Assessing nutrient critical load exceedances requires complete and accurate atmospheric deposition budgets for reactive nitrogen (N-r). The exceedance is the total amount of N-r deposited to the ecosystem in excess of the critical load, which is the amount of Nr input below which harmful effects do not occur. Total deposition includes all forms of N-r (i.e., organic and inorganic) deposited to the ecosystem by wet and dry pathways. Here we present results from the Southern Appalachian Nitrogen Deposition Study (SANDS), in which a combination of measurements and field-scale modeling was used to develop a complete annual Nr deposition budget for a deciduous forest at the Coweeta Hydrologic Laboratory. Wet deposition of ammonium, nitrate, nitrite, and bulk organic N were measured directly. The dry deposited N(r )fraction was estimated using a bidirectional resistance-based model driven with speciated measurements of N-r air concentrations (e.g., ammonia, ammonium aerosol, nitric acid, nitrate aerosol, bulk organic N in aerosol, total alkyl nitrates, and total peroxy nitrates), micrometeorology, canopy structure, and biogeochemistry. Total annual deposition was similar to 6.7 kg N ha(-1) yr(-1), which is on the upper end of Nr critical load estimates recently developed for similar ecosystems in the nearby Great Smoky Mountains National Park. Of the total (wet + dry) budget, 51.1 % was contributed by reduced forms of N-r (NHx = ammonia + ammonium), with oxidized and organic forms contributing similar to 41.3 % and 7.6 %, respectively. Our results indicate that reductions in NHx deposition would be needed to achieve the lowest estimates (similar to 3.0 kg N ha(-1) yr(-1)) of Nr critical loads in southern Appalachian forests.

Autotrophic and heterotrophic organisms require resources in stoichiometrically balanced ratios of carbon (C) to nutrients, the demand for which links organismal and ecosystem-level biogeochemical cycles. In soils, the relative availability of C and nitrogen (N) also defines the strength of competition for ammonium between autotrophic nitrifiers and heterotrophic decomposers, which may influence the coupled dynamics between N mineralization and nitrification. Here, we use data from the publicly available US National Science Foundation funded Long Term Ecological Research (LTER) network to evaluate the influence of soil C concentration on the relationship between net nitrification and net N mineralization. We found that soil C availability constrains the fraction of mineralized N that is ultimately nitrified across the continental gradient, contributing to reduced rates of nitrification in soils with high C concentrations. Nitrate, which is produced by nitrification, is a highly mobile ion that easily leaches to aquatic ecosystems or denitrifies into the greenhouse gas nitrous oxide (N 2 O). Understanding the connection between soil C concentration and soil N transformations is thus important for managing potential ecosystem N losses, understanding the biogeochemical constraints of these losses, and accurately representing coupled C-N dynamics in ecosystem models.

Most studies of land use effects on solute concentrations in streams have focused on smaller streams with watersheds dominated by a single land‐use type. Using land cover as a proxy for land use, the objective of this study was to determine whether the hydrologically‐driven response of solutes to land use in small streams could be scaled up to predict concentrations in larger receiving streams and rivers in the rural area of the Little Tennessee River basin. We measured concentrations of typically limiting nutrients (nitrogen, phosphorus), abundant anions (chloride, sulfate), and base cations in 17 small streams and four larger river sites. In the small streams, total solute concentration was strongly related to land cover ‐‐ highest in streams with developed watersheds, lowest in streams with forested watersheds, and streams with agricultural watersheds were in between. In general, the best predictor of solute concentrations in the small streams was forest land cover. We then predicted solute concentrations for the river sites based on the solute‐‐land cover relationships of the small streams using multiple linear regressions. Results were mixed ‐‐ some of the predicted river concentrations were close to measured values, others were greater or less than measured concentrations. In general, river concentrations did not scale with land cover‐solute relationships found in small tributaries. Measured values of nitrogen solutes in the river sites were greater than predicted, perhaps due to the presence of waste water treatment plants. We attributed other differences between measured and predicted river concentrations to the heterogeneous geochemistry of this mountainous region. The combined complexity of hydrology, geochemistry, and human land‐use of this mountainous region make it difficult to scale up from small streams to larger river basins.

