The changing nitrogen landscape of United States streams: Declining deposition and increasing organic nitrogen.

Air quality regulations have led to decreased nitrogen (N) and sulfur deposition across the conterminous United States (CONUS) during the last several decades, particularly in the eastern parts. But it is unclear if declining deposition has altered stream N at large scales. We compared watershed N inputs with N chemistry from over 2,000 CONUS streams where deposition was the largest N input to the watershed. Weighted change analysis showed that deposition declined across most watersheds, especially in the Eastern CONUS. Nationally, declining N deposition was not associated with significant large-scale declines in stream nitrate concentration. Instead, significant increases in stream dissolved organic carbon (DOC) and total organic N (TON) were widespread across regions. Possible mechanisms behind these increases include declines in acidity and/or ionic strength drivers, changes in carbon availability, and/or climate variables. Our results also reveal a declining trend of DOC/TON ratio over the entire study period, primarily influenced by the trend in the Eastern region, suggesting the rate of increase in stream TON exceeded the rate of increase in DOC concentration during this period. Our results illustrate the complexity of nutrient cycling that links long-term atmospheric deposition to water quality. More research is needed to understand how increased dissolved organic N could affect aquatic ecosystems and downstream riverine nutrient export.

Disentangling natural and anthropogenic effects on benthic macroinvertebrate assemblages in western US streams.

Stream macroinvertebrate assemblages are shaped by natural and human-related factors that operate through complex hierarchical pathways. Quantifying these relationships can provide additional insights into stream ecological assessment. We applied a structural equation modeling framework to evaluate hypothesized pathways by which watershed, riparian, and in-stream factors affect benthic macroinvertebrate condition in the Western Mountains (WMT) and Xeric (XER) ecoregions in the United States. We developed a conceptual model grounded in theory, empirical evidence, and expert opinion to evaluate the following hypotheses: (1) macroinvertebrate assemblages are primarily driven by proximal, in-stream factors (e.g., water quality and physical habitat); (2) anthropogenic land uses affect macroinvertebrates indirectly by altering in-stream characteristics; and (3) riparian vegetation cover attenuates land use effects. We tested our model separately on three measures of benthic macroinvertebrate assemblage condition: ratio of observed-to-expected taxonomic richness (O/E); a multimetric index (MMI); and richness of Ephemeroptera, Plecoptera, and Trichoptera taxa (EPT). In the WMT, site-level riparian cover, in-stream physical habitat (relative bed stability), and water chemistry (total nitrogen) were the top three predictors of macroinvertebrate assemblages, each having over two times the magnitude of effect on macroinvertebrates compared with watershed-level predictors. In the arid XER, annual precipitation and stream flow characteristics were top predictors of macroinvertebrate assemblages and had similar magnitudes of effect as in-stream water chemistry. Path analyses revealed that land use activities in the watershed and at the stream site degraded macroinvertebrate assemblages indirectly by altering relative bed stability, water quality, and riparian cover/complexity. Increased riparian cover was associated with greater macroinvertebrate condition by reducing land use impacts on stream flow, streambed substrate, and water quality, but the pathways differed among ecoregions. In the WMT, site-level riparian cover affected macroinvertebrate assemblages partly through indirect pathways associated with greater streambed stability and reduced total nitrogen concentrations. In contrast, in the XER, watershed-level riparian cover affected macroinvertebrate assemblages through greater specific stream power. Identifying the relative effects of and pathways by which natural and anthropogenic factors affect macroinvertebrates can serve as a framework for prioritizing management and conservation efforts.

Natural and anthropogenic controls on lake water-level decline and evaporation-to-inflow ratio in the conterminous United States.

