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Objectives and Results

This project sought to develop an optimized macroinvertebrate IBI for perennial, wadeable streams of the NWGP ecoregion of South Dakota (Chapter 2). In addition, component metrics of the NGP and NWGP IBIs were compared and differences in IBI scores were evaluated among level IV ecoregions within each level III ecoregion. The NWGP and NGP IBIs were expected to significantly differ in over half the component metrics, and all metrics comprising each IBI differed. This indicated that the two ecoregions differ biologically, as identical methods were used to sample, process, and analyze the macroinvertebrate assemblages in each ecoregion. The two IBIs displayed a similar range of scores and successfully differentiated between impaired and non-

impaired sites. No significant differences in IBI scores could be detected among level IV ecoregions for either the NWGP or NGP.

This project also sought to identify the most influential statewide in-stream drivers of biotic integrity and evaluate the importance of regions in predicting in-stream driver values (Chapter 3). Drivers associated with cultivation were expected to be most influential in the NGP, while drivers related to rangeland and soil erosion were expected to be most influential in the NWGP. Regional variables were predicted to be important for determining in-stream habitat. Cluster analysis and NMDS results indicated distinct separation between NGP and NWGP macroinvertebrate assemblages. Nitrogen loading, human use, and pasture/hay were more positively correlated with NGP sites and riparian and watershed herbaceous cover and turbidity more positively correlated with NWGP sites. These results aligned with the hypothesized major drivers for each level III

ecoregion. Random forest modeling indicated that IBI scores were related to salinity, specific conductance, total riffle length, gravel and fine substrates, and stream discharge at the time of sampling. However, variation in each of the component IBI metrics was explained by a unique number and combination of drivers. Identified drivers of component metrics included measures of substrates, stream discharge, salinity, total length of riffles, specific conductance, nutrient concentrations, dissolved oxygen, turbidity, suspended solids, channel morphology, and bank vegetation. This knowledge may assist in causal analysis, by allowing managers to evaluate changes in individual metrics as well as overall IBI scores and determine which aspects of biotic integrity are impacted by impairment. Management and mitigation efforts can then be tailored more specifically to expected causes of impairment. Important drivers of in-stream habitat were nitrogen loading, human use, and herbaceous cover in the watershed and riparian zone. This falls in line with the land use gradient identified in the NMDS plot. Regional variables, especially level IV ecoregion and river basin, improved models of in-stream habitat and were also found to be associated with different responses in in-stream habitat.

Implications for Biological Assessments in South Dakota

The United States Environmental Protection Agency (US EPA) has encouraged the development of biological monitoring and assessment tools by state water resource agencies (USEPA 1987a). In addition, states have been encouraged to link the new biological tools to existing physical and chemical assessments (USEPA 1987b). In response, South Dakota developed a narrative statement to protect the biological integrity of waters, and has used beneficial use assessment to meet Clean Water Act requirements

and protect aquatic life uses (SDDENR 2016, USEPA 2017). However, no numeric biological criteria had been defined and the use of biological assemblages in impairment decisions was limited to the NGP ecoregion and the IBI developed by the previous study (Bertrand and Troelstrup 2013, USEPA 2017). As a result of this project, South Dakota state water resource managers have an expanded biological assessment toolbox with regional IBIs for perennial, wadeable streams across 80% of the state. Random forest modeling results also demonstrate that statewide biotic integrity is related to salinity and specific conductance, and that component IBI metrics are also related to total suspended solids, dissolved oxygen, and water temperature. These stream characteristics are

important stressors identified by the South Dakota DENR as part of beneficial use assessment for the state (SDDENR 2016). This project has nearly fulfilled US EPA recommendations by helping develop tools for state biological assessments and linking them to current physical and chemical assessments.

Another major implication of this study for water resource managers of South Dakota is the importance of regional stratification in monitoring and assessments, as shown by the analyses conducted in this study. Macroinvertebrate assemblages appear to differ by level III ecoregion as a result of the land use gradient that distinguishes the NGP from the NWGP. Level IV ecoregion and river basins were important correlates of in- stream habitat. For State water resource managers, this indicates that biological

assessments in South Dakota should be conducted separately between level III ecoregions and take into account level IV ecoregion and river basin differences in abiotic

disturbed condition should be developed in each level III ecoregion and used to re- evaluate differences between level IV ecoregions (Stoddard 2005, MPCA 2014).

