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Plant species loss from European semi-natural grasslands following nutrient enrichment – is it nitrogen or is it phosphorus?


Tobias Ceulemans, Division Plant Ecology and Systematics, Department Biology, University of Leuven, Kasteelpark Arenberg 31, B-3001 Leuven, Belgium. E-mail:


Aim  Although many studies support the prevailing paradigm of nitrogen (N)-driven biodiversity loss, some have argued that phosphorus (P) may be the main culprit. This questions the generality of the global threat through N enrichment. The major objective here was to quantify the relative importance of soil N and P in explaining patterns of plant species richness, under different levels of N and P limitation.

Location  North-western Europe.

Methods  We collected soil, productivity and plant species data from 132 semi-natural grasslands located along a gradient of nutrient availability and atmospheric N deposition. We used linear mixed models to investigate the relation between soil nutrients, acidity, limitation and productivity on one side, and indices for plant species richness on the other.

Results  Mixed models explained between 38 and 50% of the total variation in species numbers, forbs and endangered species. Soil P was significantly negatively related to total species number, forbs and endangered species. Soil N was only significantly negatively related to number of forbs and endangered species. Compared with soil P, the explained variation attributed to soil N was between five- and twenty-fold lower. P-limited grasslands exhibited higher species richness, numbers of forbs and endangered species. Species richness and number of forbs decreased with lower soil acidity. N deposition was negatively related to the number of forbs and endangered species, as well as to soil acidity. Productivity was weakly positively related to soil P and negatively to species and forb numbers. We found no interaction factors between the explanatory variables.

Main conclusions  P enrichment can present a greater threat to biodiversity than N enrichment in at least some terrestrial ecosystems. However, as N- and P-driven species loss appeared independent, our results suggest that simultaneously reducing N and P inputs is a prerequisite for maintaining maximum plant diversity.