Effects of prey quantity and quality on predatory wasps

1. The simultaneous effects of prey quantity and prey quality on fitness correlates of the predatory wasp Polistes fuscatus were examined in a glasshouse study. Prey quantity was manipulated by providing prey in excess (high quantity) or one‐third of that (low quantity). Prey quality was manipulated by providing either palatable (Manduca sexta) caterpillars or unpalatable (Junonia coenia) caterpillars. 2. The effect of prey quality on wasp production depended on prey quantity. Nests given unpalatable prey produced few wasps whereas nests given palatable prey increased wasp production with increased prey. 3. The low production of nests given unpalatable prey reflected the low acceptability of those prey. The wasps preferred the palatable prey and learned to reject the unpalatable prey. With no choice of prey, they took only enough unpalatable prey to develop a small nest or colony. 4. A diet of unpalatable prey also resulted in smaller wasps and reduced the proportion of males produced, from about 40% to just 8–14%, depending on the year.     

Do native predators benefit from non‐native prey?

Despite knowledge on invasive species’ predatory effects, we know little of their influence as prey. Non‐native prey should have a neutral to positive effect on native predators by supplementing the prey base. However, if non‐native prey displace native prey, then an invader's net influence should depend on both its abundance and value relative to native prey. We conducted a meta‐analysis to quantify the effect of non‐native prey on native predator populations. Relative to native prey, non‐native prey similarly or negatively affect native predators, but only when studies employed a substitutive design that examined the effects of each prey species in isolation from other prey. When native predators had access to non‐native and native prey simultaneously, predator abundance increased significantly relative to pre‐invasion abundance. Although non‐native prey may have a lower per capita value than native prey, they seem to benefit native predators by serving as a supplemental prey resource.     

Estimating piscine prey size from partial remains: testing for shifts in foraging mode by juvenile bluefish

Knowledge of prey sizes consumed by a predator aids in the estimation of predation impact. Young-of-the-year bluefish, Pomatomus saltatrix, attack their prey tail-first and often bite their prey in half; this poses a unique problem in determining prey sizes from stomach content analysis. We developed a series of linear regressions to estimate original prey lengths from measurements of eye diameter and caudal peduncle depth for striped bass, Morone saxatilis, bay anchovy, Anchoa mitchilli, American shad, Alosa sapidissima, blueback herring, Alosa aestivalis, Atlantic silverside, Menidia menidia, and white perch, Morone americana. We then used these regressions to estimate original prey sizes from pieces of prey found in stomachs of bluefish collected in the Hudson River estuary from 1990–1993. Lengths of prey that were swallowed whole were compared to estimated lengths of prey that were consumed in pieces. Lengths of prey that were consumed in pieces were larger than prey that were consumed whole. We determined the prey length/predator length ratio at which bluefish began shifting from swallowing their prey whole to partial consumption. Shifting occurred at a ratio of approximately 0.35 irrespective of prey species, suggesting that prey length plays an important role in predator foraging decisions and may contribute to gape limitations. Shifts in foraging mode effectively reduce gape limitation and allow bluefish to consume larger prey sizes which may increase their effect on prey populations.     

Intraspecific variation in prey quality: a comparison of nutrient presence in prey and nutrient extraction by predators

Prey quality can have large impacts on the survival, growth and behavior of predators. A number of studies have examined how different species of prey vary in quality. However, far less is known about intraspecific variation in the quality of prey for predators and even less about what nutrients are extracted from prey by predators. We examined how the sex, feeding level and developmental status of prey affected the quantities of nutrients present in prey bodies and the quantities of nutrients that could be extracted from prey by spiders. Female and well‐fed prey were larger and had more nutrients than male and food‐limited prey, respectively. After taking into account differences in prey size, spiders extracted relatively more lipid and less protein from female and well‐fed prey than from male and food‐limited prey, respectively. Mealworms were of higher quality than adult mealworm beetles; spiders were able to extract more lipid, protein and other nutrients from larvae than adults. While lipid present in prey was a good predictor of lipid consumed, protein present in prey was not a reliable predictor of protein consumed. The variation in prey quality that we observed within a single species of prey (i.e. well‐fed vs food‐limited crickets) was as large as variation in quality among the three species of prey used in these experiments. Intraspecific variation in prey quality may be an important factor affecting predatory arthropods, especially in habitats or at times of year when one species of prey is abundant. Further studies are needed to examine the consequences of intraspecific variation in prey quality on the life history and behavior of predators.     

The positive effects of negative interactions: Can avoidance of competitors or predators increase resource sampling by prey?

