Predator hunting mode influences patterns of prey use from grazing and epigeic food webs

Multichannel omnivory by generalist predators, especially the use of both grazing and epigeic prey, has the potential to increase predator abundance and decrease herbivore populations. However, predator use of the epigeic web (soil surface detritus/microbe/algae consumers) varies considerably for reasons that are poorly understood. We therefore used a stable isotope approach to determine whether prey availability and predator hunting style (active hunting vs. passive web-building) impacted the degree of multichannel omnivory by the two most abundant predators on an intertidal salt marsh, both spiders. We found that carbon isotopic values of herbivores remained constant during the growing season, while values for epigeic feeders became dramatically more enriched such that values for the two webs converged in August. Carbon isotopic values for both spider species remained midway between the two webs as values for epigeic feeders shifted, indicating substantial use of prey from both food webs by both spider species. As the season progressed, prey abundance in the grazing food web increased while prey abundance in the epigeic web remained constant or declined. In response, prey consumption by the web-building spider shifted toward the grazing web to a much greater extent than did consumption by the hunting spider, possibly because passive web-capture is more responsive to changes in prey availability. Although both generalist predator species engaged in multichannel omnivory, hunting mode influenced the extent to which these predators used prey from the grazing and epigeic food webs, and could thereby influence the strength of trophic cascades in both food webs.     

Evaluating the effects of trophic complexity on a keystone predator by disassembling a partial intraguild predation food web

1. Many taxa can be found in food webs that differ in trophic complexity, but it is unclear how trophic complexity affects the performance of particular taxa. In pond food webs, larvae of the salamander Ambystoma opacum occupy the intermediate predator trophic position in a partial intraguild predation (IGP) food web and can function as keystone predators. Larval A. opacum are also found in simpler food webs lacking either top predators or shared prey. 2. We conducted an experiment where a partial IGP food web was simplified, and we measured the growth and survival of larval A. opacum in each set of food webs. Partial IGP food webs that had either a low abundance or high abundance of total prey were also simplified by independently removing top predators and/or shared prey. 3. Removing top predators always increased A. opacum survival, but removal of shared prey had no effect on A. opacum survival, regardless of total prey abundance. 4. Surprisingly, food web simplification had no effect on the growth of A. opacum when present in food webs with a low abundance of prey but had important effects on A. opacum growth in food webs with a high abundance of prey. Simplifying a partial IGP food web with a high abundance of prey reduced A. opacum growth when either top predators or shared prey were removed from the food web and the loss of top predators and shared prey influenced A. opacum growth in a non-additive fashion. 5. The non-additive response in A. opacum growth appears to be the result of supplemental prey availability augmenting the beneficial effects of top predators. Top predators had a beneficial effect on A. opacum populations by reducing the abundance of A. opacum present and thereby reducing the intensity of intraspecific competition. 6. Our study indicates that the effects of food web simplification on the performance of A. opacum are complex and depend on both how a partial IGP food web is simplified and how abundant prey are in the food web. These findings are important because they demonstrate how trophic complexity can create variation in the performance of intermediate predators that play important roles in temporary pond food webs.     

HSS revisited: multi‐channel processes mediate trophic control across a productivity gradient

Classical food web theory holds that energy channels are regulated by top‐down control with increasing productivity, arising from within‐channel processes. However, these hypotheses do not consider the existence of parallel energy channels linked by shared resource pools and which can fuel generalist predators, imposing trophic control arising from multi‐channel processes. Using 23 large marine food webs, we show that food web responses to increasing productivity are consistent with the apparent trophic cascade hypothesis (ATCH) – with rising productivity predators derive an increasing fraction of their diet from increasingly productive bottom‐up controlled detritus channels, thereby subsidising predator biomass, and in turn strengthening top‐down control in parallel grazing channels. These results testify to a fundamental role of detritus channels specifically and multi‐channel processes in general in mediating food web response to productivity and demonstrate that the ATCH provides an alternative explanation for classical predictions of top‐down control.     

