Intraguild interactions between spiders and ants and top-down control in a grassland food web

In most terrestrial ecosystems ants (Formicidae) as eusocial insects and spiders (Araneida) as solitary trappers and hunters are key predators. To study the role of predation by these generalist predators in a dry grassland, we manipulated densities of ants and spiders (natural and low density) in a two-factorial field experiment using fenced plots. The experiment revealed strong intraguild interactions between ants and spiders. Higher densities of ants negatively affected the abundance and biomass of web-building spiders. The density of Linyphiidae was threefold higher in plots without ant colonies. The abundance of Formica cunicularia workers was significantly higher in spider-removal plots. Also, population size of springtails (Collembola) was negatively affected by the presence of wandering spiders. Ants reduced the density of Lepidoptera larvae. In contrast, the abundance of coccids (Ortheziidae) was positively correlated with densities of ants. To gain a better understanding of the position of spiders, ants and other dominant invertebrate groups in the studied food web and important trophic links, we used a stable isotope analysis (¹⁵N and ¹³C). Adult wandering spiders were more enriched in ¹⁵N relative to ¹⁴N than juveniles, indicating a shift to predatory prey groups. Juvenile wandering and web-building spiders showed δ¹⁵N ratios just one trophic level above those of Collembola, and they had similar δ¹³C values, indicating that Collembola are an important prey group for ground living spiders. The effects of spiders demonstrated in the field experiment support this result. We conclude that the food resource of spiders in our study system is largely based on the detrital food web and that their effects on herbivores are weak. The effects of ants are not clear-cut and include predation as well as mutualism with herbivores. Within this diverse predator guild, intraguild interactions are important structuring forces.     

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.     

Ecosystem linkages revealed by experimental lake-derived isotope signal in heathland food webs

Cross-ecosystem movement of nutrients and biomass can have important effects on recipient systems. Emerging aquatic insects are subsidies to terrestrial ecosystems and can influence foodweb interactions in riparian systems. In a 2-year field experiment, we simulated aquatic insect deposition by adding adult midge carcasses (150?g dry mass m^?2 year^?1) to 1-m² heathland plots at a site with low natural midge deposition. We established four levels of midge-addition treatments and measured stable isotopes (??¹³C and ??¹⁵N) in plants and arthropods within each treatment. We used a multiple-source isotope Bayesian mixing model to estimate the terrestrial versus aquatic contribution to the diets of arthropods. Aquatic resources were incorporated into plant, detritivore, and predator biomass. Detritivorous Collembola showed the greatest difference in isotope values (+3??? ??¹⁵N and +4??? ??¹³C) between midge-addition and reference treatments. Isotope values of small spiders followed the same trend of enrichment as Collembola while other arthropods (mites and large spiders) were only enriched after 2?years of midge addition. Although predator diets did not change, they became isotopically enriched via their likely prey (Collembola). Plants also had elevated ??¹⁵N (+1???) in midge-addition treatments. The time required and amount of midge-derived C and N detected varied and depended on trophic position. Midge-derived nutrients were no longer present in arthropod biomass in the year following midge addition. Aquatic insect carcasses can be rapidly incorporated into terrestrial food webs in nearshore habitats, and repeated inputs can be detected at multiple trophic levels, thus highlighting the importance of the detrital pathway for aquatic to terrestrial cross-ecosystem subsidies.     

