Defining a Disturbance Gradient in a Middle-Eastern River Basin.

A physical, chemical and biological characterization of river systems is needed to evaluate their ecological quality and support restoration programs. Herein, we describe an approach using water chemistry, physical structure and land use for identification of a disturbance gradient existing in the Karun River Basin. For this purpose, at each site, physical structure and physico-chemical data were collected once in each season for a total of 4 samples during the period (October 2018 - September 2019). Principal components analysis (PCA) of 17 variables identified five variables that were influential across all seasons: conductivity, total habitat score, stream morphology, clay & silt, and sand. Of the 54 sites, 14, 26 and 14 sites were classified as least, moderate and most disturbed sites, respectively. The metric Ephemeroptera, Plecoptera and Trichoptera (EPT) taxa was used for validation of the classification. Results in different seasons showed that all the least disturbed sites (n=14) were significantly different from moderate and most disturbed sites (p < 0.01). In this study the validation process presented a good confirmation of a priori reference sites selection process, showing that the proposed criteria could be considered as appropriate tools for characterization of the existent disturbance gradient in the Karun River Basin.

Applying the index of watershed integrity to the Matanuska-Susitna basin.

The Matanuska-Susitna Borough is the fastest growing region in the State of Alaska and is impacted by a number of human activities. We conducted a multiscale assessment of the stressors facing the borough by developing and mapping the Index of Watershed Integrity (IWI) and Index of Catchment Integrity (the latter considers stressors in areas surrounding individual stream segments exclusive of upstream areas). The assessment coincided with the borough's stormwater management planning. We adapted the list of anthropogenic stressors used in the original conterminous United States IWI application to reflect the borough's geography, human activity, and data availability. This analysis also represents an early application of the NHDPlus High Resolution geospatial framework and the first use of the framework in an IWI study. We also explored how remediation of one important stressor, culverts, could impact watershed integrity at the catchment and watershed scales. Overall, we found that the integrity scores for the Matanuska-Susitna basin were high compared to the conterminous United States. Low integrity scores did occur in the rapidly developing Wasilla-Palmer core area. We also found that culvert remediation had a larger proportional impact in catchments with fewer stressors.

Adapting the Index of Watershed Integrity for Watershed Managers in the Western Balkans Region.

Sustainable development supports watershed processes and functions. To aid the sustainable development of the western Balkans' transboundary river and lake basins, the Regional Environmental Center for Central and Eastern Europe and the US Environmental Protection Agency (EPA) adapted the EPA's Index of Watershed Integrity (IWI) following the devasting 2014 floods in Albania, Bosnia and Herzegovina, Kosovo, North Macedonia, Montenegro, and Serbia. The IWI evaluates six watershed functions based on a suite of anthropogenic stressors (e.g., impervious surfaces, reservoirs). A key feature of the IWI is its ability to accumulate the impact of upstream activities of any specific location in a river network. A novel feature of the IWI, compared with other watershed assessment tools, is its capacity to provide actionable information at the local scale. IWI scores-ranging from 0 (low integrity) to 1 (high integrity)-calculated for the 1084 catchments of the study area indicated highest integrity in the Alpine geographic region (mean = 0.55, standard deviation (SD) = 0.11) and intermediate to lowest integrity within the Mediterranean (mean = 0.49, SD = 0.12) and Continental (mean = 0.40, SD = 0.10) geographic regions. The IWI results are presented hierarchically for data analysts (stressor, functional component, Index of Catchment Integrity and IWI), ecologists (stream/catchment, watershed, basin), and managers (local, national, international). We provide real-world examples for managers, and suggestions for improving the assessment.

Factors influencing social demands of aquatic ecosystems.

