Linking changes in species composition and biomass in a globally distributed grassland experiment

Global change drivers, such as anthropogenic nutrient inputs, are increasing globally. Nutrient deposition simultaneously alters plant biodiversity, species composition and ecosystem processes like aboveground biomass production. These changes are underpinned by species extinction, colonisation and shifting relative abundance. Here, we use the Price equation to quantify and link the contributions of species that are lost, gained or that persist to change in aboveground biomass in 59 experimental grassland sites. Under ambient (control) conditions, compositional and biomass turnover was high, and losses (i.e. local extinctions) were balanced by gains (i.e. colonisation). Under fertilisation, the decline in species richness resulted from increased species loss and decreases in species gained. Biomass increase under fertilisation resulted mostly from species that persist and to a lesser extent from species gained. Drivers of ecological change can interact relatively independently with diversity, composition and ecosystem processes and functions such as aboveground biomass due to the individual contributions of species lost, gained or persisting.EEA Santa CruzFil: Ladouceur, Emma. German Centre for Integrative Biodiversity Research (iDiv); AlemaniaFil: Ladouceur, Emma. Helmholtz Centre for Environmental Research – UFZ. Department of Physiological Diversity; AlemaniaFil: Ladouceur, Emma. University of Leipzig. Department of Biology; AlemaniaFil: Ladouceur, Emma. Martin Luther University Halle-Wittenberg. Institute of Computer Science; AlemaniaFil: Blowes, Shane A. German Centre for Integrative Biodiversity Research (iDiv); AlemaniaFil: Blowes, Shane A. Martin Luther University Halle-Wittenberg. Institute of Computer Science; AlemaniaFil: Chase, Jonathan M. German Centre for Integrative Biodiversity Research (iDiv); AlemaniaFil: Chase, Jonathan M. Martin Luther University Halle-Wittenberg. Institute of Computer Science; AlemaniaFil: Clark, Adam T. German Centre for Integrative Biodiversity Research (iDiv); AlemaniaFil: Clark, Adam T. Helmholtz Centre for Environmental Research – UFZ. Department of Physiological Diversity; AlemaniaFil: Clark, Adam T. Karl-Franzens University of Graz. Institute of Biology; Austria.Fil: Garbowski, Magda. German Centre for Integrative Biodiversity Research (iDiv); AlemaniaFil: Garbowski, Magda. Helmholtz Centre for Environmental Research – UFZ. Department of Physiological Diversity; AlemaniaFil: Alberti, Juan. Universidad Nacional de Mar del Plata. Instituto de Investigaciones Marinas y Costeras. Laboratorio de Ecología. Mar del Plata; Argentina.Fil: Alberti, Juan. Consejo Nacional de Investigaciones Científicas y Técnicas; Argentina.Fil: Arnillas, Carlos Alberto. University of Toronto. Department of Physical and Environmental Sciences; Canadá.Fil: Bakker, Jonathan D. University of Washington. School of Environmental and Forest Sciences; Estados UnidosFil: Barrio, Isabel C. Agricultural University of Iceland. Faculty of Environmental and Forest Sciences; IslandiaFil: Bharath, Siddharth. Atria University; India.Fil: Peri, Pablo Luis. Instituto Nacional de Tecnología Agropecuaria (INTA). Estación Experimental Agropecuaria Santa Cruz; Argentina.Fil: Peri, Pablo Luis. Universidad Nacional de la Patagonia Austral; Argentina.Fil: Peri, Pablo Luis. Consejo Nacional de Investigaciones Científicas y Técnicas; Argentina.Fil: Harpole, Stanley. German Centre for Integrative Biodiversity Research (iDiv); AlemaniaFil: Harpole, Stanley. Helmholtz Centre for Environmental Research – UFZ. Department of Physiological Diversity; AlemaniaMartin Luther University Halle-Wittenberg. Institute of Computer Science; Alemani