Effect of simulated environmental change on alpine soil arthropods

The effects of environmental change on soil animal communities are poorly known. Norwegian mountains are subject to both atmospheric nitrogen deposition and increased temperature. In a nutrient poor alpine Dryas heath in south Norway, soil arthropods were studied after 4 years of simulated environmental change by warming and/or nutrient addition. Warming alone only affected three low‐density Collembola species, while nutrient addition, with or without warming, greatly changed the dominance hierarchy of the microarthropod community. Certain Collembola species with a short (1 year) life cycle and predatory Gamasina mites increased markedly in density. These groups may have been favored by increased litter production, as plant biomass and litter producing graminoids and forbs increased significantly in plots with nutrient addition and nutrient addition combined with warming. Microarthropods with a longer life cycle, such as Oribatida and certain Collembola, were generally unaffected by nutrient addition and probably need more time to respond. The number of Oribatida taxa was, however, reduced in plots with nutrient addition, both with and without warming. A ground‐living species of Coccoidea (Homoptera) declined in plots with nutrient addition and warming compared with only warming, probably due to reduced cover of its host plant Dryas. The density of Diptera larvae (Sciaridae and Chironomidae) was unaffected by the treatments. Our results show that increased nutrient availability in nutrient poor alpine soils may have large but different effects on different taxa of soil animals. Species with short life cycles reacted first. Nutrient addition and nutrient addition combined with warming resulted in several effects below ground on microarthropods as previously shown above ground on plants: Increased biomass, high dominance of a few rapid‐growing species, contrasting responses of closely related species, and a reduction in species numbers. These short‐term responses may have profound long‐term effects in this alpine ecosystem.     

Allochthonous aquatic insects increase predation and decrease herbivory in river shore food webs

Rivers produce an abundance of aquatic insects that traverse land, where they can have bottom-up effects on predators, who, in turn, can have top-down effects on terrestrial herbivores. This effect can cascade down to plants. These trophic relationships were demonstrated in a field of stinging nettles, Urtica dioica , along a river in Germany. At the shore compared to similar microhabitats 30–60 m away the abundance and biomass of: midges were highest, spiders were also highest, while herbivorous leafhoppers were lowest. At the shore, nettle plants were less damaged by herbivores and thus had less regrowth. Spiders regularly captured both aquatic midges as well as terrestrial leafhoppers and they captured more individuals of both groups at the shore than further away. Midges supported high densities of shore spiders. This was inferred from correlation of distribution and diet in the absence of other environmental gradients. Removal of spiders from experimental plots caused leafhoppers to increase at the shore, causing more plant damage. These effects were not evident at spider-removal sites away from the shore. This demonstrated that spiders depressed leafhoppers and decreased herbivory on plants only at the shore. It is concluded that aquatic insects had a bottom-up effect on spiders and that this subsidy facilitated a top-down effect that cascaded from spiders to leafhoppers to plants. Similar effects would explain the distribution of arthropods along many rivers. Allochthony connects river food webs with shore food webs, making both components essential for each other.     

Trickle-down effects of aboveground trophic cascades on the soil food web

Trophic cascades are increasingly being regarded as important features of aboveground and belowground food webs, but the effects of aboveground cascades on soil food webs, and vice versa, remains essentially unexplored. We conducted an experiment consisting of model synthesised communities containing grassland plant and invertebrate species, in which treatments included soil only, soil+plants, soil+plants+aphids, and soil+plants+aphids+predators; predator treatments consisted of the lacewing Micromus tasmaniae and ladybird beetle Coccinella undecimpunctata added either singly or in combination. Addition of Micromus largely reversed the negative effects of aphids on plant biomass, while both of the predator species caused large changes in the relative abundances of dominant plant species. Predators of aphids also affected several components of the belowground subsystem. Micromus had positive indirect effects on the primary consumer of the soil decomposer food web (microflora), probably through promoting greater input of basal resources to the decomposer subsystem. Predator treatments also influenced densities of the tertiary consumers of the soil food web (top predatory nematodes), most likely through inducing changes in plant community composition and therefore the quality of resource input to the soil. The secondary consumers of the soil food web (microbe‐feeding nematodes) were, however, unresponsive. The fact that some trophic levels of the soil food web but not others responded to aboveground manipulations is explicable in terms of top‐down and bottom‐up forces differentially regulating different belowground trophic levels. Addition of aphids also influenced microbial community structure, promoted soil bacteria at the expense of fungi, and enhanced the diversity of herbivorous nematodes; in all cases these effects were at least partially reversed by addition of Micromus. These results in tandem point to upper level consumers in aboveground food webs as a potential driver of the belowground subsystem, and provide evidence that predator‐induced trophic cascades aboveground can have effects that trickle through soil food webs.     

