Shifting stoichiometry: Long-term trends in stream-dissolved organic matter reveal altered C:N ratios due to history of atmospheric acid deposition.

Dissolved organic carbon (DOC) and nitrogen (DON) are important energy and nutrient sources for aquatic ecosystems. In many northern temperate, freshwater systems DOC has increased in the past 50 years. Less is known about how changes in DOC may vary across latitudes, and whether changes in DON track those of DOC. Here, we present long-term DOC and DON data from 74 streams distributed across seven sites in biomes ranging from the tropics to northern boreal forests with varying histories of atmospheric acid deposition. For each stream, we examined the temporal trends of DOC and DON concentrations and DOC:DON molar ratios. While some sites displayed consistent positive or negative trends in stream DOC and DON concentrations, changes in direction or magnitude were inconsistent at regional or local scales. DON trends did not always track those of DOC, though DOC:DON ratios increased over time for ~30% of streams. Our results indicate that the dissolved organic matter (DOM) pool is experiencing fundamental changes due to the recovery from atmospheric acid deposition. Changes in DOC:DON stoichiometry point to a shifting energy-nutrient balance in many aquatic ecosystems. Sustained changes in the character of DOM can have major implications for stream metabolism, biogeochemical processes, food webs, and drinking water quality (including disinfection by-products). Understanding regional and global variation in DOC and DON concentrations is important for developing realistic models and watershed management protocols to effectively target mitigation efforts aimed at bringing DOM flux and nutrient enrichment under control.

Stream Mercury Export in Response to Contemporary Timber Harvesting Methods (Pacific Coastal Mountains, Oregon, USA).

Land-use activities can alter hydrological and biogeochemical processes that can affect the fate, transformation, and transport of mercury (Hg). Previous studies in boreal forests have shown that forestry operations can have profound but variable effects on Hg export and methylmercury (MeHg) formation. The Pacific Northwest is an important timber producing region that receives large atmospheric Hg loads, but the impact of forest harvesting on Hg mobilization has not been directly studied and was the focus of our investigation. Stream discharge was measured continuously, and Hg and MeHg concentrations were measured monthly for 1.5 years following logging in three paired harvested and unharvested (control) catchments. There was no significant difference in particulate-bound Hg concentrations or loads in the harvested and unharvested catchments which may have resulted from forestry practices aimed at minimizing erosion. However, the harvested catchments had significantly higher discharge (32%), filtered Hg concentrations (28%), filtered Hg loads (80%), and dissolved organic carbon (DOC) loads (40%) compared to forested catchments. MeHg concentrations were low (mostly <0.05 ng L-1) in harvested, unharvested, and downstream samples due to well-drained/unsaturated soil conditions and steep slopes with high energy eroding stream channels that were not conducive to the development of anoxic conditions that support methylation. These results have important implications for the role forestry operations have in affecting catchment retention and export of Hg pollution.

Drivers of nitrogen transfer in stream food webs across continents.

Studies of trophic-level material and energy transfers are central to ecology. The use of isotopic tracers has now made it possible to measure trophic transfer efficiencies of important nutrients and to better understand how these materials move through food webs. We analyzed data from thirteen 15 N-ammonium tracer addition experiments to quantify N transfer from basal resources to animals in headwater streams with varying physical, chemical, and biological features. N transfer efficiencies from primary uptake compartments (PUCs; heterotrophic microorganisms and primary producers) to primary consumers was lower (mean 11.5%, range <1% to 43%) than N transfer efficiencies from primary consumers to predators (mean 80%, range 5% to >100%). Total N transferred (as a rate) was greater in streams with open compared to closed canopies and overall N transfer efficiency generally followed a similar pattern, although was not statistically significant. We used principal component analysis to condense a suite of site characteristics into two environmental components. Total N uptake rates among trophic levels were best predicted by the component that was correlated with latitude, DIN:SRP, GPP:ER, and percent canopy cover. N transfer efficiency did not respond consistently to environmental variables. Our results suggest that canopy cover influences N movement through stream food webs because light availability and primary production facilitate N transfer to higher trophic levels.

Global synthesis of the temperature sensitivity of leaf litter breakdown in streams and rivers.

