Abstract. Atmospheric nitrogen (N) deposition is a recognized threat to plant diversity in temperate and northern parts of Europe and North America. This paper assesses evidence from field experiments for N deposition effects and thresholds for terrestrial plant diversity protection across a latitudinal range of main categories of ecosystems, from arctic and boreal systems to tropical forests. Current thinking on the mechanisms of N deposition effects on plant diversity, the global distribution of G200 ecoregions, and current and future (2030) estimates of atmospheric N-deposition rates are then used to identify the risks to plant diversity in all major ecosystem types now and in the future.This synthesis paper clearly shows that N accumulation is the main driver of changes to species composition across the whole range of different ecosystem types by driving the competitive interactions that lead to composition change and/or making conditions unfavorable for some species. Other effects such as direct toxicity of nitrogen gases and aerosols, long-term negative effects of increased ammonium and ammonia availability, soil-mediated effects of acidification, and secondary stress and disturbance are more ecosystem-and site-specific and often play a supporting role. N deposition effects in mediterranean ecosystems have now been identified, leading to a first estimate of an effect threshold. Importantly, ecosystems thought of as not N limited, such as tropical and subtropical systems, may be more vulnerable in the regeneration phase, in situations where heterogeneity in N availability is reduced by atmospheric N deposition, on sandy soils, or in montane areas.Critical loads are effect thresholds for N deposition, and the critical load concept has helped European governments make progress toward reducing N loads on sensitive ecosystems. More needs to be done in Europe and North America, especially for the more sensitive ecosystem types, including several ecosystems of high conservation importance.The results of this assessment show that the vulnerable regions outside Europe and North America which have not received enough attention are ecoregions in eastern and southern Asia (China, India), an important part of the mediterranean ecoregion (California, southern Europe), and in the coming decades several subtropical and tropical parts of Latin America and Africa. Reductions in plant diversity by increased atmospheric N deposition may be more widespread than first thought, and more targeted studies are required in low background areas, especially in the G200 ecoregions.

SummaryRatios of nitrogen (N) isotopes in leaves could elucidate underlying patterns of N cycling across ecological gradients. To better understand global-scale patterns of N cycling, we compiled data on foliar N isotope ratios (δ 15 N), foliar N concentrations, mycorrhizal type and climate for over 11 000 plants worldwide. Arbuscular mycorrhizal, ectomycorrhizal, and ericoid mycorrhizal plants were depleted in foliar δ 15 N by 2‰, 3.2‰, 5.9‰, respectively, relative to nonmycorrhizal plants. Foliar δ 15 N increased with decreasing mean annual precipitation and with increasing mean annual temperature (MAT) across sites with MAT ≥ −0.5°C, but was invariant with MAT across sites with MAT < −0.5°C. In independent landscape-level to regionallevel studies, foliar δ 15 N increased with increasing N availability; at the global scale, foliar δ 15 N increased with increasing foliar N concentrations and decreasing foliar phosphorus (P) concentrations. Together, these results suggest that warm, dry ecosystems have the highest N availability, while plants with high N concentrations, on average, occupy sites with higher N availability than plants with low N concentrations. Global-scale comparisons of other components of the N cycle are still required for better mechanistic understanding of the determinants of variation in foliar δ 15 N and ultimately global patterns in N cycling.

Abstract. Human activity in the last century has led to a significant increase in nitrogen (N) emissions and atmospheric deposition. This N deposition has reached a level that has caused or is likely to cause alterations to the structure and function of many ecosystems across the United States. One approach for quantifying the deposition of pollution that would be harmful to ecosystems is the determination of critical loads. A critical load is defined as the input of a pollutant below which no detrimental ecological effects occur over the long-term according to present knowledge.The objectives of this project were to synthesize current research relating atmospheric N deposition to effects on terrestrial and freshwater ecosystems in the United States, and to estimate associated empirical N critical loads. The receptors considered included freshwater diatoms, mycorrhizal fungi, lichens, bryophytes, herbaceous plants, shrubs, and trees. Ecosystem impacts included: (1) biogeochemical responses and (2) individual species, population, and community responses. Biogeochemical responses included increased N mineralization and nitrification (and N availability for plant and microbial uptake), increased gaseous N losses (ammonia volatilization, nitric and nitrous oxide from nitrification and denitrification), and increased N leaching. Individual species, population, and community responses included increased tissue N, physiological and nutrient imbalances, increased growth, altered root : shoot ratios, increased susceptibility to secondary stresses, altered fire regime, shifts in competitive interactions and community composition, changes in species richness and other measures of biodiversity, and increases in invasive species.The range of critical loads for nutrient N reported for U.S. ecoregions, inland surface waters, and freshwater wetlands is 1-39 kg NÁha , spanning the range of N deposition observed over most of the country. The empirical critical loads for N tend to increase in the following sequence for different life forms: diatoms, lichens and bryophytes, mycorrhizal fungi, herbaceous plants and shrubs, and trees.The critical load approach is an ecosystem assessment tool with great potential to simplify complex scientific information and communicate effectively with the policy community and the public. This synthesis represents the first comprehensive assessment of empirical critical loads of N for major ecoregions across the United States.

