Assessing tree ring δ15N of four temperate deciduous species as an indicator of N availability using independent long-term records at the Fernow Experimental Forest, WV.

Nitrogen deposition in the northeastern US changed N availability in the latter part of the twentieth century, with potential legacy effects. However, long-term N cycle measurements are scarce. N isotopes in tree rings have been used as an indicator of N availability through time, but there is little verification of whether species differ in the strength of this signal. Using long-term records at the Fernow Experimental Forest in West Virginia, we examined the relationship between soil conditions, including net nitrification rates, and wood δ15N in 2014, and tested the strength of correlation between tree ring δ15N of four species and stream water NO3- loss from 1971 to 2000. Higher soil NO3- was weakly associated with higher wood δ15N across species, and higher soil net nitrification rates were associated with higher δ15N for Quercus rubra only. The δ15N of Liriodendron tulipifera and Q. rubra, but neither Fagus grandifolia nor Prunus serotina, was correlated with stream water NO3-. L. tulipifera tree ring δ15N had a stronger association with stream water NO3- than Q. rubra. Overall, we found only limited evidence of a relationship between soil N cycling and tree ring δ15N, with a strong correlation between the wood δ15N and NO3- leaching loss through time for one of four species. Tree species differ in their ability to preserve legacies of N cycling in tree ring δ15N, and given the weak relationships between contemporary wood δ15N and soil N cycle measurements, caution is warranted when using wood δ15N to infer changes in the N cycle.

Long-Term Experimental Acidification Drives Watershed Scale Shift in Dissolved Organic Matter Composition and Flux.

Over the last several decades dissolved organic carbon concentrations (DOC) in surface waters have increased throughout much of the northern hemisphere. Several hypotheses have been proposed regarding the drivers of this phenomenon including decreased sulfur (S) deposition working via an acidity- change mechanism. Using fluorescence spectroscopy and data from two long-term (24+ years at completion of this study) whole watershed acidification experiments, that is, the Bear Brook Watershed in Maine (BBWM) and Fernow Experimental Forest in West Virginia (FEF) allowed us to control for factors other than the acidity-change mechanism (e.g., differing vegetation, shifting climate), resulting in the first study we are aware of where the acidity change mechanism could be experimentally isolated at the whole ecosystem and decadal scales as the driver of shifts in DOM dynamics. The multidecadal record of stream chemistry at BBWM demonstrates a significantly lower DOC concentration in the treated compared to the reference watershed. Additionally, at both BBWM and FEF we found significant and sustained differences in stream fluorescence index (FI) between the treated and reference watersheds, with the reference watersheds demonstrating a stronger terrestrial DOM signature. These data, coupled with evidence of pH shifts in upper soil horizons support the hypotheses that declines in S deposition are driving changes in the solubility of soil organic matter and increased flux of terrestrial DOC to water bodies.

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.

Modeled ecohydrological responses to climate change at seven small watersheds in the northeastern United States.

A cross-site analysis was conducted on seven diverse, forested watersheds in the northeastern United States to evaluate hydrological responses (evapotranspiration, soil moisture, seasonal and annual streamflow, and water stress) to projections of future climate. We used output from four atmosphere-ocean general circulation models (AOGCMs; CCSM4, HadGEM2-CC, MIROC5, and MRI-CGCM3) included in Phase 5 of the Coupled Model Intercomparison Project, coupled with two Representative Concentration Pathways (RCP 8.5 and 4.5). The coarse resolution AOGCMs outputs were statistically downscaled using an asynchronous regional regression model to provide finer resolution future climate projections as inputs to the deterministic dynamic ecosystem model PnET-BGC. Simulation results indicated that projected warmer temperatures and longer growing seasons in the northeastern United States are anticipated to increase evapotranspiration across all sites, although invoking CO2 effects on vegetation (growth enhancement and increases in water use efficiency (WUE)) diminish this response. The model showed enhanced evapotranspiration resulted in drier growing season conditions across all sites and all scenarios in the future. Spruce-fir conifer forests have a lower optimum temperature for photosynthesis, making them more susceptible to temperature stress than more tolerant hardwood species, potentially giving hardwoods a competitive advantage in the future. However, some hardwood forests are projected to experience seasonal water stress, despite anticipated increases in precipitation, due to the higher temperatures, earlier loss of snow packs, longer growing seasons, and associated water deficits. Considering future CO2 effects on WUE in the model alleviated water stress across all sites. Modeled streamflow responses were highly variable, with some sites showing significant increases in annual water yield, while others showed decreases. This variability in streamflow responses poses a challenge to water resource management in the northeastern United States. Our analyses suggest that dominant vegetation type and soil type are important attributes in determining future hydrological responses to climate change.