Lake water levels are integral to lake function, but hydrologic changes from land and water management may alter lake fluctuations beyond natural ranges. We constructed a conceptual model of multifaceted drivers of lake water-levels and evaporation-to-inflow ratio (Evap:Inflow). Using a structural equation modeling framework, we tested our model on 1) a national subset of lakes in the conterminous United States with minimal water management to describe natural drivers of lake hydrology and 2) five ecoregional subsets of lakes to explore regional variation in water management effects. Our model fit the national and ecoregional datasets and explained up to 47% of variation in Evap:Inflow, 38% of vertical water-level decline, and 79% of horizontal water-level decline (littoral exposure). For lakes with minimal water management, Evap:Inflow was related to lake depth (ß = -0.31) and surface inflow (ß = -0.44); vertical decline was related to annual climate (e.g., precipitation ß = -0.18) and water management (ß = -0.21); and horizontal decline was largely related to vertical decline (ß = 0.73) and lake morphometry (e.g., depth ß = -0.18). Anthropogenic effects varied by ecoregion and likely reflect differences in regional water management and climate. In the West, water management indicators were related to greater vertical decline (ß = 0.38), whereas in the Midwest, these indicators were related to more stable and full lake levels (ß = -0.22) even during drought conditions. National analyses show how human water use interacts with regional climate resulting in contrasting impacts to lake hydrologic variation in the US.

Deep soil nitrogen storage slows nitrate leaching through the vadose zone.

Nitrogen (N) fertilizer applications are important for agricultural yield, yet not all the applied N is taken up by crops, leading to surplus N storage in soil or leaching to groundwater and surface water. Leaching loss of fertilizer N represents a cost for farmers and has consequences for human health and the environment, especially in the southern Willamette Valley, Oregon, USA, where groundwater nitrate contamination is prevalent. While improved nutrient management and conservation practices have been implemented to minimize leaching, nitrate levels in groundwater continue to increase in many long-term monitoring wells. To elucidate controls on leaching rates and N dynamics in agricultural soils across soil depths, and in response to seasonal and annual variation in management (e.g., fertilizer input amount and summer irrigation), we intensively monitored the transport of water and nitrate every two weeks for four years through the vadose zone at three depths (0.8, 1.5, and 3.0 m) in a sweet corn (maize) field. Though nitrate leaching was highly variable among lysimeters at the same depth and across years, a strong pattern emerged: annual nitrate leaching significantly decreased with depth across the study, averaging ~104 kg N ha-1 yr-1 near the surface (0.8 m) versus ~56 kg N ha-1 yr-1 in the deep soil (3.0 m), a 54% reduction in leaching between the soil layers. Even though crops were irrigated in summer, most leaching (~72% below 3.0 m) occurred during the wet fall and winter. Based on steady state assumptions, a net equivalent of ~29% of surface N inputs leached below 3.0 m into the deeper soil and groundwater, while ~44% was removed in crop harvest, indicating considerable N retention in the soil (~27% of inputs or approximately 58 kg N ha-1 yr-1). The accumulation and long-term dynamics of deep soil N is a legacy of agricultural management that should be further studied to better manage and reduce nitrate loss to groundwater.

δ15N of Chironomidae: An index of nitrogen sources and processing within watersheds for national aquatic monitoring programs.

Nitrogen (N) removal along flowpaths to aquatic ecosystems is an important regulating ecosystem service that can help reduce N pollution in the nation's waterways, but can be challenging to measure at large spatial scales. Measurements that integrate N processing within watersheds would be particularly useful for assessing the magnitude of this vital service. Because most N removal processes cause isotopic fractionation, δ15N from basal food-chain organisms in aquatic ecosystems can provide information on both N sources and the degree of watershed N processing. As part of EPA's National Aquatic Resource Surveys (NARS), we measured δ15N of Chironomidae collected from over 2000 lakes, rivers and streams across the continental USA. Using information on N inputs to watersheds and summer total N concentrations ([TN]) in the water column, we assessed where elevated chironomid δ15N would indicate N removal rather than possible enriched sources of N. Chironomid δ15N values ranged from -4 to +20‰, and were higher in rivers and streams than in lakes, indicating that N in rivers and streams underwent more processing and cycling that preferentially removes 14N than N in lakes. Chironomid δ15N increased with watershed size, N inputs, and water chemical components, and decreased as precipitation increased. In rivers and streams with high watershed N inputs, we found lower [TN] in streams with higher chironomid δ15N values, suggesting high rates of gaseous N loss such as denitrification. At low watershed N inputs, the pattern reversed; streams with elevated chironomid δ15N had higher [TN] than streams with lower chironomid δ15N, possibly indicating unknown sources elevated in δ15N such as legacy N, or waste from animals or humans. Chironomid δ15N values can be a valuable tool to assess integrated watershed-level N sources, input rates, and processing for water quality monitoring and assessment at large scales.

