Using social-ecological models to explore stream connectivity outcomes for stakeholders and Yellowstone cutthroat trout.

Despite growing interest in conservation and re-establishment of ecological connectivity, few studies have explored its context-specific social-ecological outcomes. We aimed to explore social and ecological outcomes to changing stream connectivity for both stakeholders and native fish species impacted by habitat fragmentation and nonnative species. We (1) investigated stakeholder perceptions of the drivers and outcomes of stream connectivity, and (2) evaluated the effects of stakeholder-identified connectivity and nonnative species scenarios on Yellowstone cutthroat trout (YCT) populations. Our study was conducted in the Teton River, Idaho, USA. We integrated two modeling approaches, mental modeling and individual-based ecological modeling, to explore social-ecological outcomes for stakeholders and YCT populations. Aggregation of mental models revealed an emergent pattern of increasing complexity as more types of stakeholders were considered, as well as gaps and linkages among different stakeholder knowledge areas. These results highlight the importance of knowledge sharing among stakeholders when making decisions about connectivity. Additionally, the results from the individual-based models suggested that the potential for a large, migratory life history form of YCT, in addition to self-preference mating where they overlap with rainbow trout, had the strongest effects on outcomes for YCT. Exploring social and ecological drivers and outcomes to changing connectivity is useful for anticipating and adapting to unintended outcomes, as well as making decisions for desirable outcomes. The results from this study can contribute to the management dialogue surrounding stream connectivity in the Teton River, as well as to our understanding of connectivity conservation and its outcomes more broadly.

Riverscape approaches in practice: perspectives and applications.

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.

Magnitude and direction of stream-forest community interactions change with timescale.

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.

Reverberating effects of resource exchanges in stream-riparian food webs.

Fluxes of materials or organisms across ecological boundaries, often termed "resource subsidies," directly affect recipient food webs. Few studies have addressed how such direct responses in one ecosystem may, in turn, influence the fluxes of materials or organisms to other habitats or the potential for feedback relationships to occur among ecosystems. As part of a large-scale, multi-year experiment, we evaluated the hypothesis that the input of a marine-derived subsidy results in a complex array of resource exchanges (i.e., inputs, outputs, feedbacks) between stream and riparian ecosystems as responses disperse across ecological boundaries. Moreover, we evaluated how the physical properties of resource subsidies mediated complex responses by contrasting carcasses with a pelletized salmon treatment. We found that salmon carcasses altered stream-riparian food webs by directly subsidizing multiple aquatic and terrestrial organisms (e.g., benthic insect larvae, fishes, and terrestrial flies). Such responses further influenced food webs along indirect pathways, some of which spanned land and water (e.g., subsidized fishes reduced aquatic insect emergence, with consequences for spiders and bats). Subsidy-mediated feedbacks manifested when carcasses were removed to riparian habitats where they were colonized by carrion flies, some of which fell into the stream and acted as another prey subsidy for fishes. As the effects of salmon subsidies propagated through the stream-riparian food web, the sign of consumer responses was not always positive and appeared to be determined by the outcome of trophic interactions, such that localized trophic interactions within one ecosystem mediated the export of organisms to others.

Dry Wetlands: Nutrient Dynamics in Ephemeral Constructed Stormwater Wetlands.

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.

Global patterns and drivers of ecosystem functioning in rivers and riparian zones.

River ecosystems receive and process vast quantities of terrestrial organic carbon, the fate of which depends strongly on microbial activity. Variation in and controls of processing rates, however, are poorly characterized at the global scale. In response, we used a peer-sourced research network and a highly standardized carbon processing assay to conduct a global-scale field experiment in greater than 1000 river and riparian sites. We found that Earth's biomes have distinct carbon processing signatures. Slow processing is evident across latitudes, whereas rapid rates are restricted to lower latitudes. Both the mean rate and variability decline with latitude, suggesting temperature constraints toward the poles and greater roles for other environmental drivers (e.g., nutrient loading) toward the equator. These results and data set the stage for unprecedented "next-generation biomonitoring" by establishing baselines to help quantify environmental impacts to the functioning of ecosystems at a global scale.

