation minimization, rather than rate maximization, as the currency. Then a higher proportion of the prey is taken for each given initial prey density. This is related to the assumption that traveling between patches is the most risky activity. Compared to the optimal potential value assessment rule, the current value assessment rule performs worse, in terms of long-term intake rate achieved. The difference in performance is amplified when prey density is high or highly aggregated. These results pertain to the foraging patch spatial scale and may have consequences for the spatial distribution of prey in the environment.     

Habitat destruction in a simple predator-prey patch model: how predators enhance prey persistence and abundance

We model a metapopulation of predator-prey patches using both spatially implicit or mean-field (MF) and spatially explicit (SE) approaches. We show that in the MF model there are parameter regimes for which prey cannot persist in the absence of predators, but can in their presence. In addition, there are parameter regimes for which prey may persist in isolation, but the presence of predators will increase prey patch density. Predators may thus enhance prey persistence and overall abundance. The key mechanism responsible for this effect is the occurrence of prey dispersal from patches that are occupied by both prey and predators. In addition, these patches should be either long-lived, such as that occurs when predators keep prey from overexploiting its local resource, or the presence of a predator on a patch should significantly enhance the prey dispersal out of that patch. In the SE approach these positive effects of predators on prey persistence and abundance occur for even larger parameter ranges than in the MF model. Prey dispersal from predator-prey patches may thus be important for persistence of both species as a community, independent of the modeling framework studied. Comparison of the MF and SE approaches shows that local dispersal constraints can have the edge over global dispersal for the persistence of the metapopulation in regimes where the two species have a beneficial effect on each other. In general, our model provides an example of feedback in multiple-species metapopulations that can make the implementation of conservation schemes based on single-species arguments very risky.     

Local prey vulnerability increases with multiple attacks by a predator

Theory and empirical evidence suggest that predator activity makes prey more wary and less vulnerable to predation. However if at least some prey in the population are energetically or spatially constrained, then predators may eventually increase local prey vulnerability because of the cumulative costs of anti‐predation behaviour. We tested whether repeated attacks by a predator might increase prey vulnerability in a system where redshanks on a saltmarsh are attacked regularly by sparrowhawks from adjacent woodland. Cumulative attack number led to a reduction in redshank numbers and flock size (but had no effect on how close redshanks fed to predator‐concealing cover) because some redshanks moved to safer but less profitable habitats, leaving smaller flocks on the saltmarsh. This effect held even though numbers of redshank on the saltmarsh increased with time of day. As a result of the change in flock size, predicted attack‐success increased up to 1.6‐fold for the sparrowhawk, while individual risk of capture for the redshank increased up to 4.5‐fold among those individuals remaining on the saltmarsh. The effect did not arise simply because hawks were more likely to attack smaller flocks because attack rate was not dependent on flock size or abundance. Our data demonstrate that when some individual prey are constrained in their ability to feed on alternative, safer foraging sites, their vulnerability to predation increases as predator attacks accumulate, although those, presumably better quality individuals that leave the immediate risky area will have lower vulnerability, so that the mean vulnerability across the entire population may not have changed substantially. This suggests that the selective benefits of multiple low‐cost attacks by predators on prey could potentially lead to 1) locally heightened trait‐mediated interactions, 2) locally reduced interference among competing predators, and 3) the evolution of active prey manipulation by predators.     

Small-scale recruitment variation in a temperate fish: the roles of macrophytes and food supply

Synopsis The availability of reef-related resources, particularly food and shelter can play a significant role in determining the distribution and abundance of reef fishes. Much of the structure on temperate reefs is provided by macroalgae, and variability in the density of temperate reef fishes at large spatial scales (100's of meters) can often be explained by variation in macroalgal cover or density. In this study I investigated the role of macrophytes and associated food resources on the recruitment of a temperate fish, Tautogolabrus adspersus, at a small spatial scale (0.25 m²). No relationship between the density of new recruits and the percent cover of kelp, foliose or filamentous algae was observed. Multiple regressions revealed that less than 8% of variability in recruitment could be explained by variability in macroalgal cover. However, recruits were found in higher abundance in patches containing many functional forms of seaweeds than in patches dominated by a single form. A wide variety of prey were available for use by cunner recruits; however, crustaceans and mussels were the only common components of their diet, and crustaceans were clearly the most preferred prey. The prey composition in patches where fish were present was compared to randomly selected patches. Significantly greater numbers of isopods, amphipods and newly settled mussels were present in patches where fish were present than in randomly selected patches. The data presented in this study contradict previous work that has shown algal structure to be important in determining patterns of abundance and food supply to be of little significance. A conceptual model is proposed suggesting that settling fish select habitats in a hierarchical manner largely based on their dispersal tendencies. Hierarchical selection of habitats results in different attributes of the habitat being selected during different life-history intervals.     