Soluble soil aluminum alters the relative uptake of mineral nitrogen forms by six mature temperate broadleaf tree species: possible implications for watershed nitrate retention.

Increased availability of monomeric aluminum (Al3+) in forest soils is an important adverse effect of acidic deposition that reduces root growth and inhibits nutrient uptake. There is evidence that Al3+ exposure interferes with NO3- uptake. If true for overstory trees, the reduction in stand demand for NO3- could increase NO3- discharge in stream water. These effects may also differ between species that tolerate different levels of soil acidity. To examine these ideas, we measured changes in relative uptake of NO3- and NH4+ by six tree species in situ under increased soil Al3+ using a 15N-labeling technique, and measured soluble soil Al levels in a separate whole-watershed acidification experiment in the Fernow Experimental Forest (WV). When exposed to added Al3+, the proportion of inorganic N acquired as NO3- dropped 14% across species, but we did not detect a reduction in overall N uptake, nor did tree species differ in this response. In the long-term acidification experiment, we found that soluble soil Al was mostly in the free Al3+ form, and the concentration of Al3+ was ~65 µM higher (~250%) in the mineral soil of the acidified watershed vs. an untreated watershed. Thus, increased levels of soil Al3+ under acidic deposition cause a reduction in uptake of NO3- by mature trees. When our 15N uptake results were applied to the watershed acidification experiment, they suggest that increased Al3+ exposure could reduce tree uptake of NO3- by 7.73 kg N ha-1 year-1, and thus increase watershed NO3- discharge.

Land application of hydrofracturing fluids damages a deciduous forest stand in West Virginia.

In June 2008, 303,000 L of hydrofracturing fluid from a natural gas well were applied to a 0.20-ha area of mixed hardwood forest on the Fernow Experimental Forest, West Virginia. During application, severe damage and mortality of ground vegetation was observed, followed about 10 d later by premature leaf drop by the overstory trees. Two years after fluid application, 56% of the trees within the fluid application area were dead. Ehrh. was the tree species with the highest mortality, and L. was the least affected, although all tree species present on the site showed damage symptoms and mortality. Surface soils (0-10 cm) were sampled in July and October 2008, June and October 2009, and May 2010 on the fluid application area and an adjacent reference area to evaluate the effects of the hydrofracturing fluid on soil chemistry and to attempt to identify the main chemical constituents of the hydrofracturing fluid. Surface soil concentrations of sodium and chloride increased 50-fold as a result of the land application of hydrofracturing fluids and declined over time. Soil acidity in the fluid application area declined with time, perhaps from altered organic matter cycling. This case study identifies the need for further research to help understand the nature and the environmental impacts of hydrofracturing fluids to devise optimal, safe disposal strategies.

Pervasive interactions between ungulate browsers and disturbance regimes promote temperate forest herbaceous diversity.

Disruptions to historic disturbance and herbivory regimes have altered plant assemblages in forests worldwide. An emerging consensus suggests that these disruptions often result in impoverished forest biotas. This is particularly true for eastern U.S. deciduous forests where large gaps and understory fires were once relatively common and browsers were far less abundant. Although much research has focused on how disturbance and browsers affect tree diversity, far less attention has been devoted to forest understories where the vast majority (>75%) of the vascular species reside. Here we test the hypothesis that the reintroduction of disturbances resembling historic disturbance regimes and moderate levels of ungulate browsing enhance plant diversity. We explore whether once-common disturbances and their interaction with the top-down influence of browsers can create conditions favorable for the maintenance of a rich herbaceous layer in a region recognized as a temperate biodiversity hotspot in West Virginia, U.S.A. We tested this hypothesis via a factorial experiment whereby we manipulated canopy gaps (presence/absence) of a size typically found in old-growth stands, low-intensity understory fire (burned/unburned), and deer browsing (fenced/unfenced). We tracked the abundance and diversity of more than 140 herb species for six years. Interactions among our treatments were pervasive. The combination of canopy gaps and understory fire increased herbaceous layer richness, cover, and diversity well beyond either disturbance alone. Furthermore, we documented evidence that deer at moderate levels of abundance promote herbaceous richness and abundance by preferentially browsing fast-growing pioneer species that thrive following co-occurring disturbances (i.e., fire and gaps). This finding sharply contrasts with the negative impact browsers have when their populations reach levels well beyond those that occurred for centuries. Although speculative, our results suggest that interactions among fire, canopy gaps, and browsing provided a variable set of habitats and conditions across the landscape that was potentially capable of maintaining much of the plant diversity found in temperate forests.

Comparing decadal responses of whole-watershed manipulations at the Bear Brook and Fernow experiments.

