Cereal aphid control by generalist predators in presence of belowground alternative prey: Complementary predation as affected by prey density

Generalist predators are important antagonists of pest species in agroecosystems. Increasing populations of alternative prey through detrital subsidies is one way to foster these predators. However, alternative prey may also distract generalist predators from their prey thereby diminishing the efficiency of biological control. To develop reliable predictions for biological control, it is essential to evaluate the relative importance of generalist predators, pests, alternative prey and their respective interactions. We investigated the effects of an assemblage of generalist predators on the grain aphid Sitobion avenae in winter wheat. Treatments with 10, 100 and 1000 aphids were established inside 2 m² sized caged plots with control and reduced density of predators. Three weeks after the experiment started, samples were taken to estimate the size of aphid populations and those of alternative prey. Three prey taxa were significantly reduced by generalist predators: the grain aphid S. avenae (Aphidina), the click beetle Adrastus pallens (Elateridae) and the springtail Isotoma viridis (Collembola). Springtails were decreased by generalist predators independent of aphid densities, indicating complementary predation of springtails and aphids. At high aphid densities, grain aphid population peaks were decreased to the threshold level of economic damage, demonstrating efficient aphid suppression by the predator community. Click beetle numbers declined only at low and medium aphid densities. The results suggest that generalist predators preferentially fed on click beetles at low and medium aphid densities and switched to aphids at high aphid densities. Early-season predators likely had the greatest influence on aphid suppression. Our results indicate that alternative prey from the belowground system forms a substantial food resource for generalist predators, suggesting that the belowground subsystem modulates predator–prey interactions above the ground.     

Trophic interactions in centipedes (Chilopoda,Myriapoda) as indicated by fatty acid patterns: Variations with life stage,forest age and season

Fatty acid (FA) analysis investigates changes in the relative contribution of prey from major energy channels in decomposer food webs for predator nutrition. Adopting this approach we investigated whether the trophic niche of centipedes, as major invertebrate predators in forest soil food webs, changes with maturation, season or forest age. Generally, each of the four centipede species studied differed significantly in their FA composition suggesting trophic niche differentiation. FA profiles differed more strongly in the two geophilomorph (Strigamia acuminata and Geophilus ribauti) than in the two lithobiomorph species (Lithobius crassipes and Lithobius mutabilis) suggesting that in particular the former feed on markedly different prey. FA profiles changed during post-embryonic development in each of the four centipede species. Differences were most pronounced in the two lithobiomorph species shifting to predominantly fungal feeding prey. Further, FA profiles varied with season indicating that centipedes exploit more prey out of the bacterial channel in autumn. FA profiles of centipedes varied little with forest age suggesting that soil food webs are remarkably invariant across different forest ecosystems. The results indicate that FA composition of second order consumers closely reflects changes in diet of prey species and composition of basal resources. The study proved FA profiles as powerful tool to gain insight into critical characteristics of soil food web stability, i.e., compartmentalisation and the relative importance and variability of energy channels.     

Predator-prey relationships and responses of ungulates and their predators to the establishment of protected areas: A case study of tigers, leopards and their prey in Bardia National Park, Nepal

Interactions among sympatric large predators and their prey and how they respond to conservation measures are poorly known. This study examines predictions concerning the effects of establishing a protected area in Nepal on tigers (Panthera tigris), leopards (Panthera pardus), and their ungulate prey. Within a part of the park, after 22 years the total density of wild ungulates had increased fourfold, to ca. 200 animals/km², almost exclusively due to a remarkable increase in chital deer (Axis axis). Tiger density also increased markedly to nearly 20 animals/100 km², whereas leopard density did not and was ca. 5 animals/100 km². The prediction that grazers should increase more than browsers was only partially supported. The prediction of positive density dependence in prey selection was not supported. Instead, the most abundant species (chital and hog deer, Axis porcinus) were killed less frequently than expected, whereas the lower-density wild boar (Sus scrofa) was preferred. Predictions that (i) initially rare species suffer highest predation was partially supported, that (ii) predation is highest among the most abundant prey was not supported, and that (iii) predation is highest among the most preferred prey independently of their densities was supported. Clearly, the conservation efforts adopted in Bardia were successful, as both tigers and their natural prey base increased. However, the positive numerical response of tigers limited and depressed the abundance of some prey species. Thus, conservation activities aimed at restoring large predators are likely to change in the composition of the overall mammal community, potentially eliminating rare but preferred prey species.     