Foraging behavior of an estuarine predator, the blue crab Callinectes sapidus in a patchy environment

To define general principles of predator‐prey dynamics in an estuarine subtidal environment, we manipulated predator density (the blue crab, Callinectes sapidus) and prey (the clam, Macoma balthica) patch distribution in large field enclosures in the Rhode River subestuary of the central Chesapeake Bay. The primary objectives were to determine whether predators forage in a way that maximizes prey consumption and to assess how their foraging success is affected by density of conspecifics. We developed a novel ultrasonic telemetry system to observe behavior of individual predators with unprecedented detail. Behavior of predators was more indicative of optimal than of opportunistic foraging. Predators appeared responsive to the overall quality of prey in their habitat. Rather than remaining on a prey patch until depletion, predators appeared to vary their patch use with quality of the surrounding environment. When multiple (two) prey patches were available, residence time of predators on a prey patch was shorter than when only a single prey patch was available. Predators seemed to move among the prey patches fairly regularly, dividing their foraging time between the patches and consuming prey from each of them at a similar rate. That predators more than doubled their consumption of prey when we doubled the number of prey (by adding the second patch) is consistent with optimizing behaviors ‐ rather than with an opportunistic increase in prey consumption brought about simply by the addition of more prey. Predators at high density, however, appeared to interfere with each other's foraging success, reflected by their lower rates of prey consumption. Blue crabs appear to forage more successfully (and their prey to experience higher mortality) in prey patches located within 15–20 meters of neighboring patch, than in isolated patches. Our results are likely to apply, at least qualitatively, to other crustacean‐bivalve interactions, including those of commercial interest; their quantitative applicability will depend on the mobility of other predators and the scale of patchiness they perceive.     

Relationship between prey consumption and colony size in an orb spider

Summary I studied the relationship between prey consumption and colony size in the orb spiderPhiloponella semiplumosa. Observations of unmanipulated colonies showed that prey biomass per juvenile spider was positively correlated with colony size, indicating that prey consumption was highest in the largest colonies observed. In contrast, the relationship between prey biomass per adult female and colony size was curvilinear; prey consumption tended to be highest in intermediatesized colonies. Adult female cephalothorax width was positively correlated with colony size. Number of egg sacs per adult female tended to be highest in intermediate-sized colonies. Prey biomass per juvenile was lower in experimentally reduced colonies than in large control colonies. Aerial-arthropod abundance was not correlated with colony size, and experimental prey supplementation did not affect colony size. Thus, the relationship between prey consumption and colony size was influenced by coloniality directly, rather than by a correlation between prey abundance at a site and colony size.     

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     

Optimal foraging of sticklebacks on swarming prey

Hungry sticklebacks, Gasterosteus aculeatus, preferentially attacked the densest region of a swarm of water fleas, but with decreasing attack readiness they increasingly preferred less dense regions. Such a hunger dependent change in feeding preference has not yet been dealt with by optimal foraging theory. A model, which assumes that high swarm densities provide high feeding rates (because of small inter-prey distances) but also high costs of confusion, predicts that a predator should always choose the lowest prey density in which it can achieve a feeding rate sufficient to satisfy its hunger. Some predictions of the model were experimentally verified. Hungry fish have a higher feeding rate in a high prey density than in a lower density and less hungry fish have a higher rate in a low density than in a high density.     

