Predator‐driven elemental cycling: the impact of predation and risk effects on ecosystem stoichiometry

Empirical evidence is beginning to show that predators can be important drivers of elemental cycling within ecosystems by propagating indirect effects that determine the distribution of elements among trophic levels as well as determine the chemical content of organic matter that becomes decomposed by microbes. These indirect effects can be propagated by predator consumptive effects on prey, nonconsumptive (risk) effects, or a combination of both. Currently, there is insufficient theory to predict how such predator effects should propagate throughout ecosystems. We present here a theoretical framework for exploring predator effects on ecosystem elemental cycling to encourage further empirical quantification. We use a classic ecosystem trophic compartment model as a basis for our analyses but infuse principles from ecological stoichiometry into the analyses of elemental cycling. Using a combined analytical‐numerical approach, we compare how predators affect cycling through consumptive effects in which they control the flux of nutrients up trophic chains; through risk effects in which they change the homeostatic elemental balance of herbivore prey which accordingly changes the element ratio herbivores select from plants; and through a combination of both effects. Our analysis reveals that predators can have quantitatively important effects on elemental cycling, relative to a model formalism that excludes predator effects. Furthermore, the feedbacks due to predator nonconsumptive effects often have the quantitatively strongest impact on whole ecosystem elemental stocks, production and efficiency rates, and recycling fluxes by changing the stoichiometric balance of all trophic levels. Our modeling framework predictably shows how bottom‐up control by microbes and top‐down control by predators on ecosystems become interdependent when top predator effects permeate ecosystems.     

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.     

Top‐down and bottom‐up diversity cascades in detrital vs. living food webs

Apex predators and plant resources are both critical for maintaining diversity in biotic communities, but the indirect (‘cascading’) effects of top‐down and bottom‐up forces on diversity at different trophic levels are not well resolved in terrestrial systems. Manipulations of predators or resources can cause direct changes of diversity at one trophic level, which in turn can affect diversity at other trophic levels. The indirect diversity effects of resource and consumer variation should be strongest in aquatic systems, moderate in terrestrial systems, and weakest in decomposer food webs. We measured effects of top predators and plant resources on the diversity of endophytic animals in an understorey shrub Piper cenocladum (Piperaceae). Predators and resource availability had significant direct and indirect effects on the diversity of the endophytic animal community, but the effects were not interactive, nor were they consistent between living vs. detrital food webs. The addition of fourth trophic level beetle predators increased diversity of consumers supported by living plant tissue, whereas balanced plant resources (light and nutrients) increased the diversity of primary through tertiary consumers in the detrital resources food web. These results support the hypotheses that top‐down and bottom‐up diversity cascades occur in terrestrial systems, and that diversity is affected by different factors in living vs. detrital food webs.     

Bottom–up and top–down forces structuring consumer communities in an experimental grassland

Synthesis The interplay between bottom‐up and top‐down effects is certainly a general manifestation of any changes in both species abundances and diversity. Summary variables, such as species numbers, diversity indices or lumped species abundances provide too limited information about highly complex ecosystems. In contrast, species by species analyses of ecological communities comprising hundreds of species are inevitably only snapshot‐like and lack generality in explaining processes within communities. Our synthesis, based on species matrices of functional groups of all trophic levels, simplifies community complexity to a manageable degree while retaining full species‐specific information. Taking into account plant species richness, plant biomass, soil properties and relevant spatial scales, we decompose variance of abundance in consumer functional groups to determine the direction and the magnitude of community controlling processes. After decades of intensive research, the relative importance of top–down and bottom–up control for structuring ecological communities is still a particularly disputed issue among ecologists. In our study, we determine the relative role of bottom–up and top–down forces in structuring the composition of 13 arthropod functional groups (FG) comprising different trophic consumer levels. Based on species‐specific plant biomass and arthropod abundance data from 50 plots of a grassland biodiversity experiment, we quantified the proportions of bottom–up and top–down forces on consumer FG composition while taking into account direct and indirect effects of plant diversity, functional diversity, community biomass, soil properties and spatial arrangement of these plots. Variance partitioning using partial redundancy analysis explained 21–44% of total variation in arthropod functional group composition. Plant‐mediated bottom–up forces accounted for the major part of the explainable variation within the composition of all FGs. Predator‐mediated top–down forces, however, were much weaker, yet influenced the majority of consumer FGs. Plant functional group composition, notably legume composition, had the most important impact on virtually all consumer FGs. Compared to plant species richness and plant functional group richness, plant community biomass explained a much higher proportion of variation in consumer community composition.     

