Genus-level, trait-based multimetric diatom indices for assessing the ecological condition of rivers and streams across the conterminous United States.

Taxonomic inconsistency in species-level identifications has constrained use of diatoms as biological indicators in aquatic assessments. We addressed this problem by developing diatom multimetric indices (MMIs) of ecological condition using genus-level taxonomy and trait-based autecological information. The MMIs were designed to assess river and stream chemical, physical and biological condition across the conterminous United States. Trait-based approaches have the advantage of using both species-level and genus-level data, which require less effort and expense to acquire than traditional species-based approaches and eliminate the persistent taxonomic biases introduced over vast geographic extents. For large-extent assessment programs that require multiple taxonomic laboratories to process samples, such as the United States Environmental Protection Agency's (U.S. EPA's) National Rivers and Streams Assessment (NRSA), the trait approach can eliminate discrepancies in species-level identification or nomenclature that hinder diatom data interpretation. We developed trait-based MMIs using NRSA data for each of the three large ecoregions across the U.S. - the East, Plains, and West. All three MMIs performed well in discriminating least-disturbed from most-disturbed sites. The MMI for the East had the greatest discrimination ability, followed by MMIs for the Plains and West, respectively. The performance of the MMIs was comparable to that observed in existing NRSA fish and macroinvertebrate MMIs. Our research shows that trait-based diatom indices constructed on genus-level taxonomy can be effective for large-scale assessments, and may also allow programs such as NRSA to assess trends in freshwater condition retrospectively, by revisiting older diatom datasets. Moreover, our genus-based approach facilitates including of diatoms into other assessment programs that have limited monitoring resources.

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.

Mapping watershed integrity for the conterminous United States.

Watershed integrity is the capacity of a watershed to support and maintain the full range of ecological processes and functions essential to sustainability. Using information from EPA's StreamCat dataset, we calculated and mapped an Index of Watershed Integrity (IWI) for 2.6 million watersheds in the conterminous US with first-order approximations of relationships between stressors and six watershed functions: hydrologic regulation, regulation of water chemistry, sediment regulation, hydrologic connectivity, temperature regulation, and habitat provision. Results show high integrity in the western US, intermediate integrity in the southern and eastern US, and the lowest integrity in the temperate plains and lower Mississippi Valley. Correlation between the six functional components was high (r = 0.85-0.98). A related Index of Catchment Integrity (ICI) was developed using local drainages of individual stream segments (i.e., excluding upstream information). We evaluated the ability of the IWI and ICI to predict six continuous site-level indicators with regression analyses - three biological indicators and principal components derived from water quality, habitat, and combined water quality and habitat variables - using data from EPA's National Rivers and Streams Assessment. Relationships were highly significant, but the IWI only accounted for 1-12% of the variation in the four biological and habitat variables. The IWI accounted for over 25% of the variation in the water quality and combined principal components nationally, and 32-39% in the Northern and Southern Appalachians. We also used multinomial logistic regression to compare the IWI with the categorical forms of the three biological indicators. Results were consistent: we found positive associations but modest results. We compared how the IWI and ICI predicted the water quality PC relative to agricultural and urban land use. The IWI or ICI are the best predictors of the water quality PC for the CONUS and six of the nine ecoregions, but they only perform marginally better than agriculture in most instances. However, results suggest that agriculture would not be appropriate in all parts of the country, and the index is meant to be responsive to all stressors. The IWI in its present form (available through the StreamCat website; https://www.epa.gov/national-aquatic-resource-surveys/streamcat) could be useful for management efforts at multiple scales, especially when combined with information on site condition. The IWI could be improved by incorporating empirical or literature-derived relationships between functional components and stressors. However, limitations concerning the absence of data for certain stressors should be considered.

Continental-Scale Increase in Lake and Stream Phosphorus: Are Oligotrophic Systems Disappearing in the United States?

