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.     

Plant and soil natural abundance <Emphasis Type="Italic">δ</Emphasis><Superscript>15</Superscript>N: indicators of relative rates of nitrogen cycling in temperate forest ecosystems

Watersheds within the Catskill Mountains, New York, receive among the highest rates of nitrogen (N) deposition in the northeastern United States and are beginning to show signs of N saturation. Despite similar amounts of N deposition across watersheds within the Catskill Mountains, rates of soil N cycling and N retention vary significantly among stands of different tree species. We examined the potential use of δ ¹⁵N of plants and soils as an indicator of relative forest soil N cycling rates. We analyzed the δ ¹⁵N of foliage, litterfall, bole wood, surface litter layer, fine roots and organic soil from single-species stands of American beech (Fagus grandifolia), eastern hemlock (Tsuga canadensis), red oak (Quercus rubra), and sugar maple (Acer saccharum). Fine root and organic soil δ ¹⁵N values were highest within sugar maple stands, which correlated significantly with higher rates of net mineralization and nitrification. Results from this study suggest that fine root and organic soil δ ¹⁵N can be used as an indicator of relative rates of soil N cycling. Although not statistically significant, δ ¹⁵N was highest within foliage, wood and litterfall of beech stands, a tree species associated with intermediate levels of soil N cycling rates and forest N retention. Our results show that belowground δ ¹⁵N values are a better indicator of relative rates of soil N cycling than are aboveground δ ¹⁵N values.     

Decades of atmospheric deposition have not resulted in widespread phosphorus limitation or saturation of tree demand for nitrogen in southern New England

It is commonly assumed that nitrogen (N) is the primary mineral resource limiting the productivity of temperate forests. Sustained inputs of N via atmospheric deposition are altering the N status of temperate forests raising the possibility that nutrients such as phosphorus (P) are increasingly limiting productivity. The objective of this study was to determine whether P availability limits tree growth alone or in combination with N. This study was conducted in two forest types common throughout the New England landscape of the northeastern United States; in sugar maple and white ash dominated stands growing on base rich parent material characterized by rapid rates of N cycling and high N availability, and in red oak–beech–hemlock dominated stands growing on base-poor parent material characterized by slow rates of N cycling and low N availability. Starting in 2004, N and P were added to replicate plots in each forest type in factorial combination at a rate of 150 and 50 kg ha⁻¹ year⁻¹, respectively. Diameter growth rates of all trees >10 cm DBH were measured in 2005 and 2006 using dendrometer bands and converted into units of basal area increment (BAI) and wood production. Following 2 years of fertilization, basal area increment in the sugar maple–white ash forests remained strongly N limited. Fertilization with P did not significantly increase BAI alone, although both N and P fertilization tended (P < 0.10) to increase diameter growth in white ash. Wood production in the N-fertilized plots increased by 100 g C m⁻² year⁻¹, roughly doubling production in the non-fertilized plots. In the red oak–beech–hemlock stands, there was no overall effect of N or P fertilization on BAI or wood production because BAI in some species was stimulated by fertilization with N alone (e.g., black cherry, red oak), while in other species BAI was unaffected (e.g., red maple, beech) or negatively affected by fertilization with N or P (e.g., eastern hemlock). Given that BAI in several tree species responded to fertilization with N alone and that only one species responded to P fertilization once N was added, this study suggests that decades of atmospheric N deposition have not (yet) resulted in widespread P limitation or saturation of tree demand for N.     

Nitrogen Addition Increases Carbon Storage in Soils, But Not in Trees, in an Eastern U.S. Deciduous Forest

