Bottom-up effects of species diversity on the functioning and stability of food webs

1. The importance of species diversity for the stability of populations, communities and ecosystem functions is a central question in ecology. 2. Biodiversity experiments have shown that diversity can impact both the average and variability of stocks and rates at these levels of ecological organization in single trophic-level ecosystems. Whether these impacts hold in food webs and across trophic levels is still unclear. 3. We asked whether resource species diversity, community composition and consumer feeding selectivity in planktonic food webs impact the stability of resource or consumer populations, community biomass and ecosystem functions. We also tested the relative importance of resource diversity and community composition. 4. We found that resource diversity negatively affected resource population stability, but had no effect on consumer population stability, regardless of the consumer's feeding selectivity. Resource diversity had positive effects on most ecosystem functions and their stability, including primary production, resource biomass and particulate carbon, nitrogen and phosphorus concentrations. 5. Community composition, however, generally explained more variance in population, community and ecosystem properties than species diversity per se. This result points to the importance of the outcomes of particular species interactions and individual species' effect traits in determining food web properties and stability. 6. Among the stabilizing mechanisms tested, an increase in the average resource community biomass with increasing resource diversity had the greatest positive impact on stability. 7. Our results indicate that resource diversity and composition are generally important for the functioning and stability of whole food webs, but do not have straightforward impacts on consumer populations.     

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.     

高寒草甸植物群落中物种多样性与群落变异性的关系及其机制初探

通过对高寒草甸植物群落中采集的群落数据进行分析,探讨了物种构成的相似性、统计平均、种群变异性和净协方差等机制对形成高寒草甸植物群落中多样性与群落地上生物量变异性之间关系的影响。结果表明,地上生物量的年际变异性随着多样性的增加而减小。物种构成相似性是多样性一变异性负关系产生的原因之一,而相似性与多样性之间并无显著相关关系;统计平均效应是另一个多样性一变异性负关系的主要决定者;净协方差效应、种群变异性对多样性一变异性关系产生的影响不显著或非常有限。     

Punching above their weight: low-biomass non-native plant species alter soil properties during primary succession

Non‐native invasive plants can greatly alter community and ecosystem properties, but efforts to predict which invasive species have the greatest impacts on these properties have been generally unsuccessful. An hypothesis that has considerable promise for predicting the effects of invasive non‐native plant species is the mass ratio hypothesis (i.e. that dominant species exert the strongest effects). We tested this hypothesis using data from a four year removal experiment in which the presence of two dominant shrub species (one native and the other not), and subordinate plant species, were manipulated in factorial combinations over four years in a primary successional floodplain system. We measured the effects of these manipulations on the plant community, soil nutrient status and soil biota in different trophic levels of the soil food web. Our experiment showed that after four years, low‐biomass non‐native plant species exerted disproportionate belowground effects relative to their contribution to total biomass in the plant community, most notably by increasing soil C, soil microbial biomass, altering soil microbial community structure and increasing the abundance of microbial‐feeding and predatory nematodes. Low‐biomass, non‐native plant species had distinct life history strategies and foliar traits (higher foliar N concentrations and higher leaf area per unit mass) compared with the two dominant shrub species (97% of total plant mass). Our results have several implications for understanding species’ effects in communities and on soil properties. First, high‐biomass species do not necessarily exert the largest impacts on community or soil properties. Second, low‐biomass, inconspicuous non‐native species can influence community composition and have important trophic consequences belowground through effects on soil nutrient status or resource availability to soil biota. Our finding that low‐biomass non‐native species influence belowground community structure and soil properties more profoundly than dominant species demonstrates that the mass ratio hypothesis does not accurately predict the relative effects of different coexisting species on community‐ and ecosystem‐level properties.     