We describe continental-scale increases in lake and stream total phosphorus (TP) concentrations, identified through periodic probability surveys of thousands of water bodies in the conterminous U.S. The increases, observed over the period 2000-2014 were most notable in sites in relatively undisturbed catchments and where TP was initially low (e.g., less than 10 µg L(-1)). Nationally, the percentage of stream length in the U.S. with TP ≤ 10 µg L(-1) decreased from 24.5 to 10.4 to 1.6% from 2004 to 2009 to 2014; the percentage of lakes with TP ≤ 10 µg L(-1) decreased from 24.9 to 6.7% between 2007 and 2012. Increasing TP concentrations appear to be ubiquitous, but their presence in undeveloped catchments suggests that they cannot be entirely attributed to either point or common non-point sources of TP.

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.

Dissolved organic carbon trends resulting from changes in atmospheric deposition chemistry.

Several hypotheses have been proposed to explain recent, widespread increases in concentrations of dissolved organic carbon (DOC) in the surface waters of glaciated landscapes across eastern North America and northern and central Europe. Some invoke anthropogenic forcing through mechanisms related to climate change, nitrogen deposition or changes in land use, and by implication suggest that current concentrations and fluxes are without precedent. All of these hypotheses imply that DOC levels will continue to rise, with unpredictable consequences for the global carbon cycle. Alternatively, it has been proposed that DOC concentrations are returning toward pre-industrial levels as a result of a gradual decline in the sulphate content of atmospheric deposition. Here we show, through the assessment of time series data from 522 remote lakes and streams in North America and northern Europe, that rising trends in DOC between 1990 and 2004 can be concisely explained by a simple model based solely on changes in deposition chemistry and catchment acid-sensitivity. We demonstrate that DOC concentrations have increased in proportion to the rates at which atmospherically deposited anthropogenic sulphur and sea salt have declined. We conclude that acid deposition to these ecosystems has been partially buffered by changes in organic acidity and that the rise in DOC is integral to recovery from acidification. Over recent decades, deposition-driven increases in organic matter solubility may have increased the export of DOC to the oceans, a potentially important component of regional carbon balances. The increase in DOC concentrations in these regions appears unrelated to other climatic factors.

Shifts in the composition of nitrogen deposition in the conterminous United States are discernable in stream chemistry.

Across the conterminous United States (U.S.), the composition of atmospheric nitrogen (N) deposition is changing spatially and temporally. Previously, deposition was dominated by oxidized N, but now reduced N (ammonia [NH3] + ammonium [NH4+]) is equivalent to oxidized N when deposition is averaged across the entire nation and, in some areas, reduced N dominates deposition. To evaluate if there are effects of this change on stream chemistry at the national scale, estimates of N deposition form (oxidized or reduced) from the National Atmospheric Deposition Program Total Deposition data were coupled with stream measurements from the U.S. Environmental Protection Agency (EPA) National Rivers and Streams Assessments (three stream surveys between 2000 and 2014). A recent fine-scaled N input inventory was used to identify watersheds (<1000 km2) where atmospheric deposition is the largest N source (n = 1906). Within these more atmospherically-influenced watersheds there was a clear temporal shift from a greater proportion of sites dominated by oxidized N deposition to a greater proportion of sites dominated by reduced forms of N deposition. We found a significant positive correlation between oxidized N deposition and stream NO3- concentrations, whereas the correlation between reduced N deposition and stream NO3- concentrations were significant but weaker. Sites dominated by atmospheric inputs of reduced N forms had higher stream total organic N and total N despite lower total N deposition on average. This higher stream concentration of total N is mainly driven by the higher concentration of total organic N, suggesting an interaction between elevated reduced N in deposition and living components of the ecosystem or soil organic matter dynamics. Regardless of the proportion of reduced to oxidized N forms in deposition, stream NH4+ concentrations were generally low, suggesting that N deposited in a reduced form is rapidly immobilized, nitrified and/or assimilated by watershed processes.

Long-term rise in riverine dissolved organic carbon concentration is predicted by electrolyte solubility theory.

The riverine dissolved organic carbon (DOC) flux is of similar magnitude to the terrestrial sink for atmospheric CO2, but the factors controlling it remain poorly determined and are largely absent from Earth system models (ESMs). Here, we show, for a range of European headwater catchments, that electrolyte solubility theory explains how declining precipitation ionic strength (IS) has increased the dissolution of thermally moderated pools of soluble soil organic matter (OM), while hydrological conditions govern the proportion of this OM entering the aquatic system. Solubility will continue to rise exponentially with declining IS until pollutant ion deposition fully flattens out under clean air policies. Future DOC export will increasingly depend on rates of warming and any directional changes to the intensity and seasonality of precipitation and marine ion deposition. Our findings provide a firm foundation for incorporating the processes dominating change in this component of the global carbon cycle in ESMs.