Forest ecosystems in most industrialized and agricultural regions receive elevated rates of atmospheric nitrogen (N) deposition from air pollution. To evaluate the effects of excess N deposition on carbon (C) and N cycling, we experimentally added N (as NH₄NO₃) to naturally-occurring, single-species plots of five different tree species that are common in the Northern Hardwood forests of northeastern North America: sugar maple (Acer saccharum Marsh), American beech (Fagus grandifolia Ehrh.), yellow birch (Betula alleghaniensis Britton), eastern hemlock (Tsuga canadensis (L.) Carr), and northern red oak (Quercus rubra L.). The experiment was performed in the Catskill Mountains of southeastern New York State, USA, and used a paired-plot design with six replicate plots per species. After 6 years of treatment, most species showed increases in foliar N concentrations in N-treated plots, but only for maple and birch were those increases statistically significant. No significant effects of the N treatment were observed on woody biomass increment or aboveground net primary production (ANPP) for any species. In the oak plots, the N treatment increased acorn production in mast years. In the soils, the N treatment was associated with a significant decline in potential N mineralization and nitrification rates in the mineral horizon but not in the forest floor, and in the mineral horizon the effect of the N treatment varied among species. The N treatment caused a significant increase in C stock, N stock and C:N ratio in the forest floor, with the largest effect in the hemlock plots. Nitrate leaching increased significantly in treated plots compared to controls. Dissolved organic carbon (DOC) in soil solution was unaffected by the N treatment, but the variation in DOC across plots was correlated with the C stock in the forest floor. These results suggest that the ANPP of these forests is not limited by N availability, but that excess N may cause accumulations of C in the forest floor, particularly in hemlock stands, perhaps through inhibition of decomposition rates or by altering phenolic chemistry of the litter. The magnitude, and sometimes the direction of the N treatment responses varied among species, suggesting that predictions of forest responses to elevated N deposition should take into account spatial and temporal variation in tree species composition.     

Biotic and abiotic immobilization of ammonium, nitrite, and nitrate in soils developed under different tree species in the Catskill Mountains, New York, USA

Nitrogen retention in soil organic matter (SOM) is a key process influencing the accumulation and loss of N in forest ecosystems, but the rates and mechanisms of inorganic N retention in soils are not well understood. The primary objectives of this study were to compare ammonium (NH₄⁺), nitrite (NO₂^?), and nitrate (NO₃^?) immobilization among soils developed under different tree species in the Catskill Mountains of New York State, and to determine the relative roles of biotic or abiotic processes in soil N retention. A laboratory experiment was performed, where ¹⁵N was added as NH₄⁺, NO₂^?, or NO₃^? to live and mercury‐treated O horizon soils from three tree species (American beech, northern red oak, sugar maple), and ¹⁵N recoveries were determined in the SOM pool. Mercuric chloride was used to treat soils as this chemical inhibits microbial metabolism without significantly altering the chemistry of SOM. The recovery of ¹⁵N in SOM was almost always greater for NH₄⁺ (mean 20%) and NO₂^? (47%) than for NO₃^? (10%). Ammonium immobilization occurred primarily by biotic processes, with mean recoveries in live soils increasing from 9% at 15 min to 53% after 28 days of incubation. The incorporation of NO₂^? into SOM occurred rapidly (<15 min) via abiotic processes. Abiotic immobilization of NO₂^? (mean recovery 58%) was significantly greater than abiotic immobilization of NH₄⁺ (7%) or NO₃^? (7%). The incorporation of NO₂^? into SOM did not vary significantly among tree species, so this mechanism likely does not contribute to differences in soil NO₃^? dynamics among species. As over 30% of the ¹⁵NO₂^? label was recovered in SOM within 15 min in live soils, and the products of NO₂^? incorporation into SOM remained relatively stable throughout the 28‐day incubation, our results suggest that NO₂^? incorporation into SOM may be an important mechanism of N retention in forest soils. The importance of NO₂^? immobilization for N retention in field soils, however, will depend on the competition between incorporation into SOM and nitrification for transiently available NO₂^?. Further research is required to determine the importance of this process in field environments.     

The phosphorus status of northern hardwoods differs by species but is unaffected by nitrogen fertilization

Northern hardwood forests in the eastern US exhibit species-specific influences on nitrogen (N) cycling, suggesting that their phosphorus (P) cycling characteristics may also vary by species. These characteristics are increasingly important to understand in light of evidence suggesting that atmospheric N deposition has increased N availability in the region, potentially leading to phosphorus limitation. We examined how P characteristics differ among tree species and whether these characteristics respond to simulated N deposition (fertilization). We added NH₄NO₃ fertilizer (50 kg ha^?1 year^?1) to single-species plots of red oak (Quercus rubra L.), sugar maple (Acer saccharum Marsh.), eastern hemlock (Tsuga canadensis (L.) Carr.), American beech (Fagus grandifolia Ehrh.), and yellow birch (Betula alleghaniensis Britt.), in the Catskill Mountains, New York from 1997 to 2007. Species differences were observed in foliar, litter and root P concentrations, but all were unaffected by a cumulative N fertilization of 550 kg/ha. Similarly, measures of soil P availability and biotic P sufficiency differed by species but were unaffected by N fertilization. Results suggest species exhibit unique relationships to P as well as N cycles. We found little evidence that N fertilization leads to increased P limitation in these northern hardwood forests. However, species such as sugar maple and red oak may be sufficient in P, whereas beech and hemlock may be less sufficient and therefore potentially more sensitive to future N-stimulated P limitation.     

