Abstract. Human activities have clearly caused dramatic alterations of the terrestrial nitrogen cycle, and analyses of the extent and effects of such changes are now common in the scientific literature. However, any attempt to evaluate N cycling processes within ecosystems, as well as anthropogenic influences on the N cycle, requires an understanding of the magnitude of inputs via biological nitrogen fixation (BNF). Although there have been many studies addressing the microbiology, physiology, and magnitude of N fixation at local scales, there are very few estimates of BNF over large scales. We utilized >100 preexisting published estimates of BNF to generate biome-and global-level estimates of biological N fixation. We also used net primary productivity (NPP) and evapotranspiration (ET) estimates from the Century terrestrial ecosystem model to examine global relationships between these variables and BNF as well as to compare observed and Century-modeled BNF. Our data-based estimates showed a strong positive relationship between ecosystem ET and BNF, and our analyses suggest that while the model's simple relationships for BNF predict broad scale patterns, they do not capture much of the variability or magnitude of published rates. Patterns of BNF were also similar to patterns of ecosystem NPP. Our "best estimate" of potential nitrogen fixation by natural ecosystems is -195 Tg N yr -•, with a range of 100-290 Tg N yr '•. Although these estimates do not account for the decrease in natural N fixation due to cultivation, this would not dramatically alter our estimate, as the greatest reductions in area have occurred in systems characterized by relatively low rates of N fixation (e.g., grasslands). Although our estimate of BNF in natural ecosystems is similar to previously published estimates of terrestrial BNF, we believe that this study provides a more documented, constrained estimate of this important flux.

Conceptual and numerical models of nitrogen cycling in temperate forests assume that nitrogen is lost from these ecosystems predominantly by way of inorganic forms, such as nitrate and ammonium ions. Of these, nitrate is thought to be particularly mobile, being responsible for nitrogen loss to deep soil and stream waters. But human activities-such as fossil fuel combustion, fertilizer production and land-use change-have substantially altered the nitrogen cycle over large regions, making it difficult to separate natural aspects of nitrogen cycling from those induced by human perturbations. Here we report stream chemistry data from 100 unpolluted primary forests in temperate South America. Although the sites exhibit a broad range of environmental factors that influence ecosystem nutrient cycles (such as climate, parent material, time of ecosystem development, topography and biotic diversity), we observed a remarkably consistent pattern of nitrogen loss across all forests. In contrast to findings from forests in polluted regions, streamwater nitrate concentrations are exceedingly low, such that nitrate to ammonium ratios were less than unity, and dissolved organic nitrogen is responsible for the majority of nitrogen losses from these forests. We therefore suggest that organic nitrogen losses should be considered in models of forest nutrient cycling, which could help to explain observations of nutrient limitation in temperate forest ecosystems.

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.

Abstract. Effects of anthropogenic nitrogen (N) deposition and the ability of terrestrial ecosystems to store carbon (C) depend in part on the amount of N retained in the system and its partitioning among plant and soil pools. We conducted a meta-analysis of studies at 48 sites across four continents that used enriched 15 N isotope tracers in order to synthesize information about total ecosystem N retention (i.e., total ecosystem 15 N recovery in plant and soil pools) across natural systems and N partitioning among ecosystem pools. The greatest recoveries of ecosystem 15 N tracer occurred in shrublands (mean, 89.5%) and wetlands (84.8%) followed by forests (74.9%) and grasslands (51.8%). In the short term (,1 week after 15 N tracer application), total ecosystem 15 N recovery was negatively correlated with fine-root and soil 15 N natural abundance, and organic soil C and N concentration but was positively correlated with mean annual temperature and mineral soil C:N. In the longer term tracer were below ground in forests, shrublands, and grasslands, we conclude that growth enhancement and potential for increased C storage in aboveground biomass from atmospheric N deposition is likely to be modest in these ecosystems. Total ecosystem 15 N recovery decreased with N fertilization, with an apparent threshold fertilization rate of 46 kg NÁha À1 Áyr À1 above which most ecosystems showed net losses of applied 15 N tracer in response to N fertilizer addition.

Quantifying global patterns of terrestrial nitrogen (N) cycling is central to predicting future patterns of primary productivity, carbon sequestration, nutrient fluxes to aquatic systems, and climate forcing. With limited direct measures of soil N cycling at the global scale, syntheses of the 15N:14N ratio of soil organic matter across climate gradients provide key insights into understanding global patterns of N cycling. In synthesizing data from over 6000 soil samples, we show strong global relationships among soil N isotopes, mean annual temperature (MAT), mean annual precipitation (MAP), and the concentrations of organic carbon and clay in soil. In both hot ecosystems and dry ecosystems, soil organic matter was more enriched in 15N than in corresponding cold ecosystems or wet ecosystems. Below a MAT of 9.8°C, soil δ15N was invariant with MAT. At the global scale, soil organic C concentrations also declined with increasing MAT and decreasing MAP. After standardizing for variation among mineral soils in soil C and clay concentrations, soil δ15N showed no consistent trends across global climate and latitudinal gradients. Our analyses could place new constraints on interpretations of patterns of ecosystem N cycling and global budgets of gaseous N loss.

