Foraging activity of a dominant epigeal predator: molecular evidence for the effect of prey density on consumption

Despite the widely held assumption that ‘generalist’ predators consume most prey available to them, there is a growing body of evidence suggesting otherwise. Generalists are expected to perform well in disturbed areas because they can switch between prey pathways when one food source becomes depleted. Indeed, these predators have the potential to promote diversity by switching to prey in a frequency dependent manner and consume prey groups in relation to local abundance. It is therefore important to understand how predation rates fluctuate as local availability changes. We performed open‐field and mesocosm experiments in a corn and soybean agroecosystem to delineate the role prey density plays in determining predation frequency of a dominant epigeal predator. To track trophic pathways, molecular gut‐content analysis using enzyme‐linked immunosorbent assay (ELISA) was performed to track foraging behavior of the wolf spider Pardosa milvina feeding on dipterans, flies. Extensive monitoring of foraging activity and prey populations revealed that predation varied temporally. Importantly, the frequency of individuals testing positive for flies was lower than predicted when flies were extremely abundant but higher when they were scarce, relative to the prey community as a whole. Furthermore, isolating predators in mesocosms revealed an effect of Diptera density on the likelihood of consumption, as determined by ELISA, only when flies were at low levels (12.5% of prey provided). The molecular results suggest that these spiders do not appear to be consuming flies in a frequency‐dependent manner where the decision to switch between different prey pathways is driven by relative abundance. Rather, selectivity of prey is somewhat independent of variation of other prey groups, which is indicative of their consistent reliance on dipterans and may be related to nutritional requirements and/or capture success.     

Effects of chlorogenic acid-and tomatine-fed caterpillars on the behavior of an insect predator

If generalist insect predators are a selective force contributing to patterns of feeding specialization by insect herbivores, then predators should be deterred from eating allelochemical-fed prey. The attack and feeding behaviors of naive predators (Podisus maculiventris stinkbugs) reared on control caterpillars (Manduca sexta) fed plain diet were compared to experienced predators reared on caterpillars fed tomato allelochemicals. Tomatine-fed prey were found more quickly by both naive and tomatine-experienced predators, and chlorogenic acid-experienced predators were more stimulated to begin searching for prey. However, experienced predators were less likely to attack both chlorogenic acidfed and tomatine-fed caterpillars than were naive predators. These results indicate that allelochemical-fed prey were easier for predators to locate, but allelochemical-containing prey often deterred predation by experienced predtors.     

What you need is what you eat? Prey selection by the bat Myotis daubentonii

Optimal foraging theory predicts that predators are selective when faced with abundant prey, but become less picky when prey gets sparse. Insectivorous bats in temperate regions are faced with the challenge of building up fat reserves vital for hibernation during a period of decreasing arthropod abundances. According to optimal foraging theory, prehibernating bats should adopt a less selective feeding behaviour – yet empirical studies have revealed many apparently generalized species to be composed of specialist individuals. Targeting the diet of the bat Myotis daubentonii, we used a combination of molecular techniques to test for seasonal changes in prey selectivity and individual‐level variation in prey preferences. DNA metabarcoding was used to characterize both the prey contents of bat droppings and the insect community available as prey. To test for dietary differences among M. daubentonii individuals, we used ten microsatellite loci to assign droppings to individual bats. The comparison between consumed and available prey revealed a preference for certain prey items regardless of availability. Nonbiting midges (Chironomidae) remained the most highly consumed prey at all times, despite a significant increase in the availability of black flies (Simuliidae) towards the end of the season. The bats sampled showed no evidence of individual specialization in dietary preferences. Overall, our approach offers little support for optimal foraging theory. Thus, it shows how novel combinations of genetic markers can be used to test general theory, targeting patterns at both the level of prey communities and individual predators.     

