A method for in situ estimation of prey selectivity and predation rate in large plankton, exemplified with the jellyfish Aurelia aurita (L.)

A method to estimate predation rates of large predatory zooplankton, such as jellyfish and ctenophores, is outlined. Large plankton size allows direct visual tracking of the predator during the process of foraging. The presented method is novel in the sense that it measures predation rate of a specific individual plankton predator in situ.After prey has been evacuated from the gut of an individual predator, the predator is incubated in situ, and observed by SCUBA-divers who recapture the individual after a defined time. Given that this incubation time is shorter than prey digestion time, predation rate can be calculated as increase in gut content over time. Clearance rates for different prey can be calculated from predation rates and prey concentrations in the water, allowing accurate estimates of prey selectivity. Thus, the problem of unknown feeding history and feeding environment, which can otherwise be a problem in prey selectivity studies of in situ-captured predators, is circumvented. Benefits and limitations of the method are discussed.The method was applied to adult medusae of the common jellyfish Aurelia aurita. A large variation in number of captured prey was detected both among individual jellyfish and among the various oral arms and gastric pouches within individuals. Clearance rates varied strongly with prey type. The medusae selected large crustacean prey (cladocerans and copepods/copepodites) over echinoderm larvae and copepod nauplii. Prey distribution within the medusae indicates that both tentacles and oral arms were used as prey capturing sites. Food passage time from prey capturing organs to gastric pouches was estimated.     

Evaluation of Trail-Cameras for Analyzing the Diet of Nesting Raptors Using the Northern Goshawk as a Model

Diet studies present numerous methodological challenges. We evaluated the usefulness of commercially available trail-cameras for analyzing the diet of Northern Goshawks (Accipiter gentilis) as a model for nesting raptors during the period 2007–2011. We compared diet estimates obtained by direct camera monitoring of 80 nests with four indirect analyses of prey remains collected from the nests and surroundings (pellets, bones, feather-and-hair remains, and feather-hair-and-bone remains combined). In addition, we evaluated the performance of the trail-cameras and whether camera monitoring affected Goshawk behavior. The sensitivity of each diet-analysis method depended on prey size and taxonomic group, with no method providing unbiased estimates for all prey sizes and types. The cameras registered the greatest number of prey items and were probably the least biased method for estimating diet composition. Nevertheless this direct method yielded the largest proportion of prey unidentified to species level, and it underestimated small prey. Our trail-camera system was able to operate without maintenance for longer periods than what has been reported in previous studies with other types of cameras. Initially Goshawks showed distrust toward the cameras but they usually became habituated to its presence within 1–2 days. The habituation period was shorter for breeding pairs that had previous experience with cameras. Using trail-cameras to monitor prey provisioning to nests is an effective tool for studying the diet of nesting raptors. However, the technique is limited by technical failures and difficulties in identifying certain prey types. Our study also shows that cameras can alter adult Goshawk behavior, an aspect that must be controlled to minimize potential negative impacts.     

Orientation of the pit-building antlion larva Euroleon (Neuroptera, Myrmeleontidae) to the direction of substrate vibrations caused by prey

Pit-building antlion Euroleon nostras constructs efficient traps in sand to catch its prey. The predator is known to react to substrate vibrations produced by movements of its prey outside the pit with sand-tossing behaviour but it has not yet been ascertained if this reaction is directed towards the prey. The accuracy of the sand-tossing response in the presence of four prey species was measured using a video recording method. The sand-tossing angle was highly positively correlated with the prey angle. Sand tossing was most frequently elicited when prey was on the posterior sand surface. Covering the larval photoreceptors did not influence the antlion's localizing behaviour.     

A note on the production and use of antisera for the detection of part-digested sticklebacks in predator stomach-content samples

Preliminary experiments indicate that serological techniques provide a useful method of identifying specific prey antigens from the stomach contents of fish predators. Problems with cross-reacting antigens from different prey species can be reduced by absorbing anti-sera with tissue extracts of the cross-reacting species. Very small volumes of part-digested prey tissue are sufficient for identification of the prey species when using agarose diffusion plates (Ouchterlony tests).     