Intraguild interactions among generalist predator functional groups drive impact on herbivore and decomposer prey

Different functional groups of generalist predators may complement each other in controlling prey populations; but intraguild interactions, common among generalist predators, may also reduce the strength of top–down control. In natural communities greater alterations to ecosystem function are expected if a whole functional group declines in abundance or is lost. Therefore studying functional group diversity is important for predicting effects of predator loss. We studied the top–down impact of web‐building spiders, hunting spiders and ants, which are highly abundant generalist predators in most terrestrial ecosystems, on prey from the herbivore and decomposer system of a grassland food web. The density of the three predator groups was manipulated by continuous removal in a three‐factorial designed field experiment, which was carried out for two years. We found no positive effect of increasing predator functional group richness on prey control. However there was evidence for strong composition effects between the functional groups. The presence of ants in predator assemblages reduced the prey suppression through mostly trait‐mediated intraguild interactions, while hunting and web‐building spiders contributed additively to prey suppression and reduced the density of herbivore and decomposer prey by 50–60%. A trophic cascade on plant biomass triggered by web‐builders and hunting spiders was diminished at levels of higher predator group diversity. In conclusion, our experiments showed that intraguild interactions strongly influence the strength of top–down control by generalist predators. Among spiders there was evidence for a positive relation between functional group richness and prey suppression but the overall outcome strongly depended on the occurrence of interference, driven by trait‐mediated indirect interactions.     

Experimental evidence that aboveground predators are sustained by underground detritivores

Detrital infusion into grazing food web is considered to be important in terrestrial communities, but there is hardly any experimental evidence showing that generalist predators aboveground are sustained by belowground detritivores. We established two types of experimental plots in the forest floor, one with sheets on the ground to prevent the emergence of belowground arthropods and the other without sheet, to test the hypotheses that 1) reduced input of detrital arthropods decreases the abundance and species richness of web spiders (major generalist predators in terrestrial ecosystems) and 2) lower number of spiders increases the abundance of herbivorous arthropods. We found that spiders were less abundant in plots where the emergence of detrital arthropods was reduced, while the abundance of herbivores did not significantly increase in these plots. These results provide empirical evidence that organisms moving from underground to aboveground may be important for the maintenance of aboveground predators, although the cascading effect of predator abundance on the grazing food chain was not detected in the present study.     

The Coupling Between Grazing and Detritus Food Chains and the Strength of Trophic Cascades Across a Gradient of Nutrient Enrichment

A minimal food web model was constructed comprising one grazing and one detritus food chain coupled by nutrient cycling and generalist carnivores to investigate how prey preference by carnivores may affect the strength of trophic cascades across a gradient of nutrient enrichment. The equilibrium or mean abundance of each food web component and the magnitude of the carnivore effect on lower trophic levels were calculated for different values of the prey preference and nutrient input parameters. Our model predicts that nutrient enrichment increases the mean abundances of carnivores, autotrophs and detritus, but the magnitude of this effect is dependent on the prey preference term. On the other hand, herbivores and detritivores are relatively unaffected by enrichment but are strongly affected by carnivore preference. Carnivores have a negative effect on herbivores and a positive effect on autotrophs and detritus, whereas the effect on detritivores can be both positive and negative. At high preference for herbivores, carnivores have a positive effect on detritivores, because the positive effect of increased detritus availability due to reduced herbivore grazing outweighs the negative effect of predation. At high preference for detritivores, the balance is changed in the other direction. We argue that in systems where authochtonous primary production is the major source of detritus, herbivores can control the rates of detritus production and have indirect effects on detritivores, which may feed back into effects on herbivores through their shared enemies. This positive feedback is probably one mechanism affecting the resilience of alternative stable states in shallow lakes. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. Author Contributions  J.L.A. conceived the model and the study, J.R. wrote the Matlab programs and ran the simulations and J.L.A. wrote most of the paper.     