Nutrient enrichment increases invertebrate herbivory and pathogen damage in grasslands

1- Plant damage by invertebrate herbivores and pathogens influences the dynamics of grassland ecosystems, but anthropogenic changes in nitrogen and phosphorus availability can modify these relationships. 2- Using a globally distributed experiment, we describe leaf damage on 153 plant taxa from 27 grasslands worldwide, under ambient conditions and with experimentally elevated nitrogen and phosphorus. 3- Invertebrate damage significantly increased with nitrogen addition, especially in grasses and non-leguminous forbs. Pathogen damage increased with nitrogen in grasses and legumes but not forbs. Effects of phosphorus were generally weaker. Damage was higher in grasslands with more precipitation, but climatic conditions did not change effects of nutrients on leaf damage. On average, invertebrate damage was relatively higher on legumes and pathogen damage was relatively higher on grasses. Community-weighted mean damage reflected these functional group patterns, with no effects of N on community-weighted pathogen damage (due to opposing responses of grasses and forbs) but stronger effects of N on community-weighted invertebrate damage (due to consistent responses of grasses and forbs). 4- Synthesis. As human-induced inputs of nitrogen and phosphorus continue to increase, understanding their impacts on invertebrate and pathogen damage becomes increasingly important. Our results demonstrate that eutrophication frequently increases plant damage and that damage increases with precipitation across a wide array of grasslands. Invertebrate and pathogen damage in grasslands is likely to increase in the future, with potential consequences for plant, invertebrate and pathogen communities, as well as the transfer of energy and nutrients across trophic levels.EEA Santa CruzFil: Ebeling, Anne. University of Jena. Institute of Ecology and Evolution; AlemaniaFil: Strauss, Alex T. University of Minnesota. Department of Ecology, Evolution, and Behavior; Estados UnidosFil: Strauss, Alex T. University of Georgia. Odum School of Ecology; Estados UnidosFil: Adler, Peter B. Utah State University. Department of Wildland Resources and the Ecology Center; Estados UnidosFil: Arnillas, Carlos Alberto. University of Toronto —Scarborough. Department of Physical and Environmental Sciences; CanadáFil: Barrio, Isabel C. Agricultural University of Iceland. Faculty of Environmental and Forest Sciences; IslandiaFil: Biederman, Lori A. Iowa State University. Department of Ecology, Evolution, and Organismal Biology; Estados UnidosFil. Borer, Elizabeth T. University of Minnesota. Department of Ecology, Evolution, and Behavior; Estados UnidosFil: Bugalho, Miguel N. University of Lisbon. Centre for Applied Ecology (CEABN-InBIO). School of Agriculture; Portugal.Fil: Caldeira, Maria C. University of Lisbon. Forest Research Centre. School of Agriculture; Portugal.Fil: Cadotte, Marc W. University of Toronto Scarborough. Department of Biological Sciences; CanadáFil: Peri, Pablo Luis. Instituto Nacional de Tecnología Agropecuaria (INTA). Estación Experimental Agropecuaria Santa Cruz; Argentina.Fil: Peri, Pablo Luis. Universidad Nacional de la Patagonia Austral; Argentina.Fil: Peri, Pablo Luis. Consejo Nacional de Investigaciones Científicas y Técnicas; Argentina.Fil: Blumenthal, Dana M. USDA-ARS, Rangeland Resources & Systems Research Unit; Estados Unido

General Destabilizing Effects of Eutrophication on Grassland Productivity at Multiple Spatial Scales

