Global change progressively increases foliar nitrogen-phosphorus ratios in China's subtropical forests.

Globally increased nitrogen (N) to phosphorus (P) ratios (N/P) affect the structure and functioning of terrestrial ecosystems, but few studies have addressed the variation of foliar N/P over time in subtropical forests. Foliar N/P indicates N versus P limitation in terrestrial ecosystems. Quantifying long-term dynamics of foliar N/P and their potential drivers is crucial for predicting nutrient status and functioning in forest ecosystems under global change. We detected temporal trends of foliar N/P, quantitatively estimated their potential drivers and their interaction between plant types (evergreen vs. deciduous and trees vs. shrubs), using 1811 herbarium specimens of 12 widely distributed species collected during 1920-2010 across China's subtropical forests. We found significant decreases in foliar P concentrations (23.1%) and increases in foliar N/P (21.2%). Foliar N/P increased more in evergreen species (22.9%) than in deciduous species (16.9%). Changes in atmospheric CO2 concentrations ( P CO 2 $$ {\mathrm{P}}_{{\mathrm{CO}}_2} $$ ), atmospheric N deposition and mean annual temperature (MAT) dominantly contributed to the increased foliar N/P of evergreen species, while P CO 2 $$ {\mathrm{P}}_{{\mathrm{CO}}_2} $$ , MAT, and vapor pressure deficit, to that of deciduous species. Under future Shared Socioeconomic Pathway (SSP) scenarios, increasing MAT and P CO 2 $$ {\mathrm{P}}_{{\mathrm{CO}}_2} $$ would continuously increase more foliar N/P in deciduous species than in evergreen species, with more 12.9%, 17.7%, and 19.4% versus 6.1%, 7.9%, and 8.9% of magnitudes under the scenarios of SSP1-2.6, SSP3-7.0, and SSP5-8.5, respectively. The results suggest that global change has intensified and will progressively aggravate N-P imbalance, further altering community composition and ecosystem functioning of subtropical forests.

Nitrogen deposition accelerates soil carbon sequestration in tropical forests.

Terrestrial ecosystem carbon (C) sequestration plays an important role in ameliorating global climate change. While tropical forests exert a disproportionately large influence on global C cycling, there remains an open question on changes in below-ground soil C stocks with global increases in nitrogen (N) deposition, because N supply often does not constrain the growth of tropical forests. We quantified soil C sequestration through more than a decade of continuous N addition experiment in an N-rich primary tropical forest. Results showed that long-term N additions increased soil C stocks by 7 to 21%, mainly arising from decreased C output fluxes and physical protection mechanisms without changes in the chemical composition of organic matter. A meta-analysis further verified that soil C sequestration induced by excess N inputs is a general phenomenon in tropical forests. Notably, soil N sequestration can keep pace with soil C, based on consistent C/N ratios under N additions. These findings provide empirical evidence that below-ground C sequestration can be stimulated in mature tropical forests under excess N deposition, which has important implications for predicting future terrestrial sinks for both elevated anthropogenic CO2 and N deposition. We further developed a conceptual model hypothesis depicting how soil C sequestration happens under chronic N deposition in N-limited and N-rich ecosystems, suggesting a direction to incorporate N deposition and N cycling into terrestrial C cycle models to improve the predictability on C sink strength as enhanced N deposition spreads from temperate into tropical systems.

The University of West Florida Campus Ecosystem Study: effects of forest vegetation on light availability and soil processes.

College and university campuses with a notable arboreal component provide unique opportunities for carrying out ecological research. The University of West Florida Campus Ecosystem Study (UWF CES) was established in 2019 as interconnected research to take advantage of the extensive arborescent nature of the UWF campus, particularly concerning longleaf pine (Pinus palustris). One of these investigations established permanent plots in forested sites of two contrasting types, one dominated by longleaf pine ("pine site") and the other dominated by hardwoods ('hardwood site'). This study used these plots to examine the influence of forest vegetation on light availability and soil processes. Light was measured as photosynthetically active radiation (and expressed as photon flux density-PFD) with a handheld meter in each plot. Soil was sampled to 5 cm in each plot; texture was measured with the hydrometer method. Identical sampling methods were carried out in a persistent canopy opening to assess light and soil conditions under maximum solar radiation. Mean PFD was ~4× higher in pine stands than in hardwood stands; PFD was 12.8 and 3.5% of full light in the pine and hardwood stands, respectively. All soils were dominated by coarse-textured sands, but silt was significantly higher in pine stand soil and higher still in the canopy opening. Among forest stand plots, sand was negatively related to PFD, whereas clay was positively related to PFD. Across the three sites, silt was positively related to PFD. These relationships are consistent with the importance of solar radiation as one of many drivers of soil weathering.

