Biotic and abiotic controls of nitrogen and phosphorus cycling in Central European forests

The functioning and services of Central European forests are threatened by global change and a loss of biodiversity. Nutrient cycling as a key forest function is affected by biotic drivers (e.g., dominant tree species, understory plants, soil organisms) that interact with abiotic conditions (e.g., climate, soil properties). In contrast to grassland ecosystems, evidence for the relationship of nutrient cycles and biodiversity in forests is scarce because the structural complexity of forests limits experimental control of driving factors. Alternatively, observational studies along gradients in abiotic conditions and biotic properties may elucidate the role of biodiversity for forest nutrient cycles. This thesis aims to improve the understanding of the functional importance of biodiversity for nutrient cycles in forests by analyzing water-bound fluxes of nitrogen (N) and phosphorus (P) along gradients in biodiversity in three regions of Germany. The tested hypotheses included: (1) temperate forest canopies retain atmospheric N and retention increases with increasing plant diversity, (2) N release from organic layers increases with resource availability and population size of decomposers but N leaching decreases along a gradient in plant diversity, (3) P leaching from forest canopies increases with improved P supply from recalcitrant P fractions by a more diverse ectomycorrhizal fungal community. In the canopies of 27 forest stands from three regions, 16 % to 51 % of atmospheric N inputs were retained. Regional differences in N retention likely resulted from different in N availability in the soil. Canopy N retention was greater in coniferous than in beech forests, but this was not the case on loessderived soils. Nitrogen retention increased with increasing tree and shrub diversity which suggested complementary aboveground N uptake. The strength of the diversity effect on canopy N uptake differed among regions and between coniferous and deciduous forests. The N processing in the canopy directly coupled back to N leaching from organic layers in beech forests because throughfall-derived N flushed almost completely through the mull-type organic layers at the 12 studied beech sites. The N release from organic layers increased with stand basal area but was rather low (< 10 % of annual aboveground litterfall) because of a potentially high microbial N immobilization and intensive incorporation of litter into the mineral soil by bioturbation. Soil fauna biomass stimulated N mineralization through trophic interactions with primary producers and soil microorganisms. Both gross and net leaching from organic layers decreased with increasing plant diversity. Especially the diversity but not the cover of herbs increased N uptake. In contrast to N, P was leached from the canopy. Throughfall-derived P was also flushed quickly through the mull-type organic layers and leached P was predominantly immobilized in non directly plant-available P fractions in the mineral soil. Concentrations of plant-available phosphate in mineral soil solution were low and P leaching from the canopy increased with increasing concentrations of the moderately labile P fraction in soil and increasing ectomycorrhiza diversity while leaf C:P ratios decreased. This suggested that tree P supply benefited from complementary mining of diverse mycorrhizal communities for recalcitrant P. Canopy P leaching increased in years with pronounced spring drought which could lead to a deterioration of P supply by an increasing frequency of drought events. This thesis showed that N and P cycling in Central European forests is controlled by a complex interplay of abiotic site conditions with biological processes mediated by various groups of organisms, and that diverse plant communities contribute to tightening the N cycle in Central European forests and that diverse mycorrhizal communities improve the limited P availability. Maintaining forest biodiversity seems essential to ensure forest services in the light of environmental change.

Drivers of nitrogen leaching from organic layers in Central European beech forests

Background and Aims: The response of forest ecosystems to continuous nitrogen (N) deposition is still uncertain. We investigated imports and exports of dissolved N from mull-type organic layers to identify the controls of N leaching in Central European beech forests under continuous N deposition. Methods: Dissolved N fluxes with throughfall and through mull-type organic layers (litter leachate) were measured continuously in 12 beech forests on calcareous soil in two regions in Germany over three consecutive growing seasons. Results Mean growing season net (i.e. litter leachate – throughfall flux) fluxes of total dissolved N (TDN) from the organic layer were low (2.3 ± 5.6 kg ha −1 ) but varied widely from 12.9 kg ha −1 to –8.3 kg ha −1 . The small increase of dissolved N fluxes during the water passage through mull-type organic layers suggested that high turnover rates coincided with high microbial N assimilation and plant N uptake. Stand basal area had a positive feedback on N fluxes by providing litter for soil organic matter forma- tion. Plant diversity, especially herb diversity, reduced dissolved N fluxes. Soil fauna biomass increased NO3−-N fluxes with litter leachate by stimulating mineralization. Microbial biomass measures were not related to dissolved N fluxes. Conclusions Our results show that dissolved N exports from organic layers contain significant amounts of throughfall-derived N (mainly NO3−-N) that flushes through the organic layer but also highlight that N leaching from organic layers is driven by the complex interplay of plants, animals and microbes. Furthermore, diverse understories reduce N leaching from Central European beech forests.