Spatial variation in vegetation damage relative to primary productivity,small rodent abundance and predation

The relative importance of top‐down and bottom‐up mechanisms in shaping community structure is still a highly controversial topic in ecology. Predatory top‐down control of herbivores is thought to relax herbivore impact on the vegetation through trophic cascades. However, trophic cascades may be weak in terrestrial systems as the complexity of food webs makes responses harder to predict. Alternatively, top‐down control prevails, but the top‐level (predator or herbivore) changes according to productivity levels. Here we show how spatial variation in the occurrence of herbivores (lemmings and voles) and their predators (mustelids and foxes) relates with grazing damage in landscapes with different net primary productivity, generating two and three trophic level communities, during the 2007 rodent peak in northern Norway. Lemmings were most abundant on the unproductive high‐altitude tundra, where few predators were present and the impact of herbivores on vegetation was strong. Voles were most common on a productive, south facing slope, where numerous predators were present, and the impacts of herbivores on vegetation were weak. The impact of herbivores on the vegetation was strong only when predators were not present, and this cannot be explained by between‐habitat differences in the abundance of plant functional groups. We thus conclude that predators influence the plant community via a trophic cascade in a spatial pattern that support the exploitation ecosystems hypothesis. The responses to grazing also differed between plant functional groups, with implications for short and long‐term consequences for plant communities.     

Top‐down,bottom‐up,or side to side? Within‐trophic‐level interactions modify trophic dynamics of a salt marsh herbivore

Many factors can influence the top‐down and bottom‐up dynamics of phytophagous insects. Although interactions between herbivore species have been frequently shown to be ecologically important, the effects of such horizontal trophic interactions on the relative roles of top‐down and bottom‐up forces have gone largely unstudied. In this paper we report on the results of a factorial field experiment in which we examined the effects of within‐trophic‐level interactions on the top‐down and bottom‐up dynamics of a salt marsh planthopper.
We manipulated the bottom‐up effects of plant quality by increasing soil salinity, and manipulated top‐down effects by decreasing the intensity of parasitoid attack with yellow sticky traps that removed hymenopteran parasitoids. We applied these treatments to plots in two patches of the host plant, one with low densities of lepidopteran stem borer larvae, and one with high densities of stem borers. We maintained the treatments and monitored planthopper density for ten months, from March through December 1999. Increased salinity significantly increased planthopper density within one month of the first application of salt. The rapid response of the planthopper to salt treatments suggested a chemical mechanism, perhaps mobilization of bound nitrogen. Yellow sticky traps, although significantly reducing parasitism of planthopper eggs, had little impact on hopper density. The density of lepidopteran stem borers, however, had an even greater impact on planthopper density than did salt treatments, with high stem borer plots supporting much lower densities of hoppers. Stem borer density also reduced the response of the planthopper to other treatments, especially salt supplementation. The results of this study show that the impact of within‐trophic‐level interactions can significantly change herbivore trophic dynamics and can be even more important than either top‐down or bottom‐up effects in determining herbivore density.     

Noncrop flowering plants restore top‐down herbivore control in agricultural fields

Herbivore populations are regulated by bottom‐up control through food availability and quality and by top‐down control through natural enemies. Intensive agricultural monocultures provide abundant food to specialized herbivores and at the same time negatively impact natural enemies because monocultures are depauperate in carbohydrate food sources required by many natural enemies. As a consequence, herbivores are released from both types of control. Diversifying intensive cropping systems with flowering plants that provide nutritional resources to natural enemies may enhance top‐down control and contribute to natural herbivore regulation. We analyzed how noncrop flowering plants planted as “companion plants” inside cabbage (Brassica oleracea) fields and as margins along the fields affect the plant–herbivore–parasitoid–predator food web. We combined molecular analyses quantifying parasitism of herbivore eggs and larvae with molecular predator gut content analysis and a comprehensive predator community assessment. Planting cornflowers (Centaurea cynanus), which have been shown to attract and selectively benefit Microplitis mediator, a larval parasitoid of the cabbage moth Mamestra brassicae, between the cabbage heads shifted the balance between trophic levels. Companion plants significantly increased parasitism of herbivores by larval parasitoids and predation on herbivore eggs. They furthermore significantly affected predator species richness. These effects were present despite the different treatments being close relative to the parasitoids’ mobility. These findings demonstrate that habitat manipulation can restore top‐down herbivore control in intensive crops if the right resources are added. This is important because increased natural control reduces the need for pesticide input in intensive agricultural settings, with cascading positive effects on general biodiversity and the environment. Companion plants thus increase biodiversity both directly, by introducing new habitats and resources for other species, and indirectly by reducing mortality of nontarget species due to pesticides.     

