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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.

This dataset contains all data on which the following publication below is based. Paper Citation: Risch Anita C., Zimmermann, Stefan, Schütz, Martin, Borer, Elizabeth T., Broadbent, Arthur A.D., Caldeira, Maria C., Davies, Kendi F., Eisenhauer, Nico, Eskelinen, Anu, Fay, Philip A., Hagedorn, Frank, Knops, Johannes M.H., Lembrechts, Jonas, J., MacDougall, Andrew S., McCulley, Rebecca L., Melbourne, Brett A., Moore, Joslin L., Power, Sally A., Seabloom, Eric W., Silveira, Maria L., Virtanen, Risto, Yahdjian, Laura, Ochoa-Hueso, Raul (accepted). Drivers of the microbial metabolic quotient across global grasslands. Global Ecology and Biogeography Please cite this paper together with the citation for the datafile. The microbial metabolic quotient (MMQ; mg CO2-C mg MBC-1 h-1), defined as the amount of microbial CO2 respired (MR; mg CO2-C kg soil-1 h-1) per unit of microbial biomass C (MBC; mg C kg soil-1), is a key parameter for understanding the microbial regulation of the carbon (C) cycle, including soil C sequestration. Here, we experimentally tested hypotheses about the individual and interactive effects of multiple nutrient addition (NPK+micronutrients) and herbivore exclusion on MR, MBC, and MMQ across 23 sites (5 continents). Our sites encompassed a wide range of edaphoclimatic conditions, thus we assessed which edaphoclimatic variables affected MMQ the most and how they interacted with our treatments. Soils were collected in plots with established experimental treatments. MR was assessed in a five-week laboratory incubation without glucose addition, MBC via substrate-induced respiration. MMQ was calculated as MR/MBC and corrected for soil temperatures (MMQsoil). Using LMMs and SEMs, we analysed how edaphoclimatic characteristics and treatments interactively affected MMQsoil. MMQsoil was higher in locations with higher mean annual temperature, lower water holding capacity, and soil organic C concentration, but did not respond to our treatments across sites as neither MR nor MBC changed. We attributed this relative homeostasis to our treatments to the modulating influence of edaphoclimatic variables. For example, herbivore exclusion, regardless of fertilization, led to greater MMQsoil only at sites with lower soil organic C (-1.7%). Our results pinpoint the main variables related to MMQsoil across grasslands and emphasize the importance of the local edaphoclimatic conditions in controlling the response of the C cycle to anthropogenic stressors. By testing hypotheses about MMQsoil across global edaphoclimatic gradients, this work also helps to align the conflicting results of prior studies.

Declines in grassland diversity in response to nutrient addition are a general consequence of global change. This decline in species richness may be driven by multiple underlying processes operating at different time-scales. Nutrient addition can reduce diversity by enhancing the rate of local extinction via competitive exclusion, or by reducing the rate of colonization by constraining the pool of species able to colonize under new conditions. Partitioning net change into extinction and colonization rates will better delineate the long-term effect of global change in grasslands. We synthesized changes in richness in response to experimental fertilization with nitrogen, phosphorus and potassium with micronutrients across 30 grasslands. We quantified changes in local richness, colonization, and extinction over 8–10 years of nutrient addition, and compared these rates against control conditions to isolate the effect of nutrient addition from background dynamics. Total richness at steady state in the control plots was the sum of equal, relatively high rates of local colonization and extinction. On aggregate, 30%–35% of initial species were lost and the same proportion of new species were gained at least once over a decade. Absolute turnover increased with site-level richness but was proportionately greater at lower-richness sites relative to starting richness. Loss of total richness with nutrient addition, especially N in combination with P or K, was driven by enhanced rates of extinction with a smaller contribution from reduced colonization. Enhanced extinction and reduced colonization were disproportionately among native species, perennials, and forbs. Reduced colonization plateaued after the first few (

Causal effects of biodiversity on ecosystem functions can be estimated using experimental or observational designs — designs that pose a tradeoff between drawing credible causal inferences from correlations and drawing generalizable inferences. Here, we develop a design that reduces this tradeoff and revisits the question of how plant species diversity affects productivity. Our design leverages longitudinal data from 43 grasslands in 11 countries and approaches borrowed from fields outside of ecology to draw causal inferences from observational data. Contrary to many prior studies, we estimate that increases in plot-level species richness caused productivity to decline: a 10% increase in richness decreased productivity by 2.4%, 95% CI [−4.1, −0.74]. This contradiction stems from two sources. First, prior observational studies incompletely control for confounding factors. Second, most experiments plant fewer rare and non-native species than exist in nature. Although increases in native, dominant species increased productivity, increases in rare and non-native species decreased productivity, making the average effect negative in our study. By reducing the tradeoff between experimental and observational designs, our study demonstrates how observational studies can complement prior ecological experiments and inform future ones.

