Nitrogen addition strengthens the stabilizing effect of biodiversity on productivity by increasing plant trait diversity and species asynchrony in the artificial grassland communities

BACKGROUND AND AIMS: Nitrogen (N) enrichment usually weakens the stabilizing effect of biodiversity on productivity. However, previous studies focused on plant species richness and thus largely ignored the potential contributions of plant functional traits to stability, even though evidence is increasing that functional traits are stronger predictors than species richness of ecosystem functions. METHODS: We conducted a common garden experiment manipulating plant species richness and N addition levels to quantify effects of N addition on relations between species richness and functional trait identity and diversity underpinning the 'fast-slow' economics spectrum and community stability. RESULTS: Nitrogen addition had a minor effect on community stability but increased the positive effects of species richness on community stability. Increasing community stability was found in the species-rich communities dominated by fast species due to substantially increasing temporal mean productivity relative to its standard deviation. Furthermore, enhancement in 'fast-slow' functional diversity in species-rich communities dominated by fast species under N addition increased species asynchrony, resulting in a robust biodiversity-stability relationship under N addition the artificial grassland communities. CONCLUSION: The findings demonstrate mechanistic links between plant species richness, 'fast-slow' functional traits, and community stability under N addition, suggesting that dynamics of biodiversity-stability relations under global changes are the results of species-specific responses of 'fast-slow' traits on the plant economics spectrum

Abundance- and functional-based mechanisms of plant diversity loss with fertilization in the presence and absence of herbivores

Nutrient supply and herbivores can regulate plant species composition, biodiversity and functioning of terrestrial ecosystems. Nutrient enrichment frequently increases plant productivity and decreases diversity while herbivores tend to maintain plant diversity in productive systems. However, the mechanisms by which nutrient enrichment and herbivores regulate plant diversity remain unclear. Abundance-based mechanisms propose that fertilization leads to the extinction of rare species due to random loss of individuals of all species. In contrast, functional-based mechanisms propose that species exclusion is based on functional traits which are disadvantageous under fertilized conditions. We tested mechanistic links between fertilization and diversity loss in the presence or absence of consumers using data from a 4-year fertilization and fencing experiment in an alpine meadow. We found that both abundance- and functional-based mechanisms simultaneously affected species loss in the absence of herbivores while only abundance-based mechanisms affected species loss in the presence of herbivores. Our results indicate that an abundance-based mechanism may consistently play a role in the loss of plant diversity with fertilization, and that diversity decline is driven primarily by the loss of rare species regardless of a plant's functional traits and whether or not herbivores are present. Increasing efforts to conserve rare species in the context of ecosystem eutrophication is a central challenge for grazed grassland ecosystems

Light competition drives herbivore and nutrient effects on plant diversity

Enrichment of nutrients and loss of herbivores are assumed to cause a loss of plant diversity in grassland ecosystems because they increase plant cover, which leads to a decrease of light in the understory 1-3. Empirical tests of the role of competition for light in natural systems are based on indirect evidence, and have been a topic of debate for the last 40 years. Here we show that experimentally restoring light to understory plants in a natural grassland mitigates the loss of plant diversity that is caused by either nutrient enrichment or the absence of mammalian herbivores. The initial effect of light addition on restoring diversity under fertilization was transitory and outweighed by the greater effect of herbivory on light levels, indicating that herbivory is a major factor that controls diversity, partly through light. Our results provide direct experimental evidence, in a natural system, that competition for light is a key mechanism that contributes to the loss of biodiversity after cessation of mammalian herbivory. Our findings also show that the effects of herbivores can outpace the effects of fertilization on competition for light. Management practices that target maintaining grazing by native or domestic herbivores could therefore have applications in protecting biodiversity in grassland ecosystems, because they alleviate competition for light in the understory

Intra-annual growing season climate variability drives the community intra-annual stability of a temperate grassland by altering intra-annual species asynchrony and richness in Inner Mongolia, China

