Rainfall changes alter plant communities in Chihuahuan Desert grasslands

Variations in New Mexico’s climate have been detected over the past century and are expected to continue into the future. Some of these climatic changes result in more frequent instances of extreme drought events and alterations of monsoon seasonality, with monsoons being pushed later into the fall. Such changes could make our Chihuahuan Desert plant communities less productive, shift the abundance of the plant species, or have other destabilizing impacts. Our experiment manipulated precipitation in two desert grassland communities at the Sevilleta National Wildlife Refuge near Socorro, NM. Some plots were subjected to an extreme drought event, receiving only 33% of annual precipitation for five years, and others were subjected to a change in the timing of monsoon precipitation, delaying rainfall to September and October. We found that not only did plant production decrease but plant community assembly also changed in response to these treatments. The greatest production loss was in the extreme drought manipulation which also experienced a change in species diversity, an immediate diversity decrease that eventually recovered to levels above its initial condition. Plant diversity in the delayed monsoon treatments initially increased and eventually settled to near ambient levels, probably due to an initial propagation of bet-hedging species that had been accumulating in the seed bank. In both grasslands, the two treatments caused the dominate grasses to become subordinate which likely drove the lowered productivity and greater evenness of the plant abundances within communities of both treatments

Exploring the impact of trait number and type on functional diversity metrics in real-world ecosystems

The use of trait-based approaches to understand ecological communities has increased in the past two decades because of their promise to preserve more information about community structure than taxonomic methods and their potential to connect community responses to subsequent effects of ecosystem functioning. Though trait-based approaches are a powerful tool for describing ecological communities, many important properties of commonly-used trait metrics remain unexamined. Previous work in studies that simulate communities and trait distributions show consistent sensitivity of functional richness and evenness measures to the number of traits used to calculate them, but these relationships have yet to be studied in actual plant communities with a realistic distribution of trait values, ecologically meaningful covariation of traits, and a realistic number of traits available for analysis. Therefore, we propose to test how the number of traits used and the correlation between traits used in the calculation of functional diversity indices impacts the magnitude of eight functional diversity metrics in real plant communities. We will use trait data from three grassland plant communities in the US to assess the generality of our findings across ecosystems and experiments. We will determine how eight functional diversity metrics (functional richness, functional evenness, functional divergence, functional dispersion, kernel density estimation (KDE) richness, KDE evenness, KDE dispersion, Rao's Q) differ based on the number of traits used in the metric calculation and on the correlation of traits when holding the number of traits constant. Without a firm understanding of how a scientist's choices impact these metric, it will be difficult to compare results among studies with different metric parametrization and thus, limit robust conclusions about functional composition of communities across systems

Nutrient enrichment alters seasonal β-diversity in global grasslands

Intra-annual (i.e. seasonal) temporal niche partitioning is essential to the maintenance of biodiversity in many plant communities. However, understanding of how climate and global change drivers such as eutrophication influence seasonal niche partitioning in plant assemblages remains limited. We used early-season and late-season compositional data collected from 10 grassland sites around the world to explore relationships between climate variability and intra-annual species segregation (i.e. seasonal β-diversity) and to assess how nutrient enrichment alters seasonal β-diversity in plant communities. We then assessed whether changes in seasonal β-diversity in response to nutrient enrichment are underpinned by species turnover or nestedness and determined how specific functional groups (i.e. annual forbs, perennial forbs, C3 and C4 graminoids and legumes) respond to eutrophication within and across early and late sampling dates. We found a positive relationship between intra-annual temperature variability and seasonal β-diversity but observed no relationship between intra-annual precipitation variability and seasonal β-diversity. Nutrient enrichment increased seasonal β-diversity and increased turnover of species between early- and late-season communities. Nutrient enrichment reduced the abundance of C4 graminoids and legumes within and across sampling timepoints and eliminated intra-annual differences in these groups. In contrast, nutrient enrichment resulted in seasonal differences in C3 graminoids, which were not observed in control conditions and increased abundance of C3 graminoids and annual forbs within and across early and late sampling dates. Synthesis: Our understanding of how grasslands respond to various components of global change is primarily based on studies that document community changes at inter-annual scales. Using early-season and late-season compositional data from 10 grassland sites around the world, we show that nutrient enrichment increases seasonal β-diversity and alters intra-annual dynamics of specific functional groups in unique ways

Multiple resource limitations explain biomass-precipitation relationships in grasslands

Interannual variability in grassland primary production is strongly driven by precipitation, nutrient availability and herbivory, but there is no general consensus on the mechanisms linking these variables. If grassland biomass is limited by the single most limiting resource at a given time, then we expect that nutrient addition will not affect biomass production at arid sites. We conducted a distributed experiment manipulating nutrients and herbivores at 44 grassland sites in 8 regions around the world, spanning a broad range in aridity. We estimated the effects of 5-11 years of nutrient addition and herbivore exclusion treatments on precipitation sensitivity of biomass (proportional change in biomass relative to proportional change in rainfall among years), and the biomass in the driest year (to measure treatment effects when water was most limiting) at each site. Grazer exclusion did not interact with nutrients to influence driest year biomass or sensitivity. Nutrient addition increased driest year biomass by 74% and sensitivity by 0.12 (proportional units), and that effect did not change across the range of aridity spanned by our sites. Grazer exclusion did not interact with nutrients to influence sensitivity or driest year biomass. At almost half of our sites, the previous year\u27s rainfall explained as much variation in biomass as current year precipitation. Overall, our distributed fertilization experiment detected co-limitation between nutrients and water governing grasslands, with biomass sensitivity to precipitation being limited by nutrient availability irrespective of site aridity and herbivory. Our findings refute the classical ideas that grassland plant performance is limited by the single most limiting resource at a site. This suggests that nutrient eutrophication will destabilize grassland ecosystems through increased sensitivity to precipitation variation

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

Increasing effects of chronic nutrient enrichment on plant diversity loss and ecosystem productivity over time

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

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

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

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