Despite prolonged association in closed populations,an intertidal predator does not prefer abundant local prey to novel prey

The diets of predators should reflect interactions between their behavioural and anatomical constraints and the availability and accessibility of prey, although feeding preferences may also reflect adaptation to locally abundant prey, particularly in closed populations. On the south‐east coast of Australia, the whelk Haustrum vinosum (Lamarck, 1822) and its prey communities provide a model system in which to test the effect of variation in prey availability on diet and dietary preferences. Haustrum vinosum is a direct developing species, forming effectively closed populations, with the potential for local adaptation at local and regional scales. Here we show that populations of whelks east and west of a biogeographical barrier encounter different prey assemblages, and have different feeding patterns and apparent prey preferences. We then use a prey choice experiment to test for evidence that H. vinosum from three populations west of the barrier display an inherent preference for its most frequently encountered western prey species, the mussel Brachidontes rostratus (Dunker, 1857), over a novel prey, the barnacle Tesseropora rosea (Krauss, 1848). We detected no prey preference within any population, suggesting past association with B. rostratus did not influence prey selection. Our data support the hypothesis that predators with limited dispersal and high population differentiation are able to maintain flexible generalist foraging patterns, even when they encounter novel prey. © 2013 The Linnean Society of London     

Predation by Pterostichus melanarius (Illiger) (Coleoptera: Carabidae) on immature Rhagoletis mendax Curran (Diptera: Tephritidae) in semi-field and field conditions

Practices that enhance abundance and diversity of generalist predators are often employed with the objective of improving biological control of insect pests. Ground beetles and other predators can prey on blueberry maggot, an important pest of blueberries, when mature larvae pupate in the ground. We conducted mesocosm and field experiments to determine if Pterostichus melanarius, a common predatory ground beetle, lowers maggot numbers in compost mulch or when predator and alternative prey abundances are manipulated. At background (field) densities of alternative prey, increasing densities of P. melanarius did not significantly reduce pest numbers in mesocosms containing compost or soil. When alternative prey were removed from compost, beetles reduced pest numbers by up to 35%. In field experiments, maggot numbers were higher when beetles and other predators were excluded from soil plots, but beetle exclusion had no effect in compost plots where both predator and alternative prey numbers were high. Our results indicate that there can be some reduction of blueberry maggot by P. melanarius and other potential predators when there are few alternative prey. However, despite attracting large numbers of predators compost mulch did not lead to a significant reduction in blueberry maggot; in fact, the high abundance of alternative food associated with compost appeared to interfere with beetle predation on blueberry maggot.     

Effect of emergent aquatic insects on bat foraging in a riparian forest

1. Riparian zones serve several ecological functions for bats. They provide a source of prey and likely provide favourable structural habitats and shelter from predators. Many studies have shown that bats use the space above streams, ponds or riparian vegetation as feeding habitat. These studies, however, have never distinguished between the effects of habitat structure and prey availability on the foraging activities of bats. Such effects can only be distinguished by an experimental approach. We predicted that bat activity along a stream is influenced by the number of emerged aquatic insects. 2. We evaluated the response of terrestrial consumers, insectivorous bats, to changes in the abundance of emergent aquatic insects by conducting a manipulative field experiment. In a deciduous riparian forest in Japan, aquatic insect flux from the stream to the riparian zone was controlled with an insect-proof cover over a 1.2 km stream reach. 3. We estimated the abundance of emergent aquatic and flying terrestrial arthropods near the treatment and control reaches using Malaise traps. The foraging activity of bats was evaluated in both treatment and control reaches using ultrasonic detectors. 4. The insect-proof cover effectively reduced the flux of emergent aquatic insects to the riparian zone adjacent to the treatment reach. Adjacent to the control reach, adult aquatic insect biomass was highest in spring, and then decreased gradually. Terrestrial insect biomass increased gradually during the summer at both treatment and control reaches. 5. Foraging activity of bats was correlated with insect abundance. In spring, foraging activity of bats at the control reach was significantly greater than at the treatment reach, and increased at both sites with increasing terrestrial insect abundance. 6. Our result suggests that the flux of aquatic insects emerging from streams is one of the most important factors affecting the distribution of riparian-foraging bats. As is the case with other riparian consumers, resource subsidies from streams can directly enhance the performance or population density of riparian-dependent bats. To conserve and manage bat populations, it is important to protect not only forest ecosystems, but also adjacent aquatic systems such as streams.     

