Growth,Net Production,Litter Decomposition,and Net Nitrogen Accumulation by Epiphytic Bryophytes in a Tropical Montane Forest1

To understand the ecological roles of epiphytic bryophytes in the carbon (C) and nitrogen (N) cycles of a tropical montane forest, we used samples in enclosures to estimate rates of growth, net production, and N accumulation by shoots in the canopy, and litterbags, to estimate rates of decomposition and N dynamics of epiphytic bryophyte litter in the canopy and on the forest floor in Monteverde, Costa Rica. Growth of epiphytic bryophytes was estimated at 30.0–49.9 percent/yr, net production at 122–203 g/m²/yr, and N accumulation at 1.8–3.0 g N/m²/yr. Cumulative mass loss from litterbags after one and two years in the canopy was 17 ± 2 and 19 ± 2 percent (mean ± 1 SE) of initial sample mass, respectively, and mass loss from litter and green shoots in litterbags after one year on the forest floor was 29 ± 2 and 45 ± 3 percent, respectively. Approximately 30 percent of the initial N mass was released rapidly from litter in both locations. Nitrogen loss from green shoots on the forest floor was greater; about 47 percent of the initial N mass was lost within the first three months. There was no evidence for net N immobilization by litter or green shoots, but the remaining N in litter was apparently recalcitrant. Annual net accumulation of C and N by epiphytic bryophytes was estimated at 37–64 g C/m²/yr and 0.8–1.3 g N/m²/yr, respectively. Previous research at this site indicated that epiphytic bryophytes retain inorganic N from atmospheric deposition to the canopy. Therefore, they play a major role in transforming N from mobile to highly recalcitrant forms in this ecosystem.     

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.     

Water chemistry and nutrient budgets in an undisturbed evergreen rainforest of Southern Chile

In a pristine evergreen rainforest of Nothofagus betuloides, located at the Cordillera de los Andes in southern Chile (41?°S), concentrations and fluxes of nutrients in bulk precipitation, cloud water, throughfall water, stemflow water, soil infiltration and percolation water and runoff water were measured. The main objectives of this study were to investigate canopy–soil–atmosphere interactions and to calculate input–output budgets. From May 1999 till April 2000, the experimental watershed received 8121?mm water (86% incident precipitation, 14% cloud water), of which the canopy intercepted 16%. Runoff water volume amounted 9527?mm. Bulk deposition of inorganic (DIN) and organic (DON) nitrogen amounted 3.6?kg?ha^?1?year^?1 and 8.2?kg?ha^?1?year^?1 respectively. Occult deposition (clouds?+?fog) contributes for 40% to the atmospheric nitrogen input (bulk?+?occult deposition) of the forest. An important part of the atmospheric ammonium deposition is retained within the canopy or converted to nitrate or organic nitrogen by epiphytic bacteria or lichens. Also the export of inorganic (0.9?kg?ha^?1?year^?1) and organic (5.2?kg?ha^?1?year^?1) nitrogen via runoff is lower than the input to the forest floor via throughfall and stemflow water (3.2?kg?DIN?ha^?1?year^?1 and 5.6?kg?DON?ha^?1?year^?1). The low concentrations of NO-₃ and NH+₄ under the rooting depth suggest an effective biological immobilization by vegetation and soil microflora. Dry deposition and foliar leaching of base cations (K⁺, Ca²⁺, Mg²⁺) was estimated using a canopy budget model. Bulk deposition accounted for about 50% of the total atmospheric input. Calculated dry and occult deposition are both of equal value (about 25%). Foliar leaching of K⁺, Ca²⁺, and Mg²⁺ accounted for 45%, 38% and 6% of throughfall deposition respectively. On an annual basis, the experimental watershed was a net source for Na⁺, Ca²⁺ and Mg²⁺.     

Water chemistry and nutrient budgets in an undisturbed evergreen rainforest of southern Chile