The southern Appalachian Mountains in the United States are heavily forested with diverse forest types and vegetation structures and is generally considered a biodiversity hotspot. Fungal sporocarp investigations in this region are not new, but multi-year interseasonal investigations into sporocarp community patterns are rare. Using a 4-year (2014–2017) repeated monthly macrofungal sampling dataset (May–October) across an elevational gradient (associated with vegetation structure), we queried community, seasonal, and guild community ecological patterns. In doing so, we (1) demonstrate a temporal disconnect between saprobic and ectomycorrhizal taxa, (2) describe increased community variability across the growing season, and (3) explore individual taxa occurrences across seasons and examine co-occurrence patterns between taxa and guilds. Further, we explore congruence between sporocarp and metabarcoding fungal datasets and advocate the utility of both sampling schemes.

The water-quality effects of low-density rural land-use activities are understudied but important because of large rural land coverage. We review and synthesize spatially extensive studies of oligotrophic mountain streams in the rural Southern Appalachian Mountains, concluding that rural land-use activities significantly degrade water quality through altered and mostly enhanced landscape–stream connections, despite high forest retention. Some connections (insolation, organic inputs, root–channel interactions, stream–field connectivity, individual landowner discharges) are controlled by near-stream land-use activities, whereas others (reduced nitrogen uptake and cycling, enhanced biological nitrogen fixation, nutrient subsidy, runoff from compacted soils, road runoff delivery) are controlled by basin-wide land use. These connections merge to alter basal resources and shift fish, salamander, and invertebrate assemblages toward species tolerant of higher turbidity and summer temperatures and those more competitive in mesotrophic systems. Rural water quality problems could be mitigated substantially with well-known best management practices, raising socioecological governance questions about best management practice adoption.

Rhododendron maximum is an evergreen shrub native to the Appalachian Mountains of North America that has expanded in recent decades due to past disturbances and land management. The purpose of this study was to explore how bees and plants were affected by the experimental removal of R . maximum followed by a prescribed fire in one watershed compared to a neighboring reference watershed. Bees and plants were sampled for three years in both watersheds. Comparisons were based on the rarefaction and extrapolation sampling curves of Hill numbers as well as multivariate methods to assess effects on community composition. Bee richness, Shannon's diversity, and Simpson's diversity did not differ between watersheds in the year after removal but were all significantly higher in the removal watershed in year two, following the prescribed fire. Bee Shannon's diversity and Simpson's diversity, but not richness, remained significantly higher in the removal watershed in the third year. Similar but weaker patterns were observed for plants. Comparisons of community composition found significant differences for bees in the second and third year and significant differences for plants in all three years. For both groups, significant indicator taxa were mostly associated with the removal watershed. Because bees appeared to respond more strongly to the prescribed fire than to the removal of R . maximum and these benefits weakened considerably one year after the fire, clearing R . maximum does not appear to dramatically improve pollinator habitat in the southern Appalachians. This conclusion is underscored by the fact that about one quarter of the bee species in our study area were observed visiting R . maximum flowers. The creation of open areas with wildflowers may be a better way to benefit bees in this region judging from the high diversity of bees captured in the small roadside clearings in this study.

Soil organic matter composition controls many microbial processes in the soil matrix. How these processes interact to drive carbon cycling through greenhouse gas fluxes or carbon stabilization through biochemical transformations continues to evolve. From laboratory incubations, it is clear that low molecular weight compounds (LMWCs; e.g. dextrose, mannitol, and trehalose) can have a profound effect on microbially mediated fluxes. However, relationships between LMWCs and soil efflux have been largely unproven in field studies due to methodological constraints. In the current study we developed a novel use of Near Infrared Reflectance Spectroscopy (NIRS) to quantify LMWCs (explaining 38-51% of the variance). Paradoxically, we found dextrose concentrations to be negatively correlated with carbon dioxide fluxes in residential sites while mannitol was positively correlated with carbon dioxide fluxes in agricultural sites. Methane fluxes were strongly correlated with trehalose indicating a potential fungal interaction with bacterial methanogens. We found no clear link between LMWCs on nitrous oxide emissions suggesting inorganic nitrogen is a stronger limiting factor. The results from this study showed how a NIRS-based approach can improve mechanistic understanding of the drivers of soil greenhouse gas fluxes.

