The functional role of biodiversity in ecosystems: incorporating trophic complexity

Understanding how biodiversity affects functioning of ecosystems requires integrating diversity within trophic levels (horizontal diversity) and across trophic levels (vertical diversity, including food chain length and omnivory). We review theoretical and experimental progress toward this goal. Generally, experiments show that biomass and resource use increase similarly with horizontal diversity of either producers or consumers. Among prey, higher diversity often increases resistance to predation, due to increased probability of including inedible species and reduced efficiency of specialist predators confronted with diverse prey. Among predators, changing diversity can cascade to affect plant biomass, but the strength and sign of this effect depend on the degree of omnivory and prey behaviour. Horizontal and vertical diversity also interact: adding a trophic level can qualitatively change diversity effects at adjacent levels. Multitrophic interactions produce a richer variety of diversity‐functioning relationships than the monotonic changes predicted for single trophic levels. This complexity depends on the degree of consumer dietary generalism, trade‐offs between competitive ability and resistance to predation, intraguild predation and openness to migration. Although complementarity and selection effects occur in both animals and plants, few studies have conclusively documented the mechanisms mediating diversity effects. Understanding how biodiversity affects functioning of complex ecosystems will benefit from integrating theory and experiments with simulations and network‐based approaches

Leaf nutrients, not specific leaf area, are consistent indicators of elevated nutrient inputs

Leaf traits are frequently measured in ecology to provide a ‘common currency’ for predicting how anthropogenic pressures impact ecosystem function. Here, we test whether leaf traits consistently respond to experimental treatments across 27 globally distributed grassland sites across 4 continents. We find that specific leaf area (leaf area per unit mass)—a commonly measured morphological trait inferring shifts between plant growth strategies—did not respond to up to four years of soil nutrient additions. Leaf nitrogen, phosphorus and potassium concentrations increased in response to the addition of each respective soil nutrient. We found few significant changes in leaf traits when vertebrate herbivores were excluded in the short-term. Leaf nitrogen and potassium concentrations were positively correlated with species turnover, suggesting that interspecific trait variation was a significant predictor of leaf nitrogen and potassium, but not of leaf phosphorus concentration. Climatic conditions and pretreatment soil nutrient levels also accounted for significant amounts of variation in the leaf traits measured. Overall, we find that leaf morphological traits, such as specific leaf area, are not appropriate indicators of plant response to anthropogenic perturbations in grasslands

Environmental heterogeneity modulates the effect of biodiversity on the spatial variability of grassland biomass

Plant productivity varies due to environmental heterogeneity, and theory suggests that plant diversity can reduce this variation. While there is strong evidence of diversity effects on temporal variability of productivity, whether this mechanism extends to variability across space remains elusive. Here we determine the relationship between plant diversity and spatial variability of productivity in 83 grasslands, and quantify the effect of experimentally increased spatial heterogeneity in environmental conditions on this relationship. We found that communities with higher plant species richness (alpha and gamma diversity) have lower spatial variability of productivity as reduced abundance of some species can be compensated for by increased abundance of other species. In contrast, high species dissimilarity among local communities (beta diversity) is positively associated with spatial variability of productivity, suggesting that changes in species composition can scale up to affect productivity. Experimentally increased spatial environmental heterogeneity weakens the effect of plant alpha and gamma diversity, and reveals that beta diversity can simultaneously decrease and increase spatial variability of productivity. Our findings unveil the generality of the diversity-stability theory across space, and suggest that reduced local diversity and biotic homogenization can affect the spatial reliability of key ecosystem functions

Soil net nitrogen mineralisation across global grasslands

Soil nitrogen mineralisation (Nmin), the conversion of organic into inorganic N, is important for productivity and nutrient cycling. The balance between mineralisation and immobilisation (net Nmin) varies with soil properties and climate. However, because most global-scale assessments of net Nmin are laboratory-based, its regulation under field-conditions and implications for real-world soil functioning remain uncertain. Here, we explore the drivers of realised (field) and potential (laboratory) soil net Nmin across 30 grasslands worldwide. We find that realised Nmin is largely explained by temperature of the wettest quarter, microbial biomass, clay content and bulk density. Potential Nmin only weakly correlates with realised Nmin, but contributes to explain realised net Nmin when combined with soil and climatic variables. We provide novel insights of global realised soil net Nmin and show that potential soil net Nmin data available in the literature could be parameterised with soil and climate data to better predict realised NNational Science Foundation Research Coordination Network; Long-Term Ecological Research; Institute on the Environment at the University of Minnesota.http://www.nature.com/ncommspm2020Mammal Research InstituteZoology and Entomolog

Impacts of nutrient addition on soil carbon and nitrogen stoichiometry and stability in globally-distributed grasslands

