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Colden V. Baxter

Publications and source records attributed to Colden V. Baxter.

At least 19 recordsLinked to original sources

Bridging ecology and geosciences in riverscapes: Implications for process-based restoration

There has been a growing interest in integrating geological and ecological processes for sustainable river management and restoration. Lotic systems are shaped by diverse physical processes, including geology, geomorphology, hydrology, and interactions with terrestrial processes. However, restoration practices often prioritize specific habitats or river forms without fully considering the underlying physical processes that support biological communities and ecosystem functions. In this paper, we synthesize studies that integrally examine geological and ecological processes across different scales and components of riverscapes, including geohydrological processes, which have been less investigated in riverscape studies. We begin by examining processes at broader spatial scales, including river–watershed and river–riparian interactions, and gradually narrow our focus to the dynamics that occur among habitats within river channels, through which we highlight the significance of conceptualizing rivers as dynamic “networks” rather than linear features. Finally, we identify both scientific and practical challenges that can be addressed to bridge the gap between basic-science implications and their implementation in riverscape restorations.

Ecological Research

Food-web dynamics of a floodplain mosaic overshadow the effects of engineered logjams for Pacific salmon and steelhead

Food webs vary in space and time. The structure and spatial arrangement of food webs are theorized to mediate temporal dynamics of energy flow, but empirical corroboration in intermediate-scale landscapes is scarce. River-floodplain landscapes encompass a mosaic of aquatic habitat patches and food webs, supporting a variety of aquatic consumers of conservation concern. How the structure and productivity of these patch-scale food webs change through time, and how floodplain restoration influences their dynamics, are unevaluated. We measured productivity and food-web dynamics across a mosaic of main-channel and side-channel habitats of the Methow River, WA, USA, during two study years (2009–2010; 2015–2016) and examined how food webs that sustained juvenile anadromous salmonids responded to habitat manipulation. By quantifying temporal variation in secondary production and organic matter flow across nontreated river-floodplain habitats and comparing that variation to a side channel treated with engineered logjams, we jointly confronted spatial food-web theory and assessed whether food-web dynamics in the treated side channel exceeded natural variation exhibited in nontreated habitats. We observed that organic matter flow through the more complex, main-channel food web was similar between study years, whereas organic matter flow through the simpler, side-channel food webs changed up to ~4-fold. In the side channel treated with engineered logjams, production of benthic invertebrates and juvenile salmonids increased between study years by 2× and 4×, respectively; however, these changes did not surpass the temporal variation observed in untreated habitats. For instance, juvenile salmonid production rose 17-fold in one untreated side-channel habitat, and natural aggregation of large wood in another coincided with a shift to community and food-web dominance by juvenile salmonids. Our findings suggest that interannual dynamism in material flux across floodplain habitat mosaics is interrelated with patchiness in food-web complexity and may overshadow the ecological responses to localized river restoration. Although this dynamism may inhibit detection of the ecological effects of river restoration, it may also act to stabilize aquatic ecosystems and buffer salmon and other species of conservation concern in the long term. As such, natural, landscape-level patchiness and dynamism in food webs should be integrated into conceptual foundations of process-based, river restoration.

Washington

Human activities shape global patterns of decomposition rates in rivers

Rivers and streams contribute to global carbon cycling by decomposing immense quantities of terrestrial plant matter. However, decomposition rates are highly variable and large-scale patterns and drivers of this process remain poorly understood. Using a cellulose-based assay to reflect the primary constituent of plant detritus, we generated a predictive model (81% variance explained) for cellulose decomposition rates across 514 globally distributed streams. A large number of variables were important for predicting decomposition, highlighting the complexity of this process at the global scale. Predicted cellulose decomposition rates, when combined with genus-level litter quality attributes, explain published leaf litter decomposition rates with high accuracy (70% variance explained). Our global map provides estimates of rates across vast understudied areas of Earth and reveals rapid decomposition across continental-scale areas dominated by human activities.

Science

Food web perspectives and methods for riverine fish conservation

Food web analyses offer useful insights into understanding how species interactions, trophic relationships, and energy flow underpin important demographic parameters of fish populations such as survival, growth, and reproduction. However, the vast amount of food web literature and the diversity of approaches can be a deterrent to fisheries practitioners engaged in on-the-ground research, monitoring, or restoration. Incorporation of food web perspectives into contemporary fisheries management and conservation is especially rare in riverine systems, where approaches often focus more on the influence of physical habitat and water temperature on fish populations. In this review, we first discuss the importance of food webs in the context of several common fisheries management issues, including assessing carrying capacity, evaluating the effects of habitat change, examining species introductions or extinctions, considering bioaccumulation of toxins, and predicting the effects of climate change and other anthropogenic stressors on riverine fishes. We then examine several relevant perspectives: basic food web description, metabolic models, trophic basis of production, mass-abundance network approaches, ecological stoichiometry, and mathematical modeling. Finally, we highlight several existing and emerging methodologies including diet and prey surveys, eDNA, stable isotopes, fatty acids, and community and network analysis. Although our emphasis and most examples are focused on salmonids in riverine environments, the concepts are easily generalizable to other freshwater fish taxa and ecosystems.