Physiological responses of Douglas-fir to climate and forest disturbances as detected by cellulosic carbon and oxygen isotope ratios.

Swiss needle cast (SNC), caused by a fungal pathogen, Nothophaeocryptopus gaeumannii, is a major forest disease of Douglas-fir (Pseudotsuga menziesii) stands of the Pacific Northwest (PNW). There is mounting concern that the current SNC epidemic occurring in Oregon and Washington will continue to increase in severity, frequency and spatial extent with future warming. Nothophaeocryptopus gaeumannii occurs wherever its host is found, but very little is known about the history and spatial distribution of SNC and its effects on growth and physiological processes of mature and old-growth forests within the Douglas-fir region of the PNW. Our findings show that stem growth and physiological responses of infected Douglas-fir to climate and SNC were different between sites, growth periods and disease severity based on cellulosic stable carbon and oxygen isotope ratios and ring width data in tree rings. At a coastal Oregon site within the SNC impact zone, variations in stem growth and Δ13C were primarily influenced by disproportional reductions in stomatal conductance (gs) and assimilation (A) caused by a loss of functioning stomates through early needle abscission and stomatal occlusion by pseudothecia of N. gaeumannii. At the less severely infected inland sites on the west slopes of Oregon's Cascade Range, stem growth correlated negatively with δ18O and positively with Δ13C, indicating that gs decreased in response to high evaporative demand with a concomitant reduction in A. Current- and previous-years summer vapor pressure deficit was the principal seasonal climatic variable affecting radial stem growth and the dual stable isotope ratios at all sites. Our results indicate that rising temperatures since the mid-1970s has strongly affected Douglas-fir growth in the PNW directly by a physiological response to higher evaporative demand during the annual summer drought and indirectly by a major SNC epidemic that is expanding regionally to higher latitudes and higher elevations.

Climate impacts on source contributions and evaporation to flow in the Snake River Basin using surface water isoscapes (δ2H and δ18O).

Rising global temperatures are expected to decrease the precipitation amount that falls as snow, causing greater risk of water scarcity, groundwater overdraft, and fire in areas that rely on mountain snowpack for their water supply. Streamflow in large river basins varies with the amount, timing, and type of precipitation, evapotranspiration, and drainage properties of watersheds; however, these controls vary in time and space making it difficult to identify the areas contributing most to flow and when. In this study, we separate the evaporative influences from source values of water isotopes from the Snake River Basin in the western United States (US) to relate source area to flow dynamics. We developed isoscapes (δ2H and δ18O) for the basin and found that isotopic composition of surface water in small watersheds is primarily controlled by longitude, latitude, and elevation. To examine temporal variability in source contributions to flow, we present a six-year record of Snake River water isotopes from King Hill, Idaho after removing evaporative influences. During periods of low flow, source water values were isotopically lighter indicating a larger contribution to flow from surface waters in the highest elevation, eastern portion of the basin. River evaporation increases were evident during summer likely reflecting climate, changing water availability, and management strategies within the basin. Our findings present a potential tool for identifying critical portions of basins contributing to river flow as climate fluctuations alter flow dynamics. This tool can be applied in other continental-interior basins where evaporation may obscure source water isotopic signatures.

Context is Everything: Interacting Inputs and Landscape Characteristics Control Stream Nitrogen.

