Assessing critical load exceedances and ecosystem impacts of anthropogenic nitrogen and sulphur deposition at unmanaged forested catchments in Europe.

Anthropogenic emissions of nitrogen (N) and sulphur (S) compounds and their long-range transport have caused widespread negative impacts on different ecosystems. Critical loads (CLs) are deposition thresholds used to describe the sensitivity of ecosystems to atmospheric deposition. The CL methodology has been a key science-based tool for assessing the environmental consequences of air pollution. We computed CLs for eutrophication and acidification using a European long-term dataset of intensively studied forested ecosystem sites (n = 17) in northern and central Europe. The sites belong to the ICP IM and eLTER networks. The link between the site-specific calculations and time-series of CL exceedances and measured site data was evaluated using long-term measurements (1990-2017) for bulk deposition, throughfall and runoff water chemistry. Novel techniques for presenting exceedances of CLs and their temporal development were also developed. Concentrations and fluxes of sulphate, total inorganic nitrogen (TIN) and acidity in deposition substantially decreased at the sites. Decreases in S deposition resulted in statistically significant decreased concentrations and fluxes of sulphate in runoff and decreasing trends of TIN in runoff were more common than increasing trends. The temporal developments of the exceedance of the CLs indicated the more effective reductions of S deposition compared to N at the sites. There was a relation between calculated exceedance of the CLs and measured runoff water concentrations and fluxes, and most sites with higher CL exceedances showed larger decreases in both TIN and H+ concentrations and fluxes. Sites with higher cumulative exceedance of eutrophication CLs (averaged over 3 and 30 years) generally showed higher TIN concentrations in runoff. The results provided evidence on the link between CL exceedances and empirical impacts, increasing confidence in the methodology used for the European-scale CL calculations. The results also confirm that emission abatement actions are having their intended effects on CL exceedances and ecosystem impacts.

Logging residue piles of Norway spruce, Scots pine and silver birch in a clear-cut: Effects on nitrous oxide emissions and soil percolate water nitrogen.

We analysed how logging residue (LR) piles of common tree species in Finland, Norway spruce (Picea abies (L.) H. Karst.), Scots pine (Pinus sylvestris L.) and silver birch (Betula pendula Roth), affect nitrogen (N) losses in forest soil after final felling. A Norway spruce dominated stand was clear-cut and followed by two experimental setups to study the nitrous oxide (N2O) emissions and leaching of carbon (C) and N. Experiments consisted of four treatments: tree species treatments consisting of 40 kg m-2 of LR and a control treatment without residues. The C losses were monitored as dissolved organic carbon (DOC), the N losses as ammonium (NH4-N), nitrate (NO3-N) and dissolved organic nitrogen (DON) fluxes and concentrations in soil percolation waters and the N2O emissions as fluxes from the forest soil to the atmosphere. In addition the soil temperatures, the molecular size distribution of the DOC from the soil percolation waters and the origin of the N2O production were determined. The LR piles lowered the soil temperatures and, especially those of birch, increased the concentrations of NO3-N in the soil percolation waters already 1 year after the establishment of the piles. The LR piles increased the NH4-N concentrations. The smallest molecular size fraction (<1 kD) of DOC predominated in all treatments. The N2O fluxes peaked under the piles during the second and third growing seasons; however, the inconsistent fluxes tended to be low. The production of N2O was driven by both nitrification and denitrification processes, the proportion depending on the tree species. Our results indicate that LR piles accelerate N losses 1 year after the clear-cutting, especially NO3-N, which predominates in the soil percolation waters under the birch residues, whereas spruce residues tend to stimulate N2O emissions longer. These results have implications for sustainable forest management practices and nutrition of regrowing vegetation.

Forest mosses sensitively indicate nitrogen deposition in boreal background areas.