Interspecific synchrony among foliage-feeding forest Lepidoptera species and the potential role of generalist predators as synchronizing agents

While synchrony among geographically disjunct populations of the same species has received considerable recent attention, much less is known about synchrony between sympatric populations of two or more species. We analyzed time series of the abundance of ten species of spring foliage feeding Lepidoptera sampled over a 25-year period at 20 sites in the Slovak Republic. Six species were free-feeders and four were leaf-rollers as larvae. Twenty-nine percent of interspecific pairs were significantly synchronous and correlations were highest among species exhibiting similar feeding strategies. Similar patterns of interspecific synchrony have been previously demonstrated in several other taxa, and the synchronizing effects of weather and/or specialist predators have been proposed as mechanisms. As an alternative explanation, we explored a model in which two species within the same feeding guild were synchronized by the functional response of generalist predators. In this model, species remained unsynchronized in the complete absence of predation or when predatory pressures were applied to only one species. Pairs of prey species projecting relatively similar search images to predators were more highly synchronized than species with relatively different search images. Prey handling time only influenced synchrony when it was very high relative to the total time prey was exposed to predators. Our model's prediction of greater synchrony among species that project similar search images to predators was in agreement with our field study that showed greater synchrony among species sharing similar larval feeding strategies and morphologies.     

Annual and seasonal changes in diets of martens: evidence from stable isotope analysis

Theory predicts that generalist predators will switch to alternative prey when preferred foods are not readily available. Studies on the feeding ecology of the American marten (Martes americana) throughout North America suggest that this mustelid is a generalist predator feeding largely on voles (Microtus sp.; Clethrionomys sp.). We investigated seasonal and annual changes in diets of martens in response to the changing abundance of small rodents (Peromyscus keeni, and Microtus longicaudus) on Chichagof Island, Southeast Alaska, using stable isotope analysis. We hypothesized that martens would feed primarily on small rodents during years with high abundance of these prey species, whereas during years of low abundance of prey, martens would switch to feed primarily on the seasonally available carcasses of salmon. We also hypothesized that home-range location on the landscape (i.e., access to salmon streams) would determine the type of food consumed by martens, and martens feeding on preferred prey would exhibit better body condition than those feeding on other foods. We live-captured 75 martens repeatedly, from mid-February to mid-December 1992–1994. We also obtained marten carcasses from trappers during late autumn 1991 and 1992, from which we randomly sub-sampled 165 individuals. Using stable isotope ratios and a multiple-source mixing model, we inferred that salmon carcasses composed a large portion of the diet of martens in autumn during years of low abundance of rodents (1991 and 1992). When small rodents were available in high numbers (1993 and 1994), they composed the bulk of the diet of martens in autumn, despite salmon carcasses being equally available in all years. Selection for small rodents occurred only in seasons in which abundance of small rodents was low. Logistic regression revealed that individuals with access to salmon streams were more likely to incorporate salmon carcasses in their diet during years of low abundance of small rodents. Using stable isotope analysis on repeated samples from the same individuals, we explored some of the factors underlying feeding habits of individuals under variable ecological conditions. We were unable to demonstrate that body weights of live-captured male and female martens differed significantly between individuals feeding on marine-derived or terrestrial diets. Therefore, martens, as true generalist predators, switched to alternative prey when their principal food was not readily available on a seasonal or annual basis. Although salmon carcasses were not a preferred food for martens, they provided a suitable alternative to maintain body condition during years when small rodents were not readily available. Received: 1 May 1996 / Accepted: 24 February 1997     

Functional and aggregative responses of harbour seals to changes in salmonid abundance

There is intense debate over the potential impact of seal predation on declining salmon stocks in both the Pacific and Atlantic oceans. However, efforts to model such interactions have been constrained by a lack of data on the functional and numerical responses of these predators. Based upon theory, and data from small-scale terrestrial and freshwater systems, a type 3 functional response is expected to best describe predation by generalist pinnipeds. Similarly, theory also predicts that seal numbers should increase with salmon density in rivers following an aggregative response of predator to prey. We tested these predictions by studying the diet and local density of harbour seals in relation to seasonal variations in the abundance of salmonid in a Scottish river system. As predicted, the abundance of seals in the river was directly related to the abundance of returning salmon, and dietary data supported the type 3 functional response to changes in salmonid abundance. These studies provide empirical support for the use of type 3 response in modelling studies.     

Variable dependence on detrital and grazing food webs by generalist predators: aerial insects and web spiders

Recent studies have shown that organisms from the detritus food web subsidize generalist predators in aboveground food webs, but its significance in space and time is largely unknown. Here we report seasonal dynamics of aerial insects from grazing and detritus food webs in both forest and grassland habitats, and show how these patterns influence the dependence of web spiders on the detritus food web. Detrital insects were more abundant in spring, decreased in summer, and then increased slightly in autumn. This pattern was most conspicuous in Nematocera. Due to different seasonal activity patterns of grazing and detrital insects, the proportion of detrital insects was greater in spring and autumn. Detrital insects were relatively more abundant in the forest than in the grassland. Prey captured by web spiders generally reflected seasonal and spatial patterns of aerial insect abundance. In particular, Leucauge spiders reversed their dependence on the two food webs seasonally. Body size of spiders was negatively correlated with the proportion of detrital prey, suggesting that the detrital subsidy is essential for relatively small predators. This size effect probably resulted from interaction of the following two factors: 1) the maximum body size of prey that can be caught increased with spider body size, 2) larger body size classes of aerial insects included a higher proportion of insects from the grazing food web.     

