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The degree of coupling between the social and ecological components of social-ecological systems is seen as fundamental to understanding their functioning, interactions and trajectories. Yet, there is limited work about how to empirically understand the degree of coupling between social and ecological systems, nor the processes by which the degree of coupling could change over time. Here, we introduce a conceptual framework for characterizing trajectories over time of coupling and de-coupling in social-ecological river systems. We analyze two conceptual scenarios describing coupling and de-coupling trajectories in a social-ecological system and define a series of key concepts for understanding social-ecological system trajectories. We tested these coupling and de-coupling trajectories theory by linking these concepts to empirical case examples of two river social-ecological systems in the western United States. Finally, we propose a quantitative approach with the potential for evaluating the level of social-ecological coupling and de-coupling trajectories in other SES contexts. This paper represents an advancing on the identification of specific actions that explain current SES trajectories and immediate actions to reinforce or shift the trajectory.

Stream ecosystem metabolism contributes to global carbon cycling, yet predicting metabolism across ecosystems remains elusive. Even within stream segments, spatial variation in metabolic rates and their controls can be substantial, exhibiting sudden rather than continuous changes. We measured metabolism at 6 sites along a 75-km mainstem segment of Marsh Creek, Idaho. This agricultural stream lacks major geomorphic transitions such as tributaries or changes in valley width, but possesses patchy patterns of turbidity. We asked: (1) How variable is metabolism along this segment, and (2) How do the controls on metabolism vary along this segment? Metabolism varied several-fold along this stream segment. Average rates of gross primary production (GPP), ecosystem respiration (ER), and net ecosystem production (NEP) among sites did not correlate with water quality or aquatic macrophyte cover. Rather, reaches along this segment appear to represent different process domains that were characterized by turbidity: More turbid reaches saw negative effects of turbidity on GPP and NEP and positive effects on ER. Turbidity was associated with increased respiration and decoupled ER from GPP at turbid sites. Less turbid sites had stronger coupling between GPP and ER, and GPP was predicted by light and temperature. Heterogeneous sediment supply and transport capacity along Marsh Creek create patchy patterns of turbidity, which affects the controls on metabolism. Geomorphic controls on ecosystem processes are complex but not random. Understanding how metabolism and its controls vary across process domains is needed to scale up process rates and understand how process rates might respond to future changes.

Microbes play a critical role in plant litter decomposition and influence the fate of carbon in rivers and riparian zones. When decomposing low-nutrient plant litter, microbes acquire nitrogen (N) and phosphorus (P) from the environment (i.e., nutrient immobilization), and this process is potentially sensitive to nutrient loading and changing climate. Nonetheless, environmental controls on immobilization are poorly understood because rates are also influenced by plant litter chemistry, which is coupled to the same environmental factors. Here we used a standardized, low-nutrient organic matter substrate (cotton strips) to quantify nutrient immobilization at 100 paired stream and riparian sites representing 11 biomes worldwide. Immobilization rates varied by three orders of magnitude, were greater in rivers than riparian zones, and were strongly correlated to decomposition rates. In rivers, P immobilization rates were controlled by surface water phosphate concentrations, but N immobilization rates were not related to inorganic N. The N:P of immobilized nutrients was tightly constrained to a molar ratio of 10:1 despite wide variation in surface water N:P. Immobilization rates were temperature-dependent in riparian zones but not related to temperature in rivers. However, in rivers nutrient supply ultimately controlled whether microbes could achieve the maximum expected decomposition rate at a given temperature. Collectively, we demonstrated that exogenous nutrient supply and immobilization are critical control points for decomposition of organic matter. Key Points • Nitrogen (N) and phosphorus (P) immobilization was measured on organic matter (cotton) in 100 rivers and riparian zones representing 11 biomes • Elevated temperature in riparian zones and phosphate in rivers increased immobilization, and consequently accelerated decomposition • N and P immobilization was strongly linked by microbial stoichiometry despite widely varied surface-water nutrient ratios Plain Language Summary Bacteria and fungi contribute to the breakdown of leaf litter in rivers and floodplains. To break down leaf litter, these microbes need the nutrients nitrogen and phosphorus (P), and microbes can get nutrients either from the leaf litter itself or from the environment. Most leaf litter has low nutrient content and microbes must rely on the environment to supply nutrients. We studied microbial nutrient uptake from the environment during litter breakdown to determine whether it varies predictably across the globe and how it is influenced by changing climate and nutrient pollution. In 100 rivers and floodplains in 11 of Earth's major biomes we placed small strips of cotton as stand-ins for leaf litter. Nutrient uptake was consistently greater on cotton strips that were submerged in the river compared to cotton on the floodplain. For microbes in the river, nutrient uptake was faster in instances where there was more P in the water. For microbes in the floodplain, nutrient uptake was faster where temperatures were warmer. Faster nutrient uptake by microbes was linked with faster cotton breakdown in rivers and floodplains. Our study shows that climate change and nutrient pollution can alter the activity of microbes in rivers and floodplains.

Landscape perspectives in riverine ecology have been undertaken increasingly in the last 30 years, leading aquatic ecologists to develop a diverse set of approaches for conceptualizing, mapping and understanding ‘riverscapes’. Spatiotemporally explicit perspectives of rivers and their biota nested within the socio-ecological landscape now provide guiding principles and approaches in inland fisheries and watershed management. During the last two decades, scientific literature on riverscapes has increased rapidly, indicating that the term and associated approaches are serving an important purpose in freshwater science and management. We trace the origins and theoretical foundations of riverscape perspectives and approaches and examine trends in the published literature to assess the state of the science and demonstrate how they are being applied to address recent challenges in the management of riverine ecosystems. We focus on approaches for studying and visualizing rivers and streams with remote sensing, modelling and sampling designs that enable pattern detection as seen from above (e.g. river channel, floodplain, and riparian areas) but also into the water itself (e.g. aquatic organisms and the aqueous environment). Key concepts from landscape ecology that are central to riverscape approaches are heterogeneity, scale (resolution, extent and scope) and connectivity (structural and functional), which underpin spatial and temporal aspects of study design, data collection and analysis. Mapping of physical and biological characteristics of rivers and floodplains with high-resolution, spatially intensive techniques improves understanding of the causes and ecological consequences of spatial patterns at multiple scales. This information is crucial for managing river ecosystems, especially for the successful implementation of conservation, restoration and monitoring programs. Recent advances in remote sensing, field-sampling approaches and geospatial technology are making it increasingly feasible to collect high-resolution data over larger scales in space and time. We highlight challenges and opportunities and discuss future avenues of research with emerging tools that can potentially help to overcome obstacles to collecting, analysing and displaying these data. This synthesis is intended to help researchers and resource managers understand and apply these concepts and approaches to address real-world problems in freshwater management.

