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.     

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.     

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.     

Flexibility in assessment of prey cues: frog-eating bats and frog calls

Predators use cues associated with their prey to assess prey quality and to avoid consuming poisonous prey. Considerable attention has been given to predators' use of aposematic cues to assess prey quality, but little is known about predators that eavesdrop on prey cues that are not intended for them. Here we investigate the prey-cue/prey-quality associations of a predator that eavesdrops on the sexual advertisement signals of its prey. Stability is expected in prey-cue/prey-quality associations when mistakes in prey assessment are lethal. Conversely, flexibility is possible when mistakes are less costly. Predators that must respond to temporal and spatial fluctuations in prey availability should be more flexible in their assessment of prey quality. Given these predictions, we examined flexibility in the ability of wild-caught bats to reverse prey-cue/prey-quality associations for a preferred prey and a poisonous one. We found that the predatory bat, Trachops cirrhosus, has a heretofore undescribed ability to reverse its evaluations of the cues that signal preferred prey.     

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.     

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.     

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.     

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.     

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.     

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.     

The evolution of warning signals as reliable indicators of prey defense

It is widely argued that defended prey have tended to evolve conspicuous traits because predators more readily learn to avoid defended prey when they are conspicuous. However, a rival theory proposes that defended prey have evolved such characters because it allows them to be distinguished from undefended prey. Here we investigated how the attributes of defended (unprofitable) and undefended (profitable) computer-generated prey species tended to evolve when they were subject to selection by foraging humans. When cryptic forms of defended and undefended species were similar in appearance but their conspicuous forms were not, defended prey became conspicuous while undefended prey remained cryptic. Indeed, in all of our experiments, defended prey invariably evolved any trait that enabled them to be distinguished from undefended prey, even if such traits were cryptic. When conspicuous mutants of defended prey were extremely rare, they frequently overcame their initial disadvantage by chance. When Batesian mimicry of defended species was possible, defended prey evolved unique traits or characteristics that would make undefended prey vulnerable. Overall, our work supports the contention that warning signals are selected for their reliability as indicators of defense rather than to capitalize on any inherent educational biases of predators.     

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.     

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.     

Modulation of prey capture kinematics and the role of lingual sensory feedback in the lizard Pogona vitticeps

Most organisms feed on a variety of prey that may differ dramatically in their physical and behavioural characteristics (e.g. mobility, mass, texture, etc.). Thus the ability to modulate prey capture behaviour in accordance with the characteristics of the food appears crucial. In animals that use rapid tongue movements to capture prey (frogs and chameleons), the coordination of jaws and tongue is based on visual cues gathered prior to the prey capture event. However, most iguanian lizards have much slower tongue-based prey capture systems suggesting that sensory feedback from the tongue may play an important role in coordinating jaw and tongue movements. We investigated the modulation of prey capture kinematics in the agamid lizard Pogona vitticeps when feeding on a range of food items differing in their physical characteristics. As the lizard is a dietary generalist, we expected it to be able to modulate its prey capture kinematics as a function of the (mechanical) demands imposed by the prey. Additionally, we investigated the role of lingual sensory feedback by transecting the trigeminal sensory afferents. Our findings demonstrated that P. vitticeps modulates its prey capture kinematics according to specific prey properties (e.g. size). In addition, transection of the trigeminal sensory nerves had a strong effect on prey capture kinematics. However, significant prey type effects and prey type by transection effects suggest that other sources of sensory information are also used to modulate the prey capture kinematics in P. vitticeps.     

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.     

Prey ‘handling time’ and its importance in food selection by the 15-spined stickleback, Spinachia spinachia (L.)

The time taken to manipulate and swallow a prey item, termed ‘handling time’, increases with decreasing hunger in Spinachia spinachia (L.), Handling time is also dependent upon the size of the prey in relation to the size of the mouth. Estimates of the optimum prey size, defined as that prey which minimizes the value of the ‘cost/benefit’ ratio of handling time/mg dry weight of prey, agrees closely with the mean prey sizes of wild fish. Optimum prey size was found to be approximately half the maximum swallowing capacity of the mouth. The nature of the relationship between the cost/benefit ratio and prey weight was used to explain the facts that the size range of prey eaten is dependent upon fish length and that decreasing hunger results in fish becoming increasingly selective with respect to prey size.     