To understand the environmental and anthropogenic drivers of stream nitrogen (N) concentrations across the conterminous US, we combined summer low-flow data from 4997 streams with watershed information across three survey periods (2000-2014) of the US EPA's National Rivers and Streams Assessment. Watershed N inputs explained 51% of the variation in log-transformed stream total N (TN) concentrations. Both N source and input rates influenced stream NO3/TN ratios and N concentrations. Streams dominated by oxidized N forms (NO3/TN ratio > 0.50) were more strongly responsive to the N input rate compared to streams dominated by other N forms. NO3 proportional contribution increased with N inputs, supporting N saturation-enhanced NO3 export to aquatic ecosystems. By combining information about N inputs with climatic and landscape factors, random forest models of stream N concentrations explained 70, 58, and 60% of the spatial variation in stream concentrations of TN, dissolved inorganic N, and total organic N, respectively. The strength and direction of relationships between watershed drivers and stream N concentrations and forms varied with N input intensity. Model results for high N input watersheds not only indicated potential contributions from contaminated groundwater to high stream N concentrations but also the mitigating role of wetlands.

National framework for ranking lakes by potential for anthropogenic hydro-alteration.

Lakes face multiple anthropogenic pressures that can substantially alter their hydrology. Dams and land use in the watershed (e.g., irrigated agriculture) can modify lake water regimes beyond natural ranges, and changing climate may exacerbate anthropogenic stresses on lake hydrology. However, we lack cost-effective indicators to quantify anthropogenic hydrologic alteration potential in lakes at regional and national extents. We developed a framework to rank lakes by the potential for dams and land use to alter lake hydrology (HydrAP) that can be applied at a national scale. The HydrAP framework principles are that 1) dams are primary drivers of lake hydro-alteration, 2) land use activities are secondary drivers that alter watershed hydrology, and 3) topographic relief limits where land use and dams are located on the landscape. We ranked lakes in the United States Environmental Protection Agency National Lakes Assessment (NLA) on a HydrAP scale from zero to seven, where a zero indicates lakes with no potential for anthropogenic hydro-alteration, and a seven indicates large dams and/or intensive land use with high potential to alter lake hydrology. We inferred HydrAP population distributions in the conterminous US (CONUS) using the NLA probabilistic weights. Half of CONUS lakes had moderate to high hydro-alteration potential (HydrAP ranks 3-7), the other half had minimal to no hydro-alteration potential (HydrAP ranks 0-2). HydrAP ranks generally corresponded with natural and man-made lake classes, but >15% of natural lakes had moderate to high HydrAP ranks and ~10% of man-made lakes had low HydrAP ranks. The Great Plains, Appalachians, and Coastal Plains had the largest percentages (>50%) of high HydrAP lakes, and the West and Midwest had the lowest percentages (~30%). Water residence time (τ) and water-level change were associated with HydrAP ranks, demonstrating the framework's intended ability to differentiate anthropogenic stressors that can alter lake hydrology. Consistently across ecoregions high HydrAP lakes had shorter τ. But HydrAP relationships with water-level change varied by ecoregion. In the West and Appalachians, high HydrAP lakes experienced excessive water-level declines compared to low-ranked lakes. In contrast, high HydrAP lakes in the Great Plains and Midwest showed stable water levels compared to low-ranked lakes. These differences imply that water management in western and eastern mountainous regions may result in large water-level fluctuations, but water management in central CONUS may promote water-level stabilization. The HydrAP framework using accessible, national datasets can support large-scale lake assessments and be adapted to specific locations where data are available.

Coupling the dual isotopes of water (δ2H and δ18O) and nitrate (δ15N and δ18O): A new framework for classifying current and legacy groundwater pollution.