On the significance of belowground overyielding in temperate mixed forests: separating species identity and species diversity effects

Complementary soil exploration by the root systems of coexisting tree species has been hypothesised to result in a higher root biomass of mixed forests than of monocultures but the existing evidence for a belowground diversity effect in forests is scarce and not conclusive. In a species‐rich temperate broad‐leaved forest, we analysed the fine root biomass (roots ≤ 2 mm) and necromass in 100 plots differing in tree species diversity (one to three species) and species composition (all possible combinations of five species of the genera Acer, Carpinus, Fagus, Fraxinus and Tilia) which allowed us to separate possible species diversity and species identity effects on fine root biomass. We found no evidence of a positive diversity effect on standing fine root biomass and thus of overyielding in terms of root biomass. Root necromass decreased with increasing species diversity at marginal significance. Various lines of evidence indicate significant species identity effects on fine root biomass (10–20% higher fine root biomass in plots with presence of maple and beech than in plots with hornbeam; 100% higher fine root biomass in monospecific beech and ash plots than in hornbeam plots; differences significant). Ash fine roots tended to be over‐represented in the 2‐ and 3‐species mixed plots compared to monospecific ash plots pointing at apparent belowground competitive superiority of Fraxinus in this mixed forest. Our results indicate that belowground overyielding and spatial complementarity of root systems may be the exception rather than the rule in temperate mixed forests.     

Greater than the sum of the parts: how the species composition in different forest strata influence ecosystem function

The mechanisms underpinning forest biodiversity‐ecosystem function relationships remain unresolved. Yet, in heterogeneous forests, ecosystem function of different strata could be associated with traits or evolutionary relationships differently. Here, we integrate phylogenies and traits to evaluate the effects of elevational diversity on above‐ground biomass across forest strata and spatial scales. Community‐weighted means of height and leaf phosphorous concentration and functional diversity in specific leaf area exhibited positive correlations with tree biomass, suggesting that both positive selection effects and complementarity occur. However, high shrub biomass is associated with greater dissimilarity in seed mass and multidimensional trait space, while species richness or phylogenetic diversity is the most important predictor for herbaceous biomass, indicating that species complementarity is especially important for understory function. The strength of diversity‐biomass relationships increases at larger spatial scales. We conclude that strata‐ and scale‐ dependent assessments of community structure and function are needed to fully understand how biodiversity influences ecosystem function.     

Short‐term harvesting of biomass from conservation grasslands maintains plant diversity

High yields are a priority in managing biomass for renewable energy, but the environmental impacts of various feedstocks and production systems should be equally considered. Mixed‐species, perennial grasslands enrolled in conservation programs are being considered as a source of biomass for renewable energy. Conservation grasslands are crucial in sustaining native biodiversity throughout the US Upper Midwest, and the effects of biomass harvest on biodiversity are largely unknown. We measured the effect of late‐season biomass harvest on plant community composition in conservation grasslands in three regions of Minnesota, USA from 2009 to 2012. Temporal trends in plant species composition within harvested grasslands were compared to unharvested grasslands using mixed effects models. A before‐after control‐impact approach using effect sizes was applied to focus on pre‐ and postharvest conditions. Production‐scale biomass harvest did not affect plant species richness, species or functional group diversity, nor change the relative abundance of the main plant functional groups. Differences in the relative abundances of plant functional groups were observed across locations; and at some locations, changed through time. The proportion of non‐native species remained constant, while the proportion of noxious weeds decreased through time in both harvested and unharvested grasslands at the central location. Ordination revealed patterns in species composition due to location, but not due to harvest treatment. Therefore, habitat and bioenergy characteristics related to grassland plant communities are not expected to change due to short‐term or intermittent late‐season biomass harvest.     

