Role of six European tree species and land‐use legacy for nitrogen and water budgets in forests

Water and nutrient fluxes for single stands of different tree species have been reported in numerous studies, but comparative studies of nutrient and hydrological budgets of common European deciduous tree species are rare. Annual fluxes of water and inorganic nitrogen (N) were established in a 30‐year‐old common garden design with stands of common ash (Fraxinus excelsior), European beech (Fagus sylvatica L.), pedunculate oak (Quercus robur), small‐leaved lime (Tilia cordata Mill.), sycamore maple (Acer pseudoplatanus) and Norway spruce (Picea abies [L.] Karst.) replicated at two sites in Denmark, Mattrup and Vallø during 2 years. Mean annual percolation below the root zone (mm yr^?1±SE, n=4) ranked in the following order: maple (351±38)>lime (284±32), oak (271±25), beech (257±30), ash (307±69)? spruce (75±24). There were few significant tree species effects on N fluxes. However, the annual mean N throughfall flux (kg N ha^?1 yr^?1±SE, n=4) for spruce (28±2) was significantly larger than for maple (12±1), beech (11±1) and oak (9±1) stands but not different from that of lime (15±3). Ash had a low mean annual inorganic N throughfall deposition of 9.1 kg ha^?1, but was only present at Mattrup. Annual mean of inorganic N leaching (kg ha^?1 yr^?1±SE, n=4) did not differ significantly between species despite of contrasting tree species mean values; beech (25±9)>oak (16±10), spruce (15±8), lime (14±8)? maple (1.9±1), ash (2.0±1). The two sites had similar throughfall N fluxes, whereas the annual leaching of N was significantly higher at Mattrup than at Vallø. Accordingly, the sites differed in soil properties in relation to rates and dynamics of N cycling. We conclude that tree species affect the N cycle differently but the legacy of land use exerted a dominant control on the N cycle within the short‐term perspective (30 years) of these stands.     

Nitrogen input together with ecosystem nitrogen enrichment predict nitrate leaching from European forests

The IFEF database (Indicators of Forest Ecosystem Functioning), consisting of nitrogen deposition, nitrate leaching fluxes, and soil and ecosystem characteristics, is analysed to evaluate the C/N ratio in the organic horizon as an indicator of nitrate leaching. One hundred and eighty one forests are examined, from countries across Europe ranging from boreal to Mediterranean regions, encompassing broadleaf and coniferous sites and plot and catchment studies. N input in throughfall ranges from less than 1 kg N ha^?1 y^?1 in northern Norway and Finland to greater than 60 kg N ha^?1 y^?1 in the Netherlands and Czech Republic. The amount of NO₃^– leached covers a smaller range, between 1 and 40 kg N ha^?1 y^?1. Nitrate leaching is strongly dependent on the amount of nitrogen deposited in throughfall (N input) and simply adding the C/N ratio in the organic horizon to a regression equation does not improve this relationship. However, when the data are stratified based on C/N ratios less than or equal to 25 and greater than 25, highly significant relationships (P < 0.05) are observed between N input and NO₃^– leached. The slope of the relationship for those sites where C/N ratio is ≤ 25 (′nitrogen enriched′ sites) is twice that for those sites where C/N ratio is > 25. These empirical relationships may be used to identify which forested ecosystems are likely to show elevated rates of nitrate leaching under predicted future nitrogen deposition scenarios. Elevated NO₃^– leaching also shows a relationship with soil pH, with high rates of NO₃^– leaching only observed at sites with a pH < 4.5 and N inputs > 30 kg N ha^?1 y^?1. Tree age and species have no significant impact on the ecosystem response to N input at a regional scale.     