Mosses take up nitrogen (N) mainly from precipitation through their surfaces, which makes them competent bioindicators of N deposition. We found positive relationships between the total N concentration (mossN%) of common terrestrial moss species (feather mosses Pleurozium schreberi and Hylocomium splendens, and a group of Dicranum species) and different forms of N deposition in 11-16 coniferous forests with low N deposition load in Finland. The mosses were collected either inside (Dicranum group) or both inside and outside (feather mosses) the forests. Deposition was monitored in situ as bulk deposition (BD) and stand throughfall (TF) and detected for ammonium (NH4+-N), nitrate (NO3--N), dissolved organic N (DON), and total N (Ntot, kg ha-1yr-1). Ntot deposition was lower in TF than BD indicating that tree canopies absorbed N from deposition in N limited boreal stands. However, mossN% was higher inside than outside the forests. In regression equations, inorganic N in BD predicted best the mossN% in openings, while DON in TF explained most variation of mossN% in forests. An asymptotic form of mossN% vs. TF Ntot curves in forests and free NH4+-N accumulation in tissues in the southern plots suggested mosses were near the N saturation state already at the Ntot deposition level of 3-5 kg ha-1yr-1. N leachate from ground litterfall apparently also contributed the N supply of mosses. Our study yielded new information on the sensitivity of boreal mosses to low N deposition and their response to different N forms in canopy TF entering moss layer. The equations predicting the Ntot deposition with mossN% showed a good fit both in forest sites and openings, especially in case of P. schreberi. However, the open site mossN% is a preferable predictor of N deposition in monitoring studies to minimize the effect of tree canopies and N leachate from litterfall on the estimates.

N2 fixation associated with the bryophyte layer is suppressed by low levels of nitrogen deposition in boreal forests.

Biological fixation of atmospheric nitrogen (N2) by bryophyte-associated cyanobacteria is an important source of plant-available N in the boreal biome. Information on the factors that drive biological N2 fixation (BNF) rates is needed in order to understand the N dynamics of forests under a changing climate. We assessed the potential of several cryptogam species (the feather mosses Hylocomium splendens and Pleurozium schreberi, a group of Dicranum bryophytes, two liverworts, and Cladina lichens) to serve as associates of cyanobacteria or other N2-fixing bacteria (diazotrophs) using acetylene reduction assay (ARA). We tested the hypotheses that the legacy of chronic atmospheric N deposition reduces BNF in the three bryophyte species, sampled from 12 coniferous forests located at latitudes 60-68° N in Finland. In addition, we tested the effect of moisture and temperature on BNF. All species studied showed a BNF signal in the north, with the highest rates in feather mosses. In moss samples taken along the north-south gradient with an increasing N bulk deposition from 0.8 to 4.4 kg ha-1 year-1, we found a clear decrease in BNF in both feather mosses and Dicranum group. BNF turned off at N deposition of 3-4 kg ha-1 year-1. Inorganic N (NH4-N + NO3-N) best predicted the BNF rate among regression models with different forms of N deposition as explanatory variables. However, in southern spruce stands, tree canopies modified the N in throughfall so that dissolved organic N (DON) leached from canopies compensated for inorganic N retained therein. Here, both DON and inorganic N negatively affected BNF in H. splendens. In laboratory experiments, BNF increased with increasing temperature and moisture. Our results suggest that even relatively low N deposition suppresses BNF in bryophyte-associated diazotrophs. Further, BNF could increase in northern low-deposition areas, especially if climate warming leads to moister conditions, as predicted.

Long-term changes (1990-2015) in the atmospheric deposition and runoff water chemistry of sulphate, inorganic nitrogen and acidity for forested catchments in Europe in relation to changes in emissions and hydrometeorological conditions.

The international Long-Term Ecological Research Network (ILTER) encompasses hundreds of long-term research/monitoring sites located in a wide array of ecosystems that can help us understand environmental change across the globe. We evaluated long-term trends (1990-2015) for bulk deposition, throughfall and runoff water chemistry and fluxes, and climatic variables in 25 forested catchments in Europe belonging to the UNECE International Cooperative Programme on Integrated Monitoring of Air Pollution Effects on Ecosystems (ICP IM). Many of the IM sites form part of the monitoring infrastructures of this larger ILTER network. Trends were evaluated for monthly concentrations of non-marine (anthropogenic fraction, denoted as x) sulphate (xSO4) and base cations x(Ca+Mg), hydrogen ion (H+), inorganic N (NO3 and NH4) and ANC (Acid Neutralising Capacity) and their respective fluxes into and out of the catchments and for monthly precipitation, runoff and air temperature. A significant decrease of xSO4 deposition resulted in decreases in concentrations and fluxes of xSO4 in runoff, being significant at 90% and 60% of the sites, respectively. Bulk deposition of NO3 and NH4 decreased significantly at 60-80% (concentrations) and 40-60% (fluxes) of the sites. Concentrations and fluxes of NO3 in runoff decreased at 73% and 63% of the sites, respectively, and NO3 concentrations decreased significantly at 50% of the sites. Thus, the LTER/ICP IM network confirms the positive effects of the emission reductions in Europe. Air temperature increased significantly at 61% of the sites, while trends for precipitation and runoff were rarely significant. The site-specific variation of xSO4 concentrations in runoff was most strongly explained by deposition. Climatic variables and deposition explained the variation of inorganic N concentrations in runoff at single sites poorly, and as yet there are no clear signs of a consistent deposition-driven or climate-driven increase in inorganic N exports in the catchments.

