Piscivore efficiency and refuging prey: the importance of predator search mode

In predator-prey interactions, the efficiency of the predator is dependent on characteristics of both the predator and the prey, as well as the structure of the environment. In a field enclosure experiment, we tested the effects of a prey refuge on predator search mode, predator efficiency and prey behaviour. Replicated enclosures containing young of the year (0+) and 1-year-old (1+) perch were stocked with 3 differentially sized individuals of either of 2 piscivorous species, perch (Perca fluviatilis), pike (Esox lucius) or no piscivorous predators. Each enclosure contained an open predator area with three small vegetation patches, and a vegetated absolute refuge for the prey. We quantified the behaviour of the predators and the prey simultaneously, and at the end of the experiment the growth of the predators and the mortality and habitat use of the prey were estimated. The activity mode of both predator species was stationary. Perch stayed in pairs in the vegetation patches whereas pike remained solitary and occupied the corners of the enclosure. The largest pike individuals stayed closest to the prey refuge whereas the smallest individuals stayed farthest away from the prey refuge, indicating size-dependent interference among pike. Both size classes of prey showed stronger behavioural responses to pike than to perch with respect to refuge use, distance from refuge and distance to the nearest predator. Prey mortality was higher in the presence of pike than in the presence of perch. Predators decreased in body mass in all treatments, and perch showed a relatively stronger decrease in body mass than pike during the experiment. Growth differences of perch and pike, and mortality differences of prey caused by predation, can be explained by predator morphology, predator attack efficiency and social versus interference behaviour of the predators. These considerations suggest that pike are more efficient piscivores around prey refuges such as the littoral zones of lakes, whereas perch have previously been observed to be more efficient in open areas, such as in the pelagic zones of lakes.     

Avian prey-dropping behavior. I. The effects of prey characteristics and prey loss

Numerous species of birds break hard-shelled prey items by droppingthem from a height. This intriguing prey-extraction method providesan excellent opportunity for studying foraging behavior becausea single, easily measurable quantity—height of drop—maybe influenced by a wide variety of identifiable characteristicsof the prey (e.g., breakability, weight) and social environment(e.g., alone or in the presence of kleptoparasites). Using adynamic, state variable modeling approach, this paper presentsthe first theoretical framework for avian prey-dropping systemsthat incorporates the diversity of prey characteristics andsocial situations. The model yielded a series of qualitativepredictions about prey-dropping behavior that can be testedreadily in any prey-dropping system. In particular, the results indicatedthat quantitative and qualitative differences in item breakability andpotential kleptoparasitism should have a significant effecton the height and pattern of prey dropping.     

Prey abundance and the strength of interference in a foraging shorebird

Interference is an important component of food competition but is often difficult to detect and measure in natural animal populations. Although interference has been shown to occur between oystercatchers Haematopus ostralegus L. feeding on mussels Mytilus edulis L., four previous studies have not detected interference between oystercatchers feeding on cockles Cerastoderma edule L. In contrast, this study detected interference between cockle-feeding oystercatchers in the Baie de Somme, France. Prey stealing (kleptoparasitism), one of the main causes of interference between mussel-feeders, also occurred between oystercatchers in the Baie de Somme. The kleptoparasitism rate was related to the natural variation in the food supply, tending to be higher when cockles were rare. Feeding rate was negatively related to competitor density, so providing evidence for interference, but, as in mussel-feeders, only above a threshold density of about 50–100 birds ha⁻¹. The strength of interference at a fixed competitor density was related to the cockle food supply, usually being greater when cockles were rare. Previous studies probably failed to detect interference between cockle-feeders because competitor densities were too low, or cockles were too abundant, or because they were not conducted during late winter when interference is most intense. The study shows that natural variation in the food supply can influence the strength of interference within an animal population and provides support for those behaviour-based interference models which predict that the strength of interference will be greatest when competitor densities are high and prey scarce.     

