Bottom–up and top–down effects on plant communities: nutrients limit productivity,but insects determine diversity and composition

Top–down effects of herbivores and bottom–up effects of nutrients shape productivity and diversity across ecosystems, yet their single and combined effects on spatial and temporal beta diversity is unknown. We established a field experiment in which the abundance of insect herbivores (top–down) and soil nitrogen (bottom–up) were manipulated over six years in an existing old‐field community. We tracked plant α and β diversity – within plot richness and among plot biodiversity‐ and aboveground net primary productivity (ANPP) over the course of the experiment. We found that bottom–up factors affected ANPP while top–down factors influenced plant community structure. Across years, while N reduction lowered ANPP by 10%, N reduction did not alter ANPP relative to control plots. Further, N reduction lowered ANPP by 20% relative to N addition plots. On the other hand, the reduction of insect herbivores did not alter plant richness (α diversity) yet consistently promoted Shannon's evenness, relative to plots where insect herbivores were present. Further, insect herbivores promoted spatial‐temporal β diversity. Overall, we found that the relative importance of top–down and bottom–up controls of plant ANPP, plant α diversity, and composition (β diversity) can vary significantly in magnitude and direction. In addition, their effects varied through time, with bottom–up effects influencing ANPP quickly while the effects of top–down factors emerging only late in the experiment to influence plant community composition via shifts in plant dominance.     

Joint control by rodent herbivory and nutrient availability of plant diversity in a salt marsh–salty steppe transition zone

Questions: Do current models that predict shifting effects of herbivores on plant diversity with varying nutrient conditions apply to stressful systems like salt marshes? Do herbivores affect different components of the diversity as nutrient availability varies? Location: Salt marsh–salty steppe transition zone at the SW Atlantic Mar Chiquita coastal lagoon (37°44′52″S, 57°26′6″W), Argentina. Methods: We experimentally evaluated the separate and interactive effect of nutrients and rodent (Cavia aperea) herbivory, using exclosures and applying fertilizer (mostly nitrogen), following a factorial design in 50 cm × 50 cm plots. Results: We found a negative effect of herbivory on diversity in the resource‐poor scenario (due to a reduction in species richness), but a positive effect when nutrients were added, by reducing the abundance of the dominant plant (and hence increasing evenness). Conclusions: Our experimental results contribute to the limited factorial evidence evaluating the role of nutrients and herbivory on the diversity of terrestrial plant communities, even in highly stressful environments like salt marsh–salty steppe transition zones. Our results also support the model that predicts negative effects of herbivores on plant diversity in low‐nutrient conditions and positive effects in nutrient‐enriched scenarios, and also support the mechanism assumed to act in these situations.     

Crab herbivory regulates re‐colonization of disturbed patches in a southwestern Atlantic salt marsh

Recent work exploring the effects of physical stress and herbivory on secondary succession in estuarine plant communities agrees with basic stress models and reveal that herbivory is an important force in brackish and oligohaline marshes but negligible in physically stressful salt marshes. In these systems, herbivores are terrestrial, and thus negatively affected by the same stressful factors that affect marsh plants (i.e. frequent flooding or high salinities). We evaluated the effects of a marine herbivore (i.e. the crab Neohelice granulata) on plant secondary succession in a southwestern Atlantic salt marsh. Field surveys revealed that disturbance‐generated bare patches have harsh physical conditions and that their edges suffer higher herbivore pressure compared to the marsh matrix. A factorial experiment demonstrated that asexual expansion of the surrounding plants is the only possible mechanism to re‐colonize disturbed patches and that crab exclusion can increase this colonization rate by more than 30 times. Our results show that even in highly stressful environments, herbivores strongly impact marsh structure by regulating patch recovery. The synergism of physical stress and herbivory may make plant succession an extremely slow process and lead to the prevalence of bare areas.     

Biogeographic consequences of nutrient enrichment for plant–herbivore interactions in coastal wetlands

A major challenge in ecology is to understand broadscale trends in the impact of environmental change. We provide the first integrative analysis of the effects of eutrophication on plants, herbivores, and their interactions in coastal wetlands across latitudes. We show that fertilisation strongly increases herbivory in salt marshes, but not in mangroves, and that this effect increases with increasing latitude in salt marshes. We further show that stronger nutrient effects on plant nitrogen concentration at higher latitudes is the mechanism likely underlying this pattern. This biogeographic variation in nutrient effects on plant–herbivore interactions has consequences for vegetation, with those at higher latitudes being more vulnerable to consumer pressure fuelled by eutrophication. Our work provides a novel, mechanistic understanding of how eutrophication affects plant–herbivore systems predictably across broad latitudinal gradients, and highlights the power of incorporating biogeography into understanding large‐scale variability in the impacts of environmental change.     