Nitrate contamination of groundwater is a concern globally, particularly in agricultural regions where decades of fertilizer nitrogen (N) use has led to a legacy of N accumulation in soils and groundwater. Linkages between current management practices and groundwater nitrate dynamics are often confounded by the legacy effect, and other processes unrelated to management. A coupled analysis of dual stable isotopes of water (δH2O = δ2H and δ18O) and nitrate (δNO3 - = δ15N and δ18O) can be a powerful approach to identify sources and processes responsible for groundwater pollution. To assess how management practices impact groundwater nitrate, we interpreted behavior of δH2O and δNO3 -, together with nitrate concentrations, in water samples collected from long-term monitoring wells in the Southern Willamette Valley (SWV), Oregon. The source(s) of nitrate and water varied among wells, suggesting that the nitrate concentration patterns were not uniform across the shallow aquifer of the valley. Analyzing the stability versus variability of a well's corresponding δH2O and δNO3 - values over time revealed the mechanisms controlling nitrate concentrations. Wells with stable δH2O and δNO3 - values and nitrate concentrations were influenced by one water source with a long residence time and one nitrate source. Variable nitrate concentrations of other wells were attributed to dilution with an alternate water source, mixing of two nitrate sources, or variances in the release of legacy N from overlying soils. Denitrification was not an important process influencing well nitrate dynamics. Understanding the drivers of nitrate dynamics and interaction with legacy N is crucial for managing water quality improvement. This case study illustrates when and where such coupled stable isotope approaches might provide key insights to management on groundwater nitrate contamination issues.

Temperature Decrease along Hyporheic Pathlines in a Large River Riparian Zone.

Hyporheic zones contribute to lower temperatures in many rivers, creating a longitudinal heterogeneous array of thermal refuges. In this study, we had the unique opportunity to show temperature reduction along actual hyporheic zone pathlines in a large river system that contribute to the maintenance of refuges through discharge into off-channel habitats. Temperature was monitored in a dense network of wells that were located along pathlines in small islands, from a calibrated ground-water flow model. Temperature along one 600-m pathline was reduced about 7 °C. Among three islands that were adjacent to the river, the northern two showed exponential decrease in temperature with distance, with fitted thermal Péclet numbers of 2.7 and 6.5, while the southern island showed no significant decrease. We suggest this is due to the higher infiltration rate in the wet season in this larger, more mature island, which suppresses hyporheic flow in the wet season. Stable isotope sampling showed that values of δ2H were higher in areas where we observed lower temperatures. The overall relationship of δ2H versus temperature was significant with a slope of -0.329. This implies that lower temperatures are associated with water that has had contact with deeper groundwater or that lower temperatures have been affected by local rainfall infiltration, or water that has entered the hyporheic zone in winter. These findings are important because they allow estimation of the temperature benefit that may be achieved in similar geomorphic settings, providing implications for riparian restoration.

Lake Water Levels and Associated Hydrologic Characteristics in the Conterminous U.S.

Establishing baseline hydrologic characteristics for lakes in the U.S. is critical to evaluate changes to lake hydrology. We used the U.S. EPA National Lakes Assessment 2007 and 2012 surveys to assess hydrologic characteristics of a population of ~45,000 lakes in the conterminous U.S. based on probability samples of ~1,000 lakes/yr distributed across nine ecoregions. Lake hydrologic study variables include water-level drawdown (i.e., vertical decline and horizontal littoral exposure) and two water stable isotope-derived parameters: evaporation-to-inflow (E:I) and water residence time. We present 1) national and regional distributions of the study variables for both natural and man-made lakes and 2) differences in these characteristics between 2007 and 2012. In 2007, 59% of the population of U.S. lakes had Greater than normal or Excessive drawdown relative to water levels in ecoregional reference lakes with minimal human disturbances; while in 2012, only 20% of lakes were significantly drawn down beyond normal ranges. Water isotope-derived variables did not differ significantly between survey years in contrast to drawdown. Median E:I was 20% indicating that flow-through processes dominated lake water regimes. For 75% of U.S. lakes, water residence time was < 1 year and was longer in natural vs. man-made lakes. Our study provides baseline ranges to assess local and regional lake hydrologic status and inform management decisions in changing environmental conditions.

Trait integration and functional differentiation among co-existing plant species.