Well-informed foraging: damage-released chemical cues of injured prey signal quality and size to predators

Predators use a variety of information sources to locate potential prey, and likewise prey animals use numerous sources of information to detect and avoid becoming the meal of a potential predator. In freshwater environments, chemosensory cues often play a crucial role in such predator/prey interactions. The importance of chemosensory information to teleost fish in marine environments is not well understood. Here, we tested whether coral reef fish predators are attracted to damage-released chemical cues from already wounded prey in order to find patches of prey and minimize their own costs of obtaining food. Furthermore, we tested if these chemical cues would convey information about status of the prey. Using y-maze experiments, we found that predatory dottybacks, Pseudochromis fuscus, were more attracted to skin extracts of damselfish, Pomacentrus amboinensis, prey that were in good condition compared to prey in poor body condition. Moreover, in both the laboratory and field, we found that predators could differentiate between skin extracts from prey based on prey size, showing a greater attraction to extracts made from prey that were the appropriate size to consume. This suggests that predators are not attracted to any general substance released from an injured prey fish instead being capable of detecting and distinguishing relatively small differences in the chemical composition of the skin of their prey. These results have implications for understanding predator foraging strategies and highlights that chemical cues play a complex role in predator–prey interactions in marine fish.     

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.     

Cooperative foraging in neotropical pseudoscorpions: effects of prey changes on behavioral adjustments of colonies

The pseudoscorpion Paratemnoides nidificator is a generalist predator that captures large arthropods that live on tree trunks. Few pseudoscorpions species show some degree of sociality. We investigated how colonies of the pseudoscorpion P. nidificator adjust their cooperative capture behavior under a situation of changing prey types as a simulation of variation in prey availability. We hypothesized that colonies would be more efficient at prey capture under repeated exposure to the same prey, and that the change in the availability of prey would be followed by new behavioral adjustments to adequately exploit the new prey. Eight experimental colonies housed in the laboratory received repetitions of three different ant species as prey. The number of pseudoscorpions attacking the prey, the number of behavioral acts, and the time expended subduing prey were evaluated as measures of prey capture performance, in relation to repetitive exposure to the same prey and also in relation to prey type changes. However, only individuals’ recruitment significantly responded to prey type exposure. Prey capture behavior was heterogeneous among colonies, resulting in highly variable behavioral responses. Colonies showed a tendency toward increasing capture success through repeated prey type exposure. However, 50% of the colonies were unable to capture the new prey type and died of starvation. Although it is a generalist predator, prey capture behavior could depend on different coordination components for subduing and handling large prey. Therefore, changes in prey availability could cause the attenuation of a cooperative relationship in some colonies, making them more prone to failure during capture.     

Chromaticity in the UV/blue range facilitates the search for achromatically background-matching prey in birds

A large variety of predatory species rely on their visual abilities to locate their prey. However, the search for prey may be hampered by prey camouflage. The most prominent example of concealing coloration is background-matching prey coloration characterized by a strong visual resemblance of prey to the background. Even though this principle of camouflage was recognized to efficiently work in predator avoidance a long time ago, the underlying mechanisms are not very well known. In this study, we assessed whether blue tits (Cyanistes caeruleus) use chromatic cues in the search for prey. We used two prey types that were achromatically identical but differed in chromatic properties in the UV/blue range and presented them on two achromatically identical backgrounds. The backgrounds had either the same chromatic properties as the prey items (matching combination) or differed in their chromatic properties (mismatching combination). Our results show that birds use chromatic cues in the search for mismatching prey, whereupon chromatic contrast leads to a ‘pop-out’ of the prey item from the background. When prey was presented on a matching background, search times were significantly higher. Interestingly, search for more chromatic prey on the matching background was easier than search for less chromatic prey on the matching background. Our results indicate that birds use both achromatic and chromatic cues when searching for prey, and that the combination of both cues might be helpful in the search task.