Biodiversity mediates top–down control in eelgrass ecosystems: a global comparative‐experimental approach

Nutrient pollution and reduced grazing each can stimulate algal blooms as shown by numerous experiments. But because experiments rarely incorporate natural variation in environmental factors and biodiversity, conditions determining the relative strength of bottom–up and top–down forcing remain unresolved. We factorially added nutrients and reduced grazing at 15 sites across the range of the marine foundation species eelgrass (Zostera marina) to quantify how top–down and bottom–up control interact with natural gradients in biodiversity and environmental forcing. Experiments confirmed modest top–down control of algae, whereas fertilisation had no general effect. Unexpectedly, grazer and algal biomass were better predicted by cross‐site variation in grazer and eelgrass diversity than by global environmental gradients. Moreover, these large‐scale patterns corresponded strikingly with prior small‐scale experiments. Our results link global and local evidence that biodiversity and top–down control strongly influence functioning of threatened seagrass ecosystems, and suggest that biodiversity is comparably important to global change stressors.     

Seasonal shifts in the importance of bottom–up and top–down factors on stream periphyton community structure

We examined the importance of temporal variability in top–down and bottom–up effects on the accumulation of stream periphyton, which are complex associations of autotrophic and heterotrophic microorganisms. Periphyton contributes to primary production and nutrient cycling and serves as a food resource for herbivores (grazers). Periphyton growth is often limited by the availability of nitrogen and phosphorus, and biomass can be controlled by grazers. In this study we experimentally manipulated nutrients and grazers simultaneously to determine the relative contribution of bottom–up and top–down controls on periphyton over time. We used nutrient diffusing substrates to regulate nutrient concentrations and an underwater electric field to exclude grazing insects in three sequential 16–17 day experiments from August to October in montane Colorado, USA. We measured algal biomass, periphyton organic mass, and algal community composition in each experiment and determined densities of streambed insect species, including grazers. Phosphorus was the primary limiting nutrient for algal biomass, but it did not influence periphyton organic mass across all experiments. Effects of nutrient additions on algal biomass and community composition decreased between August and October. Grazed substrates supported reduced periphyton biomass only in the first experiment, corresponding to high benthic abundances of a dominant mayfly grazer (Rhithrogena spp.). Grazed substrates in the first experiment also showed altered algal community composition with reduced diatom relative abundances, presumably in response to selective grazing. We showed that top–down grazing effects were strongest in late summer when grazers were abundant. The effects of phosphorus additions on algal biomass likely decreased over time because temperature became more limiting to growth than nutrients, and because reduced current velocity decreased nutrient uptake rates. These results suggest that investigators should proceed with caution when extending findings based on short‐term experiments. Furthermore, these results support the need for additional seasonal‐scale field research in stream ecology.     

Nitrate reduction capacity is limited by belowground plant recovery in a 32‐year‐old created salt marsh

Human activities have decreased global salt marsh surface area with a subsequent loss in the ecosystem functions they provide. The creation of marshes in terrestrial systems has been used to mitigate this loss in marsh cover. Although these constructed marshes may rapidly recover ecosystem structure, biogeochemical processes may be slow to recover. We compared denitrification and dissimilatory nitrate reduction to ammonium (DNRA) rates between a 32‐year‐old excavation‐created salt marsh (CON‐2) and a nearby natural reference salt marsh (NAT) to assess the recovery of ecosystem function. These process rates were measured at 5 cm increments to a depth of 25 cm to assess how plant rooting depth and organic matter accumulation impact N‐cycling. We found that, for both marshes, denitrification and DNRA declined with depth with the highest rates occurring in the top 10 cm. In both systems, N‐retention by DNRA accounted for upwards of 75% of nitrate reduction, but denitrification and DNRA rates were nearly 2× and 3× higher in NAT than CON‐2, respectively. Organic matter was 6× lower in CON‐2, likely due to limited plant belowground biomass production. However, there was no response to glucose additions, suggesting that the microbial functional community, not substrate limitation, limited nitrate reduction recovery. Response ratios showed that denitrification in CON‐2 recovered in surficial sediments where belowground biomass was highest, even though biomass recovery was minimal. This indicates that although recovery of ecosystem function was constrained, it occurred on a faster trajectory than that of ecosystem structure.     