Cascading effects of belowground predators on plant communities are density‐dependent

Soil food webs comprise a multitude of trophic interactions that can affect the composition and productivity of plant communities. Belowground predators feeding on microbial grazers like Collembola could decelerate nutrient mineralization by reducing microbial turnover in the soil, which in turn could negatively influence plant growth. However, empirical evidences for the ecological significance of belowground predators on nutrient cycling and plant communities are scarce. Here, we manipulated predator density (Hypoaspis aculeifer: predatory mite) with equal densities of three Collembola species as a prey in four functionally dissimilar plant communities in experimental microcosms: grass monoculture (Poa pratensis), herb monoculture (Rumex acetosa), legume monoculture (Trifolium pratense), and all three species as a mixed plant community. Density manipulation of predators allowed us to test for density‐mediated effects of belowground predators on Collembola and lower trophic groups. We hypothesized that predator density will reduce Collembola population causing a decrease in nutrient mineralization and hence detrimentally affect plant growth. First, we found a density‐dependent population change in predators, that is, an increase in low‐density treatments, but a decrease in high‐density treatments. Second, prey suppression was lower at high predator density, which caused a shift in the soil microbial community by increasing the fungal: bacterial biomass ratio, and an increase of nitrification rates, particularly in legume monocultures. Despite the increase in nutrient mineralization, legume monocultures performed worse at high predator density. Further, individual grass shoot biomass decreased in monocultures, while it increased in mixed plant communities with increasing predator density, which coincided with elevated soil N uptake by grasses. As a consequence, high predator density significantly increased plant complementarity effects indicating a decrease in interspecific plant competition. These results highlight that belowground predators can relax interspecific plant competition by increasing nutrient mineralization through their density‐dependent cascading effects on detritivore and soil microbial communities.     

Diel and seasonal patterns of prey available to epigeal predators: Evidence for food limitation in a linyphiid spider community

Linyphiid spiders, as part of the community of natural enemies that frequent agroecosystems, can exert significant pressure on prey populations. Many aspects of linyphiid feeding ecology remain understudied, including temporal and seasonal patterns of prey utilization. To quantify the diversity, quantity, and spatial pattern of availability over diel and seasonal gradients, we monitored prey in an alfalfa crop, using web-site specific sticky traps. Although there were no differences in total number of prey between the day and night, significantly more Collembola were intercepted nocturnally rather than diurnally. Conversely, Diptera and Aphididae were significantly more abundant diurnally. Overall there were more prey at web-sites than in the general crop environment, indicating that spiders were selectively constructing webs where prey was relatively more abundant. However, spiders still may have experienced a high degree of prey limitation because the abundance of Collembola, an important detrital prey, was low over much of the season and below levels calculated as being required to sustain growth and reproductive output in linyphiid spiders. In conclusion, our data suggests that linyphiid spiders, in order to avoid food limitation, may benefit by foraging throughout the diel cycle, consuming herbivorous prey in addition to their core diet of detrital prey. This link between above and below ground food webs indicates the importance of understanding the trophic interactions between epigean spiders and their prey if they are to be integrated into future biological control programs.     

Importance of habitat structure to the arthropod food-web in Douglas-fir canopies

This study tested the hypothesis that habitat structure dictates the distribution and community composition of arboreal arthropods. A diverse arthropod assemblage of Douglas-fir canopies, which included Araneae, Psocoptera, Collembola and Homoptera, was chosen as a model system. Habitat structural diversity, defined as needle density and branching complexity of Douglas-fir branches, was manipulated in a four-month experiment by needle removal, thinning and tying of branches. Abundance of canopy spiders declined significantly following needle density reduction and branch thinning, branch tying significantly increased spider abundance. Distinct habitat utilization patterns were found among individual spider guilds. Orb weavers (Araneidae) dominated spider assemblages in structurally simple habitats, whereas tied branches were colonized primarily by sheet-web weavers (Linyphiidae) and nocturnal hunting spiders (Anyphaenidae and Clubionidae). Spider species richness and average body size of several spider species increased in structurally more complex habitats. Arboreal spiders appeared to be limited by strong bottom-up effects in the form of habitat quality and, to a lesser degree, prey availability. Habitat manipulations did not affect densities or biomass of flying arthropod colonists in the branch vicinity. Needle removal and branch thinning led to a significant decline in the abundance of Psocoptera and Collembola. Tying of branches resulted in an eight-fold increase in Collembola numbers, organisms most sensitive to habitat alterations. Canopy habitat structure modified vertical dispersal of Collembola from forest litter, which may have significant implications for arboreal consumers. Our results lend strong support to the importance of habitat structural diversity in explaining general patterns of arthropod abundance and diversity on plants.     