In a pristine evergreen rainforest of Nothofagus betuloides, located at the Cordillera de los Andes in southern Chile (41 °S), concentrations and fluxes of nutrients in bulk precipitation, cloud water, throughfall water, stemflow water, soil infiltration and percolation water and runoff water were measured. The main objectives of this study were to investigate canopy-soil-atmosphere interactions and to calculate input-output budgets. From May 1999 till April 2000, the experimental watershed received 8121 mm water (86% incident precipitation, 14% cloud water), of which the canopy intercepted 16%. Runoff water volume amounted 9527 mm. Bulk deposition of inorganic (DIN) and organic (DON) nitrogen amounted 3.6 kg ha^–1 year^–1 and 8.2 kg ha^–1 year^–1 respectively. Occult deposition (clouds + fog) contributes for 40% to the atmospheric nitrogen input (bulk + occult deposition) of the forest. An important part of the atmospheric ammonium deposition is retained within the canopy or converted to nitrate or organic nitrogen by epiphytic bacteria or lichens. Also the export of inorganic (0.9 kg ha^–1 year^–1) and organic (5.2 kg ha^–1 year^–1) nitrogen via runoff is lower than the input to the forest floor via throughfall and stemflow water (3.2 kg DIN ha–1 year^–1 and 5.6 kg DON ha^–1 year^–1). The low concentrations of NO ₃ ^– and NH ₄ ⁺ under the rooting depth suggest an effective biological immobilization by vegetation and soil microflora. Dry deposition and foliar leaching of base cations (K⁺, Ca²⁺, Mg²⁺) was estimated using a canopy budget model. Bulk deposition accounted for about 50% of the total atmospheric input. Calculated dry and occult deposition are both of equal value (about 25%). Foliar leaching of K⁺, Ca²⁺, and Mg²⁺ accounted for 45%, 38% and 6% of throughfall deposition respectively. On an annual basis, the experimental watershed was a net source for Na⁺, Ca²⁺ and Mg²⁺.     

The chemistry of wet deposition for a tallgrass prairie ecosystem: inputs and interactions with plant canopies

The chemistry and nutrient inputs of wet deposition, and the N chemistry of throughfall, were characterized for a tallgrass prairie in north-central Kansas. Dominant ions in wetfall were NH ₄ ⁺ , Ca²⁺, H⁺, NO ₃ ^- , and SO ₄ ^2- ; weighted mean pH was 4.79. Principal sources of ions appeared to be natural emissions and wind-blown soils. Concentrations of NO ₃ ^- -N, NH ₄ ⁺ -N, and organic N in wet deposition were 0.31, 0.30, and 0.17 mg/L, respectively, resulting in N inputs of 2.5, 2.5, and 1.4 kg · ha^-1 · yr^-1. Comparisons with bulk precipitation suggested that at least 50% of atmospheric N inputs were from dry deposition. Concentrations of NO ₃ ^- -N, NH ₄ ⁺ -N, and organic N in unburned prairie throughfall were 0.27, 0.28, and 1.28 mg/L, and in burned prairie throughfall were 0.33, 0.37, and 0.91 mg/L, respectively. The prairie canopy intercepted up to 48% of incident precipitation. Lower inorganic N and higher organic N concentrations in throughfall relative to wet deposition probably resulted from leaf uptake of N and immobilization by microbes associated with the standing dead plant materials of the prairie canopy. The removal of these materials by fire is important in maintaining N availability for tallgrass prairie. Much of the N immobilization appeared to have been of N that was supplied to the prairie canopy by dry deposition.     

Natural 15N abundance of epiphytes depends on the position within the forest canopy: source signals and isotope fractionation

The natural ¹⁵N abundance (δ¹⁵N) of epiphytes and its N sources were studied in the canopy of a lowland rainforest in Costa Rica. Vascular and non‐vascular epiphytes and canopy soils were collected from four canopy zones and analysed for N contents and δ¹⁵N signals. In addition, the N concentrations and δ¹⁵N signatures of bulk precipitation, throughfall and stemflow were measured during the wet and the dry season. The δ¹⁵N values of epiphyte leaves decreased significantly from the lower zones (means of ?3·9 and ?4·3‰) to the upper zones (means of ?5·4 and ?6·1‰) of the canopy. In contrast, δ¹⁵N signatures of canopy soils (average ?0·3‰) differed little between the zones. Bulk deposition was enriched in ¹⁵N (+4·3‰) compared to all other potential N sources and was higher than throughfall and stemflow (+0·5 to ?1·3‰). δ¹⁵N values of atmospheric deposition were inversely related to those of the epiphyte leaves, whereas N isotopic composition of canopy soils did not vary significantly. Consequently, it is concluded that the variations in foliar N isotope composition of epiphytes were not simply caused by utilization of isotopically different N sources, but by different ¹⁵N discrimination during N acquisition.     