Contrasting effects of diversity on the temporal stability of plant populations

Recent theoretical and empirical work suggests that diversity enhances the temporal stability of a community. However, the effect of diversity on the stability of the individual populations within the community remains unclear. Some models predict a decrease of population stability with diversity, whereas others suggest that diversity has a stabilizing effect on populations. Empirical evidence for either relationship between population stability and diversity is weak. The few studies that directly assessed the stability of populations reported contradicting results. We used a six-year data-set from a plant diversity experiment to examine the relationships between diversity and temporal stability of plant biomass. Our results show that stability increased with diversity at the community-level, while the stability of populations, averaged over all species, decreased with diversity. However, when examining species separately we found positive, negative and neutral relationships between population stability and diversity. Our findings suggest that diversity may contribute to the stability of ecosystem services at the community level, but the effect of diversity on the stability of the individual populations within the community are generally negative. However, different species within the community may show strikingly different relationships between diversity and stability.     

Drivers of productivity and its temporal stability in a tropical tree diversity experiment

There is increasing evidence that mixed‐species forests can provide multiple ecosystem services at a higher level than their monospecific counterparts. However, most studies concerning tree diversity and ecosystem functioning relationships use data from forest inventories (under noncontrolled conditions) or from very young plantation experiments. Here, we investigated temporal dynamics of diversity–productivity relationships and diversity–stability relationships in the oldest tropical tree diversity experiment. Sardinilla was established in Panama in 2001, with 22 plots that form a gradient in native tree species richness of one‐, two‐, three‐ and five‐species communities. Using annual data describing tree diameters and heights, we calculated basal area increment as the proxy of tree productivity. We combined tree neighbourhood‐ and community‐level analyses and tested the effects of both species diversity and structural diversity on productivity and its temporal stability. General patterns were consistent across both scales indicating that tree–tree interactions in neighbourhoods drive observed diversity effects. From 2006 to 2016, mean overyielding (higher productivity in mixtures than in monocultures) was 25%–30% in two‐ and three‐species mixtures and 50% in five‐species stands. Tree neighbourhood diversity enhanced community productivity but the effect of species diversity was stronger and increased over time, whereas the effect of structural diversity declined. Temporal stability of community productivity increased with species diversity via two principle mechanisms: asynchronous responses of species to environmental variability and overyielding. Overyielding in mixtures was highest during a strong El Niño‐related drought. Overall, positive diversity–productivity and diversity–stability relationships predominated, with the highest productivity and stability at the highest levels of diversity. These results provide new insights into mixing effects in diverse, tropical plantations and highlight the importance of analyses of temporal dynamics for our understanding of the complex relationships between diversity, productivity and stability. Under climate change, mixed‐species forests may provide both high levels and high stability of production.     

Comparison of size-structured and species-level trophic networks reveals antagonistic effects of temperature on vertical trophic diversity at the population and species level

It is predicted that warmer conditions should lead to a loss of trophic levels, as larger bodied consumers, which occupy higher trophic levels, experience higher metabolic costs at high temperature. Yet, it is unclear whether this prediction is consistent with the effect of warming on the trophic structure of natural systems. Furthermore, effects of temperature at the species level, which arise through a change in species composition, may differ from those at the population level, which arise through a change in population structure. We investigate this by building species-level trophic networks, and size-structured trophic networks, as a proxy for population structure, for 18 648 stream fish communities, from 4 145 234 individual fish samples, across 7024 stream locations in France from 1980 to 2008. We estimated effects of temperature on total trophic diversity (total number of nodes), vertical trophic diversity (mean and maximum trophic level) and distribution of biomass across trophic level (correlation between trophic level and biomass) in these networks. We found a positive effect of temperature on total trophic diversity in both species- and size-structured trophic networks. We found that maximum trophic level and biomass distribution decreased in species-level and size-structured trophic networks, but the mean trophic level decreased only in size-structured trophic networks. These results show that warmer temperatures associate with a lower vertical trophic diversity in size-structured networks, and a higher one in species-level networks. This suggests that vertical trophic diversity is shaped by antagonistic effects of temperature on population structure and on species composition. Our results hence demonstrate that effects of temperature do not only differ across trophic levels, but also across levels of biological organisation, from population to species level, implying complex changes in network structure and functioning with warming.     

