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Atmospheric deposition of nitrogen (N) influences forest demographics and carbon (C) uptake through multiple mechanisms that vary among tree species. Prior studies have estimated the effects of atmospheric N deposition on temperate forests by leveraging forest inventory measurements across regional gradients in deposition. However, in the United States (U.S.), these previous studies were limited in the number of species and the spatial scale of analysis, and did not include sulfur (S) deposition as a potential covariate. Here, we present a comprehensive analysis of how tree growth and survival for 71 species vary with N and S deposition across the conterminous U.S. Our analysis of 1,423,455 trees from forest plots inventoried between 2000 and 2016 reveals that the growth and/or survival of the vast majority of species in the analysis (n = 66, or 93%) were significantly affected by atmospheric deposition. Species co-occurred across the conterminous U.S. that had decreasing and increasing relationships between growth (or survival) and N deposition, with just over half of species responding negatively in either growth or survival to increased N deposition somewhere in their range (42 out of 71). Averaged across species and conterminous U.S., however, we found that an increase in deposition above current rates of N deposition would coincide with a small net increase in tree growth (1.7% per Δ kg N ha-1 yr-1), and a small net decrease in tree survival (-0.22% per Δ kg N ha-1 yr-1), with substantial regional and among-species variation. Adding S as a predictor improved the overall model performance for 70% of the species in the analysis. Our findings have potential to help inform ecosystem management and air pollution policy across the conterminous U.S., and suggest that N and S deposition have likely altered forest demographics in the U.S.

Human activity has significantly increased the deposition of nitrogen (N) on terrestrial ecosystems over pre‐industrial levels leading to a multitude of effects including losses of biodiversity, changes in ecosystem functioning, and impacts on human well‐being. It is challenging to explicitly link the level of deposition on an ecosystem to the cascade of ecological effects triggered and ecosystem services affected, because of the multitude of possible pathways in the N cascade. To address this challenge, we report on the activities of an expert workshop to synthesize information on N‐induced terrestrial eutrophication from the published literature and to link critical load exceedances with human beneficiaries by using the STressor–Ecological Production function–final ecosystem Services Framework and the Final Ecosystem Goods and Services Classification System (FEGS‐CS). We found 21 N critical loads were triggered by N deposition (ranging from 2 to 39 kg N·ha−1·yr−1), which cascaded to distinct beneficiary types through 582 individual pathways in the five ecoregions examined (Eastern Temperate Forests, Marine West Coast Forests, Northwestern Forested Mountains, North American Deserts, Mediterranean California). These exceedances ultimately affected 66 FEGS across a range of final ecosystem service categories (21 categories, e.g., changes in timber production, fire regimes, and native plant and animal communities) and 198 regional human beneficiaries of different types. Several different biological indicators were triggered in different ecosystems, including grasses and/or forbs (33% of all pathways), mycorrhizal communities (22%), tree species (21%), and lichen biodiversity (11%). Ecoregions with higher deposition rates for longer periods tended to have more numerous and varied ecological impacts (e.g., Eastern Temperate Forests, eight biological indicators) as opposed to other ecoregions (e.g., North American Deserts and Marine West Coast Forests each with one biological indicator). Nonetheless, although ecoregions differed by ecological effects from terrestrial eutrophication, the number of FEGS and beneficiaries impacted was similar across ecoregions. We found that terrestrial eutrophication affected all ecosystems examined, demonstrating the widespread nature of terrestrial eutrophication nationally. These results highlight which people and ecosystems are most affected according to present knowledge, and identify key uncertainties and knowledge gaps to be filled by future research.

In the western United States vast acreages of land are exposed to low levels of atmospheric nitrogen (N) deposition, with interspersed hotspots of elevated N deposition downwind of large, expanding metropolitan centers or large agricultural operations. Biological response studies in western North America demonstrate that some aquatic and terrestrial plant and microbial communities are significantly altered by N deposition. Greater plant productivity is counterbalanced by biotic community changes and deleterious effects on sensitive organisms (lichens and phytoplankton) that respond to low inputs of N (3 to 8 kilograms N per hectare per year). Streamwater nitrate concentrations are elevated in high-elevation catchments in Colorado and are unusually high in southern California and in some chaparral catchments in the southwestern Sierra Nevada. Chronic N deposition in the West is implicated in increased fire frequency in some areas and habitat alteration for threatened species. Between hotspots, N deposition is too low to cause noticeable effects or has not been studied.