Arbuscular Mycorrhizal Fungi Shift Soil Bacterial Community Composition and Reduce Soil Ammonia Volatilization and Nitrous Oxide Emissions.

Arbuscular mycorrhizal fungi (AMF) establish mutualistic relationships with the majority of terrestrial plants, increasing plant uptake of soil nitrogen (N) in exchange for photosynthates. And may influence soil ammonia (NH3) volatilization and nitrous oxide (N2O) emissions directly by improving plant N uptake, and/or indirectly by modifying soil bacterial community composition for the soil C availability increasing. However, the effects of AMF on soil NH3 volatilization and N2O emissions and their underlying mechanisms remain unclear. We carried out two independent experiments using contrasting methods, one with a compartmental box device (in 2016) and the other with growth pot experiment (in 2020) to examine functional relationships between AMF and soil NH3 volatilization and N2O emissions under varying N input. The presence of AMF significantly reduced soil NH3 volatilization and N2O emissions while enhancing plant biomass and plant N acquisition, and reducing soil NH4+ and NO3-, even with high N input. The presence of AMF also significantly reduced the relative abundance within the bacterial orders Sphingomonadales and Rhizobiales. Sphingomonadales correlated significantly and positively with soil NH3 volatilization in 2016 and N2O emissions, whereas Rhizobiales correlated positively with soil N2O emissions. High N input significantly increased soil NH3 volatilization and N2O emissions with increasing relative abundance of Sphingomonadales and Rhizobiales. These findings demonstrate the contribution of AMF in regulating NH3 and N2O emission by improving plant N uptake and altering soil bacterial communities. They also suggest that altering the rhizosphere microbiome might offer additional potential for restoration of N-enriched agroecosystems.

Drivers of foliar 15 N trends in southern China over the last century.

Foliar stable nitrogen (N) isotopes (δ15 N) generally reflect N availability to plants and have been used to infer about changes thereof. However, previous studies of temporal trends in foliar δ15 N have ignored the influence of confounding factors, leading to uncertainties on its indication to N availability. In this study, we measured foliar δ15 N of 1811 herbarium specimens from 12 plant species collected in southern China forests from 1920 to 2010. We explored how changes in atmospheric CO2 , N deposition and global warming have affected foliar δ15 N and N concentrations ([N]) and identified whether N availability decreased in southern China. Across all species, foliar δ15 N significantly decreased by 0.82‰ over the study period. However, foliar [N] did not decrease significantly, implying N homeostasis in forest trees in the region. The spatiotemporal patterns of foliar δ15 N were explained by mean annual temperature (MAT), atmospheric CO2 ( P CO 2 ), atmospheric N deposition, and foliar [N]. The spatiotemporal trends of foliar [N] were explained by MAT, temperature seasonality, P CO 2 , and N deposition. N deposition within the rates from 5.3 to 12.6 kg N ha-1  year-1 substantially contributed to the temporal decline in foliar δ15 N. The decline in foliar δ15 N was not accompanied by changes in foliar [N] and therefore does not necessarily reflect a decline in N availability. This is important to understand changes in N availability, which is essential to validate and parameterize biogeochemical cycles of N.

Arbuscular Mycorrhizae Shift Community Composition of N-Cycling Microbes and Suppress Soil N2O Emission.

Mycorrhizae are ubiquitous symbiotic associations between arbuscular mycorrhizal fungi (AMF) and terrestrial plants, in which AMF receive photosynthates from and acquire soil nutrients for their host plants. Plant uptake of soil nitrogen (N) reduces N substrate for microbial processes that generate nitrous oxide (N2O), a potent greenhouse gas. However, the underlying microbial mechanisms remain poorly understood, particularly in agroecosystems with high reactive N inputs. We examined how plant roots and AMF affect N2O emissions, N2O-producing (nirK and nirS) and N2O-consuming (nosZ) microbes under normal and high N inputs in conventional (CONV) and organically managed (OM) soils. Here, we show that high N input increased soil N2O emissions and the ratio of nirK to nirS microbes. Roots and AMF did not affect the (nirK + nirS)/nosZ ratio but significantly reduced N2O emissions and the nirK/nirS ratio. They reduced the nirK/nirS ratio by reducing nirK-Rhodobacterales but increasing nirS-Rhodocyclales in the CONV soil while decreasing nirK-Burkholderiales but increasing nirS-Rhizobiales in the OM soil. Our results indicate that plant roots and AMF reduced N2O emission directly by reducing soil N and indirectly through shifting the community composition of N2O-producing microbes in N-enriched agroecosystems, suggesting that harnessing the rhizosphere microbiome through agricultural management might offer additional potential for N2O emission mitigation.