Prey size, prey nutrition, and food handling by shrews of different body sizes

We tested some predictions relating metabolic constraints offoraging behavior and prey selection by comparing food handlingand utilization in four sympatric shrew species: Sorex minutus(mean body mass = 3.0 g), S. araneus (8.0 g), Neomys anomalus(10.0 g), and N. fodiens (14.4 g). Live fly larvae, mealwormlarvae, and aquatic arthropods were offered to shrews as smallprey (body mass <0.1 g). Live earthworms, snails, and smallfish were offered as large prey (>0.3 g). The larvae werethe high-nutrition food (>8 kJ/g), and the other prey werethe low-nutrition food (<4 kJ/g). The smallest shrew, S.minutus, utilized (ate + hoarded) <30% of offered food,and the other species utilized >48% of food. The largerthe shrew, the more prey it ate per capita. However, highlyenergetic insect larvae composed 75% of food utilized by S.minutus and only >40% of the food utilized by the other species. Thus, inverse relationships appeared between shrewbody mass and mass-specific food mass utilization and betweenshrew body mass and mass-specific food energy utilization:the largest shrew, N. fodiens, utilized the least food massand the least energy quantity per 1 g of its body mass. Also,the proportion of food hoarded by shrews decreased with increase in size of shrew. With the exception of S. araneus, the sizeof prey hoarded by the shrews was significantly larger thanthe size of prey eaten. Tiny S. minutus hoarded and ate smallerprey items than the other shrews, and large N. fodiens hoardedlarger prey than the other shrews.     

The dynamics of prey choice in fish: the importance of prey size and satiation

Over a number of decades the process of prey choice has been investigated using fishes as model predators. Using fishes for the model has allowed the proximate factors that determine how a mobile predator finds and chooses to eat the prey encountered within a variable 3‐D environment to be estimated. During prey choice a number of constraints exist, in particular most fish predators will eat their prey whole thus their jaws and gut create functional limitations once a prey has been attacked. By considering the relationship between the size of the prey and the predator's feeding apparatus and feeding motivation this study explores the link between mechanistic studies and theoretical, optimal foraging based predictions. How the prediction of prey choices made by the fish following prey encounter can be reconciled with what is likely to be found in the fish's stomach is discussed. This study uses a progression of empirical examples to illustrate how the limits of functional constraints and prey choice at different stages of motivation to feed can be taken into account to improve predictions of predator prey choice.     

Limited attention: the constraint underlying search image

Recent models of predator search behavior integrate proximate neurobiological constraints with ultimate economic considerations.These models are based on two assumptions, which we have criticallyexamined in experiments with blue jays searching for artificialprey images presented on a computer monitor. We found, first,that when jays had to switch between searching for two distinctprey types, they showed no reduction in detection rates comparedto no-switching to no-switching conditions, and second, that when jays divided attention between searching for two prey typesat the same time, they had lower detection rates than whenthey focused attention on one prey type at a time. Our resultssuggest that limited attention strongly affects predator searchpatterns and diet choice, including the ubiquitous tendencyto form search images.     

Predator size and the suitability of a common prey

 Although a predator’s mass should influence the suitability of its prey, this subject has received little direct attention. We studied the capture and processing of an abundant syrphid fly Toxomerus marginatus (c. 4 mg) by 0.6- to 40-mg juvenile crab spiders Misumena vatia (Thomisidae) to determine how profitability, relative profitability (profitability/predator mass), overall gain in mass, and relative gain in mass differed with predator mass, and whether foraging changed concurrently. In multi-prey experiments, the smallest successful spiders (0.6–3.0 mg) extracted less mass from flies, and did so more slowly, than large spiders. This gain was proportionately similar to that of 10- to 40-mg spiders with access to many Toxomerus. However, many small spiders failed to capture flies. When we gave spiders only a single Toxomerus, the smallest ones again extracted mass more slowly than the large ones and increased in mass less than the large ones, but increased in mass proportionately more than large ones. Relative gain in mass from a single prey decreased with increasing spider mass. Spiders larger than 10 mg all extracted similar amounts of mass from a single Toxomerus at similar rates, but varied in time spent between captures. Thus, Toxomerus changes with spider mass from a large, hard-to-capture bonanza to a small, easy-to-capture item of low per capita value. However, Toxomerus is common enough that large spiders can capture it en masse, thereby compensating for its decline in per capita value. Received: 7 May 1996 / Accepted: 23 September 1996     