There is increasing evidence of environmental change impacts on freshwater ecosystem services especially through land use and climate change. However, little is known about how land conservation could help mitigate adverse water-sustainability impacts. In this paper, we utilized the InVEST tool and the Residual Trends method to assess the joint effects and relative contributions of climate change and land conservation on freshwater ecosystem services in the Portneuf River catchment in Idaho, USA. We developed five hypothesized scenarios regarding gain and loss in the enrollment of Conservation Reserve Program (CRP), the largest agricultural land-retirement program in the U.S., plus riparian buffer and assessed their interactions with climate change. Results suggest that the realized water yield in the Portneuf River catchment would possibly be 56% less due to climate change and 24% less due to the decline of CRP enrollment. On the contrary, if CRP enrollment is promoted by ~30% and riparian buffer protection is implemented, the water supply reduction in the year 2050 could be changed from 56% to 26%, the total phosphorus (TP) and total nitrogen (TN) export would be reduced by 10% and 11%, and the total suspended sediment (TSS) reduced by 17%. This study suggests that increasing implementation of the CRP would likely preserve key freshwater ecosystem services and assist proactive mitigation, especially for semiarid regions vulnerable to changing climate conditions.

Stream drying and wildfire are projected to increase with climate change in the western United States, and both are likely to impact stream chemistry patterns and processes. To investigate drying and wildfire effects on stream chemistry (carbon, nutrients, anions, cations, and isotopes), we examined seasonal drying in two intermittent streams in southwestern Idaho, one stream that was unburned and one that burned 8 months prior to our study period. During the seasonal recession following snowmelt, we hypothesized that spatiotemporal patterns of stream chemistry would change due to increased evaporation, groundwater dominance, and autochthonous carbon production. With increased nutrients and reduced canopy cover, we expected greater shifts in the burned stream. To capture spatial chemistry patterns, we sampled surface water for a suite of analytes along the length of each stream with a high spatial scope (50-m sampling along ~2,500 m). To capture temporal variation, we sampled each stream in April (higher flow), May, and June (lower flow) in 2016. Seasonal patterns and processes influencing stream chemistry were generally similar in both streams, but some were amplified in the burned stream. Mean dissolved inorganic carbon (DIC) concentrations increased with drying by 22% in the unburned and by 300% in the burned stream. In contrast, mean total nitrogen (TN) concentrations decreased in both streams, with a 16% TN decrease in the unburned stream and a 500% TN decrease (mostly nitrate) in the burned stream. Contrary to expectations, dissolved organic carbon (DOC) concentrations varied more in space than in time. In addition, we found the streams did not become more evaporative relative to the Local Meteoric Water Line (LMWL) and we found weak evidence for evapoconcentration with drying. However, consistent with our expectations, strontium-DIC ratios indicated stream water shifted toward groundwater-dominance, especially in the burned stream. Fluorescence and absorbance measurements showed considerable spatial variation in DOC sourcing each month in both streams, and mean values suggested a temporal shift from allochthonous toward autochthonous carbon sources in the burned stream. Our findings suggest that the effects of fire may magnify some chemistry patterns but not the biophysical controls that we tested with stream drying.

Networks of direct and indirect biotic interactions underpin the complex dynamics and stability of ecological systems, yet experimental and theoretical studies often yield conflicting evidence regarding the direction (positive or negative) or magnitude of these interactions. We revisited pioneering data sets collected at the deciduous forested Horonai Stream and conducted ecosystem-level syntheses to demonstrate that the direction of direct and indirect interactions can change depending on the timescale of observation. Prior experimental studies showed that terrestrial prey that enter the stream from the adjacent forest caused positive indirect effects on aquatic invertebrates during summer by diverting fish consumption. Seasonal and annual estimates of secondary production and organic matter flows along food web pathways demonstrate that this seasonal input of terrestrial invertebrate prey increases production of certain fish species, reversing the indirect effect on aquatic invertebrates from positive at the seasonal timescale to negative at the annual timescale. Even though terrestrial invertebrate prey contributed 54% of the annual organic matter flux to fishes, primarily during summer, fish still consumed 98% of the aquatic invertebrate annual production, leading to top-down control that is not revealed in short-term experiments and demonstrating that aquatic prey may be a limiting resource for fishes. Changes in the direction or magnitude of interactions may be a key factor creating nonlinear or stabilizing feedbacks in complex systems, and these dynamics can be revealed by merging experimental and comparative approaches at different scales.

Species traits and trophic interactions drive contaminant flux and fate in linked river-riparian food webs., Mercury (Hg) biomagnification in aquatic food webs is a global concern; yet, the ways species traits and interactions mediate these fluxes remain poorly understood. Few pathways dominated Hg flux in the Colorado River despite large spatial differences in food web complexity, and fluxes were mediated by one functional trait, predation resistance. New Zealand mudsnails are predator resistant and a trophic dead end for Hg in food webs we studied. Fishes preferred blackflies, which accounted for 56 to 80% of Hg flux to fishes, even where blackflies were rare. Food web properties, i.e., match/mismatch between insect production and fish consumption, governed amounts of Hg retained in the river versus exported to land. An experimental flood redistributed Hg fluxes in the simplified tailwater food web, but not in complex downstream food webs. Recognizing that species traits, species interactions, and disturbance mediate contaminant exposure can improve risk management of linked aquatic-terrestrial ecosystems.