Response of epiphytic bryophytes to simulated N deposition in a subtropical montane cloud forest in southwestern China

A field manipulation experiment was conducted in a subtropical montane cloud forest in southwestern China to determine the possible responses of epiphytic bryophytes to increasing nitrogen (N) deposition from community to physiology level, and to find sensitive epiphytic bryophytes that may be used as indicators for assessing the degree of N pollution. N addition had significantly negative effects on species richness and cover of the epiphytic bryophyte community. Harmful effects of high N loads were recorded for chlorophyll, growth, and vitality of the species tested. The decline of some epiphytic bryophytes may result from detrimental effects on degradation to photosynthetic pigments. Bazzania himalayana (Mitt.) Schiffn., Bazzania ovistipula (Steph.) Mizut., and Homaliodendron flabellatum (Sm.) Fleisch. are candidates in atmospheric nitrogen monitoring. Epiphytic bryophytes in the montane cloud forest are very sensitive to increasing N deposition and often difficult to recover once they have been destroyed, providing early detection of enhanced N pollution for trees or even the whole forest ecosystem. The inference that increasing N pollution may lead to loss of biodiversity is a concern to the developing economy in western China, and should alert the government to the adverse impacts caused by increased industrial pollution during the process of China’s West Development.     

Water inputs across a tropical montane landscape in Veracruz,Mexico: synergistic effects of land cover,rain and fog seasonality,and interannual precipitation variability

Land‐cover change can alter the spatiotemporal distribution of water inputs to mountain ecosystems, an important control on land‐surface and land‐atmosphere hydrologic fluxes. In eastern Mexico, we examined the influence of three widespread land‐cover types, montane cloud forest, coffee agroforestry, and cleared areas, on total and net water inputs to soil. Stand structural characteristics, as well as rain, fog, stemflow, and throughfall (water that falls through the canopy) water fluxes were measured across 11 sites during wet and dry seasons from 2005 to 2008. Land‐cover type had a significant effect on annual and seasonal net throughfall (NTF <0=canopy water retention plus canopy evaporation; NTF >0=fog water deposition). Forest canopies retained and/or lost to evaporation (i.e. NTF<0) five‐ to 11‐fold more water than coffee agroforests. Moreover, stemflow was fourfold higher under coffee shade than forest trees. Precipitation seasonality and phenological patterns determined the magnitude of these land‐cover differences, as well as their implications for the hydrologic cycle. Significant negative relationships were found between NTF and tree leaf area index (R²=0.38, P<0.002), NTF and stand basal area (R²=0.664, P<0.002), and stemflow and epiphyte loading (R²=0.414, P<0.001). These findings indicate that leaf and epiphyte surface area reductions associated with forest conversion decrease canopy water retention/evaporation, thereby increasing throughfall and stemflow inputs to soil. Interannual precipitation variability also altered patterns of water redistribution across this landscape. Storms and hurricanes resulted in little difference in forest‐coffee wet season NTF, while El Niño Southern Oscillation was associated with a twofold increase in dry season rain and fog throughfall water deposition. In montane headwater regions, changes in water delivery to canopies and soils may affect infiltration, runoff, and evapotranspiration, with implications for provisioning (e.g. water supply) and regulating (e.g. flood mitigation) ecosystem services.     

The nutrient status of epiphytes and their host trees along an elevational gradient in Costa Rica

Vascular epiphytes are a conspicuous and highly diverse group in tropical wet forests; yet, we understand little about their mineral nutrition across sites. In this study, we examined the mineral nutrition of three dominant vascular epiphyte groups: ferns, orchids, and bromeliads, and their host trees from samples collected along a 2600 m elevational gradient in the tropical wet forests of Costa Rica. We predicted that the mineral nutrition of ferns, orchids, and bromeliads would differ because of their putative differences in nutrient acquisition mechanisms and nutrient sources—atmospherically dependent, foliar feeding bromeliads would have lower nitrogen (N) and phosphorous (P) concentrations and more depleted δ¹⁵N values than those in canopy soil-rooted ferns because canopy soil is higher in available N, and more enriched in δ¹⁵N than the atmospheric sources of precipitation and throughfall. We also predicted that epiphyte foliar chemistry would mirror that of host trees because of the likely contribution of host trees to the nutrient cycle of epiphytes via foliar leaching and litter contributions to canopy soil. In the same vein, we predicted that epiphyte and host tree foliar chemistry would vary with elevation reflecting ecosystem-level nutrients—soil N availability increases and P availability decreases with increasing elevation. Our results confirmed that canopy soil-rooted epiphytes had higher N concentrations than atmospheric epiphytes; however, our predictions were not confirmed with respect to P which did not vary among groups indicating fixed P availability within sites. In addition, foliar δ¹⁵N values did not match our prediction in that canopy soil-rooted as well as atmospheric epiphytes had variable signatures. Discriminant function analysis (DFA) on foliar measurements determined that ferns, orchids, and bromeliads are statistically distinct in mineral nutrition. We also found that P concentrations of ferns and orchids, but not bromeliads, were significantly correlated with those of host trees indicating a possible link in their mineral nutrition’s via canopy soil. Interestingly, we did not find any patterns of epiphyte foliar chemistry with elevation. These data indicate that the mineral nutrition of the studied epiphyte groups are distinct and highly variable within sites and the diverse uptake mechanisms of these epiphyte groups enhance resource partitioning which may be a mechanism for species richness maintenance in tropical forest canopies.     