Understanding the factors that regulate the functioning of our ecosystems in response to environmental changes can help to maintain the stable provisioning of ecosystem services to mankind. This is especially relevant given the increased variability of environmental conditions due to human activities. In particular, maintaining a stable production and plant biomass during the growing season (intra-annual stability) despite pervasive and directional changes in temperature and precipitation through time can help to secure food supply to wild animals, livestock, and humans. Here, we conducted a 29-year field observational study in a temperate grassland to explore how the intra-annual stability of primary productivity is influenced by biotic and abiotic variables through time. We found that intra-annual precipitation variability in the growing season indirectly influenced the community intra-annual biomass stability by its negative effect on intra-annual species asynchrony. While the intra-annual temperature variability in the growing season indirectly altered community intra-annual biomass stability through affecting the intra-annual species richness. At the same time, although the intra-annual biomass stability of the dominant species and the dominant functional group were insensitive to climate variability, they also promoted the stable community biomass to a certain extent. Our results indicate that ongoing intra-annual climate variability affects community intra-annual biomass stability in the temperate grassland, which has important theoretical significance for us to take active measures to deal with climate change

The effect of livestock grazing on plant diversity and productivity of mountainous grasslands in South America: A meta-analysis

Mountainous grasslands in South America, characterized by their high diversity, provide a wide range of contributions to people, including water regulation, soil erosion prevention, livestock feed provision, and preservation of cultural heritage. Prior research has highlighted the significant role of grazing in shaping the diversity and productivity of grassland ecosystems, especially in highly productive, eutrophic systems. In such environments, grazing has been demonstrated to restore grassland plant diversity by reducing primary productivity. However, it remains unclear whether these findings are applicable to South American mountainous grasslands, where plants are adapted to different environmental conditions. To address this uncertainty, we conducted a meta-analysis of experiments excluding livestock grazing to assess its impact on plant diversity and productivity across mountainous grasslands in South America. In alignment with studies in temperate grasslands, our findings indicated that herbivore exclusion resulted in increased aboveground biomass but reduced species richness and Shannon diversity. The effects of grazing exclusion became more pronounced with longer durations of exclusion; nevertheless, they remained resilient to various climatic conditions, including mean annual precipitation and mean annual temperature, as well as the evolutionary history of grazing. In contrast to results observed in temperate grasslands, the reduction in species richness due to herbivore exclusion was not associated with increased aboveground biomass. This suggests that the processes governing (sub)tropical grassland plant diversity may differ from those in temperate grasslands. Consequently, further research is necessary to better understand the specific factors influencing plant diversity and productivity in South American montane grasslands and to elucidate the ecological implications of herbivore exclusion in these unique ecosystems

Environmental heterogeneity modulates the effect of plant diversity 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.EEA Santa CruzFil: Daleo, Pedro. Consejo Nacional de Investigaciones Científicas y Técnicas. Instituto de Investigaciones Marinas y Costeras (IIMyC); Argentina.Fil: Alberti, Juan. Consejo Nacional de Investigaciones Científicas y Técnicas. Instituto de Investigaciones Marinas y Costeras (IIMyC); Argentina.Fil: Chaneton, Enrique J. Universidad de Buenos Aires. Facultad de Agronomía; Argentina.Fil: Chaneton, Enrique J. Consejo Nacional de Investigaciones Científicas y Técnicas. Instituto de Investigaciones Fisiológicas y Ecológicas Vinculadas a la Agricultura (IFEVA); Argentina.Fil: Iribarne, Oscar. Consejo Nacional de Investigaciones Científicas y Técnicas. Instituto de Investigaciones Marinas y Costeras (IIMyC); Argentina.Fil: Tognetti, Pedro M. Universidad de Buenos Aires. Facultad de Agronomía; Argentina.Fil: Tognetti, Pedro M. Consejo Nacional de Investigaciones Científicas y Técnicas. Instituto de Investigaciones Fisiológicas y Ecológicas Vinculadas a la Agricultura (IFEVA); Argentina.Fil: Bakker, Jonathan D. University of Washington. School of Environmental and Forest Sciences; Estados UnidosFil: Borer, Elizabeth T. University of Minnesota. Department of Ecology, Evolution & Behavior; Estados UnidosFil: Bruschetti, Martín. Consejo Nacional de Investigaciones Científicas y Técnicas. Instituto de Investigaciones Marinas y Costeras (IIMyC); Argentina.Fil: MacDougall, Andrew S. University of Guelph.Department of Integrative Biology; CanadáFil: Pascual, Jesús. Consejo Nacional de Investigaciones Científicas y Técnicas. Instituto de Investigaciones Marinas y Costeras (IIMyC); Argentina.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: Hautier, Yann. Utrecht University. Department of Biology. Ecology and Biodiversity Group; Países Bajo