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.     

The role of light availability and herbivory on algal responses to nutrient enrichment in a riparian wetland,Alaska

We investigated how the relative availability of solar radiation in the presence or absence of grazing alters the ability of benthic algae to respond to nutrient enrichment in an Alaskan marsh. We used a factorial mesocosm experiment that included nutrient enrichment (enriched or control), grazing (grazed or ungrazed), and light (unshaded or shaded) to simulate shading by macrophytes early and late in the growing season, respectively. We found stronger effects of grazers and nutrients compared to light on benthic algal biomass and taxonomic composition. Algal biomass increased in nutrient‐enriched treatments and was reduced by grazing. Shading did not have an effect on algal biomass or taxonomic composition, but the concentration of chl a per algal biovolume increased with shading, demonstrating the ability of algae to compensate for changes in light availability. Algal taxonomic composition was more affected by grazer presence than nutrients or light. Grazer‐resistant taxa (basal filaments of Stigeoclonium) were replaced by diatoms (Nitzschia) and filamentous green algae (Ulothrix) when herbivores were removed. The interacting and opposing influences of nutrients and grazing indicate that the algal community is under dual control from the bottom‐up (nutrient limitation) and from the top‐down (consumption by herbivores), although grazers had a stronger influence on algal biomass and taxonomic composition than nutrient enrichment. Our results suggest that low light availability will not inhibit the algal response to elevated nutrient concentrations expected with ongoing climate change, but grazers rapidly consume algae following enrichment, masking the effects of elevated nutrients on algal production.     

The natural regeneration of salt marsh on formerly reclaimed land

Question: Does the vegetation of restored salt marshes increasingly resemble natural reference communities over time? Location: The Essex estuaries, southeast England. Methods: Abandoned reclamations, where coastal defences had been breached in storm events, and current salt marsh recreation schemes were surveyed giving a chronosequence of salt marsh regeneration from 2 to 107 years. The presence, abundance and height of plant species were recorded and comparisons were made with adjacent reference salt marsh communities at equivalent elevations. Results: Of the 18 paired sites surveyed, 13 regenerated marshes had fewer species than their adjacent reference marsh, three had an equal number and two had more. The plant communities of only two de‐embankment sites matched that of the reference community. 0–50 year old sites and 51–100 year old sites had fewer species per quadrat than the 101+ year sites and the reference salt marshes. There was a weak relationship between differences in species richness for regenerated and reference marshes and the time since sites were first re‐exposed to tidal inundation. Cover values for the invasive and recently evolved Spartina anglica were greater within regenerated than reference marshes. Conclusions: Salt marsh plants will colonise formerly reclaimed land relatively quickly on resumption of tidal flooding. However, even after 100 years regenerated salt marshes differ in species richness, composition and structure from reference communities.     

Biogeography of salt marsh plant zonation on the Pacific coast of South America

Aim

The aim of this study was to investigate the biogeography of plant zonation in salt marshes on the Pacific coast of South America; to examine whether salt marsh plant zonation varies with latitude; and to explore the relative importance of climatic, tidal, edaphic and disturbance factors in explaining large‐scale variation in salt marsh plant community structure.

Location

A 2,000‐km latitudinal gradient on the Pacific coast in Chile, with a climate shift from hyper‐arid at low to hyper‐humid at high latitudes.

Methods

Plant zonation was quantified in field surveys of ten marshes. Climate, tidal regimes, edaphic factors and disturbances (tsunami and rainfall floods) were determined. We used multivariate analyses to explore their relative importance in explaining large‐scale variation in salt marsh plant community structure.

Results

Across latitude, marshes were dominated by distinct plant communities in different climate regions, especially at the extreme dry and wet latitudes. Intertidal plant species zonation was present in hyper‐arid and semi‐arid climates, but not in arid, humid and hyper‐humid climates. Latitudinal variation in low‐marsh plant communities (regularly flooded at high tide) was largely a function of precipitation, while at high marshes (never flooded at high tide) latitudinal variation was explained with precipitation, temperature, tidal cycles, soil salinity and disturbances.