Land conservation can mitigate freshwater ecosystem services degradation due to climate change in a semiarid catchment: The case of the Portneuf River catchment, Idaho, USA.

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.

Spatial complexity reduces interaction strengths in the meta-food web of a river floodplain mosaic.

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.

Mercury and selenium accumulation in the Colorado River food web, Grand Canyon, USA.

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.

A Critical Assessment of the Ecological Assumptions Underpinning Compensatory Mitigation of Salmon-Derived Nutrients.

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.

The floodplain food web mosaic: a study of its importance to salmon and steelhead with implications for their recovery.

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.

Assessing the potential for salmon recovery via floodplain restoration: a multitrophic level comparison of dredge-mined to reference segments.

Pre-restoration studies typically focus on physical habitat, rather than the food-base that supports aquatic species. However, both food and habitat are necessary to support the species that habitat restoration is frequently aimed at recovering. Here we evaluate if and how the productivity of the food-base that supports fish production is impaired in a dredge-mined floodplain within the Yankee Fork Salmon River (YFSR), Idaho (USA); a site where past restoration has occurred and where more has been proposed to help recover anadromous salmonids. Utilizing an ecosystem approach, we found that the dredged segment had comparable terrestrial leaf and invertebrate inputs, aquatic primary producer biomass, and production of aquatic invertebrates relative to five reference floodplains. Thus, the food-base in the dredged segment did not necessarily appear impaired. On the other hand, we observed that off-channel aquatic habitats were frequently important to productivity in reference floodplains, and the connection of these habitats in the dredged segment via previous restoration increased invertebrate productivity by 58%. However, using a simple bioenergetic model, we estimated that the invertebrate food-base was at least 4× larger than present demand for food by fish in dredged and reference segments. In the context of salmon recovery efforts, this observation questions whether additional food-base productivity provided by further habitat restoration would be warranted in the YFSR. Together, our findings highlight the importance of studies that assess the aquatic food-base, and emphasize the need for more robust ecosystem models that evaluate factors potentially limiting fish populations that are the target of restoration.

Ecosystem ecology meets adaptive management: food web response to a controlled flood on the Colorado River, Glen Canyon.

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.

Quantity and quality: unifying food web and ecosystem perspectives on the role of resource subsidies in freshwaters.

Although the study of resource subsidies has emerged as a key topic in both ecosystem and food web ecology, the dialogue over their role has been limited by separate approaches that emphasize either subsidy quantity or quality. Considering quantity and quality together may provide a simple, but previously unexplored, framework for identifying the mechanisms that govern the importance of subsidies for recipient food webs and ecosystems. Using a literature review of > 90 studies of open-water metabolism in lakes and streams, we show that high-flux, low-quality subsidies can drive freshwater ecosystem dynamics. Because most of these ecosystems are net heterotrophic, allochthonous inputs must subsidize respiration. Second, using a literature review of subsidy quality and use, we demonstrate that animals select for high-quality food resources in proportions greater than would be predicted based on food quantity, and regardless of allochthonous or autochthonous origin. This finding suggests that low-flux, high-quality subsidies may be selected for by animals, and in turn may disproportionately affect food web and ecosystem processes (e.g., animal production, trophic energy or organic matter flow, trophic cascades). We then synthesize and review approaches that evaluate the role of subsidies and explicitly merge ecosystem and food web perspectives by placing food web measurements in the context of ecosystem budgets, by comparing trophic and ecosystem production and fluxes, and by constructing flow food webs. These tools can and should be used to address future questions about subsidies, such as the relative importance of subsidies to different trophic levels and how subsidies may maintain or disrupt ecosystem stability and food web interactions.

Species replacement by a nonnative salmonid alters ecosystem function by reducing prey subsidies that support riparian spiders.