Modelling study of soil C, N and pH response to air pollution and climate change using European LTER site observations.

Current climate warming is expected to continue in coming decades, whereas high N deposition may stabilize, in contrast to the clear decrease in S deposition. These pressures have distinctive regional patterns and their resulting impact on soil conditions is modified by local site characteristics. We have applied the VSD+ soil dynamic model to study impacts of deposition and climate change on soil properties, using MetHyd and GrowUp as pre-processors to provide input to VSD+. The single-layer soil model VSD+ accounts for processes of organic C and N turnover, as well as charge and mass balances of elements, cation exchange and base cation weathering. We calibrated VSD+ at 26 ecosystem study sites throughout Europe using observed conditions, and simulated key soil properties: soil solution pH (pH), soil base saturation (BS) and soil organic carbon and nitrogen ratio (C:N) under projected deposition of N and S, and climate warming until 2100. The sites are forested, located in the Mediterranean, forested alpine, Atlantic, continental and boreal regions. They represent the long-term ecological research (LTER) Europe network, including sites of the ICP Forests and ICP Integrated Monitoring (IM) programmes under the UNECE Convention on Long-range Transboundary Air Pollution (LRTAP), providing high quality long-term data on ecosystem response. Simulated future soil conditions improved under projected decrease in deposition and current climate conditions: higher pH, BS and C:N at 21, 16 and 12 of the sites, respectively. When climate change was included in the scenario analysis, the variability of the results increased. Climate warming resulted in higher simulated pH in most cases, and higher BS and C:N in roughly half of the cases. Especially the increase in C:N was more marked with climate warming. The study illustrates the value of LTER sites for applying models to predict soil responses to multiple environmental changes.

Adaptive root foraging strategies along a boreal-temperate forest gradient.

The tree root-mycorhizosphere plays a key role in resource uptake, but also in the adaptation of forests to changing environments. The adaptive foraging mechanisms of ectomycorrhizal (EcM) and fine roots of Picea abies, Pinus sylvestris and Betula pendula were evaluated along a gradient from temperate to subarctic boreal forest (38 sites between latitudes 48°N and 69°N) in Europe. Variables describing tree resource uptake structures and processes (absorptive fine root biomass and morphology, nitrogen (N) concentration in absorptive roots, extramatrical mycelium (EMM) biomass, community structure of root-associated EcM fungi, soil and rhizosphere bacteria) were used to analyse relationships between root system functional traits and climate, soil and stand characteristics. Absorptive fine root biomass per stand basal area increased significantly from temperate to boreal forests, coinciding with longer and thinner root tips with higher tissue density, smaller EMM biomass per root length and a shift in soil microbial community structure. The soil carbon (C) : N ratio was found to explain most of the variability in absorptive fine root and EMM biomass, root tissue density, N concentration and rhizosphere bacterial community structure. We suggest a concept of absorptive fine root foraging strategies involving both qualitative and quantitative changes in the root-mycorrhiza-bacteria continuum along climate and soil C : N gradients.

Spatially valid data of atmospheric deposition of heavy metals and nitrogen derived by moss surveys for pollution risk assessments of ecosystems.