Molecular detection of nematode predation and scavenging in oribatid mites: Laboratory and field experiments

Recent stable isotope analyses indicate that a number of putative detritivorous soil microarthropods is not typical detritivores but rather live as predators or scavengers. Using molecular gut content analyses the present study investigates if nematodes indeed form part of the diet of oribatid mites. First, in a no-choice laboratory feeding experiment two nematode species (Phasmarhabditis hermaphrodita and Steinernema feltiae) were offered to eight species of oribatid mites and one gamasid mite. Second, after feeding for 4 and 48 h on each nematode species the detection time of prey DNA in the oribatid mite species Steganacarus magnus was investigated. Third, in a field experiment nematode prey (P. hermaphrodita and S. feltiae) in the diet of microarthropods was investigated distinguishing between scavenging and predation. In the no-choice laboratory experiment not only the gamasid mite but also several of the studied oribatid mite species consumed nematodes. After feeding on nematodes for 4 h prey DNA was detectable in S. magnus for only 4 h, but after feeding for 48 h prey DNA was detectable for 128 h, indicating that the duration of feeding on prey is an important determinant for prey DNA detection. The field experiment confirmed that oribatid mite species including Liacarus subterraneus, Platynothrus peltifer and S. magnus intensively prey on nematodes. Interestingly, DNA of dead P. hermaphrodita was detectable to a similar degree as that of living individuals indicating that scavenging is of significant importance in decomposer food webs. Results of our study indicate that predation and scavenging on nematodes by “detritivorous” microarthropods in soil food webs need to be reconsidered.     

Compartmentalization of the soil animal food web as indicated by dual analysis of stable isotope ratios (N/N and C/C)

The soil animal food web has become a focus of recent ecological research but trophic relationships still remain enigmatic for many taxa. Analysis of stable isotope ratios of N and C provides a powerful tool for disentangling food web structure. In this study, animals, roots, soil and litter material from a temperate deciduous forest were analysed. The combined measurement of δ¹⁵N and δ¹³C provided insights into the compartmentalization of the soil animal food web. Leaf litter feeders were separated from animals relying mainly on recent belowground carbon resources and from animals feeding on older carbon. The trophic pathway of leaf litter-feeding species appears to be a dead end, presumably because leaf litter feeders (mainly diplopods and oribatid mites) are unavailable to predators due to large size and/or strong sclerotization. Endogeic earthworms that rely on older carbon also appear to exist in predator-free space. The data suggest that the largest trophic compartment constitutes of ectomycorrhizal feeders and their predators. Additionally, there is a smaller trophic compartment consisting of predators likely feeding on enchytraeids and potentially nematodes.     

Scavenging and active predation in generalist predators: A mesocosm study employing DNA-based gut content analysis

Scavenging presumably is common among invertebrate generalist predators, however, only few studies have specifically dealt with the consumption of dead prey. Here consumption of dead and living prey within a guild of generalist predators was investigated using mesocosm experiments and DNA-based gut content analysis. A community of predators including carabids, staphylinids and spiders was released into 0.2 m² mesocosms, planted with wheat and infested with grain aphids Sitobion avenae. At predator release, freshly killed bird cherry-oat aphids, Rhopalosiphum padi, were provided as carrion prey at the soil surface. Two days after predator release, predators were collected and their gut content screened for DNA of both aphid species using multiplex PCR. Comparing detection frequencies of DNA from living and dead prey, we found that all predators frequently consumed dead aphids, except for the lycosid spider Trochosa ruricola which exclusively fed on S. avenae. In contrast, in the tetragnathid spider Pachygnatha degeeri, supposed to feed mainly on living prey, detection rates of R. padi DNA were high. Our findings support previous assumptions that many generalist predators are facultative scavengers. This needs to be considered for the interpretation of field-derived data using approaches of gut content analyses as well as when assessing predators’ pest control potential.     

