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.     

Effects of prey density,prey size and predator size on rates of feeding by an intertidal predatory gastropod Morula marginalba Blainville (Muricidae), on several species of prey

As a prerequisite for models of foraging behaviour of the whelk, Morula marginalba Blainville (Muricidae), the effects of variation in density of prey on the rate of feeding of the predator were examined in field conditions for three coexisting species of prey. Densities of prey used were those at which the prey, two limpets and a barnacle, occurred naturally in the rocky intertidal habitat.Large limpets, Cellana tramoserica (Sowerby) can resist attacks by predatory gastropods by raising the mantle over the outside of the shell. These experiments showed that no C. tramoserica were killed by Morula marginalba even at very great densities and with no alternative prey present. For the small limpet Patelloida latistrigata (Angas), one of the whelk's most highly preferred prey, juveniles were eaten 1.4 times as fast as adults. Fitting the random predator equation gave greater attack coefficients and shorter handling times for juvenile than adult limpets.Sizes of both predator and prey affected rates of eating barnacles, Tesseropora rosea (Krauss), but not in a simple way. Whelks of 15-mm aperture length ate adult barnacles 4.2 times faster than did 12-mm whelks, but there was no significant difference in the rates at which the two sizes of snail ate juvenile barnacles.Rates of feeding on T. rosea and Patelloida latistrigata increased significantly with prey density. These results form a basis for including the density of prey in models of spatial dispersion of the predatory gastropod Morula marginalba.     

Prey size,prey perishability and group foraging in a social spider

Summary Selection might favor group foraging and social feeding when prey are distributed in patches that do not last long enough for a solitary individual to consume more than a small fraction of them (Pulliam and Millikan 1982; Pulliam and Caraco 1984). Here we considered the foraging behavior of a social spider, Anelosimus eximius, in light of this ephemeral resource hypothesis. This species builds large webs in which members cooperate to capture a wide variety of different sizes and types of prey, many of which are very large. The capture success of this species was very high across all prey sizes, presumably due to the fact that they foraged in groups. Group consumption times in natural colonies for all prey larger than five mm were less than the time that dead insects remained on the plastic sheets that we used as artificial webs. Solitary consumption estimates, calculated from the rate at which laboratory individuals extracted insect biomass while feeding, were the same as the residence times of insects on artificial webs in the field for insects between 6 and 15 mm in length and were significantly longer than the persistence of insects on plastic sheets for all larger insects. Large prey, that contribute substantially to colony energy supplies, appeared to be ephemeral resources for these spiders that could not be consumed by a single spider in the time they were available. These factors made the food intake of one spider in a group less sensitive to scavenging by others and could act to reinforce the social system of this species.     

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.     

Prey size spectra and prey availability of larval and small juvenile cod

The aim of the present study is to describe the prey preference characteristics of cod larvae and assess preference variability in relation to species and size composition of copepod prey. A further aim is to examine the hypothesis that dietary prey size spectra remain the same during the larval stage when viewed on a relative predator/prey size scale. The study is based on stomach analysis of larval/juvenile cod in the size range 10–35 mm from nursery grounds in the North Sea. Stomach contents (species, size) were compared to environmental composition and preference indices were calculated. Prey size spectra had the expected relationship to larval cod size, and preference for given copepod species could be ascribed to their relative size. Additional species-specific preferences were evident, for example the larger Pseudocalanus and the larger Calanus spp. were highly preferred. Available prey biomass was highest in the areas of a hydrographic front, where larvae have been shown to concentrate. Changes in prey availability with larval growth depend strongly on specific prey biomass spectra at a given location. Both increasing and decreasing prey availability at increasing larval size were indicated, dependent on location. The findings illustrate the usefulness of coupling dietary prey size spectra and biomass spectra of available prey sizes during studies of ichthyoplankton feeding ecology.     