Nitrogen addition stimulates soil aggregation and enhances carbon storage in terrestrial ecosystems of China: A meta-analysis.

China is experiencing a high level of atmospheric nitrogen (N) deposition, which greatly affects the soil carbon (C) dynamics in terrestrial ecosystems. Soil aggregation contributes to the stability of soil structure and to soil C sequestration. Although many studies have reported the effects of N enrichment on bulk soil C dynamics, the underlying mechanisms explaining how soil aggregates respond to N enrichment remain unclear. Here, we used a meta-analysis of data from 76N manipulation experiments in terrestrial ecosystems in China to assess the effects of N enrichment on soil aggregation and its sequestration of C. On average, N enrichment significantly increased the mean weight diameter of soil aggregates by 10%. The proportion of macroaggregates and silt-clay fraction were significantly increased (6%) and decreased (9%) by N enrichment, respectively. A greater response of macroaggregate C (+15%) than of bulk soil C (+5%) to N enrichment was detected across all ecosystems. However, N enrichment had minor effects on microaggregate C and silt-clay C. The magnitude of N enrichment effect on soil aggregation varied with ecosystem type and fertilization regime. Additionally, soil pH declined consistently and was correlated with soil aggregate C. Overall, our meta-analysis suggests that N enrichment promotes particulate organic C accumulation via increasing macroaggregate C and acidifying soils. In contrast, increases in soil aggregation could inhibit microbially mediated breakdown of soil organic matter, causing minimal change in mineral-associated organic C. Our findings highlight that atmospheric N deposition may enhance the formation of soil aggregates and their sequestration of C in terrestrial ecosystems in China.

Plant acclimation to long-term high nitrogen deposition in an N-rich tropical forest.

Anthropogenic nitrogen (N) deposition has accelerated terrestrial N cycling at regional and global scales, causing nutrient imbalance in many natural and seminatural ecosystems. How added N affects ecosystems where N is already abundant, and how plants acclimate to chronic N deposition in such circumstances, remains poorly understood. Here, we conducted an experiment employing a decade of N additions to examine ecosystem responses and plant acclimation to added N in an N-rich tropical forest. We found that N additions accelerated soil acidification and reduced biologically available cations (especially Ca and Mg) in soils, but plants maintained foliar nutrient supply at least in part by increasing transpiration while decreasing soil water leaching below the rooting zone. We suggest a hypothesis that cation-deficient plants can adjust to elevated N deposition by increasing transpiration and thereby maintaining nutrient balance. This result suggests that long-term elevated N deposition can alter hydrological cycling in N-rich forest ecosystems.

Conditional vulnerability of plant diversity to atmospheric nitrogen deposition 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(-1)⋅y(-1), 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.

Global environmental change effects on plant community composition trajectories depend upon management legacies.

The contemporary state of functional traits and species richness in plant communities depends on legacy effects of past disturbances. Whether temporal responses of community properties to current environmental changes are altered by such legacies is, however, unknown. We expect global environmental changes to interact with land-use legacies given different community trajectories initiated by prior management, and subsequent responses to altered resources and conditions. We tested this expectation for species richness and functional traits using 1814 survey-resurvey plot pairs of understorey communities from 40 European temperate forest datasets, syntheses of management transitions since the year 1800, and a trait database. We also examined how plant community indicators of resources and conditions changed in response to management legacies and environmental change. Community trajectories were clearly influenced by interactions between management legacies from over 200 years ago and environmental change. Importantly, higher rates of nitrogen deposition led to increased species richness and plant height in forests managed less intensively in 1800 (i.e., high forests), and to decreases in forests with a more intensive historical management in 1800 (i.e., coppiced forests). There was evidence that these declines in community variables in formerly coppiced forests were ameliorated by increased rates of temperature change between surveys. Responses were generally apparent regardless of sites' contemporary management classifications, although sometimes the management transition itself, rather than historic or contemporary management types, better explained understorey responses. Main effects of environmental change were rare, although higher rates of precipitation change increased plant height, accompanied by increases in fertility indicator values. Analysis of indicator values suggested the importance of directly characterising resources and conditions to better understand legacy and environmental change effects. Accounting for legacies of past disturbance can reconcile contradictory literature results and appears crucial to anticipating future responses to global environmental change.