Old trees contribute bio-available nitrogen through canopy bryophytes

Symbiotic cyanobacteria??bryophyte associations on the forest floor are shown to contribute significantly to stand-level nitrogen budgets through the process of biological nitrogen fixation (BNF), but few studies have considered the role of canopy bryophytes. Given the high biomass of epiphytic bryophytes in many tree species of the North American temperate rain forest, we suggest that canopy bryophytes may contribute substantially to stand-level N dynamics. We confirm the presence of cyanobacteria and measure rates of BNF at three heights (0, 15 and 30 m) in Sitka spruce trees across three watershed estuaries of Clayoquot Sound, British Columbia, Canada. This study is the first to report BNF by cyanobacteria associated with epiphytic and forest floor bryophytes in the coastal temperate rain forest of North America. Cyanobacteria density was significantly greater in epiphytic bryophytes compared to mosses on the forest floor, and rates of BNF were highest at 30 m in the canopy. The majority of total stand-level BNF (0.76 kg N · ha^-1 · yr^-1) occurs in the canopy, rather than on the forest floor (0.26 kg N · ha^-1 · yr^-1). We suggest that BNF by cyanobacterial-bryophyte associations in the canopy of coastal temperate rain forests is a unique source of ecosystem N, which is dependent on large, old trees with high epiphytic bryophyte biomass.     

Roads as nitrogen deposition hot spots

Mobile sources are the single largest source of nitrogen emissions to the atmosphere in the US. It is likely that a portion of mobile-source emissions are deposited adjacent to roads and thus not measured by traditional monitoring networks, which were designed to measure long-term and regional trends in deposition well away from emission sources. To estimate the magnitude of near-source nitrogen deposition, we measured concentrations of both dissolved inorganic nitrogen (DIN) and total dissolved nitrogen (inorganic + organic) (TDN) in throughfall (i.e., the nitrogen that comes through the forest canopy) along transects perpendicular to two moderately trafficked roads on Cape Cod in Falmouth MA, coupled with measurements of both DIN and TDN in bulk precipitation made in adjacent open fields at the same transect distances. We used the TDN throughfall data to estimate total nitrogen deposition, including dry gaseous nitrogen deposition in addition to wet deposition and dry particle deposition. There was no difference in TDN in the bulk collectors along the transects at either site; however TDN in the throughfall collectors was always higher closest to the road and decreased with distance. These patterns were driven primarily by differences in the inorganic N and not the organic N. Annual throughfall deposition was 8.7 (±0.4) and 6.8 (±0.5) TDN kg N ha^?1 year^?1 at sites 10 and 150 m away from the road respectively. We also characterized throughfall away from a non-road edge (power line right-of-way) to test whether the increased deposition observed near road edges was due to deposition near emission sources or due to a physical, edge effect causing higher deposition. The increased deposition we observed near roads was due to increases in inorganic N especially NH₄ ⁺. This increased deposition was not the result of an edge effect; rather it is due to near source deposition of mobile source emissions. We scaled these results to the entire watershed and estimate that by not taking into account the effects of increased gaseous N deposition from mobile sources we are underestimating the amount of N deposition to the watershed by 13–25 %.     