Prey capture and processing behaviors vary with prey size and shape in Australian and subantarctic fur seals

When hunting at sea, pinnipeds should adapt their foraging behaviors to suit the prey they are targeting. We performed captive feeding trials with two species of otariid seal, Australian fur seals (Arctocephalus pusillus doriferus) and subantarctic fur seals (Arctocephalus tropicalis). This allowed us to record detailed observations of how their foraging behaviors vary when presented with prey items that cover the full range of body shapes and sizes encountered in the wild. Small prey were captured using suction alone, while larger prey items were caught in the teeth using raptorial biting. Small fish and long skinny prey items could then be swallowed whole or processed by shaking, while all prey items with body depths greater than 7.5 cm were processed by shaking at the water's surface. This matched opportunistic observations of feeding in wild Australian fur seals. Use of “shake feeding” as the main prey processing tactic also matches predictions that this method would be one of the only tactics available to aquatic tetrapods that are unable to secure prey using their forelimbs.     

Prey Selection by the Lady Beetle Harmonia axyridis: The Influence of Prey Mobility and Prey Species

The influence of prey mobility and species on prey selection by the coccinellid Harmonia axyridis Pallas was determined under laboratory conditions for two prey species, Hyaliodes vitripennis (Say) and Tetranychus urticae Koch. Prey selection was influenced by prey mobility. In the presence of active prey, the coccinellid selected T. urticae while in presence of immobilized prey, H. vitripennis was preferred. Harmonia axyridis searching time was longer in the presence of active H. vitripennis than in the presence of active T. urticae. Moreover, the coccinellid capture rate was lower for active H. vitripennis caused by effective defensive mechanisms. Prey suitability was affected by prey mobility and species. Immobilized H. vitripennis were the most profitable prey, i.e. induced a shorter developmental time and no mortality. However, active H. vitripennis were not a suitable food source for H. axyridis. Our results suggested that three factors are involved in prey selection by H. axyridis: (i) prey mobility; (ii) prey defensive mechanisms; and (iii) prey species.     

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.     

Prey selection by a spider wasp,Batozonellus lacerticida (Hymenoptera: Pompilidae): Effects of seasonal variation in prey species,size and density

Prey selection by a spider wasp,Batozonellus lacerticida, was investigated at the riverside of the Toyohira River in Misumai, Sapporo, Hokkaido over 7 years from 1981 to 1987. Seventeen species of araneid spiders were found in the study area and six of them were hunted by the wasps. Most (97.3%) of the 223 prey records obtained were of three species,Araneus macacus, A. marmoreus andA. pinguis. Nesting activity of the wasps continued until mid-August in 1981, 1982 and 1986, but stopped in July in 1984, 1985 and 1987. The length of nesting period was related to whether the wasps successfully switched prey species fromA. macacus toA. marmoreus andA. pinguis andA. pinguis or not Switching seemed to occur at the phase when density ofA. macacus remained high, depending on the density of largerA. marmorus andA. pinguis. This switching had large effects on species composition and size distribution of actual prey. Chesson's index α calculated at every 10 day period revealed that the wasps preferredA. macacus to the two other species and large prey to medium and small ones. The effect of prey density on preference depended on whetherA. macacus was present or absent. In the presence of it there were only slight differences in preference among various conditions of prey density. However, whenA. macacus was absent, densities of bothA. marmoreus andA. pinguis had some complementary effects on preference. The importance of prey size selection by pompilid wasps, and implications of density effects on preference are discussed.     

Prolonging the arctic pulse: long-term exploitation of cached eggs by arctic foxes when lemmings are scarce