Ecological models predict that the effects of mammalian herbivore exclusion on plant diversity depend on resource availability and plant exposure to ungulate grazing over evolutionary time. Using an experiment replicated in 57 grasslands on six continents, with contrasting evolutionary history of grazing, we tested how resources (mean annual precipitation and soil nutrients) determine herbivore exclusion effects on plant diversity, richness and evenness. Here we show that at sites with a long history of ungulate grazing, herbivore exclusion reduced plant diversity by reducing both richness and evenness and the responses of richness and diversity to herbivore exclusion decreased with mean annual precipitation. At sites with a short history of grazing, the effects of herbivore exclusion were not related to precipitation but differed for native and exotic plant richness. Thus, plant species’ evolutionary history of grazing continues to shape the response of the world’s grasslands to changing mammalian herbivory.

In the version of this Article originally published, there were unit conversion errors in the dataset when calculating specific leaf area (SLA) values at 10 of the 27 sites; errors were made when converting SLA from cm 2 g –1 to mm 2 g –1 and from mm 2 mg –1 to mm 2 g –1. This resulted in two incorrect data points: Phleum pratense in plot 27 at the Frue.ch site (SLA >300,000 mm 2 g –1) and Poa secunda in plot 31 at the shps.us site (SLA >60 mm 2 g –1). These two values were changed to ‘NA’ in the dataset, and therefore some reported estimates were changed, resulting in changes to the data points in Fig. 2a, the top bar of Fig. 3 and several values in the text.

Human activities are enriching many of Earth's ecosystems with biologically limiting mineral nutrients such as nitrogen (N) and phosphorus (P). In grasslands, this enrichment generally reduces plant diversity and increases productivity. The widely demonstrated positive effect of diversity on productivity suggests a potential negative feedback, whereby nutrient-induced declines in diversity reduce the initial gains in productivity arising from nutrient enrichment. In addition, plant productivity and diversity can be inhibited by accumulations of dead biomass, which may be altered by nutrient enrichment. Over longer time frames, nutrient addition may increase soil fertility by increasing soil organic matter and nutrient pools. We examined the effects of 5-11 yr of nutrient addition at 47 grasslands in 12 countries. Nutrient enrichment increased aboveground live biomass and reduced plant diversity at nearly all sites, and these effects became stronger over time. We did not find evidence that nutrient-induced losses of diversity reduced the positive effects of nutrients on biomass; however, nutrient effects on live biomass increased more slowly at sites where litter was also increasing, regardless of plant diversity. This work suggests that short-term experiments may underestimate the long-term nutrient enrichment effects on global grassland ecosystems.

Aim: Climate variability threatens to destabilize production in many ecosystems. Asynchronous species dynamics may buffer against such variability when a decrease in performance by some species is offset by an increase in performance of others. However, high climatic variability can eliminate species through stochastic extinctions or cause similar stress responses among species that reduce buffering. Local conditions, such as soil nutrients, can also alter production stability directly or by influencing asynchrony. We test these hypotheses using a globally distributed sampling experiment. Location: Grasslands in North America, Europe and Australia. Time period: Annual surveys over 5 year intervals occurring between 2007 and 2014. Major taxa studied: Herbaceous plants. Methods: We sampled annually the per species cover and aboveground community biomass [net primary productivity (NPP)], plus soil chemical properties, in 29 grasslands. We tested how soil conditions, combined with variability in precipitation and temperature, affect species richness, asynchrony and temporal stability of primary productivity. We used bivariate relationships and structural equation modelling to examine proximate and ultimate relationships. Results: Climate variability strongly predicted asynchrony, whereas NPP stability was more related to soil conditions. Species richness was structured by both climate variability and soils and, in turn, increased asynchrony. Variability in temperature and precipitation caused a unimodal asynchrony response, with asynchrony being lowest at low and high climate variability. Climate impacted stability indirectly, through its effect on asynchrony, with stability increasing at higher asynchrony owing to lower inter-annual variability in NPP. Soil conditions had no detectable effect on asynchrony but increased stability by increasing the mean NPP, especially when soil organic matter was high. Main conclusions: We found globally consistent evidence that climate modulates species asynchrony but that the direct effect on stability is low relative to local soil conditions. Nonetheless, our observed unimodal responses to variability in temperature and precipitation suggest asynchrony thresholds, beyond which there are detectable destabilizing impacts of climate on primary productivity.