N deposition, N transformation and N leaching in acid forest soils

Nitrogen deposition, mineralisation, uptake and leaching were measured on a monthly basis in the field during 2 years in six forested stands on acidic soils under mountainous climate. Studies were conducted in three Douglas-fir [Pseudotsuga menziesii (Mirb.) Franco] plantations (D20: 20 year; D40: 40 yr; D60: 60 yr) on abandoned croplands in the Beaujolais Mounts; and two spruce (Picea abies Karst.) plantations (S45: 45 yr; S90: 90 yr) and an old beech (Fagus sylvatica L.) stand (B150: 150 yr) on ancient forest soils in a small catchment in the Vosges Mountains. N deposition in throughfall varied between 7–8 kg ha^–1 year^–1 (D20, B150, S45) and 15–21 kg ha^–1 yr^–1 (S90, D40, D60). N in annual litterfall varied between 20–29 kg ha^–1 (D40, D60, S90), and 36–43 kg ha^–1 (D20, S45, B150). N leaching below root depth varied among stands within a much larger range, between 1–9 kg ha^–1 yr^–1 (B150, S45, D60) and 28–66 kg ha^–1 yr^–1 (D40, S90, D20), with no simple relationship with N deposition, or N deposition minus N storage in stand biomass. N mineralisation was between 57–121 kg ha^–1 yr^–1 (S45, D40, S90) and between 176–209 kg ha^–1 yr^–1 in (B150, D60 and D20). The amounts of nitrogen annually mineralised and nitrified were positively related. Neither general soil parameters, such as pH, soil type, base saturation and C:N ratio, nor deposition in throughfall or litterfall were simply related to the intensity of mineralisation and/or nitrification. When root uptake was not allowed, nitrate leaching increased by 11 kg ha^–1 yr^–1 at S45, 36 kg ha^–1 yr^–1 at S90 and between 69 and 91 kg ha^–1 yr^–1 at D20, D40, B150 and D60, in relation to the nitrification rates of each plot. From this data set and recent data from the literature, we suggest that: high nitrification and nitrate leaching in Douglas-fir soils was likely related to the former agricultural land use. High nitrification rate but very low nitrate leaching in the old beech soil was related to intense recycling of mineralised N by beech roots. Medium nitrification and nitrate leaching in the old spruce stand was related to the average level of N deposition and to the deposition and declining health of the stand. Very low nitrification and N leaching in the young spruce stand were considered representative of fast growing spruce plantations receiving low N deposition on acidic soils of ancient coniferous forests. Consequently, we suggest that past land use and fine root cycling (which is dependent on to tree species and health) should be taken into account to explain the variability in the relation between N deposition and leaching in forests.     

Changes in soil properties after afforestation of former intensively managed soils with oak and Norway spruce

In many European countries, surplus agricultural production and ecological problems due to intensive soil cultivation have increased the interest in afforestation of arable soils. Many environmental consequences which might rise from this alternative land-use are only known from forest establishment on less intensively managed or marginal soils. The present study deals with changes in soil properties following afforestation of nutrient-rich arable soils. A chronosequence study was carried out comprising seven Norway spruce (Picea abies (Karst.) L.) and seven oak (Quercus robur L.) stands established from 1969 to 1997 on former horticultural and agricultural soils in the vicinity of Copenhagen, Denmark. For comparison, a permanent pasture and a ca. 200-year-old mixed deciduous forest were included. This paper reports on changes in pH values, base saturation (BS_eff), exchangeable calcium, soil N pools (N_min contents), and C/N ratios in the Ap-horizon (0–25 cm) and the accumulated forest floor. The results suggest that afforestation slowly modifies soil properties of former arable soils. Land-use history seems to influence soil properties more than the selected tree species. An effect of tree species was only found in the forest floor parameters. Soil acidification was the most apparent change along the chronosequence in terms of a pH decrease from 6 to 4 in the upper 5 cm soil. Forest floor pH varied only slightly around 5. Nitrogen storage in the Ap-horizon remained almost constant at 5.5 Mg N ha^–1. This was less than in the mineral soil of the ca. 200-year-old forest. In the permanent pasture, N storage was somewhat higher in 0–15 cm depth than in afforested stands of comparable age. Nitrogen storage in the forest floor of the 0–30-year-old stands increased in connection with the build-up of forest floor mass. The increase was approximately five times greater under spruce than oak. Mineral soil C/N ratios ranged from 10 to 15 in all stands and tended to increase in older stands only in 0–5 cm depth. Forest floor C/N ratios were higher in spruce stands (26.4) as compared to oak stands (22.7). All stands except the youngest within a single tree species had comparable C/N ratios.     

