Drivers of the microbial metabolic quotient across global grasslands

Aim: The microbial metabolic quotient (MMQ; mg CO2-C/mg MBC/h), defined as the amount of microbial CO2 respired (MR; mg CO2-C/kg soil/h) per unit of microbial biomass C (MBC; mg C/kg soil), 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 (nitrogen + phosphorus + potassium + micronutrients) and herbivore exclusion on MR, MBC and MMQ across 23 sites (five 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. Location: Australia, Asia, Europe, North/South America. Time period: 2015–2016. Major taxa: Soil microbes. Methods: Soils were collected from plots with established experimental treatments. MR was assessed in a 5-week laboratory incubation without glucose addition, MBC via substrate-induced respiration. MMQ was calculated as MR/MBC and corrected for soil temperatures (MMQsoil). Using linear mixed effects models (LMMs) and structural equation models (SEMs), we analysed how edaphoclimatic characteristics and treatments interactively affected MMQsoil. Results: MMQsoil was higher in locations with higher mean annual temperature, lower water holding capacity and lower 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%). Main conclusions: 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

Nitrogen but not phosphorus addition affects symbiotic N2 fixation by legumes in natural and semi‑natural grasslands located on four continents

The amount of nitrogen (N) derived from symbiotic N2 fixation by legumes in grasslands might be affected by anthropogenic N and phosphorus (P) inputs, but the underlying mechanisms are not known. Methods We evaluated symbiotic N2 fixation in 17 natural and semi-natural grasslands on four continents that are subjected to the same full-factorial N and P addition experiment, using the 15N natural abundance method. Results N as well as combined N and P (NP) addition reduced aboveground legume biomass by 65% and 45%, respectively, compared to the control, whereas P addition had no significant impact. Addition of N and/or P had no significant effect on the symbiotic N2 fixation per unit legume biomass. In consequence, the amount of N fixed annually per grassland area was less than half in the N addition treatments compared to control and P addition, irrespective of whether the dominant legumes were annuals or perennials. Conclusion Our results reveal that N addition mainly impacts symbiotic N2 fixation via reduced biomass of legumes rather than changes in N2 fixation per unit legume biomass. The results show that soil N enrichment by anthropogenic activities significantly reduces N 2 fixation in grasslands, and these effects cannot be reversed by additional P amendment.EEA Santa CruzFil: Vázquez, Eduardo. University of Bayreuth. Department of Soil Ecology. Bayreuth Center of Ecology and Environmental Research (BayCEER); AlemaniaFil: Vázquez, Eduardo. Swedish University of Agricultural Sciences. Department of Soil and Environment; SueciaFil: Schleuss, Per‑Marten. University of Bayreuth. Department of Soil Ecology. Bayreuth Center of Ecology and Environmental Research (BayCEER); AlemaniaFil: Borer, Elizabeth T. University of Minnesota. Department of Ecology, Evolution, and Behavior; Estados UnidosFil: Bugalho, Miguel N. University of Lisbon. Centre for Applied Ecology “Prof. Baeta Neves” (CEABN-InBIO). School of Agriculture; Portugal.Fil: Caldeira, Maria. C. University of Lisbon. Forest Research Centre. School of Agriculture; Portugal.Fil: Eisenhauer, Nico. German Centre for Integrative Biodiversity Research; AlemaniaFil: Eisenhauer, Nico. Leipzig University. Institute of Biology; AlemaniaFil: Eskelinen, Anu. German Centre for Integrative Biodiversity Research; AlemaniaFil: Eskelinen, Anu. Physiological Diversity, Helmholtz Centrefor Environmental Research; AlemaniaFil: Eskelinen, Anu. University of Oulu. Ecology & Genetics; FinlandiaFil: Fay, Philip A. Grassland Soil and Water Research Laboratory (USDA-ARS); Estados UnidosFil: Haider, Sylvia. German Centre for Integrative Biodiversity Research; AlemaniaFil: Haider, Sylvia. Martin Luther University. Institute of Biology. Geobotany and Botanical Garden; AlemaniaFil: Jentsch, Anke. University of Bayreuth. Department of Soil Ecology. Bayreuth Center of Ecology and Environmental Research (BayCEER); AlemaniaFil: Kirkman, Kevin P. University of KwaZulu-Natal. School of Life Sciences; SudáfricaFil: McCulley, Rebecca L. University of Kentucky. Department of Plant and Soil Sciences; Estados UnidosFil: 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: Price, Jodi. Charles Sturt University. Institute for Land, Water and Society; Australia.Fil: Richards, Anna E. CSIRO Land and Water. Northern Territory; Australia.Fil: Risch, Anita C. Swiss Federal Institute for Forest, Snow and Landscape Research WSL; SuizaFil: Roscher, Christiane. German Centre for Integrative Biodiversity Research; AlemaniaFil: Roscher, Christiane. Physiological Diversity, Helmholtz Centre for Environmental Research; AlemaniaFil: Schütz, Martin. Swiss Federal Institute for Forest, Snow and Landscape Research WSL; SuizaFil: Seabloom, Eric William. University of Minnesota. Dept. of Ecology, Evolution, and Behavior; Estados UnidosFil: Standish, Rachel J. Murdoch University. Harry Butler Institute; Australia.Fil: Stevens, Carly J. Lancaster University. Lancaster Environment Centre; Reino UnidoFil: Tedder, Michelle J. University of KwaZulu-Natal. School of Life Sciences; SudáfricaFil: Virtanen, Risto. University of Oulu. Ecology & Genetics; Finlandia.Fil: Spohn, Marie. University of Bayreuth. Department of Soil Ecology. Bayreuth Center of Ecology and Environmental Research (BayCEER); AlemaniaFil: Spohn, Marie. Swedish University of Agricultural Sciences. Department of Soil and Environment; Sueci