Non-random species loss in a forest herbaceous layer following nitrogen addition.

Nitrogen (N) additions have decreased species richness (S) in hardwood forest herbaceous layers, yet the functional mechanisms for these decreases have not been explicitly evaluated. We tested two hypothesized mechanisms, random species loss (RSL) and non-random species loss (NRSL), in the hardwood forest herbaceous layer of a long-term, plot-scale, fertilization experiment in the central Appalachian Mountains, USA. Using a random thinning algorithm, we simulated changes in species densities under RSL and compared the simulated densities to the observed densities among N-fertilized (+N), N-fertilized and limed (+N+L), and reference (REF) plots in regenerating forest stands. We found a lower S in the +N treatment across all survey years and determined that the reduction in S was a function of NRSL. Furthermore, non-random effects were observed in certain species, as they occurred at densities that were either higher or lower than expected due to RSL. Differential advantages were also observed among species between +N and +N+L treatments, suggesting that species responded to either the fertilization or acidification effects of N, though no consistent pattern emerged. Species nitrophily status was not a useful trait for predicting specific species losses, but was a significant factor when averaged across all treatments and sampling years. Our results provide strong evidence that declines in S in the forest herbaceous layer under N fertilization are due largely to NRSL and not simply a function of species rarity.

Forest ecosystems of temperate climatic regions: from ancient use to climate change.

871 I. 871 II. 874 III. 875 IV. 878 V. 882 884 References 884 SUMMARY: Humans have long utilized resources from all forest biomes, but the most indelible anthropogenic signature has been the expanse of human populations in temperate forests. The purpose of this review is to bring into focus the diverse forests of the temperate region of the biosphere, including those of hardwood, conifer and mixed dominance, with a particular emphasis on crucial challenges for the future of these forested areas. Implicit in the term 'temperate' is that the predominant climate of these forest regions has distinct cyclic, seasonal changes involving periods of growth and dormancy. The specific temporal patterns of seasonal change, however, display an impressive variability among temperate forest regions. In addition to the more apparent current anthropogenic disturbances of temperate forests, such as forest management and conversion to agriculture, human alteration of temperate forests is actually an ancient phenomenon, going as far back as 7000 yr before present (bp). As deep-seated as these past legacies are for temperate forests, all current and future perturbations, including timber harvesting, excess nitrogen deposition, altered species' phenologies, and increasing frequency of drought and fire, must be viewed through the lens of climate change.

Combining community resurvey data to advance global change research.

More and more ecologists have started to resurvey communities sampled in earlier decades to determine long-term shifts in community composition and infer the likely drivers of the ecological changes observed. However, to assess the relative importance of, and interactions among, multiple drivers joint analyses of resurvey data from many regions spanning large environmental gradients are needed. In this paper we illustrate how combining resurvey data from multiple regions can increase the likelihood of driver-orthogonality within the design and show that repeatedly surveying across multiple regions provides higher representativeness and comprehensiveness, allowing us to answer more completely a broader range of questions. We provide general guidelines to aid implementation of multi-region resurvey databases. In so doing, we aim to encourage resurvey database development across other community types and biomes to advance global environmental change research.

Microclimate moderates plant responses to macroclimate warming.

Recent global warming is acting across marine, freshwater, and terrestrial ecosystems to favor species adapted to warmer conditions and/or reduce the abundance of cold-adapted organisms (i.e., "thermophilization" of communities). Lack of community responses to increased temperature, however, has also been reported for several taxa and regions, suggesting that "climatic lags" may be frequent. Here we show that microclimatic effects brought about by forest canopy closure can buffer biotic responses to macroclimate warming, thus explaining an apparent climatic lag. Using data from 1,409 vegetation plots in European and North American temperate forests, each surveyed at least twice over an interval of 12-67 y, we document significant thermophilization of ground-layer plant communities. These changes reflect concurrent declines in species adapted to cooler conditions and increases in species adapted to warmer conditions. However, thermophilization, particularly the increase of warm-adapted species, is attenuated in forests whose canopies have become denser, probably reflecting cooler growing-season ground temperatures via increased shading. As standing stocks of trees have increased in many temperate forests in recent decades, local microclimatic effects may commonly be moderating the impacts of macroclimate warming on forest understories. Conversely, increases in harvesting woody biomass--e.g., for bioenergy--may open forest canopies and accelerate thermophilization of temperate forest biodiversity.