Changes in nitrogen cycling and retention processes in soils under spruce forests along a nitrogen enrichment gradient in Germany

A network of long-term monitoring sites on nitrogen (N) input and output of forests across Germany showed that a number of Germany's forests are subject to or are experiencing N saturation and that spruce (Picea abies) stands have high risk. Our study was aimed at (1) quantifying the changes in gross rates of microbial N cycling and retention processes in forest soils along an N enrichment gradient and (2) relating the changes in soil N dynamics to N losses. We selected spruce sites representing an N enrichment gradient (indicated by leaching : throughfall N ratios) ranging from 0.04–0.13 (low N),≤0.26 (intermediate N enrichment) to≥0.42 (highly N enriched). To our knowledge, our study is the first to report on mechanistic changes in gross rates of soil N cycling and abiotic NO₃⁻ retention under ambient N enrichment gradient. Gross N mineralization, NH₄⁺ immobilization, gross nitrification, and NO₃⁻ immobilization rates increased up to intermediate N enrichment level and somewhat decreased at highly N-enriched condition. The turnover rates of NH₄⁺ and microbial N pools increased while the turnover rates of the NO₃⁻ pool decreased across the N enrichment gradient. Abiotic immobilization of NH₄⁺ did not differ across sites and was lower than that of NO₃⁻. Abiotic NO₃⁻ immobilization decreased across the N enrichment gradient. Microbial assimilation and turnover appeared to contribute largely to the retention of NH₄⁺. The increasing NO₃⁻ deposition and decreasing turnover rates of the NO₃⁻ pool, combined with decreasing abiotic NO₃⁻ retention, possibly contributed to increasing NO₃⁻ leaching and gaseous emissions across the N enrichment gradient. The empirical relationships of changes in microbial N cycling across the N enrichment gradient may be integrated in models used to predict responses of forest ecosystems (e.g. spruce) to increasing N deposition.     

Exchange of Proton and Major Elements in Two-Layer Canopies Under Acid Rain in a Subtropical Evergreen Forest in Central-South China

Canopy exchanges of H^＋ and N （NH4^＋-N, NO3^--N） and other major ions were evaluated and quantified In twolayer canopies based on throughfall measurements in Shaoshan Forest during the period 2000-2002, central-south China, The collected annual rainfall, throughfall, and sub-throughfall were 1 401, 1 191, and 1 084 mm/year, respectively. Fifteen percent and 8% of rainfall （or 9% of throughfall） were intercepted by the top canopy and sub-canopy layers, respectively, The foliar leaching of base cations from the top canopy was significantly higher than that from the sub-canopy, and the latter accounted for 25% of the former. The uptake of H^＋ and NH4^＋ was significantly higher in the top canopy than in the sub-canopy, indicating that the canopy buffering capacity in the top canopy was stronger than the sub-canopy; Mg^2＋ can be absorbed from water flux on the sub-canopy foliar surfaces to compensate for the Mg deficit in the forest soil during the growing season,     

Nitrogen cycling in canopy soils of tropical montane forests responds rapidly to indirect N and P fertilization