It is frequently assumed that nitrogen (N2) fixation and denitrification do not co-occur in streams because each process should be favored under different concentrations of dissolved inorganic nitrogen (DIN), and therefore these processes are rarely quantified together. We asked if these processes could co-exist by conducting a spatial survey of N2 fixation using acetylene reduction and denitrification using acetylene block [with and without amendments of carbon (C) as glucose and nitrogen (N) as nitrate]. Rates were measured on rocks and sediment in 8 southeastern Idaho streams encompassing a DIN gradient of 26–615 µg L−1. Sampling at each site was repeated in summer 2015 and 2016. We found that both denitrification and N2 fixation occurred across the gradient of DIN concentrations, with N2 fixation occurring primarily on rocks and denitrification occurring in sediment. N2 fixation rates on rocks significantly decreased 100× across the DIN gradient in 1 year of the study, and amended (with N and C) denitrification rates increased 10× across the DIN gradient in both years. Multiple linear regression and partial least squares models with environmental characteristics measured at the scale of entire stream reaches showed that C and phosphorus were positive predictors of amended and unamended denitrification rates, but no significant model could explain N2 fixation rates across all streams and years. This, coupled with the observation that detectable rates of N2 fixation occurred primarily on rocks and denitrification occurred primarily on sediment, suggests that microhabitat scale factors may better predict the co-occurrence of these processes within stream reaches. Overlooking the potential co-occurrence of N2 fixation and denitrification in stream ecosystems will impede understanding by oversimplifying the contribution of each process to the N cycle.

Traditionally, exploration of ecosystems in the context of undergraduate education has been restricted to connections within conventionally defined habitats (i.e., within a stream, within a forest). Further, instruction regarding the aquatic-terrestrial interface has emphasized directional inputs from land to water. However, a relatively new body of research has characterized reciprocal interactions and draws attention to fluxes from water to land, including the emergence of aquatic insects that serve as prey for terrestrial predators. We present a guide to an inquiry-based lesson for undergraduate biology that explores interactions and connections across aquatic and terrestrial habitat boundaries. The focus is on cross-habitat linkages within ecosystems, specifically addressing the question, What is the role of insect emergence in connecting the web of life linking aquatic and terrestrial habitats and organisms? Students (1) engage with a documentary film, (2) explore insect emergence and make observations of riparian insectivores, (3) explain the collected data, (4) elaborate on alternative study designs and a measure of ecosystem health, and (5) evaluate their new understanding. This lesson addresses core concepts and competencies for undergraduate biology education, as identified in the Vision and Change report.

Pervasive environmental degradation has altered biodiversity at a global scale. At smaller scales, species extirpations, invasions, and replacements have greatly influenced how ecosystems function and interact by affecting the exchanges of energy, materials, and organisms. In this chapter, we examine how a variety of environmental stressors, and associated species losses and gains, change the exchange of resources (materials or organisms) within and among ecosystems. We specifically consider how changes that occur within an ecosystem may trigger effects that reverberate (e.g., directly, indirectly, and via feedbacks) back and forth across ecological boundaries and propagate to other ecosystems connected via exchanges of materials and organisms. Our synthesis provides cursory overviews of ecosystem “openness” as it has been addressed by community ecologists and the conceptual development of ecological frameworks used to examine resource exchanges between ecosystems. We then describe four case studies and examine how species losses and gains affect food web structure via resource exchanges between ecosystems, with particular emphasis on effects spanning land-water boundaries. Finally, we discuss the need for more complex conceptual treatment of the interconnectedness of food webs among ecosystems.

Identifying and understanding functional process domains (sensu Montgomery, 1999 ) in rivers is paramount for linking the physical habitat template to ecosystem structure and function. To date, efforts to do this have been rare, especially in large rivers, as they require appropriate tools for quantifying habitat heterogeneity with fine-scale resolution across broad spatial extents. In this study, we used side-scan sonar technology to map riverbed substrate at six sites in the Yellowstone and Missouri rivers. Substrate maps were then analyzed and visualized using geospatial analysis to relate fine-grained spatial substrate patterns to process domain structure. Our findings revealed two distinct nested domains of substrate patchiness, suggesting that different factors are responsible for shaping patterns of substrate at different scales. Although small-scale patchiness in substrate was likely driven by internal, or autogenic, physical processes, patterns at larger segment extents (>3 km) were often driven by abrupt transitions in habitat related to exogenous factors such as lateral erosion of talus, tributary inputs, and bank armoring. Additionally, we found that heterogeneity in benthic substrate increased with spatial extent at all of our study sites; however, this relationship was lower in the Missouri River, which is altered by impoundment. Our study represents one of the first efforts to relate benthic habitat heterogeneity to nested process domain structure in large riverscapes, and offers a unique perspective for linking landscape processes, geomorphological habitat heterogeneity, and biological structure and function in large rivers.

The River Continuum Concept (RCC) predicts that food webs (and, in particular, invertebrates) of rivers in temperate, forested drainages should exhibit a longitudinal gradient from reliance on terrestrially derived organic matter (e.g., seasonally shed leaves) in the headwaters to autochthonous sources (e.g., algae) in the mid-orders, to suspended material in larger rivers. This prediction has been evaluated by longitudinal comparisons of macroinvertebrate communities in terms of functional feeding groups (FFGs), but such an approach yields only indirect evidence regarding actual food use. To retest this prediction, we investigated invertebrate diets from the Salmon River, Idaho, by examining the gut contents of archived specimens from the longitudinal set of sites sampled in the original 1976 RCC study and by collecting invertebrates from these same sites in 2009. We detected no apparent shifts in diets over the ∼30-y time span. The importance of allochthonous materials in invertebrate diets diff...