Prey spectra of two swarm-raiding army ant species in East Africa

Although swarm-raiding army ants are considered keystone predators in tropical forest ecosystems, information on the prey spectra of most species is based on anecdotal reports and not on systematic studies with extensive sampling. We analysed prey samples of 18 colonies of the two afrotropical species Dorylus ( Anomma ) molestus and Dorylus ( Anomma ) wilverthi (4289 prey items in total) to examine the prey composition variation within and between species and to determine the best methodology to obtain reliable prey spectrum estimates for a given species, site and season. Variation in prey composition was substantial for D. molestus even within a single site and season, with the biomass proportion of the most important prey type differing by a factor of 12. Conclusions from studies using small samples sizes may thus be misleading. We demonstrate that the method of assessing prey spectra in terms of relative prey item numbers often produces biased results, and therefore recommend relative prey biomass as the more useful parameter. The near absence of earthworms, which always constituted a substantial part of the D. molestus prey, in the diet of D. wilverthi is interpreted to result from subtle differences in swarm-raiding behaviour between these two species, but could alternatively also be due to low availability. Similar studies recording prey composition as biomass proportions and analysing large samples sizes from many colonies are needed to understand the effect of army ant swarm raids on invertebrate communities in afrotropical forests.     

Fresh,frozen or fake: A comparison of predation rates measured by various types of sentinel prey

Arthropod predators and parasitoids support the health and functioning of the world's ecosystems, most notably by supplying biological control services to agricultural landscapes. Quantifying the impact that these organisms have on their prey can be challenging, as direct observation and measurement of arthropod predation is difficult. The use of sentinel prey is one method to measure predator impact; however, despite widespread use, few studies have compared predation on different prey types within a single experiment. This study evaluated the predation rates on four sentinel prey items in grass and wheat fields in south-east Queensland, Australia. Attack rates on live and dead Helicoverpa armigera eggs, and dead H. armigera larvae and artificial plasticine larvae, were compared and the predators that were attracted to each prey type were documented with the use of field cameras. There was no significant difference in predation rates between sentinel eggs, while dead larvae were significantly more attacked than artificial larvae. Prey were attacked by a diverse range of predators, including ants, beetles, various nymph and juvenile insects and small mammals. Different predators were active in grass and crop fields, with predator activity peaking around dawn and dusk. The same trends were observed within and between the two habitats studied, providing a measure of confidence in the sentinel prey method. A range of different sentinel prey types could be suitable for use in most comparative studies; however, each prey type has its own benefits and limitations, and these should be carefully evaluated to determine which is most suitable to address the research questions.     

Feeding behaviour of free-ranging penguins determined by oesophageal temperature

Sea birds play a major role in marine food webs, and it is important to determine when and how much they feed at sea. A major advance has been made by using the drop in stomach temperature after ingestion of ectothermic prey. This method is less sensitive when birds eat small prey or when the stomach is full. Moreover, in diving birds, independently of food ingestion, there are fluctuations in the lower abdominal temperature during the dives. Using oesophageal temperature, we present here a new method for detecting the timing of prey ingestion in free-ranging sea birds, and, to our knowledge, report the first data obtained on king penguins (Aptenodytes patagonicus). In birds ashore, which were hand-fed 2-15 g pieces of fish, all meal ingestions were detected with a sensor in the upper oesophagus. Detection was poorer with sensors at increasing distances from the beak. At sea, slow temperature drops in the upper oesophagus and stomach characterized a diving effect per se. For the upper oesophagus only, abrupt temperature variations were superimposed, therefore indicating prey ingestions. We determined the depths at which these occurred. Combining the changes in oesophageal temperatures of marine predators with their diving pattern opens new perspectives for understanding their foraging strategy, and, after validation with concurrent applications of classical techniques of prey survey, for assessing the distribution of their prey.     

Simple order of prey preference technique for modelling the predator functional response

The predator functional response to several prey types and densities may be conceptualized as a multi-dimensional version of the one-dimensional Holling functional-response curves; however, this empirical approach requires inordinate amounts of data to develop and test. A simulation method of modelling this functional response is to consider the behavior of a predator faced with the choice of several prey types. In this model, when all prey are available the predator’s selection will depend on the absolute abundance of the most-preferred prey type, irrespective of the abundances of the less-preferred prey types. Consequently, the predator will consume only the most-preferred prey types while that type is available in sufficient numbers. When abundance of the most-preferred type declines below a certain level, the predator will begin to include in its diet the second-most-preferred prey type along with the most-preferred prey type. This order-of-preference technique holds up well when the model is compared to population data fromOligonychus pratensis (Acarina: Tetranychidae)/Neoseiulus fallacis (Acarina: Phytoseiidae), and is consistent with optimal foraging theory. Implementation is simple, and the data requirements are reduced to determining the predator’s order of preference and normalizing the nutritional values of the prey types to a single type.     