Aquatic ecosystems provide services essential to human health and economies. Therefore, resource management programs aim to ensure the sustainable flow of these services. Stakeholder engagement is often a critical tool in learning what services are of priority to the public and may be integral to the success of aquatic ecosystem management because public participation in planning and decision making can generate broader support, e.g., financial, intellectual, and labor, for the management plan. The collection of such information may even be statutorily mandated, such as in the Clean Water Act of the United States, which requires that water bodies be classified for the beneficial uses, e.g., fisheries, drinking water, or recreation, they provide. Past evaluations of stakeholder engagement with aquatic ecosystems have considered a wide range of factors influencing engagement. We conducted a critical review of the literature on characteristics of stakeholders and characteristics of the environment that influence stakeholder engagement and participation with aquatic ecosystems. Our objective was to identify factors that should be considered in the creation of surveys to help encourage the inclusion of ecological and social beneficial uses data in large-scale water monitoring programs. Factors identified in our review were, extent and influence of place-based knowledge; proximity to, and frequency of visitation of the resource(s) being considered; basic demographics such as age, gender, education, and income; home community type; aesthetic appeal of the resource; and primary reason for engagement with the resource. We propose these factors, with subfactors, as a template for survey development.

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.

Characterization of River Networks: A GIS Approach and Its Applications.

Fluvial geomorphology provides the basis for characterizing complex river networks and evaluating biophysical processes within watersheds. Understanding the spatial organization of morphological features, their influencing processes, and resultant geomorphic diversity in stream networks are important for efficient restoration, river health assessment, and improving our knowledge of the resilience of riverine landscapes. River characterization is a means to determine the biophysical character of river networks but many methods are fraught with pitfalls, such as the use of incorrect variables and limited acknowledgment of the hierarchical organization of rivers. In this paper, a top-down geographic information system-based approach for determining the physical typology of river networks is outlined. A suite of multivariate analyses are used to develop a nomenclature for functional process zones (FPZs) - large tracts of the river network with similar hydro-geomorphological character. Applied to the Little Miami River, Ohio, six distinct FPZs emerged, which had a nonuniform distribution along the river network. Some FPZs repeated downstream; others were rare in terms of total length and number of FPZ segments. The physical structure of the Little Miami River network was analyzed using a series of community metrics. Application of this approach for river monitoring, establishing reference conditions, as well as management of threatened and endangered species and asset trading is highlighted.

Benthic macroinvertebrate field sampling effort required to produce a sample adequate for the assessment of rivers and streams of Neuquén Province, Argentina.

This multi-year pilot study evaluated a proposed field method for its effectiveness in the collection of a benthic macroinvertebrate sample adequate for use in the condition assessment of streams and rivers in the Neuquén Province, Argentina. A total of 13 sites, distributed across three rivers, were sampled. At each site, benthic macroinvertebrates were collected at 11 transects. Each sample was processed independently in the field and laboratory. Based on a literature review and resource considerations, the collection of 300 organisms (minimum) at each site was determined to be necessary to support a robust condition assessment, and therefore, selected as the criterion for judging the adequacy of the method. This targeted number of organisms was collected at all sites, at a minimum, when collections from all 11 transects were combined. Subsequent bootstrapping analysis of data was used to estimate whether collecting at fewer transects would reach the minimum target number of organisms for all sites. In a subset of sites, the total number of organisms frequently fell below the target when fewer than 11 transects collections were combined.Site conditions where < 300 organisms might be collected are discussed. These preliminary results suggest that the proposed field method results in a sample that is adequate for robust condition assessment of the rivers and streams of interest. When data become available from a broader range of sites, the adequacy of the field method should be reassessed.

Evaluation of an alternate method for sampling benthic macroinvertebrates in low-gradient streams sampled as part of the National Rivers and Streams Assessment.

Benthic macroinvertebrates are sampled in streams and rivers as one of the assessment elements of the US Environmental Protection Agency's National Rivers and Streams Assessment. In a 2006 report, the recommendation was made that different yet comparable methods be evaluated for different types of streams (e.g., low gradient vs. high gradient). Consequently, a research element was added to the 2008-2009 National Rivers and Streams Assessment to conduct a side-by-side comparison of the standard macroinvertebrate sampling method with an alternate method specifically designed for low-gradient wadeable streams and rivers that focused more on stream edge habitat. Samples were collected using each method at 525 sites in five of nine aggregate ecoregions located in the conterminous USA. Methods were compared using the benthic macroinvertebrate multimetric index developed for the 2006 Wadeable Streams Assessment. Statistical analysis did not reveal any trends that would suggest the overall assessment of low-gradient streams on a regional or national scale would change if the alternate method was used rather than the standard sampling method, regardless of the gradient cutoff used to define low-gradient streams. Based on these results, the National Rivers and Streams Survey should continue to use the standard field method for sampling all streams.