Effects of intra-annual precipitation patterns on grassland productivity moderated by the dominant species phenology

Phenology and productivity are important functional indicators of grassland ecosystems. However, our understanding of how intra-annual precipitation patterns affect plant phenology and productivity in grasslands is still limited. Here, we conducted a two-year precipitation manipulation experiment to explore the responses of plant phenology and productivity to intra-annual precipitation patterns at the community and dominant species levels in a temperate grassland. We found that increased early growing season precipitation enhanced the above-ground biomass of the dominant rhizome grass, Leymus chinensis, by advancing its flowering date, while increased late growing season precipitation increased the above-ground biomass of the dominant bunchgrass, Stipa grandis, by delaying senescence. The complementary effects in phenology and biomass of the dominant species, L. chinensis and S. grandis, maintained stable dynamics of the community above-ground biomass under intra-annual precipitation pattern variations. Our results highlight the critical role that intra-annual precipitation and soil moisture patterns play in the phenology of temperate grasslands. By understanding the response of phenology to intra-annual precipitation patterns, we can more accurately predict the productivity of temperate grasslands under future climate change

Herbivore exclusion stabilizes alpine grassland biomass production across spatial scales

There is growing evidence that land-use management practices such as livestock grazing can strongly impact the local diversity, functioning, and stability of grassland communities. However, whether these impacts depend on environmental condition and propagate to larger spatial scales remains unclear. Using an 8-year grassland exclosure experiment conducted at nine sites in the Tibetan Plateau covering a large precipitation gradient, we found that herbivore exclusion increased the temporal stability of alpine grassland biomass production at both the local and larger (site) spatial scales. Higher local community stability was attributed to greater stability of dominant species, whereas higher stability at the larger scale was linked to higher spatial asynchrony of productivity among local communities. Additionally, sites with higher mean annual precipitation had lower dominant species stability and lower grassland stability at both the spatial scales considered. Our study provides novel evidence that livestock grazing can impair grassland stability across spatial scales and climatic gradients

Biodiversity promotes ecosystem functioning despite environmental change

Three decades of research have demonstrated that biodiversity can promote the functioning of ecosystems. Yet, it is unclear whether the positive effects of biodiversity on ecosystem functioning will persist under various types of global environmental change drivers. We conducted a meta-analysis of 46 factorial experiments manipulating both species richness and the environment to test how global change drivers (i.e. warming, drought, nutrient addition or CO2 enrichment) modulated the effect of biodiversity on multiple ecosystem functions across three taxonomic groups (microbes, phytoplankton and plants). We found that biodiversity increased ecosystem functioning in both ambient and manipulated environments, but often not to the same degree. In particular, biodiversity effects on ecosystem functioning were larger in stressful environments induced by global change drivers, indicating that high-diversity communities were more resistant to environmental change. Using a subset of studies, we also found that the positive effects of biodiversity were mainly driven by interspecific complementarity and that these effects increased over time in both ambient and manipulated environments. Our findings support biodiversity conservation as a key strategy for sustainable ecosystem management in the face of global environmental change

Experimental impacts of grazing on grassland biodiversity and function are explained by aridity

Grazing by domestic herbivores is the most widespread land use on the planet, and also a major global change driver in grasslands. Yet, experimental evidence on the long-term impacts of livestock grazing on biodiversity and function is largely lacking. Here, we report results from a network of 10 experimental sites from paired grazed and ungrazed grasslands across an aridity gradient, including some of the largest remaining native grasslands on the planet. We show that aridity partly explains the responses of biodiversity and multifunctionality to long-term livestock grazing. Grazing greatly reduced biodiversity and multifunctionality in steppes with higher aridity, while had no effects in steppes with relatively lower aridity. Moreover, we found that long-term grazing further changed the capacity of above- and below-ground biodiversity to explain multifunctionality. Thus, while plant diversity was positively correlated with multifunctionality across grasslands with excluded livestock, soil biodiversity was positively correlated with multifunctionality across grazed grasslands. Together, our cross-site experiment reveals that the impacts of long-term grazing on biodiversity and function depend on aridity levels, with the more arid sites experiencing more negative impacts on biodiversity and ecosystem multifunctionality. We also highlight the fundamental importance of conserving soil biodiversity for protecting multifunctionality in widespread grazed grasslands