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.     

Nitrogen cycling in a northern hardwood forest: Do species matter?

To investigate the influence of individual tree species on nitrogen (N) cycling in forests, we measured key characteristics of the N cycle in small single-species plots of five dominant tree species in the Catskill Mountains of New York State. The species studied were sugar maple (Acer saccharum), American beech (Fagus grandifolia), yellow birch (Betula alleghaniensis), eastern hemlock (Tsuga canadensis), and red oak (Quercus rubra). The five species varied markedly in N cycling characteristics. For example, hemlock plots consistently showed characteristics associated with "slow" N cycling, including low foliar and litter N, high soil C:N, low extractable N pools, low rates of potential net N mineralization and nitrification and low NO ₃ ^– amounts trapped in ion-exchange resin bags buried in the mineral soil. Sugar maple plots had the lowest soil C:N, and the highest levels of soil characteristics associated with NO ₃ ^– production and loss (nitrification, extractable NO ₃ ^– , and resin bag NO ₃ ^– ). In contrast, red oak plots had near-average net mineralization rates and soil C:N ratios, but very low values of the variables associated with NO ₃ ^– production and loss. Correlations between soil N transformations and litter concentrations of N, lignin, lignin:N ratio, or phenolic constituents were generally weak. The inverse correlation between net nitrification rate and soil C:N that has been reported in the literature was present in this data set only if red oak plots were excluded from the analysis. This study indicates that tree species can exert a strong control on N cycling in forest ecosystems that appears to be mediated through the quality of soil organic matter, but that standard measures of litter quality cannot explain the mechanism of control.     

Soil amino-acid availability across a temperate-forest fertility gradient

Despite increasing recognition that free amino acids can be an important source of N for plant uptake, we have a poor understanding of environmental variation in the availability of amino-acid N in soils outside of arctic, alpine and boreal regions. I investigated patterns of amino-acid availability along a temperate forest fertility gradient ranging from low mineral N availability, oak-dominated forests to high mineral N availability, maple-basswood forests (5 sites). I measured standing pools of free amino acids, soluble peptides, ammonium and nitrate, rates of amino acid production (native proteolysis activity) and rates of consumption of a ¹⁵N-labeled leucine tracer. Standing pools of amino acid N decreased consistently along the fertility gradient from the low fertility black oak/white oak system to the high fertility sugar maple/basswood system, with a 25-fold difference in pool sizes between the poorest and richest sites. Standing pools of soluble peptides varied little among sites, instead, the relationship between free amino acids and peptides changed markedly across the gradient. At low fertility sites free amino acids were positively correlated with soluble peptides, whereas free amino acid pools were universally low at high fertility sites, regardless of peptide pools. Assays for native proteolysis activity indicated that amino acid production did not vary significantly among sites. Recovery of leucine tracer in inorganic (NH₄ ⁺ and NO₃ ⁻) pools and in residual soil organic matter both increased with increasing soil fertility; however, total consumption of the added amino-acid tracer did not vary among sites. Results from this study demonstrate that free amino acids can make an important contribution to potentially plant-available N pools in temperate forest soils, particularly at low fertility sites.     