Long-term (LT) stream chemistry studies are used to examine changes in and responses to the environment. Much of the data collected over long periods of time goes through changes in instrumentation, methods, and personnel potentially resulting in changing values. A data user must understand these measures of data quality through quality control (QC) results to know with certainty if trends are real or attributable to other factors. We used the Web of Science database search engine to search for LT stream chemistry studies. For each study, we then determined: record or study length; if QC was reported; and if QC was used. We found that 33% of papers reported QC in the method, and 12% presented the QC in the results. Next, we conducted a case study on 46 years of stream chemistry data to evaluate the data with and without the application of QC protocols from two watersheds (WS) at Coweeta Hydrologic Laboratory; WS 7; clear-cut in 1967-77 and adjacent WS 2 which serves as a reference. We focused on nitrogen and sulfur due to their importance in understanding the forest ecosystem response to disturbance (NO3) and acid deposition (SO4). We determined average annual dissolved inorganic nitrogen (DIN) export (NH4 + NO3 = DIN) using three methods for censoring values below the method detection limit (mdl): (1) the found value, (2) the value of zero, and (3) one-half the mdl value. We found that DIN export for WS 2/WS 7 was (1) 66.9/831.4 (g ha(-1) yr(-1)), (2) 45.4/808.0 (g ha(-1) yr(-1)), and (3) 72.1/823.2 (g ha(-1) yr(-1)) using the three censoring methods, and that the export estimate was significantly different for WS 2 but not for WS 7 (P = 0.001). We found that on average stream NH4 concentrations were below the mdl 58% of the time until an instrument change in 1994 resulted in improved mdls resulting in fewer data points below detection. We found increased bias for stream SO4 concentration following an instrumentation change from segmented flow analysis to ion chromatography. As a result, stream SO4 concentration data that were bias-corrected declined more rapidly in WS 2 compared with non-bias-corrected data, but not in WS 7. We conclude that including QC results with LT data is essential to verify data validity and give the data user a full understanding of the results.

Subsurface flow dominates water movement from hillslopes to streams in most forested headwater catchments. Hewlett and Hibbert (1963, https://doi.org/10.1029/JZ068i004p01081 ) constructed an idealized hillslope model (0.91 × 0.91 × 15.0 m; 21.8°) using reconstituted C horizon soil to investigate importance of interflow, a type of subsurface flow. They saturated the model, covered it to prevent evaporation, and allowed free drainage for 145 days. The resulting recession drainage curve suggested two phases: fast drainage of saturated soil in the first 1.5 days and then slow drainage of unsaturated soil. Hydrologists interpreted the latter as evidence interflow could sustain baseflow, even during extended drought. Since that experiment, typical forest vegetation grew in the model, providing root and litter inputs for 55 years. We removed all aboveground live biomass and repeated the experiment physically and numerically (HYDRUS‐2D), hypothesizing that pedogenesis would change the drainage curve and further elucidate the role of unsaturated flow from hillslopes. Contrary to this hypothesis, drainage curves in our twice‐repeated physical experiments and numerical simulation were unchanged for the first ~10 days, indicating pedogenesis and biological processes had not largely altered bulk hydraulic conductivities or soil moisture release characteristics. However, drainage unexpectedly ceased after about 2 weeks (14.3 ± 2.5 days), an order of magnitude sooner than in the original experiment, due to an apparent leak in the hillslope analogous to commonly observed bedrock fractures in natural systems. Thus, our results are a more natural recession behavior that highlight how incorporation of alternative hydrologic outputs can reduce drainage duration and volume from soils to baseflow.

In the Sand County Almanac (Leopold 1949), Aldo Leopold writes of the odyssey of element X and thus of the circulation of all nutrient elements as they cycle through the Earth’s forests, rangelands, lakes, and oceans. In mid-century, G.E. Hutchinson, while always one to praise aesthetic values, commented disparagingly about the quantitative science of element cycling, specifically that ecosystem carbon (C) data were “wretchedly inadequate” (Hutchinson 1954). Hutchinson’s comment was not only a complaint but also a challenge to all ecosystem scientists who followed to quantify the Earth’s biogeochemical cycles, for he understood that the resilience and functioning of ecosystems was entirely dependent on how plants, animals, and decomposers used and reused the chemical elements that we call nutrients.

Wildfires are landscape scale disturbances that can significantly affect hydrologic processes such as runoff generation and sediment and nutrient transport to streams. In Fall 2016, multiple large drought‐related wildfires burned forests across the southern Appalachian Mountains. Immediately after the fires, we identified and instrumented eight 28.4–344 ha watersheds (four burned and four unburned) to measure vegetation, soil, water quantity, and water quality responses over the following two years. Within burned watersheds, plots varied in burn severity with up to 100% tree mortality and soil O‐horizon loss. Watershed scale high burn severity extent ranged from 5% to 65% of total watershed area. Water quantity and quality responses among burned watersheds were closely related to the high burn severity extent. Total water yield (Q) was up to 39% greater in burned watersheds than unburned reference watersheds. Total suspended solids (TSS) concentration during storm events were up to 168 times greater in samples collected from the most severely burned watershed than from a corresponding unburned reference watershed, suggesting that there was elevated risk of localized erosion and sedimentation of streams. NO3‐N concentration, export, and concentration dependence on streamflow were greater in burned watersheds and increased with increasing high burn severity extent. Mean NO3‐N concentration in the most severely burned watershed increased from 0.087 mg L−1 in the first year to 0.363 mg L−1 (+317%) in the second year. These results suggest that the 2016 wildfires degraded forest condition, increased Q, and had negative effects on water quality particularly during storm events.