Insectivorous bats selectively source moths and eat mostly pest insects on dryland and irrigated cotton farms

Insectivorous bats are efficient predators of pest arthropods in agroecosystems. This pest control service has been estimated to be worth billions of dollars to agriculture globally. However, few studies have explicitly investigated the composition and abundance of dietary prey items consumed or assessed the ratio of pest and beneficial arthropods, making it difficult to evaluate the quality of the pest control service provided. In this study, we used metabarcoding to identify the prey items eaten by insectivorous bats over the cotton‐growing season in an intensive cropping region in northern New South Wales, Australia. We found that seven species of insectivorous bat (n = 58) consumed 728 prey species, 13 of which represented around 50% of total prey abundance consumed. Importantly, the identified prey items included major arthropod pests, comprising 65% of prey relative abundance and 13% of prey species recorded. Significant cotton pests such as Helicoverpa punctigera (Australian bollworm) and Achyra affinitalis (cotton webspinner) were detected in at least 76% of bat fecal samples, with Teleogryllus oceanicus (field crickets), Helicoverpa armigera (cotton bollworm), and Crocidosema plebejana (cotton tipworm) detected in 55% of bat fecal samples. Our results indicate that insectivorous bats are selective predators that exploit a narrow selection of preferred pest taxa and potentially play an important role in controlling lepidopteran pests on cotton farms. Our study provides crucial information for farmers to determine the service or disservice provided by insectivorous bats in relation to crops, for on‐farm decision making.     

Diet choice and predator—prey dynamics

Summary We compare the dynamics of predator-prey systems with specialist predators or adaptive generalist predators that base diet choice on energy-maximizing criteria. Adaptive predator behaviour leads to functional responses that are influenced by the relative abundance of alternate prey. This results in the per capita predation risk being positively density-dependent near points of diet expansion. For a small set of parameter values, systems with adaptive predators can be locally stable whereas systems with specialist predators would be unstable. This occurs mainly when alternate prey have low enough profitability that predators cannot sustain themselves indefinitely when feeding on alternate prey. Local stability of systems with adaptive predator behaviour is inversely related to the goodness of fit to optimal diet choice criteria. Hence, typical patterns of partial prey preference are more stabilizing than perfect optimal diet selection. Locally stable systems with adaptive predators are often globally unstable, converging on limit cycles for many initial population densities. The small range of parameter combinations and initial population densities leading to stable equilibria suggest that adaptive diet selection is unlikely to be a ubiquitous stabilizing factor in trophic interactions.     

Of mice and mallards: positive indirect effects of coexisting prey on waterfowl nest success

Coexisting prey species interact indirectly via their shared predators when one prey type influences predation rates of the second prey type. In a temperate system where the predominant shared predator is a generalist, I studied the indirect effects of rodent populations on waterfowl nest success, both within the nesting season among sites and among years. Among six to ten upland fields (14 to 27 ha), mallard ( Anas platyrhynchos ) nest success was positively correlated with rodent abundance in all three years of the study. After removing year effects, mallard nest success remained positively correlated with the relative abundance of rodents. Of the rodent species present, California voles ( Microtus californicus ) were the most important coexisting prey type influencing nest success. Among years, mallard nest success was positively correlated with vole abundance; the asymptotic relationship suggests a threshold response to vole abundance, beyond which predators become satiated and additional voles do little to affect nest success. I tested and rejected three alternative explanations for the observed positive correlation between mallard nest success and rodent abundance that do not involve an indirect effect of coexisting prey populations. The influences of dense nesting cover, nesting density, and predator activity did not explain the observed patterns of nest success. These results suggest that rodent populations buffer predation on waterfowl nests, both within and among years, via the behavioral responses of shared predators to coexisting prey.     