Nitrogen but not phosphorus addition affects symbiotic N-2 fixation by legumes in natural and semi-natural grasslands located on four continents

Background and aims: The amount of nitrogen (N) derived from symbiotic N-2 fixation by legumes in grasslands might be affected by anthropogenic N and phosphorus (P) inputs, but the underlying mechanisms are not known.Methods: We evaluated symbiotic N-2 fixation in 17 natural and semi-natural grasslands on four continents that are subjected to the same full-factorial N and P addition experiment, using the N-15 natural abundance method.Results: N as well as combined N and P (NP) addition reduced aboveground legume biomass by 65% and 45%, respectively, compared to the control, whereas P addition had no significant impact. Addition of N and/or P had no significant effect on the symbiotic N-2 fixation per unit legume biomass. In consequence, the amount of N fixed annually per grassland area was less than half in the N addition treatments compared to control and P addition, irrespective of whether the dominant legumes were annuals or perennials.Conclusion: Our results reveal that N addition mainly impacts symbiotic N-2 fixation via reduced biomass of legumes rather than changes in N-2 fixation per unit legume biomass. The results show that soil N enrichment by anthropogenic activities significantly reduces N-2 fixation in grasslands, and these effects cannot be reversed by additional P amendment

Multidimensional responses of grassland stability to eutrophication

Eutrophication usually impacts grassland biodiversity, community composition, and biomass production, but its impact on the stability of these community aspects is unclear. One challenge is that stability has many facets that can be tightly correlated (low dimensionality) or highly disparate (high dimensionality). Using standardized experiments in 55 grassland sites from a globally distributed experiment (NutNet), we quantify the effects of nutrient addition on five facets of stability (temporal invariability, resistance during dry and wet growing seasons, recovery after dry and wet growing seasons), measured on three community aspects (aboveground biomass, community composition, and species richness). Nutrient addition reduces the temporal invariability and resistance of species richness and community composition during dry and wet growing seasons, but does not affect those of biomass. Different stability measures are largely uncorrelated under both ambient and eutrophic conditions, indicating consistently high dimensionality. Harnessing the dimensionality of ecological stability provides insights for predicting grassland responses to global environmental change

Local Loss and Spatial Homogenization of Plant Diversity Reduce Ecosystem Multifunctionality