Landuse Change in Savannas Disproportionately Reduces Functional Diversity of Invertebrate Predators at the Highest Trophic Levels: Spiders as an Example

Predators play a disproportionately positive role in ensuring integrity of food webs, influencing ecological processes and services upon which humans rely. Predators tend to be amongst the first species to be affected by anthropogenic disturbance, however. Spiders impact invertebrate population dynamics and stabilise food webs in natural and agricultural systems (potentially mitigating against crop pests and reduced yields). Africa’s savannas are undergoing continent-wide conversion from low-density rangelands to villages and croplands, as human populations burgeon. Despite limited research, and evidence of deleterious impacts to biodiversity, African savannas are earmarked by prominent international organisations for conversion to cropland. Given the key role of spiders in food webs, they can have beneficial impacts in agroecosystems. Furthermore, functional diversity (FD) reflects ecosystem pattern and processes better than species diversity, so we evaluated impacts of large-scale landuse change on both species richness and FD. We surveyed spiders using pitfall traps at 42 sites (14 replicates each in rangeland, cropland, and villages) in South African savannas, investigating effects of landuse, season, and habitat variables on spider species diversity and FD. Species richness was lowest in villages. FD was lowest in cropland, however, with reduced representation of traits associated with hunting of larger invertebrates. Furthermore, there were fewer specialists in croplands. These findings suggest that even when cropland does not impact species diversity, loss of FD can still occur. As savanna systems transform, impacts on invertebrate population dynamics may increase the possibility of a breakdown in pest control in natural and agricultural systems, given changes in FD of invertebrate predators.     

Richness and species composition of arboreal arthropods affected by nutrients and predators: a press experiment

A longstanding goal for ecologists is to understand the processes that maintain biological diversity in communities, yet few studies have investigated the combined effects of predators and resources on biodiversity in natural ecosystems. We fertilized nutrient limited plots and excluded insectivorous birds in a randomized block design, and examined the impacts on arthropods associated with the dominant tree in the Hawaiian Islands, Metrosideros polymorpha (Myrtaceae). After 33 months, the species load (per foliage mass) of herbivores and carnivores increased with fertilization, but rarified richness (standardized to abundance) did not change. Fertilization depressed species richness of arboreal detritivores, and carnivore richness dropped in caged, unfertilized plots, both because of the increased dominance of common, introduced species with treatments. Herbivore species abundance distributions were more equitable than other trophic levels following treatments, and fertilization added specialized native species without changing relativized species richness. Overall, bird removal and nutrient addition treatments on arthropod richness acted largely independently, but with countervailing influences that obscured distinct top-down and bottom-up effects on different trophic levels. This study demonstrates that species composition, biological invasions, and the individuality of species traits may complicate efforts to predict the interactive effects of resources and predation on species diversity in food webs.     

Complex responses to invasive grass litter by ground arthropods in a Mediterranean scrub ecosystem