High retention of 15N‐labeled nitrogen deposition in a nitrogen saturated old‐growth tropical forest

The effects of increased reactive nitrogen (N) deposition in forests depend largely on its fate in the ecosystems. However, our knowledge on the fates of deposited N in tropical forest ecosystems and its retention mechanisms is limited. Here, we report the results from the first whole ecosystem ¹⁵N labeling experiment performed in a N‐rich old‐growth tropical forest in southern China. We added ¹⁵N tracer monthly as ¹⁵NH₄¹⁵NO₃ for 1 year to control plots and to N‐fertilized plots (N‐plots, receiving additions of 50 kg N ha^?1 yr^?1 for 10 years). Tracer recoveries in major ecosystem compartments were quantified 4 months after the last addition. Tracer recoveries in soil solution were monitored monthly to quantify leaching losses. Total tracer recovery in plant and soil (N retention) in the control plots was 72% and similar to those observed in temperate forests. The retention decreased to 52% in the N‐plots. Soil was the dominant sink, retaining 37% and 28% of the labeled N input in the control and N‐plots, respectively. Leaching below 20 cm was 50 kg N ha^?1 yr^?1 in the control plots and was close to the N input (51 kg N ha^?1 yr^?1), indicating N saturation of the top soil. Nitrogen addition increased N leaching to 73 kg N ha^?1 yr^?1. However, of these only 7 and 23 kg N ha^?1 yr^?1 in the control and N‐plots, respectively, originated from the labeled N input. Our findings indicate that deposited N, like in temperate forests, is largely incorporated into plant and soil pools in the short term, although the forest is N‐saturated, but high cycling rates may later release the N for leaching and/or gaseous loss. Thus, N cycling rates rather than short‐term N retention represent the main difference between temperate forests and the studied tropical forest.     

Tree growth and soil acidification in response to 30 years of experimental nitrogen loading on boreal forest

Relations among nitrogen load, soil acidification and forest growth have been evaluated based on short‐term (<15 years) experiments, or on surveys across gradients of N deposition that may also include variations in edaphic conditions and other pollutants, which confound the interpretation of effects of N per se. We report effects on trees and soils in a uniquely long‐term (30 years) experiment with annual N loading on an un‐polluted boreal forest. Ammonium nitrate was added to replicated (N=3) 0.09 ha plots at two doses, N1 and N2, 34 and 68 kg N ha^?1 yr^?1, respectively. A third treatment, N3, 108 kg N ha^?1 yr^?1, was terminated after 20 years, allowing assessment of recovery during 10 years. Tree growth initially responded positively to all N treatments, but the longer term response was highly rate dependent with no gain in N3, a gain of 50 m³ ha^?1 stemwood in N2 and a gain of 100 m³ ha^?1 stemwood in excess of the control (N0) in N1. High N treatments caused losses of up to 70% of exchangeable base cations (Ca²⁺, Mg²⁺, K⁺) in the mineral soil, along with decreases in pH and increases in exchangeable Al³⁺. In contrast, the organic mor‐layer (forest floor) in the N‐treated plots had similar amounts per hectare of exchangeable base cations as in the N0 treatment. Magnesium was even higher in the mor of N‐treated plots, providing evidence of up‐lift by the trees from the mineral soil. Tree growth did not correlate with the soil Ca/Al ratio (a suggested predictor of effects of soil acidity on tree growth). A boron deficiency occurred on N‐treated plots, but was corrected at an early stage. Extractable NH₄⁺ and NO₃^?were high in mor and mineral soils of on‐going N treatments, while NH₄+ was elevated in the mor only in N3 plots. Ten years after termination of N addition in the N3 treatment, the pH had increased significantly in the mineral soil; there were also tendencies of higher soil base status and concentrations of base cations in the foliage. Our data suggest the recovery of soil chemical properties, notably pH, may be quicker after removal of the N‐load than predicted. Our long‐term experiment demonstrated the fundamental importance of the rate of N application relative to the total amount of N applied, in particular with regard to tree growth and C sequestration. Hence, experiments adding high doses of N over short periods do not mimic the long‐term effects of N deposition at lower rates.     