New insights into the source of decadal increases of dissolved organic matter in acid-sensitive lakes of the northeastern United States.

The last several decades have seen decreases in SO(4)(2-) deposition across the northeastern United States. As a result, SO(4)(2-) concentrations in lakes and streams have also decreased and many surface water bodies have become less acidic. During the same time period, there has been a concurrent increase in dissolved organic carbon (DOC) concentrations in many lakes and streams. We used fluorescence spectroscopy to characterize the dissolved organic matter (DOM) quality of archived samples from nine acid-sensitive lakes in Maine collected between 1993 and 2009, and determined that increased DOM contributions to lakes were primarily derived from litter and soil. All five lakes with increasing DOC trends demonstrated significant decreasing (i.e., more terrestrial) trends in fluorescence index (FI) and significant positive correlations between SO(4)(2-) and FI. This study used the chemical signature of terrestrial DOM to support the hypothesis that increased DOC concentrations in lakes and streams are driven by declining acid deposition and increased solubility of soil organic matter across a large area of the landscape.

Cleaner air reveals growing influence of climate on dissolved organic carbon trends in northern headwaters.

Surface water browning, the result of increasing concentrations of dissolved organic matter (DOM), has been widespread in northern ecosystems in recent decades. Here, we assess a database of 426 undisturbed headwater lakes and streams in Europe and North America for evidence of trends in DOM between 1990 and 2016. We describe contrasting changes in DOM trends in Europe (decelerating) and North America (accelerating), which are consistent with organic matter solubility responses to declines in sulfate deposition. While earlier trends (1990-2004) were almost entirely related to changes in atmospheric chemistry, climatic and chemical drivers were equally important in explaining recent DOM trends (2002-2016). We estimate that riverine DOM export from northern ecosystems increased by 27% during the study period. Increased summer precipitation strengthened upward dissolved organic carbon trends while warming apparently damped browning. Our results suggest strong but changing influences of air quality and climate on the terrestrial carbon cycle, and on the magnitude of carbon export from land to water.

Setting expectations for the ecological condition of streams: the concept of reference condition.

An important component of the biological assessment of stream condition is an evaluation of the direct or indirect effects of human activities or disturbances. The concept of a "reference condition" is increasingly used to describe the standard or benchmark against which current condition is compared. Many individual nations, and the European Union as a whole, have codified the concept of reference condition in legislation aimed at protecting and improving the ecological condition of streams. However, the phrase "reference condition" has many meanings in a variety of contexts. One of the primary purposes of this paper is to bring some consistency to the use of the term. We argue the need for a "reference condition" term that is reserved for referring to the "naturalness" of the biota (structure and function) and that naturalness implies the absence of significant human disturbance or alteration. To avoid the confusion that arises when alternative definitions of reference condition are used, we propose that the original concept of reference condition be preserved in this modified form of the term: "reference condition for biological integrity," or RC(BI). We further urge that these specific terms be used to refer to the concepts and methods used in individual bioassessments to characterize the expected condition to which current conditions are compared: "minimally disturbed condition" (MDC); "historical condition" (HC); "least disturbed condition" (LDC); and "best attainable condition" (BAC). We argue that each of these concepts can be narrowly defined, and each implies specific methods for estimating expectations. We also describe current methods by which these expectations are estimated including: the reference-site approach (condition at minimally or least-disturbed sites); best professional judgment; interpretation of historical condition; extrapolation of empirical models; and evaluation of ambient distributions. Because different assumptions about what constitutes reference condition will have important effects on the final classification of streams into condition classes, we urge that bioassessments be consistent in describing the definitions and methods used to set expectations.

Using relative risk to compare the effects of aquatic stressors at a regional scale.

