Adaptive superparasitism and patch time allocation in solitary parasitoids : the influence of pre-patch experience

Summary Dynamic optimal diet models predict that host selection decisions and patch time allocation are influenced by the resource value of the habitat. We tested these predictions using the solitary' parasitoid Leptopilina heterotoma. Assuming that travel times between patches, the quality of previously visited patches and parasitoid density affect the parasitoids' estimation of the resource value of the habitat, different treatments were given before introducing parasitoids singly to a patch containing 5 unparasitized and 15 parasitized hosts. The decision to superparasitize is only slightly influenced by the rate of patch encounter. The quality of the previously visited patch has a strong influence. When a poor patch has been visited on the previous day, more superparasitism is observed in the partly depleted patch than when a rich patch has been searched. More superparasitism is also observed when the parasitoids are kept with conspecifics before the experiment than when they are kept alone. Increasing patch residence times are observed as the quality of the previously presented patch decreases. Host selection decisions and patch time allocation are thus clearly influenced by the pre-patch experience of the parasitoid, as predicted by dynamic optimal diet models. This can also explain why females that have never oviposited in unparasitized hosts will superparasitize readily. Correspondence to: M.E. Visser     

Travel duration,energetic expenditure,and patch exploitation in the parasitic wasp <Emphasis Type="Italic">Venturia canescens</Emphasis>

Foraging animals usually keep track of how costly it is to reach new resource patches and adjust patch residence time and exploitation rate accordingly. There are at least two potential factors, which are not necessarily closely linked, that animals could measure to estimate costs of traveling: the time the forager needs to reach the next patch and the amount of energy it has to invest until arrival. In the parasitoid wasp Venturia canescens, females forage for hosts from which their offspring can develop. Two different types of this parasitoid exist. The thelytokous type lives in anthropogenic habitats where flight is not necessarily linked with foraging. The arrhenotokous type lives under field conditions and shows frequent flight activity. We tested whether the wasps would use time or energy needed to assess patch availability, by either confining them into vials or letting them travel actively in a flight mill between patch visits. Our results show that in thelytokous lines, time is a sufficient cue influencing patch exploitation and an additional effect of the energy needed was not visible. In the arrhenotokous wasps, however, only the number of rounds flown in the mill influenced subsequent behavior, while mere time spent traveling did not. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

The loading effect in central place foraging wheatears (Oenanthe oenanthe L.)

Summary The behaviour of five adult wheatears (Oe. oenanthe) delivering prey (maggots and mealworms) from artificial patches to the nestlings was studied. The existence of a so-called loading effect was confirmed. Handling times for subsequent prey items increased with the number held in the beak, which led to a decrease in collecting rates with patch time and, in most cases, to positively accelerating loading functions when collecting times for prey items are plotted as a function of load size. All birds seemed to become more efficient at loading prey in the experimental patches during the course of the nestling period. Loading functions for maggots and mealworms were slightly different. When forcing the birds to visit several cups (only one item in each cup) and remove a layer of moss before reaching the prey (low-density patches), all birds took fewer prey and two of them stayed longer in them than in highdensity patches (one cup filled with prey items). This was due to an increase in search times with the number of prey held in the beak.From knowledge of the loading functions and travel times to the nest, it is possible to predict the optimal load sizes according to a mathematical solution of the delivery rate model of Orians and Pearson (1979). By transforming collecting and travel times to energy expenditures, it is also possible to derive predictions from an energy efficiency model (maximizing energy delivery per unit energy expended in a round-trip). The observed average load sizes did not differ significantly from those predicted by the delivery rate model, but they were significantly smaller in all cases than those predicted by the energy efficiency model. For birds feeding nestlings, it may be more important to sacrifice efficiency in energy expenditure in favour of greater delivery rates, thereby maximizing the growth rate of the young.     

