Anthropogenic activities are increasing nutrient availability and altering precipitation regimes. This may lead to critical changes in grasslands functioning. This is particularly important for grasslands in the Mediterranean Basin that have evolved in nutrient poor soils, and where more frequent and prolonged droughts are projected to occur. However, there is limited knowledge regarding the interacting effects of multiple nutrient inputs and rainfall variability on the plant functional structure and diversity of Mediterranean grasslands. We conducted a nutrient addition experiment in a Mediterranean grassland during four contrasting precipitation years. We established four treatments that varied in the number of added nutrients, from no added nutrients (control), to one added nutrient (Nitrogen-N, Phosphorus-P, or Potassium-K), two added nutrients, (NP, NK, or PK) and three added nutrients (NPK). We assessed the effect of increasing nutrient addition in wet, normal, dry, and very dry years on plant species functional traits at the community level. We determined the community functional structure (e.g., Community Weighted Mean, CWM) and functional diversity (e.g., Functional Dispersion, FDis) for eight key functional traits indicators of nutrient and water use strategies. We also assessed if CWM, FDis, and species richness were related to the aggregate grassland functioning property, i.e., productivity. We found that CWM was affected by nutrient addition and precipitation and, for some traits, by their interaction. However, FDis of most traits was affected by precipitation. The very dry year had a negative effect on FDis of most traits (e.g., dispersal modes, nutrient uptake strategies) and interacted with three added nutrients to decrease FDis of growth-forms. Conversely, FDis of reproductive traits decreased during the wet year. Species richness and FDis were not related to grassland productivity, whereas CWM was the main determinant of grassland productivity supporting the importance of species functional traits in determining ecosystem functioning. Our results highlight drought as a critical factor determining a decrease in the functional diversity of Mediterranean grasslands. Moreover, drought can also interact with nutrient addition changing the dominance of many traits and further decreasing functional diversity. This may have important implications for grasslands functioning in the context of global changes.

Document Type


Publication Date


Notes/Citation Information

Published in Frontiers in Ecology and Evolution, v. 6, 155, p. 1-15.

© 2018 Nogueira, Nunes, Bugalho, Branquinho, McCulley and Caldeira.

This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner(s) are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.

Digital Object Identifier (DOI)


Funding Information

The authors acknowledge the financial support of the Portuguese Science Foundation (Fundação para a Ciência e Tecnologia- FCT), through the project IF/740/2014 and PTDC/AAG-GLO/0045/2014 and fellowships to CN (SFRH/BD/88650/2012) and MB (IF/1171/2014). Centro de Estudos Florestais (CEF) a research unit funded by FCT, Portugal (UID/AGR/00239/2013). We are also grateful to FLAD/NSF for funding (A1/Proj 124/12).

Related Content

The raw data supporting the conclusions of this manuscript will be made available by the authors, without undue reservation, to any qualified researcher.