Although the canopy can play an important role in forest nutrient cycles, canopy‐based processes are often overlooked in studies on nutrient deposition. In areas of nitrogen (N) and phosphorus (P) deposition, canopy soils may retain a significant proportion of atmospheric inputs, and also receive indirect enrichment through root uptake followed by throughfall or recycling of plant litter in the canopy. We measured net and gross rates of N cycling in canopy soils of tropical montane forests along an elevation gradient and assessed indirect effects of elevated nutrient inputs to the forest floor. Net N cycling rates were measured using the buried bag method. Gross N cycling rates were measured using ¹⁵N pool dilution techniques. Measurements took place in the field, in the wet and dry season, using intact cores of canopy soil from three elevations (1000, 2000 and 3000 m). The forest floor had been fertilized biannually with moderate amounts of N and P for 4 years; treatments included control, N, P, and N + P. In control plots, gross rates of NH₄⁺ transformations decreased with increasing elevation; gross rates of NO₃^? transformations did not exhibit a clear elevation trend, but were significantly affected by season. Nutrient‐addition effects were different at each elevation, but combined N + P generally increased N cycling rates at all elevations. Results showed that canopy soils could be a significant N source for epiphytes as well as contributing up to 23% of total (canopy + forest floor) mineral N production in our forests. In contrast to theories that canopy soils are decoupled from nutrient cycling in forest floor soil, N cycling in our canopy soils was sensitive to slight changes in forest floor nutrient availability. Long‐term atmospheric N and P deposition may lead to increased N cycling, but also increased mineral N losses from the canopy soil system.     

The natural abundance of 15N in litter and soil profiles under six temperate tree species: N cycling depends on tree species traits and site fertility

Aims

We investigated the influence of tree species on the natural ¹⁵N abundance in forest stands under elevated ambient N deposition.

Methods

We analysed δ¹⁵N in litter, the forest floor and three mineral soil horizons along with ecosystem N status variables at six sites planted three decades ago with five European broadleaved tree species and Norway spruce.

Results

Litter δ¹⁵N and ¹⁵N enrichment factor (δ¹⁵N_litter–δ¹⁵N_soil) were positively correlated with N status based on soil and litter N pools, nitrification, subsoil nitrate concentration and forest growth. Tree species differences were also significant for these N variables and for the litter δ¹⁵N and enrichment factor. Litter from ash and sycamore maple with high N status and low fungal mycelia activity was enriched in ¹⁵N (+0.9 delta units) relative to other tree species (European beech, pedunculate oak, lime and Norway spruce) even though the latter species leached more nitrate.

Conclusions

The δ¹⁵N pattern reflected tree species related traits affecting the N cycling as well as site fertility and former land use, and possibly differences in N leaching. The tree species δ¹⁵N patterns reflected fractionation caused by uptake of N through mycorrhiza rather than due to nitrate leaching or other N transformation processes.     

Control of Nitrous Oxide Emissions in European Beech,Norway Spruce and Scots Pine Forests

Elevated nitrogen deposition has increased tree growth, the storage of soil organic matter, and nitrate leaching in many European forests, but little is known about the effect of tree species and nitrogen deposition on nitrous oxide emission. Here we report soil N₂O emission from European beech, Scots pine and Norway spruce forests in two study areas of Germany with distinct climate, N deposition and soils. N₂O emissions and throughfall input of nitrate and ammonium were measured biweekly during growing season and monthly during dormant season over a 28 months period. Annual N₂O emission rates ranged between 0.4 and 1.3 kg N ha^?1 year^?1 among the stands and were higher in 1998 than in 1999 due to higher precipitation during the growing season of 1998. A 2-way-ANOVA revealed that N₂O fluxes were significantly higher (p<0.001) at Solling than at Unterlüß while tree species had no effect on N₂O emissions. Soil texture and the amount of throughfall explained together 94% of the variance among the stands, indicating that increasing portions of silt and clay may promote the formation of N₂O in wet forest soils. Moreover, cumulative N₂O fluxes were significantly correlated (r² = 0.60, p<0.001) with cumulative NO ₃ ^? fluxes at 10 cm depth as an indicator of N saturation, however, the slope of the regression curve indicates a rather weak effect of NO ₃ ^? fluxes on N₂O emissions. N input by throughfall was not correlated with N₂O emissions and only 1.6–3.2% of N input was released as N₂O to the atmosphere. Our results suggest that elevated N inputs have little effect on N₂O emissions in beech, spruce and pine forests.     