Nitrogen retention and recycling are topics of enduring interest in ecosystem ecology, yet we lack a mechanistic field-tested model of how these processes work in unpolluted, old-growth temperate forests. Forests of the Cordillera Piuchué Ecosystem Study (CPES) in southern Chile provide an opportunity to examine nitrogen cycling and retention in a forest that is virtually free of human disturbance. We applied 15 N pool dilution and pulse-chase tracer techniques as complementary approaches within small plots to understand flows of inorganic nitrogen in the surface soil of an evergreen mixed-angiosperm forest. We also followed separate pulses of 15 NH 4 ϩ and 15 NO 3 Ϫ for two years to gain insights into how short-term mechanisms of inorganic nitrogen cycling translate into long-term patterns of ecosystem nitrogen retention.Strong consumption appears to limit losses of NH 4 ϩ and NO 3 Ϫ from this forest, and predominantly by the same mechanisms for both forms of nitrogen. As a result, the extent of 15 NH 4 ϩ and 15 NO 3 Ϫ retention were also similar, yet ϳ44-fold higher rates of gross NH 4 ϩ production leads to the dominance of NH 4 ϩ over NO 3 Ϫ in soil and stream waters. Microbial biomass played a key role in the short-term assimilation of 15 N tracers, but retention was only transient. Turnover of 15 N through microbial biomass was rapid and appeared to be only weakly retained in soil exchangeable pools, fine roots, and soil organic matter, resulting in substantial losses of 15 N from soils within weeks of tracer additions. Assimilation of 15 N into fine roots was a much larger sink (13%) than has been reported for other forested ecosystems (1-3%), and the transport of 15 N from microbial biomass to aboveground sinks in vegetation may explain the observed loss of 15 N from surface soils over time. Losses of 15 N from microbial biomass did not enter the extractable pool of dissolved organic nitrogen (DON), suggesting that DON losses do not originate directly from active microbial turnover, and also that microbial activity may not exert as much control over hydrologic losses of DON as compared to losses of NH 4 ϩ and NO 3 Ϫ . Our results also suggest an additional rapid and extremely transient (1 d) mechanism of NO 3 Ϫ retention via incorporation into extractable-DON.The long-term retention of 15 N at the whole-plot level did not differ significantly between 15 NH 4 ϩ and 15 NO 3 Ϫ treatments, and averaged 65% after two years. The lack of an appreciable change in 15 N recovery for ϳ1.5 yr following the initial assimilation, redistribution, and loss of 15 N suggests that the majority of 15 N was not recycled over the long term through inorganic nitrogen pools and microbial biomass via mineralization/immobilization pathways. Instead, long-term retention of inorganic 15 N appeared to be dominated by rapid and possibly direct assimilation into a slow-turnover pool of soil organic matter. Elevated 15 N contents in fine-root and microbial pools for up to two years after 15 N additions, however, also indicated sustained ...

Few high‐elevation tropical catchments worldwide are gauged, and even fewer are studied using combined hydrometric and isotopic data. Consequently, we lack information needed to understand processes governing rainfall–runoff dynamics and to predict their influence on downstream ecosystem functioning. To address this need, we present a combination of hydrometric and water stable isotopic observations in the wet Andean páramo ecosystem of the Zhurucay Ecohydrological Observatory (7.53 km2). The catchment is located in the Andes of south Ecuador between 3400 and 3900 m a.s.l. Water samples for stable isotopic analysis were collected during 2 years (May 2011–May 2013), while rainfall and runoff measurements were continuously recorded since late 2010. The isotopic data reveal that andosol soils predominantly situated on hillslopes drain laterally to histosols (Andean páramo wetlands) mainly located at the valley bottom. Histosols, in turn, feed water to creeks and small rivers throughout the year, establishing hydrologic connectivity between wetlands and the drainage network. Runoff is primarily composed of pre‐event water stored in the histosols, which is replenished by rainfall that infiltrates through the andosols. Contributions from the mineral horizon and the top of the fractured bedrock are small and only seem to influence discharge in small catchments during low flow generation (non‐exceedance flows < Q35). Variations in source contributions are controlled by antecedent soil moisture, rainfall intensity, and duration of rainy periods. Saturated hydraulic conductivity of the soils, higher than the year‐round low precipitation intensity, indicates that Hortonian overland flow rarely occurs during high‐intensity precipitation events. Deep groundwater contributions to discharge seem to be minimal. These results suggest that, in this high‐elevation tropical ecosystem, (1) subsurface flow is a dominant hydrological process and (2) (histosols) wetlands are the major source of stream runoff. Our study highlights that detailed isotopic characterization during short time periods provides valuable information about ecohydrological processes in regions where very few basins are gauged. Copyright © 2016 John Wiley & Sons, Ltd.

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