Published by Oxford University Press on behalf of American Institute of Biological Sciences 2016. This work is written by (a) US Government employee(s) and is in the public domain in the US. The published article can be found at: http://bioscience.oxfordjournals.org/content/66/7/561

Constructed stormwater wetlands (CSWs) are used to address contaminants in urban stormwater such as nitrogen (N) and phosphorus (P), but their performance is variable. Ephemeral CSWs tend to be less effective than perennial CSWs at removing N and P. We asked: How does wetland vegetation and sediment affect nutrient cycling/release from sediment and vegetation in ephemeral CSWs? We focused on two ephemeral urban CSWs in Pocatello, ID, USA, one densely vegetated and the other nearly bare. We rewetted intact cores of dry wetland sediments and, separately, senesced vegetation for 1 week at the end of the summer dry period to assess whether wetland sediments and vegetation acted as sources or sinks of N and P. For both CSWs, there was a pulse of nutrients immediately following rewetting, but the magnitude of that pulse and subsequent changes in nutrient concentrations suggest different processes dominate at each wetland, driven by differences in wetland vegetation and associated sediment characteristics. There was evidence of denitrification between and during events at the vegetated wetland, but larger fluxes of P at this site suggests a tradeoff between denitrification and P release. While the experimental results suggested specific biogeochemical controls, CSW nutrient concentrations during three events were more dynamic and suggested more biogeochemical complexity than that represented in our experiment, both within events and seasonally. Ephemeral CSWs may create unique biogeochemical conditions and require careful design to ensure N and P retention. Managers will also need to consider whether perennial water sources would improve CSW function.

Foodweb analyses can provide insight as to how species persist in naturally dynamic environments and how these communities may respond to anthropogenic alteration. In the remaining undammed rivers of the arid southwestern United States, it is thought that plasticity in feeding behavior allows native fishes to thrive in the context of natural fluctuations in temperature, sediment, and resource availability associated with extreme swings in season and hydrology. There have been few evaluations of this hypothesis, in part owing to the rarity of rivers that retain their natural environmental dynamics and complement of native species. We investigated the feeding ecology of the fish assemblage of one such river, the Little Colorado River, Arizona, focusing on four native species: bluehead sucker (Catostomus discobolus), flannelmouth sucker (Catostomus latipinnis), speckled dace (Rhinichthys osculus), and endangered humpback chub (Gila cypha). Over two years, we sampled fish diets in July, September, January, March, and June, encompassing turbid and clear‐water conditions across seasons. We quantified food resource contributions to fish diets in fish‐centered food webs, comparing diversity of trophic linkages and linkage‐strength distribution across time. We used ordination analyses to investigate differences in diets among species and changes in diet composition across seasons and conducted seasonal comparisons of gut fullness. We observed a high diversity of dietary items (as absolute numbers and consumer–resource ratios) within a fish assemblage that was apparently resistant to extreme and recurrent environmental variability. Fish diet diversity was ~2× that of the nearby dammed Colorado River's tailwater, and over 50% of fish‐resource linkages were weak (

Facing the challenges of environmental and social changes, sustainable management of ecosystem services is a worldwide priority. The Programme on Ecosystem Change and Society (PECS) approach provides a unique opportunity for promoting transdisciplinary place-based comparative research for social-ecological systems (SES) management. As part of the PECS-sponsored WaterSES project, we used four place-based SES research sites to analyze patterns in perceptions of ecosystem services. Our data come from about 1,500 face-to-face surveys conducted in southern Spain, the south-central Great Plains of Oklahoma (USA), and the Portneuf and Treasure Valleys, Idaho (USA). Specifically, this study aimed to (1) describe and compare perceptions of ecosystem services within and across SES sites, (2) explore how perceptions of ecosystem services vary among local respondents and by sociodemographic factors, and (3) evaluate the overall relationship between place-based SES contexts and ecosystem service perceptions. Our results revealed that cultural ecosystem services were the most highly mentioned among those surveyed across all four sites. However, we found differences in how ecosystem services were perceived among the four SES contexts. For instance, both, social (e.g., gender, education) and local ecological (e.g., land use and climate) characteristics play roles in influencing people's perceptions of which services are important. Overall, our findings suggest the relationship between people's perceptions of ecosystem services and their social-environmental context is complex, which highlights the value of the PECS approach for crafting more effective and inclusive landscape management strategies.

Ecosystems are connected through fluxes of nutrients, organic matter, and organisms. Disturbances that alter structure and function of one ecosystem may have consequences for other linked ecosystems. We investigated how wildfire and subsequent debris flows altered fluxes of invertebrates from tributaries in the Salmon River Basin, Idaho, USA. We compared fluxes of invertebrates downstream through drift and laterally via insect emergence from streams with varying disturbance histories (unburned, burned, and burned + debris flow) during two summers (3–4 years post fire and 2–3 years post debris flow). We observed that the combination of wildfire and debris flow increased the biomass export of invertebrates from tributaries to main-stem ecosystems 2–3 × compared to other streams. In contrast, aquatic insect emergence did not differ in magnitude among streams of different disturbance histories, but instead diverged in timing. Underwater surveys indicated trout in the main-stem river selected confluence habitats, with a tendency for stronger selection of confluences with burned streams. In a behavioral comparison between confluence and non-confluence habitats, rates of agonistic behavior were 4–6 × higher in confluence areas, indicating that confluences may be worth defending. Abundances of web-spinning spiders that depend on emerging insects did not vary with disturbance history in early-mid summer, but tended to be highest in riparian areas along burned streams by August. Because wildfire and debris flows are predicted to increase, our results elucidate potential pathways by which altered disturbance regimes may affect coupled aquatic-terrestrial ecosystems.

Postfire disturbances, such as debris flows, may alter the recovery of stream ecosystems from wildfire. Furthermore, changes to productivity and the dynamics of nutrients and organic matter (OM) in tributaries may affect fluxes of materials and organisms to mainstem habitats. We investigated the consequences of watershed wildfire and debris flows for exports from tributaries to mainstem habitats in the South Fork Salmon River Basin of central Idaho, USA. We compared fluxes from 15 streams (n = 5 unaffected [UA], n = 5 fire-affected [FA], and n = 5 fire + debris flow-affected [FDA]) in summer 2011 (4 y postfire, 3 y postdebris flow). In early June, FDA streams exported 650% higher concentrations of sediment to downstream habitats than did UA streams (mean, FDA: 18.7 g/m3, UA: 1.9 g/m3). Overall, concentrations of dissolved organic C were 75% greater in FDA than in UA streams (FDA: 2.8, UA: 1.6 mg/L), and the quality of OM was higher (i.e., more chlorophyll a [chl a] and higher chl a:OM) in FDA than...

