Seeing the trees for the forest: Nitrogen deposition alters tree growth and survival across the northeastern U.S., responses vary by species.

Assessments of regional forest carbon (C) balance have long speculated a role for atmospheric nitrogen (N) deposition. To date, however, evidence for an N effect has been restricted to plot-level fertilization and 15N experiments, biogeochemical modeling studies, and recent and controversial correlations between nitrogen deposition and C balance across 20 intensive C monitoring sites. These studies have yielded widely varying conclusions on the magnitude and even the direction of response (positive or negative effects on C storage). We assessed the effects of N deposition on forest growth and survival using forest inventory data from > 160,000 trees, spanning a 19-state region of the northeastern United States, and estimates of total (wet and plot-specific dry) inorganic N inputs (N deposition to the plots ranged from 3.2 to 11 kg N ha-1 y-1). The growth rates of 14 of the region's 24 most common tree species responded to N deposition. Nine species showed a monotonic increase in aboveground growth rates across the range of N deposition and some of these species experienced a doubling of growth rates. Four species showed humped-shaped responses. A single species, Pinus resinosa (red pine), showed a monotonic decline in growth across the range of deposition. Species responsiveness to N deposition increased with mean growth rate of the species. In the analysis of survival, 13 of the 24 species responded to N: eight species showed a monotonic decrease, 3 showed a monotonic increase and 1 showed a humped-shape response in survival. The stand-level analysis showed a distinctly humped-shaped relationship between N deposition and aboveground C increment, peaking at 6 kg N ha-1 yr-1, which can likely be explained by the distribution of species and their individual growth and mortality responses. Overall, our data suggest that moderate levels of N deposition have enhanced aboveground C accumulation in temperate forests of the northeastern U.S., but the shape and trajectory of the response is species-specific.