Human activities are transforming grassland biomass via changing climate, elemental nutrients, and herbivory. Theory predicts that food-limited herbivores will consume any additional biomass stimulated by nutrient inputs (‘consumer-controlled’). Alternatively, nutrient supply is predicted to increase biomass where herbivores alter community composition or are limited by factors other than food (‘resource-controlled’). Using an experiment replicated in 58 grasslands spanning six continents, we show that nutrient addition and vertebrate herbivore exclusion each caused sustained increases in aboveground live biomass over a decade, but consumer control was weak. However, at sites with high vertebrate grazing intensity or domestic livestock, herbivores consumed the additional fertilization-induced biomass, supporting the consumer-controlled prediction. Herbivores most effectively reduced the additional live biomass at sites with low precipitation or high ambient soil nitrogen. Overall, these experimental results suggest that grassland biomass will outstrip wild herbivore control as human activities increase elemental nutrient supply, with widespread consequences for grazing and fire risk.

Background and aims The diverse flora of the Brazilian Cerrado is threatened by agricultural expansion, nutrient enrichment and invasion of alien plants. We performed a fertilization experiment to investigate the nature of nutrient limitation in Cerrado vegetation, and evaluate whether native and alien invasive species are limited by the same or different nutrients. Methods We applied various combinations of nutrients (phosphorus (P), nitrogen (N), and a mixture of other macro-and micro-nutrients ('cations treatment')) to six types of Cerrado vegetation. We then studied over a 3year period how these treatments affected the aboveground biomass of native forbs, native C 3 and C 4 grasses, and invasive C 4 grasses. ResultsThe full nutrient treatment (N+P+ 'cations') significantly increased total community biomass across our sites, but P alone had no effect. The nutrient treatments also affected the relative abundance of functional plant groups in the six vegetation types. P addition, either alone or in combination with other nutrients, increased the biomass of alien C 4 grasses, where present, whereas the cations treatment stimulated growth of the native C 4 grasses. Addition of N+P reduced the biomass of native C 3 grasses. Conclusions Our results indicate co-limitation by several nutrients, including P, perhaps N, and at least one other nutrient. Further research is needed to determine what the other nutrient (or nutrients) may be. Native and invasive species appear to be limited by different nutrients, with P alone stimulating growth of African C 4 grasses. This should be considered in managing both natural and invaded communities.

SummaryAlthough 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.

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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.

Cerrado is the second largest biome in Brazil and is classified as a biodiversity hotspot. The establishment of hydroelectric power stations in Brazil originated degraded areas due to the removal of soil for construction of dams, in which native vegetation sometimes fails to reestablish due to the bad soil conditions. Sheep excrete most of the nutrients they ingest, such as phosphorus, calcium, potassium and nitrogen. This study aimed at investigating whether sheep dung contributes to the improvement of soil quality and stimulates the process of ecological succession in areas affected by the construction of the hydroelectric power station of Ilha Solteira. Four areas were selected, located at the Experimental Farm of Unesp Ilha Solteira/SP, and ten plots of 1 m2 each were established. From October 2014 to December 2015, five replicates received 150 g fresh sheep dung every 15 days. In December 2015 soil sample from top 0.10 m was collected for the determination of soil attributes. Aboveground biomass was also collected, separated into four functional groups, oven-dried and weighed. Data were analysed to check the effect of dung addition upon soil and vegetation variables. Dung addition stimulated the development of several functional groups and changed soil nutrient concentrations in all four studied areas.

Nutrients are known to limit productivity of plant communities around the world. In the Brazilian Cerrado, indirect evidences point to phosphorus as the main limiting nutrient, but some fertilization experiments suggest that one or more micronutrients might play this role. Boron is one of the essential micronutrients for plants. Agronomically, it received some attention, but it has mostly been neglected in ecological studies assessing the effects of nutrients on plant growth. Through field fertilization and mesocosm experiments in a degraded area in the Cerrado, we show that boron addition increased biomass production of herbaceous vegetation. This could be related to a lower aluminum uptake in the boron fertilized plants. Even considering that plant growth was promoted by boron addition due to aluminum toxicity alleviation, this is the first study reporting boron limitation in natural, noncultivated plant communities and also the first report of this kind in vegetative grasses. These results contribute to disentangling patterns of nutrient limitation among plant species of the species‐rich, aluminum‐rich, and nutrient‐poor Cerrado biome and highlight the potential role of micronutrients, such as boron, for growth of noncrop plants. Understanding how nutrient limitation differs among functional groups in the highly biodiverse areas founded on ancient tropical soils may help managing these plant communities in a changing world.

Brazil has extensive degraded areas, where vegetation fails to establish due to harsh soil conditions. However, some invasive species such as Urochloa decumbens are successful pioneers in such areas, but the reasons deserve investigation. Mycorrhizal fungi are abundant in Cerrado soils, and their association with plants are beneficial for their establishment in natural and degraded areas. This study investigated whether arbuscular mycorrhizal colonization of native and exotic plants in the Cerrado differs between pristine and degraded areas. We collected 135 plants from four functional groups in 68 areas and determined the percentage of mycorrhizal colonization. The invasive grass Urochloa decumbens had significantly higher colonization rates in degraded than in native areas, and higher colonization than the native species. These results are important for soil management since Urochloa decumbens is widely used in early soil restoration efforts, and for nature conservation concerning the management of invasive plants in restoration areas.