Physical habitat in conterminous US streams and rivers, Part 1: Geoclimatic controls and anthropogenic alteration.

Anthropogenic alteration of physical habitat structure in streams and rivers is increasingly recognized as a major cause of impairment worldwide. As part of their assessment of the status and trends in the condition of rivers and streams in the U.S., the U.S. Environmental Protection Agency's (USEPA) National Aquatic Resource Surveys (NARS) quantify and monitor channel size and slope, substrate size and stability, instream habitat complexity and cover, riparian vegetation cover and structure, anthropogenic disturbance activities, and channel-riparian interaction. Like biological assemblages and water chemistry, physical habitat is strongly controlled by natural geoclimatic factors that can obscure or amplify the influence of human activities. We developed a systematic approach to estimate the deviation of observed river and stream physical habitat from that expected in least-disturbed reference conditions. We applied this approach to calculate indices of anthropogenic alteration of three aspects of physical habitat condition in the conterminous U.S. (CONUS): streambed sediment size and stability, riparian vegetation cover, and instream habitat complexity. The precision and responsiveness of these indices led the USEPA to use them to evaluate physical habitat condition in CONUS rivers and streams. The scores of these indices systematically decreased with greater anthropogenic disturbance at river and stream sites in the CONUS and within ecoregions, which we interpret as a response of these physical habitat indices to anthropogenic influences. Although anthropogenic activities negatively influenced all three physical habitat indices in the least-disturbed sites within most ecoregions, natural geoclimatic and geomorphic factors were the dominant influences. For sites over the full range of anthropogenic disturbance, analyses of observed/expected sediment characteristics showed augmented flood flows and basin and riparian agriculture to be the leading predictors of streambed instability and excess fine sediments. Similarly, basin and riparian agriculture and non-agricultural riparian land uses were the leading predictors of reduced riparian vegetation cover complexity in the CONUS and within ecoregions. In turn, these reductions in riparian vegetation cover and complexity, combined with reduced summer low flows, were the leading predictors of instream habitat simplification. We conclude that quantitative measures of physical habitat structure are useful and important indicators of the impacts of human activities on stream and river condition.

Physical habitat in conterminous US streams and Rivers, part 2: A quantitative assessment of habitat condition.

Rigorous assessments of the ecological condition of water resources and the effect of human activities on those waters require quantitative physical, chemical, and biological data. The U.S. Environmental Protection Agency's river and stream surveys quantify river and stream bed particle size and stability, instream habitat complexity and cover, riparian vegetation cover and structure, and anthropogenic disturbance activities. Physical habitat is strongly controlled by natural geoclimatic factors that co-vary with human activities. We expressed the anthropogenic alteration of physical habitat as O/E ratios of observed habitat metric values divided by values expected under least-disturbed reference conditions, where site-specific expected values vary given their geoclimatic and geomorphic context. We set criteria for good, fair, and poor condition based on the distribution of O/E values in regional least-disturbed reference sites. Poor conditions existed in 22-24% of the 1.2 million km of streams and rivers in the conterminous U.S. for riparian human disturbance, streambed sediment and riparian vegetation cover, versus 14% for instream habitat complexity. Based on the same four indicators, the percentage of stream length in poor condition within 9 separate U.S. ecoregions ranged from 4% to 42%. Associations of our physical habitat indices with anthropogenic pressures demonstrate the scope of anthropogenic habitat alteration; habitat condition was negatively related to the level of anthropogenic disturbance nationally and in nearly all ecoregions. Relative risk estimates showed that streams and rivers with poor sediment, riparian cover complexity, or instream habitat cover conditions were 1.4 to 2.6 times as likely to also have fish or macroinvertebrate assemblages in poor condition. Our physical habitat condition indicators help explain deviations in biological conditions from those observed among least-disturbed sites and inform management actions for rehabilitating impaired waters and mitigating further ecological degradation.

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.

Correspondence between a recreational fishery index and ecological condition for U.S.A. streams and rivers.

Sport fishing is an important recreational and economic activity, especially in Australia, Europe and North America, and the condition of sport fish populations is a key ecological indicator of water body condition for millions of anglers and the public. Despite its importance as an ecological indicator representing the status of sport fish populations, an index for measuring this ecosystem service has not been quantified by analyzing actual fish taxa, size and abundance data across the U.S.A. Therefore, we used game fish data collected from 1,561 stream and river sites located throughout the conterminous U.S.A. combined with specific fish species and size dollar weights to calculate site-specific recreational fishery index (RFI) scores. We then regressed those scores against 38 potential site-specific environmental predictor variables, as well as site-specific fish assemblage condition (multimetric index; MMI) scores based on entire fish assemblages, to determine the factors most associated with the RFI scores. We found weak correlations between RFI and MMI scores and weak to moderate correlations with environmental variables, which varied in importance with each of 9 ecoregions. We conclude that the RFI is a useful indicator of a stream ecosystem service, which should be of greater interest to the USA public and traditional fishery management agencies than are MMIs, which tend to be more useful for ecologists, environmentalists and environmental quality agencies.

Sampling Efforts for Estimating Fish Species Richness in Western USA River Sites.