Different modes of resource variation provide a critical test of ideal free distribution models

Ideal free distribution (IFD) models are perhaps the group of mathematical models of behavior that have been the most widely and successfully applied by empiricists. These models can be applied to nearly any situation in which consumers compete—by any mechanism—for resources that are patchily distributed in their environment. Although IFD models have come to be broadly accepted, experiments that simultaneously test more than a single prediction are rare. Instead, investigators normally either test (1) for a relationship between the distribution of consumers and the distribution of resources or (2) whether average fitnesses are equal across resource patches. We conducted experiments with pea aphids (Acyrthosiphon pisum Harris) feeding on two patches of fava beans (Vicia faba L.) to fully independently parameterize an IFD model with interference competition and then test quantitative predictions about aphid spatial distributions and the payoffs of patch choice. We found a precise fit between aphids’ predicted and observed reproductive successes. Furthermore, by varying patch “quality” in two ways, we were able to show that aphid distributions vary with the mode of resource variation in the predicted manner: aphids (1) matched resources when patches varied in size but not quality and (2) overmatched the good patch when patches varied in quality but not size (predicted as a consequence of weak interference). The close correspondence between quantitative predictions of the model with observed behaviors suggests that IFD theory is a framework with more explanatory power than is generally appreciated.     

Group foraging, patch exploitation time and the finder's advantage

 We use a combination of the marginal value theorem (MVT) of Charnov (1976), and a group foraging model featuring information sharing to address patch residence in an environment where food occurs in discrete patches. We shall show that among equal competitors the optimal patch time for the individual that finds the food patch is shorter than that for the non-finder among equal competitors, T _E < T _N. This is the case if the patch-finder commences food harvesting in the patch earlier and manages to monopolise a fraction of the prey items (finder's advantage) before the other individuals come to take their benefit. When individuals differ in their food-searching abilities so that some of them (producers) contribute proportionally more to food-searching than others (scroungers), and differ in ability to compete for the food found, a difference emerges between producer and scrounger individuals in the optimal patch time. Within a patch we always have the finder's advantage (T _E < T _N) regardless of phenotype. Between patches a suite of optimal patch times for encountering individuals emerges depending on the performance of producers and scroungers when changing from solitary feeding to feeding in a group. The optimal patch time for individuals that are affected more severely by competition is shorter than that for individuals of the phenotype with better competitive ability. When both phenotypes are affected similarly no difference in optimal patch times emerges. Received: 13 February 1996 / Accepted after revision: 28 September 1996     

Prey ecology and behaviour affect foraging strategies in the Great Cormorant

The fine link between a particular dive pattern and a specific prey item represents a challenging task in the analysis of marine predator–prey relationships. There is growing evidence that prey type affects diving seabirds’ foraging strategies, dive shapes and underwater activity costs. This study investigates whether a generalist diver, the Great Cormorant Phalacrocorax carbo, modifies the time budget allocated to prey-capture behaviour and breathing strategies (reactive vs. anticipatory) with respect to the prey type (pelagic vs. benthic). Video recordings of 91 Great Cormorants show how the ecology and behaviour of their main prey, Mullets (Mugilidae) and Flounders Platichthys flesus, affect dive/surface durations and the diving pattern. The demersal habit and the low mobility of Flounders leads to an easy access to prey with an anticipatory strategy. Moreover, the patchy distribution of this fish species increases prey-capture rates. Conversely, Mullets exploit the whole water column and are highly mobile, and this is reflected in the need of performing two sequential dives to capture a prey, both longer and likely more expensive, with a consequent switch of strategy from reactive in the searching phase to anticipatory breathing during prey-capture events. This study provides evidence that a generalist diver may switch between different foraging strategies, and it shows how each of them may be optimal under particular ecological conditions. These constraints influence the dynamics that operate within the marine food chains and have relevant implications in managing lagoon areas, including fish ponds.     

Optimal patch time allocation for time-limited foragers

The Charnov Marginal Value Theorem (MVT) predicts the optimal foraging duration of animals exploiting patches of resources. The predictions of this model have been verified for various animal species. However, the model is based on several assumptions that are likely too simplistic. One of these assumptions is that animals are living forever (i.e., infinite horizon). Using a simple dynamic programming model, we tested the importance of this assumption by analysing the optimal strategy for time-limited foragers. We found that, for time-limited foragers, optimal patch residence times should be greater than those predicted from the classic, static MVT, and the deviation should increase when foragers are approaching the end of their life. These predictions were verified for females of the parasitoid Anaphes victus (Hymenoptera: Mymaridae) exploiting egg patches of its host, the carrot weevil Listronotus oregonensis (Coleoptera: Curculionidae). As predicted by the model, females indeed remained for a longer time on host patches when they approached the end of their life. Experimental results were finally analysed with a Cox regression model to identify the patch-leaving decision rules females used to behave according to the model’s predictions.     