Adaptive evolution of anti-predator ability promotes the diversity of prey species: critical function analysis

In this paper, with the method of adaptive dynamics and critical function analysis, we investigate the evolutionary diversification of prey species. We assume that prey species can evolve safer strategies such that it can reduce the predation risk, but this has a cost in terms of its reproduction. First, by using the method of critical function analysis, we identify the general properties of trade-off functions that allow for continuously stable strategy and evolutionary branching in the prey strategy. It is found that if the trade-off curve is globally concave, then the evolutionarily singular strategy is continuously stable. However, if the trade-off curve is concave-convex-concave and the prey's sensitivity to crowding is not strong, then the evolutionarily singular strategy may be an evolutionary branching point, near which the resident and mutant prey can coexist and diverge in their strategies. Second, we find that after branching has occurred in the prey strategy, if the trade-off curve is concave-convex-concave, the prey population will eventually evolve into two different types, which can coexist on the long-term evolutionary timescale. The algebraical analysis reveals that an attractive dimorphism will always be evolutionarily stable and that no further branching is possible for the concave-convex-concave trade-off relationship.     

Imaging Beads-Retained Prey Assay for Rapid and Quantitative Protein-Protein Interaction

Conventional Western blot based pull-down methods involve lengthy and laborious work and the results are generally not quantitative. Here, we report the imaging beads-retained prey (IBRP) assay that is rapid and quantitative in studying protein-protein interactions. In this assay, the bait is immobilized onto beads and the prey is fused with a fluorescence protein. The assay takes advantage of the fluorescence of prey and directly quantifies the amount of prey binding to the immobilized bait under a microscope. We validated the assay using previously well studied interactions and found that the amount of prey retained on beads could have a relative linear relationship to both the inputs of bait and prey. IBRP assay provides a universal, fast, quantitative and economical method to study protein interactions and it could be developed to a medium- or high-throughput compatible method. With the availability of fluorescence tagged whole genome ORFs in several organisms, we predict IBRP assay should have wide applications.     

Chew,shake, and tear: Prey processing in Australian sea lions (Neophoca cinerea)

Pinnipeds generally target relatively small prey that can be swallowed whole, yet often include larger prey in their diet. To eat large prey, they must first process it into pieces small enough to swallow. In this study we explored the range of prey‐processing behaviors used by Australian sea lions (Neophoca cinerea) when presented with large prey during captive feeding trials. The most common methods were chewing using the teeth, shaking prey at the surface, and tearing prey held between the teeth and forelimbs. Although pinnipeds do not masticate their food, we found that sea lions used chewing to create weak points in large prey to aid further processing and to prepare secured pieces of prey for swallowing. Shake feeding matches the processing behaviors observed in fur seals, but use of forelimbs for “hold and tear” feeding has not been previously reported for other otariids. When performing this processing method, prey was torn by being stretched between the teeth and forelimbs, where it was secured by being squeezed between the palms of their flippers. These results show that Australian sea lions use a broad repertoire of behaviors for prey processing, which matches the wide range of prey species in their diet.     

Effect of prey mass and selection on predator carrying capacity estimates

The ability to determine the prey-specific biomass intake of large predators is fundamental to their conservation. In the absence of actual prey data, researchers generally use a “unit mass” method (estimated as 3/4 adult female mass) to calculate the biomass intake of predators. However, differences in prey preference and range across geographic regions are likely to have an influence on biomass calculations. Here we investigate the influence of estimated prey mass on leopard biomass calculations, and subsequent carrying capacity estimates, in an understudied mountain population. Potential leopard feeding sites were identified using global positioning system (GPS) location clusters obtained from GPS collars. We investigated 200 potential leopard feeding sites, of which 96 were actual feeding sites. Jaw bones, horns, hooves, and other indicative bones were used to determine gender and age of prey items, which were subsequently used to calculate mass of each prey item based on previously published values. There were significant differences in the biomass values calculated using the traditional unit mass method and the calculated prey masses obtained from leopard feeding sites. However, there were no considerable differences in the carrying capacity estimates using the preferred prey species model and leopard density estimates calculated using a non-biased spatial approach, which suggests that estimating carnivore carrying capacity based on 3/4 adult female masses is a reliable method also for the mountain population in this study.     