Experimental and observational evidence reveals that predators in natural environments do not regulate their prey: They are passengers,not drivers

Among both ecologists and the wider community there is a tacit assumption that predators regulate populations of their prey. But there is evidence from a wide taxonomic and geographic range of studies that predators that are adapted to co-evolved prey generally do not regulate their prey. This is because predators either cannot reproduce as fast as their prey and/or are inefficient hunters unable to catch enough prey to sustain maximum reproduction. The greater capacity of herbivores to breed successfully is, however, normally restricted by a lack of enough food of sufficient quality to support reproduction. But whenever this shortage is alleviated by a large pulse of food, herbivores increase their numbers to outbreak levels. Their predators are unable to contain this increase, but their numbers, too, surge in response to this increase in food. Eventually both their populations will crash once the food supply runs out, first for the herbivores and then for the predators. Then an “over-run” of predators will further depress the already declining prey population, appearing to be controlling its abundance. This latter phenomenon has led many ecologists to conclude that predators are regulating the numbers of their prey. However, it is the same process that is revealed during outbreaks that limits populations of both predator and prey in “normal” times, although this is usually not readily apparent. Nevertheless, as all the diverse cases discussed here attest, the abundance of predators and their co-evolved prey are both limited by their food: the predators are passengers, not drivers.     

Diet of the snake Philodryas nattereri (Steindachner, 1870) suggests a high plasticity of foraging in north-eastern Brazil

The knowledge about interactions between predators and prey is essential for understanding the natural history of animals, especially snakes, which are cryptic organisms that are difficult to visualize in the wild. This article reports on the predation of lizards, frogs, bats and venomous snakes by the snake Philodryas nattereri, evidencing its generalist feeding habits.     

Dietary overlap and seasonality in three species of mormoopid bats from a tropical dry forest

Competing hypotheses explaining species’ use of resources have been advanced. Resource limitations in habitat and/or food are factors that affect assemblages of species. These limitations could drive the evolution of morphological and/or behavioural specialization, permitting the coexistence of closely related species through resource partitioning and niche differentiation. Alternatively, when resources are unlimited, fluctuations in resources availability will cause concomitant shifts in resource use regardless of species identity. Here, we used next‐generation sequencing to test these hypotheses and characterize the diversity, overlap and seasonal variation in the diet of three species of insectivorous bats of the genus Pteronotus. We identified 465 prey (MOTUs) in the guano of 192 individuals. Lepidoptera and Diptera represented the most consumed insect orders. Diet of bats exhibited a moderate level of overlap, with the highest value between Pteronotus parnellii and Pteronotus personatus in the wet season. We found higher dietary overlap between species during the same seasons than within any single species across seasons. This suggests that diets of the three species are driven more by prey availability than by any particular predator‐specific characteristic. P. davyi and P. personatus increased their dietary breadth during the dry season, whereas P. parnellii diet was broader and had the highest effective number of prey species in all seasons. This supports the existence of dietary flexibility in generalist bats and dietary niche overlapping among groups of closely related species in highly seasonal ecosystems. Moreover, the abundance and availability of insect prey may drive the diet of insectivores.     

Species on the menu of a generalist predator, the eastern red bat (Lasiurus borealis): using a molecular approach to detect arthropod prey

One of the most difficult interactions to observe in nature is the relationship between a predator and its prey. When direct observations are impossible, we rely on morphological classification of prey remains, although this is particularly challenging among generalist predators whose faeces contain mixed and degraded prey fragments. In this investigation, we used a polymerase chain reaction and sequence-based technique to identify prey fragments in the guano of the generalist insectivore, the eastern red bat ( Lasiurus borealis ), and evaluate several hypotheses about prey selection and prey defences. The interaction between bats and insects is of significant evolutionary interest because of the adaptive nature of insect hearing against echolocation. However, measuring the successes of predator tactics or particular prey defences is limited because we cannot normally identify these digested prey fragments beyond order or family. Using a molecular approach, we recovered sequences from 89% of the fragments tested, and through comparison to a reference database of sequences, we were able to identify 127 different species of prey. Our results indicate that despite the robust jaws of L. borealis , most prey taxa were softer-bodied Lepidoptera. Surprisingly, more than 60% of the prey species were tympanate, with ears thought to afford protection against these echolocating bats. Moths of the family Arctiidae, which employ multiple defensive strategies, were not detected as a significant dietary component. Our results provide an unprecedented level of detail for the study of predator–prey relationships in bats and demonstrate the advantages which molecular tools can provide in investigations of complex ecological systems and food-web relationships.     