Implications of intraguild predation for sea turtle nest protection

In the United States, raccoons Procyon lotor are often removed from sea turtle nesting beaches to decrease egg mortality. However, raccoons also consume ghost crabs Ocypode quadrata, another common egg predator. Reducing predator populations can benefit secondary predators, inflating total predation pressure and leading to a decline in prey species. We used track and burrow counts to compare raccoon and ghost crab abundance at four beaches in Florida, USA, that differ in management activity and determined predation rates on loggerhead Caretta caretta nests by each predator. Mean raccoon abundance (range 0.12-0.46 tracks plot⁻¹ night⁻¹) and ghost crab density (0.09-0.19 burrows m⁻²) were inversely correlated. Ghost crabs were largest at the site with the fewest raccoons. The stable nitrogen isotope ratios of ghost crabs (mean 9.8‰) were positively correlated with body mass, indicating larger ghost crabs feed at a higher trophic level and suggesting large ghost crabs may consume more loggerhead eggs. The highest rates of egg predation by both predators (31%) occurred where raccoon abundance was lowest and ghost crab abundance was highest, suggesting ghost crab burrows may facilitate predation by raccoons. Our data suggest that predation by raccoons limits ghost crabs and that removing raccoons can increase ghost crab abundance and sea turtle egg mortality. Although predator removal can be effective when nest predation rates are quite high, maintaining moderate raccoon densities may be important for controlling ghost crabs. These results highlight the importance of understanding food web connectivity in developing management strategies to achieve conservation goals, especially when the species of concern are threatened or facing extinction.     

Trophic structure of a macroarthropod litter food web in managed coniferous forest stands: a stable isotope analysis with δN and δC

We studied the composition of a litter detrital community in a temperate coniferous forest using stable isotopes of nitrogen and carbon. Samples of mineral soil, bulk litter material, macroarthropods and understory plants were collected from ten experimental forest stands. Half of the stands were previously thinned 17–42 years ago, the other half served as controls. Values of δ¹⁵N and δ¹³C were based on the analysis of almost 500 individuals of at least 22 species in 11 arthropod families. The isotopic analysis showed a significant increase in δ¹⁵N and δ¹³C values with soil depth. Isotopic signatures of macroarthropods ranged from −26.51‰ to −20.52‰ for δ¹³C and −2.85‰ to 5.10‰ for δ¹⁵N. All consumers showed levels of ¹³C enrichment substantially higher than those of primary producers and litter. Predators were generally significantly more ¹⁵N enriched than detritivores and herbivores, but their δ¹³C levels were similar to those of primary consumers. Our data indicate that this community consists of at least 2–3 trophic levels with a considerable amount of variation in the ¹⁵N enrichment among detritivores and predators. We suggest that the spread of δ¹⁵N values of predators likely reflects the diversity of potential prey among detritivores and a varying degree of intraguild predation among different species. Our findings generally agree closely with the results of similar studies from other forest litter communities. Thinning did not appear to influence the overall isotopic composition of the detrital food web. Extensive omnivory and intraguild predation among litter consumers may buffer long-term effects of thinning on the trophic structure of these species-rich communities.     

Taking it to the next level: Trophic transfer of marker fatty acids from basal resource to predators

Fatty acid (FA) analysis is used increasingly to investigate the trophic structure of soil animal food webs as the technique allows separation of the role of detrital resources such as bacteria, fungi and plant material for consumer nutrition. The applicability of FAs as biomarkers for different diets has been verified for Collembola and Nematoda. However, for the analysis of whole food webs it is crucial to know whether marker FA are valid for different taxa and whether they are transferred along the food chain to higher trophic levels, i.e. predators. Top-predators are integrators of lower level energy fluxes in food webs; therefore analysis of their FAs may allow to identify trophic pathways and to separate bacterial vs. fungal based energy channels. Chilopoda and Arachnida are among the main predators in soil food webs. Our aim was to test the applicability of marker FAs for these two predator taxa and to verify the trophic transfer of marker FAs of different basal resources via first order consumers into predators, i.e. over three trophic levels. Therefore, we investigated the transfer of FAs from different basal resources [fungi (Chaetomium globosum), plant leaf litter (Tilia europaea), Gram-positive (Bacillus amyloliquefaciens) and Gram-negative bacteria (Stenotrophomonas maltophilia)] via Collembola (Heteromurus nitidus) as first order consumers into predators [Lithobius forficatus (Chilopoda) and Pardosa lugubris (Arachnida)]. Fatty acid profiles of predators of food chains with different basal resources differed significantly. Marker FAs of basal resources were clearly detectable in predators, suggesting that FA analysis allows to separate trophic channels of soil food webs. By reflecting basal resources, FAs of predators allow tracking energy/resource fluxes through the food web and thereby clarifying the relative importance of bacterial vs. fungal vs. plant resources for soil animal food webs.     