Rainfall partitioning into throughfall and stemflow and associated nutrient fluxes: land use impacts in a lower montane tropical region of Panama

Land-use change alters catchment hydrology by influencing the quality and quantity of partitioned rainfall. We compared rainfall partitioning (throughfall, stemflow and interception) and nutrient concentrations in rainfall, throughfall and stemflow in three land-use types [primary forest (PF), secondary forest (SF) and agriculture (A)] in Panama. Measurements of throughfall were highly variable which may have masked seasonal and land use differences but it was clear that throughfall at agricultural sites made up a larger proportion of gross precipitation than at forest sites. Of incident precipitation, 94% became throughfall in agriculture sites while 83 and 81% of gross precipitation became throughfall in PF and SF, respectively. The size of the precipitation event was the main driver of variation in throughfall and stemflow. Consistent patterns in nutrient cycling were also difficult to identify. Vegetation has a vital role in delivering nutrients as throughfall deposition of K was often larger than precipitation deposition. A canopy budget model indicated that canopy exchange was often more dominant than dry deposition. Throughfall was generally enriched with nutrients, especially K and Mg, with enrichment factors of up to 17 and 5 for K and Mg, respectively, in PF. In contrast, Ca was sometimes taken up by the canopy. Values of nutrient deposition were high (with up to 15, 3, 30 and 15 kg ha^?1 month^?1 in stand deposition of Ca, Mg, K and Na, respectively in PF), possibly due to the slash-and-burn agricultural practices in the area or marine inputs. Throughfall and stemflow are vital sources of nutrients in these ecosystems.     

Effects of forest clear-cutting on the carbon and nitrogen fluxes through podzolic soil horizons

Effects of clear-cutting on the dissolved fluxes of organic C (DOC), organic N (DON), NO₃ ^– and NH₄ ⁺ through surface soil horizons were studied in a Norway spruce dominated mixed boreal forest in eastern Finland. Bulk deposition, total throughfall and soil water from below the organic (including understorey vegetation and, after clear-cutting, also logging residues), eluvial and illuvial horizons were sampled weekly from 1993 to 1999. Clear-cutting was carried out in September 1996. The removal of the tree canopy decreased the deposition of DOC and DON to the forest floor and increased that of NH₄ ⁺ and NO₃ ^– but did not affect the deposition of total N (DTN, <3 kg ha^–1 a^–1). The leaching of DOC and DON from the organic horizon increased over twofold after clear-cutting (fluxes were on an average 168 kg C and 3.3 kg N ha^–1 a^–1), but the increased outputs were effectively retained in the surface mineral soil horizons. Inorganic N deposition was mainly retained by the logging residues and organic horizon indicating microbial immobilization. Increased NO₃ ^– formation reflected as elevated concentrations in the percolate from below the mineral soil horizons were observed especially in the third year after clear-cutting. However, the changes were small and the increased leaching of DTN from below the illuvial horizon remained small (<0.4 kg ha^–1 a^–1) and mainly DON. Effects of clear-cutting on the transport of C and N to surface waters will probably be negligible.     

Afforestation effects on SOC in former cropland: oak and spruce chronosequences resampled after 13 years

Chronosequences are commonly used to assess soil organic carbon (SOC) sequestration after land‐use change, but SOC dynamics predicted by this space‐for‐time substitution approach have rarely been validated by resampling. We conducted a combined chronosequence/resampling study in a former cropland area (Vestskoven) afforested with oak (Quercus robur) and Norway spruce (Picea abies) over the past 40 years. The aims of this study were (i) to compare present and previous chronosequence trends in forest floor and top mineral soil (0–25 cm) C stocks; (ii) to compare chronosequence estimates with current rates of C stock change based on resampling at the stand level; (iii) to estimate SOC changes in the subsoil (25–50 cm); and (iv) to assess the influence of two tree species on SOC dynamics. The two chronosequence trajectories for forest floor C stocks revealed consistently higher rates of C sequestration in spruce than oak. The chronosequence trajectory was validated by resampling and current rates of forest floor C sequestration decreased with stand age. Chronosequence trends in topsoil SOC in 2011 did not differ significantly from those reported in 1998, however, there was a shift from a negative rate (1998: ?0.3 Mg C ha^?1 yr^?1) to no change in 2011. In contrast SOC stocks in the subsoil increased with stand age, however, not significantly (P = 0.1), suggesting different C dynamics in and below the former plough layer. Current rates of C change estimated by repeated sampling decreased with stand age in forest floors but increased in the topsoil. The contrasting temporal change in forest floor and mineral soil C sequestration rates indicate a shift in C source‐sink strength after approximately 40 years. We conclude that afforestation of former cropland within the temperate region may induce soil C loss during the first decades followed by a recovery phase of yet unknown duration.     