Droughts, fires, and floods are natural disturbances influencing aquatic ecosystems. If drought is accompanied by fires, and if fires are closely followed by floods, teasing apart their distinctive and potentially interactive responses can be difficult. We compared the responses of macroinvertebrate communities to fire via comparisons of streams in burned and unburned catchments in 3 fire-prone biomes that differ biogeographically and climatically (northwestern Mediterranean, southeastern Australia, and northwestern intermountain USA). The responses of macroinvertebrate communities in streams in burned catchments were similar in all biogeographic regions, but the magnitude of these responses varied. Fire combined with high seasonal stream flows, flooding, or drought was associated with reduced measures of taxonomic richness and increased abundance, especially of r-strategist taxa. Differences between sites in burned and unburned catchments were consistently stronger in southeastern Australia than ...

Mercury (Hg) and selenium (Se) biomagnify in aquatic food webs and are toxic to fish and wildlife. The authors measured Hg and Se in organic matter, invertebrates, and fishes in the Colorado River food web at sites spanning 387 river km downstream of Glen Canyon Dam (AZ, USA). Concentrations were relatively high among sites compared with other large rivers (mean wet wt for 6 fishes was 0.17-1.59 μg g(-1) Hg and 1.35-2.65 μg g(-1) Se), but consistent longitudinal patterns in Hg or Se concentrations relative to the dam were lacking. Mercury increased (slope = 0.147) with δ(15) N, a metric of trophic position, indicating biomagnification similar to that observed in other freshwater systems. Organisms regularly exceeded exposure risk thresholds for wildlife and humans (6-100% and 56-100% of samples for Hg and Se, respectfully, among risk thresholds). In the Colorado River, Grand Canyon, Hg and Se concentrations pose exposure risks for fish, wildlife, and humans, and the findings of the present study add to a growing body of evidence showing that remote ecosystems are vulnerable to long-range transport and subsequent bioaccumulation of contaminants. Management of exposure risks in Grand Canyon will remain a challenge, as sources and transport mechanisms of Hg and Se extend far beyond park boundaries.

We critically evaluate some of the key ecological assumptions underpinning the use of nutrient replacement as a means of recovering salmon populations and a range of other organisms thought to be linked to productive salmon runs. These assumptions include: (1) nutrient mitigation mimics the ecological roles of salmon, (2) mitigation is needed to replace salmon-derived nutrients and stimulate primary and invertebrate production in streams, and (3) food resources in rearing habitats limit populations of salmon and resident fishes. First, we call into question assumption one because an array of evidence points to the multi-faceted role played by spawning salmon, including disturbance via redd-building, nutrient recycling by live fish, and consumption by terrestrial consumers. Second, we show that assumption two may require qualification based upon a more complete understanding of nutrient cycling and productivity in streams. Third, we evaluate the empirical evidence supporting food limitation of fish populations and conclude it has been only weakly tested. On the basis of this assessment, we urge caution in the application of nutrient mitigation as a management tool. Although applications of nutrients and other materials intended to mitigate for lost or diminished runs of Pacific salmon may trigger ecological responses within treated ecosystems, contributions of these activities toward actual mitigation may be limited.

Theory states that both the spatial complexity of landscapes and the strength of interactions between consumers and their resources are important for maintaining biodiversity and the balance of nature. Spatial complexity is hypothesized to promote biodiversity by reducing the potential for competitive exclusion; whereas, models show that weak trophic interactions can enhance stability and maintain biodiversity by dampening destabilizing oscillations associated with strong interactions. Here, we show that spatial complexity can reduce the strength of consumer-resource interactions in natural food webs. By sequentially aggregating food webs of individual aquatic habitat patches across a floodplain mosaic, we found that increasing spatial complexity resulted in decreases in the strength of interactions between predators and prey, owing to a greater proportion of weak interactions and a reduced proportion of strong interactions in the meta-food web. The main mechanism behind this pattern was that some patches provided predation refugia for species that were often strongly preyed upon in other patches. If weak trophic interactions do indeed promote stability, then our findings may signal an additional mechanism by which complexity and stability are linked in nature. In turn, this may have implications for how the values of landscape complexity, and the costs of biophysical homogenization, are assessed.

Nearly all ecosystems have been altered by human activities, and most communities are now composed of interacting species that have not co-evolved. These changes may modify species interactions, energy and material flows, and food-web stability. Although structural changes to ecosystems have been widely reported, few studies have linked such changes to dynamic food-web attributes and patterns of energy flow. Moreover, there have been few tests of food-web stability theory in highly disturbed and intensely managed freshwater ecosystems. Such synthetic approaches are needed for predicting the future trajectory of ecosystems, including how they may respond to natural or anthropogenic perturbations. We constructed flow food webs at six locations along a 386-km segment of the Colorado River in Grand Canyon (Arizona, USA) for three years. We characterized food-web structure and production, trophic basis of production, energy efficiencies, and interaction-strength distributions across a spatial gradient of pertu...