Spatial overlap between predators and prey is key to predicting their interaction strength and population dynamics. We constructed a spatially-explicit simulation model to explore how predator and prey behavioral traits and patterns of resource distribution influence spatial overlap between predators, prey, and prey resources. Predator and prey spatial association primarily followed the ideal free distribution. Departures from this model were intriguing, especially from the interactions of predator and prey behavior. When prey weakly avoided conspecifics, they associated more highly with resources when predators were present. Predators increased the rate of prey movement between patches, which increased their ability to sample their environment and aggregate in patches with high resources. When prey strongly avoided each other, predators decreased prey association with resources. That is, an increased rate of prey movement increased the probability that prey would interact and avoid each other without regard to the distribution of resources. More generally, a more highly clumped distribution of resources acted as a spatial anchor that generally increased prey, predator, and resource association. Prey tended to congregate with resources and predators generally congregated with prey.     

Predator-prey shell games: large-scale movement and its implications for decision-making by prey

Many classical models of food patch use under predation risk assume that predators impose patch-specific predation risks independent of prey behavior. These models predict that prey should leave a chosen patch only if and when the food depletes below some critical level. In nature, however, prey individuals may regularly move among food patches, even in the apparent absence of food depletion. We suggest that such prey movement is part of a predator-prey "shell game", in which predators attempt to learn prey location, and the prey attempt to be unpredictable in space. We investigate this shell game using an individual-based model that allows predators to update information about prey location, and permits prey to move with some random component among patches, but with reduced energy intake. Our results show the best prey strategy depends on what the predator does. A non-learning (randomly moving) predator favors non-moving prey – moving prey suffer higher starvation and predation. However, a learning predator favors prey movement. In general, the best prey strategy involves movement biased toward, but not completely committed to, the richer food patch. The strategy of prey movement remains beneficial even in combination with other anti-predator defenses, such as prey vigilance.     

Effect of prey density on behaviour and development of the predatory mosquito,Toxorhynchites towadensis

The effect of prey density on feeding behaviour, killing behaviour, and development of the predatory mosquito,Toxorhynchites towadensis, was investigated in the laboratory. The number of prey consumed per larva increased toward an upper asymptote as prey density increased. Prey consumption curves during fourth instar were concave at low prey densities but convex at high prey densities. This phenomenon was not observed during other instars. Killing without consuming any part of prey occurred at prey densities of 20 per container and over. The number of prey killed but not consumed increased linearly with the number of unconsumed prey in the container. Prey acquisition behaviour was not affected by prey densities during the prepupal period. Developmental time from first instar to adult emergence decreased with increasing prey densities, but remained constant at densities of 10 per container and over. Adult size increased with increasing prey densities but there was no effect at prey densities of 20 and over.     

Effects of prey density and spatial distribution on prey consumption of the adult predatory ladybird beetle

The effects of prey density and spatial distribution on prey consumption of the adult predatory ladybird, Harmonia axyridis , were investigated by using a 2 × 2 factorial design in large scale cages. Prey density influenced prey consumption of the ladybirds, and the frequency with which predation occurred was quite different between the prey distributions. The ladybirds consumed a relatively constant and small number of aphids when the prey were uniformly distributed, whereas the number of prey consumed per day when predation occurred was large and much more variable when the prey were contagiously distributed. At high prey density, the number of prey consumed was highest during the first day of the experiment; thereafter, only 10–20 aphids were consumed during the following 3 days. However, these patterns of prey consumption were not observed at low prey density. The percentage of aphids that remained on the host plants when the experiments were terminated was higher at low prey density than at high prey density, suggesting that predator foraging efficiency at low prey density was lower than at high prey density. Ladybirds foraging for high prey density were more frequently observed on the plants with aphids than ladybirds foraging for low prey density. Prey distribution also influenced the frequency of residence of ladybirds on the plants. The different predation patterns observed in the two spatial distributions, in which prey consumption was much more variable for the contagious distribution, might be explained by the difference in prey encounter rate of the predator between the distributions. This study indicated that the ladybirds had limited ability to search out prey over large spatial scales.     

Predators target rare prey in coral reef fish assemblages

Predation can result in differing patterns of local prey diversity depending on whether predators are selective and, if so, how they select prey. A recent study comparing the diversity of juvenile fish assemblages among coral reefs with and without predators concluded that decreased prey diversity in the presence of predators was most likely caused by predators actively selecting rare prey species. We used several related laboratory experiments to explore this hypothesis by testing: (1) whether predators prefer particular prey species, (2) whether individual predators consistently select the same prey species, (3) whether predators target rare prey, and (4) whether rare prey are more vulnerable to predation because they differ in appearance/colouration from common prey. Rare prey suffered greater predation than expected and were not more vulnerable to predators because their appearance/colouration differed from common prey. Individual predators did not consistently select the same prey species through time, suggesting that prey selection behaviour was flexible and context dependent rather than fixed. Thus, selection of rare prey was unlikely to be explained by simple preferences for particular prey species. We hypothesize that when faced with multiple prey species predators may initially focus on rare, conspicuous species to overcome the sensory confusion experienced when attacking aggregated prey, thereby minimizing the time required to capture prey. This hypothesis represents a community-level manifestation of two well-documented and related phenomena, the “confusion effect” and the “oddity effect”, and may be an important, and often overlooked, mechanism by which predators influence local species diversity.     