Bottom‐up vs. top‐down effects on terrestrial insect herbivores: a meta‐analysis

Primary consumers are under strong selection from resource (‘bottom‐up’) and consumer (‘top‐down’) controls, but the relative importance of these selective forces is unknown. We performed a meta‐analysis to compare the strength of top‐down and bottom‐up forces on consumer fitness, considering multiple predictors that can modulate these effects: diet breadth, feeding guild, habitat/environment, type of bottom‐up effects, type of top‐down effects and how consumer fitness effects are measured. We focused our analyses on the most diverse group of primary consumers, herbivorous insects, and found that in general top‐down forces were stronger than bottom‐up forces. Notably, chewing, sucking and gall‐making herbivores were more affected by top‐down than bottom‐up forces, top‐down forces were stronger than bottom‐up in both natural and controlled (cultivated) environments, and parasitoids and predators had equally strong top‐down effects on insect herbivores. Future studies should broaden the scope of focal consumers, particularly in understudied terrestrial systems, guilds, taxonomic groups and top‐down controls (e.g. pathogens), and test for more complex indirect community interactions. Our results demonstrate the surprising strength of forces exerted by natural enemies on herbivorous insects, and thus the necessity of using a tri‐trophic approach when studying insect‐plant interactions.     

Top–down limits on prey populations may be more severe in larger prey species,despite having fewer predators

Variation in the vulnerability of herbivore prey to predation is linked to body size, yet whether this relationship is size‐nested or size‐partitioned remains debated. If size‐partitioned, predators would be focused on prey within their preferred prey size range. If size‐nested, smaller prey species should become increasingly more vulnerable because increasingly more predators are capable of catching them. Yet, whether either of these strategies manifests in top–down prey population limitation would depend both on the number of potential predator species as well as the total mortality imposed. Here we use a rare ecosystem scale ‘natural experiment’ comparing prey population dynamics between a period of intense predator persecution and hence low predator densities and a period of active predator protection and population recovery. We use three decades of data on herbivore abundance and distribution to test the role of predation as a mechanism of population limitation among prey species that vary widely in body size. Notably, we test this within one of the few remaining systems where a near‐full suite of megaherbivores occur in high density and are thus able to include a thirtyfold range in herbivore body size gradient. We test whether top–down limitation on prey species of particular body size leads to compositional shifts in the mammalian herbivore community. Our results support both size‐nested and size‐partitioning predation but suggest that the relative top–down limiting impact on prey populations may be more severe for intermediate sized species, despite having fewer predators than small species. In addition we show that the gradual recovery of predator populations shifted the herbivore community assemblage towards large‐bodied species and has led to a community that is strongly dominated by large herbivore biomass.     

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.     

Community‐wide effects of below‐ground rhizobia on above‐ground arthropods

1. Plants take nutrients for their growth and reproduction from not only soil but also symbiotic microbes in the rhizosphere, and therefore below‐ground microbes may indirectly influence the above‐ground arthropod community through changes in the quality and quantity of plants. 2. Rhizobia are root‐nodulating bacteria that provide NH₄⁺ to legume plants. We examined bottom‐up effects of rhizobia on the community properties of the arthropods on host plants, using a root‐nodulating soybean strain (R+) and a non‐nodulating strain (R?) in a common garden. 3. R+ plants grew larger and produced a greater number of leaves than R? plants. We observed 28 species of herbivores and three taxonomic groups of predators on R+ and R? plants. The herbivorous species were classified into sap feeders (12 species) and chewers (16 species). 4. The species richness of overall herbivores, sap feeders, and chewers on R+ plants was greater than that on R? plants. Rhizobia positively affected the abundance of chewers. 5. The community composition of herbivores was significantly different between R? and R+ plants, although species diversity and evenness did not differ. 6. Rhizobia‐induced bottom‐up effects were transmitted to the third trophic level. The abundance, taxonomic richness, and diversity of the predators on R+ plants were greater but evenness was lower than those on R? plants. The community composition of predators was not affected by rhizobia. 7. These results indicate that the below‐ground microbes initiated bottom‐up effects on above‐ground herbivores and predators through trophic levels.     