Plant invasions have tremendous potential to alter food webs by changing basal resources. Recent studies document how plant invasions may contribute to increased arthropod abundances in detritus-based food webs. An obvious mechanism for this phenomenon—a bottom-up effect resulting from elevated levels of detritus from the invasive plant litter—has not been explicitly studied. We examined the effects of an annual grass invasion on ground arthropod assemblages in the coastal sage scrub (CSS) of southern California. Bottom-up food web theory predicts that the addition of detritus would increase generalist-feeding arthropods at all trophic levels; accordingly, we expected increases in fungi, Collembola, and common predators such as mites and spiders. For the common ant taxa, habitat alteration may also be important for predicting responses. Thus we expected that Forelius mccooki and Pheidole vistana, the most common ant species, would decline because of changes in soil temperature (F. mccooki) and habitat structure (P. vistana) associated with litter. We studied trends observationally and conducted a 3-year experiment in which we manipulated litter quantity. In contrast to other published studies, most detritus-based arthropod taxa declined in areas of high grass invasion, and, within trophic levels, responses often varied idiosyncratically. For the two most common taxa, a native ant (F. mccooki), and predatory mites in the Anystidae, we experimentally linked declines in abundance to increased levels of invasive grass litter. Such declines, especially those exhibited by the most common ant taxa, could have cascading effects on the CSS ecosystem, where ants are numerically dominant and thus may have broad influences on food web and ecosystem properties. Our results highlight that accurately predicting arthropod responses to invasive plant litter requires careful consideration of the structural and food resources provided by detritus to each particular food web.     

Evidence of exploitative competition among young stages of the wolf spider Schizocosa ocreata

Summary Previous research by many investigators has demonstrated food limitation in both web-building and wandering spiders. Field experiments have tested for exploitative competition for prey in web-building, but not wandering species. As a first step to examining the question of whether spiders without webs exhibit exploitative competition, we manipulated densities of young stages of a common wolf spider, Schizocos ocreata, and measured (1) spider growth rate and (2) numbers of Collembola, a potential prey organism. Replicate populations of recently hatched S. ocreata were established in 1-m² fenced plots at four levels: 0×, 0.25×, 1× and 4× natural density. Increasing spider density had a negative effect on spider growth rate, defined as increase in weight or cephalothorax width. Early in the experiment spider density had a weak negative effect on Collembola numbers [p(F)=0.08]. Taken together, this probable response by Collembola and the clear effect of spider density on growth rate constitute the first experimental evidence of intraspecific exploitative competition for prey in a species of wandering spider. We discuss (1) the strength of this evidence given the constraints of the experiment's design, and (2) the implications of the strong convergence in spider densities that had occurred after 2.5 months.     

The impact of altered precipitation on spatial stratification and activity-densities of springtails (Collembola) and spiders (Araneae)

Abstract.  1. A field experiment was conducted to determine how short-term changes in moisture can alter activity-densities of spiders and springtails.
2. In a Kentucky forest 10 unfenced 4-m² plots were divided into two rainfall treatments. A clear roof over five plots excluded rainfall to simulate severe drought conditions ( drought treatment). Water was sprayed on the five uncovered plots at a rate equal to two times the long-term mean in order to establish the high-rainfall treatment. Activity-densities of Collembola and spiders were measured using pitfall traps designed to sample the top, middle, and bottom layers of leaf litter. The experiment ran from 20 July to 23 September 2001.
3. Overall (i.e. litter layers pooled) activity-density (mean number trapped each sampling date) of Collembola was ≈ 60% lower in drought plots than in plots receiving increased precipitation. Surprisingly, overall spider activity-density was ≈ 1.6 times greater in the drought plots.
4. Differences in rainfall affected the spatial stratification of Collembola and spiders in strikingly different ways. Activity-densities of neither group differed between drought and high-rainfall treatments in the bottom litter layer. Collembola activity-density was three times greater in the top and middle litter layers in high-rainfall plots than in drought plots. In contrast, spider activity-density did not differ between treatments in the top layer, but activity-density was decreased by 50% in the middle layer of high-rainfall plots compared with drought plots.
5. Three Collembola families (Sminthuridae, Tomoceridae, and Entomobryidae) accounted for most of the Collembola pattern. The spider response was due to altered activity-density of one family of wandering spider, the Gnaphosidae.     