Streams and rivers are important conduits of terrestrially derived carbon (C) to atmospheric and marine reservoirs. Leaf litter breakdown rates are expected to increase as water temperatures rise in response to climate change. The magnitude of increase in breakdown rates is uncertain, given differences in litter quality and microbial and detritivore community responses to temperature, factors that can influence the apparent temperature sensitivity of breakdown and the relative proportion of C lost to the atmosphere vs. stored or transported downstream. Here, we synthesized 1025 records of litter breakdown in streams and rivers to quantify its temperature sensitivity, as measured by the activation energy (Ea , in eV). Temperature sensitivity of litter breakdown varied among twelve plant genera for which Ea could be calculated. Higher values of Ea were correlated with lower-quality litter, but these correlations were influenced by a single, N-fixing genus (Alnus). Ea values converged when genera were classified into three breakdown rate categories, potentially due to continual water availability in streams and rivers modulating the influence of leaf chemistry on breakdown. Across all data representing 85 plant genera, the Ea was 0.34 ± 0.04 eV, or approximately half the value (0.65 eV) predicted by metabolic theory. Our results indicate that average breakdown rates may increase by 5-21% with a 1-4 °C rise in water temperature, rather than a 10-45% increase expected, according to metabolic theory. Differential warming of tropical and temperate biomes could result in a similar proportional increase in breakdown rates, despite variation in Ea values for these regions (0.75 ± 0.13 eV and 0.27 ± 0.05 eV, respectively). The relative proportions of gaseous C loss and organic matter transport downstream should not change with rising temperature given that Ea values for breakdown mediated by microbes alone and microbes plus detritivores were similar at the global scale.

Stream denitrification across biomes and its response to anthropogenic nitrate loading.

Anthropogenic addition of bioavailable nitrogen to the biosphere is increasing and terrestrial ecosystems are becoming increasingly nitrogen-saturated, causing more bioavailable nitrogen to enter groundwater and surface waters. Large-scale nitrogen budgets show that an average of about 20-25 per cent of the nitrogen added to the biosphere is exported from rivers to the ocean or inland basins, indicating that substantial sinks for nitrogen must exist in the landscape. Streams and rivers may themselves be important sinks for bioavailable nitrogen owing to their hydrological connections with terrestrial systems, high rates of biological activity, and streambed sediment environments that favour microbial denitrification. Here we present data from nitrogen stable isotope tracer experiments across 72 streams and 8 regions representing several biomes. We show that total biotic uptake and denitrification of nitrate increase with stream nitrate concentration, but that the efficiency of biotic uptake and denitrification declines as concentration increases, reducing the proportion of in-stream nitrate that is removed from transport. Our data suggest that the total uptake of nitrate is related to ecosystem photosynthesis and that denitrification is related to ecosystem respiration. In addition, we use a stream network model to demonstrate that excess nitrate in streams elicits a disproportionate increase in the fraction of nitrate that is exported to receiving waters and reduces the relative role of small versus large streams as nitrate sinks.

Nitrous oxide emission from denitrification in stream and river networks.

Nitrous oxide (N(2)O) is a potent greenhouse gas that contributes to climate change and stratospheric ozone destruction. Anthropogenic nitrogen (N) loading to river networks is a potentially important source of N(2)O via microbial denitrification that converts N to N(2)O and dinitrogen (N(2)). The fraction of denitrified N that escapes as N(2)O rather than N(2) (i.e., the N(2)O yield) is an important determinant of how much N(2)O is produced by river networks, but little is known about the N(2)O yield in flowing waters. Here, we present the results of whole-stream (15)N-tracer additions conducted in 72 headwater streams draining multiple land-use types across the United States. We found that stream denitrification produces N(2)O at rates that increase with stream water nitrate (NO(3)(-)) concentrations, but that <1% of denitrified N is converted to N(2)O. Unlike some previous studies, we found no relationship between the N(2)O yield and stream water NO(3)(-). We suggest that increased stream NO(3)(-) loading stimulates denitrification and concomitant N(2)O production, but does not increase the N(2)O yield. In our study, most streams were sources of N(2)O to the atmosphere and the highest emission rates were observed in streams draining urban basins. Using a global river network model, we estimate that microbial N transformations (e.g., denitrification and nitrification) convert at least 0.68 Tg·y(-1) of anthropogenic N inputs to N(2)O in river networks, equivalent to 10% of the global anthropogenic N(2)O emission rate. This estimate of stream and river N(2)O emissions is three times greater than estimated by the Intergovernmental Panel on Climate Change.

Stream nitrate enrichment and increased light yet no algal response following forest harvest and experimental manipulation of headwater riparian zones.