Atmospheric nitrogen (N) deposition has been shown to decrease plant species richness along regional deposition gradients in Europe and in experimental manipulations. However, the general response of species richness to N deposition across different vegetation types, soil conditions, and climates remains largely unknown even though responses may be contingent on these environmental factors. We assessed the effect of N deposition on herbaceous richness for 15,136 forest, woodland, shrubland, and grassland sites across the continental United States, to address how edaphic and climatic conditions altered vulnerability to this stressor. In our dataset, with N deposition ranging from 1 to 19 kg N·ha, we found a unimodal relationship; richness increased at low deposition levels and decreased above 8.7 and 13.4 kg N·ha −1 ·y−1 in open and closed-canopy vegetation, respectively. N deposition exceeded critical loads for loss of plant species richness in 24% of 15,136 sites examined nationwide. There were negative relationships between species richness and N deposition in 36% of 44 community gradients. Vulnerability to N deposition was consistently higher in more acidic soils whereas the moderating roles of temperature and precipitation varied across scales. We demonstrate here that negative relationships between N deposition and species richness are common, albeit not universal, and that fine-scale processes can moderate vegetation responses to N deposition. Our results highlight the importance of contingent factors when estimating ecosystem vulnerability to N deposition and suggest that N deposition is affecting species richness in forested and nonforested systems across much of the continental United States.nitrogen deposition | plant species richness | diversity | soil pH | climate

N saturation induced by atmospheric N deposition can have serious consequences for forest health in many regions. In order to evaluate whether foliar d 15 N may be a robust, regional-scale measure of the onset of N saturation in forest ecosystems, we assembled a large dataset on atmospheric N deposition, foliar and root d 15 N and N concentration, soil C:N, mineralization and nitrification. The dataset included sites in northeastern North America, Colorado, Alaska, southern Chile and Europe. Local drivers of N cycling (net nitrification and mineralization, and forest floor and soil C:N) were more closely coupled with foliar d 15 N than the regional driver of N deposition. Foliar d 15 N increased non-linearly with nitrification:mineralization ratio and decreased with forest floor C:N. Foliar d 15 N was more strongly related to nitrification rates than was foliar N concentration, but concentration was more strongly correlated with N deposition. Root d 15 N was more tightly coupled to forest floor properties than was foliar d 15 N. We observed a pattern of decreasing foliar d 15 N values across the following species: American beech>yellow birch>sugar maple. Other factors that affected foliar d 15 N included species composition and climate. Relationships between foliar d 15 N and soil variables were stronger when analyzed on a species by species basis than when many species were lumped. European sites showed distinct patterns of lower foliar d 15 N, due to the importance of ammonium deposition in this region. Our results suggest that examining d 15 N values of foliage may improve understanding of how forests respond to the cascading effects of N deposition.

Increased losses of nitrate from watersheds may accelerate the depletion of nutrient cations and affect the acidification and trophic status of surface waters. Patterns of nitrate concentrations and losses were evaluated in four forested watersheds (East Bear Brook Watershed, Lead Mountain, ME; Watershed 6, Hubbard Brook Experimental Forest, White Mountains, NH; Arbutus Watershed, Huntington Forest, Adirondack Mountains, NY; Biscuit Brook, Catskill Mountains, NY) located across the northeastern United States. A synchronous pattern was observed in nitrate concentrations of drainage waters from these four sites from 1983 through 1993. Most notably, high concentrations and high drainage water losses followed an anomalous cold period (mean daily temperature −11.4 to −16 °C in December 1989) for all four sites. After high nitrate losses during the snowmelt of 1990, nitrate concentrations and fluxes decreased at all sites. These results suggest that climatic variation can have a major effect on nitrogen flux and cycling and may influence temporal patterns of nitrate loss in a region.

Abstract:The natural abundance of nitrogen and oxygen isotopes in nitrate can be a powerful tool for identifying the source of nitrate in streamwater in forested watersheds, because the two main sources of nitrate, atmospheric deposition and microbial nitrification, have distinct υ18 O values. Using a simple mixing model, we estimated the relative fractions in streamwater derived from these sources for two forested watersheds with markedly different streamwater nitrate outputs. In this study, we monitored υ 15 N and υ 18

Export of microbially produced nitrate from an ecosystem is expected to increase δ15N in the remaining soil organic matter and NH4+. To test the hypothesis that nitrification and nitrate loss induced by clear-cutting cause an increase in soil and foliar δ15N, we measured δ15N in a clear-cut watershed at the Hubbard Brook Experimental Forest, New Hampshire. δ15N ranged from –0.02‰ in the Oie horizon to 7.7‰ in the Bs2 horizon prior to clear-cutting and increased significantly by 1.3‰ in the Oie horizon and 0.9‰ in the Oa horizon 3 years after clear-cutting. Fifteen years after clear-cutting, δ15N in both O horizons decreased to near-initial values. No significant temporal changes in the Bs2 and C horizons δ15N were observed. Foliar δ15N was highest (1.7‰) the first 2 years after clear-cutting and was significantly higher than in the reference watershed (mean δ15N = –1.2‰), decreasing to 0.0‰ 3–5 years after clear-cutting and to –1.3‰ 9–11 years after clear-cutting. Increased foliar δ15N coincided with increased stream-water nitrate concentration, suggesting that the increased nitrification responsible for elevated stream-water nitrate may also have caused an enrichment of the plant-available ammonium pool. The response observed in this catchment also suggests that sampling of soil or foliar δ15N may provide a practical alternative to long time series of stream chemistry for evaluating nitrogen saturation of forested ecosystems.