Main conclusions

Salt marshes on the Pacific coast of South America belong to Dry Coast and Temperate biogeographic types. Salt marsh plant zonation varies across latitude, and is explained by climatic, tidal, edaphic and disturbance factors. These patterns appear to be mechanistically explained by extrapolating experimentally generated community assembly models and have implications for predicting responses to climate change.     

Abiotic stress mediates top-down and bottom-up control in a Southwestern Atlantic salt marsh

Increasing evidence has shown that nutrients and consumers interact to control primary productivity in natural systems, but how abiotic stress affects this interaction is unclear. Moreover, while herbivores can strongly impact zonation patterns in a variety of systems, there are few examples of this in salt marshes. We evaluated the effect of nutrients and herbivores on the productivity and distribution of the cordgrass Spartina densiflora along an intertidal stress gradient, in a Southwestern Atlantic salt marsh. We characterized abiotic stresses (salinity, ammonium concentration, and anoxia) and manipulated nutrients and the presence of the herbivorous crab Neohelice (Chasmagnathus) granulata, at different tidal heights with a factorial experiment. Abiotic stress increased at both ends of the tidal gradient. Salinity and anoxia were highest at the upper and lower edge of the intertidal, respectively. Nutrients and herbivory interacted to control cordgrass biomass, but their relative importance varied with environmental context. Herbivory increased at lower tidal heights to the point that cordgrass transplants onto bare mud substrate were entirely consumed unless crabs were excluded, while nutrients were most important where abiotic stress was reduced. Our results show how the impact of herbivores and nutrients on plant productivity can be dependent on environmental conditions and that the lower intertidal limits of marsh plants can be controlled by herbivory.     

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.     

Indirect and direct controls of macroinvertebrates and small fish by abiotic factors and trophic interactions in the Florida Everglades

1. The roles of nutrients, disturbance and predation in regulating consumer densities have long been of interest, but their indirect effects have rarely been quantified in wetland ecosystems. The Florida Everglades contains gradients of hydrological disturbance (marsh drying) and nutrient enrichment (phosphorus), often correlated with densities of macroinvertebrate infauna (macroinvertebrates inhabiting periphyton), small fish and larger invertebrates, such as snails, grass shrimp, insects and crayfish. However, most causal relationships have yet to be quantified. 2. We sampled periphyton (content and community structure) and consumer (small omnivores, carnivores and herbivores, and infaunal macroinvertebrates inhabiting periphyton) density at 28 sites spanning a range of hydrological and nutrient conditions and compared our data to seven a priori structural equation models. 3. The best model included bottom‐up and top‐down effects among trophic groups and supported top‐down control of infauna by omnivores and predators that cascaded to periphyton biomass. The next best model included bottom‐up paths only and allowed direct effects of periphyton on omnivore density. Both models suggested a positive relationship between small herbivores and small omnivores, indicating that predation was unable to limit herbivore numbers. Total effects of time following flooding were negative for all three consumer groups even when both preferred models suggested positive direct effects for some groups. Total effects of nutrient levels (phosphorus) were positive for consumers and generally larger than those of hydrological disturbance and were mediated by changes in periphyton content. 4. Our findings provide quantitative support for indirect effects of nutrient enrichment on consumers, and the importance of both algal community structure and periphyton biomass to Everglades food webs. Evidence for top‐down control of infauna by omnivores was noted, though without substantially greater support than a competing bottom‐up‐only model.     

Role of biotic interactions and physical factors in determining the distribution of marsh species along an estuarine salinity gradient