Interactions among young stages of the wolf spiders Pardosamoesta and P. mackenziana (Araneae: Lycosidae)

Young stages of the wolf spiders Pardosa moesta and P . mackenziana coexist on the forest floor in a deciduous forest in central Alberta, yet nothing is known of the potential interactions between and among these ubiquitous species. An experiment, using a series of 0.25-m² arenas, was designed to test for inter- and intraspecific exploitative competition. Experimental treatments were established in the arenas in late July. Treatments involved stocking recently hatched spiderlings of each species alone, and with both species together at natural (10 spiderlings per arena) and two times the natural densities. In September the arenas were searched for Pardosa spiderlings, and the survival and mass gain of each species were determined. Other predatory arthropods, and Collembola, a common prey of young wolf spiders, were counted in the arenas before the experimental treatments were established and at the end of the experiment. No evidence for competitive interactions was uncovered, but spiderling survival was 29% for P . moesta and 20% for P . mackenziana , suggesting other mortality factors (e.g., cannibalism, intraguild predation) may be important. Top-down effects (predatory arthropods) had no effect on Pardosa survival. Bottom-up effects, however, were significant as treatments containing spiderlings had fewer Collembola compared to control arenas, further supporting the importance of spiders in leaf-litter food webs.     

Grazing simplifies soil micro‐food webs and decouples their relationships with ecosystem functions in grasslands

Livestock grazing often alters aboveground and belowground communities of grasslands and their mediated carbon (C) and nitrogen (N) cycling processes at the local scale. Yet, few have examined whether grazing‐induced changes in soil food webs and their ecosystem functions can be extrapolated to a regional scale. We investigated how large herbivore grazing affects soil micro‐food webs (microbes and nematodes) and ecosystem functions (soil C and N mineralization), using paired grazed and ungrazed plots at 10 locations across the Mongolian Plateau. Our results showed that grazing not only affected plant variables (e.g., biomass and C and N concentrations), but also altered soil substrates (e.g., C and N contents) and soil environment (e.g., soil pH and bulk density). Grazing had strong bottom‐up effects on soil micro‐food webs, leading to more pronounced decreases at higher trophic levels (nematodes) than at lower trophic levels (microbes). Structural equation modeling showed that changes in plant biomass and soil environment dominated grazing effects on microbes, while nematodes were mainly influenced by changes in plant biomass and soil C and N contents; the grazing effects, however, differed greatly among functional groups in the soil micro‐food webs. Grazing reduced soil C and N mineralization rates via changes in plant biomass, soil C and N contents, and soil environment across grasslands on the Mongolian Plateau. Spearman's rank correlation analysis also showed that grazing reduced the correlations between functional groups in soil micro‐food webs and then weakened the correlation between soil micro‐food webs and soil C and N mineralization. These results suggest that changes in soil micro‐food webs resulting from livestock grazing are poor predictors of soil C and N processes at regional scale, and that the relationships between soil food webs and ecosystem functions depend on spatial scales and land‐use changes.     

Soil acidification reduces the effects of short‐term nutrient enrichment on plant and soil biota and their interactions in grasslands

Soil nitrogen (N) and phosphorus (P) contents, and soil acidification have greatly increased in grassland ecosystems due to increased industrial and agricultural activities. As major environmental and economic concerns worldwide, nutrient enrichment and soil acidification can lead to substantial changes in the diversity and structure of plant and soil communities. Although the separate effects of N and P enrichment on soil food webs have been assessed across different ecosystems, the combined effects of N and P enrichment on multiple trophic levels in soil food webs have not been studied in semiarid grasslands experiencing soil acidification. Here we conducted a short‐term N and P enrichment experiment in non‐acidified and acidified soil in a semiarid grassland on the Mongolian Plateau. We found that net primary productivity was not affected by N or P enrichment alone in either non‐acidified or acidified soil, but was increased by combined N and P enrichment in both non‐acidified and acidified soil. Nutrient enrichment decreased the biomass of most microbial groups in non‐acidified soil (the decrease tended to be greatest with combined N and P enrichment) but not in acidified soil, and did not affect most soil nematode variables in non‐acidified or acidified soil. Nutrient enrichment also changed plant and microbial community structure in non‐acidified but not in acidified soil, and had no effect on nematode community structure in non‐acidified or acidified soil. These results indicate that the responses to short‐term nutrient enrichment were weaker for higher trophic groups (nematodes) than for lower trophic groups (microorganisms) and primary producers (plants). The findings increase our understanding of the effects of nutrient enrichment on multiple trophic levels of soil food webs, and highlight that soil acidification, as an anthropogenic stressor, reduced the responses of plants and soil food webs to nutrient enrichment and weakened plant–soil interactions.     