Partial consumption of prey: the significance of prey water loss on estimates of biomass intake

Summary A number of studies on the feeding behaviour of sucking predators have estimated the weight of biomass the predator extracts from the prey by measuring the weight change occurring in the prey. This method does not consider that a proportion of the prey weight change is lost to the immediate environment. I examined the spider Diaea sp. feeding on the fruit fly Drosophila immigrans and found that the prey lost approximately 28% more weight than the predator gained. This difference was largely explained by water loss from the prey. My results suggest that water loss, which is not available to the predator, is an important part of prey weight loss. To avoid overestimating predator biomass gain it is necessary to measure the predator weight gain directly or take into account water loss as a component of prey weight change.     

Multiple species‐specific molecular markers using nanofluidic array as a tool to detect prey DNA from carnivore scats

Large carnivore feeding ecology plays a crucial role for management and conservation for predators and their prey. One of the keys to this kind of research is to identify the species composition in the predator diet, for example, prey determination from scat content. DNA‐based methods applied to detect prey in predators’ scats are viable alternatives to traditional macroscopic approaches, showing an increased reliability and higher prey detection rate. Here, we developed a molecular method for prey species identification in wolf (Canis lupus) scats using multiple species‐specific marker loci on the cytochrome b gene for 18 target species. The final panel consisted of 80 assays, with a minimum of four markers per target species, and that amplified specifically when using a high‐throughput Nanofluidic array technology (Fluidigm Inc.). As a practical example, we applied the method to identify target prey species DNA in 80 wolf scats collected in Sweden. Depending on the number of amplifying markers required to obtain a positive species call in a scat, the success in determining at least one prey species from the scats ranged from 44% to 92%. Although we highlight the need to evaluate the optimal number of markers for sensitive target species detection, the developed method is a fast and cost‐efficient tool for prey identification in wolf scats and it also has the potential to be further developed and applied to other areas and large carnivores as well.     

Testing the validity of functional response models using molecular gut content analysis for prey choice in soil predators

Analysis of predator–prey interactions is a core concept of animal ecology, explaining structure and dynamics of animal food webs. Measuring the functional response, i.e. the intake rate of a consumer as a function of prey density, is a powerful method to predict the strength of trophic links and assess motives of prey choice, particularly in arthropod communities. However, due to their reductionist set‐up, functional responses, which are based on laboratory feeding experiments, may not display field conditions, possibly leading to skewed results. Here, we tested the validity of functional responses of centipede predators and their prey by comparing them with empirical gut content data from field‐collected predators. Our predator–prey system included lithobiid and geophilomorph centipedes, abundant and widespread predators of forest soils and their soil‐dwelling prey. First, we calculated the body size‐dependent functional responses of centipedes using a published functional response model in which we included natural prey abundances and animal body masses. This allowed us to calculate relative proportions of specific prey taxa in the centipede diet. In a second step, we screened field‐collected centipedes for DNA of eight abundant soil‐living prey taxa and estimated their body size‐dependent proportion of feeding events. We subsequently compared empirical data for each of the eight prey taxa, on proportional feeding events with functional response‐derived data on prey proportions expected in the gut, showing that both approaches significantly correlate in five out of eight predator–prey links for lithobiid centipedes but only in one case for geophilomorph centipedes. Our findings suggest that purely allometric functional response models, which are based on predator–prey body size ratios are too simple to explain predator–prey interactions in a complex system such as soil. We therefore stress that specific prey traits, such as defence mechanisms, must be considered for accurate predictions.     

Laboratory experiments on prey selection by perch fry (Perca fluviatilis)

SUMMARY. In laboratory experiments, prey selection of zooplankters by starved perch fry ( Perca fluviatilis ) from Windermere, English Lake District, is temporally variable and there is usually, in the short-term, a persistent succession from one actively selected prey species to another. Provided that the prey is visible, the patterns of prey selection are maintained irrespective of the accessibility of the prey. The fish do not optimize their energy intake per effort. The method of observing predation activity by fish shows qualitative and quantitative prey selection as time progresses in great detail.     