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.     

The Role of gastric evacuation rate in handling time of equal-mass rations of different prey sizes in northern pike

In a laboratory experiment, northern pike Esox lucius gastric evacuation rates did not differ between equal-mass rations of small and large prey. In a comparison between intermediate and large prey, the pike were unable to fit two intermediate prey completely into the stomach at the same time, resulting in two consecutive evacuations, and changes in patterns of gastric evacuation. Thus, total gastric evacuation time was not affected by prey size composition in equal-mass rations, but patterns in evacuation rate may depend on the size ratio between predator and prey. Cumulative manipulation time between strike and complete swallowing of prey differed between equal-mass rations of small, intermediate and large prey, in that small prey took the shortest time to manipulate. Pike had problems striking and redirecting intermediate prey to swallow them head first, and the manipulation times for intermediate prey were as long as for large prey. Since an increased time manipulating prey in the mouth increases risk of predation and intraspecific interactions in pike, it is concluded that risks associated with long manipulation times, and not only energy per total handling time, determine prey value and prey size preference in this piscivore.     

Response of predators to prey abundance: separating the effects of prey density and patch size

We tested the relative and combined effects of prey density and patch size on the functional response (number of attacks per unit time and duration of attacks) of a predatory reef fish (Cheilodactylus nigripes (Richardson)) to their invertebrate prey. Fish attacked prey at a greater rate and for longer time in large than small patches of prey, but large patches had naturally greater densities of prey. We isolated the effects of patch size and prey density by reducing the density of prey in larger patches to equal that of small patches; thereby controlling for prey density. We found that the intensity at which fish attacked prey (combination of attack rate and duration) was primarily a response to prey density rather than the size of patch they occupied. However, there was evidence that fish spent more time foraging in larger than smaller patches independent of prey density; presumably because of the greater total number of prey available. These experimental observations suggest that fish can distinguish between different notions of prey abundance in ways that enhance their rate of consumption. Although fish may feed in a density dependent manner, a critical issue is whether their rate of consumption outstrips the rate of increase in prey abundance to cause density dependent mortality of prey.     

Preference and consumption of Macrolophus pygmaeus preying on mixed instar assemblages of Myzus persicae

This study examines the effects of changes in the prey frequency and abundance on prey selection among the four instars of Myzus persicae by the predator Macrolophus pygmaeus under laboratory conditions. The central hypothesis was that M. pygmaeus will become more selective as prey density increases. It was also observed that M. pygmaeus can occasionally abandon a prey item that had already been killed (non-consumptive prey mortality). It was assumed that the frequency of this behavior would increase with the prey size and prey density. For these purposes prey selection was evaluated by simultaneously presenting all instars of M. persicae to the predator in equal proportions and at increasing densities. M. pygmaeus showed a higher predation rate and a higher preference for smaller prey instars at all prey densities. However, if the predation rate by the predator is expressed in terms of biomass consumed, then biomass gain was higher when feeding on the larger instars of M. persicae. The prey selectivity was indicated by the total prey mortality (consumptive plus non-consumptive prey mortality) as well as by the non-consumptive prey mortality, was associated with relatively high prey densities, depending on the prey instar. Therefore, we argued that the predatory impact of M. pygmaeus on the various instars of the aphid depends not only on prey traits but also on their relative abundance in a patch. Observed decreases in biomass gain from larger prey were likely the result of high prey availability at densities before saturation, which might have caused confusion in the predator’s prey selection.     

No evidence of nonlinear effects of predator density,refuge availability,or body size of prey on prey mortality rates

Predator density, refuge availability, and body size of prey can all affect the mortality rate of prey. We assume that more predators will lead to an increase in prey mortality rate, but behavioral interactions between predators and prey, and availability of refuge, may lead to nonlinear effects of increased number of predators on prey mortality rates. We tested for nonlinear effects in prey mortality rates in a mesocosm experiment with different size classes of western mosquitofish (Gambusia affinis) as the prey, different numbers of green sunfish (Lepomis cyanellus) as the predators, and different levels of refuge. Predator number and size class of prey, but not refuge availability, had significant effects on the mortality rate of prey. Change in mortality rate of prey was linear and equal across the range of predator numbers. Each new predator increased the mortality rate by about 10% overall, and mortality rates were higher for smaller size classes. Predator–prey interactions at the individual level may not scale up to create nonlinearity in prey mortality rates with increasing predator density at the population level.     