Using functional trait diversity to evaluate the contribution of multiple ecological processes to community assembly during succession

Successional chronosequences provide a unique opportunity to study the effects of multiple ecological processes on plant community assembly. Using a series of 0.5 × 0.5 m² plots (n = 30) from five successional sub‐alpine meadow plant communities (ages 3, 5, 9, 12, and undisturbed) in the Qinghai‐Tibetan Plateau, we investigated whether community assembly is stochastic or deterministic for species and functional traits. We tested directional change in species composition, functional trait composition, and then functional trait diversity measured by Rao's quadratic entropy for four traits – plant height, leaf dry matter content, specific leaf area, and seed mass – along two comparable successional chronosequences. We then evaluated the importance of species interactions, habitat filtering and stochasticity by comparing with random communities and partitioning the environmental and spatial components of Rao's quadratic entropy. We found no directional change in species composition, but clear directionality in functional trait composition. None of the abiotic environmental variables (except P) showed linear change with successional age, but soil moisture and nitrogen were positively related to functional diversity within meadows. Functional trait diversity increased significantly with the increase in successional age. Comparison with random communities showed a significant shift from trait divergence in early stages of succession (3‐ and 5‐yr) to convergence in the later stages of succession 9‐, 12‐yr and undisturbed). The relative importance of abiotic variables and spatial structure for functional trait diversity changed in a predictable manner with successional age. Stochasticity at the species level may indicate dispersal limitation, but deterministic effects on functional trait distributions show the role of both habitat effects and biotic interactions.     

Resource–consumer diversity: testing the effects of leaf litter species diversity on stream macroinvertebrate communities

1. Understanding relationships between resource and consumer diversity is essential to predicting how changes in resource diversity might affect several trophic levels and overall ecosystem functioning. 2. We tested for the effects of leaf litter species diversity (i.e. litter mixing) on litter mass remaining and macroinvertebrate communities (taxon diversity, abundance and biomass) during breakdown in a detritus‐based headwater stream (North Carolina, U.S.A.). We used full‐factorial analyses of single‐ and mixed‐species litter from dominant riparian tree species with distinct leaf chemistries [red maple (Acer rubrum), tulip poplar (Liriodendron tulipifera), chestnut oak (Quercus prinus) and rhododendron (Rhododendron maximum)] to test for additivity (single‐species litter presence/absence effects) and non‐additivity (emergent effects of litter species interactions). 3. Significant non‐additive effects of litter mixing on litter mass remaining were explained by species composition, but not richness, and litter‐mixing effects were variable throughout breakdown. Specifically, small differences in observed versus expected litter mass remaining were measured on day 14; whereas observed litter mass remaining in mixed‐species leaf packs was significantly higher on day 70 and lower on day 118 than expected from data for single‐species leaf packs. 4. Litter mixing had non‐additive effects on macroinvertebrate community structure. The number of species in litter mixtures (two to four), but not litter species composition, was a significant predictor of the dominance of particular macroinvertebrates (i.e. indicator taxa) within mixed‐species packs. 5. In addition, the presence/absence of high‐ (L. tulipifera) and low‐quality (R. maximum) litter had additive effects on macroinvertebrate taxon richness, abundance and biomass. The presence of L. tulipifera litter had both positive (synergistic) and negative (antagonistic) effects on invertebrate taxon richness, that varied during breakdown but were not related to litter chemistry. In contrast, the presence/absence of L. tulipifera had a negative relationship with total macroinvertebrate biomass (due to low leaf mass remaining when L. tulipifera was present and higher condensed and hydrolysable tannins associated with leaf packs lacking L. tulipifera). Macroinvertebrate abundance was consistently lower when R. maximum was present, which was partially explained by litter chemistry [e.g., high concentrations of lignin, condensed tannins, hydrolysable tannins and total phenolics and high carbon to nutrient (N and P) ratios]. 6. The bottom‐up effects of litter species diversity on stream macroinvertebrates and litter breakdown are different, which suggests that structural attributes of macroinvertebrate communities may only partially explain the effects of litter‐mixing on organic matter processing in streams. In addition, stream macroinvertebrates colonising decomposing litter are influenced by resource diversity as well as resource availability. Broad‐scale shifts in riparian tree species composition will alter litter inputs to streams, and our results suggest that changes in the diversity and availability of terrestrial litter may alter stream food webs and organic matter processing.     