The regional-scale importance of an aquatic stressor depends both on its regional extent (i.e., how widespread it is) and on the severity of its effects in ecosystems where it is found. Sample surveys, such as those developed by the U.S. Environmental Protection Agency's Environmental Monitoring and Assessment Program (EMAP), are designed to estimate and compare the extents, throughout a large region, of elevated conditions for various aquatic stressors. In this article, we propose relative risk as a complementary measure of the severity of each stressor's effect on a response variable that characterizes aquatic ecological condition. Specifically, relative risk measures the strength of association between stressor and response variables that can be classified as either "good" (i.e., reference) or "poor" (i.e., different from reference). We present formulae for estimating relative risk and its confidence interval, adapted for the unequal sample inclusion probabilities employed in EMAP surveys. For a recent EMAP survey of streams in five Mid-Atlantic states, we estimated the relative extents of eight stressors as well as their relative risks to aquatic macroinvertebrate assemblages, with assemblage condition measured by an index of biotic integrity (IBI). For example, a measure of excess sedimentation had a relative risk of 1.60 for macroinvertebrate IBI, with the meaning that poor IBI conditions were 1.6 times more likely to be found in streams having poor conditions of sedimentation than in streams having good sedimentation conditions. We show how stressor extent and relative risk estimates, viewed together, offer a compact and comprehensive assessment of the relative importances of multiple stressors.

Use of ecological regions in aquatic assessments of ecological condition.

Ecological regions are areas of similar climate, landform, soil, potential natural vegetation, hydrology, or other ecologically relevant variables. The makeup of aquatic biological assemblages (e.g., fish, macroinvertebrates, algae, riparian birds, etc.) varies dramatically over the landscape, as do the environmental stresses that affect the condition of those assemblages. Ecoregions delineate areas where similar assemblages are likely to occur and, therefore, where similar expectations can be established. For this reason, ecological regions have proven to be an important tool for use in the process of ecological assessment. This article describes four examples of the use of ecological regions in important aspects of environmental monitoring and assessment: (1) design of monitoring networks; (2) estimating expected conditions (criteria development); (3) reporting of results; (4) setting priorities for future monitoring and restoration. By delineating geographic areas with similar characteristics, ecological regions provide a framework for developing relevant indicators, setting expectations through the use of regional reference sites, establishing ecoregion-specific criteria and/or standards, presenting results, focusing models based on relationships between landscape and surface water metrics, and setting regional priorities for management and restoration. The Environmental Protection Agency and many state environmental departments currently use ecoregions to aid the development of environmental criteria, to illustrate current environmental condition, and to guide efforts to maintain and restore physical, chemical and biological integrity in lakes, streams, and rivers.

Development and evaluation of a Macroinvertebrate Biotic Integrity Index (MBII) for regionally assessing Mid-Atlantic Highlands Streams.

The Macroinvertebrate Biotic Integrity Index (MBII) was developed from data collected at 574 wadeable stream reaches in the Mid-Atlantic Highlands region (MAHR) by the U.S. Environmental Protection Agency's (USEPA) Environmental Monitoring and Assessment Program (EMAP). Over 100 candidate metrics were evaluated for range, precision, responsiveness to various disturbances, relationship to catchment area, and redundancy. Seven metrics were selected, representing taxa richness (Ephemeroptera richness, Plecoptera richness, Trichoptera richness), assemblage composition (percent non-insect individuals, percent 5 dominant taxa), pollution tolerance [Macroinvertebrate Tolerance Index (MTI)], and one functional feeding group (collector-filterer richness). We scored metrics and summed them, then ranked the resulting index through use of independently evaluated reference stream reaches. Although sites were classified into lowland and upland ecoregional groups, we did not need to develop separate scoring criteria for each ecoregional group. We were able to use the same metrics for pool and riffle composite samples, but we had to score them differently. Using the EMAP probability design, we inferred the results, with known confidence bounds, to the 167,797 kilometers of wadeable streams in the Mid-Atlantic Highlands. We classified 17% of the target stream length in the MAHR as good, 57% as fair, and 26% as poor. Pool-dominated reaches were relatively rare in the MAHR, and the usefulness of the MBII was more difficult to assess in these reaches. The process used for developing the MBII is widely applicable and resulted in an index effective in evaluating region-wide conditions and distinguishing good and impaired reaches among both upland and lowland streams dominated by riffle habitat.