The geomorphic template of streams and rivers exerts strong controls on the structure and function of aquatic ecosystems. However, relationships between stream geomorphology and ecosystem structure and function are not always clear and have not been investigated equally across spatial scales. In montane regions, rivers often alternate between canyon-confined segments and unconfined floodplain segments. Yet, few studies have evaluated how this pattern influences aquatic ecosystems. Here, we compared five confined river segments to five paired floodplain segments in terms of allochthonous inputs, aquatic primary producer and invertebrate production, stream retentive capacity, and aquatic invertebrate community composition. We found that floodplains had a higher (up to 4×) retentive capacity, a greater richness (58%) of aquatic invertebrates, and a distinctly different invertebrate community, relative to confined segments. Contrary to our expectations, allochthonous inputs were approximately 2× greater for confined segments, and aquatic primary and invertebrate production exhibited no consistent differences between segment types. However, results did indicate that floodplains had greater overall community respiration than confined segments. Together, these findings suggest that floodplain and confined segments do indeed differ in terms of aquatic ecosystem structure and function but not entirely as expected. Confined segments had greater allochthonous inputs but a lower capacity to retain those inputs, whereas floodplains had a high capacity to retain transported organic matter and also a more diverse community of invertebrates and higher overall community respiration to ‘digest’ retained organic matter. If these findings are generalizable, then they would indicate that confined segments are sources for organic matter within river networks, whereas floodplains act as filters, removing and processing organic matter transported from upstream confined segments. Copyright © 2013 John Wiley & Sons, Ltd.

Summary Non-native species can affect food web and community structure, including the flow of resources from one habitat to another. In many streams of western North America, non-native brook trout (Salvelinus fontinalis) have replaced native cutthroat trout (Oncorhynchus clarkii). Because brook trout naturally occur at higher densities and exhibit different feeding habits, this replacement may have consequences for a range of organisms in stream-riparian food webs. We conducted a large-scale, 2-month field experiment to test whether, compared with cutthroat trout, brook trout reduce benthic insects, cause an increase in stream algae, and reduce emerging adult aquatic insects as well as riparian spiders that rely on emergence as prey. Twenty enclosed reaches from which trout were removed were treated by adding age-1 and older: (i) cutthroat trout at natural density (0.15 fish m−2); (ii) brook trout at a natural density (0.40 fish m−2); (iii) brook trout at a low density equal to the cutthroat trout treatment (0.15 fish m−2) or (iv) no trout added. Brook trout reduced the flux of emerging insects by 55% compared with cutthroat trout, but, surprisingly, only at the lower density. This reduction in emergent insects was predicted to reduce spider abundance by 20%, provided that sufficient branches were available to support riparian spider webs. We also detected an effect of trout on large-bodied benthic and emerging insects, but not on the entire assemblage of benthic insects, or periphyton. We hypothesise that our results were influenced by trout foraging behaviour mediated by their density and size structure. Regardless of the mechanisms responsible, our results indicate that non-native brook trout can reduce the flux of emerging insects, and further analysis showed that this could reduce riparian spiders and birds that prey on them. Similar effects of non-native fish are likely to occur in lotic and lentic waters beyond the western United States.

Climate change (CC) is projected to increase the frequency and severity of natural disturbances (wildfires, insect outbreaks, and debris flows) and shift distributions of terrestrial ecosystems on a global basis. Although such terrestrial changes may affect stream ecosystems, they have not been incorporated into predictions of stream responses to CC. Here, we introduce a conceptual framework to evaluate to what extent responses of streams to CC will be driven by not only changes in thermal and hydrologic regimes, but also alterations of terrestrial processes. We focused on forested watersheds of western North America because this region is projected to experience CC-induced alteration of terrestrial processes. This provided a backdrop for investigating interactive effects of climate and terrestrial responses on streams. Because stream responses to terrestrial processes have been well-studied in contexts largely independent of CC research, we synthesized this knowledge to demonstrate how CC-induced alterations of terrestrial ecosystems may affect streams. Our synthesis indicated that altered terrestrial processes will change terrestrial–aquatic linkages and autotrophic production, potentially yielding greater sensitivity of streams to CC than would be expected based on shifts in temperature and precipitation regime alone. Despite uncertainties that currently constrain predictions regarding stream responses to these additional pathways of change, this synthesis highlighted broader effects of CC that require additional research. Based on widespread evidence that CC is linked to changing terrestrial processes, we conclude that accurate predictions of CC effects on streams may be coupled to the accuracy of predictions for long-term changes in terrestrial ecosystems.

This chapter introduces key concepts and sampling methods concerning drift of benthic stream invertebrates, and the emergence and oviposition behavior of lotic insects, with a focus on the growing array of methods for their quantification being applied in food web and ecosystem studies. Methods are provided to measure the rate of net clogging (which informs the design of drift studies), compare drift among habitat types in wadeable streams and between day and night, and quantify the vertical distribution of drift in large rivers. Methods are also described for sampling emerging adults and estimating emergence flux, examining dispersal distances of adult insects laterally from a stream, and quantifying emergence or postemergent adults as prey available to terrestrial insectivores. Finally, methods are presented regarding a phenomenon closely linked to both emergence and drift, oviposition, including observing behavior and searching for insect eggs, quantifying egg densities and oviposition-site selection, rearing insect eggs, and measuring hatch success.

River regulation may mediate the interactions among native and nonnative species, potentially favoring nonnative species and contributing to the decline of native populations. We examined food resource use and diet overlap among small-bodied fishes in the Grand Canyon section of the Colorado River as a first step in evaluating potential resource competition. We compared the diets of the predominant small-bodied fishes (native Speckled Dace Rhinichthys osculus, juvenile Flannelmouth Sucker Catostomus latipinnis, and juvenile Bluehead Sucker C. discobolus, and nonnative Fathead Minnow Pimephales promelas) across seasons at four sites downstream of Glen Canyon Dam using nonmetric multidimensional scaling and Schoener's similarity index. The diets of these fishes included diatoms, amorphous detritus, aquatic invertebrates (especially simuliid and chironomid larvae), terrestrial invertebrates, and terrestrial vegetation. Diets varied with season and were affected by high turbidity. Fish consumed more amorphous detritus and terrestrial vegetation during the summer monsoon season (July–September), when turbidity was higher. The diets of all species overlapped, but there was large variation in the degree of overlap. The diets of juvenile suckers and Fathead Minnows were most similar, while Speckled Dace had relatively distinct diets. The differences took the form of higher proportions of diatoms and amorphous detritus in the diets of Bluehead Suckers and Fathead Minnows and higher proportions of simuliids and chironomids in those of Speckled Dace. If food resources are or become limiting, diet overlap suggests that competition may occur among native and nonnative species, which could have implications for the population dynamics of these fishes and for the management of the Colorado River ecosystem in Grand Canyon. Received September 4, 2013; accepted February 11, 2014