Health food versus fast food: the effects of prey quality and mobility on prey selection by a generalist predator and indirect interactions among prey species

1. In order to understand the relative importance of prey quality and mobility in indirect interactions among alternative prey that are mediated by a shared natural enemy, the nutritional quality of two common prey for a generalist insect predator along with the predator's relative preference for these prey was determined. 2. Eggs of the corn earworm Helicoverpa zea (Lepidoptera: Noctuidae) were nutritionally superior to pea aphids Acyrthosiphum pisum (Homoptera: Aphididae) as prey for big‐eyed bugs Geocoris punctipes (Heteroptera: Geocoridae). Big‐eyed bugs survived four times as long when fed corn earworm eggs than when fed pea aphids. Furthermore, only big‐eyed bugs fed corn earworm eggs completed development and reached adulthood. 3. In two separate choice experiments, however, big‐eyed bugs consistently attacked the nutritionally inferior prey, pea aphids, more frequently than the nutritionally superior prey, corn earworm eggs. 4. Prey mobility, not prey nutritional quality, seems to be the most important criterion used by big‐eyed bugs to select prey. Big‐eyed bugs attacked mobile aphids preferentially when given a choice between mobile and immobilised aphids. 5. Prey behaviour also mediated indirect interactions between these two prey species. The presence of mobile pea aphids as alternative prey benefited corn earworms indirectly by reducing the consumption of corn earworm eggs by big‐eyed bugs. The presence of immobilised pea aphids, however, did not benefit corn earworms indirectly because the consumption of corn earworm eggs by big‐eyed bugs was not reduced when they were present. 6. These results suggest that the prey preferences of generalist insect predators mediate indirect interactions among prey species and ultimately affect the population dynamics of the predator and prey species. Understanding the prey preferences of generalist insect predators is essential to predict accurately the efficacy of these insects as biological control agents.     

Kleptoparasitic prey competition in shoaling fish: effects of familiarity and prey distribution

Familiarity is thought to stabilize dominance hierarchies andreduce aggressive interactions within groups of socially livinganimals. Though familiarity has been widely studied in shoalingfish, few studies have investigated changes in prey competitionas a function of time spent together within groups of initiallyunfamiliar individuals. In this study, we created shoals ofthree-spined stickleback (Gasterosteus aculeatus) and monitoredchanges in foraging rates and related competitive behaviorswithin shoals over a 4-week period in experimental series whereprey was spatially and temporally concentrated or dispersed.Prey share was unequal under both prey distribution modes, anddisparity in prey share was not seen to change as trials progressed.Interestingly, the contest rate for prey items fell over timewhen individuals were competing for dispersed prey but not whenprey were concentrated. We found no evidence that fish showedassociation preferences for either group members that had consumeda greater or lesser proportion of prey during trials. Thoughthe intensity of competition may be reduced by increased groupstability in nature, this is likely to be strongly dependenton the way prey resources are distributed through space andtime.     