Diet of Neotropic cormorant (Phalacrocorax brasilianus) in an estuarine environment

The diet of the Neotropic cormorant (Phalacrocorax brasilianus) was studied by analysing 289 regurgitated pellets collected from a roosting site at Lagoa dos Patos estuary, southern Brazil, between November 2001 and October 2002 (except April to June). In total, 5,584 remains of prey items from 20 food types were found. Fish composed the bulk of the diet representing 99.9% by mass and 99.7% by number. The main food items were White croaker (Micropogonias furnieri) (73.7% by frequency of occurrence, 48.9% by mass and 41.2% by number), followed by Catfish (Ariidae) and anchovies (Engraulididae). In Lagoa dos Patos estuary the generalist Neotropic cormorant fed mainly on the two most abundant demersal fishes (White croaker and Catfish), which accounted for the low niche breadth calculated. The total length of all fish preyed varied from 27.2 to 318.3 mm (113.5 ± 48.0 mm), and preyed White croakers’ size differed between months. Neotropic cormorants seem to prey on most abundant class sizes of White croaker instead of selecting similar prey size throughout the time. However, temporary changes in diet in terms of food items, abundance and prey size were detected, revealing a high ecological plasticity of the species. Individual daily food intake of Neotropic cormorants estimated by pellets and metabolic equations corresponded to 23.7 and 27.1% of their body mass, falling in the range of other cormorant species. Annual food consumption of the population estimated by both methods was 73.4 and 81.9 tonnes, comprising mainly immature and subadult White croaker and Catfish which are commercially important. Temporal variations in diet composition and fish size preyed by Neotropics cormorants, a widespread and generalist species, suggest shifts according to fluctuations in the abundance of prey. The plasticity of this cormorant is also revealed by their ability to adjust feeding behaviour in response to temporal or local changes in the environment, from a generalist at the species level to a specialist at the individual or local population level.     

Recognition of individual-specific marked parasitized hosts by the solitary parasitoid Epidinocarsis lopezi

Summary In parasitoid wasps, self-superparasitism (oviposition into a host already parasitized by the female herself) often contributes less to the reproductive success of the parasitoid than oviposition into a host previously parasitized by a conspecific (conspecific superparasitism). It could therefore often be profitable for parasitoids to avoid self-superparasitism. This requires a mechanism for either (1) the avoidance of previously searched areas and/or (2) the rejection of hosts containing eggs laid by the searching female. We investigated whether the solitary parasitoid Epidinocarsis lopezi is able to avoid self-superparasitism. We show that visits to previously searched patches were shorter than visits to unsearched patches and conclude that E. lopezi females leave a trail odour on patches they have searched. No differences were found between the time on patches previously searched by the wasp itself and on patches visited by conspecifics. However, E. lopezi superparasitizes fewer hosts previously parasitized by itself than hosts parasitized by a conspecific. Thus, they recognize an individual-specific mark in or on the host. We discuss how patch marking and host marking enable E. lopezi to avoid self-superparasitism.     

Patch leaving decision rules in parasitoids: do they use sequential decisional sampling?

Within the framework of optimal foraging theory, models assume that parasitoid insects are able to evaluate the quality of the patch in which they are currently searching for hosts and the travel time between patches. They can adjust their residence time in consequence. Simple and more realistic decision mechanisms that induce behavior compatible with the predictions of these models have been proposed for a number of species. Most of these decision mechanisms only take into account the presence of unparasitized hosts. Here, we studied the consequences for leaving patches containing different proportions of unparasitized and parasitized hosts. We support the hypothesis that parasitoids sample their environment and we propose a binomial sequential model, based on the type of host encountered (unparasitized or parasitized) instead of on the time spent in a patch, to explain the giving-up behavior of a parasitoid in a patch. A motivational incremental/decremental stochastic process is proposed to explain a possible mechanism of the apparent sampling scheme followed by the insect. The empirical data support the hypothesis of a sequential, decisional, binomial sampling scheme performed with a limited memory. This memory is, in fact, more an effect of habituation than the "true memory" of the parasitoid. The theoretical model was applied to real data obtained with an encyrtid parasitoid. These data were also compared to realizations of the incremental/decremental process.     