Disturbances to forested watersheds often result in increases of nutrients and light to nearby streams. Such changes are generally expected to produce a shift to a more autotrophic aquatic ecosystem, with measurable increases in algae, and associated implications for food webs and fisheries. Although this paradigm is widely established, results from our 10-year study (2007-2016) in 12 headwater streams and four sites downstream in the Trask River Watershed (Oregon, USA), did not concur. In 2012, one watershed was thinned, three were clearcut harvested with variable buffers and three with uniform riparian buffers. After harvest, light to the stream surface significantly increased at the three watersheds with variable buffers while dissolved inorganic nitrogen (DIN) significantly increased in all of the clearcut harvested streams. Despite the increase in DIN and light, algal standing stocks and chlorophyll a concentrations did not significantly increase. The common assumption of increased autotrophic responses in stream food webs following increases of nitrogen and light was not supported here. We postulate the co-limitation of nutrients, driven by low phosphorus concentrations, which unlike DIN did not increase post-harvest, and the characteristics of the algal community, which were dominated by low light adapted diatoms rather than green algae, contributed to our findings of no responses for standing stocks of epilithic algae or concentrations of chlorophyll a. The inclusion of multiple statistical analyses provided more certainty around our findings. This study documents responses to current forest practices and provides cautionary information for management and restoration activities aiming to increase fish abundance and standing stocks by opening riparian canopies and adding nutrients.

UPRLIMET: UPstream Regional LiDAR Model for Extent of Trout in stream networks.

Predicting the edges of species distributions is fundamental for species conservation, ecosystem services, and management decisions. In North America, the location of the upstream limit of fish in forested streams receives special attention, because fish-bearing portions of streams have more protections during forest management activities than fishless portions. We present a novel model development and evaluation framework, wherein we compare 26 models to predict upper distribution limits of trout in streams. The models used machine learning, logistic regression, and a sophisticated nested spatial cross-validation routine to evaluate predictive performance while accounting for spatial autocorrelation. The model resulting in the best predictive performance, termed UPstream Regional LiDAR Model for Extent of Trout (UPRLIMET), is a two-stage model that uses a logistic regression algorithm calibrated to observations of Coastal Cutthroat Trout (Oncorhynchus clarkii clarkii) occurrence and variables representing hydro-topographic characteristics of the landscape. We predict trout presence along reaches throughout a stream network, and include a stopping rule to identify a discrete upper limit point above which all stream reaches are classified as fishless. Although there is no simple explanation for the upper distribution limit identified in UPRLIMET, four factors, including upstream channel length above the point of uppermost fish, drainage area, slope, and elevation, had highest importance. Across our study region of western Oregon, we found that more of the fish-bearing network is on private lands than on state, US Bureau of Land Mangement (BLM), or USDA Forest Service (USFS) lands, highlighting the importance of using spatially consistent maps across a region and working across land ownerships. Our research underscores the value of using occurrence data to develop simple, but powerful, prediction tools to capture complex ecological processes that contribute to distribution limits of species.

Beyond Streamflow: Call for a National Data Repository of Streamflow Presence for Streams and Rivers in the United States.

Observations of the presence or absence of surface water in streams are useful for characterizing streamflow permanence, which includes the frequency, duration, and spatial extent of surface flow in streams and rivers. Such data are particularly valuable for headwater streams, which comprise the vast majority of channel length in stream networks, are often non-perennial, and are frequently the most data deficient. Datasets of surface water presence exist across multiple data collection groups in the United States but are not well aligned for easy integration. Given the value of these data, a unified approach for organizing information on surface water presence and absence collected by diverse surveys would facilitate more effective and broad application of these data and address the gap in streamflow data in headwaters. In this paper, we highlight the numerous existing datasets on surface water presence in headwater streams, including recently developed crowdsourcing approaches. We identify the challenges of integrating multiple surface water presence/absence datasets that include differences in the definitions and categories of streamflow status, data collection method, spatial and temporal resolution, and accuracy of geographic location. Finally, we provide a list of critical and useful components that could be used to integrate different streamflow permanence datasets.

Experimental nitrogen and phosphorus enrichment stimulates multiple trophic levels of algal and detrital-based food webs: a global meta-analysis from streams and rivers.