Divergent responses of soil buffering capacity to long-term N deposition in three typical tropical forests with different land-use history.

Elevated anthropogenic nitrogen (N) deposition has become an important driver of soil acidification at both regional and global scales. It remains unclear, however, how long-term N deposition affects soil buffering capacity in tropical forest ecosystems and in ecosystems of contrasting land-use history. Here, we expand on a long-term N deposition experiment in three tropical forests that vary in land-use history (primary, secondary, and planted forests) in Southern China, with N addition as NH4NO3 of 0, 50, 100, and 150 kg N ha(-1) yr(-1), respectively. Results showed that all three forests were acid-sensitive ecosystems with poor soil buffering capacity, while the primary forest had higher base saturation and cation exchange capacity than others. However, long-term N addition significantly accelerated soil acidification and decreased soil buffering capacity in the primary forest, but not in the degraded secondary and planted forests. We suggest that ecosystem N status, influenced by different land-use history, is primarily responsible for these divergent responses. N-rich primary forests may be more sensitive to external N inputs than others with low N status, and should be given more attention under global changes in the future, because lack of nutrient cations is irreversible.

A reference-based approach for estimating leaf area and cover in the forest herbaceous layer.

Cover data are used to assess vegetative response to a variety of ecological factors. Estimating cover in the herbaceous layer of forests presents a problem because the communities are structurally complex and rich in species. The currently employed techniques for estimating cover are less than optimal for measuring such rich understories because they are inaccurate, slow, or impracticable. A reference-based approach to estimating cover is presented that compares the area of foliar surfaces to the area of an observer's hand. While this technique has been used to estimate cover in prior studies, its accuracy has not been tested. We tested this hand-area method at the individual plant, population, and community scales in a deciduous forest herbaceous layer, and in a separate farm experiment. The precision, accuracy, observer bias, and species bias of the method were tested by comparing the hand-estimated leaf area index values with actual leaf area index, measured using a leaf area meter. The hand-area method was very precise when regressed against actual leaf area index at the plant, population, and community scales (R(2) of 0.97, 0.93, and 0.87). Among the deciduous sites, the hand-area method overestimated leaf area index consistently by 39.1% at all scales. There was no observer bias detected at any scale, but plant overestimation bias was detected in one species at the population scale. The hand-area method is a rapid and reliable technique for estimating leaf area index or cover in the forest herbaceous layer and should be useful to field ecologists interested in answering questions at the plant, population, or community level.

Nitrogen deposition contributes to soil acidification in tropical ecosystems.

Elevated anthropogenic nitrogen (N) deposition has greatly altered terrestrial ecosystem functioning, threatening ecosystem health via acidification and eutrophication in temperate and boreal forests across the northern hemisphere. However, response of forest soil acidification to N deposition has been less studied in humid tropics compared to other forest types. This study was designed to explore impacts of long-term N deposition on soil acidification processes in tropical forests. We have established a long-term N-deposition experiment in an N-rich lowland tropical forest of Southern China since 2002 with N addition as NH4 NO3 of 0, 50, 100 and 150 kg N ha(-1)  yr(-1) . We measured soil acidification status and element leaching in soil drainage solution after 6-year N addition. Results showed that our study site has been experiencing serious soil acidification and was quite acid-sensitive showing high acidification (pH(H2O) <4.0), negative water-extracted acid neutralizing capacity (ANC) and low base saturation (BS,< 8%) throughout soil profiles. Long-term N addition significantly accelerated soil acidification, leading to depleted base cations and decreased BS, and further lowered ANC. However, N addition did not alter exchangeable Al(3+) , but increased cation exchange capacity (CEC). Nitrogen addition-induced increase in SOC is suggested to contribute to both higher CEC and lower pH. We further found that increased N addition greatly decreased soil solution pH at 20 cm depth, but not at 40 cm. Furthermore, there was no evidence that Al(3+) was leaching out from the deeper soils. These unique responses in tropical climate likely resulted from: exchangeable H(+) dominating changes of soil cation pool, an exhausted base cation pool, N-addition stimulating SOC production, and N saturation. Our results suggest that long-term N addition can contribute measurably to soil acidification, and that shortage of Ca and Mg should receive more attention than soil exchangeable Al in tropical forests with elevated N deposition in the future.