Using community and population approaches to understand how contemporary and historical factors have shaped species distribution in river ecosystems

Aim To examine how the employment of both community‐ and population‐level approaches can provide a wider view of the importance of contemporary and historical factors on current species distribution. We posit that community ecology should provide more information about contemporary factors, whereas population genetics should provide better information about historical factors. Location Rivers of the western Mediterranean Basin, including four subregions differing in geological history: the Iberian Plate, Transitional, Betic and Rif. Methods For a community‐level approach, Trichoptera richness and community composition were compared between subregions using species accumulation curves and a correspondence analysis. For a population‐level approach, the mtDNA cytochrome C oxidase subunit I (COI) gene of specimens of the Trichoptera midstream‐lowland species Chimarra marginata (L.) was sequenced and analysed using phylogeographical methods. Results The community approach revealed that historical events had more influence on headwater communities than contemporary ecological factors, whereas historical events had negligible influence on midstream‐lowland communities. In midstream‐lowland sites, however, the population approach showed that the genetic structure of C. marginata differed significantly between subregions and revealed patterns of historical gene migration. In terms of species richness, the Rif subregion had the lowest value per basin due to local climatic features and isolation. Main conclusions Both community‐ and population‐level approaches yielded information about the effects of historical factors on species distribution. However, the importance of historical events on current Trichoptera communities depends on the river zonation. Unlike headwater sites, midstream‐lowland sites showed signs of historical events at the population level but not at the community level at the scale used, indicating that both approaches should be employed together in biogeographical studies. Lack of detection of historical events at the community level does not necessarily mean that they are negligible. Most likely, the organizational level used is not appropriate. We also stress the importance of implementing conservation measures for rivers in the western Mediterranean, especially under future scenarios of climate change and human disturbances in the Mediterranean Basin.     

Dispersal rates affect species composition in metacommunities of Sarracenia purpurea inquilines

Dispersal among local communities can have a variety of effects on species composition and diversity at local and regional scales. Local conditions (e.g., resource and predator densities) can have independent effects, as well as interact with dispersal, to alter these patterns. Based on metacommunity models, we predicted that local diversity would show a unimodal relationship with dispersal frequency. We manipulated dispersal frequencies, resource levels, and the presence of predators (mosquito larvae) among communities found in the water-filled leaves of the pitcher plant Sarracenia purpurea. Diversity and abundance of species of the middle trophic level, protozoa and rotifers, were measured. Increased dispersal frequencies significantly increased regional species richness and protozoan abundance while decreasing the variance among local communities. Dispersal frequency interacted with predation at the local community scale to produce patterns of diversity consistent with the model. When predators were absent, we found a unimodal relationship between dispersal frequency and diversity, and when predators were present, there was a flat relationship. Intermediate dispersal frequencies maintained some species in the inquiline communities by offsetting extinction rates. Local community composition and the degree of connectivity between communities are both important for understanding species diversity patterns at local and regional scales.     

Species richness, environmental fluctuations, and temporal change in total community biomass

Theory and empirical results suggest that high biodiversity should often cause lower temporal variability in aggregate community properties such as total community biomass. We assembled microbial communities containing 2 to 8 species of competitors in aquatic microcosms and found that the temporal change in total community biomass was positively but insignificantly associated with diversity in a constant temperature environment. There was no evidence of any trend in variable temperature environments. Three non-exclusive mechanisms might explain the lack of a net stabilising effect of species richness on temporal change. (1) A direct destabilising effect of diversity on population level variances caused some populations to vary more when embedded in more diverse communities. (2) Similar responses of the different species to environmental variability might have limited any insurance effect of increased species richness. (3) Large differences in the population level variability of different species (i.e., unevenness) could weaken the relation between species richness and community level stability. These three mechanisms may outweigh the stabilising effects of increases in total community biomass with diversity, statistical averaging, and slightly more negative covariance in more diverse communities. Our experiment and analyses advocate for further experimental investigations of diversity-variability relations.     