Abundance,diversity and connectance of soil food web channels along environmental gradients in an agricultural landscape

Soil food webs respond to anthropogenic and natural environmental variables and gradients. We studied abundance, connectance (a measure of the trophic interactions within each channel), and diversity in three different channels of the soil food web, each comprised of a resource-consumer pair: the microbivore channel (microbes and their nematode grazers), the plant–herbivore channel (plants and plant-feeding nematodes), and the predator–prey channel (predatory nematodes and their nematode prey), and their associations with different gradients in a heterogeneous agricultural landscape that consisted of intensive row crop agriculture and grazed non-irrigated grasslands in central California. Samples were taken at three positions in relation to water channels: water’s edge, bench above waterway, and the adjacent arable or grazed field. Nematode communities, phospholipid fatty acid (PLFA) biomarkers, and soil properties (NH₄⁺-N, NO₃⁻-N, total N, total C, pH, P, bulk density and soil texture) were measured, and riparian health ratings were scored. Environmental variables were obtained from publicly-available data sources (slope, elevation, available water capacity, erodability, hydraulic conductivity, exchangeable cation capacity, organic matter, clay and sand content and pH).The abundance and richness in most food web components were higher in grazed grasslands than in intensive agricultural fields. Consumers contributed less than their resources to the abundance and richness of the community in all channels. The association between richness and abundance for each component was strongest for the lowest trophic links (microbes, as inferred by PLFA) and weakest for the highest (predatory nematodes). The trophic interactions for the predator–prey and plant–herbivore channels were greater in the grassland than in the cropland. Fields for crops or grazing supported more interactions than the water’s edge in the plant–herbivore and microbivore channels. Connectance increased with the total richness of each community. Higher connectance within the microbivore and predator–prey soil food web channels were associated with soil NO₃⁻-N and elevation respectively, which served as surrogate indicators of high and low agricultural intensification.     

Preservation of wide-ranging top predators by site-protection: Black and red kites in Doñana National Park

We used data from a long-term study on two medium-sized generalist predators, the black kite (Milvus migrans) and the red kite (Milvus milvus), to illustrate the complexity of managing wide-ranging top predators by site-protection. The study was conducted between 1989 and 2000 in the Reserva Biológica de Doñana, located at the core of Doñana National Park. Both species occurred at high density and showed high productivity. Black kites were slightly increasing, and red kites showed a non-significant trend. The main cause of breeding failure was nest predation, mainly by mammalian predators, most likely Iberian lynx (Lynx pardinus) or common genet (Genetta genetta). The main cause of mortality was illegal poisoning, which sharply increased in the late 1990s. Nest-site selection models showed that: (1) black kites selected areas near the seasonal marshes, rich in their main prey species and (2) red kites selected areas rich in marshland and far from the park border, possibly in response to illegal poisoning episodes. Productivity was positively related to the availability of marshland and of open terrestrial habitats for black and red kites, respectively, and negatively related for both species to the density of black kites within 200 m of the nest. Our results showed that: (1) intraguild predation and competition among predators make outcomes of interspecific interactions extremely complex, leading to unpredictable side-effects of priority actions targeting one species at a time and (2) though the park was fenced and patrolled by wardens, the wide ranging behaviour of the two species made park borders permeable to negative human pressures, such as poisoning, leading to extreme edge effects and absence of a true, safe core of the park. Park management needs to be better integrated with its regional socio-economic and cultural context. Education programmes against illegal poisoning should be urgently promoted.     