Arbuscular mycorrhizal fungi reduce soil N2O emissions by altering root traits and soil denitrifier community composition.

Arbuscular mycorrhizal fungi (AMF) increase the ability of plants to obtain nitrogen (N) from the soil, and thus can affect emissions of nitrous oxide (N2O), a long-lived potent greenhouse gas. However, the mechanisms underlying the effects of AMF on N2O emissions are still poorly understood, particularly in agroecosystems with different forms of N fertilizer inputs. Utilizing a mesocosm experiment in field, we examined the effects of AMF on N2O emissions via their influence on maize root traits and denitrifying microorganisms under ammonia and nitrate fertilizer input using 15N isotope tracer. Here we show that the presence of AMF alone or both maize roots and AMF increased maize biomass and their 15N uptake, root length, root surface area, and root volume, but led to a reduction in N2O emissions under both N input forms. Random forest model showed that root length and surface area were the most important predictors of N2O emissions. Additionally, the presence of AMF reduced the (nirK + nirS)/nosZ ratio by increasing the relative abundance of nirS-Bradyrhizobium and Rubrivivax with ammonia input, but reducing nosZ-Azospirillum, Cupriavidus and Rhodopseudomonas under both fertilizer input. Further, N2O emissions were significantly and positively correlated with the nosZ-type Azospirillum, Cupriavidus and Rhodopseudomonas, but negatively correlated with the nirS-type Bradyrhizobium and Rubrivivax. These results indicate that AMF reduce N2O emissions by increasing root length to explore N nutrients and altering the community composition of denitrifiers, suggesting that effective management of N fertilizer forms interacting with the rhizosphere microbiome may help mitigate N2O emissions under future N input scenarios.

Estimated losses of plant biodiversity in the United States from historical N deposition (1985-2010).

Although nitrogen (N) deposition is a significant threat to herbaceous plant biodiversity worldwide, it is not a new stressor for many developed regions. Only recently has it become possible to estimate historical impacts nationally for the United States. We used 26 years (1985-2010) of deposition data, with ecosystem-specific functional responses from local field experiments and a national critical loads (CL) database, to generate scenario-based estimates of herbaceous species loss. Here we show that, in scenarios using the low end of the CL range, N deposition exceeded critical loads over 0.38, 6.5, 13.1, 88.6, and 222.1 million ha for the Mediterranean California, North American Desert, Northwestern Forested Mountains, Great Plains, and Eastern Forest ecoregions, respectively, with corresponding species losses ranging from < 1% to 30%. When we ran scenarios assuming ecosystems were less sensitive (using a common CL of 10 kg x ha(-1) x yr(-1), and the high end of the CL range) minimal losses were estimated. The large range in projected impacts among scenarios implies uncertainty as to whether current critical loads provide protection to terrestrial plant biodiversity nationally and urge greater research in refining critical loads for U.S. ecosystems.

Divergent roles of herbivory in eutrophying forests.

Ungulate populations are increasing across Europe with important implications for forest plant communities. Concurrently, atmospheric nitrogen (N) deposition continues to eutrophicate forests, threatening many rare, often more nutrient-efficient, plant species. These pressures may critically interact to shape biodiversity as in grassland and tundra systems, yet any potential interactions in forests remain poorly understood. Here, we combined vegetation resurveys from 52 sites across 13 European countries to test how changes in ungulate herbivory and eutrophication drive long-term changes in forest understorey communities. Increases in herbivory were associated with elevated temporal species turnover, however, identities of winner and loser species depended on N levels. Under low levels of N-deposition, herbivory favored threatened and small-ranged species while reducing the proportion of non-native and nutrient-demanding species. Yet all these trends were reversed under high levels of N-deposition. Herbivores also reduced shrub cover, likely exacerbating N effects by increasing light levels in the understorey. Eutrophication levels may therefore determine whether herbivory acts as a catalyst for the "N time bomb" or as a conservation tool in temperate forests.