The Bear Brook Watershed in Maine (BBWM), USA, and the Fernow Experimental Forest in West Virginia, USA, represent unique, long-term, paired, whole watershed, experimental manipulations focusing on the effects of nitrogen (N) and sulfur (S) deposition on temperate forests. Both watersheds began whole-ecosystem additions of N and S as (NH(4))(2)SO(4) in the fall of 1989, and both are entering their third decade of chronic enrichment of the treated watersheds, while the reference watersheds offer unique opportunities to evaluate forest watershed responses to recovery. Differences between BBWM and Fernow in the history of atmospheric deposition, soil properties, and forest composition all contribute to different response trajectories in stream chemical exports over time. The four watersheds represent a spectrum of N enrichment and retention, ranging from ≈98% N retention in the reference watershed in Maine, to ≈20% N retention in the treated watershed in West Virginia. Despite these differences, there is evidence that mechanisms of response in base cation leaching and other processes are similar among all four watersheds. In both cases, the history to date of two decades of research and monitoring has provided new insights into ecosystem response not evident in more traditional short-term research.

Response of soil fertility to 25 years of experimental acidification in a temperate hardwood forest.

The effects of enhanced acid deposition from the atmosphere, and associated elevated inputs of N, are widely evident, especially for forests where excess N has led to a variety of deleterious effects. These include declines in biodiversity, a response that will likely require considerable time for recovery. The purpose of this study was to determine responses of plant nutrient availability in surface mineral soil to 25 yr of experimental acidification and N addition in a central Appalachian hardwood forest ecosystem. We hypothesized that chronic additions of (NH4 )2 SO4 will increase mineral N, decrease soil pH, P, and base cations, increase micronutrients (Mn2+ and Fe2+ ), and increase levels of Al3+ . Results supported these predictions, although Mn2+ did not vary significantly. Earlier work on these plots found no response of any of the extractable nutrients to 3 yr of treatment, yet after 25 yr, our results suggest that impacts are apparent in the top 5 cm of the A horizon. We surmise that impacts in these soils may have lagged behind the onset of acidification treatments or that several years of treatment were required to overcome preexisting differences in soil ions. Generally, current findings confirm that (NH4 )2 SO4 treatments have lowered the pH, enhanced levels of exchangeable Al3+ , and increased stream-water exports of NO3 - and base cations-a process that further acidifies soil. The combination of these changes in surface soils, with their high proportion of fine roots, may contribute to the reduced growth and competitiveness of some hardwood species at the acidified site.

Ecological issues related to N deposition to natural ecosystems: research needs.

There has and continues to be concern about the effects of elevated nitrogen (N) deposition on natural ecosystems. In this paper, research on natural ecosystems, including wetlands, heathlands, grasslands, steppe, naturally regenerated forests and deserts, is evaluated to determine what is known about nitrogen cycling in these ecosystems, the effects of elevated nitrogen on them and to identify research gaps. Aquatic ecosystems are not included in this review, except as they are part of the larger ecosystem. Research needs fall into several categories: (1) improved understanding and quantification of the N cycle, particularly relatively unstudied processes such as dry deposition, N fixation and decomposition/mineralization; (2) carbon cycling as affected by increased N deposition; (3) effects on arid ecosystems and other "neglected" ecosystems; (4) effects on complex ecosystems and interactions with other pollutants; (5) indicators and assessment tools for natural ecosystems.

Wetfall Deposition and Precipitation Chemistry for a Central Appalachian Forest.

Although extensive research on acidic deposition has been directed toward spruce-fir forests, less research has been done on the impacts of air pollution on eastern montane hardwood forests. The purpose of this study was to describe precipitation chemistry for several Appalachian hardwood forest sites at or near the Fernow Experimental Forest (FEF) to assess the potential for problems associated with acidic deposition. Emphasis was placed on seasonal patterns of ionic concentrations (H+, Ca++, NH4+; NO3-, and SO4=) and spatial variability of ionic concentrations and deposition among sites. Seasonal patterns of most ions showed highest concentrations during the summer months and deposition of H+ was especially pronounced during this time. Deposition of all ions was generally greater (related to greater precipitation) at three montane forested sample sites compared to a nonforested riverbottom site. Precipitation chemistry at FEF was similar to other sites throughout the eastern United States and contrasted sharply with mid-western and western sites. Eastern sites, including means for FEF sites, were uniformly 3-4 times higher in H+ and SO4= concentration than the mid-western and western sites. Precipitation at FEF was chronically acidic, more so during the growing season, and highest at higher elevations where environmental stresses can be most severe. Furthermore, there were occasional large discrepancies between the low-elevation site and the higher-elevation forested sites for precipitation chemistry and acidic deposition. These results suggest that synoptic-scale (network) data may greatly underestimate the pollutant conditions to which highelevation forest trees are exposed, since network data rarely take elevation into account and typically are based on annual ionic concentration and deposition means that may be considerably lower than those of the growing season.