Global changes will modify future nutrient availability with implications for grassland biogeochemistry. Soil organic matter (SOM) is central to grasslands for both provision of nutrients and climate mitigation through carbon (C) storage. While we know that C and nitrogen (N) in SOM can be influenced by greater nutrient availability, we lack understanding of nutrient effects on C and N coupling and stability in soil. Different SOM fractions have different functional relevance and mean residence times, i.e., mineral-associated organic matter (MAOM) has a higher mean residence time than particulate organic matter (POM). By separating effects of nutrient supply on the different SOM fractions, we can better evaluate changes in soil C and N coupling and stability and associated mechanisms. To this end, we studied responses of C and N ratios and distributions across POM and MAOM to 6–10 years of N, phosphorus (P), potassium and micronutrients (K+µ), and combined NPK+µ additions at 11 grassland sites spanning 3 continents and globally relevant environmental gradients in climate, plant growth, soil texture, and nutrient availability. We found addition of N and NPK+µ generally reduced C:N in MAOM and POM. However, at low fertility and at warm, sandy sites, nutrient addition promoted higher MAOM and POM C:N, respectively. Addition of NPK+µ also promoted C storage in POM relative to MAOM, and this was consistent across sites. Our results suggest that addition of macro- and micronutrients consistently decrease SOM stabilization, whereas responses of soil C:N stoichiometry were contingent on SOM fraction and environmental conditions

Spatial turnover of multiple ecosystem functions is more associated with plant than soil microbial β-diversity

Biodiversity—both above- and belowground—influences multiple functions in terrestrial ecosystems. Yet, it is unclear whether differences in above- and belowground species composition (β-diversity) are associated with differences in multiple ecosystem functions (e.g., spatial turnover in ecosystem function). Here, we partitioned the contributions of above- and belowground β-diversity and abiotic factors (geographic distance, differences in environments) on the spatial turnover of multiple grassland ecosystem functions. We compiled a dataset of plant and soil microbial communities and six indicators of grassland ecosystem functions (i.e., plant aboveground live biomass, plant nitrogen [N], plant phosphorus [P], root biomass, soil total N, and soil extractable P) from 18 grassland sites on four continents contributing to the Nutrient Network experiment. We used Mantel tests and structural equation models to disentangle the relationship between above- and belowground β-diversity and spatial turnover in grassland ecosystem functions. We found that the effects of abiotic factors on the spatial turnover of ecosystem functions were largely indirect through their influences on above- and belowground β-diversity, and that spatial turnover of ecosystem function was more strongly associated with plant β-diversity than with soil microbial β-diversity. These results indicate that changes in above- and belowground species composition are one mechanism that interacts with environmental change to determine variability in multiple ecosystem functions across spatial scales. As grasslands face global threats from shrub encroachment, conversion to agriculture, or are lost to development, the functions and services they provide will more strongly converge with increased aboveground community homogenization than with soil microbial community homogenization

Opposing community assembly patterns for dominant and jonnondominant plant species in herbaceous ecosystems globally

Biotic and abiotic factors interact with dominant plants—the locally most frequent or with the largest coverage—and nondominant plants differently, partially because dominant plants modify the environment where nondominant plants grow. For instance, if dominant plants compete strongly, they will deplete most resources, forcing nondominant plants into a narrower niche space. Conversely, if dominant plants are constrained by the environment, they might not exhaust available resources but instead may ameliorate environmental stressors that usually limit nondominants. Hence, the nature of interactions among nondominant species could be modified by dominant species. Furthermore, these differences could translate into a disparity in the phylogenetic relatedness among dominants compared to the relatedness among nondominants. By estimating phylogenetic dispersion in 78 grasslands across five continents, we found that dominant species were clustered (e.g., co-dominant grasses), suggesting dominant species are likely organized by environmental filtering, and that nondominant species were either randomly assembled or overdispersed. Traits showed similar trends for those sites (<50%) with sufficient trait data. Furthermore, several lineages scattered in the phylogeny had more nondominant species than expected at random, suggesting that traits common in nondominants are phylogenetically conserved and have evolved multiple times. We also explored environmental drivers of the dominant/nondominant disparity. We found different assembly patterns for dominants and nondominants, consistent with asymmetries in assembly mechanisms. Among the different postulated mechanisms, our results suggest two complementary hypotheses seldom explored: (1) Nondominant species include lineages adapted to thrive in the environment generated by dominant species. (2) Even when dominant species reduce resources to nondominant ones, dominant species could have a stronger positive effect on some nondominants by ameliorating environmental stressors affecting them, than by depleting resources and increasing the environmental stress to those nondominants. These results show that the dominant/nondominant asymmetry has ecological and evolutionary consequences fundamental to understand plant communities