Prey preference of top predators manipulates the functioning and stability of multi-trophic ecosystems

The relationship between biodiversity and ecosystem functioning, and the mechanisms underpinning the food web stability, have been intensively investigated in ecological research. The ubiquities of generalists in natural food webs and its important role in dictating these ecosystem properties have been generally recognized. However, how competition between multiple top predators shape these ecosystem properties and determine the success of invasive predators remain largely unexplored. Here, we use a well-developed food web model to investigate the effects of prey preference of top predators on ecosystem functioning and food web stability in both local and invasive conditions. We design several modeling scenarios to mimic combinations of different types of top predators (specialist/generalist) and their origins (local/invasive). Our model theoretically shows that lower exploitation competition for prey between top predators (with distinct prey preferences featured by higher attack rates) would be beneficial for the ecosystem functioning and food web stability. We also demonstrate that the success of top predator invasion depends on the prey preference of both local and invasive top predators. Sensitivity analysis on the model further supports our findings. Our results highlight the importance of prey preference of multiple top predators in manipulating the properties of multi-trophic ecosystems. Our findings may have important implications because the current ongoing global changes profoundly change the phenology of many biological systems and create trophic mismatch, which may manipulate prey preference of top predators and in turn deteriorate ecosystem functioning and food web stability.     

Lake size and fish diversity determine resource use and trophic position of a top predator in high‐latitude lakes

Prey preference of top predators and energy flow across habitat boundaries are of fundamental importance for structure and function of aquatic and terrestrial ecosystems, as they may have strong effects on production, species diversity, and food‐web stability. In lakes, littoral and pelagic food‐web compartments are typically coupled and controlled by generalist fish top predators. However, the extent and determinants of such coupling remains a topical area of ecological research and is largely unknown in oligotrophic high‐latitude lakes. We analyzed food‐web structure and resource use by a generalist top predator, the Arctic charr Salvelinus alpinus (L.), in 17 oligotrophic subarctic lakes covering a marked gradient in size (0.5–1084 km²) and fish species richness (2–13 species). We expected top predators to shift from littoral to pelagic energy sources with increasing lake size, as the availability of pelagic prey resources and the competition for littoral prey are both likely to be higher in large lakes with multispecies fish communities. We also expected top predators to occupy a higher trophic position in lakes with greater fish species richness due to potential substitution of intermediate consumers (prey fish) and increased piscivory by top predators. Based on stable carbon and nitrogen isotope analyses, the mean reliance of Arctic charr on littoral energy sources showed a significant negative relationship with lake surface area, whereas the mean trophic position of Arctic charr, reflecting the lake food‐chain length, increased with fish species richness. These results were supported by stomach contents data demonstrating a shift of Arctic charr from an invertebrate‐dominated diet to piscivory on pelagic fish. Our study highlights that, because they determine the main energy source (littoral vs. pelagic) and the trophic position of generalist top predators, ecosystem size and fish diversity are particularly important factors influencing function and structure of food webs in high‐latitude lakes.     

Effects of trophic skewing of species richness on ecosystem functioning in a diverse marine community

Widespread overharvesting of top consumers of the world's ecosystems has "skewed" food webs, in terms of biomass and species richness, towards a generally greater domination at lower trophic levels. This skewing is exacerbated in locations where exotic species are predominantly low-trophic level consumers such as benthic macrophytes, detritivores, and filter feeders. However, in some systems where numerous exotic predators have been added, sometimes purposefully as in many freshwater systems, food webs are skewed in the opposite direction toward consumer dominance. Little is known about how such modifications to food web topology, e.g., changes in the ratio of predator to prey species richness, affect ecosystem functioning. We experimentally measured the effects of trophic skew on production in an estuarine food web by manipulating ratios of species richness across three trophic levels in experimental mesocosms. After 24 days, increasing macroalgal richness promoted both plant biomass and grazer abundance, although the positive effect on plant biomass disappeared in the presence of grazers. The strongest trophic cascade on the experimentally stocked macroalgae emerged in communities with a greater ratio of prey to predator richness (bottom-rich food webs), while stronger cascades on the accumulation of naturally colonizing algae (primarily microalgae with some early successional macroalgae that recruited and grew in the mesocosms) generally emerged in communities with greater predator to prey richness (the more top-rich food webs). These results suggest that trophic skewing of species richness and overall changes in food web topology can influence marine community structure and food web dynamics in complex ways, emphasizing the need for multitrophic approaches to understand the consequences of marine extinctions and invasions.     