Dietary switching of collembola in grassland soil food webs

Soil food webs are characterised by complex direct and indirect effects among the organisms. Consumption of microorganisms by soil animals is considered as an important factor that contributes to the stability of communities, though cascading effects within the food web can be difficult to detect. In a greenhouse experiment, an addition of a high number the fungal feeding collembola Folsomia quadrioculata was applied to grassland soil food webs in monocultures of three plant species: Plantago lanceolata (forb), Lotus corniculatus (legume) and Holcus lanatus (grass). The abundance of microorganisms, determined as the abundances of phospholipid fatty acids (PLFAs) and the abundances of resident invertebrates, nematodes and collembolans, did not change due to the addition of F. quadrioculata. Trophic positions of collembolans were determined by analyses of natural abundances of ¹⁵N stable isotopes. The use of food resources by microorganisms and collembolans was determined by ¹³C analysis of microbial PLFAs and solid samples of collembolans. δ¹³C values of the resident collembola Folsomia fimetaria were lower in the presence of F. quadrioculata than in the control food webs indicating a use of more depleted ¹³C food resources by F. fimetaria. The δ¹⁵N values of F. fimetaria did not change at the addition of F. quadrioculata thus no change of trophic levels was detected. The switch of F. fimetaria to a different food resource could be due to indirect interactions in the food web as the two collembolan species were positioned on different trophic positions, according to different δ¹⁵N values.     

Free-ranging domestic dogs (Canis familiaris) as predators and prey in rural Zimbabwe: threats of competition and disease to large wild carnivores

Domestic dogs (Canis familiaris) arrived in Zimbabwe ca. 1000 years ago. Numbers of free-ranging dogs have reached unprecedented levels in communal lands (agro-pastoralist rural areas), and interact with large wild carnivores along boundaries with wildlife reserves as predators and prey. This study examined a population of 236 dogs in a 33-km² section of Gokwe Communal Land (GCL) bordering the Sengwa Wildlife Research Area (SWRA) in north-western Zimbabwe in 1995-1996. Dogs were found up to 6 km within the SWRA, and were the most common carnivore on the GCL-SWRA boundary. Observations of 16 radio-collared dogs showed that they were inefficient predators. Only 20 kills were recorded amongst the remaining dog population, of which three were wild ungulates. Dogs were unsuccessful predators due to their small group size (mean 1.7) and body mass (mean 14.7 kg), and the abundance of alternative food. It is therefore unlikely that they compete with large carnivores for wild prey. However, leopards (Panthera pardus), lions (P. leo) and spotted hyaenas (Crocuta crocuta) preyed on dogs in GCL, removing ?6% of the dog population in 1993. Such predation provides ideal circumstances for disease transmission. Canid disease was prevalent in the study area; including rabies and probably distemper. The risk of infection is greatest during the dry season (May-October), when peaks in rates of disease, carnivore incursions into GCL, and predation on dogs coincided. The role of jackals (Canis adustus and Canis mesomelas) and spotted hyaena predation of dogs is discussed in relation to disease epidemics within wildlife reserves. With a dog population growth rate of 6.5% per annum, and the prevalence of canid diseases, the conservation threat posed by dogs is escalating on communal land-wildlife reserve boundaries in Zimbabwe. Measures to control dog numbers and improve vaccination coverage of dogs are discussed.     

Resolving the urban nest predator paradox: The role of alternative foods for nest predators

Urbanization is a leading cause of species endangerment in the United States; however, certain species thrive in urban habitats. The loss of key predators or the addition of new predators in urban areas could alter the structure of urban communities. A reduction in nest predation is hypothesized to explain the high density of urban birds, yet urban areas typically have increased populations of avian nest predators. The loss of important nest predators in urban habitats, prey switching of urban predators, or successful nest defense against avian nest predators could explain this urban nest predator paradox. To assess these hypotheses I compared nest predation rates of Northern mockingbirds (Mimus polyglottos) in parking lots and residential neighborhoods to populations in pastures and wildlife preserves during 2007–2009 in Florida, USA and placed video cameras on a subset of nests in 2008–2009. Data do not support the hypothesis that urban nest predation rates are consistently lower than non-urban nest predation rates. Of the 56 nest predation events recorded, cats were the dominant urban predator and Cooper’s hawks (Accipiter cooperii) were the dominant non-urban predator. There was no evidence for a loss of important nest predators in urban habitats; however, prey switching by Cooper’s hawks likely occurred. There was also indirect evidence for the importance of nest defense. Furthermore, some of the cats recorded as nest predators in residential neighborhoods were owned cats and all but one cat predation event occurred at night. To reduce nest predation rates, cat owners should keep their cats indoors at night.     