Eutrophication is a widespread environmental change that usually reduces the stabilizing effect of plant diversity on productivity in local communities. Whether this effect is scale dependent remains to be elucidated. Here, we determine the relationship between plant diversity and temporal stability of productivity for 243 plant communities from 42 grasslands across the globe and quantify the effect of chronic fertilization on these relationships. Unfertilized local communities with more plant species exhibit greater asynchronous dynamics among species in response to natural environmental fluctuations, resulting in greater local stability (alpha stability). Moreover, neighborhood communities that have greater spatial variation in plant species composition within sites (higher beta diversity) have greater spatial asynchrony of productivity among communities, resulting in greater stability at the larger scale (gamma stability). Importantly, fertilization consistently weakens the contribution of plant diversity to both of these stabilizing mechanisms, thus diminishing the positive effect of biodiversity on stability at differing spatial scales. Our findings suggest that preserving grassland functional stability requires conservation of plant diversity within and among ecological communities

Linking changes in species composition and biomass in a globally distributed grassland experiment

Global change drivers, such as anthropogenic nutrient inputs, are increasing globally. Nutrient deposition simultaneously alters plant biodiversity, species composition and ecosystem processes like aboveground biomass production. These changes are underpinned by species extinction, colonisation and shifting relative abundance. Here, we use the Price equation to quantify and link the contributions of species that are lost, gained or that persist to change in aboveground biomass in 59 experimental grassland sites. Under ambient (control) conditions, compositional and biomass turnover was high, and losses (i.e. local extinctions) were balanced by gains (i.e. colonisation). Under fertilisation, the decline in species richness resulted from increased species loss and decreases in species gained. Biomass increase under fertilisation resulted mostly from species that persist and to a lesser extent from species gained. Drivers of ecological change can interact relatively independently with diversity, composition and ecosystem processes and functions such as aboveground biomass due to the individual contributions of species lost, gained or persisting.Fil: Ladouceur, Emma. Martin Luther University Halle-Wittenberg; Alemania. Universitat Leipzig; Alemania. German Centre for Integrative Biodiversity Research (iDiv) Leipzig-Halle-Jena; AlemaniaFil: Blowes, Shane A.. Martin Luther University Halle-Wittenberg; Alemania. German Centre for Integrative Biodiversity Research (iDiv) Leipzig-Halle-Jena; AlemaniaFil: Chase, Jonathan M.. German Centre for Integrative Biodiversity Research (iDiv) Leipzig-Halle-Jena; Alemania. Martin Luther University Halle-Wittenberg; AlemaniaFil: Clark, Adam T.. Martin Luther University Halle-Wittenberg; Alemania. German Centre for Integrative Biodiversity Research (iDiv) Leipzig-Halle-Jena; Alemania. University of Graz; AustriaFil: Garbowski, Magda. German Centre for Integrative Biodiversity Research (iDiv) Leipzig-Halle-Jena; Alemania. Universitat Leipzig; AlemaniaFil: Alberti, Juan. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Mar del Plata. Instituto de Investigaciones Marinas y Costeras. Universidad Nacional de Mar del Plata. Facultad de Ciencias Exactas y Naturales. Instituto de Investigaciones Marinas y Costeras; ArgentinaFil: Arnillas, Carlos Alberto. University of Toronto; CanadáFil: Bakker, Jonathan. University of Washington; Estados UnidosFil: Barrio, Isabel C.. Agricultural University of Iceland; IslandiaFil: Bharath, Siddharth. Atria University; IndiaFil: Borer, Elizabeth. University of Minnesota; Estados UnidosFil: Brudvig, Lars A.. Michigan State University; Estados UnidosFil: Cadotte, Marc W.. University of Toronto; CanadáFil: Chen, Qingqing. Peking University; ChinaFil: Collins, Scott L.. University of New Mexico; Estados UnidosFil: Dickman, Christopher R.. The University Of Sydney; AustraliaFil: Donohue, Ian. Trinity College Dublin; IrlandaFil: Du, Guozhen. Lanzhou University; ChinaFil: Ebeling, Anne. Universitat Jena; AlemaniaFil: Eisenhauer, Nico. Martin Luther University Halle—Wittenberg; Alemania. German Centre For Integrative Biodiversity Research (idiv) Halle-jena-leipzig; AlemaniaFil: Fay, Philip A.. USDA-ARS Grassland Soil and Water Research Lab; Estados UnidosFil: Hagenah, Nicole. University Of Pretoria; SudáfricaFil: Hautier, Yann. University of Utrecht; Países BajosFil: Jentsch, Anke. University of Bayreuth; AlemaniaFil: Jónsdóttir, Ingibjörg S.. University of Iceland; IslandiaFil: Komatsu, Kimberly J.. Smithsonian Environmental Research Center; Estados UnidosFil: MacDougall, Andrew. University of Guelph; CanadáFil: Martina, Jason P.. Texas State University; Estados UnidosFil: Moore, Joslin L.. Arthur Rylah Institute For Environmental Research; Australia. Monash University; AustraliaFil: Morgan, John W.. La Trobe University; AustraliaFil: Peri, Pablo Luis. Instituto Nacional de Tecnología Agropecuaria; Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas; Argentin