Environmental impact assessment and conservation planning of a Middle-Eastern River basin using a fish-based tolerance index.

The tolerance of aquatic organisms to stressors is widely used to monitor and evaluate the condition of freshwater ecosystems. Tolerance values (TV) derived from analyses of the relationship between species and their environment are considered to be more objective than those that rely on expert opinion. We used principal component analysis (PCA) to derive a generalized stressor gradient based on physicochemical characteristics and physical habitat quality and structure. Scores of the first principal component axis (PC1) were used to estimate TV for 37 fish species collected from 54 sites in the Karun River basin, Iran. PCA of 17 variables identified stressors that were influential such as total phosphate, total nitrogen, total coliform, and habitat and morphological score. The species were separated into three categories on the stressor gradient: sensitive (18.9%), semi-tolerant (48.6%), and tolerant species (32.4%). Based on these results we developed the Karun Fish Tolerance Index (KFTI) and demonstrated that it performed well in separating the least, moderate, and most disturbed sites in the study area. The discrimination efficiency of the KFTI was 82.5%, which makes it a robust management tool for the protection and conservation of streams and rivers in the Karun River watershed. TV developed here reflect objective characteristics of the sensitivity of fish species to the predominant stressors in the Karun and similar systems.

Improving ecosystem health in highly altered river basins: a generalized framework and its application to the Mississippi-Atchafalaya River Basin.

Continued large-scale public investment in declining ecosystems depends on demonstrations of "success". While the public conception of "success" often focuses on restoration to a pre-disturbance condition, the scientific community is more likely to measure success in terms of improved ecosystem health. Using a combination of literature review, workshops and expert solicitation we propose a generalized framework to improve ecosystem health in highly altered river basins by reducing ecosystem stressors, enhancing ecosystem processes and increasing ecosystem resilience. We illustrate the use of this framework in the Mississippi-Atchafalaya River Basin (MARB) of the central United States (U.S.), by (i) identifying key stressors related to human activities, and (ii) creating a conceptual ecosystem model relating those stressors to effects on ecosystem structure and processes. As a result of our analysis, we identify a set of landscape-level indicators of ecosystem health, emphasizing leading indicators of stressor removal (e.g., reduced anthropogenic nutrient inputs), increased ecosystem function (e.g., increased water storage in the landscape) and increased resilience (e.g., changes in the percentage of perennial vegetative cover). We suggest that by including these indicators, along with lagging indicators such as direct measurements of water quality, stakeholders will be better able to assess the effectiveness of management actions. For example, if both leading and lagging indicators show improvement over time, then management actions are on track to attain desired ecosystem condition. If, however, leading indicators are not improving or even declining, then fundamental challenges to ecosystem health remain to be addressed and failure to address these will ultimately lead to declines in lagging indicators such as water quality. Although our model and indicators are specific to the MARB, we believe that the generalized framework and the process of model and indicator development will be valuable in an array of altered river basins.

Critical Role for hierarchical geospatial analyses in the design of fluvial research, assessment, and management.