Spatiotemporal food web dynamics along a desert riparian–upland transition

Increased awareness of spatiotemporal variation in species interactions has motivated the study of temporally-resolved food web dynamics at the landscape level. Empiricists have focused attention on cross-habitat flows of materials, nutrients, and prey, largely ignoring the movement of predators between habitats that differ in productivity (and how predators integrate pulses in resource availability over time). We set out to study seasonal variation in food web interactions between mammalian carnivores and their rodent prey along a riparian–upland gradient in semi-arid southeastern Arizona which features both spatial and temporal heterogeneity in resource availability. Specifically, we tested whether mammalian carnivores spill over from productive, near-river habitats into adjacent, desert-scrub habitats; and if they do, to document the effects of this spillover on rodent communities. Furthermore, we examined seasonal variation in top-down effects by measuring changes in carnivore diet and distribution patterns and rodent populations over time. The results indicate that carnivores track seasonally-abundant resources across the landscape, varying both their diet and movement patterns. In turn, desert-scrub rodent population dynamics track seasonal shifts in carnivore habitat use but not resource availability, suggesting that predation plays a role in structuring rodent communities along the San Pedro River. Further evidence comes from data on rodent community composition, which differs between desert-scrub habitats near and far from the river, despite similarities in resource availability. Our data also suggest that seasonal omnivory helps predators survive lean times, increasing their effects on prey populations. Taken together, these results underscore the importance of spatiotemporal variation in species interactions, highlighting the complexity of natural systems and the need for further detailed studies of food web dynamics.     

Numerical and dietary responses of a predator community in a temperate zone of Europe

The generalist predation hypothesis predicts that the functional responses of generalist predator species should be quicker than those of specialist predators and have a regulating effect on vole populations. New interpretations of their role in temperate ecosystems have, however, reactivated a debate suggesting generalist predators may have a destabilizing effect under certain conditions (e.g. landscape homogeneity, low prey diversity, temporary dominance of 1 prey species associated with a high degree of dietary specialization). We studied a rich predator community dominated by generalist carnivores ( Martes spp., Vulpes vulpes, Felis catus ) over a 6 yr period in farmland and woodland in France. The most frequent prey were small rodents (mostly Microtus arvalis , a grassland species, and Apodemus spp., a woodland species). Alternative prey were diverse and dominated by lagomorphs ( Oryctolagus cuniculus, Lepus europeus ). We detected a numerical response among specialist carnivores but not among generalist predators. The dietary responses of generalist predators were fairly complex and most often dependent on variation in density of at least 1 prey species. These results support the generalist predation hypothesis. We document a switch to alternative prey, an increase of diet diversity, and a decrease of diet overlap between small and medium-sized generalists during the low density phase of M. arvalis . In this ecosystem, the high density phases of small mammal species are synchronous and cause a temporary specializing of several generalist predator species. This rapid functional response may indicate the predominant role of generalists in low amplitude population cycles of voles observed in some temperate areas.     

Stable isotope analyses document intraguild predation in wolf spiders (Araneae: Lycosidae) and underline beneficial effects of alternative prey and microhabitat structure on intraguild prey survival

Intraguild predation (IGP) is common among generalist predators and an important issue in food web theory, because IGP may destabilise communities by increasing extinction of species. Also, IGP may interfere with the effectiveness of generalist predators as biological control agents. In general, occurrence of IGP in laboratory or field studies is inferred from abundance data or direct observations only. We investigated if tracing stable isotopes allows distinction between different types of predation and confirmation of IGP. Wolf spiders were chosen as model organisms for generalist predators; IGP of third instar A. cuneata on second instar P. palustris was investigated in a laboratory experiment. The availability of alternative prey and the complexity of the microhabitat were manipulated, since both factors are thought to facilitate coexistence of predators.
Stable isotope analysis documented predation of A. cuneata on P. palustris and predation on alternative prey by both juveniles. Both the presence of alternative prey and the availability of shelter reduced mortality of juvenile P. palustris during the first week. During the second week mortality increased in complex structure without alternative prey presumably due to enhanced activity and cannibalism among starving P. palustris . Thus, microhabitat complexity and prey abundance may foster coexistence of wolf spiders in the field.
In conclusion, stable isotope analysis was proven a powerful tool to investigate animal behaviour without direct observation. The method allowed disentangling predator feeding behaviour when more than one type of prey was present.