Anthropogenic increases in nitrogen (N) and phosphorus (P) concentrations can strongly influence the structure and function of ecosystems. Even though lotic ecosystems receive cumulative inputs of nutrients applied to and deposited on land, no comprehensive assessment has quantified nutrient-enrichment effects within streams and rivers. We conducted a meta-analysis of published studies that experimentally increased concentrations of N and/or P in streams and rivers to examine how enrichment alters ecosystem structure (state: primary producer and consumer biomass and abundance) and function (rate: primary production, leaf breakdown rates, metabolism) at multiple trophic levels (primary producer, microbial heterotroph, primary and secondary consumers, and integrated ecosystem). Our synthesis included 184 studies, 885 experiments, and 3497 biotic responses to nutrient enrichment. We documented widespread increases in organismal biomass and abundance (mean response = +48%) and rates of ecosystem processes (+54%) to enrichment across multiple trophic levels, with no large differences in responses among trophic levels or between autotrophic or heterotrophic food-web pathways. Responses to nutrient enrichment varied with the nutrient added (N, P, or both) depending on rate versus state variable and experiment type, and were greater in flume and whole-stream experiments than in experiments using nutrient-diffusing substrata. Generally, nutrient-enrichment effects also increased with water temperature and light, and decreased under elevated ambient concentrations of inorganic N and/or P. Overall, increased concentrations of N and/or P altered multiple food-web pathways and trophic levels in lotic ecosystems. Our results indicate that preservation or restoration of biodiversity and ecosystem functions of streams and rivers requires management of nutrient inputs and consideration of multiple trophic pathways.

Short-term reservoir draining to streambed for juvenile salmon passage and non-native fish removal.

Fish passage out of reservoirs is a critical issue for downstream movement of juvenile salmonids and other migratory species. Reservoirs can delay downstream migrations by juvenile salmon for months or years. Here, we examine whether a novel management activity implementing annual short-term draining of a reservoir to streambed improves timely downstream migration of juvenile salmonids. We analyse 12 years of fish capture data from a screw trap located downstream of Fall Creek Reservoir (Oregon, USA) to examine changes in timing of passage out of the reservoir and to compare fish species composition pre- and post-draining. We observed a contraction in the timing of downstream migration for juvenile Chinook Salmon and reduction of yearlings in years following draining. We suggest that briefly draining the reservoir to streambed leads to reduced abundance of warm-water invasive fishes in the reservoir after it refills. These changes could decrease predation and shift competition between invasive and resident riverine-adapted native fishes in the reservoir. Collectively, our findings suggest that this low-cost reservoir management option may improve passage and connectivity for juvenile Chinook Salmon while also decreasing the abundance of invasive fish species in the reservoir. This case study underscores the crucial need for further evaluations of reservoir draining in other systems and contexts.

Evidence for lasting alterations to aquatic food webs with short-duration reservoir draining.

Large dams and their respective reservoirs provide renewable energy and water security, but also can profoundly alter riverine ecosystems. Here, we present evidence of changing aquatic food web structure in the seasons following short-duration, extreme manipulation of water levels in a reservoir (i.e., draining of the reservoir to the original riverbed during fall to assist outmigration of juvenile Chinook Salmon). We find unintended and lagged consequences of transitioning from a lake to a river, even temporarily, that resulted in trophic shifts away from piscivory and towards feeding at lower trophic levels for two common piscivorous fishes in reservoirs. Using natural abundances of nitrogen stable isotopes, we observed lower trophic level of feeding for invasive Largemouth Bass (Micropterus salmoides) and native Rainbow Trout (Oncorhynchus mykiss) during the summers following reservoir refilling than in nearby reference reservoirs that were not temporarily drained during fall. Declines in trophic levels of aquatic top predators have been rarely documented outside of controlled laboratory conditions. While useful for assisting outmigration of juvenile salmonids, the temporary draining of a reservoir to riverbed can also result in novel shifts in foodweb dynamics including reduced piscivory. As large dams continue to be operated and constructed worldwide, increased understanding of the community and ecosystem-level effects of reservoir management will be critical to evaluating trade-offs between human water needs, conservation of high value species, and ecosystem services impacted by river fragmentation.

Projections of water, carbon, and nitrogen dynamics under future climate change in an old-growth Douglas-fir forest in the western Cascade Range using a biogeochemical model.

Statistically downscaled climate change scenarios from four General Circulation Models for two Representative Concentration Pathways (RCP) were applied as inputs to a biogeochemical model, PnET-BGC, to examine potential future dynamics of water, carbon, and nitrogen in an old-growth Douglas-fir forest in the western Cascade Range. Projections show 56% to 77% increases in stomatal conductance throughout the year from 1986-2010 to 2076-2100, and 65% to 104% increases in leaf carbon assimilation between October and June over the same period. However, future dynamics of water and carbon under the RCP scenarios are affected by a 49% to 86% reduction in foliar biomass resulting from severe air temperature and humidity stress to the forest in summer. Important implications of future decreases in foliar biomass include 1) 20% to 71% decreases in annual transpiration which increase soil moisture by 7% to 15% in summer and fall; 2) decreases in photosynthesis by 77% and soil organic matter by 62% under the high radiative forcing scenario; and 3) altered foliar and soil carbon to nitrogen stoichiometry. Potential carbon dioxide fertilization effects on vegetation are projected to 1) amplify decreases in transpiration by 4% to 9% and increases in soil moisture in summer and fall by 1% to 2%; and 2) alleviate decreases in photosynthesis by 4%; while 3) having negligible effects on the dynamics of nitrogen. Our projections suggest that future decrease in transpiration and moderate water holding capacity may mitigate soil moisture stress to the old-growth Douglas-fir forest. Future increases in nitrogen concentration in soil organic matter are projected to alleviate the decrease in net nitrogen mineralization despite a reduction in decomposition of soil organic matter by the end of the century.