Temporal rarity is a better predictor of local extinction risk than spatial rarity

Spatial rarity is often used to predict extinction risk, but rarity can also occur temporally. Perhaps more relevant in the context of global change is whether a species is core to a community (persistent) or transient (intermittently present), with transient species often susceptible to human activities that reduce niche space. Using 5–12 yr of data on 1,447 plant species from 49 grasslands on five continents, we show that local abundance and species persistence under ambient conditions are both effective predictors of local extinction risk following experimental exclusion of grazers or addition of nutrients; persistence was a more powerful predictor than local abundance. While perturbations increased the risk of exclusion for low persistence and abundance species, transient but abundant species were also highly likely to be excluded from a perturbed plot relative to ambient conditions. Moreover, low persistence and low abundance species that were not excluded from perturbed plots tended to have a modest increase in abundance following perturbance. Last, even core species with high abundances had large decreases in persistence and increased losses in perturbed plots, threatening the long-term stability of these grasslands. Our results demonstrate that expanding the concept of rarity to include temporal dynamics, in addition to local abundance, more effectively predicts extinction risk in response to environmental change than either rarity axis predicts alone.Fil: Wilfahrt, Peter A.. University of Minnesota; Estados UnidosFil: Asmus, Ashley L.. University of Minnesota; Estados UnidosFil: Seabloom, Eric. University of Minnesota; Estados UnidosFil: Henning, Jeremiah A.. University of Minnesota; Estados UnidosFil: Adler, Peter. State University of Utah; Estados UnidosFil: Arnillas, Carlos A.. University of Toronto Scarborough; CanadáFil: Bakker, Jonathan. University of Washington; Estados UnidosFil: Biederman, Lori. University of Iowa; Estados UnidosFil: Brudvig, Lars A.. Michigan State University; Estados UnidosFil: Cadotte, Marc W.. University of Toronto Scarborough; CanadáFil: Daleo, Pedro. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Mar del Plata. Instituto de Investigaciones Marinas y Costeras. Universidad Nacional de Mar del Plata. Facultad de Ciencias Exactas y Naturales. Instituto de Investigaciones Marinas y Costeras; ArgentinaFil: Eskelinen, Anu. German Centre for Integrative Biodiversity Research; AlemaniaFil: Firn, Jennifer. University of Queensland; AustraliaFil: Harpole, W. Stanley. German Centre for Integrative Biodiversity Research; Alemania. Helmholtz Centre for Environmental Research; Alemania. Martin Luther University Halle-Wittenberg; AlemaniaFil: Hautier, Yann. Utrecht University; Países BajosFil: Kirkman, Kevin P.. University of KwaZulu-Natal; SudáfricaFil: Komatsu, Kimberly J.. Smithsonian Environmental Research Center; Estados UnidosFil: Laungani, Ramesh. Doane University; Estados UnidosFil: MacDougall, Andrew. University of Guelph; CanadáFil: McCulley, Rebecca L.. University of Kentucky; Estados UnidosFil: Moore, Joslin L.. Monash University; AustraliaFil: Morgan, John W.. La Trobe University; AustraliaFil: Mortensen, Brent. Benedictine College; Estados UnidosFil: Ochoa Hueso, Raul. Universidad de Cádiz; EspañaFil: Ohlert, Timothy. University of New Mexico; Estados UnidosFil: Power, Sally A.. University of Western Sydney; AustraliaFil: Price, Jodi. Charles Sturt University; AustraliaFil: Risch, Anita C.. Swiss Federal Institute for Forest, Snow and Landscape Research; SuizaFil: Schuetz, Martin. Swiss Federal Institute for Forest, Snow and Landscape Research; SuizaFil: Shoemaker, Lauren. University of Wyoming; Estados UnidosFil: Stevens, Carly. Lancaster University; Reino UnidoFil: Strauss, Alexander T.. University of Minnesota; Estados Unidos. University of Georgia; Estados UnidosFil: Tognetti, Pedro Maximiliano. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Instituto de Investigaciones Fisiológicas y Ecológicas Vinculadas a la Agricultura. Universidad de Buenos Aires. Facultad de Agronomía. Instituto de Investigaciones Fisiológicas y Ecológicas Vinculadas a la Agricultura; ArgentinaFil: Virtanen, Risto. University of Oulu; FinlandiaFil: Borer, Elizabeth. University of Minnesota; Estados Unido