River science and management can be conducted at a range of spatiotemporal scales from reach to basin levels as long as the project goals and questions are matched correctly with the study design's spatiotemporal scales and dependent variables. These project goals should also incorporate information on the hydrogeomorphically patchy nature of riverine macrosystems which is only partially predictable in type and location from a river's headwaters to its terminus. This patchiness significantly affects a river's habitat template, and thus community structure, ecosystem function, and responses to perturbations. Our manuscript is designed for use by senior administrators at government agencies through entry-level river scientists. It analyzes common challenges in project design and recommends solutions based partially on hierarchical analyses that combine geographic information systems and multivariate statistical analysis to enable self-emergence of a stream's patchy structure. These approaches are useful at all spatial levels and can vary from primary reliance on geospatial techniques at the valley level to a greater dependence on field-based measurements and expert opinion at the reach level. Comparative uses of functional process zones (FPZs = valley-scale hydrogeomorphic patches), ecoregions, hydrologic unit codes, and reaches in project designs are discussed along with other comparative approaches for stream classification and analysis of species distributions (e.g., GAP analysis). Use of hierarchical classification of patch structure for sample stratification, reference site selection, ecosystem services, rehabilitation, and mitigation are briefly explored.

Automated riverine landscape characterization: GIS-based tools for watershed-scale research, assessment, and management.

River systems consist of hydrogeomorphic patches (HPs) that emerge at multiple spatiotemporal scales. Functional process zones (FPZs) are HPs that exist at the river valley scale and are important strata for framing whole-watershed research questions and management plans. Hierarchical classification procedures aid in HP identification by grouping sections of river based on their hydrogeomorphic character; however, collecting data required for such procedures with field-based methods is often impractical. We developed a set of GIS-based tools that facilitate rapid, low cost riverine landscape characterization and FPZ classification. Our tools, termed RESonate, consist of a custom toolbox designed for ESRI ArcGIS®. RESonate automatically extracts 13 hydrogeomorphic variables from readily available geospatial datasets and datasets derived from modeling procedures. An advanced 2D flood model, FLDPLN, designed for MATLAB® is used to determine valley morphology by systematically flooding river networks. When used in conjunction with other modeling procedures, RESonate and FLDPLN can assess the character of large river networks quickly and at very low costs. Here we describe tool and model functions in addition to their benefits, limitations, and applications.

Conservation of blackwater rivers and streams of the coastal plains of United States: Knowledge and research needs.

Blackwater rivers and streams are a distinct resource type in multiple ecoregions of the United States, including the Coastal Plains. Given the unique nature of blackwaters, they may not be fully protected if using typical water quality standards. Information provided by State agencies was used to identify specific research needs that would directly enhance current assessment approaches. Protection of blackwater rivers and streams would be aided by (1) development of an operational definition; (2) development of a classification framework that distinguishes among blackwater system types; (3) identification of undisturbed or minimally disturbed blackwater systems; (4) identification of stressors that challenge blackwater condition and resilience; and (5) development of criteria that are more suited to the unique conditions of blackwater rivers and streams. Benefits stemming from addressing these research needs may include identification of sites that are unrecognized as impaired and the re-evaluation of sites that may be erroneously listed as impaired when they are not. This would improve reporting by states for Clean Water Act (CWA) Section 305(b) and, for some states, could represent a significant resource savings by reducing the number of sites on CWA Section 303d lists.

Distribution and Expansion of Alien Fish Species in the Karun River Basin, Iran.

We assessed the distribution of alien fishes in the Karun River Basin, Iran. Fish were collected from 39 sites during the November-December 2018 low-flow period. In total, 39 fish species from nine orders and 14 families were documented. Among these, 10 species were alien to the basin (986 individuals; 15.7%). Four species were the most abundant alien species and primarily in impounded, downstream reaches. Redundancy analysis (RDA) was conducted to identify the extent of changes in alien fish assemblages with environmental parameters. RDA1 and RDA2 accounted for 36.24% and 25.33% of the variation of alien species, respectively. Altitude, depth, electrical conductivity, water temperature, turbidity, dissolved oxygen, and river width were the most significant parameters affecting alien species distributions. We present a dual-pathway cause-and-effect hypothesis proposing that alien fish species presence causes declines in the ecological status of native fish communities. We then explore how human-induced aquatic ecosystem degradation creates opportunities for alien species to invade new ecosystems, further impacting native fish communities. Our study contributes insight into the cause and effect of the presence of alien fish species in the Karun River Basin and emphasizes the urgency of conservation measures to protect this critically endangered watershed.