PREMISE: Determining which traits characterize strategies of coexisting species is important to developing trait-based models of plant communities. First, global dimensions may not exist locally. Second, the degree to which traits and trait spectra constitute independent dimensions of functional variation at various scales continues to be refined. Finally, traits may be associated with existing categorical groupings. METHODS: We assessed trait integration and differentiation across 57 forest understory plant species in Douglas-fir forests of western Oregon, United States. We combined measurements for a range of traits with literature-based estimates of seed mass and species groupings. We used network analysis and nonmetric multidimensional scaling ordination (NMS) to determine the degree of integration. RESULTS: We observed a strong leaf economics spectrum (LES) integrated with stem but not root traits. However, stem traits and intrinsic water-use efficiency integrated LES and root traits. Network analyses indicated a modest grouping of a priori trait dimensions. NMS indicated that multivariate differences among species were related primarily to (1) rooting depth and plant height vs. specific root length, (2) the LES, and (3) leaf size vs. seed mass. These differences were related to species groupings associated with growth and life form, leaf lifespan and seed dispersal mechanisms. CONCLUSIONS: The strategies of coexisting understory plant species could not be reduced to a single dimension. Yet, species can be characterized efficiently and effectively for trait-based studies of plant communities by measuring four common traits: plant height, specific leaf area, leaf size, and seed mass.

Investigating old-growth ponderosa pine physiology using tree-rings, δ13 C, δ18 O, and a process-based model.

In dealing with predicted changes in environmental conditions outside those experienced today, forest managers and researchers rely on process-based models to inform physiological processes and predict future forest growth responses. The carbon and oxygen isotope ratios of tree-ring cellulose (δ13 Ccell , δ18 Ocell ) reveal long-term, integrated physiological responses to environmental conditions. We incorporated a submodel of δ18 Ocell into the widely used Physiological Principles in Predicting Growth (3-PG) model for the first time, to complement a recently added δ13 Ccell submodel. We parameterized the model using previously reported stand characteristics and long-term trajectories of tree-ring growth, δ13 Ccell , and δ18 Ocell collected from the Metolius AmeriFlux site in central Oregon (upland trees). We then applied the parameterized model to a nearby set of riparian trees to investigate the physiological drivers of differences in observed basal area increment (BAI) and δ13 Ccell trajectories between upland and riparian trees. The model showed that greater available soil water and maximum canopy conductance likely explain the greater observed BAI and lower δ13 Ccell of riparian trees. Unexpectedly, both observed and simulated δ18 Ocell trajectories did not differ between the upland and riparian trees, likely due to similar δ18 O of source water isotope composition. The δ18 Ocell submodel with a Peclet effect improved model estimates of δ18 Ocell because its calculation utilizes 3-PG growth and allocation processes. Because simulated stand-level transpiration (E) is used in the δ18 O submodel, aspects of leaf-level anatomy such as the effective path length for transport of water from the xylem to the sites of evaporation could be estimated.

Performance of National Maps of Watershed Integrity at Watershed Scales.

Watershed integrity, the capacity of a watershed to support and maintain ecological processes essential to the sustainability of services provided to society, can be influenced by a range of landscape and in-stream factors. Ecological response data from four intensively monitored case study watersheds exhibiting a range of environmental conditions and landscape characteristics across the United States were used to evaluate the performance of a national level Index of Watershed Integrity (IWI) at regional and local watershed scales. Using Pearson's correlation coefficient (r), and Spearman's rank correlation coefficient (rs ), response variables displayed highly significant relationships and were significantly correlated with IWI and ICI (Index of Catchment Integrity) values at all watersheds. Nitrogen concentration and flux-related watershed response metrics exhibited significantly strong negative correlations across case study watersheds, with absolute correlations (|r|) ranging from 0.48 to 0.97 for IWI values, and 0.31 to 0.96 for ICI values. Nitrogen-stable isotope ratios measured in chironomids and periphyton from streams and benthic organic matter from lake sediments also demonstrated strong negative correlations with IWI values, with |r| ranging from 0.47 to 0.92, and 0.35 to 0.89 for correlations with ICI values. This evaluation of the performance of national watershed and catchment integrity metrics and their strong relationship with site level responses provides weight-of-evidence support for their use in state, local and regionally focused applications.