Opposing community assembly patterns for dominant and nondominant 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 (\u3c50%) 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

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.Fil: Arnillas, Carlos Alberto. University of Toronto Scarborough; CanadáFil: Borer, Elizabeth. University of Minnesota; Estados UnidosFil: Seabloom, Eric. University of Minnesota; Estados UnidosFil: Alberti, Juan. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Mar del Plata. Instituto de Investigaciones Marinas y Costeras. Universidad Nacional de Mar del Plata. Facultad de Ciencias Exactas y Naturales. Instituto de Investigaciones Marinas y Costeras; ArgentinaFil: Baez, Selene. Escuela Politécnica Nacional; EcuadorFil: Bakker, Jonathan. University of Washington; Estados UnidosFil: Boughton, Elizabeth H.. Archbold Biological Station; Estados UnidosFil: Buckley, Yvonne M.. Trinity College Dublin; IrlandaFil: Bugalho, Miguel Nuno. Universidad de Lisboa; PortugalFil: Donohue, Ian. Trinity College Dublin; IrlandaFil: Dwyer, John. University of Queensland; AustraliaFil: Firn, Jennifer. The University of Queensland; AustraliaFil: Gridzak, Riley. Queens University; CanadáFil: Hagenah, Nicole. University of Pretoria; SudáfricaFil: Hautier, Yann. Utrecht University; Países BajosFil: Helm, Aveliina. University of Tartu; EstoniaFil: Jentsch, Anke. University of Bayreuth; AlemaniaFil: Knops, Johannes M. H.. Xi'an Jiaotong Liverpool University; China. University of Nebraska; Estados UnidosFil: Komatsu, Kimberly J.. Smithsonian Environmental Research Center; Estados UnidosFil: Laanisto, Lauri. Estonian University of Life Sciences; EstoniaFil: Laungani, Ramesh. Poly Prep Country Day School; Estados UnidosFil: McCulley, Rebecca. University of Kentucky; Estados UnidosFil: Moore, Joslin L.. Monash University; AustraliaFil: Morgan, John W.. La Trobe University; AustraliaFil: Peri, Pablo Luis. Universidad Nacional de la Patagonia Austral; Argentina. Instituto Nacional de Tecnología Agropecuaria. Centro Regional Patagonia Sur. Estación Experimental Agropecuaria Santa Cruz. Agencia de Extensión Rural Río Gallegos; ArgentinaFil: Power, Sally A.. University of Western Sydney; AustraliaFil: Price, Jodi. Charles Sturt University; AustraliaFil: Sankaran, Mahesh. National Centre for Biological Sciences; IndiaFil: Schamp, Brandon. Algoma University; CanadáFil: Speziale, Karina Lilian. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Patagonia Norte. Instituto de Investigaciones en Biodiversidad y Medioambiente. Universidad Nacional del Comahue. Centro Regional Universidad Bariloche. Instituto de Investigaciones en Biodiversidad y Medioambiente; ArgentinaFil: Standish, Rachel. Murdoch University; AustraliaFil: Virtanen, Risto. University of Oulu; FinlandiaFil: Cadotte, Marc W.. University of Toronto Scarborough; Canadá. University of Toronto; Canad

