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.

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.

Nitrogen and phosphorus fertilization consistently favor pathogenic over mutualistic fungi in grassland soils.

Ecosystems across the globe receive elevated inputs of nutrients, but the consequences of this for soil fungal guilds that mediate key ecosystem functions remain unclear. We find that nitrogen and phosphorus addition to 25 grasslands distributed across four continents promotes the relative abundance of fungal pathogens, suppresses mutualists, but does not affect saprotrophs. Structural equation models suggest that responses are often indirect and primarily mediated by nutrient-induced shifts in plant communities. Nutrient addition also reduces co-occurrences within and among fungal guilds, which could have important consequences for belowground interactions. Focusing only on plots that received no nutrient addition, soil properties influence pathogen abundance globally, whereas plant community characteristics influence mutualists, and climate influence saprotrophs. We show consistent, guild-level responses that enhance our ability to predict shifts in soil function related to anthropogenic eutrophication, which can have longer-term consequences for plant communities.

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.

Manipulating plant phylogenetic diversity for green roof ecosystem service delivery.

Plant species and functional trait diversity have each been shown to improve green roof services. Species and trait differences that contribute to ecosystem services are the product of past evolutionary change and phylogenetic diversity (PD), which quantifies the relatedness among species within a community. In this study, we present an experimental framework to assess the contribution of plant community PD for green roof ecosystem service delivery, and data from one season that support our hypotheses that PD would be positively correlated with two services: building cooling and rainwater management. Using 28 plant species in 12 families, we created six community combinations with different levels of PD. Each of these communities was replicated at eight green roofs along an elevation gradient, as well as a ground level control. We found that the minimum and mean roof temperature decreased with increasing PD in the plant community. Increasing PD also led to an increase in the volume of rainwater captured, but not the proportion of water lost via evapotranspiration 48 hr following the rain event. Our findings suggest that considering these evolutionary relationships could improve functioning of green infrastructure and we recommend that understanding how to make PD (and other measures of diversity) serviceable for plant selection by practitioners will improve the effectiveness of design and ecosystem service delivery. Lastly, since no two green roof sites are the same and can vary tremendously in microclimate conditions, our study illustrates the importance of including multiple independent sites in studies of green roof performance.

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.

Predicting communities from functional traits.

Species traits influence where species live and how they interact. While there have been many advances in describing the functional composition and diversity of communities, only recently do researchers have the ability to predict community composition and diversity. This predictive ability can offer fundamental insights into ecosystem resilience and restoration.