Independent and combined effects of multiple predators across ontogeny of a dominant grazer

Ecosystems host multiple coexisting predator species whose interactions may strengthen or weaken top–down control of grazers. Grazer populations often exhibit size‐structure, but the nature of multiple predator effects on suppression of size‐structured prey has seldom been explicitly considered. In a southeastern US salt‐marsh, we used both field (additive design) and mesocosm (additive‐substitutive design) experiments to test the independent and combined effects of two species of predatory crab on the survival and predator‐avoidance behavior (i.e. a non‐consumptive effect) of both juveniles and adults of a dominant grazing snail. Results showed: 1) juvenile snails were more vulnerable to predation; 2) consumptive impacts of predators were hierarchically nested, i.e. the larger predator consumed both juvenile and adult snails, while the smaller‐bodied predator consumed only juvenile snails; 3) there were no emergent multiple predator effects on snail consumption; and 4) non‐consumptive effects differed from consumptive effects, with only the large predator inducing predator‐avoidance behavior of individuals within either snail ontogenetic class. The smaller predator therefore played a functionally redundant trophic role across the prey classes considered, augmenting and potentially stabilizing trophic regulation of juvenile snails. Meanwhile, the larger predator played a complementary and functionally unique role by both expanding the size‐spectrum of prey trophic regulation and non‐consumptively altering prey behavior. While our study suggests that nestedness of consumptive interactions determined by predator and prey body sizes may allow prediction of the functional redundancy of particular predator species, it also shows that traits beyond predator body size (e.g. habitat domain) may be required to predict potentially cascading non‐consumptive effects. Future studies of multiple predators (and predator biodiversity) should continue to strive towards greater realism by incorporating not only size‐structured prey, but also other aspects of resource and environmental heterogeneity typical of natural ecosystems.     

Mechanisms of coexistence in diverse herbivore–carnivore assemblages: demographic,temporal and spatial heterogeneities affecting prey vulnerability

Simple models coupling the dynamics of single predators to single prey populations tend to generate oscillatory dynamics of both predator and prey, or extirpation of the prey followed by that of the predator. In reality, such oscillatory dynamics may be counteracted by prey refugia or by opportunities for prey switching by the predator in multi‐prey assemblages. How these mechanisms operate depends on relative prey vulnerability, a factor ignored in simple interactive models. I outline how compositional, temporal, demographic and spatial heterogeneities help explain the contrasting effects of top predators on large herbivore abundance and population dynamics in species‐rich African savanna ecosystems compared with less species‐diverse northern temperate or subarctic ecosystems. Demographically, mortality inflicted by predation depends on the relative size and life history stage of the prey. Because all animals eventually die and are consumed by various carnivores, the additive component of the mortality inflicted is somewhat less than the predation rate. Prey vulnerability varies annually and seasonally, and between day and night. Spatial variation in the risk of predation depends on vegetation cover as well as on the availability of food resources. During times of food shortage, herbivores become prompted to occupy more risky habitats retaining more food. Predator concentrations dependent on the abundance of primary prey species may restrict the occurrence of other potential prey species less resistant to predation. The presence of multiple herbivore species of similar size in African savannas allows the top predator, the lion, to shift its prey selection flexibly dependent on changing prey vulnerability. Hence top–down and bottom–up influences on herbivore populations are intrinsically entangled. Models coupling the population dynamics of predators and prey need to accommodate the changing influences of prey demography, temporal variation in environmental conditions, and spatial variation in the relative vulnerability of alternative prey species to predation. Synthesis While re‐established predators have had major impacts on prey populations in northern temperate regions, multiple large herbivore species typically coexist along with diverse carnivores in African savanna ecosystems. In order to explain these contrasting outcomes, certain functional heterogeneities must be recognised, including relative vulnerability of alternative prey, temporal variation in the risk of predation, demographic differences in susceptibility to predation, and spatial contrasts in exposure to predation. Food shortfalls prompt herbivores to exploit more risky habitats, meaning that top–down and bottom–up influences on prey populations are intrinsically entangled. Models coupling the interactive dynamics of predator and prey populations need to incorporate these varying influences on relative prey vulnerability.     