In mountainous headwater catchments, downslope flow of subsurface water could buffer downslope forest communities from soil moisture stress during drought. Here we investigated changes in landscape-scale vegetation patterns at five forested headwater catchments in the Coweeta Hydrologic Laboratory in the southern Appalachians. We used a ca. 30-year Landsat Thematic Mapper (TM) image record of normalized difference vegetation index (NDVI), spanning a period of recorded warming since the mid-1970. We then, related spatial and temporal canopy patterns to seasonal water balance, streamflow recession behavior, and low flow dynamics from the long-term hydrologic records. All hydrologic metrics indicated increasing evapotranspiration, decreasing streamflow given precipitation, and potentially decreasing downslope subsidy at the watershed scale over time, especially during low-flow periods. Contrary to expectations, leaf area index (LAI) and basal area increased more upslope compared to downslope over time, coincident with warming. Trends in the ratio of NDVI in upslope and downslope topographic positions were also supported by long-term tree basal area increment, litterfall, and sap flux data in one of the reference watersheds. Mesophytic trees downslope appeared to respond more to frequent droughts and experience lower growth than xerophytic trees upslope, closely mediated by the isohydric/anisohydric continuum along hydrologic flow paths. Considering ongoing forest “mesophication” under a history of fire suppression across the eastern United States deciduous forests, this study suggests that mesophytic trees downslope may be more vulnerable than xerophytic trees upslope under ongoing climate change due to an apparent dependence on upslope water subsidy.

Human history is intricately linked to the soil. As human populations increased agricultural land use intensified. Wood availability was also important and resulted in the management of forested land for fuel, fiber and food. Soil mapping was an essential tool in planning future expansion of agriculture and resulted in an increased understanding in the factors that regulate soil formation. This understanding eventually lead to the formation of forest soil science as a separate area of research. Early forest soil research focused on silviculture, harvesting, soil chemistry and acid rain. As the science progressed areas of research broadened to include specialized forest harvest methods and eventually to ecosystem-related research designed to understand how soils function as an integral part of the forest.

Understanding the dominant soil nitrogen (N) cycling processes in southern Appalachian forests is crucial for predicting ecosystem responses to changing N deposition and climate. The role of anaerobic nitrogen cycling processes in well-aerated soils has long been questioned, and recent N cycling research suggests it needs to be re-evaluated. We assessed gross and potential rates of soil N cycling processes, including mineralization, nitrification, denitrification, nitrifier denitrification, and dissimilatory nitrate reduction to ammonium (DNRA) in sites representing a vegetation and elevation gradient in the U.S. Department of Agriculture (USDA) Forest Service Experimental Forest, Coweeta Hydrologic Laboratory in southwestern North Carolina, USA. N cycling processes varied among sites, with gross mineralization and nitrification being greatest in high-elevation northern hardwood forests. Gaseous N losses via nitrifier denitrification were common in all ecosystems but were greatest in northern hardwood. Ecosystem N retention via DNRA (nitrification-produced NO3 reduced to NH4) ranged from 2% to 20% of the total nitrification and was highest in the mixed-oak forest. Our results suggest the potential for gaseous N losses through anaerobic processes (nitrifier denitrification) are prevalent in well-aerated forest soils and may play a key role in ecosystem N cycling.

Long-term research studies are critical to understanding soil productivity and the sustainability of forest and woodland ecosystems around the world. They inform management decisions about best harvest techniques, soil property impacts and recovery, anthropogenic stressors (e.g., forest management, acid rain, climate change), and the influence if governmental policies, guidelines, and regulations. Forest ecosystems represent a major source of drinking water in much of the world making the interaction of atmospheric chemistry, vegetation, land use, soil and water one of global importance. This intimate connection among interacting variables also make forests a logical system to study ecosystem processes. In this chapter we discuss the challenges and benefits of establishing and maintaining long-term studies, the utility of these studies for informing decisions about how to manage forest soil to sustain the delivery of ecosystem services and lessons learned for forest research.