Nitrogen fixation of epiphytic plants enwrapping trees in Ailao Mountain cloud forests, Yunnan, China

Epiphytic plants play an important role in the nutrient cycle of forest ecosystems. There had been fewer studies in subtropical regions than in other climate zones. Prior research showed that the canopy epiphyte could fix nitrogen combined with microorganism in tropical forest. The epiphytic plants enwrapping trees in canopy layer are very abundant in the subtropical mountainous cloud forest of Ailao Mountain (central and southern Yunnan Province, SW China). This forest lacks widespread nitrogen-fixing plants, and the nitrogen origin is elusive. Maybe there also exist such nitrogen-fixing systems in epiphyte community. Nitrogen-fixing potentials of canopy epiphytes increased greatly from dry season to wet season. There occurred an obvious difference on the epiphytic nitrogen fixation abilities between upper canopy layer and sub-canopy layer in alternant period between wet season and dry season. Epiphytic nitrogen-fixing potentials for the subtropical moist forest in Ailao Mountains ranged between 0.027 and 2.24 kg ha^?1?year^?1. Our results indicate that the canopy epiphytes in the subtropical moist forest of Ailao Mountains can fix a significant amount of atmospheric nitrogen. This finding suggests a new nitrogen source for the subtropical forest ecosystem, thus can have profound impact on the studies of nitrogen cycling.     

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.     

The effect of forest type on throughfall deposition and seepage flux: a review

Converting deciduous forests to coniferous plantations and vice versa causes environmental changes, but till now insight into the overall effect is lacking. This review, based on 38 case studies, aims to find out how coniferous and deciduous forests differ in terms of throughfall (+stemflow) deposition and seepage flux to groundwater. From the comparison of coniferous and deciduous stands at comparable sites, it can be inferred that deciduous forests receive less N and S via throughfall (+stemflow) deposition on the forest floor. In regions with relatively low open field deposition of atmospheric N (<10 kg N ha⁻¹ year⁻¹), lower NH₄⁺ mean throughfall (+stemflow) deposition was, however, reported under conifers compared to deciduous forest, while in regions with high atmospheric N pollution (>10 kg N ha⁻¹ year⁻¹), the opposite could be concluded. The higher the open field deposition of NH₄⁺, the bigger the difference between the coniferous and deciduous throughfall (+stemflow) deposition. Furthermore, it can be concluded that canopy exchange of K⁺, Ca²⁺ and Mg²⁺ is on average higher in deciduous stands. The significantly higher stand deposition flux of N and S in coniferous forests is reflected in a higher soil seepage flux of NO₃⁻, SO₄²⁻, K⁺, Ca²⁺, Mg²⁺ and Al(III). Considering a subset of papers for which all necessary data were available, a close relationship between throughfall (+stemflow) deposition and seepage was found for N, irrespective of the forest type, while this was not the case for S. This review shows that the higher input flux of N and S in coniferous forests clearly involves a higher seepage of NO₃⁻ and SO₄²⁻ and accompanying cations K⁺, Ca²⁺, Mg²⁺ and Al(III) into the groundwater, making this forest type more vulnerable to acidification and eutrophication compared to the deciduous forest type.     

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,     

Changes in the composition of rainwater upon passage through the canopies of trees and of ground vegetation in a Dutch oak-birch forest

The composition of canopy throughfall water in an oak-birch woodland, heavily affected by atmospheric deposition of N and S, changed markedly upon contact with the above-ground parts of the ground vegetation, which consisted mainly of bracken. The fluxes of nitrate and H⁺ decreased, simultaneously with an increase in the flux of bicarbonate, indicative of above-ground uptake of nitrate by the ground vegetation. This above-ground assimilation takes place in spite of abundant availability of inorganic nitrogen in the root zone of the ground vegetation. Fluxes of phosphate were somewhat lower, and those of ammonium somewhat higher in throughfall of the ground vegetation than in that of the tree layer. Although those differences were not statistically significant, they do suggest assimilation of some P and extra net dry deposition of atmospheric ammonia below the tree canopies.     

Modeling and Mapping of Atmospheric Mercury Deposition in Adirondack Park,New York

The Adirondacks of New York State, USA is a region that is sensitive to atmospheric mercury (Hg) deposition. In this study, we estimated atmospheric Hg deposition to the Adirondacks using a new scheme that combined numerical modeling and limited experimental data. The majority of the land cover in the Adirondacks is forested with 47% of the total area deciduous, 20% coniferous and 10% mixed. We used litterfall plus throughfall deposition as the total atmospheric Hg deposition to coniferous and deciduous forests during the leaf-on period, and wet Hg deposition plus modeled atmospheric dry Hg deposition as the total Hg deposition to the deciduous forest during the leaf-off period and for the non-forested areas year-around. To estimate atmospheric dry Hg deposition we used the Big Leaf model. The average atmospheric Hg deposition to the Adirondacks was estimated as 17.4 g m yr with a range of −3.7–46.0 g m yr. Atmospheric Hg dry deposition (370 kg yr) was found to be more important than wet deposition (210 kg yr) to the entire Adirondacks (2.4 million ha). The spatial pattern showed a large variation in atmospheric Hg deposition with scattered areas in the eastern Adirondacks having total Hg deposition greater than 30 μg m⁻² yr⁻¹, while the southwestern and the northern areas received Hg deposition ranging from 25–30 μg m⁻² yr⁻¹.     