Biodiversity effects of the predation gauntlet

The ubiquity of trophic downgrading has led to interest in the consequences of mesopredator release on prey communities and ecosystems. This issue is of particular concern for reef-fish communities, where predation is a key process driving ecological and evolutionary dynamics. Here, we synthesize existing experiments that have isolated the effects of mesopredators to quantify the role of predation in driving changes in the abundance and biodiversity of recently settled reef fishes. On average, predators reduced prey abundance through generalist foraging behavior, which, through a statistical sampling artifact, caused a reduction in alpha diversity and an increase in beta diversity. Thus, the synthesized experiments provide evidence that predation reduces overall abundance within prey communities, but—after accounting for sampling effects—does not cause disproportionate effects on biodiversity.

     

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.     

Structure of tropical river food webs revealed by stable isotope ratios

Fish assemblages in tropical river food webs are characterized by high taxonomic diversity, diverse foraging modes, omnivory, and an abundance of detritivores. Feeding links are complex and modified by hydrologic seasonality and system productivity. These properties make it difficult to generalize about feeding relationships and to identify dominant linkages of energy flow. We analyzed the stable carbon and nitrogen isotope ratios of 276 fishes and other food web components living in four Venezuelan rivers that differed in basal food resources to determine 1) whether fish trophic guilds integrated food resources in a predictable fashion, thereby providing similar trophic resolution as individual species, 2) whether food chain length differed with system productivity, and 3) how omnivory and detritivory influenced trophic structure within these food webs. Fishes were grouped into four trophic guilds (herbivores, detritivores/algivores, omnivores, piscivores) based on literature reports and external morphological characteristics. Results of discriminant function analyses showed that isotope data were effective at reclassifying individual fish into their pre-identified trophic category. Nutrient-poor, black-water rivers showed greater compartmentalization in isotope values than more productive rivers, leading to greater reclassification success. In three out of four food webs, omnivores were more often misclassified than other trophic groups, reflecting the diverse food sources they assimilated. When fish δ¹⁵N values were used to estimate species position in the trophic hierarchy, top piscivores in nutrient-poor rivers had higher trophic positions than those in more productive rivers. This was in contrast to our expectation that productive systems would promote longer food chains. Although isotope ratios could not resolve species-level feeding pathways, they did reveal how top consumers integrate isotopic variability occurring lower in the food web. Top piscivores, regardless of species, had carbon and nitrogen profiles less variable than other trophic groups.     

Imperfect prey selectivity of predators promotes biodiversity and irregularity in food webs

Ecological communities are often characterised by many species occupying the same trophic level and competing over a small number of vital resources. The mechanisms maintaining high biodiversity in such systems are still poorly understood. Here, we revisit the role of prey selectivity by generalist predators in promoting biodiversity. We consider a generic tri‐trophic food web, consisting of a single limiting resource, a large number of primary producers and a generalist predator. We suggest a framework to describe the predator functional response, combining food selectivity for distinctly different functional prey groups with proportion‐based consumption of similar prey species. Our simulations reveal that intermediate levels of prey selectivity can explain a high species richness, functional biodiversity, and variability among prey species. In contrast, perfect food selectivity or purely proportion‐based food consumption leads to a collapse of prey functional biodiversity. Our results are in agreement with empirical phytoplankton rank‐abundance curves in lakes.     

Seasonal Trophic Niche Shift and Cascading Effect of a Generalist Predator Fish

Few studies have examined how foraging niche shift of a predator over time cascade down to local prey communities. Here we examine patterns of temporal foraging niche shifts of a generalist predator (yellow catfish, Pelteobagrus fulvidraco) and the abundance of prey communities in a subtropical lake. We predicted that the nature of these interactions would have implications for patterns in diet shifts and growth of the predator. Our results show significant decreases in planktivory and benthivory from late spring to summer and autumn, whereas piscivory increased significantly from mid-summer until late autumn and also increased steadily with predator body length. The temporal dynamics in predator/prey ratios indicate that the predation pressure on zooplankton and zoobenthos decreased when the predation pressure on the prey fish and shrimps was high. Yellow catfish adjusted their foraging strategies to temporal changes in food availability, which is in agreement with optimal foraging theory. Meanwhile the decrease in planktivory and benthivory of yellow catfish enabled primary consumers, such as zooplankton and benthic invertebrates, to develop under low grazing pressure via trophic cascading effects in the local food web. Thus, yellow catfish shifts its foraging niche to intermediate consumers in the food web to benefit the energetic demand on growth and reproduction during summer, which in turn indirectly facilitate the primary consumers. In complex food webs, trophic interactions are usually expected to reduce the strength and penetrance of trophic cascades. However, our study demonstrates strong associations between foraging niche of piscivorous fish and abundance of prey. This relationship appeared to be an important factor in producing top-down effects on both benthic and planktonic food webs.     