Multiyear fate of a 15N tracer in a mixed deciduous forest: retention,redistribution, and differences by mycorrhizal association

The impact of atmospheric nitrogen deposition on forest ecosystems depends in large part on its fate. Past tracer studies show that litter and soils dominate the short‐term fate of added ¹⁵N, yet few have examined its longer term dynamics or differences among forest types. This study examined the fate of a ¹⁵N‐ tracer over 5–6 years in a mixed deciduous stand that was evenly composed of trees with ectomycorrhizal and arbuscular mycorrhizal associations. The tracer was expected to slowly mineralize from its main initial fate in litter and surface soil, with some ¹⁵N moving to trees, some to deeper soil, and some net losses. Recovery of added ¹⁵N in trees and litterfall totaled 11.3% both 1 and 5–6 years after the tracer addition, as ¹⁵N redistributed from fine and especially coarse roots into cumulative litterfall and small accumulations in woody tissues. Estimates of potential carbon sequestration from tree ¹⁵N recovery amounted to 12–14 kg C per kg of N deposition. Tree ¹⁵N acquisition occurred within the first year after the tracer addition, with no subsequent additional net transfer of ¹⁵N from detrital to plant pools. In both years, ectomycorrhizal trees gained 50% more of the tracer than did trees with arbuscular mycorrhizae. Much of the ¹⁵N recovered in wood occurred in tree rings formed prior to the ¹⁵N addition, demonstrating the mobility of N in wood. Tracer recovery rapidly decreased over time in surface litter material and accumulated in both shallow and deep soil, perhaps through mixing by earthworms. Overall, results showed redistribution of tracer ¹⁵N through trees and surface soils without any losses, as whole‐ecosystem recovery remained constant between 1 and 5–6 years at 70% of the ¹⁵N addition. These results demonstrate the persistent ecosystem retention of N deposition even as it redistributes, without additional plant uptake over this timescale.     

Seedling and Sapling Dynamics of Treefall Pits in Puerto Rico1

Seedling and sapling dynamics in a Puerto Rican rain forest were compared between forest understory and soil pits created by the uprooting of 27 trees during Hurricane Hugo. Soil N and P, organic matter, and soil moisture were lower and bulk densities were higher in the disturbed mineral soils of the pits than in undisturbed forest soils ten months after the hurricane. No differences in N and P levels were found in pit or forest soils under two trees with N–fixing symbionts (Inga laurina and Ormosia krugii) compared to soils under a tree species without N–fixing sym–bionts (Casearia arborea), but other soil variables (Al, Fe, K) did vary by tree species. Forest plots had greater species richness of seedlings (<10 cm tall) and saplings (10–100 cm tall) than plots in the soil pits (and greater sapling densities), but seedling densities were similar between plot types. Species richness and seedling densities did not vary among plots associated with the three tree species, but some saplings were more abundant under trees of the same species. Pit size did not affect species richness or seedling and sapling densities. Recruitment of young Cecropia schreberiana trees (>5 m tall) 45 months after the hurricane was entirely from the soil pits, with no tree recruitment from forest plots. Larger soil pits had more tree recruitment than smaller pits. Defoliation of the forest by the hurricane created a large but temporary increase in light availability. Recruitment of C. schreberiana to the canopy occurred in gaps created by the treefall pits that had lower soil nutrients but provided a longer–term increase in light availability. Treefall pits also significantly altered the recruitment and mortality of many understory species in the Puerto Rican rain forest but did not alter species richness.     

Litter decomposition contrasts in second- and old-growth Douglas-fir forests of the Pacific Northwest, USA

Litterfall and its subsequent decomposition are important feedback mechanisms in the intrasystem cycling of nutrients in forest ecosystems. The amount of litterfall and the rate of decomposition are expected to vary with stand age and climate. Over a 2-year period, decomposition of five litter types were measured in two second-growth forest stands and one old-growth stand in the Cascade Mountains of southern Washington state, USA. Both second-growth stands were dominated by Douglas-fir [Pseudotsuga menziesii (Mirb.,) Franco] but one had a significant proportion of red alder (Alnus rubra Bong.), a nitrogen (N) fixer. The old-growth stand was dominated by Douglas-fir and western hemlock [Tsuga heterophylla (Raf.) Sarg.]. All stands had a relatively shallow layer of forest floor mass. The five litter types were placed in each stand to evaluate decomposition patterns. Despite significant differences in stand age, microclimate and mean residence times for carbon (C) and N, the rates of litter mass loss varied only slightly between sites. The relative order of species litter mass loss was: vine maple ≫ salal = western hemlock > Douglas-fir (from the youngest stand) > Douglas-fir (from the N rich stand with red alder). The initial litter lignin concentration, not lignin:N, was the primary determinant of decomposition rates, although the initial N concentration was the predictor for mass loss after 2 years in the N rich Douglas-fir-alder stand. All litter types showed immobilization of N for nearly 2 years. Data for Douglas-fir litter suggest that higher levels of N may retard decomposition of tissues with greater amounts of lignified material. The retention of N by the litter appeared influenced by the nutrient capital of the stands as well as the forest floor C:N ratio. Decomposition was minimal during the cold winter months, but displayed a definitive peak period during early Fall with wet weather, warm soils, and fungal activity. Thus, long-term climatic change effects on forest floor C storage may depend more on changes in seasonality of precipitation changes than just temperature changes.     