Fish Species Composition, Distribution and Community Structure in Relation to Environmental Variation in a Semi-Arid Mountainous River Basin, Iran.

We analyzed spatial variation in fish species richness and community composition in the Karun River basin, Iran. Knowledge about fish diversity in the basin is incomplete and varies widely along spatial and temporal scales: The Karun is the longest river in Iran (950 km) with the largest drainage area (about 67,000 km2). Fish samples were collected from 54 sites from July through August 2019 using a backpack electro-fisher. Physico-chemical and habitat parameter data collected at each site included pH, conductivity (µS/cm), dissolved oxygen (mg/L), water temperature (°C), turbidity (NTU), stream width (m), stream depth (m), water velocity (m/s) and elevation (m). In total, 37 species were collected (5241 individuals weighing 110.67 kg). The species collected represented 12 families and 27 genera. A total of 13 endemic species (35.14%), 16 native species (43.24%), and eight non-native species (21.62%) were recorded. Diversity indices were calculated and used to measure the spatial variation in community composition. Relationships between native and endemic species assemblage structure and environmental descriptors were assessed using canonical correspondence analysis (CCA). The first two axes of the canonical correspondence analysis explained 62.57% of the variation in the data. Of the nine environmental descriptors analyzed, eight significantly affected species distribution; however, electrical conductivity and elevation were most influential. Our study provides up-to-date status information on the distribution of freshwater fishes in the Karun River basin. This information is essential for developing conservation and management strategies to support the long-term sustainability of fish populations in the Karun River basin.

A fish-based multi-metric assessment index in the Karun River basin, Iran.

Large river systems are one of the most important water resources for human societies. However, the ecological integrity of large rivers has been altered greatly by human activities. To monitor and manage these ecosystems, multimetric indices (MMI) are considered as efficient tools. This study aimed to develop and validate a fish-based multimetric index for the Karun River basin, Iran (Karun fish-based multi-metric index [KFMMI]). Eighteen rivers and 54 sites in the basin were sampled in July-August 2019, and physico-chemical and habitat characteristics were used to identify reference sites based on principal components analysis (PCA). Of the 54 sites, 14, 26, and 14 sites were classified as least, moderate, and most disturbed sites, respectively. Fifty-four candidate metrics were evaluated for range, responsiveness, and redundancy with other metrics. This resulted in the identification of eight metrics (relative abundance of native and endemic taxa, relative richness of migratory taxa, relative richness of Leuciscidae taxa, relative richness of herbivorous taxa, relative abundance of cyprinid taxa, relative richness of vegetative inhabitant taxa, relative abundance of slow water flow, and relative richness of edge inhabitant taxa) that informed on species richness and composition, migratory status, functional feeding groups, and habitat preferences. The KFMMI showed excellent performance in separating least, moderate, and most disturbed sites in our study area. Regarding water quality, the KFMMI was classified 16, 5, and 29 sites as good, moderate, and bad, respectively. The discrimination efficiency of KFMMI was 81.6%, which makes it an effective management tool for directing restoration actions at most disturbed sites and intensifying protection of least disturbed sites.

Biological assessment of western USA sandy bottom rivers based on modeling historical and current fish and macroinvertebrate data.

Biological monitoring is important for assessing the ecological condition of surface waters. However, there are challenges in determining what constitutes reference conditions, what assemblages should be used as indicators, and how assemblage data should be converted into quantitative indicator scores. In this study, we developed and applied biological condition gradient (BCG) modeling to fish and macroinvertebrate data previously collected from large, sandy bottom southwestern USA rivers. Such rivers are particularly vulnerable to altered flow regimes resulting from dams, water withdrawals and climate change. We found that sensitive ubiquitous taxa for both fish and macroinvertebrates had been replaced by more tolerant taxa, but that the condition assessment ratings based on fish and macroinvertebrate assemblages differed. We conclude that the BCG models based on both macroinvertebrate and fish assemblage condition were useful for classifying the condition of southwestern USA sandy bottom rivers. However, our fish BCG model was slightly more sensitive than the macroinvertebrate model to anthropogenic disturbance, presumably because we had historical fish data, and because fish may be more sensitive to dams and altered flow regimes than are macroinvertebrates.