Sensory cues of a top‐predator indirectly control a reef fish mesopredator

Behavioural trophic cascades highlight the importance of indirect/risk effects in the maintenance of healthy trophic‐level links in complex ecosystems. However, there is limited understanding on how the loss of indirect top–down control can cascade through the food‐web to modify lower level predator–prey interactions. Using a reef fish food‐web, our study examines behavioural interactions among predators to assess how fear elicited by top‐predator cues (visual and chemical stimuli) can alter mesopredator behaviour and modify their interaction with resource prey. Under experimental conditions, the presence of any cue (visual, chemical, or both) from the top‐predator (coral trout Plectropomus leopardus) strongly restricted the distance swum, area explored and foraging activity of the mesopredator (dottyback Pseudochromis fuscus), while indirectly triggering a behavioural release of the resource prey (recruits of the damselfish Pomacentrus chrysurus). Interestingly, the presence of a large non‐predator species (thicklip wrasse Hemigymnus melapterus) also mediated the impact of the mesopredator on prey, as it provoked mesopredators to engage in an ‘inspection’ behaviour, while significantly reducing their feeding activity. Our study describes for the first time a three‐level behavioural cascade of coral reef fish and stresses the importance of indirect interactions in marine food‐webs.     

Mesopredator suppression by an apex predator alleviates the risk of predation perceived by small prey

Predators can impact their prey via consumptive effects that occur through direct killing, and via non-consumptive effects that arise when the behaviour and phenotypes of prey shift in response to the risk of predation. Although predators'' consumptive effects can have cascading population-level effects on species at lower trophic levels there is less evidence that predators'' non-consumptive effects propagate through ecosystems. Here we provide evidence that suppression of abundance and activity of a mesopredator (the feral cat) by an apex predator (the dingo) has positive effects on both abundance and foraging efficiency of a desert rodent. Then by manipulating predators'' access to food patches we further the idea that apex predators provide small prey with refuge from predation by showing that rodents increased their habitat breadth and use of ‘risky′ food patches where an apex predator was common but mesopredators rare. Our study suggests that apex predators'' suppressive effects on mesopredators extend to alleviate both mesopredators'' consumptive and non-consumptive effects on prey.     

Non‐consumptive effects of a top‐predator decrease the strength of the trophic cascade in a four‐level terrestrial food web

The fear of predators can strongly impact food web dynamics and ecosystem functioning through effects on herbivores morphology, physiology or behaviour. While non‐consumptive predator effects have been mostly studied in three‐level food chains, we lack evidence for the propagation of non‐consumptive indirect effects of apex predators in four level food‐webs, notably in terrestrial ecosystems. In experimental mesocosms, we manipulated a four‐level food chain including top‐predator cues (snakes), mesopredators (lizards), herbivores (crickets), and primary producers (plants). The strength of the trophic cascade induced by mesopredators through the consumption of herbivores decreased in the presence of top‐predator cues. Specifically, primary production was higher in mesocosms where mesopredators were present relative to mesocosms with herbivores only, and this difference was reduced in presence of top‐predator cues, probably through a trait‐mediated effect on lizard foraging. Our study demonstrates that non‐consumptive effects of predation risk can cascade down to affect both herbivores and plants in a four‐level terrestrial food chain and emphasises the need to quantify the importance of such indirect effects in natural communities.     