Anthropogenic nitrogen (N) deposition is known to reduce plant diversity in ecosystems worldwide; however, effects on the diversity of Mediterranean‐type ecosystems—global hotspots of biodiversity—are relatively unexplored. In California, elevated N deposition due to air pollution has a multitude of ecological effects including the facilitation of nonnative plant invasion and altered ecosystem functioning, but impacts on plant richness have been inadequately quantified. We addressed this research gap by evaluating patterns of plant richness in coastal sage scrub (CSS), a severely threatened, highly diverse Mediterranean‐type shrubland, across the Santa Monica Mountains National Recreation Area. This is the largest urban national park in the United States and experiences a strong gradient of N deposition due to its proximity to urban Los Angeles. We measured soil N, plant cover, and richness at 30 CSS sites across this gradient and used regression analyses to explore relationships between richness, N deposition, and other environmental variables. We observed significant declines in plant richness across a steep gradient of soil N availability that paralleled patterns of N deposition, primarily due to decreases in native forb species. Our analyses identified soil N as the best predictor of patterns of native forb richness, but other factors, including nonnative plant cover and aridity, may also drive reduced richness. In addition to the marked decline in the number of native forb species, increasing N deposition was also associated with lower native shrub richness per area and increased cover of nonnatives. These results highlight the threat posed by N deposition to the conservation of this already imperiled ecosystem under continued environmental change.

[This corrects the article DOI: 10.1371/journal.pone.0205296.].

Wet plus dry deposition of both reduced (i.e., ammonia, ammonium) and oxidized (i.e., nitric acid vapor, various nitrogen oxides, nitrate) N forms should be included in any assessment of N deposition effects on water quality, either as a nutrient effect or when considering acidification effects. [...]the use of the sum of nitrate and sulfate in wet deposition expressed on a mass basis (kilograms per hectare) as a measure of deposition inputs (Brown and Froemke 2012) is also highly unconventional.

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 N·ha −1 ·yr −1 , 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.

We review the mechanisms of deleterious nitrogen (N) deposition impacts on temperate forests, with a particular focus on trees and lichens. Elevated anthropogenic N deposition to forests has varied effects on individual organisms depending on characteristics both of the N inputs (form, timing, amount) and of the organisms (ecology, physiology) involved. Improved mechanistic knowledge of these effects can aid in developing robust predictions of how organisms respond to either increases or decreases in N deposition. Rising N levels affect forests in micro‐ and macroscopic ways from physiological responses at the cellular, tissue, and organism levels to influencing individual species and entire communities and ecosystems. A synthesis of these processes forms the basis for the overarching themes of this paper, which focuses on N effects at different levels of biological organization in temperate forests. For lichens, the mechanisms of direct effects of N are relatively well known at cellular, organismal, and community levels, though interactions of N with other stressors merit further research. For trees, effects of N deposition are better understood for N as an acidifying agent than as a nutrient; in both cases, the impacts can reflect direct effects on short time scales and indirect effects mediated through long‐term soil and belowground changes. There are many gaps on fundamental N use and cycling in ecosystems, and we highlight the most critical gaps for understanding potential deleterious effects of N deposition. For lichens, these gaps include both how N affects specific metabolic pathways and how N is metabolized. For trees, these gaps include understanding the direct effects of N deposition onto forest canopies, the sensitivity of different tree species and mycorrhizal symbionts to N, the influence of soil properties, and the reversibility of N and acidification effects on plants and soils. Continued study of how these N response mechanisms interact with one another, and with other dimensions of global change, remains essential for predicting ongoing changes in lichen and tree populations across North American temperate forests.

Nitrogen (N) deposition in the western United States ranges from 1 to 4 kilograms (kg) per hectare (ha) per year over much of the region to as high as 30 to 90 kg per ha per year downwind of major urban and agricultural areas. Primary N emissions sources are transportation, agriculture, and industry. Emissions of N as ammonia are about 50% as great as emissions of N as nitrogen oxides. An unknown amount of N deposition to the West Coast originates from Asia. Nitrogen deposition has increased in the West because of rapid increases in urbanization, population, distance driven, and large concentrated animal feeding operations. Studies of ecological effects suggest that emissions reductions are needed to protect sensitive ecosystem components. Deposition rates are unknown for most areas in the West, although reasonable estimates are available for sites in California, the Colorado Front Range, and central Arizona. National monitoring networks provide long-term wet deposition data and, more recently, estimated dry deposition data at remote sites. However, there is little information for many areas near emissions sources.

Net fluxes of nitrogen (N) mineralization and nitrification were measured in situ on a monthly basis for 3 years at a high (HN) and low (LN) N deposition site in the San Bernardino Mountains, California. Mean N mineralization fluxes in the forest floor and top 10 cm of mineral soil were 19.0 and 59.8 kg N.ha(-1).year(-1) at LN and HN, respectively. Mean net nitrification fluxes were 11.2 and 55.9 kg N.ha(-1).year(-1) at LN and HN, respectively. Relative nitrification (the percent N mineralized that was nitrified) was generally lower under Pinus ponderosa Dougl. ex P. & C. Laws. (or Pinus jeffreyi Grev. & Balf.) canopies than under Quercus kelloggii Newb. or open canopies. The rate of net N mineralization was the key factor for predicting the rate of net nitrification. Fertilization with 50 and 150 kg N.ha(-1) at LN significantly increased the rates of net mineralization and net nitrification. At HN fertilization had no significant effect on net nitrification. We conclude that at low-deposition sites increased nitrification occurs in the short term in response to added N, but that sustained elevated net nitrification is driven by the accumulation of N-enriched litter and soil organic matter in conjunction with chronic throughfall N deposition inputs.