Foliar and soil concentrations and stoichiometry of N and P across European Pinus sylvestris forests : relationships with climate, N deposition and tree growth

1. This study investigated the factors underlying the variability of needle and soil elemental composition and stoichiometry and their relationships with growth in Pinus sylvestris forests throughout the species' distribution in Europe by analyzing data from 2245 forest stands. 2. Needle N concentrations and N:P ratios were positively correlated with total atmospheric N deposition, whereas needle P concentrations were negatively correlated. These relationships were especially pronounced at sites where high levels of N deposition coincided with both higher mean annual temperature and higher mean annual precipitation. Trends toward foliar P deficiency were thus more marked when high N deposition coincided with climatic conditions favorable to plant production. 3. Atmospheric N deposition was positively correlated with soil-solution NO3-, SO42-, K+, P, and Ca2+ concentrations, the soil-solution NO3-:P ratio, total soil N, and the total soil N:Olsen P ratio, and negatively correlated with soil Olsen P concentration. 4. Despite these nutrient imbalances, during the period studied (1990-2006), N deposition was positively related with Pinus sylvestris absolute basal diameter (BD) growth, although only accounting for the 10% of the total variance. However, neither N deposition nor needle N concentration were related with relative annual BD growth. In contrast, needle P concentration was positively related with both absolute and relative annual BD growth. 5. These results thus indicate a tendency of European P. sylvestris forests to store N in trees and soil in response to N deposition and unveil a trend toward increased nutrient losses in runoff as a consequence of higher soil-solution N concentrations. Overall, the data show increasing ecosystem nutrient imbalances with increasingly limiting roles of P and other nutrients such as K in European P. sylvestris forests, especially in the center of their distribution where higher levels of N deposition are observed. Thus, although the data show that N deposition has had an overall positive effect on P. sylvestris growth, the effect of continuous N deposition, associated with decreasing P and K and increasing N:P in leaves and in soil, may in the future become detrimental for the growth and competitive ability of P. Sylvestris trees