Although numerous studies have attempted to place species of interest within the context of food webs, such efforts have generally occurred at small scales or disregard potentially important spatial heterogeneity. If food web approaches are to be employed to manage species, studies are needed that evaluate the multiple habitats and associated webs of interactions in which these species participate. Here, we quantify the food webs that sustain rearing salmon and steelhead within a floodplain landscape of the Methow River, Washington, USA, a location where restoration has been proposed to restore side channels in an attempt to recover anadromous fishes. We combined year-long measures of production, food demand, and diet composition for the fish assemblage with estimates of invertebrate prey productivity to quantify food webs within the main channel and five different, intact, side channels; ranging from channels that remained connected to the main channel at low flow to those reduced to floodplain ponds. Although we found that habitats within the floodplain had similar invertebrate prey production, these habitats hosted different local food webs. In the main channel, 95% of total prey consumption flowed to fishes that are not the target of proposed restoration. These fishes consumed 64% and 47% of the prey resources that were found to be important to fueling chinook and steelhead production in the main channel, respectively. Conversely, in side channels, a greater proportion of prey was consumed by anadromous salmonids. As a result, carrying capacity estimates based on food were 251% higher, on average, for anadromous salmonids in side channels than the main channel. However, salmon and steelhead production was generally well below estimated capacity in both the main and side channels, suggesting these habitats are under-seeded with respect to food, and that much larger populations could be supported. Overall, this study demonstrates that floodplain heterogeneity is associated with the occurrence of a mosaic of food webs, all of which were utilized by anadromous salmonids, and all of which may be important to their recovery and persistence. In the long term, these and other fishes would'likely benefit from restoring the processes that maintain floodplain complexity.

Artificial additions of nutrients of differing forms such as salmon carcasses and analog pellets (i.e. pasteurized fishmeal) have been proposed as a means of stimulating aquatic productivity and enhancing populations of anadromous and resident fishes. Nutrient mitigation to enhance fish production in stream ecosystems assumes that the central pathway by which effects occur is bottom‐up, through aquatic primary and secondary production, with little consideration of reciprocal aquatic‐terrestrial pathways. The net outcome (i.e. bottom‐up vs. top‐down) of adding salmon‐derived materials to streams depend on whether or not these subsidies indirectly intensify predation on in situ prey via increases in a shared predator or alleviate such predation pressure. We conducted a 3‐year experiment across nine tributaries of the N. Fork Boise River, Idaho, USA, consisting of 500‐m stream reaches treated with salmon carcasses (n = 3), salmon carcass analog (n = 3), and untreated control reaches (n = 3). We observed 2–8 fold increases in streambed biofilms in the 2–6 weeks following additions of both salmon subsidy treatments in years 1 and 2 and a 1.5‐fold increase in standing crop biomass of aquatic invertebrates to carcass additions in the second year of our experiment. The consumption of benthic invertebrates by stream fishes increased 110–140% and 44–66% in carcass and analog streams in the same time frame, which may have masked invertebrate standing crop responses in years 3 and 4. Resident trout directly consumed 10.0–24.0 g·m−2·yr−1 of salmon carcass and

Understanding the role of species as drivers of ecosystem processes is imperative to preserve, utilize, and sustain ecosystems globally. Addition of species through invasion and loss of species through extirpation or extinction can have profound effects on ecosystem structure and function (Zavaleta et al. 2009). This is especially true for freshwater ecosystems in which a preponderance of native species are threatened with extinction and where nonnative species are frequently introduced (Dudgeon and Smith 2006). Commonly, anthropogenic activities result in the loss of biodiversity and enhance the ability of exotic species to invade and persist in novel habitats (Dudgeon and Smith 2006). Because these activities are expected to increase through time, advances in understanding the consequences of species loss and addition on ecosystem function are needed to guide appropriate management and conservation decisions. The loss and addition of organisms may render habitats functionally impaired (Covich et al. 2004); therefore, understanding the consequences of such change is imperative to manage, mitigate, and restore freshwater ecosystems.

We examined the effect of agricultural land use on whole-stream nutrient availability, biofilm standing crop, and biofilm nutrient limitation and uptake in 21 stream locations in southeastern Idaho. Higher stream water concentrations of dissolved organic carbon (DOC), but not nitrate (NO3-N) or phosphate (PO4-P), were associated with % agriculture in the watershed. Streambed chlorophyll a (Chl a) and ash free dry weight (AF dry wt) also increased with agricultural land use. Nutrient diffusing substrate (NDS) bioassays, which determine biofilm nutrient limitation, showed that Chl a was NO3-N limited and suppressed by labile DOC, whereas AF dry wt did not respond to either NO3-N or labile DOC additions. Together, the different responses of Chl a and AF dry wt suggest potential competition between biofilm autotrophs and heterotrophs for nutrients or other resources. Nutrient uptake, determined with short-term releases of NO3-N, PO4-P, and DOC (as glucose) showed that NO3-N uptake velocity (Vf) increased with agricultural land use. Despite the N-limited status of biofilms indicated by the NDS results, uptake of NO3-N could not be consistently detected at all locations. The differences in response of biofilm to organic carbon enrichment suggests a difference in DOC quality, with labile DOC added with NDS compared to perhaps less-labile DOC found in-stream. Linking the interactive responses of biofilm communities to altered nutrient availability is an important step toward understanding whole ecosystem responses to land-use change.

A disturbance, such as species invasion, can alter the exchange of materials and organisms between ecosystems, with potential consequences for the function of both ecosystems. Russian olive (Elaeagnus angustifolia) is an exotic tree invading riparian corridors in the western United States, and may alter stream organic matter budgets by increasing allochthonous litter and by reducing light via shading, in turn decreasing in-stream primary production. We used a before-after invasion comparison spanning 35 years to show that Russian olive invasion increased allochthonous litter nearly 25-fold to an invaded vs. a control reach of a stream, and we found that this litter decayed more slowly than native willow. Despite a mean 50% increase in canopy cover by Russian olive and associated shading, there were no significant changes in gross primary production. Benthic organic matter storage increased fourfold after Russian olive invasion compared to pre-invasion conditions, but there were no associated changes in stream ecosystem respiration or organic matter export. Thus, estimated stream ecosystem efficiency (ratio of ecosystem respiration to organic matter input) decreased 14%. These findings show that invasions of nonnative plant species in terrestrial habitats can alter resource fluxes to streams with consequences for whole-ecosystem functions.