Factors influencing perceptions of aquatic ecosystems.

While factors influencing perceptions of drinking water have been well studied, those of aquatic ecosystems have been to lesser extent. We conducted a review to improve awareness of these factors. Environmental factors found to influence public perceptions of aquatic ecosystems were presence/absence of water plants and algae, presence/absence of floating debris, the odor, movement (for flowing waters) and clarity/turbidity of the water, and the type, condition, setting, naturalness, and overall aesthetic appeal of the ecosystem. Sociocultural factors found to influence public perceptions of aquatic ecosystems included age, education, gender, and place-based knowledge. We provide perspectives of how managers can better meet the diverse social demands placed on aquatic ecosystems. The importance and benefits of considering these perspectives may be especially beneficial where significant multi-generational and culturally relevant place-based knowledge exist.

Understanding rivers and their social relations: A critical step to advance environmental water management.

River flows connect people, places, and other forms of life, inspiring and sustaining diverse cultural beliefs, values, and ways of life. The concept of environmental flows provides a framework for improving understanding of relationships between river flows and people, and for supporting those that are mutually beneficial. Nevertheless, most approaches to determining environmental flows remain grounded in the biophysical sciences. The newly revised Brisbane Declaration and Global Action Agenda on Environmental Flows (2018) represents a new phase in environmental flow science and an opportunity to better consider the co-constitution of river flows, ecosystems, and society, and to more explicitly incorporate these relationships into river management. We synthesize understanding of relationships between people and rivers as conceived under the renewed definition of environmental flows. We present case studies from Honduras, India, Canada, New Zealand, and Australia that illustrate multidisciplinary, collaborative efforts where recognizing and meeting diverse flow needs of human populations was central to establishing environmental flow recommendations. We also review a small body of literature to highlight examples of the diversity and interdependencies of human-flow relationships-such as the linkages between river flow and human well-being, spiritual needs, cultural identity, and sense of place-that are typically overlooked when environmental flows are assessed and negotiated. Finally, we call for scientists and water managers to recognize the diversity of ways of knowing, relating to, and utilizing rivers, and to place this recognition at the center of future environmental flow assessments. This article is categorized under: Water and Life > Conservation, Management, and Awareness Human Water > Water Governance Human Water > Water as Imagined and Represented.

A patchy continuum? Stream processes show varied responses to patch- and continuum-based analyses.

Many conceptual syntheses in ecology and evolution are undergirded by either a patch- or continuum-based model. Examples include gradualism and punctuated equilibrium in evolution, and edge effects and the theory of island biogeography in ecology. In this study, we sought to determine how patch- or continuum-based analyses could explain variation in concentrations of stream macronutrients and system metabolism, represented by measures of productivity and respiration rates, at the watershed scale across the Kanawha River Basin, USA. Using Strahler stream order (SSO; continuum) and functional process zone (FPZ; patch) as factors, we produced statistical models for each variable and compared model performance using likelihood ratio tests. Only one nutrient (i.e., PO43- ) responded better to patch-based analysis. Both models were significantly better than a null model for ecosystem respiration; however, neither outperformed the other. Importantly, in most cases, a combination model, including both SSO and FPZ, best described observed variation in the system. Our findings suggest that several patch- and continuum-based processes may simultaneously influence the concentration of macronutrients and system metabolism. Nutrient spiral- ing along a continuum and the patch mosaic of land cover may both alter macronutrients, for example. Similarly, increases in temperature and discharge associated with increasing SSO, as well as the differences in light availability and channel morphology associated with different FPZs, may influence system metabolism. For these reasons, we recommend a combination of patch- and continuum-based analyses when modeling, analyzing, and interpreting patterns in stream ecosystem parameters.