Large Canopy Exchange Fluxes of Inorganic and Organic Nitrogen and Preferential Retention of Nitrogen by Epiphytes in a Tropical Lowland Rainforest

Little is known about how tropical forest canopies interact with atmospheric nitrogen deposition and how this affects the internal nutrient dynamics and the processing of external nutrient inputs. The objectives of this study therefore were (1) to investigate gross and net canopy nitrogen (N) fluxes (retention and leaching) and (2) the effect of canopy components on net canopy N retention. Tracers were applied on detached branches in a tropical wet lowland rainforest, Costa Rica. A novel ¹⁵N pool dilution method showed that gross canopy fluxes (retention and leaching) of NO₃ ^?, NH₄ ⁺, and dissolved organic nitrogen (DON) were remarkably higher than net throughfall fluxes. Gross fluxes of NH₄ ⁺ and NO₃ ^? resulted in a negligible net flux whereas DON showed net uptake by the canopy. The highest quantity of ¹⁵N was recovered in epiphytic bryophytes (16.4%) although the largest biomass fraction was made up of leaves. The study demonstrates that tracer applications allow investigation of the dynamic and complex canopy exchange processes and that epiphytic communities play a major role in solute fluxes in tree canopies and therefore in the nutrient dynamics of tropical rain forests.     

Deterioration of Norway spruce vitality despite a sharp decline in acid deposition: a long‐term integrated perspective

Since the late 1970s, several long‐term ecological studies were conducted to better understand the biogeochemical functioning of Norway spruce stands in the Ardennes as these nutrient‐poor ecosystems were subject to high levels of acid deposition and exhibited symptoms of tree health decline. Between 1978 and 2009, acid deposition declined sharply, especially sulfur and to a lesser extent nitrogen deposition. The aim of this study was (i) to determine if the Norway spruce stands recovered after the reduction of acid deposition and (ii) to explain why such a recovery occurred or not. Therefore, we collected data from different projects carried out in the Ardennes to characterize the long‐term temporal trends in soil solution chemistry, foliar nutrition, and crown condition. In parallel, a model describing the nutrient cycling in forests (NuCM) was calibrated and used to check the consistency of the observed temporal trends and to explain them. The soil solution concentration of most of the elements decreased between 1978 and 2002, which was ascribed to a decrease in atmospheric deposition. For potassium, a decline in the exchangeable pool was also showed based on the simulation carried out with NuCM. As nitrogen (N) deposition remained at an elevated level, Norway spruce stands were progressively saturated in N and mineral nutrition became more and more unbalanced. Except the foliar N and Al concentration that remained constant and increased respectively, the foliar concentration of all other nutrients decreased between 1993 and 2009, which can be explained by the decrease in ion concentration in solution. These nutritional disorders weakened trees and were probably exacerbated during the 2003 summer drought, after which symptoms of vitality loss progressively appeared. In these N‐saturated ecosystems, the N cycle was disrupted by this health decline, which increased NO₃^? leaching reinforcing soil acidification and risk of aluminum (Al) toxicity.     