Biodiversity is declining in many local communities while also becoming increasingly homogenized across space. Experiments show that local plant species loss reduces ecosystem functioning and services, but the role of spatial homogenization of community composition and the potential interaction between diversity at different scales in maintaining ecosystem functioning remains unclear, especially when many functions are considered (ecosystem multifunctionality). We present an analysis of eight ecosystem functions measured in 65 grasslands worldwide. We find that more diverse grasslands—those with both species-rich local communities (α-diversity) and large compositional differences among localities (β-diversity)—had higher levels of multifunctionality. Moreover, α- and β-diversity synergistically affected multifunctionality, with higher levels of diversity at one scale amplifying the contribution to ecological functions at the other scale. The identity of species influencing ecosystem functioning differed among functions and across local communities, explaining why more diverse grasslands maintained greater functionality when more functions and localities were considered. These results were robust to variation in environmental drivers. Our findings reveal that plant diversity, at both local and landscape scales, contributes to the maintenance of multiple ecosystem services provided by grasslands. Preserving ecosystem functioning therefore requires conservation of biodiversity both within and among ecological communities

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 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

Long-term N-addition alters the community structure of functionally important N-cycling soil microorganisms across global grasslands

Anthropogenic nitrogen (N) input is known to alter the soil microbiome, but how N enrichment influences the abundance, alpha-diversity and community structure of N-cycling functional microbial communities in grasslands remains poorly understood. Here, we collected soils from plant communities subjected to up to 9 years of annual N-addition (10 g N m−2 per year using urea as a N-source) and from unfertilized plots (control) in 30 grasslands worldwide spanning a large range of climatic and soil conditions. We focused on three key microbial groups responsible for two essential processes of the global N cycle: N2 fixation (soil diazotrophs) and nitrification (AOA: ammonia-oxidizing archaea and AOB: ammonia-oxidizing bacteria). We targeted soil diazotrophs, AOA and AOB using Illumina MiSeq sequencing and measured the abundance (gene copy numbers) using quantitative PCR. N-addition shifted the structure of the diazotrophic communities, although their alpha-diversity and abundance were not affected. AOA and AOB responded differently to N-addition. The abundance and alpha-diversity of AOB increased, and their community structure shifted with N-addition. In contrast, AOA were not affected by N-addition. AOA abundance outnumbered AOB in control plots under conditions of low N availability, whereas N-addition favoured copiotrophic AOB. Overall, N-addition showed a low impact on soil diazotrophs and AOA while effects for AOB communities were considerable. These results reveal that long-term N-addition has important ecological implications for key microbial groups involved in two critical soil N-cycling processes. Increased AOB abundance and community shifts following N-addition may change soil N-cycling, as larger population sizes may promote higher rates of ammonia oxidation and subsequently increase N loss via gaseous and soil N-leaching. These findings bring us a step closer to predicting the responses and feedbacks of microbial-mediated N-cycling processes to long-term anthropogenic N-addition in grasslands

Global change effects on plant communities are magnified by time and the number of global change factors imposed

Global change drivers (GCDs) are expected to alter community structure and consequently, the services that ecosystems provide. Yet, few experimental investigations have examined effects of GCDs on plant community structure across multiple ecosystem types, and those that do exist present conflicting patterns. In an unprecedented global synthesis of over 100 experiments that manipulated factors linked to GCDs, we show that herbaceous plant community responses depend on experimental manipulation length and number of factors manipulated. We found that plant communities are fairly resistant to experimentally manipulated GCDs in the short term (<10 y). In contrast, long-term (≥10 y) experiments show increasing community divergence of treatments from control conditions. Surprisingly, these community responses occurred with similar frequency across the GCD types manipulated in our database. However, community responses were more common when 3 or more GCDs were simultaneously manipulated, suggesting the emergence of additive or synergistic effects of multiple drivers, particularly over long time periods. In half of the cases, GCD manipulations caused a difference in community composition without a corresponding species richness difference, indicating that species reordering or replacement is an important mechanism of community responses to GCDs and should be given greater consideration when examining consequences of GCDs for the biodiversity–ecosystem function relationship. Human activities are currently driving unparalleled global changes worldwide. Our analyses provide the most comprehensive evidence to date that these human activities may have widespread impacts on plant community composition globally, which will increase in frequency over time and be greater in areas where communities face multiple GCDs simultaneously

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 (<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.National Science Foundation; Natural Sciences and Engineering Research Council of Canada; Institute on the Environment, University of Minnesota and Portuguese Science Foundation.http://www.ecolevol.orghj2022Mammal Research InstituteZoology and Entomolog