Ecological influence of the entomopathogenic nematode,Steinernema carpocapsae,on pistachio orchard soil arthropods

The entomopathogenic nematode, Steinernema carpocapsae, can reduce pesticide reliance in pistachios by controlling overwintering larvae of the navel orangeworm, Amyelois transitella (Lepidoptera: Pyralidae). But, beyond this, their influence in pistachio soil food webs is unclear. Given soil food webs’ complexity, S. carpocapsae likely interact with more species than just their intended target, infecting alternate hosts or providing food for native predators. This study quantifies the nematodes’ effects on soil arthropod and surface arthropod diversity in two large orchards in Madera County, California. We found significantly more isotomid collembolans, predatory anystid mites and gnaphosid spiders under trees where nematodes were applied indicating either direct predation or indirect trophic effects. Significantly fewer Forficula auricularia (Dermaptera: Forficulidae) and Blapstinus discolor (Coleoptera: Tenebrionidae) were found under treated trees, suggesting a possible non-target infection. Nematode persistence was limited but positively correlated with pitfall catches of the tenebrionid beetles, Nyctoporis cristata and B. discolor.     

Abundance, diversity and connectance of soil food web channels along environmental gradients in an agricultural landscape

Soil food webs respond to anthropogenic and natural environmental variables and gradients. We studied abundance, connectance (a measure of the trophic interactions within each channel), and diversity in three different channels of the soil food web, each comprised of a resource-consumer pair: the microbivore channel (microbes and their nematode grazers), the plant–herbivore channel (plants and plant-feeding nematodes), and the predator–prey channel (predatory nematodes and their nematode prey), and their associations with different gradients in a heterogeneous agricultural landscape that consisted of intensive row crop agriculture and grazed non-irrigated grasslands in central California. Samples were taken at three positions in relation to water channels: water’s edge, bench above waterway, and the adjacent arable or grazed field. Nematode communities, phospholipid fatty acid (PLFA) biomarkers, and soil properties (NH₄⁺-N, NO₃⁻-N, total N, total C, pH, P, bulk density and soil texture) were measured, and riparian health ratings were scored. Environmental variables were obtained from publicly-available data sources (slope, elevation, available water capacity, erodability, hydraulic conductivity, exchangeable cation capacity, organic matter, clay and sand content and pH).The abundance and richness in most food web components were higher in grazed grasslands than in intensive agricultural fields. Consumers contributed less than their resources to the abundance and richness of the community in all channels. The association between richness and abundance for each component was strongest for the lowest trophic links (microbes, as inferred by PLFA) and weakest for the highest (predatory nematodes). The trophic interactions for the predator–prey and plant–herbivore channels were greater in the grassland than in the cropland. Fields for crops or grazing supported more interactions than the water’s edge in the plant–herbivore and microbivore channels. Connectance increased with the total richness of each community. Higher connectance within the microbivore and predator–prey soil food web channels were associated with soil NO₃⁻-N and elevation respectively, which served as surrogate indicators of high and low agricultural intensification.     

The soil food web: structure and perspectives

This review outlines directions for future research in soil food web ecology. Two lines of research are considered to be most important: the adoption of new methodologies to investigate food relationships and the strengthening of experimentation to investigate the interaction strength between food web components. For a better understanding of food relationships molecular methods, particularly fluorescence in situ hybridization, and stable isotope methodology, including the analysis of variations in the abundance of ¹³C and ¹⁵N, are thought to be most promising. Implications of results of the studies which employed these methodologies for the structure and function of soil food webs are highlighted.     