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.     

Frequency-dependent predation and maintenance of prey polymorphism

In positive frequency-dependent predation, predation risk of an individual prey correlates positively with the frequency of that prey type. In a number of small-scale experiments individual predators have shown frequency-dependent behaviour, often leading to the conclusion that a population of such predators could maintain prey polymorphism. Using simulations, I studied the dynamics of frequency-dependent predation and prey polymorphism. The model suggests that persistence of prey polymorphism decreases with increasing number of predators that show frequency-dependent behaviour, questioning conclusions about polymorphism based on experiments with few predators. In addition, prey population size, prey crypsis, difference in crypsis between prey morphs and the way the behaviour was adjusted affected the persistence of polymorphism. Under some circumstances prey population remained polymorphic for a shorter time under frequency-dependent than under frequency-independent predation. This suggests that although positive frequency-dependent predator behaviour may maintain prey polymorphism, it is not a sufficient condition for persistent prey polymorphism.     

Craniodental indicators of prey size preference in the Felidae

In the present study, we used linear morphometrics of the crania, mandible and dentition to explore the association between craniodental shape and prey size among 35 species of living felids. To accomplish this, felids were divided into three prey-size groups: (1) large prey specialists; (2) small prey specialists; and (3) mixed prey feeders. From these linear measurements, large prey specialist felids can be distinguished from small and mixed prey feeders by their relatively robust canines and incisors and relatively wide muzzles. These cranial characters are advantageous when dispatching large prey, due to the stranglehold that cats employ during this activity. Robust canines resist the bending and torsional forces applied by struggling prey and a wider muzzle helps to stabilize grip and distribute bite forces more evenly during the killing bite. Small prey specialists had smaller canines, narrower muzzles and slightly longer jaws for a speed advantage when catching small, quick prey. Mixed prey feeders were intermediate between large and small prey specialists, indicating they are adapted to killing both sizes of prey. Given the success of this ecomorphological analysis of living felids that specialize on different prey sizes, we look forward to applying this same approach to extinct species.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 96 , 784–799.     

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.     

Separating the effects of prey size and speed on the kinematics of prey capture in the omnivorous lizard Gerrhosaurus major

Feeding behavior is known to be modulated as prey properties change. During prey capture, external prey properties, including size and mobility, are likely some of the most important components in predator–prey interactions. Whereas prey size has been demonstrated to elicit modulation of jaw movements during capture, how prey speed affects the approach and capture of prey remains unknown. We quantified the kinematics associated with movements of both the feeding and locomotor systems during prey capture in a lizard, Gerrhosaurus major, while facing prey differing in size and mobility (newborn mice, grasshoppers, and mealworms). Our data show that the feeding and locomotor systems were recruited differently in response to changes in the size or speed of the prey. The timing of jaw movements and of the positioning of the head are affected by changes in prey size—and speed, to a lesser extent. Changes in prey speed resulted in concomitant changes in the speed of strike and an early and greater elevation of the neck. External prey properties, and prey mobility in particular, are relevant in predator–prey interactions and elicit specific responses in different functional systems.     

Effects of Structural Refuge and Density on Foraging Behaviour and Mortality of Hungry Tadpoles Subject to Predation Risk

Theoretical models of prey behaviour predict that food‐limited prey engage in risk‐prone foraging and thereby succumb to increased mortality from predation. However, predation risk also may be influenced by factors including prey density and structural cover, such that the presumed role of prey hunger on predation risk may be obfuscated in many complex predator–prey systems. Using a tadpole (prey) – dragonfly larva (predator) system, we determined relative risk posed to hungry vs. sated prey when both density and structural cover were varied experimentally. Overall, prey response to perceived predation risk was primarily restricted to increased cover use, and hungry prey did not exhibit risk‐prone foraging. Surprisingly, hungry prey showed lower activity than sated prey when exposed to predation risk, perhaps indicating increased effort in search of refuge or spatial avoidance of predator cues among sated animals. An interaction between hunger level and predation risk treatments indicated that prey state affected sensitivity to perceived risk. We also examined the lethal implications of prey hunger by allowing predators to select directly between hungry and sated prey. Although predators qualitatively favoured hungry prey when density was elevated and structural cover was sparse, the overall low observed variation in mortality risk between hunger treatments suggests that preferential selection of hungry prey was weak. This implies that hunger effects on prey mortality risk may not be readily observed in complex landscapes with additional factors influencing risk. Thus, current starvation‐predation trade‐off theory may need to be broadened to account for other mechanisms through which undernourished prey may cope with predation risk.