For analysing element input into ecosystems and associated risks due to atmospheric deposition, element concentrations in moss provide complementary and time-integrated data at high spatial resolution every 5 years since 1990. The paper reviews (1) minimum sample sizes needed for reliable, statistical estimation of mean values at four different spatial scales (European and national level as well as landscape-specific level covering Europe and single countries); (2) trends of heavy metal (HM) and nitrogen (N) concentrations in moss in Europe (1990-2010); (3) correlations between concentrations of HM in moss and soil specimens collected across Norway (1990-2010); and (4) canopy drip-induced site-specific variation of N concentration in moss sampled in seven European countries (1990-2013). While the minimum sample sizes on the European and national level were achieved without exception, for some ecological land classes and elements, the coverage with sampling sites should be improved. The decline in emission and subsequent atmospheric deposition of HM across Europe has resulted in decreasing HM concentrations in moss between 1990 and 2010. In contrast, hardly any changes were observed for N in moss between 2005, when N was included into the survey for the first time, and 2010. In Norway, both, the moss and the soil survey data sets, were correlated, indicating a decrease of HM concentrations in moss and soil. At the site level, the average N deposition inside of forests was almost three times higher than the average N deposition outside of forests.

Relevance of canopy drip for the accumulation of nitrogen in moss used as biomonitors for atmospheric nitrogen deposition in Europe.

High atmospheric deposition of nitrogen (N) impacts functions and structures of N limited ecosystems. Due to filtering and related canopy drip effects forests are particularly exposed to N deposition. Up to now, this was proved by many studies using technical deposition samplers but there are only some few studies analysing the canopy drip effect on the accumulation of N in moss and related small scale atmospheric deposition patterns. Therefore, we investigated N deposition and related accumulation of N in forests and in (neighbouring) open fields by use of moss sampled across seven European countries. Sampling and chemical analyses were conducted according to the experimental protocol of the European Moss Survey. The ratios between the measured N content in moss sampled inside and outside of forests were computed and used to calculate estimates for non-sampled sites. Potentially influencing environmental factors were integrated in order to detect their relationships to the N content in moss. The overall average N content measured in moss was 20.0mgg(-1) inside and 11.9mgg(-1) outside of forests with highest N values in Germany inside of forests. Explaining more than 70% of the variance, the multivariate analyses confirmed that the sampling site category (site with/without canopy drip) showed the strongest correlation with the N content in moss. Spatial variances due to enhanced dry deposition in vegetation stands should be considered in future monitoring and modelling of atmospheric N deposition.

Relationship between site-specific nitrogen concentrations in mosses and measured wet bulk atmospheric nitrogen deposition across Europe.

To assess the relationship between nitrogen concentrations in mosses and wet bulk nitrogen deposition or concentrations in precipitation, moss tissue and deposition were sampled within a distance of 1 km of each other in seven European countries. Relationships for various forms of nitrogen appeared to be asymptotic, with data for different countries being positioned at different locations along the asymptotic relationship and saturation occurring at a wet bulk nitrogen deposition of ca. 20 kg N ha(-1) yr(-1). The asymptotic behaviour was more pronounced for ammonium-N than nitrate-N, with high ammonium deposition at German sites being most influential in providing evidence of the asymptotic behaviour. Within countries, relationships were only significant for Finland and Switzerland and were more or less linear. The results confirm previous relationships described for modelled total deposition. Nitrogen concentration in mosses can be applied to identify areas at risk of high nitrogen deposition at European scale.

Long-term changes in acidity and DOC in throughfall and soil water in Finnish forests.

The main objective of this study was to examine if any detectable trends in dissolved organic carbon (DOC), sulphate (SO4-S) concentrations and acid neutralizing capacity (ANC) in throughfall (TF) and soil water (SW) could be found during 1990-2010 and to relate them to recent changes in decreased acid deposition. The study was conducted in seven boreal coniferous forest sites: four of which are managed and three unmanaged forests sites. Generally, temporal trend showed a significant decrease in SO4-S concentrations in bulk precipitation (BP), TF and SW. At some of the sites, there was an increasing tendency in BP and TF in the DOC concentrations. This feature coincides with decreasing SO4-S concentration, indicating that SO4-S may be an important driver of DOC release from the canopy. However, a slightly increased temperature, larger senescing needle mass and consequently increased decaying activity in the canopy may partly explain the increasing trend in DOC. In SW, no consistent DOC trend was seen. At some sites, the decreased base cation concentrations mostly account for the decrease in the ANC values in SW and TF.

Artificial recharge of groundwater through sprinkling infiltration: impacts on forest soil and the nutrient status and growth of Scots pine.