Evidence of indirect impacts of introduced trout on native amphibians via facilitation of a shared predator

Hyperpredation occurs when non-native prey facilitate invasive predators, which then suppress native prey. Direct impacts of introduced fish on amphibians are well studied, but the role of fish in supporting shared predators has not been considered. We present evidence for indirect effects of trout on amphibians through snake predation. Analyses of the diet, distribution and density of the Pacific coast aquatic garter snake (Thamnophis atratus) relative to the sympatric common garter snake (Thamnophis sirtalis) in the Klamath Mountains of California suggest that trout introductions facilitated expansion of T. atratus by providing alternative prey. T. atratus diet included trout and amphibians whereas T. sirtalis preyed solely upon amphibians. The distribution and density of T. atratus matched that of introduced trout instead of native amphibians. Populations of T. atratus could reach high densities in the absence of high densities of amphibians. When the snakes opportunistically prey upon amphibians whose numbers are already directly impacted by trout, they can cause significant additional declines. When T. atratus was present in lake basins, native Cascades frogs (Rana cascadae) were rarer than in basins without T. atratus. This case differs from other hyperpredation studies because the two prey species also interact via intraguild predation. Given the worldwide practice of stocking fish into aquatic habitats, it is important to understand the consequences of the practice on food-web structure and ecosystem functioning. Bottom-up impacts of introduced predators should be considered as well as top-down so that managers can incorporate the range of ecosystem-level effects into conservation goals and decisions.     

Effects of food quality, starvation and life stage on stable isotope fractionation in Collembola

Naturally occurring stable isotopes of carbon and nitrogen are powerful tools to investigate food webs, where the ratio of ¹⁵N/¹⁴N is used to assign trophic levels and of ¹³C/¹²C to determine the food source. A shift in δ¹⁵N value of 3‰ is generally suggested as mean difference between two trophic levels, whereas the carbon isotope composition of a consumer is assumed to reflect the signal of its diet. This study investigates the effects of food quality, starvation and life stage on the stable isotope fractionation in fungal feeding Collembola. The fractionation of nitrogen was strongly affected by food quality, i.e. the C/N ratio of the fungal diet. Collembola showed enrichment in the heavier isotope with increasing N concentration of the food source. Δ¹⁵N varied between 2.4‰, which assigns a shift in one trophic level, and 6.3‰, suggesting a shift in two trophic levels. Starvation up to 4 weeks resulted in an increase in the total δ¹⁵N value from 2.8‰ to 4.0‰. Different life stages significantly affected the isotope discrimination by Collembola with juveniles showing a stronger enrichment (Δ¹⁵N=4.9‰) compared to adults (Δ¹⁵N=3.5‰). Δ¹³C varied between −2.1‰ and −3.3‰ depending on the food quality, mainly due to compensational feeding on low quality diet. During starvation δ¹³C value decreased by 1.1‰, whereas the life stage of Collembola had no significant effect on isotopic ratios. The results indicate that the food resource and the physiological status of the consumer have important impact on stable isotope discrimination. They may cause differences in fractionation rate comparable to trophic level shifts, a fact to consider when analysing food web structure.     

The feeding ecology of elaterid larvae in central European arable land: New perspectives based on naturally occurring stable isotopes

To understand soil food webs, empirically generated data on the trophic connections and the feeding ecology of the major below-ground animal taxa are needed. Here we used stable isotope analysis to assess the trophic ecology of wireworms, the larvae of click beetles, in Central European arable land. Wireworms are amongst the major soil macroinvertebrates and are of practical importance in arable soils. Besides feeding on crops, they are thought to feed on weeds, soil organic matter (SOM), and even animal prey, but their feeding ecology is poorly studied under natural conditions. Elaterid larvae and their putative feeding substrates—plant roots, SOM, and litter—were sampled at 17 locations in Austria, Germany, and Italy and their isotope ratios of carbon (¹²C/¹³C) and nitrogen (¹⁴N/¹⁵N) measured to determine the wireworms’ trophic level, the importance of SOM and weeds within the diet of Agriotes larvae, as well as the individual diet variation in Agriotes obscurus larvae. δ¹⁵N signatures suggested that Agriotes larvae are predominately herbivorous, whereas the other wireworm species primarily fed on animal prey. In contrast to SOM, weeds were readily eaten by Agriotes larvae: their dietary contribution ranged between 28% and 67% in weedy maize fields. Most A. obscurus larvae fed on a mixed diet of weeds and maize, although ～15% of the larvae fed primarily on one of the two food sources only. δ¹⁵N signatures indicated that ～10% of the “herbivorous” A. obscurus larvae fed primarily on animal prey, revealing high intraspecific trophic plasticity in these soil insects. Wireworm feeding behaviour is apparently complex at the individual level: the population consists of types A and B generalists, a phenomenon which needs further assessment.