Ecological and evolutionary implications of diet in monitor lizards

A survey of 35 species indicates that monitor lizards (Varanus) typically hunt over large areas, search in particular microhabitats, and feed frequently on a wide variety of prey, many of which are relatively small. There is ontogenetic, seasonal, and geographic variation in diet. With some exceptions, invertebrates are the predominant prey, but rare predation on vertebrates is often energetically significant. A few monitors specialize on prey types that occur as occasional items in the diet of species with more generalized diets; these include crabs, snails, orthopterans, lizards, and large mammals. For most species, prey specialization occurs via habitat selection and a variety of prey types and sizes are eaten, as expected for widely searching predators. Comparisons with other anguimorphans suggest that derived features of Varanus are associated with high body temperature and activity levels; specialized chemoreception; and rapid, skillful capture of hidden and/or potentially hard to catch prey. Occasional ingestion of moderately large prey is primitive for Varanoidca (Helodermatidae +Varanidae), accentuating a trend that is perhaps primitive for anguimorphan lizards. Reduction of very large prey prior to ingestion is a derived attribute within Varanus , seen infrequently in several larger species and commonly in V. komodoensis. This study illustrates the synthesis of comparative natural history in a phylogenetic context, a method that addresses the history of organismal change.     

A review of the sentinel prey method as a way of quantifying invertebrate predation under field conditions

Sentinel prey can provide a direct, quantitative measure of predation under field conditions. Live sentinel prey provides more realistic data but rarely allows the partitioning of the total predation pressure; artificial prey is less natural but traces left by different predators are identifiable, making it suitable for comparative studies. We reviewed the available evidence of the use of both types of invertebrate sentinel prey. Fifty‐seven papers used real prey, usually measuring predation on a focal (often pest) species, with studies overwhelmingly from North America. The median predation was 25.8% d^?1. Artificial sentinel prey (45 papers) were used in both temperate and tropical areas, placed more above ground than at ground level. The most commonly used artificial prey imitated a caterpillar. Up to 14 predator groups were identified, registering a median of 8.8% d^?1 predation; half the studies reported only bird predation. Predation on real prey was higher than on artificial ones, but invertebrate predation was not higher than vertebrate predation. Invertertebrate but not vertebrate predation was negatively related to prey size. Predation near the Equator was not higher than at higher latitudes, nor in cultivated than noncultivated habitats. The use of sentinel prey is not yet standardised in terms of prey size, arrangement, exposure period or data reporting. Due to the simplicity and ease of use of the method, such standardisation may increase the usefulness of comparative studies, contributing to the understanding of the importance and level of predation in various habitats worldwide.     

Analysis of otter (Lutra lutra) spraints: Part 1: Comparison of methods to estimate prey proportions; Part 2: Estimation of the size of prey fish

The accuracy of five different methods used in spraint analysis for extimating prey proportions is tested through a feeding experiment with three otters in captivity. Estimated prey proportions are copared to the actual diet of the otters. The results of analysing 362 spraints and the time budget indicated a score-bulk estimate to be the most appropriate method of estimating prey diet and it is concluded that dietary studies using different methods can be compared in broad outlines. The relatins between the methods are confirmed in the analysis of 978 spriaints, collected in the field.     

Diet choice of large carnivores in northern Cameroon

The diet of African wild dogs, lions and spotted hyenas was studied in the woodland savannah of Faro National Park and surrounding hunting zones in northern Cameroon during the dry season. Faecal analysis revealed that the diet of the three large carnivores overlaps considerably. Frequencies of prey remains of large and medium‐sized antelopes were highest, with Buffon's kob being by far the most common prey species. Carnivores seem to consume most abundant prey species. No prey item of domestic livestock was found in the faeces. Faecal analysis has proved to be a useful method to reveal diet choice for large carnivores in northern Cameroon. Detailed information on the distribution and density of large carnivores and their prey is needed to give a better picture of the status of carnivores in northern Cameroon and to help resolve conflicts with livestock. Further investigations into the seasonal variation of predation and other factors structuring the large carnivore community in northern Cameroon are recommended.