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.     

温带两种林型对氮沉降的再分配及其生长季动态与影响因子

氮(N)沉降对森林生态系统的结构与功能具有重要的影响,而森林对到达林地的N沉降量及其分配格局的影响尚不清楚。量化了2012—2013年5—10月两个生长季蒙古栎林和杂木林的林内树干径流和穿透雨及其林外大气降雨总氮(TN)、可溶性氮(DN)和颗粒态氮(PN)沉降通量的季节动态,旨在比较两种林型对N沉降的再分配格局及其季节变化,分析影响其变异的主要因子。结果表明:林外大气降雨、蒙古栎林林内、杂木林林内(树干径流+穿透雨)TN沉降平均值分别为:8.49、15.97、13.39 kg hm~(-2)a~(-1),其中DN分别占其TN的76.35%、82.79%和75.02%,PN分别占其TN的26.35%、17.21%和24.98%,蒙古栎林和杂木林林内穿透雨TN沉降量分别占其TN的95.5%和94.5%。蒙古栎林和杂木林冠层淋溶TN沉降量分别为7.48kg hm~(-2)a~(-1)和4.90 kg hm~(2)a~(-1);其中,前者的DN高于后者,但PN呈相反趋势。两种林型的N沉降组分具有明显的季节动态:沉降量均集中在生长季中期(6—8月),生长季前期和末期较低。林外降雨量分别与林外大气降雨、蒙古栎林和杂木林林内的树干径流和穿透雨中的TN、DN浓度呈显著负指数函数关系(P0.001)。连续降雨天数对蒙古栎林、杂木林林内TN、DN浓度的影响表现为连续降雨2 d以内为富集作用,之后为稀释作用。研究表明林冠对大气氮沉降有显著富集作用,其富集强度及时间动态与森林类型和降雨特征有关,建议氮沉降试验应考虑林冠的富集效应。     

Bottomland hardwood productivity: case study in a rapidly subsiding,Louisiana, USA,watershed

The Verret basin was formerly an overflow area between the Mississippi and Atchafalaya rivers and contains about 41,000ha of forested wetlands. Water levels are rising at the rate of over 1 cm/year in this area, and the forests are subjected to longer and deeper flooding. Tree growth, litterfall, and species composition were monitored across a flooding gradient during January 1985–December 1986. The driest area was only 20 cm higher in elevation than the wettest area, but the structure of the forest changes greatly over this range. The drier area was dominated by sweetgum (Liquidambar styraciflua L.), oaks (Quercus spp.), and sugarberry (Celtis laevigata Willd.), while green ash (Fraxinus pennsylvanica Marsh.), red maple (Acer rubrum L.), and baldcypress (Taxodium distichum (L.) Rich.) were dominant in the wetter area. Green ash and bitter pecan (Carya aquatica (Michaux. f.) Nutt.) were found in all plots, but these two species are under severe stress in the more flooded area as evidenced by dead and dying trees. Stem wood production increased from 1985 to 1986 in the driest (392 to 473 g/m²/yr) and wettest (199 to 399 g/m²/yr) plots, but remained relatively unchanged in the transitional area (386 to 380g/m²/yr). Leaf litter production decreased across the gradient from dry to flooded plots during both years. Over 40% of the litterfall in the drier plot was from flood-tolerant shrub species. In the flooded plots, red maple and baldcypress were major contributors to total litterfall. Increased flooding of dry bottomland forests in the future could lead to decreased litterfall and increased tree death over the entire watershed.     