Applying the principles of isotope analysis in plant and animal ecology to forensic science in the Americas.

The heart of forensic science is application of the scientific method and analytical approaches to answer questions central to solving a crime: Who, What, When, Where, and How. Forensic practitioners use fundamentals of chemistry and physics to examine evidence and infer its origin. In this regard, ecological researchers have had a significant impact on forensic science through the development and application of a specialized measurement technique-isotope analysis-for examining evidence. Here, we review the utility of isotope analysis in forensic settings from an ecological perspective, concentrating on work from the Americas completed within the last three decades. Our primary focus is on combining plant and animal physiological models with isotope analyses for source inference. Examples of the forensic application of isotopes-including stable isotopes, radiogenic isotopes, and radioisotopes-span from cotton used in counterfeit bills to anthrax shipped through the U.S. Postal Service and from beer adulterated with cheap adjuncts to human remains discovered in shallow graves. Recent methodological developments and the generation of isotope landscapes, or isoscapes, for data interpretation promise that isotope analysis will be a useful tool in ecological and forensic studies for decades to come.

Inferring the source of evaporated waters using stable H and O isotopes.

Stable isotope ratios of H and O are widely used to identify the source of water, e.g., in aquifers, river runoff, soils, plant xylem, and plant-based beverages. In situations where the sampled water is partially evaporated, its isotope values will have evolved along an evaporation line (EL) in δ2H/δ18O space, and back-correction along the EL to its intersection with a meteoric water line (MWL) has been used to estimate the source water's isotope ratios. Here, we review the theory underlying isotopic estimation of source water for evaporated samples (iSWE). We note potential for bias from a commonly used regression-based approach for EL slope estimation and suggest that a model-based approach may be preferable if assumptions of the regression approach are not valid. We then introduce a mathematical framework that eliminates the need to explicitly estimate the EL-MWL intersection, simplifying iSWE analysis and facilitating more rigorous uncertainty estimation. We apply this approach to data from the US EPA's 2007 National Lakes Assessment. We find that data for most lakes are consistent with a water source similar to annual runoff, estimated from monthly precipitation and evaporation within the lake basin. Strong evidence for both summer- and winter-biased sources exists, however, with winter bias pervasive in most snow-prone regions. The new analytical framework should improve the rigor of iSWE in ecohydrology and related sciences, and our initial results from US lakes suggest that previous interpretations of lakes as unbiased isotope integrators may only be valid in certain climate regimes.

Polyploidy influences plant-environment interactions in quaking aspen (Populus tremuloides Michx.).

Quaking aspen (Populus tremuloides Michx.), a widespread and keystone tree species in North America, experienced heat and drought stress in the years 2002 and 2003 in the southwestern United States. This led to widespread aspen mortality that has altered the composition of forests, and is expected to occur again if climate change continues. Understanding interactions between aspen and its environments is essential to understanding future mortality risk in forests. Polyploidy, which is common in aspen, can modify plant structure and function and therefore plant-environment interactions, but the influence of polyploidy on aspen physiology is still not well understood. Furthermore, the ploidy types of aspen have different biogeographies, with triploids being most frequent at lower latitudes in generally warmer and drier climates, while the northerly populations are virtually 100% diploid. This suggests that ploidy-environment interactions differ, and could mean that the ploidy types have different vulnerabilities to environmental stress. In this study, to understand aspen ploidy-environment interactions, we measured 38 different traits important to carbon uptake, water loss and water-use efficiency in diploid and triploid aspen in Colorado. We found that triploid aspen had lower stand density, and greater leaf area, leaf mass, leaf mass per area, percent nitrogen content, chlorophyll content and stomatal size. These differences corresponded to greater potential net carbon assimilation (A, measured using A/Ci curves, and chlorophyll fluorescence) and stomatal conductance (gs) in triploids than diploids. While triploid aspen had higher intrinsic water-use efficiency (iWUE, calculated from measurements of δ13C in leaf tissue), they also had greater potential water loss from higher measured gs and lower stomatal sensitivity to increasing vapor pressure deficit. Therefore, despite greater iWUE, triploids may have lower resilience to climate-induced stress. We conclude that ploidy type strongly influences physiological traits and function, and mediates drought stress responses in quaking aspen.