Responses of plants and invertebrate trophic groups to contrasting herbicide regimes in the Farm Scale Evaluations of genetically modified herbicide-tolerant crops

Effects of genetically modified herbicide-tolerant (GMHT) and conventional crop management on invertebrate trophic groups (herbivores, detritivores, pollinators, predators and parasitoids) were compared in beet, maize and spring oilseed rape sites throughout the UK. These trophic groups were influenced by season, crop species and GMHT management. Many groups increased twofold to fivefold in abundance between early and late summer, and differed up to 10-fold between crop species. GMHT management superimposed relatively small (less than twofold), but consistent, shifts in plant and insect abundance, the extent and direction of these effects being dependent on the relative efficacies of comparable conventional herbicide regimes. In general, the biomass of weeds was reduced under GMHT management in beet and spring oilseed rape and increased in maize compared with conventional treatments. This change in resource availability had knock-on effects on higher trophic levels except in spring oilseed rape where herbivore resource was greatest. Herbivores, pollinators and natural enemies changed in abundance in the same directions as their resources, and detritivores increased in abundance under GMHT management across all crops. The result of the later herbicide application in GMHT treatments was a shift in resource from the herbivore food web to the detritivore food web. The Farm Scale Evaluations have demonstrated over 3 years and throughout the UK that herbivores, detritivores and many of their predators and parasitoids in arable systems are sensitive to the changes in weed communities that result from the introduction of new herbicide regimes.     

Augmentation of soil detritus affects the spider community and herbivory in a soybean agroecosystem

If soil detritivores provide a significant prey source for predators in the vegetation, then augmentation of the soil community could affect the grazing food web. Specifically, increases in predator density could enhance any top‐down effects and reduce herbivory. We tested this hypothesis by providing detrital subsidies in the form of composted vegetable matter to 36 m² plots in soybean, Glycine max (L.) Merr. (Fabales: Fabaceae), fields that were managed using either conventional or conservation tillage practices. The foliage‐dwelling spiders, insect predators, and leaf‐chewing insects were censused and the body size of one large spider species, Argiope trifasciata (Forskål) (Araneae: Araneidae), was measured. In addition, the density and size of the plants were assessed and leaf damage was quantified. Any effects of treatments on the palatability of soybean plants to herbivores were determined in two laboratory experiments. Compost increased the density of foliage dwelling spiders and the abdomen size of A. trifasciata. We uncovered no treatment effects on insect predators, herbivorous insects, or plant characteristics except that compost addition reduced leaf damage. In addition, there was a negative correlation across plots between spider abundance and soybean leaf damage and abdomen width of A. trifasciata and weed herbivory levels across plots. These results suggest a connection between the soil community and the foliage food web, but the spiders appear to have exerted a top‐down effect without a shift in herbivore abundance. Further study of the specific seasonality of the herbivores and their behavior in the presence of spiders are needed to uncover the underlying mechanism. Nevertheless, these results provide evidence for complex linkage between the soil and grazing food webs that may be important to biological control.     

Variations in prey consumption of centipede predators in forest soils as indicated by molecular gut content analysis

Predation is an important ecological factor driving animal population structures, community assemblages and consequently ecosystem stability and biodiversity. Many environmental factors influence direction and intensity of predation, suggesting that trophic linkages between animals vary between different habitats. This in consequence has particular relevance in anthropogenically altered habitats such as managed forests, where disturbance regime, tree composition and stand age may change the natural food web structure. We investigated how prey consumption of three common centipede predators (Lithobius spp., Chilopoda), representing two body sizes varies between four differently managed forest types in two regions across Germany. We hypothesized that prey preference of these generalist predators is independent of forest type but rather driven by habitat structure, prey abundance and predator body size. Applying specific PCR assays to test for DNA of three abundant prey groups, i.e. Collembola, Diptera and Lumbricidae, in the predators’ guts, we tracked trophic interactions. The results showed that management type indeed has no influence on centipede prey consumption but depth of litter layer and soil pH. Trophic interactions varied between the two sampled forests regions mainly due to changes in the detection of Lumbricidae and Diptera. Also, effect of litter layer and prey abundance significantly differed between the smaller L. crassipes and the larger L. mutabilis, indicating a body size effect. The results complement food web analyses using fatty acids and stable isotopes by elucidating trophic interactions in soil in unprecedented detail.