We studied the chemical changes in forest soil and the effects on Scots pine trees caused by continuous sprinkling infiltration over a period of two years, followed by a recovery period of two years. Infiltration increased the water input onto the forest soil by a factor of approximately 1000. After one year of infiltration, the pH of the organic layer had risen from about 4.0 to 6.7. The NH(4)-N concentration in the organic layer increased, most probably due to the NH(4) ions in the infiltration water, as the net N mineralization rate did not increase. Sprinkling infiltration initiated nitrification in the mineral soil. Macronutrient concentrations generally increased in the organic layer and mineral soil. An exception, however, was the concentration of extractable phosphorus, which decreased strongly during the infiltration period and did not show a recovery within two years. The NO(3)-N and K concentrations had reverted back to their initial level during the two-year recovery period, while the concentrations of Ca, Mg and NH(4)-N were still elevated. Nutrient concentrations in the pine needles increased on the infiltrated plots. However, the needle P concentration increased, despite the decrease in plant-available P in the soil. Despite the increase in the nutrient status, there were some visible signs of chlorosis in the current-year needles after two years of infiltration. The radial growth of the pines more than doubled on the infiltrated plots, which suggests that the very large increase in the water input onto the forest floor had no adverse effect on the functioning of the trees. However, a monitoring period of four years is not sufficient for detecting potential long term detrimental effects on forest trees.

Ectomycorrhizal root tips in relation to site and stand characteristics in Norway spruce and Scots pine stands in boreal forests.

Variations in ectomycorrhizal (EcM) short root tips of Norway spruce (Picea abies (L.) Karst.) and Scots pine (Pinus sylvestris L.) in 16 stands throughout Finland were studied, and their relationships with latitude, organic layer C:N ratio, temperature sum and foliage biomass were determined. There were no significant differences in EcM root tip frequency (number per milligram of fine roots) or root tip mass between tree species or between northern and southern sites. The EcM root tip number per unit area of the organic layer plus the 0-30 cm mineral soil layer varied between 0.8 and 2.4 million per m(2) for Norway spruce and between 0.7 and 2.9 million per m(2) for Scots pine, and it was higher in the northern Scots pine stands than in the southern Scots pine stands. Over 80% of the EcM root tips of both species were in the organic layer and the upper 0-10 cm mineral soil layer. We related EcM root tips to foliage mass because these two components are the most important functional units in boreal tree physiology. Both species, especially the Scots pine trees, had more EcM root tips in relation to foliage mass in northern Finland than in southern Finland. Scots pine trees had more EcM root tips in relation to foliage mass than Norway spruce in the same climatic region. The EcM root tip:foliage biomass ratio of Norway spruce was positively related to the C:N ratio in the organic layer, whereas that of Scots pine was negatively related to the temperature sum. The number of EcM root tips per milligram of fine root biomass was constant, implying that trees of both species increase nutrient uptake by increasing fine root production and hence their total number of EcM tips and the area of soil occupied by mycelia. Both tree species responded to nitrogen (N) deficiency by maintaining more EcM tips per foliage unit, and this may be related to a higher proportion of N uptake in an organic form.

Changes in dissolved organic carbon during artificial recharge of groundwater in a forested esker in Southern Finland.

Sprinkling infiltration in a forested esker leading to artificial recharge of groundwater was studied in Southern Finland. Changes in dissolved organic carbon (DOC) and the molecular size distribution and chemical properties of the organic carbon were investigated during the infiltration process. Artificial groundwater was produced using sprinkling infiltration directly onto the forest floor. One result of lake water infiltration through the organic horizon and I m thick mineral soil layer was a slight net increase in the DOC concentrations from 9.4 mg/L in the infiltration water to 13.2 mg/ L in percolation water. This indicates that the forest soil represents a potential input of organic matter into infiltration water. However, the DOC concentrations decreased by 27-38% as the infiltration water percolated down through the unsaturated soil layer into the groundwater zone. At a distance of 1450 m from the infiltration area, the mean DOC concentration in the groundwater was below the recommended value for drinking water in Finland of 2.0 mg/L. There was a strong reduction in the concentrations of hydrophilic and hydrophobic acids, but only a slight decrease in hydrophilic neutral organic compounds during the infiltration process. The DOC in the production well consisted of low molecular size fractions. Larger molecular size fractions were removed effectively from the water during the infiltration process.