Non-native brook trout (Salvelinus fontinalis) have replaced native cutthroat trout (Oncorhynchus clarkii) in streams across the western USA. Brook trout have been assumed to be functionally equivalent to cutthroat trout, which could argue against extensive control of this species by government agencies. This assumption was assessed by comparing the diets of the two trout species and their effects on benthic invertebrates in a large-scale field experiment. Within a Rocky Mountain stream where brook trout had already become established, trout species and density were manipulated in 20 fenced sections to create four treatments: cutthroat trout at a density natural for un-invaded streams, brook trout at two densities, natural for invaded streams or reduced to match the natural density of cutthroat trout, and areas depleted of most trout. By the end of the experiment, populations of brook trout at natural density consumed seven times more benthic invertebrates than cutthroat trout. Brook trout also consumed more Apatania caddisflies and Heterlimnius riffle beetles per capita than cutthroat trout. However, the greater consumption did not cause detectable reductions in benthic invertebrate densities. A large-scale experiment of trout removal like this may be inherently limited in its ability to detect effects on benthic invertebrate densities. However, this and several companion studies indicate that stream ecosystems would not function in the same way if cutthroat trout are replaced by brook trout. Sustaining these functions, as well as biodiversity and socioeconomic values, are important reasons to conserve native cutthroat trout. Copyright © 2012 John Wiley & Sons, Ltd.

Frequently nonnative species invade habitats occupied by a similar, even closely related, species, and it is often assumed that the two have functionally redundant ecological roles. However, the consequences of these types of invasions are rarely studied. Eastern brook trout are the most widely distributed and abundant nonnative fish throughout western North America and are assumed to be analogous in terms of ecological function to the native cutthroat trout they replace. However, based on results of previous studies conducted on a few streams, we hypothesized that the two species differ ecologically, and we predicted that brook trout would exhibit greater density, biomass, and annual production than cutthroat trout at the onset of invasion (i.e., in sympatry) and once replacement has occurred (i.e., in allopatry). We compiled data sets (sympatry: n = 169; allopatry: n = 687) from across the historical range of inland cutthroat trout and made comparisons for all sites combined (western scale) and among regions. In sympatry, at the western scale we observed that brook trout exhibited 3.1 times greater density and 2.5 times greater biomass than cutthroat trout, but no difference in production. In allopatry, density and biomass were 1.5 times greater and annual production 1.9 times higher for brook trout than cutthroat trout. Results of comparisons by region were similar, though more variable. In addition, we found that trout density and biomass were correlated with landscape features such as elevation and gradient, but such watershed variables did not account for the differences we observed between the trout species. These results suggest a general pattern of greater density, biomass, and production when brook trout invade and replace cutthroat trout, and highlight the need for more studies that elucidate the mechanisms responsible for this pattern and that investigate the broader consequences of a nonnative species replacing a closely related (and often presumed to be similar) native species.

Large dams have been constructed on rivers to meet human demands for water, electricity, navigation, and recreation. As a consequence, flow and temperature regimes have been altered, strongly affecting river food webs and ecosystem processes. Experimental high-flow dam releases, i.e., controlled floods, have been implemented on the Colorado River, U.S.A., in an effort to reestablish pulsed flood events, redistribute sediments, improve conditions for native fishes, and increase understanding of how dam operations affect physical and biological processes. We quantified secondary production and organic matter flows in the food web below Glen Canyon dam for two years prior and one year after an experimental controlled flood in March 2008. Invertebrate biomass and secondary production declined significantly following the flood (total biomass, 55% decline; total production, 56% decline), with most of the decline driven by reductions in two nonnative invertebrate taxa, Potamopyrgus antipodarum and Gammarus lacustris. Diatoms dominated the trophic basis of invertebrate production before and after the controlled flood, and the largest organic matter flows were from diatoms to the three most productive invertebrate taxa (P. antipodarum, G. lacustris, and Tubificida). In contrast to invertebrates, production of rainbow trout (Oncorhynchus mykiss) increased substantially (194%) following the flood, despite the large decline in total secondary production of the invertebrate assemblage. This counterintuitive result is reconciled by a post-flood increase in production and drift concentrations of select invertebrate prey (i.e., Chironomidae and Simuliidae) that supported a large proportion of trout production but had relatively low secondary production. In addition, interaction strengths, measured as species impact values, were strongest between rainbow trout and these two taxa before and after the flood, demonstrating that the dominant consumer-resource interactions were not necessarily congruent with the dominant organic matter flows. Our study illustrates the value of detailed food web analysis for elucidating pathways by which dam management may alter production and strengths of species interactions in river food webs. We suggest that controlled floods may increase production of nonnative rainbow trout, and this information can be used to help guide future dam management decisions.

Mountain whitefish (Prosopium williamsoni) are among the most abundant native fishes in western North America, yet their role in ecosystems is largely unknown. We investigated their ecology in Big Creek, a 7th-order wilderness watershed in central Idaho. Based on underwater visual counts and hook and line surveys conducted during summer months, as well as tissue analysis, we estimated mountain whitefish distribution and abundance, total biomass, production, and energy and nutrient contributions along a 60 km segment of Big Creek. We observed that abundance decreased with distance upstream, and 93% of mountain whitefish surveyed in Big Creek were greater than 200 mm (≥ 3 years); few juveniles were observed, suggesting that rearing occurs in habitats downstream. Mountain whitefish were the dominant fish in Big Creek, in terms of both abundance (33% of fish observed) and biomass (57% of salmonid biomass). We estimated their production averaged across the study segment as 0.51 g m-2 yr-1, though it r...