Spatial models reveal the microclimatic buffering capacity of old-growth forests.

Climate change is predicted to cause widespread declines in biodiversity, but these predictions are derived from coarse-resolution climate models applied at global scales. Such models lack the capacity to incorporate microclimate variability, which is critical to biodiversity microrefugia. In forested montane regions, microclimate is thought to be influenced by combined effects of elevation, microtopography, and vegetation, but their relative effects at fine spatial scales are poorly known. We used boosted regression trees to model the spatial distribution of fine-scale, under-canopy air temperatures in mountainous terrain. Spatial models predicted observed independent test data well (r = 0.87). As expected, elevation strongly predicted temperatures, but vegetation and microtopography also exerted critical effects. Old-growth vegetation characteristics, measured using LiDAR (light detection and ranging), appeared to have an insulating effect; maximum spring monthly temperatures decreased by 2.5°C across the observed gradient in old-growth structure. These cooling effects across a gradient in forest structure are of similar magnitude to 50-year forecasts of the Intergovernmental Panel on Climate Change and therefore have the potential to mitigate climate warming at local scales. Management strategies to conserve old-growth characteristics and to curb current rates of primary forest loss could maintain microrefugia, enhancing biodiversity persistence in mountainous systems under climate warming.

Local Variability Mediates Vulnerability of Trout Populations to Land Use and Climate Change.

Land use and climate change occur simultaneously around the globe. Fully understanding their separate and combined effects requires a mechanistic understanding at the local scale where their effects are ultimately realized. Here we applied an individual-based model of fish population dynamics to evaluate the role of local stream variability in modifying responses of Coastal Cutthroat Trout (Oncorhynchus clarkii clarkii) to scenarios simulating identical changes in temperature and stream flows linked to forest harvest, climate change, and their combined effects over six decades. We parameterized the model for four neighboring streams located in a forested headwater catchment in northwestern Oregon, USA with multi-year, daily measurements of stream temperature, flow, and turbidity (2007-2011), and field measurements of both instream habitat structure and three years of annual trout population estimates. Model simulations revealed that variability in habitat conditions among streams (depth, available habitat) mediated the effects of forest harvest and climate change. Net effects for most simulated trout responses were different from or less than the sum of their separate scenarios. In some cases, forest harvest countered the effects of climate change through increased summer flow. Climate change most strongly influenced trout (earlier fry emergence, reductions in biomass of older trout, increased biomass of young-of-year), but these changes did not consistently translate into reductions in biomass over time. Forest harvest, in contrast, produced fewer and less consistent responses in trout. Earlier fry emergence driven by climate change was the most consistent simulated response, whereas survival, growth, and biomass were inconsistent. Overall our findings indicate a host of local processes can strongly influence how populations respond to broad scale effects of land use and climate change.

Toward efficient riparian restoration: integrating economic, physical, and biological models.

This paper integrates economic, biological, and physical models to explore the efficient combination and spatial allocation of conservation efforts to protect water quality and increase salmonid populations in the Grande Ronde basin, Oregon. We focus on the effects of shade on water temperatures and the subsequent impacts on endangered juvenile salmonid populations. The integrated modeling system consists of a physical model that links riparian conditions and hydrological characteristics to water temperature; a biological model that links water temperature and riparian conditions to salmonid abundance, and an economic model that incorporates both physical and biological models to estimate minimum cost allocations of conservation efforts. Our findings indicate that conservation alternatives such as passive and active riparian restoration, the width of riparian restoration zones, and the types of vegetation used in restoration activities should be selected based on the spatial distribution of riparian characteristics in the basin. The relative effectiveness of passive and active restoration plays an important role in determining the efficient allocations of conservation efforts. The time frame considered in the restoration efforts and the magnitude of desired temperature reductions also affect the efficient combinations of restoration activities. If the objective of conservation efforts is to maximize fish populations, then fishery benefits should be directly targeted. Targeting other criterion such as water temperatures would result in different allocations of conservation efforts, and therefore are not generally efficient.