Fish species richness is an indicator of river ecological condition but it is particularly difficult to estimate in large unwadeable rapidly flowing rivers. Intensive multi-gear sampling is time consuming, logistically complex and expensive. However, insufficient sampling effort underestimates species richness and yields inaccurate data about the ecological condition of river sites. We raft-electrofished 10 river sites in 10 different ecoregions and six western USA states for distances equal to 300 times their mean wetted channel widths (MCWs) to estimate the effort needed to approach asymptotes in fish species richness. To collect 90% of the observed fish species at the sites, we found that an average of 150 MCWs (ranging 80-210 MCWs) were needed, with the number of MCWs increasing in rivers with a higher proportion of spatially rare species. Frequently, the second or third additional 100 MCWs produced only one or two additional singletons or doubletons (species occurring only once or twice at a site). Before initiating sampling programs for adequately estimating species richness, we recommend assessing sampling effort, particularly if rare or uncommon species are expected or desired.

The Relation of Lotic Fish and Benthic Macroinvertebrate Condition Indices to Environmental Factors Across the Conterminous USA.

National and regional ecological assessments are essential for making rational decisions concerning water body conservation and management at those spatial extents. We analyzed data from 4597 samples collected from 3420 different sites across the conterminous USA during the U.S. Environmental Protection Agency's 2008-2009 and 2013-2014 National Rivers and Streams Assessment. We evaluated the relationship between both fish and macroinvertebrate multimetric index (MMI) condition scores and 38 environmental factors to assess the relative importance of natural versus anthropogenic predictors, contrast site-scale versus watershed-scale predictors, and examine ecoregional and assemblage differences. We found that most of the environmental factors we examined were related to either fish and/or macroinvertebrate MMI scores in some fashion and that the factors involved, and strength of the relationship, varied by ecoregion and between assemblages. Factors more associated with natural conditions were usually less important in explaining MMI scores than factors more directly associated with anthropogenic disturbances. Local site-scale factors explained more variation than watershed-scale factors. Random forest and multiple regression models performed similarly, and the fish MMI-environment relationships were stronger than macroinvertebrate MMI-environment relationships. Among ecoregions, the strongest environmental relationships were observed in the Northern Appalachians and the weakest in the Southern Plains. The fish and macroinvertebrate MMIs were only weakly correlated with each other, and they generally responded more strongly to different groups of variables. These results support the use of multiple assemblages and the sampling of multiple environmental indicators in ecological assessments across large spatial extents.

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.

Patterns in and predictors of stream and river macroinvertebrate genera and fish species richness across the conterminous USA.

Both native and non-native taxa richness patterns are useful for evaluating areas of greatest conservation concern. To determine those patterns, we analyzed fish and macroinvertebrate taxa richness data obtained at 3475 sites collected by the USEPA's National Rivers and Streams Assessment. We also determined which natural and anthropogenic variables best explained patterns in regional richness. Macroinvertebrate and fish richness increased with the number of sites sampled per region. Therefore, we determined residual taxa richness from the deviation of observed richness from predicted richness given the number of sites per region. Regional richness markedly exceeded average site richness for both macroinvertebrates and fish. Predictors of macroinvertebrate-genus and fish-species residual-regional richness differed. Air temperature was an important predictor in both cases but was positive for fish and negative for macroinvertebrates. Both natural and land use variables were significant predictors of regional richness. This study is the first to determine mean site and regional richness of both fish and aquatic macroinvertebrates across the conterminous USA, and the key anthropogenic drivers of regional richness. Thus, it offers important insights into regional USA biodiversity hotspots.

How might selenium moderate the toxic effects of mercury in stream fish of the western U.S.?

The ability of selenium (Se) to moderate mercury (Hg) toxicity is well established in the literature. Mercury exposures that might otherwise produce toxic effects are counteracted by Se, particularly when Se:Hg molar ratios approach or exceed 1. We analyzed whole body Se and Hg concentrations in 468 fish representing 40 species from 137 sites across 12 western U.S. states. The fish samples were evaluated relative to a published wildlife protective Hg threshold (0.1 sg Hg x g(-1) wet wt.), the currenttissue based methylmercury (MeHg) water quality criterion (WQC) for the protection of humans (0.3 microg Hg x g(-1) wet wt) and to presumed protections against Hg toxicity when Se:Hg molar ratios are >1. A large proportion (56%) of our total fish sample exceeded the wildlife Hg threshold, whereas a smaller, but significant proportion (12%), exceeded the MeHg WQC. However, 97.5% of the total fish sample contained more Se than Hg (molar ratio >1) leaving only 2.5% with Se: Hg ratios <1. All but one of the fish with Se:Hg <1, were of the genus Ptychochelius (pikeminnow). Scientific literature on Se counteracting Hg toxicity and our finding that 97.5% of the freshwater fish in our survey have sufficient Se to potentially protect them and their consumers against Hg toxicity suggests that Se in fish tissue (Se:Hg molar ratio) must be considered when assessing the potential toxic effects of Hg.

Mercury concentration in frozen whole-fish homogenates is insensitive to holding time.

Current recommended holding times for the analysis of total mercury (Hg) in fish tissue ranges from 28 to 180 days. In 2006, we evaluated the effect of an extended holding time on Hg concentrations by reanalyzing whole-fish wet homogenates that were analyzed originally in 2002 and had been stored frozen at -20 degrees C since that time. Seven species, 13-15 samples each, were reanalyzed. Comparisons of concentration differences between 2006 and 2002 indicated that no statistically significant changes in Hg concentrations occurred in any of the seven fish species. These results indicate that wet fish tissue homogenates can be held frozen for at least four years without affecting analytical results, thus extending holding times far beyond those currently recommended.

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.