1. Many ecosystems are characterized by pulses of dramatically higher than normal levels of foods (pulsed resources) to which animals often respond by caching foods for future use. However, the extent to which animals use cached foods and how this varies in relation to fluctuations in other foods is poorly understood in most animals. 2. Arctic foxes Alopex lagopus (L.) cache thousands of eggs annually at large goose colonies where eggs are often superabundant during the nesting period by geese. We estimated the contribution of cached eggs to arctic fox diets in spring and autumn, when geese were not present in the study area, by comparing stable isotope ratios (delta(13)C and delta(15)N) of fox tissues with those of their foods using a multisource mixing model in Program IsoSource. 3. The contribution of cached eggs to arctic fox diets was inversely related to collared lemming Dicrostonyx groenlandicus (Traill) abundance; the contribution of cached eggs to overall fox diets increased from < 28% in years when collared lemmings were abundant to 30-74% in years when collared lemmings were scarce. 4. Further, arctic foxes used cached eggs well into the following spring (almost 1 year after eggs were acquired) - a pattern that differs from that of carnivores generally storing foods for only a few days before consumption. 5. This study showed that long-term use of eggs that were cached when geese were superabundant at the colony in summer varied with fluctuations in collared lemming abundance (a key component in arctic fox diets throughout most of their range) and suggests that cached eggs functioned as a buffer when collared lemmings were scarce.     

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.     

Measurement of body size and abundance in tests of macroecological and food web theory

1. Mean body mass (W) and mean numerical (N) or biomass (B) abundance are frequently used as variables to describe populations and species in macroecological and food web studies. 2. We investigate how the use of mean W and mean N or B, rather than other measures of W and/or accounting for the properties of all individuals, can affect the outcome of tests of macroecological and food web theory. 3. Theoretical and empirical analyses demonstrate that mean W, W at maximum biomass (W(mb)), W when energy requirements are greatest (W(me)) and the W when a species uses the greatest proportion of the energy available to all species in a W class (W(mpe)) are not consistently related. 4. For a population at equilibrium, relationships between mean W and W(me) depend on the slope b of the relationship between trophic level and W. For marine fishes, data show that b varies widely among species and thus mean W is an unreliable indicator of the role of a species in the food web. 5. Two different approaches, 'cross-species' and 'all individuals' have been used to estimate slopes of abundance-body mass relationships and to test the energetic equivalence hypothesis and related theory. The approaches, based on relationships between (1) log(10) mean W and log(10) mean N or B, and (2) log(10) W and log(10) N or B of all individuals binned into log(10) W classes (size spectra), give different slopes and confidence intervals with the same data. 6. Our results show that the 'all individuals' approach has the potential to provide more powerful tests of the energetic equivalence hypothesis and role of energy availability in determining slopes, but new theory and empirical analysis are needed to explain distributions of species relative abundance at W. 7. Biases introduced when working with mean W in macroecological and food web studies are greatest when species have indeterminate growth, when relationships between W and trophic level are strong and when the range of species'W is narrow.     

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.     

Size-dependent predation by snakes: selective foraging or differential prey vulnerability?

I staged replicate encounters between unrestrained lizards andsnakes in outdoor enclosures to examine size-dependent predationwithin the common garden skink (Lampropholis guichenoti). Yellow-facedwhip snakes (Demansia psammophis) forage widely for activeprey and most often consumed large skinks, whereas death adders(Acanthophis antarcticus) ambush active prey and most oftenconsumed small skinks. Small-eyed snakes (Rhinoplocephalusnigrescens) forage widely for inactive prey and consumed bothsmall and large skinks equally often. Differential predationmay reflect active choice by the predator, differential preyvulnerability, or both. To test for active choice, I presentedforaging snakes with an inert small lizard versus an inertlarge lizard. They did not actively select lizards of a particularbody size. To test for differential prey vulnerability, I quantifiedvariation between small and large lizards in behavior thatis important for determining the outcome of predator—prey interactions. Snakes did not differentiate between integumentarychemicals from small and large lizards. Large lizards tendto flee from approaching predators, thereby eliciting attackby the visually oriented whip snakes. Small lizards were moremobile than large lizards and therefore more likely to passby sedentary death adders. Additionally, small skinks were more effectively lured by this sit-and-wait species and less likelyto avoid its first capture attempt. In contrast, overnightretreat site selection (not body size) determined a lizard'schances of being detected by small-eyed snakes. Patterns ofsize-dependent predation by elapid snakes may arise not becauseof active choice but as a function of species-specific predatortactics and prey behavior.     