Response of predators to loss and fragmentation of prey habitat: a review of theory

Despite extensive empirical research and previous reviews, no clear patterns regarding the effects of habitat loss and fragmentation on predator-prey interactions have emerged. We suggest that this is because empirical researchers do not design their studies to test specific hypotheses arising from the theoretical literature. In fact, theoretical work is almost completely ignored by empirical researchers, perhaps because it may be inaccessible to them. The purpose of this paper is to review theoretical work on the effects of habitat loss and fragmentation on predator-prey interactions. We provide a summary of clear, testable theoretical predictions for empirical researchers. To test one or more of these predictions, an empiricist will need certain information on the predator and prey species of interest. This includes: (1) whether the predator is a specialist on one prey species or feeds on many kinds of prey (omnivore and generalist); (2) whether the predator is restricted to the same habitat type as the focal prey (specialist), can use a variety of habitats but has higher survival in the prey habitat (omnivore), or lives primarily outside of the focal prey's habitat (generalist); (3) whether prey-only patches have lower prey extinction rates than predator-prey patches; and (4) whether the prey emigrate at higher rates from predator-prey patches than from prey-only patches. Empiricists also need to be clear on whether they are testing a prediction about habitat loss or habitat fragmentation and need to conduct empirical studies at spatial scales appropriate for testing the theoretical prediction(s). We suggest that appropriate use of the theoretical predictions in future empirical research will resolve the apparent inconsistencies in the empirical literature on this topic.     

Predators choosing between patches with standing crop: the influence of switching rules and input types

Where prey arriving in a patch are not consumed immediately, they will accumulate. Predators are then presented with a prey density or standing crop that increases through further input, and decreases through the consumption by predators. Firstly, I show that the switching rule of predators has a significant influence on the expected predator equilibrium distribution in such a dynamic system. Three rules are compared; for all rules, analytical solutions are calculated (where possible). To test their plausibility for natural situations, predator distributions are simulated given the assumption that each predator obtains individual patch profitability estimates by using a common learning rule. As long as prey arrive in the patches in constant numbers per time unit, the first rule leads to input matching because predators stop switching when consumption in the two patches is equal. The other two rules, where predators continue to sample both patches even in the equilibrium state, lead to predator distributions where the more profitable patch is underused. The final equilibrium depends on the exact assumptions of the switching rule; however, it is independent of interference. But if the input delivered into a patch is a function of the current prey standing crop (for example in a reproducing prey population), predator and prey distributions will not reach an equilibrium in most cases: either standing crops increase indefinitely, or they approach zero, with all predators concentrating on the better patch. Only a small number of parameter sets show intermediate crops that are reasonably stable. With this input type, only up to 54% of the simulations reach the expected distribution. In a system with competition for dynamic standing crop, it is therefore essential to know the type of input and the switching-rule used by predators to be able to predict equilibrium predator distributions. Received: 17 March 1995/Accepted after revision: 5 November 1995     

Larval behavior of predacious sister-species: orientation,molting site,and survival in Chrysopa

Field and laboratory studies compared two features of larval behavior in a pair of predacious sisterspecies of green lacewings: one (Chrysopa slossonae) a specialist on a single species of colonial aphids (the woolly alder aphid) that occur on branches and trunks of alder trees, the other (C. quadripunctata) a general aphid feeder whose primary prey is dispersed on foliage of diverse types of trees. First, a few hours after hatching, larvae of the two species develop significantly different phototactic responses; the differences correspond well with the spatial distributions of their prey. Most C. slossonae exhibited negative phototaxis, a response that helps move hatchlings inward on alder trees toward the woolly alder aphid colonies, whereas most C. quadripunctata hatchlings showed positive orientation to light, a response that tends to keep them in tree canopies with their prey. Second, in greenhouse experiments, a significantly greater proportion of C. slossonae larvae (second instars) molted within woolly alder aphid colonies and remained with the aphids than did C. quadripunctata larvae. These differences indicate that the specialist larvae have evolved a high degree of behavioral fidelity to their prey. However, larvae (second instars) of the two species that were released near ant-tended woolly alder aphid colonies in the field had similar recovery (= survival) rates. Consequently, natural selection may not act on behavioral traits that influence larval fidelity to prey during the late second and early third instars.     