The nitrogen cycle of tropical montane forest in Ecuador turns inorganic under environmental change

Water-bound nitrogen (N) cycling in temperate terrestrial ecosystems of the Northern Hemisphere is today mainly inorganic because of anthropogenic release of reactive N to the environment. In little-industrialized and remote areas, in contrast, a larger part of N cycling occurs as dissolved organic N (DON). In a north Andean tropical montane forest in Ecuador, the N cycle changed markedly during 1998–2010 along with increasing N deposition and reduced soil moisture. The DON concentrations and the fractional contribution of DON to total N significantly decreased in rainfall, throughfall, and soil solutions. This inorganic turn of the N cycle was most pronounced in rainfall and became weaker along the flow path of water through the system until it disappeared in stream water. Decreasing organic contributions to N cycling were caused not only by increasing inorganic N input but also by reduced DON production and/or enhanced DON decomposition. Accelerated DON decomposition might be attributable to less waterlogging and higher nutrient availability. Significantly increasing NO3-N concentrations and NO3-N/NH4-N concentration ratios in throughfall and litter leachate below the thick organic layers indicated increasing nitrification. In mineral soil solutions, in contrast, NH4-N concentrations increased and NO3-N/NH4-N concentration ratios decreased significantly, suggesting increasing net ammonification. Our results demonstrate that the remote tropical montane forests on the rim of the Amazon basin experienced a pronounced change of the N cycle in only one decade. This change likely parallels a similar change which followed industrialization in the temperate zone of the Northern Hemisphere more than a century ago.

Transient simulations of the carbon and nitrogen dynamics in northern peatlands: from the Last Glacial Maximum to the 21st century

The development of northern high-latitude peatlands played an important role in the carbon (C) balance of the land biosphere since the Last Glacial Maximum (LGM). At present, carbon storage in northern peatlands is substantial and estimated to be 500 ± 100 Pg C (1 Pg C = 1015 g C). Here, we develop and apply a peatland module embedded in a dynamic global vegetation and land surface process model (LPX-Bern 1.0). The peatland module features a dynamic nitrogen cycle, a dynamic C transfer between peatland acrotelm (upper oxic layer) and catotelm (deep anoxic layer), hydrology- and temperature-dependent respiration rates, and peatland specific plant functional types. Nitrogen limitation down-regulates average modern net primary productivity over peatlands by about half. Decadal acrotelm-to-catotelm C fluxes vary between −20 and +50 g C m−2 yr−1 over the Holocene. Key model parameters are calibrated with reconstructed peat accumulation rates from peat-core data. The model reproduces the major features of the peat core data and of the observation-based modern circumpolar soil carbon distribution. Results from a set of simulations for possible evolutions of northern peat development and areal extent show that soil C stocks in modern peatlands increased by 365–550 Pg C since the LGM, of which 175–272 Pg C accumulated between 11 and 5 kyr BP. Furthermore, our simulations suggest a persistent C sequestration rate of 35–50 Pg C per 1000 yr in present-day peatlands under current climate conditions, and that this C sink could either sustain or turn towards a source by 2100 AD depending on climate trajectories as projected for different representative greenhouse gas concentration pathways.

Land use change and nitrogen feedbacks constrain the trajectory of the land carbon sink

Our understanding of Earth's carbon climate system depends critically upon interactions between rising atmospheric CO2, changing land use, and nitrogen limitation on vegetation growth. Using a global land model, we show how these factors interact locally to generate the global land carbon sink over the past 200 years. Nitrogen constraints were alleviated by N2 fixation in the tropics and by atmospheric nitrogen deposition in extratropical regions. Nonlinear interactions between land use change and land carbon and nitrogen cycling originated from three major mechanisms: (i) a sink foregone that would have occurred without land use conversion; (ii) an accelerated response of secondary vegetation to CO2 and nitrogen, and (iii) a compounded clearance loss from deforestation. Over time, these nonlinear effects have become increasingly important and reduce the present-day net carbon sink by ~40% or 0.4 PgC yr−1.

Evidence from the real world: N-15 natural abundances reveal enhanced nitrogen use at high plant diversity in Central European grasslands

Complementarity that leads to more efficient resource use is presumed to be a key mechanism explaining positive biodiversity–productivity relationships but has been described solely for experimental set-ups with controlled environmental settings or for very short gradients of abiotic conditions, land-use intensity and biodiversity. Therefore, we analysed plant diversity effects on nitrogen dynamics across a broad range of Central European grasslands. The 15N natural abundance in soil and plant biomass reflects the net effect of processes affecting ecosystem N dynamics. This includes the mechanism of complementary resource utilization that causes a decrease in the 15N isotopic signal. We measured plant species richness, natural abundance of 15N in soil and plants, above-ground biomass of the community and three single species (an herb, grass and legume) and a variety of additional environmental variables in 150 grassland plots in three regions of Germany. To explore the drivers of the nitrogen dynamics, we performed several analyses of covariance treating the 15N isotopic signals as a function of plant diversity and a large set of covariates. Increasing plant diversity was consistently linked to decreased δ15N isotopic signals in soil, above-ground community biomass and the three single species. Even after accounting for multiple covariates, plant diversity remained the strongest predictor of δ15N isotopic signals suggesting that higher plant diversity leads to a more closed nitrogen cycle due to more efficient nitrogen use. Factors linked to increased δ15N values included the amount of nitrogen taken up, soil moisture and land-use intensity (particularly fertilization), all indicators of the openness of the nitrogen cycle due to enhanced N-turnover and subsequent losses. Study region was significantly related to the δ15N isotopic signals indicating that regional peculiarities such as former intensive land use could strongly affect nitrogen dynamics. Synthesis. Our results provide strong evidence that the mechanism of complementary resource utilization operates in real-world grasslands where multiple external factors affect nitrogen dynamics. Although single species may differ in effect size, actively increasing total plant diversity in grasslands could be an option to more effectively use nitrogen resources and to reduce the negative environmental impacts of nitrogen losses.