Wiley Interdisciplinary Reviews (WIREs): Water

Dietary composition and fatty acid content of giant salmonflies (Pteronarcys californica) in two Rocky Mountain rivers

Many aquatic invertebrates are declining or facing extinction from stressors that compromise physiology, resource consumption, reproduction, and phenology. However, the influence of these common stressors specifically on consumer–resource interactions for aquatic invertebrate consumers is only beginning to be understood. We conducted a field study to investigate Pteronarcys californica (i.e., the “giant salmonfly”), a large-bodied insect that is ecologically and culturally significant to rivers throughout the western United States. We sampled gut contents and polyunsaturated fatty acid composition of salmonflies to compare resource consumption across river (Madison or Gallatin, Montana), sex (male or female), and habitat (rock or woody debris). We found that allochthonous detritus comprised the majority of salmonfly diets in the Gallatin and Madison Rivers, making up 68% of the gut contents on average, followed by amorphous detritus, diatoms, and filamentous algae. Diets showed little variation across river, sex, or length. Minor differences in diets were detected by habitat type, with a higher proportion of diatoms in the diets of salmonflies collected from rocky habitat compared to woody debris. Fatty acid composition generally supported the results of gut content analysis but highlighted the importance of primary producers. The presence of eicosapentaenoic acid (20:5n-3) and alpha linolenic acid (18:3n-3) indicated consumption of diatoms and filamentous green algae, respectively. Our research underscores the importance of a healthy riparian zone that provides allochthonous detritus for invertebrate nutrition as well as the role of algae as an important source of fatty acids.

Montana

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.

Biological Reviews

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.

Ecology

Food web controls on mercury fluxes and fate in the Colorado River, Grand Canyon

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.

Arizona, Nevada

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.

Idaho

Flow management for hydropower extirpates aquatic insects, undermining river food webs

Dams impound the majority of rivers and provide important societal benefits, especially daily water releases that enable on-peak hydroelectricity generation. Such “hydropeaking” is common worldwide, but its downstream impacts remain unclear. We evaluated the response of aquatic insects, a cornerstone of river food webs, to hydropeaking using a life history–hydrodynamic model. Our model predicts that aquatic-insect abundance will depend on a basic life-history trait—adult egg-laying behavior—such that open-water layers will be unaffected by hydropeaking, whereas ecologically important and widespread river-edge layers, such as mayflies, will be extirpated. These predictions are supported by a more-than-2500-sample, citizen-science data set of aquatic insects from the Colorado River in the Grand Canyon and by a survey of insect diversity and hydropeaking intensity across dammed rivers of the Western United States. Our study reveals a hydropeaking-related life history bottleneck that precludes viable populations of many aquatic insects from inhabiting regulated rivers.

BioScience

Production of resident fish benefits from experimental salmon subsidies via direct and indirect pathways across stream-riparian boundaries

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&ndash;8 fold increases in streambed biofilms in the 2&ndash;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&ndash;140% and 44&ndash;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&ndash;24.0 g&middot;m &minus;2 &middot;yr &minus;1 of salmon carcass and <1&ndash;11.0 g&middot;m &minus;2 &middot;yr &minus;1 of analog material, which resulted in 1.2&ndash;2.9 g&middot;m &minus;2 &middot;yr &minus;1 and 0.03&ndash;1.4 g&middot;m &minus;2 &middot;yr &minus;1 of tissue produced. In addition, a feedback flux of terrestrial maggots to streams contributed 0.0&ndash;2.0 g&middot;m &minus;2 &middot;yr &minus;1 to trout production. Overall, treatments increased annual trout production by 2&ndash;3 fold, though density and biomass were unaffected. Our results indicate the strength of bottom-up and top-down responses to subsidy additions was asymmetrical, with top-down forces masking bottom-up effects that required multiple years to manifest. The findings also highlight the need for nutrient mitigation programs to consider multiple pathways of energy and nutrient flow to account for the complex effects of salmon subsidies in stream-riparian ecosystems.

Ecosphere

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.

Environmental Management

High diet overlap between native small-bodied fishes and nonnative fathead minnow in the Colorado River, Grand Canyon, Arizona

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.