Impact of sheep grazing on nutrient budgets of dry heathlands

Questions: What effect does sheep grazing have on the nutrient budgets of heathlands? Can grazing compensate for atmospheric nutrient loads in heathland ecosystems? What are the conclusions for heathland management? Location: Lüneburg Heath, NW Germany. Methods: During a one-year grazing experiment (stocking rate 1.1 sheep/ha) nutrient balances for N, Ca, K, Mg and P were calculated by quantifying input rates (atmospheric deposition, sheep excrement) and output rates (biomass removal, leaching). Results: Atmospheric nutrient deposition amounted to 22.8 kg.ha⁻¹.a⁻¹ for N and < 0.2 kg.ha⁻¹.a⁻¹ for P. Sheep excrement increased the inputs for N and P by ca. 3.5 and 0.2 kg.ha⁻¹.a⁻¹, respectively. Grazing reduced N- and P-stores in the above-ground biomass by 25.6 and 1.9 kg.ha⁻¹.a⁻¹, respectively. N-and P-losses via leaching amounted to 2.2 and < 0.2 kg.ha⁻¹.a⁻¹. Output:input ratios for P were high, indicating that grazing severely affected P-budgets of heaths. Conclusions: Our results suggest that sheep grazing has the potential to compensate for atmospheric nutrient loads (particularly for current N deposition rates). However, in the long term the combination of elevated N-deposition and P-loss due to grazing may cause a shift from N-(co-) limited to more P-(co-) limited plant growth. To counteract an aggravation of P-deficiency in the long term, grazing may be combined with management measures that affect P-budgets to a lesser extent (e.g. prescribed burning).     

On the tracks of Nitrogen deposition effects on temperate forests at their southern European range – an observational study from Italy

We studied forest monitoring data collected at permanent plots in Italy over the period 2000–2009 to identify the possible impact of nitrogen (N) deposition on soil chemistry, tree nutrition and growth. Average N throughfall (N‐NO₃+N‐NH₄) ranged between 4 and 29 kg ha^?1 yr^?1, with Critical Loads (CLs) for nutrient N exceeded at several sites. Evidence is consistent in pointing out effects of N deposition on soil and tree nutrition: topsoil exchangeable base cations (BCE) and pH decreased with increasing N deposition, and foliar nutrient N ratios (especially N : P and N : K) increased. Comparison between bulk openfield and throughfall data suggested possible canopy uptake of N_, levelling out for bulk deposition >4–6 kg ha^?1 yr^?1. Partial Least Square (PLS) regression revealed that ‐ although stand and meteorological variables explained the largest portion of variance in relative basal area increment (BAI_rel 2000–2009) ‐ N‐related predictors (topsoil BCE, C : N, pH; foliar N‐ratios; N deposition) nearly always improved the BAI_rel model in terms of variance explained (from 78.2 to 93.5%) and error (from 2.98 to 1.50%). N deposition was the strongest predictor even when stand, management and atmosphere‐related variables (meteorology and tropospheric ozone) were accounted for. The maximal annual response of BAI_rel was estimated at 0.074–0.085% for every additional kgN. This corresponds to an annual maximal relative increase of 0.13–0.14% of carbon sequestered in the above‐ground woody biomass for every additional kgN, i.e. a median value of 159 kgC per kgN ha^?1 yr^?1 (range: 50–504 kgC per kgN, depending on the site). Positive growth response occurred also at sites where signals of possible, perhaps recent N saturation were detected. This may suggest a time lag for detrimental N effects, but also that, under continuous high N input, the reported positive growth response may be not sustainable in the long‐term.     

Deposition and soil leaching in stands of Norway spruce and European Beech: Results from the Höglwald research in comparison with other European case studies

Stands of Norway spruce (Picea abies K.) and European beech (Fagus sylvatica L.) were investigated at the Höglwald research area, Southern Germany from 1985–1988 and from 1994–1997 in order to determine the effects of tree species on deposition and soil solution fluxes. The results were compared to 15 European case studies representing different deposition levels and site conditions. At the Höglwald site, which is characterised by a high nitrogen and a moderate sulphur load, throughfall deposition of nitrogen and sulphur compounds was about two-fold higher in spruce stands compared to beech stands. The differences in elemental input were clearly reflected in soil solution chemistry with a higher leaching of nitrate and sulphate in the spruce stands. The turnover of sulphur and nitrogen compounds induced a strong soil internal production of protons especially in the spruce stands. These results are in accordance with the other European case studies. Throughfall deposition and soil leaching of nitrogen and sulphur compounds was generally higher for spruce stands compared to beech stands. The species-related differences were mainly caused by dry deposition and were relatively small in remote areas. The consequences for the forest ecosystem itself and for the hydrosphere are discussed.     