A sprinkling experiment to quantify celerity-velocity differences at the hillslope scale.

Few studies have quantified the differences between celerity and velocity of hillslope water flow and explained the processes that control these differences. Here, we asses these differences by combining a 24-day hillslope sprinkling experiment with a spatially explicit hydrologic model analysis. We focused our work on Watershed 10 at the H. J. Andrews Experimental Forest in western Oregon. Celerities estimated from wetting front arrival times were generally much faster than average vertical velocities of δ2H. In the model analysis, this was consistent with an identifiable effective porosity (fraction of total porosity available for mass transfer) parameter, indicating that subsurface mixing was controlled by an immobile soil fraction, resulting in the attenuation of the δ2H input signal in lateral subsurface flow. In addition to the immobile soil fraction, exfiltrating deep groundwater that mixed with lateral subsurface flow captured at the experimental hillslope trench caused further reduction in the δ2H input signal. Finally, our results suggest that soil depth variability played a significant role in the celerity-velocity responses. Deeper upslope soils damped the δ2H input signal, while a shallow soil near the trench controlled the δ2H peak in lateral subsurface flow response. Simulated exit time and residence time distributions with our hillslope hydrologic model showed that water captured at the trench did not represent the entire modeled hillslope domain; the exit time distribution for lateral subsurface flow captured at the trench showed more early time weighting.

Basal area growth, carbon isotope discrimination, and intrinsic water use efficiency after fertilization of Douglas-fir in the Oregon Coast Range.

Many hectares of intensively managed Douglas-fir (Pseudotsuga menziesii Mirb. Franco) stands in western North America are fertilized with nitrogen (N) to increase growth rates, but only about ⅔ of all stands respond. Understanding the mechanisms of response facilitates prioritization of stands for treatment. The primary objective of this study was to test the hypothesis that the short-term basal area growth response to a single application of 224 kg N ha-1 as urea was associated with reduced stable carbon isotope discrimination (Δ13C) and increased intrinsic water use efficiency (iWUE) in a 20-yr-old plantation of Douglas-fir in the Oregon Coast Range, USA. Increment cores were measured to estimate earlywood, latewood, and total basal area increment over a time series from 1997 to 2015. Stable carbon isotope discrimination and iWUE were estimated using earlywood and latewood stable carbon isotope concentrations in tree-ring holocellulose starting seven years before fertilization in early 2009 and ending seven years after treatment. A highly significant (p<0.01) interaction effect between fertilization treatment and year was found for total basal area growth and earlywood basal area increment. Specifically, fertilized trees showed significant responses (p<0.05) in total basal area growth and earlywood basal area increment in the first (2009) and second (2010) growing seasons after fertilization in 2009. A marginally significant (p<0.10) fertilization effect was found for latewood basal area increment only in the first growing season after treatment. A significant treatment x year interaction was also found for Δ13C and iWUE in earlywood and latewood. Fertilization significantly reduced earlywood Δ13C and increased earlywood iWUE in the first and second growing seasons after fertilization. Only a marginally significant fertilization effect was detected for latewood Δ13C and iWUE in the second growing season after treatment. Previous studies of N fertilization of Douglas-fir forests have reported consistently increased growth and iWUE on low productivity sites treated with relatively high fertilization rates. This study suggested that these responses can also be observed on highly productive sites despite their lower frequency and apparently shorter duration. Other key mechanisms driving growth responses appear less important than iWUE, including an increase in LAI and shift from belowground to aboveground carbon allocation.