Nitrogen Cycling and Mass Balance for a Forested Catchment in the Italian Alps. Assessment of Nitrogen Status

During 1999–2001 the chemical composition and fluxes were measured in rainfall, throughfall, soil solution and stream water in a remote forested site in the Italian Alps. The analysis of temporal patterns revealed the differential behaviour of nitrogen and sulphur and suggested that different mechanisms controlled their flux. No important changes in sulphate concentration and fluxes emerged as the solution passed through the various components of the forest ecosystem, and temporal variations of SO₄ in the soil solution and stream were likely driven by the physical process of dilution. The availability of nitrate and ammonia, by contrast, was drastically reduced as throughfall water entered the soil and passed through the mineral layers, irrespective of season. The calculated hydrochemical budget based on throughfall and soil solution N fluxes revealed that ~80% N retention in the forest soil, corresponding to 12 kg ha⁻¹ yr⁻¹, despite a relatively high N deposition loading (15 kg ha⁻¹ yr⁻¹). Most of the leached nitrogen (90%) was in the organic form. Indicators of the N status of this ecosystem, such as C/N ratio in solid and solution phase of the soil and N foliage content as well as land use history were examined. Despite the strong N retention in the forested part of the catchment, the stream water N–NO₃ levels were consistently above 10 μg l⁻¹ suggesting that the Val Masino catchment as a whole was less efficient in processing atmospheric N inputs. This contrasting N behaviour illustrates the role of landscape features, such as the soil cover and vegetation type, that is characteristic of an alpine catchment.     

Changes in Canopy Processes Following Whole-Forest Canopy Nitrogen Fertilization of a Mature Spruce-Hemlock Forest

Abstract Most experimental additions of nitrogen to forest ecosystems apply the N to the forest floor, bypassing important processes taking place in the canopy, including canopy retention of N and/or conversion of N from one form to another. To quantify these processes, we carried out a large-scale experiment and determined the fate of nitrogen applied directly to a mature coniferous forest canopy in central Maine (18–20 kg N ha⁻¹ y⁻¹ as NH₄NO₃ applied as a mist using a helicopter). In 2003 and 2004 we measured NO₃ ⁻, NH₄ ⁺, and total dissolved N (TDN) in canopy throughfall (TF) and stemflow (SF) events after each of two growing season applications. Dissolved organic N (DON) was greater than 80% of the TDN under ambient inputs; however NO₃ ⁻ accounted for more than 50% of TF N in the treated plots, followed by NH₄ ⁺ (35%) and DON (15%). Although NO₃ ⁻ was slightly more efficiently retained by the canopy under ambient inputs, canopy retention of NH₄ ⁺as a percent of inputs increased markedly under fertilization. Recovery of less than 30% of the fertilizer N in TF suggested that the forest canopy retained more than 70% of the applied N (>80% when corrected for N which bypassed tree surfaces at the time of fertilizer addition). Results from plots receiving ¹⁵N enriched NO₃ ⁻ and NH₄ ⁺ confirmed bulk N estimations that more NO₃ ⁻ than NH₄ ⁺ was washed from the canopy by wet deposition. The isotope data did not show evidence of canopy nitrification, as has been reported in other spruce forests receiving much higher N inputs. Conversions of fertilizer-N to DON were observed in TF for both ¹⁵NH₄ ⁺ and ¹⁵NO₃ ⁻ additions, and occurred within days of the application. Subsequent rain events were not significantly enriched in ¹⁵N, suggesting that canopy DON formation was a rapid process related to recent N inputs to the canopy. We speculate that DON may arise from lichen and/or microbial N cycling rather than assimilation and re-release by tree tissues in this forest. Canopy retention of experimentally added N may meet and exceed calculated annual forest tree demand, although we do not know what fraction of retained N was actually physiologically assimilated by the plants. The observed retention and transformation of DIN within the canopy demonstrate that the fate and ecosystem consequences of N inputs from atmospheric deposition are likely influenced by forest canopy processes, which should be considered in N addition studies. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

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.