Constraints on prey size selection by the three-spined stickleback: energy requirements and the capacity and fullness of the gut

An experiment was designed to study how gut fullness and encounter with 5-mm Asellus aquaticus influenced acceptance or rejection of less profitable 8-mm Asellus . 45-mm sticklebacks were found to always accept 5-mm prey whereas 8-mm prey were accepted with an initial probability of about 0.9. This probability decreased as the gut filled. Fish of differing sizes and sex had similar daily energy intakes per unit body size, however the acceptance of 8-mm prey was related to fish size. Whenever a fish orientated to a prey it was followed by pursuit and manipulation independently of prey size. The decision to accept or reject prey occurred after one manipulation, a criterion that was more variable for the larger prey. For one feeding session per day the total energy intake was almost constant despite the changing combination of prey sizes eaten. The fish ate prey with long handling times if the energetic contents of the stomach had not reached 450 J. Calculations were made of how many of each millimetre prey size group would satisfy the 450 J demand and how long the estimated number would take to handle. This showed that the best option is to consume 5-mm prey if given the choice.     

Behavioral interference and facilitation in the foraging cycle shape the functional response

Individual forager behaviors should affect per capita intakerates and thereby population and consumer-resource properties.We consider and incorporate conspecific facilitation and interferenceduring the separate foraging-cycle stages in a functional responsemodel that links individual behavioral interactions with consumer-resourceprocesses. Our analyses suggest that failing to properly considerand include all effects of behavioral interactions on foraging-cyclestage performances may either over- or underestimate effectsof interactions on the shape of both functional responses andpredator zero-growth isoclines. Incorporation of prey- and predator-dependentinteractions among foragers in the model produces predator isoclineswith potentials for highly complex consumer-resource dynamics.Facilitation and interference during the foraging cycle aretherefore suggested as potent behavioral mechanisms to causepatterns of community dynamics. We emphasize that correct estimationsof interaction-mediated foraging-cycle efficiencies should beconsidered in empirical and theoretical attempts to furtherour understanding of the mechanistic link between social behaviorsand higher order processes.     

When cormorants go fishing: the differing costs of hunting for sedentary and motile prey

Cormorants hunt both benthic (sedentary) and pelagic (motile) prey but it is not known if the energy costs of foraging on these prey differ. We used respirometry to measure the costs of diving in double-crested cormorants (Phalacrocorax auritus) foraging either for sedentary (fish pieces) or motile (juvenile salmon) prey in a deep dive tank. Short dives for sedentary prey were more expensive than dives of similar duration for motile prey (e.g. 20% higher for a 10s dive) whereas the reverse was true for long dives (i.e. long dives for motile prey were more expensive than for sedentary prey). Across dives of all durations, the foraging phase of the dive was more expensive when the birds hunted motile prey, presumably due to pursuit costs. The period of descent in all the dives undertaken appears to have been more expensive when the birds foraged on sedentary prey, probably due to a higher swimming speed during this period.     

The efficiency of adaptive search tactics for different prey distribution patterns: a simulation model based on the behaviour of juvenile plaice

Juvenile plaice Pleuronectes platessa are particularly useful for studying forager search behaviour because their search paths are essentially two dimensional, and punctuated by natural stops. Their prey occur in a range of natural distributions from highly aggregated to over‐dispersed. Juvenile plaice use area‐restricted search near aggregated prey and extensive search, consisting of longer moves and fewer turns, between aggregations and when searching for dispersed prey. They search for less conspicuous prey items mainly in the pauses between movements. This saltatory search behaviour contrasts with the continuous search that is usually assumed in search models. A simulation model of saltatory search behaviour showed that a strategy combining extensive and intensive search allows the efficient exploitation of a range of natural prey distribution patterns, and that it is particularly effective when the search behaviour is controlled by perceived prey density. This allows the predator to respond to the localized aggregations which often occur in nature. The selective use of intensive search was more efficient than the continuous use of extensive search even in prey distribution patterns that were statistically over‐dispersed.     