Describing termite assemblage structure in a Peruvian lowland tropical rain forest: a comparison of two alternative methods

Termites are frequently dominant invertebrate decomposers and bioturbators in lowland tropical forests and therefore strongly influence ecosystem processes favouring soil stability, porosity and nutrient retention. In this study, we provide the first spatially replicated dataset on termite assemblage composition, abundance and biomass in a Peruvian rainforest by sampling six separate plots. In addition, two alternative sampling methods (transect method-TM and quadrat method-QM), providing termite species density data, were compared among the plots. The relationships between a range of environmental and spatial variables and species composition were examined using canonical correspondence analysis variation partitioning. We found that the TM captured a higher proportion of the known species in the site (82 %) compared with the QM (66 %). In addition, 56 % of the species sampled by TM were common between the plots while only 18 % of species overlapped using the QM. The QM may therefore potentially have undersampled the species pool. Environmental variables were shown to explain a larger proportion of the species patterns than the spatial variables with elevation, soil temperature and distance to the river being the most important. We discuss the impacts of the environmental and spatial variables on termite species composition.     

Habitat structure mediates top–down effects of spiders and ants on herbivores

Differences in structural complexity of habitats have been suggested to modify the extent of top–down forces in terrestrial food webs. In order to test this hypothesis, we manipulated densities of generalist invertebrate predators and the complexity of habitat structure in a two-factorial design. We conducted two field experiments in order to study predation effects of ants and spiders and, in particular, of the wasp spider Argiope bruennichi on herbivorous arthropods such as grasshoppers, plant- and leafhoppers in a grassland. Predator densities were manipulated by removal in habitats of higher and lower structural diversity, and the effects on herbivore densities were assessed by suction sampling. Habitat structure was changed by cutting the vegetation to half its height and removing leaf litter.We found a significant negative effect of this assemblage of generalist predators on plant- and leafhoppers, which were 1.6 times more abundant in predator removal plots. This effect was stronger in low-structured (cut) than in uncut vegetation. Densities of the most abundant planthopper Ribautodelphax pungens (Delphacidae) were 2.2 times higher in predator removal plots. Furthermore, adult plant- and leafhoppers responded more strongly than juveniles and epigeic species more strongly than hypergeic species. The presence of predators had a positive effect on plant- and leafhopper species diversity. In a second field experiment, we tested the exclusive impact of Argiope bruennichi on its prey, and found that its effect was also significant, although weaker than the effect of the predator assemblage. This effect was stronger in grass-dominated vegetation compared to structurally more complex mixed vegetation of grasses and herbs. We conclude that habitat structure and in particular vegetation height and architectural complexity strongly modify the strength of top–down forces and indirectly affect the diversity of herbivorous arthropods.     

Complex food webs prevent competitive exclusion among producer species

Herbivorous top-down forces and bottom-up competition for nutrients determine the coexistence and relative biomass patterns of producer species. Combining models of predator-prey and producer-nutrient interactions with a structural model of complex food webs, I investigated these two aspects in a dynamic food-web model. While competitive exclusion leads to persistence of only one producer species in 99.7% of the simulated simple producer communities without consumers, embedding the same producer communities in complex food webs generally yields producer coexistence. In simple producer communities, the producers with the most efficient nutrient-intake rates increase in biomass until they competitively exclude inferior producers. In food webs, herbivory predominantly reduces the biomass density of those producers that dominated in producer communities, which yields a more even biomass distribution. In contrast to prior analyses of simple modules, this facilitation of producer coexistence by herbivory does not require a trade-off between the nutrient-intake efficiency and the resistance to herbivory. The local network structure of food webs (top-down effects of the number of herbivores and the herbivores' maximum consumption rates) and the nutrient supply (bottom-up effect) interactively determine the relative biomass densities of the producer species. A strong negative feedback loop emerges in food webs: factors that increase producer biomasses also increase herbivory, which reduces producer biomasses. This negative feedback loop regulates the coexistence and biomass patterns of the producers by balancing biomass increases of producers and biomass fluxes to herbivores, which prevents competitive exclusion.