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.     

The effects of prey relative abundance and chemical cues on prey selection in rainbow trout

An optimal foraging model was used to predict prey selection based on both energy maximization and number maximization strategies. The influence of chemical cues and relative abundance on rainbow trout diet selection was examined under laboratory conditions.
In most fish, diet composition was strongly influenced by chemical cues. No fish followed an energy maximization strategy, and selection of prey based on taste persisted despite large caloric penalties associated with these choices. In the absence of chemical cues, diet composition was based on relative abundance of prey (a number maximization strategy). Within the feasible constraints of the optimal foraging model lie a large number of possible diet combinations which would provide sufficient energy for growth and reproduction. This provides a wide scope for feeding flexibility. Response to prey chemical cues may be the basis for observations of individual diet variability in trout.     

Body size, prey handling efficiency and choice of diet in three coniferous forest tits (Paridae)

The main purpose of this study was to link morphological differences between great tit ( Parus major ), willow tit ( P. montanus ) and coal tit ( P. ater ) and their rate of energy acquisition and choice of diet in order to explore the potential for competitive relations between them more directly. Handling times were measured in the laboratory by presenting mealworms of different sizes to the birds. Great tits were more efficient in handling large prey than were the smaller-bodied willow- and coal tits; for small prey sizes the coal tit was the least efficient species. Using the ratio of prey mass to the handling time value, a utility function for each species was constructed. These results suggests a potential for a segregation of the species on the food axis. However, results from the prey choice experiment show that despite considerable differences in functional morphology between the three species they do not differ significantly in the range of prey size exploited. My results suggest that the alleged importance of prey size partitioning is not likely to play the major role for the coexistence of these coniferous forests tits.     

Optimal foraging constrains macroecological patterns: body size and dietary niche breadth in lizards

Aim  To explore and identify probable mechanisms contributing to the relationships among body size, dietary niche breadth and mean, minimum, maximum and range of prey size in predaceous lizards.
Location  Our data set includes species from tropical rainforests, semi-arid regions of Brazil, and from deserts of the south-western United States, Australia and the Kalahari of Africa.
Methods  We calculated phylogenetic and non-phylogenetic regressions among predator body size, dietary breath and various prey size measures.
Results  We found a negative association between body size and dietary niche breadth in 159 lizard species sampled across most evolutionary lineages of squamate reptiles and across major continents and habitats. We also show that mean, minimum, maximum and range of prey size were positively associated with body size.
Main conclusions  Our results suggest not only that larger lizards tend to eat larger prey, but in doing so offset their use of smaller prey. Reduction of dietary niche breadth with increased body size in these lizards suggests that large predators target large and more profitable prey. Consequently, the negative association between body size and niche breadth in predators most likely results from optimal foraging. Though this result may appear paradoxical and runs counter to previous studies, resources for predators may be predictably more limited than resources for herbivores, thus driving selection for more profitable prey.     

Effects of prey density, prey instar, and patch size on the development of the predatory mosquito, Toxorhynchites towadensis

Effects of prey density, prey instar, and patch size on the development of the predatory mosquito larva, Toxorhynchites towadensis, were investigated in the laboratory. Survivors of T. towadensis showed different developmental patterns in relation to prey age structure. All predatory larvae in containers with only second instar prey developed into the third instar. However, in several containers with fourth instar prey, mortality of predators was observed. During the third instar, no predatory larva died, but both prey density and prey instar significantly affected the survival of predators during their fourth instar. Large prey size promoted large predator adults, and predatory larvae which grew up in small surface containers responded by developing to large sizes than those in large containers. Larval developmental time of the predators differed in each treatment. During first and second instars, faster predator development was observed in containers with small surface areas and containing young prey individuals. However, when development was enhanced by the presence of old prey individuals, no surface effect was observed. The fastest predator development was observed with prey of mixed instars and high density. This study suggests that a small surface container containing prey of mixed instars and high density is suitable for development of predators.