Latitude and protection affect decadal trends in reef trophic structure over a continental scale

The relative roles of top‐down (consumer‐driven) and bottom‐up (resource‐driven) forcing in exploited marine ecosystems have been much debated. Examples from a variety of marine systems of exploitation‐induced, top‐down trophic forcing have led to a general view that human‐induced predator perturbations can disrupt entire marine food webs, yet other studies that have found no such evidence provide a counterpoint. Though evidence continues to emerge, an unresolved debate exists regarding both the relative roles of top‐down versus bottom‐up forcing and the capacity of human exploitation to instigate top‐down, community‐level effects. Using time‐series data for 104 reef communities spanning tropical to temperate Australia from 1992 to 2013, we aimed to quantify relationships among long‐term trophic group population density trends, latitude, and exploitation status over a continental‐scale biogeographic range. Specifically, we amalgamated two long‐term monitoring databases of marine community dynamics to test for significant positive or negative trends in density of each of three key trophic levels (predators, herbivores, and algae) across the entire time series at each of the 104 locations. We found that trophic control tended toward bottom‐up driven in tropical systems and top‐down driven in temperate systems. Further, alternating long‐term population trends across multiple trophic levels (a method of identifying trophic cascades), presumably due to top‐down trophic forcing, occurred in roughly fifteen percent of locations where the prerequisite significant predator trends occurred. Such alternating trophic trends were significantly more likely to occur at locations with increasing predator densities over time. Within these locations, we found a marked latitudinal gradient in the prevalence of long‐term, alternating trophic group trends, from rare in the tropics (<5% of cases) to relatively common in temperate areas (~45%). Lastly, the strongest trends in predator and algal density occurred in older no‐take marine reserves; however, exploitation status did not affect the likelihood of alternating long‐term trophic group trends occurring. Our data suggest that the type and degree of trophic forcing in this system are likely related to one or more covariates of latitude, and that ecosystem resiliency to top‐down control does not universally vary in this system based on exploitation level.     

The effect of predation pressure and predator adaptive foraging on the relative importance of consumptive and non‐consumptive predator net effects in a freshwater model system

An important challenge in community ecology is identifying the functional characteristics capable of predicting the nature and strength of predator effects on food webs. We developed an individual‐based model, based on a shallow lake model system, to evaluate the total, consumptive, and non‐consumptive indirect effect that predators have on basal resources when the predators differ in their foraging types (active adaptive foraging or sedentary foraging). Overall, both predator types caused similar total indirect effects on lower trophic levels. However, the nature net effects of predators diverged between predator foraging types. Active predators caused larger non‐consumptive effects, relative to the total indirect effect, irrespective of predation pressure levels. On the other hand, sedentary predators caused larger non‐consumptive effects for lower predation pressure levels, but consumptive effects became more important as predation pressure increased. Our simulations showed that the reliance on a particular mechanism driving consumer–resource interactions is altered by predator foraging behavior and highlight the importance of both prey and predator foraging behaviors to predict the causes and consequences of cascading effects observed in food webs.     

Overfishing of top predators eroded the resilience of the Black Sea system regardless of the climate and anthropogenic conditions

It is well known that human activities, such as harvesting, have had major direct effects on marine ecosystems. However, it is far less acknowledged that human activities in the surroundings might have important effects on marine systems. There is growing evidence suggesting that major reorganization (i.e., a regime shift) is a common feature in the temporal evolution of a marine system. Here we show, and quantify, the interaction of human activities (nutrient upload) with a favourable climate (run‐off) and its contribution to the eutrophication of the Black Sea in the 1980s. Based on virtual analysis of the bottom‐up (eutrophication) vs. top‐down (trophic cascades) effects, we found that an earlier onset of eutrophication could have counteracted the restructuring of the trophic regulation at the base of the food web that resulted from the depletion of top predators in the 1970s. These enhanced bottom‐up effects would, however, not propagate upwards in the food web beyond the zooplankton level. Our simulations identified the removal of apex predators as a key element in terms of loss of resilience that inevitably leads to a reorganization. Once the food web has been truncated, the type and magnitude of interventions on the group replacing the apex predator as the new upper trophic level have no effect in preventing the trophic cascade. By characterizing the tipping point at which increased bottom‐up forcing exactly counteracts the top‐down cascading effects, our results emphasize the importance of a comprehensive analysis that take into account all structuring forces at play (including those beyond the marine system) at a given time.     