Anthropogenic activities are increasing nutrient inputs to ecosystems worldwide, with consequences for global carbon and nutrient cycles. Recent meta-analyses show that aboveground primary production is often co-limited by multiple nutrients; however, little is known about how root production responds to changes in nutrient availability. At twenty-nine grassland sites on four continents, we quantified shallow root biomass responses to nitrogen (N), phosphorus (P) and potassium plus micronutrient enrichment and compared below- and aboveground responses. We hypothesized that optimal allocation theory would predict context dependence in root biomass responses to nutrient enrichment, given variation among sites in the resources limiting to plant growth (specifically light versus nutrients). Consistent with the predictions of optimal allocation theory, the proportion of total biomass belowground declined with N or P addition, due to increased biomass aboveground (for N and P) and decreased biomass belowground (N, particularly in sites with low canopy light penetration). Absolute root biomass increased with N addition where light was abundant at the soil surface, but declined in sites where the grassland canopy intercepted a large proportion of incoming light. These results demonstrate that belowground responses to changes in resource supply can differ strongly from aboveground responses, which could significantly modify predictions of future rates of nutrient cycling and carbon sequestration. Our results also highlight how optimal allocation theory developed for individual plants may help predict belowground biomass responses to nutrient enrichment at the ecosystem scale across wide climatic and environmental gradients.

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. Fil: Firn, Jennifer. Queensland University of Technology; Australia Fil: McGree, James M.. Queensland University of Technology; Australia Fil: Harvey, Eric. University of Montreal; Canadá Fil: Flores Moreno, Habacuc. University of Minnesota; Estados Unidos Fil: Schütz, Martin. Swiss Federal Institute for Forest, Snow and Landscape Research; Suiza Fil: Buckley, Yvonne M.. Trinity College; Irlanda Fil: Borer, Elizabeth T.. University of Minnesota; Estados Unidos Fil: Seabloom, Eric W.. University of Minnesota; Estados Unidos Fil: La Pierre, Kimberly J.. Smithsonian Environmental Research Center; Estados Unidos Fil: MacDougall, Andrew M.. University of Guelph; Canadá Fil: Prober, Suzanne M.. No especifíca; Fil: Stevens, Carly J.. Universidad de Lancaster; Reino Unido Fil: Sullivan, Lauren L.. University of Minnesota; Estados Unidos Fil: Porter, Erica. Queensland University of Technology; Australia Fil: Ladouceur, Emma. Helmholtz Center for Environmental Research; Alemania Fil: Allen, Charlotte. Queensland University of Technology; Australia Fil: Moromizato, Karine H.. Queensland University of Technology; Australia Fil: Morgan, John W.. La Trobe University; Australia Fil: Harpole, W. Stanley. Helmholtz Center for Environmental Research; Alemania. Universidad de Halle-Wittenberg ; Alemania Fil: Hautier, Yann. University of Utrecht; Países Bajos Fil: Eisenhauer, Nico. Universitat Leipzig; Alemania Fil: Wright, Justin P.. University of Duke; Estados Unidos Fil: Adler, Peter B.. State University of Utah; Estados Unidos Fil: Arnillas, Carlos Alberto. University of Toronto; Canadá Fil: Bakker, Jonathan D.. University of Washington; Estados Unidos Fil: Biederman, Lori. University of Iowa; Estados Unidos Fil: Broadbent, Arthur A. D.. University of Manchester; Reino Unido Fil: Brown, Cynthia S.. State University of Colorado at Boulder; Estados Unidos Fil: Bugalho, Miguel N.. Universidade Nova de Lisboa; Portugal Fil: Caldeira, Maria C.. Universidade Nova de Lisboa; Portugal Fil: Cleland, Elsa E.. University of California; Estados Unidos Fil: Ebeling, Anne. Universitat Jena; Alemania Fil: Fay, Philip A.. No especifíca; Fil: Hagenah, Nicole. Universidad de Pretoria; Sudáfrica Fil: Kleinhesselink, Andrew R.. University of California at Los Angeles; Estados Unidos Fil: Mitchell, Rachel. Arizona State University; Estados Unidos Fil: Moore, Joslin L.. Monash University; Australia Fil: Nogueira, Carla. Universidade Nova de Lisboa; Portugal Fil: Peri, Pablo Luis. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro de Investigaciones y Transferencia de Santa Cruz. Universidad Tecnológica Nacional. Facultad Regional Santa Cruz. Centro de Investigaciones y Transferencia de Santa Cruz. Universidad Nacional de la Patagonia Austral. Centro de Investigaciones y Transferencia de Santa Cruz; Argentina Fil: Roscher, Christiane. Helmholtz Center for Environmental Research; Alemania Fil: Smith, Melinda D.. State University of Colorado at Boulder; Estados Unidos Fil: Wragg, Peter D.. University of Minnesota; Estados Unidos Fil: Risch, Anita C.. No especifíca