Clarifying the effect of biodiversity on productivity in natural ecosystems with longitudinal data and methods for causal inference

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

Negative effects of nitrogen override positive effects of phosphorus on grassland legumes worldwide

Anthropogenic nutrient enrichment is driving global biodiversity decline and modifying ecosystem functions. Theory suggests that plant functional types that fix atmospheric nitrogen have a competitive advantage in nitrogen-poor soils, but lose this advantage with increasing nitrogen supply. By contrast, the addition of phosphorus, potassium, and other nutrients may benefit such species in low-nutrient environments by enhancing their nitrogen-fixing capacity. We present a global-scale experiment confirming these predictions for nitrogen-fixing legumes (Fabaceae) across 45 grasslands on six continents. Nitrogen addition reduced legume cover, richness, and biomass, particularly in nitrogen-poor soils, while cover of non–nitrogen-fixing plants increased. The addition of phosphorous, potassium, and other nutrients enhanced legume abundance, but did not mitigate the negative effects of nitrogen addition. Increasing nitrogen supply thus has the potential to decrease the diversity and abundance of grassland legumes worldwide regardless of the availability of other nutrients, with consequences for biodiversity, food webs, ecosystem resilience, and genetic improvement of protein-rich agricultural plant species.DATA AVAILABILITY : Plant, PAR, climate, and soil nitrogen data have been deposited in the Environmental Data Initiative (EDI) repository (https://portal.edirepository.org/nis/mapbrowse?packageid=edi.838.1) (83). Source data are provided with this paper.This work was generated using data from the Nutrient Network (https://nutnet.org/) experiment, funded at the site scale by individual researchers. Coordination and data management were supported by funding to E.T.B. and E.W.S. from the NSF Research Coordination Network (NSF-DEB-1042132) and Long-Term Ecological Research (NSF-DEB-1234162 to Cedar Creek Long-Term Ecological Research) programs, and the Institute on the Environment (DG-0001-13). We also thank the Minnesota Supercomputer Institute for hosting project data and the Institute of the Environment for hosting Network meetings. P.M.T. was supported by an Argentine Research Council fellowship (Consejo Nacional de Investigaciones Científicas y Técnicas) and the Australian Endeavour Programme.https://www.pnas.orghj2022Mammal Research InstituteZoology and Entomolog

Spatial heterogeneity in species composition constrains plant community responses to herbivory and fertilisation

Environmental change can result in substantial shifts in community composition. The associated immigration and extinction events are likely constrained by the spatial distribution of species. Still, studies on environmental change typically quantify biotic responses at single spatial (time series within a single plot) or temporal (spatial beta diversity at single time points) scales, ignoring their potential interdependence. Here, we use data from a global network of grassland experiments to determine how turnover responses to two major forms of environmental change – fertilisation and herbivore loss – are affected by species pool size and spatial compositional heterogeneity. Fertilisation led to higher rates of local extinction, whereas turnover in herbivore exclusion plots was driven by species replacement. Overall, sites with more spatially heterogeneous composition showed significantly higher rates of annual turnover, independent of species pool size and treatment. Taking into account spatial biodiversity aspects will therefore improve our understanding of consequences of global and anthropogenic change on community dynamics