Seasonal variation in the strength of interference competition among headwater stream predators

1. Vertebrate communities in headwater streams are assumed to be regulated through competitive and predatory interactions. Although documented predation is rare, studies regularly report competitive dominance by fish that, as larger competitors reliant on aquatic habitat, exclude semi-aquatic salamanders to marginal stream habitat. However, it is unclear whether fish interact with stream-breeding salamanders through indirect effects such as competition for resources (e.g. food or cover) or fear (i.e. threat of predation) nor is it known whether these interactions are consistent through time.
2. This study used a novel caging approach to determine if competitive outcomes between a headwater fish and salamanders were regulated primarily through resource depletion (exploitative competition) or behavioural avoidance (interference competition).
3. We paired banded sculpin (Cottus carolinae) and larval red salamanders (Pseudotriton ruber) of similar body size in independent flow through mesocosms with intra- and inter-specific pairs allowed to interact physically or non-physically. The experiment was repeated in the autumn and in the spring when stream salamander larvae begin to transform into terrestrial juveniles.
4. Banded sculpin negatively influenced growth of red salamanders regardless of whether they were allowed to physically interact, suggesting interference competition and behavioural avoidance. This asymmetrical effect was strongest in the spring when salamanders underwent metamorphosis at higher rates in the presence of fish. However, in the autumn, the effects were more balanced between the two species with salamanders impacting fish through exploitative competition.
5. By studying the temporal relationships between two competitors and using a caging method novel to competition studies, we established that the outcomes of competition are dependent on season and may vary in type relative to the timing of life-history events. For this community, these results suggest that outcomes of competition are highly dependent on season and could indicate a biotic mechanism maintaining headwater salamander distributions through source–sink dynamics. Our results also suggest that, in this species interaction, it may be unwarranted to assume that the outcomes of competition at one time represent the complex relationships regulating community interactions.

     

Foraging behaviour and prey size of the painted stork

The foraging behaviour of painted stork Mycteria leucocephala was studied during 2004–2006 at 14 different sites in the Delhi region, India. Observations were recorded on 131 individuals, including 29 juvenile birds using a video camera. Recordings were also made at the nesting colony in Delhi zoo to study the prey sizes regurgitated to nestlings. The results confirm that the painted stork is a tactile forager and exclusively piscivorous. Foraging group size ranged from 1 to 18 individuals. Per 5 min foot stirring rates in the vegetated habitats were significantly higher than in non-vegetated habitats. The attempt rate and feeding rate in the breeding season were significantly higher than that in the non-breeding season. Prey sizes taken in the breeding season were significantly smaller than those taken in the non-breeding season. About 80% fish fed to the chicks were smaller than 10 cm. Young chicks were offered smaller prey compared with older chicks. The variations in foraging parameters are discussed in relation to habitats and their conservation in the Delhi region.     

The influence of prey size and abundance, and individual phenotype on prey choice by the three-spined stickleback, Gasterosteus aculeatm L.

Prey selection behaviour of three-spined sticklebacks, Gasterosteus aculeatus L., was studied in two experiments. Where possible, the experimental apparatus satisfied the assumptions of the simplest optimal diet model (the basic prey model); prey were presented sequentially, the fish could not search for and handle prey at the same time, and net energy gain, handling time and encounter rate were fixed. Experiment 1 presented fish with a range of Asellus sizes so that pursuit ( p ) and handling ( h ) time could be related to prey size. Published energy values of Asellus together with pursuit and handling times were used to calculate E /( p+h ) for Asellus measuring 3,4,5,6,7 and 9 mm. Pursuit times did not differ with prey size but handling times did. E /( p+h ) was very variable particularly at the larger prey sizes. Experiment 2 presented fish with two sequences of prey differing in the encounter rate with the most profitable prey sizes. Fish did not select the diet predicted by the basic prey model tending to always ignore the largest prey even when net energy gain would have been maximized by including them in the diet. Further analysis showed that the probability of a prey size being taken was a function of prey size, fish stomach fullness and encounter rate. It is concluded that the basic prey model is too simple to capture the behaviour of the fish. One of its main faults is that the changing state of the fish through the feeding bout is ignored.