This study also provides managers with baseline information and visualizations of expected interactions between measures of biotic integrity and in-stream habitat, and between in-stream habitat and watershed characteristics and regions in perennial,

wadeable streams of South Dakota. Managers now have a means of evaluating the current condition of these streams and detecting impairment, and a method to evaluate

differences in condition. Overall, this study supplies water resource managers in South Dakota with new tools to evaluate biotic integrity of stream systems and identify drivers and pathways of biological impairment. The NGP IBI is already in use by SD DENR for assessments and undergoing continual calibration (SDDENR 2016); the NWGP IBI developed here will likewise be adopted.

Implications for the North American Great Plains

The results of this study fit within the broader patterns and drivers of biotic integrity in the Great Plains. The finding of different component IBI metrics in individual ecoregions is consistent with work done in Minnesota and Wyoming (Hargett 2011, MPCA 2014). Individual IBI metrics were generally consistent with those found elsewhere in the central United States (Griffith et al. 2005, Weigel and Dimick 2011, Larsen 2013, MPCA 2014), although the combination of metrics remains unique to each ecoregion. This study also highlighted a major land use gradient driven by changes in herbaceous cover, human use, and nitrogen loading, which corresponded to a shift from rangeland/grassland to cultivated agriculture. These landscape drivers coincide with the

major types of land use/land cover in the Great Plains (Taylor, J.L. et al. 2015), which are in turn known to affect water quality and aquatic and terrestrial biodiversity (Allan 2004, Foley et al. 2005, Lenhart et al. 2011, Newbold et al. 2015, Tanaka et al. 2016).

Results of this effort contribute to assessment and management elsewhere in the Great Plains. Data collected in this study improve general knowledge of prairie stream macroinvertebrate assemblages and the response of biotic and abiotic aspects stream components to natural and anthropogenic pressures, which are areas of concern for water resource managers (Matthews 1988, Mullen et al. 2011, Roberts et al. 2016). A total of 222 taxa representing 201 genera and 80 families were identified in this study. Voucher specimens have been submitted to the South Dakota Aquatic Invertebrate Collection housed at South Dakota State University to supplement sparse records west of the Missouri River. The importance of ecoregions and river basins as drivers of biotic integrity and in-stream habitat may reflect the natural gradients in geology, precipitation, and temperature that define the Great Plains (Trimble 1980, Ojima et al. 2012, Basara et al. 2013), which in turn result in patterns of hydrology, turbidity, substrates, land use, and other habitat characteristics that determine the structure and composition of biotic

assemblages (Matthews 1988, Dodds et al. 2004, Phillips et al. 2016). The present study indicates the need for regional stratification of biological assessment efforts in light of these patterns; this should be evaluated in other Great Plains states where the importance of stratification may not have been tested. The methods used in this study are capable of identifying major drivers of biotic integrity and their relationships to in-stream habitat in environmentally harsh prairie streams and may be applied by other water resource managers in the Great Plains.

Additional Considerations for Future Work

Additional work is needed to ensure that the full range of stream conditions in South Dakota are being evaluated. This includes extending IBI development to other ecoregions in the State, such as the Northwestern Glaciated Plains and the Middle

Rockies of the Black Hills. Additionally, other abiotic variables and possible interactions should be evaluated. Natural gradients like precipitation, elevation, or geology might be incorporated into analyses of landscape drivers to more thoroughly evaluate underlying drivers of in-stream habitat. For instance, direct measurements of geology may improve substrate models. Additional anthropogenic variables should also be measured, such as pesticides or other toxins, as these are known to have effects on freshwater invertebrate biodiversity (Beketov et al. 2013).

Finally, the tools developed here for macroinvertebrate assemblages should be applicable to other biological assemblages. Fish IBIs have been developed for the NGP (Krause et al. 2013) and the NWGP (Kaiser 2017), and statewide driver analysis can be conducted using the same methodology as for the macroinvertebrates. Several studies in stream and river systems across Iowa (Quist and Schultz 2014), Michigan (Lammert and Allan 1999), Colorado (Griffith et al. 2005), New York (Johnson and Ringler 2014), and the Southeast United States (Paller et al. 2017) have found that separate biological assemblages are influenced by different environmental drivers and respond differently to disturbance. These findings have also been reported globally in Korea (Bae et al. 2014), New Zealand (Clapcott et al. 2012), Germany and Austria (Dahm et al. 2013), France (Marzin et al. 2013), and Brazil (Tanaka et al. 2016). Taken together, multiple

assemblages may provide a more holistic evaluation of biotic integrity than can be achieved by examining only one assemblage.

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