Differential use of conspecific-derived information by sexual and asexual parasitic wasps exploiting partially depleted host patches

When foraging partially depleted patches (i.e., a fraction of hosts are already parasitized), female parasitoids must decide: 1—whether to superparasitize, and 2—whether to stay in their current patch (thus missing the opportunity of finding a better patch elsewhere). To make these decisions, parasitoids may rely on different cues, produced both by the environment and by conspecifics. Animals thriving in different environments may differ in cues they use. In the solitary parasitoid Venturia canescens, thelytokous (asexual) and arrhenotokous (sexual) individuals are found in two contrasting environments. Thelytokous females, from anthropogenic conditions, are known to cope with superparasitism in an adaptive way. On the other hand, little is known about superparasitism by arrhenotokous females. We compared the host exploitation strategies of thelytokous and arrhenotokous females in partially depleted patches. Hosts parasitized by thelytokous females were more frequently avoided than those parasitized by arrhenotokous females, suggesting a stronger chemical marking of the former. Only thelytokous females used information from conspecifics for patch-leaving decisions. The conformity of the differences in the behavior of thelytokous and arrhenotokous females with the environmental conditions they experience in their habitat is discussed.     

Inadvertent errors and error-constrained optimization: fallible foraging by bluegill sunfish

Summary When the expected reward rate is continuously reduced by foraging in a patch, foragers may adjust their patch persistence times to maximize the average long-term reward rate. The marginal-value model predicts the optimal persistence time for this situation. But real foragers may be unable consistently to achieve a precise persistence time. If the costs of under- and over-persistence differ, or if the resulting distribution of persistence times is skewed, a sufficiently broad persistence-time distribution can substantially shift the actual optimum. Moreover, this error-constrained optimum depends on the variable used by the forager to decide when to leave the patch (e.g., on persistence time per se, cumulative number of prey eaten, or instantaneous feeding rate). Here, we analyze laboratory data from bluegill sunfish (Lepomis macrochirus) foraging on larval-midge prey (Chironomus riparius) in patches of artificial vegetation, and we explore some wider implications of a model that seems to fit the data. The bluegills stayed 4%–157% longer in patches than predicted by the marginal value theorem. This behavior closely matched numerical solutions based on the observed variability of persistence times and the assumption that departures were cued by instantaneous feeding rate. On the other hand, the other two mechanisms that we investigated (i.e., persistence time per se and cumulative number of prey eaten) predict weak to moderate underpersistence relative to the marginal-value predictions, patterns quite unlike those observed. Surprisingly, the instantaneous-rate mechanism yields roughly a 10% lower over-all maximal reward rate than would either of the other two departure-cuing mechanisms. The modeling analysis documents the considerable sensitivity of our results to (1) the departure-cuing mechanism, (2) the shape of the frequency distribution of the departure-cuing variable, (3) the way that the shape of this distribution shifts as its mean changes, and (4) the magnitudes of the foraging parameters. Offprint requests to: P.H. Crowley     

Specificity of odour-mediated avoidance of competition in Drosophila parasitoids

Although there are many examples of the role of volatile infochemicals in interactions between trophic levels of insect communities, surprisingly little is known of volatile interactions between species within the third trophic level. Recently it was found that Leptopilina heterotoma, an endoparasitoid that attacks Drosophila larvae, avoids one type of patches (decaying stinkhorn mushrooms) when parasitoids of another species (L. clavipes) are present on these patches. L. heterotoma is able to smell the presence of L. clavipes from a distance (Fig. 1). In this paper we investigate the source of the odour that induces avoidance behaviour, by varying the host species and parasitoid species present on stinkhorn mushrooms, and by using another type of patch (sap-fluxes of wounded trees). L. heterotoma was found to avoid stinkhorn patches with conspecific as well as heterospecific parasitoids (Fig. 2). Hosts had to be present in the patch to elicit avoidance, but avoidance behaviour was also found with another host species present in the patch (Fig. 3). No avoidance behaviour was found with sap-flux patches with hosts and parasitoids on them (Fig. 4). Avoidance of stinkhorn patches only occurred when the parasitoids present on the patch were able to contact hosts (Figs. 5 and 6). The exact source of the odour that elicits avoidance is still unclear, so that one can only speculate on the function of the signal. However, there is a clear benefit to the receiver, because it is able to avoid superior competitors. Avoidance can lead to non-aggregated parasitoid distributions. The importance of avoidance behaviour for population dynamics and stability of parasitoid-host systems is discussed.     

