Ammonium as a Driving Force of Plant Diversity and Ecosystem Functioning: Observations Based on 5 Years' Manipulation of N Dose and Form in a Mediterranean Ecosystem

Enhanced nitrogen (N) availability is one of the main drivers of biodiversity loss and degradation of ecosystem functions. However, in very nutrient-poor ecosystems, enhanced N input can, in the short-term, promote diversity. Mediterranean Basin ecosystems are nutrient-limited biodiversity hotspots, but no information is available on their medium- or long-term responses to enhanced N input. Since 2007, we have been manipulating the form and dose of available N in a Mediterranean Basin maquis in south-western Europe that has low ambient N deposition (<4 kg N ha⁻¹ yr⁻¹) and low soil N content (0.1%). N availability was modified by the addition of 40 kg N ha⁻¹ yr⁻¹ as a 1∶1 NH₄Cl to (NH₄)₂SO₄ mixture, and 40 and 80 kg N ha⁻¹ yr⁻¹ as NH₄NO₃. Over the following 5 years, the impacts on plant composition and diversity (richness and evenness) and some ecosystem characteristics (soil extractable N and organic matter, aboveground biomass and % of bare soil) were assessed. Plant species richness increased with enhanced N input and was more related to ammonium than to nitrate. Exposure to 40 kg NH₄ ⁺-N ha⁻¹ yr⁻¹ (alone and with nitrate) enhanced plant richness, but did not increase aboveground biomass; soil extractable N even increased under 80 kg NH₄NO₃-N ha⁻¹ yr⁻¹ and the % of bare soil increased under 40 kg NH₄ ⁺-N ha⁻¹ yr⁻¹. The treatment containing less ammonium, 40 kg NH₄NO₃-N ha⁻¹ yr⁻¹, did not enhance plant diversity but promoted aboveground biomass and reduced the % of bare soil. Data suggest that enhanced NH_y availability affects the structure of the maquis, which may promote soil erosion and N leakage, whereas enhanced NO_x availability leads to biomass accumulation which may increase the fire risk. These observations are relevant for land use management in biodiverse and fragmented ecosystems such as the maquis, especially in conservation areas.