Understanding the mechanisms that shape plant distribution patterns is a major goal in ecology. We investigated the role of biotic interactions (competition and facilitation) and abiotic factors in creating horizontal plant zonation along salinity gradients in the Elbe estuary. We conducted reciprocal transplant experiments with four dominant species from salt and tidal freshwater marshes at two tidal elevations. Ten individuals of each species were transplanted as sods to the opposing marsh type and within their native marsh (two sites each). Transplants were placed at the centre of 9‐m² plots along a line parallel to the river bank. In order to disentangle abiotic and biotic influences, we set up plots with and without neighbouring vegetation, resulting in five replicates per site. Freshwater species (Bolboschoenus maritimus and Phragmites australis) transplanted to salt marshes performed poorly regardless of whether neighbouring vegetation was present or not, although 50–70% of the transplants did survive. Growth of Phragmites transplants was impaired also by competition in freshwater marshes. Salt marsh species (Spartina anglica and Puccinellia maritima) had extremely low biomass when transplanted to freshwater marshes and 80–100% died in the presence of neighbours. Without neighbours, biomass of salt marsh species in freshwater marshes was similar to or higher than that in salt marshes. Our results indicate that salt marsh species are precluded from freshwater marshes by competition, whereas freshwater species are excluded from salt marshes by physical stress. Thus, our study provides the first experimental evidence from a European estuary for the general theory that species boundaries along environmental gradients are determined by physical factors towards the harsh end and by competitive ability towards the benign end of the gradient. We generally found no significant impact of competition in salt marshes, indicating a shift in the importance of competition along the estuarine gradient.     

Ecosystem consequences of diversity depend on food chain length in estuarine vegetation

Biodiversity and food chain length each can strongly influence ecosystem functioning, yet their interactions rarely have been tested. We manipulated grazer diversity in seagrass mesocosms with and without a generalist predator and monitored community development. Changing food chain length altered biodiversity effects: higher grazer diversity enhanced secondary production, epiphyte grazing, and seagrass biomass only with predators present. Conversely, changing diversity altered top‐down control: predator impacts on grazer and seagrass biomass were weaker in mixed‐grazer assemblages. These interactions resulted in part from among‐species trade‐offs between predation resistance and competitive ability. Despite weak impact on grazer abundance at high diversity, predators nevertheless enhanced algal biomass through a behaviourally mediated trophic cascade. Moreover, predators influenced every measured variable except total plant biomass, suggesting that the latter is an insensitive metric of ecosystem functioning. Thus, biodiversity and trophic structure interactively influence ecosystem functioning, and neither factor's impact is predictable in isolation.     

The influence of legacy effects and recovery from perturbations in a tritrophic salt marsh complex

Abstract. 1. Much has been learned in recent years regarding the influence of environmental conditions on top‐down and bottom‐up effects acting on insect herbivores. Temporal variation in environmental conditions, however, has gone largely unstudied in spite of undoubtedly strong influences in most systems. 2. A 2‐year study was conducted to examine the legacy effects of previous manipulations of host plant quality and parasitism pressure on the top‐down and bottom‐up effects influencing population densities of the salt marsh planthopper Pissonotus quadripustulatus. 3. For 10 months in 1998, a 2 × 2 factorial experiment was carried out, in which host plant quality was increased by the addition of nitrogen fertiliser, and parasitism pressure was decreased through the use of yellow sticky traps. This was followed by 2 months in the winter with no treatment applications. Treatments were then reversed in 1999 for a further 10 months. 4. In 1998, fertilisation treatments increased plant quality significantly, which resulted in strong effects on P. quadripustulatus density. Parasitism reduction treatments had weaker and time‐dependent effects on the herbivore, increasing planthopper density in late summer and autumn. 5. After 2 months without treatments, previous fertilisation treatments were still influencing all response variables measured significantly. The legacy effects of fertilisation persisted for at least 7 months for the host plant, and at least 5 months for the herbivore and parasitoid. 6. Fertilisation treatments in 1999 increased P. quadripustulatus density by approximately the same percentage as in 1998, suggesting that previous reductions in parasitism had no influence on herbivore responses to increased nutrients. Parasitism reduction treatments in 1999, however, resulted in greater increases in herbivore density than in 1998, suggesting that previous increases in nutrients enabled greater responses to reductions in top‐down pressure. 7. The results show that the top‐down effects of parasitism attenuated more quickly than did the bottom‐up effects of increased plant quality through greater nutrient availability. They also suggest that the recent history of nutrient status in an ecosystem may be important in determining the relative strengths of top‐down and bottom‐up forces.     