Long-term changes of ecosystem services at Solling,Germany: Recovery from acidification,but increasing nitrogen saturation?

Time series of values of ingenious parameters indicating ecosystem services from European beech and Norway spruce ecosystems at Solling, Germany, were evaluated with respect to resilient or adaptive behaviour. Studied indicators comprise the use of monitoring data with up to more than 40 years of observation on deposition of potential acidity, sulphate (SO₄²⁻) budgets, exchangeable base cation pools, Bc/Al ratio in soil solution, nitrogen (N) budgets, foliar nutrition as indicated by the foliar Bc/N ratio, and defoliation. Deposition of potential acidity decreased considerably at both ecosystems. SO₄²⁻ budgets reveal retention of sulphur in the soils affecting acid/base budgets. Exchangeable base cation pools decreased at both ecosystems by about 60%. Bc/Al ratio in soil solution in the mineral soil was mostly below critical limits indicating potential toxic stress to tree roots. N retention in the soils decreased from about 40 kg ha⁻¹ yr⁻¹ in the 1970s to currently very low rates of 0–20 kg ha⁻¹ yr⁻¹ indicating increasing N saturation. Foliar Bc/N ratio decreased at the spruce ecosystem indicating possible nutrient imbalances. Defoliation at both Solling ecosystems is on a high level compared to other forests in Germany, but reveals no distinct relation to soil acidification or N saturation. From the selected indicators, SO₄²⁻ and N budgets reveal resilient behaviour, whereas indicators related to the acid/base status tend to adaptive behaviour.     

Topography strongly affects atmospheric deposition and canopy exchange processes in different types of wet lowland rainforest,Southwest Costa Rica

Bulk precipitation and throughfall were collected in a wet lowland rainforest in SW Costa Rica on an event basis to allow modelling the contributions of dry deposition and canopy exchange to nutrient inputs and internal cycling of nutrients. Estimates based on bulk precipitation underestimated total atmospheric deposition to tropical rainforests by up to 10-fold ignoring the contributions of dry deposition. Canopy exchange contributed most of the aboveground inputs to the forest soil of Na⁺, about half for K⁺, 10% for P and Mg²⁺ and negligible for N, C and other elements. Tree species composition did not account for the differences found in net throughfall between forest sites, and vegetation structure (plant area index) had only a small effect on net throughfall. Forest regrowth affected net throughfall through reduced soil fertility and differences in leaf traits. Topography most significantly affected net throughfall via increased dry deposition at sites of higher elevation and via soil fertility and increased canopy exchange at down slope sites.     

Potential canopy influences on the isotopic composition of nitrogen and sulphur in atmospheric deposition

Isotopic studies of nitrogen and sulphur inputs to plant/soil systems commonly rely on limited published data for the ¹⁵N/¹⁴N and ³⁴S/³²S ratios of nitrate, ammonium and sulphate in rainfall. For systems with well-developed plant canopies, however, inputs of these ions from dry deposition or particulates may be more important than rainfall. The manner in which isotopic fractionation between ions and gases may lead to dry deposition and particulates having ¹⁵N/¹⁴N or ³⁴S/³²S ratios different from those of rainfall is considered. Data for rainfall and throughfall in coniferous plantations are then discussed, and suggest that: (1) in line with expectations, nitrate washed from the canopy has ¹⁵N/¹⁴N ratios higher than those in rainfall; (2) the ¹⁵N/¹⁴N ratios of ammonium washed from the canopy are variable, with high ratios being found for canopies of higher pH in conditions of elevated ambient ammonia gas concentrations; and (3) in accord with expectations and previous work, ³⁴S/³²S ratios of sulphate washed from the canopy are not substantially different from those in rainfall. The study suggests that if atmospheric inputs are relevant to isotopic studies of the sources of nitrogen for canopied systems, then confident interpretation will require analysis of these inputs. Received: 3 March 1996 / Accepted: 28 September 1996