Herbivory and nutrients shape grassland soil seed banks.

Anthropogenic nutrient enrichment and shifts in herbivory can lead to dramatic changes in the composition and diversity of aboveground plant communities. In turn, this can alter seed banks in the soil, which are cryptic reservoirs of plant diversity. Here, we use data from seven Nutrient Network grassland sites on four continents, encompassing a range of climatic and environmental conditions, to test the joint effects of fertilization and aboveground mammalian herbivory on seed banks and on the similarity between aboveground plant communities and seed banks. We find that fertilization decreases plant species richness and diversity in seed banks, and homogenizes composition between aboveground and seed bank communities. Fertilization increases seed bank abundance especially in the presence of herbivores, while this effect is smaller in the absence of herbivores. Our findings highlight that nutrient enrichment can weaken a diversity maintaining mechanism in grasslands, and that herbivory needs to be considered when assessing nutrient enrichment effects on seed bank abundance.

Evolutionary history of grazing and resources determine herbivore exclusion effects on plant diversity.

Ecological models predict that the effects of mammalian herbivore exclusion on plant diversity depend on resource availability and plant exposure to ungulate grazing over evolutionary time. Using an experiment replicated in 57 grasslands on six continents, with contrasting evolutionary history of grazing, we tested how resources (mean annual precipitation and soil nutrients) determine herbivore exclusion effects on plant diversity, richness and evenness. Here we show that at sites with a long history of ungulate grazing, herbivore exclusion reduced plant diversity by reducing both richness and evenness and the responses of richness and diversity to herbivore exclusion decreased with mean annual precipitation. At sites with a short history of grazing, the effects of herbivore exclusion were not related to precipitation but differed for native and exotic plant richness. Thus, plant species' evolutionary history of grazing continues to shape the response of the world's grasslands to changing mammalian herbivory.

Species richness both impedes and promotes alien plant invasions in the Brazilian Cerrado.

Worldwide, alien plant invasions have been intensively studied in the past decades, but mechanisms controlling the invasibility of native communities are not fully understood yet. The stochastic niche hypothesis predicts that species-rich plant communities are less prone to alien plant invasions than species-poor communities, which is supported by some but not all field studies, with some very species-rich communities such as the Brazilian Cerrado becoming heavily invaded. However, species-rich communities potentially contain a greater variety of facilitative interactions in resource exploitation than species-poor communities, from which invasive plants might benefit. This alternative hypothetical mechanism might explain why nutrient-poor, species-rich ecosystems are prone to invasion. Here we show that a high species richness both impedes and promotes invasive plants in the Brazilian Cerrado, using structural equation modelling and data from 38 field sites. We found support for the stochastic niche hypothesis through an observed direct negative influence of species richness on abundance of alien invasive species, but an indirect positive effect of species richness on invasive alien plants through soil phosphatase activity that enhances P availability was also found. These field observations were supported with results from a mesocosm experiment. Root phosphatase activity of plants increased with species richness in the mesocosms, which was associated with greater community P and N uptake. The most prominent alien grass species of the region, Melinis minutiflora, benefited most from the higher N and P availability in the species mixtures. Hence, this study provides a novel explanation of why species-richness may sometimes promote rather than impede invasion, and highlights the need to perform facilitation experiments in multi-species communities.

Alien and endangered plants in the Brazilian Cerrado exhibit contrasting relationships with vegetation biomass and N : P stoichiometry.

Although endangered and alien invasive plants are commonly assumed to persist under different environmental conditions, surprisingly few studies have investigated whether this is the case. We examined how endangered and alien species are distributed in relation to community biomass and N : P ratio in the above-ground community biomass in savanna vegetation in the Brazilian Cerrado. For 60 plots, we related the occurrence of endangered (Red List) and alien invasive species to plant species richness, vegetation biomass and N : P ratio, and soil variables. Endangered plants occurred mainly in plots with relatively low above-ground biomass and high N : P ratios, whereas alien invasive species occurred in plots with intermediate to high biomass and low N : P ratios. Occurrences of endangered or alien plants were unrelated to extractable N and P concentrations in the soil. These contrasting distributions in the Cerrado imply that alien species only pose a threat to endangered species if they are able to invade sites occupied by these species and increase the above-ground biomass and/or decrease the N : P ratio of the vegetation. We found some evidence that alien species do increase above-ground community biomass in the Cerrado, but their possible effect on N : P stoichiometry requires further study.