Long‐term nutrient addition alters arthropod community composition but does not increase total biomass or abundance

A simple bottom–up hypothesis predicts that plant responses to nutrient addition should determine the response of consumers: more productive and less diverse plant communities, the usual result of long‐term nutrient addition, should support greater consumer abundances and biomass and less consumer diversity. We tested this hypothesis for the response of an aboveground arthropod community to an uncommonly long‐term (24‐year) nutrient addition experiment in moist acidic tundra in arctic Alaska. This experiment altered plant community composition, decreased plant diversity and increased plant production and biomass as a deciduous shrub, Betula nana, became dominant. Consistent with strong effects on the plant community, nutrient addition altered arthropod community composition, primarily through changes to herbivore taxa in the canopy‐dwelling arthropod assemblage and detritivore taxa in the ground assemblage. Surprisingly, however, the loss of more than half of plant species was accompanied by negligible changes to diversity (rarefied richness) of arthropod taxa (which were primarily identified to family). Similarly, although long‐term nutrient addition in this system roughly doubles plant production and biomass, arthropod abundance was either unchanged or decreased by nutrient addition, and total arthropod biomass was unaffected. Our findings differ markedly from the handful of terrestrial studies that have found bottom‐up diversity cascades and productivity responses by consumers to nutrient addition. This is probably because unlike grasslands and salt marshes (where such studies have historically been conducted), this arctic tundra community becomes less palatable, rather than more so, after many years of nutrient addition due to increased dominance of B. nana. Additionally, by displacing insulating mosses and increasing the cover of shrubs that cool and shade the canopy microenvironment, fertilization may displace arthropods keenly attuned to microclimate. These results indicate that terrestrial arthropod assemblages may be more constrained by producer traits (i.e. palatability, structure) than they are by total primary production or producer diversity.     

Environmental and plant genetic effects on tri‐trophic interactions

The effects of plant genotype and environmental factors on tri‐trophic interactions have usually been investigated separately, limiting our ability to compare the relative strength of these effects as well as their potential to interactively shape arthropod communities. We studied the interactions among the herb Ruellia nudiflora, a seed predator, and its parasitoids using 14 maternal plant families grown in a common garden. By fertilizing half of the plants of each family and subsequently recording fruit number, seed predator number, and parasitoid number per plant, we sought to compare the strength of plant genetic effects with those of soil fertility, and determine if these factors interactively shape tri‐trophic interactions. Furthermore, we evaluated if these bottom–up factors influenced higher trophic levels through changes in abundance across trophic levels (density‐mediated) or changes in the function of species interactions (trait‐mediated). Plant genetic effects on seed predators and parasitoids were stronger than fertilization effects. Moreover, we did not find plant genetic variation for fertilization effects on fruit, seed predator, or parasitoid abundance, showing that each factor acted independently on plant resources and higher trophic levels. Both bottom–up forces were transmitted via density‐mediated effects where increased fruit number from fertilization and plant genetic effects increased seed predator and parasitoid abundance; however, seed predator attack was density‐dependent, while parasitoid attack was density‐independent. Importantly, there was evidence (marginally significant in one case) that fertilization modified the function of plant‐seed predator and seed predator–parasitoid interactions by increasing the number of seed predators per fruit and decreasing the number of parasitoids per seed predator, respectively. These findings show that plant genetic and soil fertility effects cascaded up this simple food chain, that plant genetic effects were stronger across all trophic levels, and that these effects were transmitted independently and through contrasting mechanisms.     

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.     

Effects of environmental warming and drought on size‐structured soil food webs

Soil systems maintain important ecosystem processes crucial for plant life and food production. Especially agricultural systems are strongly affected by climate change due to low vegetation cover associated with high temperatures and drought. Nevertheless, the response of soil systems to climate change is little explored. We used microcosms with a simplified soil community to address effects of climate change using independent temperature and dryness gradients and addressed their effects on top–down control and litter decomposition. The community consisted of maize litter as a basal resource, fungi, springtails and as top predators mites and centipedes. As the body‐size structure is of high importance for communities, we included differently‐sized springtails and predator species. After seven weeks, the experiment was terminated, and the impact of climate change on direct feeding interactions and indirect effects across trophic levels was analysed. With increasing temperature and dryness, consumption rates increased, thereby amplifying the negative influence of consumer populations on their resources. Hence, these climate‐change variables increased the top–down control of 1) predators (mainly mites) on springtails and 2) fungi on litter decomposition. In addition, we found that the climate‐change variables strengthened trophic cascades from predators on fungi whose density was thus increasingly decoupled from top–down control by their springtail consumers. Their increased decomposition rates are of high importance for carbon cycling and may result in accelerated nutrient turnover. In conclusion, our results suggest that climate change may strongly influence the structure and functioning of soil systems by strengthening consumption rates and trophic cascades, which will have far reaching consequences for the nutrient turnover and productivity of agricultural ecosystems.