Replacement of a native species by a nonnative can have strong effects on ecosystem function, such as altering nutrient cycling or disturbance frequency. Replacements may cause shifts in ecosystem function because nonnatives establish at different biomass, or because they differ from native species in traits like foraging behavior. However, no studies have compared effects of wholesale replacement of a native by a nonnative species on subsidies that support consumers in adjacent habitats, nor quantified the magnitude of these effects. We examined whether streams invaded by nonnative brook trout (Salvelinus fontinalis) in two regions of the Rocky Mountains, USA, produced fewer emerging adult aquatic insects compared to paired streams with native cutthroat trout (Oncorhynchus clarkii), and whether riparian spiders that depend on these prey were less abundant along streams with lower total insect emergence. As predicted, emergence density was 36% lower from streams with the nonnative fish. Biomass of brook trout was higher than the cutthroat trout they replaced, but even after accounting for this difference, emergence was 24% lower from brook trout streams. More riparian spiders were counted along streams with greater total emergence across the water surface. Based on these results, we predicted that brook trout replacement would result in 6-20% fewer spiders in the two regions. When brook trout replace cutthroat trout, they reduce cross-habitat resource subsidies and alter ecosystem function in stream-riparian food webs, not only owing to increased biomass but also because traits apparently differ from native cutthroat trout.

Predicting effects of hydrologic alteration and climate change on ecosystem metabolism in a western U.S. river.

We estimated past and future hydrographs and patterns of ecosystem metabolism in a fifth-order river of the western United States, where water use and climate change are both expected to alter hydrology in the immediate future. We first reconstructed the unregulated hydrograph to estimate how the current hydrograph has been altered. Due to consumptive use, 95% as irrigation, current discharge during summer (July-September) was 70% lower than would occur if the river was unregulated. We then predicted a future hydrograph including effects of consumptive use and climate change; the magnitude of flow changes were minor under this regime relative to those already manifested by consumptive uses. We used time-series regression and a six-year continuous record of open-water metabolism to demonstrate that, under the current hydrologic regime, gross primary production (GPP) was dependent on both water temperature and flow and that ecosystem respiration (ER) was most dependent on temperature. Monte Carlo simulations under the three hydrologic regimes and three temperature scenario's (current, +2 degrees C, +4 degrees C) suggested that flow, but not temperature, may have profound effects on the magnitude of metabolism. Linking temporally detailed analyses of ecological function and hydrology may lead to better understanding and management of changes due to basin-scale water use and/or global-scale climate change.

Invading rainbow trout usurp a terrestrial prey subsidy from native charr and reduce their growth and abundance.

Movements of prey organisms across ecosystem boundaries often subsidize consumer populations in adjacent habitats. Human disturbances such as habitat degradation or non-native species invasions may alter the characteristics or fate of these prey subsidies, but few studies have measured the direct effects of this disruption on the growth and local abundance of predators in recipient habitats. Here we present evidence, obtained from a combined experimental and comparative study in northern Japan, that an invading stream fish usurped the flux of allochthonous prey to a native fish, consequently altering the diet and reducing the growth and abundance of the native species. A large-scale field experiment showed that excluding terrestrial invertebrates that fell into the stream with a mesh greenhouse reduced terrestrial prey in diets of native Dolly Varden charr (Salvelinus malma) by 46-70%, and reduced their growth by 25% over six weeks. However, when nonnative rainbow trout (Oncorhynchus mykiss) were introduced, they monopolized these prey and caused an even greater reduction of terrestrial prey in charr diets of 82-93%, and reduced charr growth by 31% over the same period. Adding both greenhouse and rainbow trout treatments together produced similar results to adding either alone. Results from a comparative field study of six other stream sites in the region corroborated the experimental findings, showing that at invaded sites rainbow trout usurped the terrestrial prey subsidy, causing a more than 75% decrease in the biomass of terrestrial invertebrates in Dolly Varden diets and forcing them to shift their foraging to insects on the stream bottom. Moreover, at sites with even low densities of rainbow trout, biomass of Dolly Varden was more than 75% lower than at sites without rainbow trout. Disruption of resource fluxes between habitats may be a common, but unidentified, consequence of invasions, and an additional mechanism contributing to the loss of native species.