Ectomycorrhizal fungal diversity and saprotrophic fungal diversity are linked to different tree community attributes in a field‐based tree experiment

Exploring the link between above‐ and belowground biodiversity has been a major theme of recent ecological research, due in large part to the increasingly well‐recognized role that soil microorganisms play in driving plant community processes. In this study, we utilized a field‐based tree experiment in Minnesota, USA, to assess the effect of changes in plant species richness and phylogenetic diversity on the richness and composition of both ectomycorrhizal and saprotrophic fungal communities. We found that ectomycorrhizal fungal species richness was significantly positively influenced by increasing plant phylogenetic diversity, while saprotrophic fungal species richness was significantly affected by plant leaf nitrogen content, specific root length and standing biomass. The increasing ectomycorrhizal fungal richness associated with increasing plant phylogenetic diversity was driven by the combined presence of ectomycorrhizal fungal specialists in plots with both gymnosperm and angiosperm hosts. Although the species composition of both the ectomycorrhizal and saprotrophic fungal communities changed significantly in response to changes in plant species composition, the effect was much greater for ectomycorrhizal fungi. In addition, ectomycorrhizal but not saprotrophic fungal species composition was significantly influenced by both plant phylum (angiosperm, gymnosperm, both) and origin (Europe, America, both). The phylum effect was caused by differences in ectomycorrhizal fungal community composition, while the origin effect was attributable to differences in community heterogeneity. Taken together, this study emphasizes that plant‐associated effects on soil fungal communities are largely guild‐specific and provides a mechanistic basis for the positive link between plant phylogenetic diversity and ectomycorrhizal fungal richness.     

Disentangling direct and indirect effects of a legume shrub on its understorey community

Direct and indirect interactions among plants contribute to shape community composition through above‐ and belowground processes. However, we have not disentangled yet the direct and indirect soil and canopy effects of dominants on understorey species. We addressed this issue in a semi‐arid system from southeast Spain dominated by the legume shrub Retama sphaerocarpa. During a year with an exceptionally dry spring, we removed the shrub canopy to quantify aboveground effects and compared removed‐canopy plots to open plots between shrubs to quantify soil effects, both with and without watering. We added a grass removal treatment in order to separate direct from indirect shrub effects and quantified biomass, abundance, richness and composition of the forb functional group. With watering, changes in forb biomass were primarily driven by indirect shrub effects, with contrasting negative soil and positive aboveground indirect effects; changes in forb abundance and composition were more influenced by direct shrub soil effects with contrasting species composition between open and Retama patches. As community composition was different between open and Retama patches the indirect effects of Retama on forb species did not concern forbs from the open community but forbs from Retama patches. Indirect effects are, thus, important at the functional group level rather than at the species level. Without watering, there were no significant interactions. Changes in species richness between treatments were weak and seldom significant. We conclude that shrub effects on understorey forbs are primarily due to their influence on soil properties, directly affecting forb species composition but indirectly affecting the biomass of the forbs of the Retama patches, and only with sufficient water.     