Catchments with minimal disturbance usually have low dissolved inorganic nitrogen (DIN) export, but disturbances and anthropogenic inputs result in elevated DIN concentration and export and eutrophication of downstream ecosystems. We studied streams in the southern Appalachian Mountains, USA, an area dominated by hardwood deciduous forest but with areas of valley agriculture and increasing residential development. We collected weekly grab samples and storm samples from nine small catchments and three river sites. Most discharge occurred at baseflow, with baseflow indices ranging from 69% to 95%. We identified three seasonal patterns of baseflow DIN concentration. Streams in mostly forested catchments had low DIN with bimodal peaks, and summer peaks were greater than winter peaks. Streams with more agriculture and development also had bimodal peaks; however, winter peaks were the highest. In streams draining catchments with more residential development, DIN concentration had a single peak, greatest in winter and lowest in summer. Three methods for estimating DIN export produced consistent results. Annual DIN export ranged from less than 200 g ha−1 year−1 for the less disturbed catchments to over 2,000 g ha−1 year−1 in the catchments with the least forest area. Land cover was a strong predictor of DIN concentration but less significant for predicting DIN export. The two forested reference catchments appeared supply limited, the most residential catchment appeared transport limited, and export for the other catchments was significantly related to discharge. In all streams, baseflow DIN export exceeded stormflow export. Morphological and climatological variation among watersheds created complexities unexplainable by land cover. Nevertheless, regression models developed using land cover data from the small catchments reasonably predicted concentration and export for receiving rivers. Our results illustrate the complexity of mechanisms involved in DIN export in a region with a mosaic of climate, geology, topography, soils, vegetation, and past and present land use.

In Appalachian ecosystems, forest disturbance has long-term effects on microbially driven biogeochemical processes such as nitrogen (N) cycling. However, little is known regarding long-term responses of forest soil microbial communities to disturbance in the region. We used 16S and ITS sequencing to characterize soil bacterial (16S) and fungal (ITS) communities across forested watersheds with a range of past disturbance regimes and adjacent reference forests at the Coweeta Hydrologic Laboratory in the Appalachian mountains of North Carolina. Bacterial communities in previously disturbed forests exhibited consistent responses, including increased alpha diversity and increased abundance of copiotrophic (e.g., Proteobacteria) and N-cycling (e.g., Nitrospirae) bacterial phyla. Fungal community composition also showed disturbance effects, particularly in mycorrhizal taxa. However, disturbance did not affect fungal alpha diversity, and disturbance effects were not consistent at the fungal class level. Co-occurrence networks constructed for bacteria and fungi showed that disturbed communities were characterized by more connected and tightly clustered network topologies, indicating that disturbance alters not only community composition but also potential ecological interactions among taxa. Although bacteria and fungi displayed different long-term responses to forest disturbance, our results demonstrate clear responses of important bacterial and fungal functional groups (e.g., nitrifying bacteria and mycorrhizal fungi), and suggest that both microbial groups play key roles in the long-term alterations to biogeochemical processes observed following forest disturbance in the region.

The interactions of terrestrial C pools and fluxes with spatial and temporal variation in climate are not well understood. We conducted this study in the southern Appalachian Mountains where complex topography provides variability in temperature, precipitation, and forest communities. In 1990, we established five large plots across an elevation gradient allowing us to study the regulation of C and N pools and cycling by temperature and water, in reference watersheds in Coweeta Hydrologic Laboratory, a USDA Forest Service Experimental Forest, in western NC, USA. Communities included mixed-oak pine, mixed-oak, cove hardwood, and northern hardwood. We examined 20-year changes in overstory productivity and biomass, leaf litterfall C and N fluxes, and total C and N pools in organic and surface mineral soil horizons, and coarse wood, and relationships with growing season soil temperature and precipitation. Productivity increased over time and with precipitation. Litterfall C and N flux increased over time and with increasing temperature and precipitation, respectively. Organic horizon C and N did not change over time and were not correlated to litterfall inputs. Mineral soil C and N did not change over time, and the negative effect of temperature on soil pools was evident across the gradient. Our data show that increasing temperature and variability in precipitation will result in altered aboveground productivity. Variation in surface soil C and N is related to topographic variation in temperature which is confounded with vegetation community. Data suggest that climatic changes will result in altered aboveground and soil C and N sequestration and fluxes.