Arizona

Nutrient additions to mitigate for loss of Pacific salmon: consequences for stream biofilm and nutrient dynamics

Mitigation activities designed to supplement nutrient and organic matter inputs to streams experiencing decline or loss of Pacific salmon typically presuppose that an important pathway by which salmon nutrients are moved to fish (anadromous and/or resident) is via nutrient incorporation by biofilms and subsequent bottom-up stimulation of biofilm production, which is nutrient-limited in many ecosystems where salmon returns have declined. Our objective was to quantify the magnitude of nutrient incorporation and biofilm dynamics that underpin this indirect pathway in response to experimental additions of salmon carcasses and pelletized fish meal (a.k.a., salmon carcass analogs) to 500-m reaches of central Idaho streams over three years. Biofilm standing crops increased 2&ndash;8-fold and incorporated marine-derived nutrients (measured using 15 N and 13 C) in the month following treatment, but these responses did not persist year-to-year. Biofilms were nitrogen (N) limited before treatments, and remained N limited in analog, but not carcass-treated reaches. Despite these biofilm responses, in the month following treatment total N load was equal to 33&ndash;47% of the N added to the treated reaches, and N spiraling measurements suggested that as much as 20%, but more likely 2&ndash;3% of added N was taken up by microbes. Design of biologically and cost-effective strategies for nutrient addition will require understanding the rates at which stream microbes take up nutrients and the downstream distance traveled by exported nutrients.

Idaho

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.

Washington

Food-web dynamics in a large river discontinuum

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 perturbation (i.e., distance from Glen Canyon Dam), as well as before and after an experimental flood. We found strong longitudinal patterns in food-web characteristics that strongly correlated with the spatial position of large tributaries. Above tributaries, food webs were dominated by nonnative New Zealand mudsnails (62% of production) and nonnative rainbow trout (100% of fish production). The simple structure of these food webs led to few dominant energy pathways (diatoms to few invertebrate taxa to rainbow trout), large energy inefficiencies (i.e., <20% of invertebrate production consumed by fishes), and right-skewed interaction-strength distributions, consistent with theoretical instability. Below large tributaries, invertebrate production declined ∼18-fold, while fish production remained similar to upstream sites and comprised predominately native taxa (80–100% of production). Sites below large tributaries had increasingly reticulate and detritus-based food webs with a higher prevalence of omnivory, as well as interaction strength distributions more typical of theoretically stable food webs (i.e., nearly twofold higher proportion of weak interactions). Consistent with theory, downstream food webs were less responsive to the experimental flood than sites closest to the dam. We show how human-induced shifts to food-web structure can affect energy flow and interaction strengths, and we show that these changes have consequences for food-web function and response to perturbations.

Arizona

Macroinvertebrate diets reflect tributary inputs and turbidity-driven changes in food availability in the Colorado River downstream of Glen Canyon Dam

Physical changes to rivers associated with large dams (e.g., water temperature) directly alter macroinvertebrate assemblages. Large dams also may indirectly alter these assemblages by changing the food resources available to support macroinvertebrate production. We examined the diets of the 4 most common macroinvertebrate taxa in the Colorado River through Glen and Grand Canyons, seasonally, at 6 sites for 2.5 y. We compared macroinvertebrate diet composition to the composition of epilithon (rock and cliff faces) communities and suspended organic seston to evaluate the degree to which macroinvertebrate diets tracked downstream changes in resource availability. Diets contained greater proportions of algal resources in the tailwater of Glen Canyon Dam and more terrestrial-based resources at sites downstream of the 1st major tributary. As predicted, macroinvertebrate diets tracked turbidity-driven changes in resource availability, and river turbidity partially explained variability in macroinvertebrate diets. The relative proportions of resources assimilated by macroinvertebrates ranged from dominance by algae to terrestrial-based resources, despite greater assimilation efficiencies for algal than terrestrial C. Terrestrial resources were most important during high turbidity conditions, which occurred during the late-summer monsoon season (July–October) when tributaries contributed large amounts of organic matter to the mainstem and suspended sediments reduced algal production. Macroinvertebrate diets were influenced by seasonal changes in tributary inputs and turbidity, a result suggesting macroinvertebrate diets in regulated rivers may be temporally dynamic and driven by tributary inputs.

Arizona

Native and nonnative fish populations of the Colorado River are food limited--evidence from new food web analyses

Fish populations in the Colorado River downstream from Glen Canyon Dam appear to be limited by the availability of high-quality invertebrate prey. Midge and blackfly production is low and nonnative rainbow trout in Glen Canyon and native fishes in Grand Canyon consume virtually all of the midge and blackfly biomass that is produced annually. In Glen Canyon, the invertebrate assemblage is dominated by nonnative New Zealand mudsnails, the food web has a simple structure, and transfers of energy from the base of the web (algae) to the top of the web (rainbow trout) are inefficient. The food webs in Grand Canyon are more complex relative to Glen Canyon, because, on average, each species in the web is involved in more interactions and feeding connections. Based on theory and on studies from other ecosystems, the structure and organization of Grand Canyon food webs should make them more stable and less susceptible to large changes following perturbations of the flow regime relative to food webs in Glen Canyon. In support of this hypothesis, Grand Canyon food webs were much less affected by a 2008 controlled flood relative to the food web in Glen Canyon.

Colorado River