Parasites determine a predator's optimal feeding strategy

Summary The diet selected by three-spined sticklebacks (Gasterosteus aculeatus) depends on the degree of parasitization by one or both of two parasite species (Schistocephalus solidus, Glugea anomala). Uninfested fish prefer the more profitable of two different size classes of prey (Daphinia magna). Fish parasitized by Glugea or Schistocephalus attack both prey types equally often, whereas sticklebacks infested by both parasite species prefer to attack the less profitable prey. The diet selected is optimal under the condition that parasites decrease their host's competitive ability.     

The influence of prey distribution on the foraging strategy of the lizard Oligosoma grande (Reptilia: Scincidae)

The grand skink, Oligosoma grande, is a diurnal rock-dwelling lizard from the tussock grasslands of Central Otago, New Zealand, whose diet includes a variety of arthropods and fruit. We conducted a field experiment to examine the influence of prey distribution on foraging behavior and spacing patterns. On sites where prey distribution was unaltered (control sites), males and females differed in diet and foraging behavior. Most male feeding attempts were directed at large strong-flying insects, and males used a saltatory search pattern that involved relatively infrequent moves of long duration. Females spent more effort catching small weak-flying insects and visiting fruiting plants. Their search behavior involved frequent moves of short duration. The placement of meat-bait on experimental sites led to a redistribution of large flies without influencing other prey types. Experimental females switched foraging strategy by adopting a search pattern of relatively infrequent moves of long duration, increasing the frequency of attempts to capture large prey, and reducing the importance of fruit in their diet. The experimental manipulation appeared to influence space use. On control sites, both sexes had comparably sized home ranges. On experimental sites, male home ranges were significantly larger than female home ranges. Received: 3 November 1997 / Accepted after revision: 13 December 1998     

Effects of within- and among-patch experiences on the patch-leaving decision rules in an insect parasitoid

The present study aimed to address how an insect parasitoid makes patch-departure decisions from various types of host patches and how previous patch experiences in the environment modify this decision-making process. Experiments were done with the parasitic wasp Aphidius rhopalosiphi attacking the grain aphid Sitobion avenae. In the experiments, wasps were observed in a laboratory environment containing several patches of various host densities, and behavioural records were analysed using a Coxs proportional hazards model. Consideration of the effect of the within-patch experience gave a classic pattern of patch-leaving decision rules in parasitoids: A. rhopalosiphi used local information on host quality (i.e. numbers of ovipositions or rejections) and availability (i.e. patch density) to determine departure decision. However, consideration of previous patch experiences provided evidence that these departure rules are fundamentally dynamic, responding to the physiological state of the animal and the information it has about its environment. Results showed that A. rhopalosiphi decreased its tendency to leave the visited patch after an oviposition. However, when a female has already laid several other eggs in the environment, such an incremental mechanism gradually switched to a decremental one. Hence, A. rhopalosiphi responded to egg-load depletion by leaving the visited patches sooner and by depositing a smaller number of eggs in those patches, which probably led to a decreased level of superparasitism. Results also indicated that previous experiences enabled wasps to estimate spatial host distribution and then to adjust their behavioural decisions accordingly. Thus, A. rhopalosiphi was shown to adjust its patch residence time according to the quality and the number of the patches previously visited. These proximate mechanistic rules adopted by A. rhopalosiphi females are discussed in the context of general predictions from optimality models.Communicated by D. Gwynne     

Dynamic response to danger in a parasitoid wasp

Despite the multitude of work on patch time allocation and the huge number of studies on patch choice in the face of danger, the patch leaving response of foragers perceiving cues of danger has received relatively little attention. We investigated the response of parasitoid insects to cues of danger both theoretically and experimentally. Using stochastic dynamic theory, we demonstrate that patch-leaving responses in response to the detection of danger should be seen as a dynamic decision that depends upon reproductive options on the current host patch and on alternative patches that might be found after leaving the current patch. Our theory predicts a sigmoidal response curve of parasitoids, where they should accept the danger and stay on the patch when patch quality is high and should increasingly avoid the risk and emigrate from the patch with decreasing patch quality and decreasing costs of traveling to an alternative host patch. Experiments with females of the drosophilid parasitoid Asobara tabida that were exposed to a puff of formic acid (a danger cue) at different times through their patch exploitation confirmed the theoretical predictions (i.e., a sigmoid response curve); however, the predicted curve was significantly steeper than observed. We discuss the impact of dynamic patch-exit decisions of individual foragers on population and community dynamics.