Cycling and retention of nitrogen in European beech (<i>Fagus sylvatica</i> L.) ecosystems under elevated fructification frequency

Abstract

Atmospheric deposition of nitrogen (N) has exceeded its demand for plant increment in forest ecosystems in Germany. High N inputs increased plant growth, the internal N cycling within the ecosystem, the retention of N in soil and plant compartments, and the N output by seepage water. But the processes involved are not fully understood, notably the effect of fructification in European beech (Fagus sylvatica L.) on N fluxes. The frequency of fructification has increased together with air temperature and N deposition, but its impact on N fluxes and the sequestration of carbon (C) and N in soils have been hardly studied. A field experiment using 15N-labeled leaf litter exchange was carried out over a 5.5-year period at seven long-term European beech (Fagus sylvatica L.) monitoring sites to study the impact of current mast frequency on N cycling. Mean annual leaf litterfall contained 35 kg N ha−1, but about one-half of that was recovered in the soil 5.5 years after the establishment of the leaf litter 15N exchange experiment. In these forests, fructification occurred commonly at intervals of 5 to 10 years, which has now changed to every 2 years as observed during this study period. Seed cupules contributed 51 % to the additional litterfall in mast years, which creates a high nutrient demand during their decomposition due to the very high ratios of C to N and C to phosphorus (P). Retention of leaf litter 15N in the soil was more closely related to the production of total litterfall than to the leaf litterfall, indicating the role of seed cupules in the amount of leaf N retained in the soil. Higher mast frequency increased the mass of mean annual litterfall by about 0.5 Mg ha−1 and of litterfall N by 8.7 kg ha−1. Mean net primary production (NPP) increased by about 4 %. Mean total N retention in soils calculated by input and output fluxes was unrelated to total litterfall, indicating that mast events were not the primary factor controlling total N retention in soils. Despite reduced N deposition since the 1990s, about 5.7 out of 20.7 kg N ha−1 deposited annually between 1994 and 2008 was retained in soils, notably at acid sites with high N/P and C/P ratios in the organic layers and mineral soils, indicating P limitation for litter decomposition. Trees retained twice as much N compared to soils by biomass increment, particularly in less acidic stands where the mineral soils had low C/N ratios. These results have major implications for our understanding of the C and N cycling and N retention in forest ecosystems. In particular the role of mast products in N retention needs more research in the future.