Last-century increases in intrinsic water-use efficiency of grassland communities have occurred over a wide range of vegetation composition, nutrient inputs and soil pH

Last-century climate change has led to variable increases of the intrinsic water-use efficiency (Wi; the ratio of net CO2 assimilation to stomatal conductance for water vapor) of trees and C3 grassland ecosystems, but the causes of the variability are not well understood. Here, we address putative drivers underlying variable Wi responses in a wide range of grassland communities. Wi was estimated from carbon isotope discrimination in archived herbage samples from 16 contrasting fertilizer treatments in the Park Grass Experiment, Rothamsted, England, for the 1915 to 1929 and 1995 to 2009 periods. Changes in Wi were analyzed in relation to nitrogen input, soil pH, species richness, and functional group composition. Treatments included liming as well as
phosphorus and potassium additions with or without ammonium or nitrate fertilizer applications at three levels. Wi increased between 11% and 25% (P , 0.001) in the different treatments between the two periods. None of the fertilizers had a direct effect on the change of Wi (DWi). However, soil pH (P , 0.05), species richness (P , 0.01), and percentage grass content (P , 0.01) were significantly related to DWi
. Grass-dominated, species-poor plots on acidic soils showed the largest DWi (+14.7 mmol mol21).
The DWi response of these acidic plots was probably related to drought effects resulting from aluminum toxicity on root
growth. Our results from the Park Grass Experiment show that Wi in grassland communities consistently increased over a
wide range of nutrient inputs, soil pH, and plant community compositions during the last century.