Differential effects of two dominant plant species on community structure and invasibility in an old-field ecosystem

Aims In this study, we examined the effects of Solidago altissima (hereafter Solidago) and two species in the genus Verbesina, Verbesina virginica and Verbesina occidentalis (hereafter Verbesina), on the structure of an old-field plant community and establishment by an invasive plant species, Lespedeza cuneata (hereafter Lespedeza).Methods We removed Solidago, Verbesina and both Solidago and Verbesina from 4-m 2 plots in an intact old-field community during two growing seasons. We then quantified the effects of these removals on richness, evenness, diversity and composition of the subdominant plant community. We also measured the total aboveground biomass and the aboveground biomass of the subdominant community. To assess how these removals affected establishment by Lespedeza, we planted 20 seeds in each plot and tracked seedling emergence and survival for one growing season.Important findings Subdominant community evenness and Shannon diversity were higher in plots from which Solidago and Verbesina were removed relative to control plots. However, there were no effects of dominant species removal on species richness or composition of the subdominant community. Total aboveground biomass was not affected by dominant species removal, suggesting that the community of subdominant species exhibited compensation. In fact, subdominant community biomass was greater when Solidago, but not Verbesina, was removed. Light availability was also greater in plots where Solidago was removed relative to control plots throughout the growing season. In addition, removal of dominant species, in particular Solidago, indirectly reduced the emergence, but not survival, of Lespedeza seedlings by directly promoting subdominant community biomass. Taken together, our results suggest that dominant old-field plant species affect subdominant community structure and indirectly promote establishment by Lespedeza .     

Differential predation drives overyielding of prey species in a patchy environment

In environments characterized by regional heterogeneity among patches, competitor diversity can enhance ecosystem functions such as biomass production. Studies that have addressed the strength of diversity effects in heterogeneous environments have primarily considered a patchy distribution of resources. However, in many systems, top–down effects influence competitor productivity and composition. We use a three‐trophic level consumer–resource model to ask how differential responses to predation influence consumer diversity effects at two scales; 1) in patches with and without predator populations, and 2) at a ‘regional’ scale, consisting of one patch with‐ and one patch without a predator population. At the local scale, the strength and direction of consumer diversity effects depended on the strength of the differential response to predation. Positive or negative influences of consumer richness on equilibrium consumer biomass were the result of a selection effect of diversity. At the regional scale, we observed transgressive overyielding driven by a positive complementarity effect for parameters that define a strong differential response to predation. Given the prevalence of spatially and temporally heterogeneous top–down effects on competitor composition in many ecosystems and trophic levels, we advocate consideration of differential predation as an important step towards incorporating realistic trophic complexity into diversity–function studies.     

The response of a three trophic level soil food web to the identity and diversity of plant species and functional groups

Despite considerable recent interest in how biodiversity may influence ecosystem properties, the issue of how plant diversity and composition may affect multiple trophic levels in soil food webs remains essentially unexplored. We conducted a glasshouse experiment in which three plant species of each of three functional groups (grasses, N‐fixing legumes and forbs) were grown in monoculture and in mixtures of three species (with the three species being in the same or different functional groups) and all nine species. Plant species identity had important effects on the biomasses or population densities of belowground primary consumers (microbial biomass, herbivorous nematodes) and two groups of secondary consumers (microbe‐feeding nematodes and enchytraeids); the third consumer trophic level (predatory nematodes) was marginally not significantly affected at P=0.05. Plant species also influenced the relative importance of the bacterial‐based and fungal‐based energy channels for both the primary and secondary consumer trophic levels. Within‐group diversity of only the soil microflora and herbivorous nematodes (both representing the basal consumer trophic level) were affected by plant species identity. However, community composition within all trophic groupings considered (herbivorous nematodes, microbes, microbe‐feeding nematodes, predatory nematodes) was strongly influenced by what plant species were present. Despite the strong responses of the soil biota to plant species identity, there were few effects of plant species or functional group richness on any of the belowground response variables measured. Further, net primary productivity (NPP) was unaffected by plant diversity. Since some belowground response variables were correlated with NPP across treatments, it is suggested that belowground responses to plant diversity might become more apparent in situations when NPP itself responds to plant diversity. Our results point to plant species identity as having important multitrophic effects on soil food webs, both at the whole trophic group and within‐group levels of resolution, and suggest that differences in plant traits across species may be important in driving the decomposer subsystem.