Development of Tree Regeneration in Fern-dominated Forest Understories After Reduction of Deer Browsing

Dennstaedtia punctilobula (hay‐scented fern) can act as a native invasive species in forests in eastern North America where prolonged deer browsing occurs in stands with partially open overstory canopies. Ferns dominate the understory with a 60‐cm tall canopy, with little regeneration of native tree species. It has been hypothesized that, once established, ferns may continue to inhibit tree regeneration after deer browsing has been reduced. To test this hypothesis, we documented the pattern of recovery of the tree seedling understory in plantations of Pinus strobus (white pine) and Pinus resinosa (red pine) on the Quabbin Reservation watershed protection forest in central Massachusetts, where after 40 years of intensive deer browsing the deer herd was rapidly reduced through controlled hunting. Dense fern understories occur on nearly 4,000 ha of the predominantly oak–pine forest. Three years after deer herd reduction, stands with the highest density fern cover (77% of plots with>90% cover) had significantly fewer seedlings at least 30 cm in height, compared with stands with lower fern density, and those seedlings consisted almost entirely of Betula lenta (black birch) and white pine. Height growth analysis showed that black birch and white pine grew above the height of the fern canopy in 3 and 6 years, respectively. In contrast, two common species, Fraxinus americana (white ash) and Quercus rubra (red oak), grew beneath the dense fern cover for 5 years with height growth less than 5 cm/yr after the first year. A study of spring phenology indicated that the ability of black birch to grow through the fern canopy might have been due to its early leaf development in spring before the fern canopy was formed, in contrast to oak and ash with delayed leaf development. Thus, the ferns showed differential interference among species with seedling development after reduction of deer browse.     

Resistance of European tree species to drought stress in mixed versus pure forests: evidence of stress release by inter‐specific facilitation

While previous studies focused on tree growth in pure stands, we reveal that tree resistance and resilience to drought stress can be modified distinctly through species mixing. Our study is based on tree ring measurement on cores from increment boring of 559 trees of Norway spruce (Picea abies [L.] Karst.), European beech (Fagus sylvatica [L.]) and sessile oak (Quercus petraea (Matt.) Liebl.) in South Germany, with half sampled in pure, respectively, mixed stands. Indices for resistance, recovery and resilience were applied for quantifying the tree growth reaction on the episodic drought stress in 1976 and 2003. The following general reaction patterns were found. (i) In pure stands, spruce has the lowest resistance, but the quickest recovery; oak and beech were more resistant, but recover was much slower and they are less resilient. (ii) In mixture, spruce and oak perform as in pure stands, but beech was significantly more resistant and resilient than in monoculture. (iii) Especially when mixed with oak, beech is facilitated. We hypothesise that the revealed water stress release of beech emerges in mixture because of the asynchronous stress reaction pattern of beech and oak and a facilitation of beech by hydraulic lift of water by oak. This facilitation of beech in mixture with oak means a contribution to the frequently reported overyield of beech in mixed versus pure stands. We discuss the far‐reaching implications that these differences in stress response under intra‐ and inter‐specific environments have for forest ecosystem dynamics and management under climate change.     

Intact amino acid uptake by northern hardwood and conifer trees

Empirical and modeling studies of the N cycle in temperate forests of eastern North America have focused on the mechanisms regulating the production of inorganic N, and assumed that only inorganic forms of N are available for plant growth. Recent isotope studies in field conditions suggest that amino acid capture is a widespread ecological phenomenon, although northern temperate forests have yet to be studied. We quantified fine root biomass and applied tracer-level quantities of U–¹³C₂–¹⁵N-glycine, ¹⁵NH₄ ⁺ and ¹⁵NO₃ ⁻ in two stands, one dominated by sugar maple and white ash, the other dominated by red oak, beech, and hemlock, to assess the importance of amino acids to the N nutrition of northeastern US forests. Significant enrichment of ¹³C in fine roots 2 and 5 h following tracer application indicated intact glycine uptake in both stands. Glycine accounted for up to 77% of total N uptake in the oak–beech–hemlock stand, a stand that produces recalcitrant litter, cycles N slowly and has a thick, amino acid-rich organic horizon. By contrast, glycine accounted for only 20% of total N uptake in the sugar maple and white ash stand, a stand characterized by labile litter and rapid rates of amino acid production and turnover resulting in high rates of mineralization and nitrification. This study shows that amino acid uptake is an important process occurring in two widespread, northeastern US temperate forest types with widely differing rates of N cycling.