Drought effects on growth and density of temperate tree regeneration under different levels of nitrogen deposition

Temperate forests in Central Europe suffer from climate change-induced productivity and vitality reductions and increased tree mortality. Most field work assessing climate change effects in forests refers to mature trees and does not cover interaction between climate change and nitrogen deposition. Here we show in a study of 54 forest sites representing different combinations of climatic conditions and atmospheric nitrogen deposition across Germany that nitrogen significantly affects the drought tolerance of tree regeneration in the field. We compared shoot length increment and regeneration density of Central Europe's naturally most dominant tree species, European beech (Fagus sylvatica) with three species, which are discussed as potentially more drought-tolerant replacement tree species for climate change adaptation of forestry. Growth of beech was reduced with increasing nitrogen deposition identifying high reactive nitrogen loads as an additional threat for this species, but between-site variation of beech growth was not dependent on climatic parameters despite growth reductions after the drought summer 2018 across all study sites. Remarkably, the only species in the between-site comparisons where low growth was attributable to dry climate was Douglas fir (Pseudotsuga menziesii), as shoot length increment was reduced at sites with dry spring climate. Silver fir (Abies alba) showed increasing growth with increasing nitrogen deposition in combination with increasing drought, probably due to reduced ammonium uptake. Shoot length increment in sessile oak (Quercus petraea) was not affected by the between-site variation of either drought or nitrogen. Our results (combined with results of regeneration density) indicate that high atmospheric nitrogen deposition, which is primarily found in regions with intensive livestock farming in Central Europe, modifies the climate change response of temperate tree species. Interaction of drought and nitrogen deposition should thus be addressed in the debate on climate change adaptation of forestry.