Emergence of nutrient co‐limitation through movement in stoichiometric meta‐ecosystems

Evidence that ecosystems and primary producers are limited in their productivity by multiple nutrients has caused the traditional nutrient limitation framework to include multiple limiting nutrients. The models built to mimic these responses have invoked local mechanisms at the level of the primary producers. In this paper, we explore an alternative explanation for the emergence of co‐limitation by developing a simple, stoichiometrically explicit meta‐ecosystem model with two limiting nutrients, autotrophs and herbivores. Our results show that differences in movement rates for the nutrients, autotrophs and herbivores can allow for nutrient co‐limitation in biomass response to emerge despite no local mechanisms of nutrient co‐limitation. Furthermore, our results provide an explanation to why autotrophs show positive growth responses to nutrients despite ‘nominal’ top‐down control by herbivores. These results suggest that spatial processes can be mechanisms for nutrient co‐limitation at local and regional scales, and can help explain anomalous results in the co‐limitation literature.     

Bottom‐up and top‐down effects of tree species diversity on leaf insect herbivory

The diversity of plant neighbors commonly results in direct, bottom‐up effects on herbivore ability to locate their host, and in indirect effects on herbivores involving changes in plant traits and a top‐down control by their enemies. Yet, the relative contribution of bottom‐up and top‐down forces remains poorly understood. We also lack knowledge on the effect of abiotic constraints such as summer drought on the strength and direction of these effects. We measured leaf damage on pedunculate oak (Quercus robur), alone or associated with birch, pine or both in a long‐term tree diversity experiment (ORPHEE), where half of the plots were irrigated while the other half remained without irrigation and received only rainfall. We tested three mechanisms likely to explain the effects of oak neighbors on herbivory: (1) Direct bottom‐up effects of heterospecific neighbors on oak accessibility to herbivores, (2) indirect bottom‐up effects of neighbors on the expression of leaf traits, and (3) top‐down control of herbivores by predators. Insect herbivory increased during the growth season but was independent of neighbor identity and irrigation. Specific leaf area, leaf toughness, and thickness varied with neighbor identity while leaf dry matter content or C:N ratio did not. When summarized in a principal component analysis (PCA), neighbor identity explained 87% of variability in leaf traits. PCA axes partially predicted herbivory. Despite greater rates of attack on dummy caterpillars in irrigated plots, avian predation, and insect herbivory remained unrelated. Our study suggests that neighbor identity can indirectly influence insect herbivory in mixed forests by modifying leaf traits. However, we found only partial evidence for these trait‐mediated effects and suggest that more attention should be paid to some unmeasured plant traits such as secondary metabolites, including volatile organic compounds, to better anticipate the effects of climate change on plant‐insect interactions in the future.     

Above- and belowground insect herbivores differentially affect soil nematode communities in species-rich plant communities

Interactions between above‐ and belowground invertebrate herbivores alter plant diversity, however, little is known on how these effects may influence higher trophic level organisms belowground. Here we explore whether above‐ and belowground invertebrate herbivores which alter plant community diversity and biomass, in turn affect soil nematode communities. We test the hypotheses that insect herbivores 1) alter soil nematode diversity, 2) stimulate bacterial‐feeding and 3) reduce plant‐feeding nematode abundances. In a full factorial outdoor mesocosm experiment we introduced grasshoppers (aboveground herbivores), wireworms (belowground herbivores) and a diverse soil nematode community to species‐rich model plant communities. After two years, insect herbivore effects on nematode diversity and on abundance of herbivorous, bacterivorous, fungivorous and omni‐carnivorous nematodes were evaluated in relation to plant community composition. Wireworms did not affect nematode diversity despite enhanced plant diversity, while grasshoppers, which did not affect plant diversity, reduced nematode diversity. Although grasshoppers and wireworms caused contrasting shifts in plant species dominance, they did not affect abundances of decomposer nematodes at any trophic level. Primary consumer nematodes were, however, strongly promoted by wireworms, while community root biomass was not altered by the insect herbivores. Overall, interaction effects of wireworms and grasshoppers on the soil nematodes were not observed, and we found no support for bottom‐up control of the nematodes. However, our results show that above‐ and belowground insect herbivores may facilitate root‐feeding rather than decomposer nematodes and that this facilitation appears to be driven by shifts in plant species composition. Moreover, the addition of nematodes strongly suppressed shoot biomass of several forb species and reduced grasshopper abundance. Thus, our results suggest that nematode feedback effects on plant community composition, due to plant and herbivore parasitism, may strongly depend on the presence of insect herbivores.