The effects of plant dispersion and prey density on parasitism rates in a naturally patchy habitat

Despite extensive research on parasitoid-prey interactions and especially the effects of heterogeneity in parasitism on stability, sources of heterogeneity other than prey density have been little investigated. This research examines parasitism rates by three parasitoid species in relationship to prey density and habitat spatial pattern. The herbivore Itame andersoni (Geometridae) inhabits a subdivided habitat created by patches of its host plant, Dryas drummondii, in the Wrangell Mountains of Alaska. Dryas colonizes glacial moraines and spreads clonally to form distinct patches. Habitat subdivision occurs both on the patch scale and on the larger spatial scale of sites due to patchy successional patterns. Itame is attacked by three parasitoids: an ichneumonid wasp (Campoletis sp.), a braconid wasp (Aleiodes n. sp.), and the tachinid fly (Phyrxe pecosensis). I performed a large survey study at five distinct sites and censused Itame density and parasitism rates in 206 plant patches for 1–3 years. Parasitism rates varied with both plant patch size and isolation and also between sites, and the highest rates of overall parasitism were in the smallest patches. However, the effects of both small- and large-scale heterogeneity on parasitism differed for the three parasitoid species. There was weak evidence that Itame density was positively correlated with parasitism for the braconid and tachinid at the patch scale, but density effects differed for different patch sizes, patch isolations, and sites. At the site scale, there was no evidence of positive, but some indication of negative density-dependent parasitism. These patterns do not appear to be driven by negative interactions between the three parasitoid species, but reflect, rather, individual differences in habitat use and response to prey density. Finally, there was no evidence that parasitism strongly impacted the population dynamics of Itame. These results demonstrate the importance of considering habitat pattern when examining spatial heterogeneity of parasitism and the impacts of parasitoids. Received: 3 June 1999 / Accepted: 4 October 1999     

Variation in the effect of profitability on prey size selection by the lacertid lizard Psammodromus algirus

Maximizing the average rate of energy intake (profitability) may not always be the optimal foraging strategy for ectotherms with relatively low energy requirements. To test this hypothesis, we studied the feeding behaviour of captive insectivorous lizards Psammodromus algirus, and we obtained experimental estimates of prey mass, handling time, profitability, and attack distance for several types of prey. Handling time increased linearly with prey mass and differed significantly among prey types when prey size differences were controlled for, and mean profitabilities differed among prey taxa, but profitability was independent of prey size. The attack distance increased with prey length and with the mobility of prey, but it was unrelated to profitability. Thus, lizards did not seem to take account of the rate of energy intake per second as a proximate cue eliciting predatory behavior. This information was combined with pitfall-trap censuses of prey (in late April, mid-June and late July) that allowed us to compare the mass of the prey captured in the environment with that of the arthropods found in the stomachs of sacrificed free-living lizards. In April, when food abundance was low and lizards were reproducing, profitability had a pronounced effect on size selection and lizards selected prey larger than average from all taxa except the least profitable ones. As the active season progressed, and with a higher availability of food, the number of prey per stomach decreased and their mean ize increased. The effect of profitability on size selection decreased (June) and eventually vanished (July–August). This variation is probably related to seasonal changes in the ecology of lizards, e.g. time minimization in the breeding season as a means of saving time for nonforaging activities versus movement minimization by selecting fewer (but larger) prey in the postbreeding season. Thus, the hypothesis that maximizing profitability could be just an optional strategy for a terrestrial ectothermic vertebrate was supported by our data.     

The oddity effect drives prey choice but not necessarily attack time

The tendency of predators to preferentially attack phenotypically odd prey in groups (the oddity effect) is a clear example of how predator cognition can impact behaviour and morphology in prey. Through targeting phenotypically odd prey, predators are thought to avoid the cognitive constraints that delay and limit the success of attacks on homogenous prey groups (the confusion effect). In addition to influencing which prey a predator will attack, the confusion and oddity effects would also predict that attacks on odd prey can occur more rapidly than attacking the majority prey type, as odd prey are more easily targeted, but this prediction has yet to be tested. Here, we used kerri tetra fish, Inpaichthys kerri, presented with mixed phenotypic groups of Daphnia dyed red or black to investigate whether odd prey in groups are preferentially attacked and whether these attacks were faster than those on the majority prey type. In agreement with previous work, odd prey were targeted and attacked more often than expected from their frequency in the prey groups, regardless of whether the odd prey was red in a group of black prey or vice versa. However, no difference was found in the time taken to attack odd vs. majority prey items, contrary to our predictions. Our results suggest that the time taken to make an attack is determined by a wider range of factors or is subject to greater variance than the choice of which prey is selectively targeted in a group.