Theodore A. Kennedy

The role of discharge variation in scaling of drainage area and food chain length in rivers

River Food Chains Food chain length, or the number of organisms involved in any eat-or-be-eaten relationship, is a key property of aquatic habitats that regulates nutrient cycling, energy flow, and carbon exchange between aquatic ecosystems and the atmosphere. Investigating river food webs in North American watersheds, Sabo et al. (p. 965, published online 14 October) found discharge variation governed the link between ecosystem size (watershed area) and food chain length. River-drying truncated food chain length because intermittent streams have much higher discharge variation than perennial rivers. These results could be important for the management of rivers as drought and climate change may increase the frequency of river drying and discharge variability. The lengths of river food chains are indirectly related to drainage area and directly affected by flow and discharge. Food chain length (FCL) is a fundamental component of food web structure. Studies in a variety of ecosystems suggest that FCL is determined by energy supply, environmental stability, and/or ecosystem size, but the nature of the relationship between environmental stability and FCL, and the mechanism linking ecosystem size to FCL, remain unclear. Here we show that FCL increases with drainage area and decreases with hydrologic variability and intermittency across 36 North American rivers. Our analysis further suggests that hydrologic variability is the mechanism underlying the correlation between ecosystem size and FCL in rivers. Ecosystem size lengthens river food chains by integrating and attenuating discharge variation through stream networks, thereby enhancing environmental stability in larger river systems.

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.

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 pertu...

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

Abstract.  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.

Air–water oxygen exchange in a large whitewater river

Air – water gas exchange governs fluxes of gas into and out of aquatic ecosystems. Knowing this flux is necessary to calculate gas budgets (i.e., O2) to estimate whole-ecosystem metabolism and basin-scale carbon budgets. Empirical data on rates of gas exchange for streams, estuaries, and oceans are readily available. However, there are few data from large rivers and no data from whitewater rapids. We measured gas transfer velocity in the Colorado River, Grand Canyon, as decline in O2 saturation deficit, 7 times in a 28-km segment spanning 7 rapids. The O2 saturation deficit exists because of hypolimnetic discharge from Glen Canyon Dam, located 25 km upriver from Lees Ferry. Gas transfer velocity (k600) increased with slope of the immediate reach. k600 was , 10 cm h -1 in flat reaches, while k600 for the steepest rapid ranged 3600 – 7700 cm h -1 , an extremely high value ofk600. Using the rate of gas exchange per unit length of water surface elevation (Kdrop ,m -1 ), segment-integrated k600 varied between 74 and 101 cm h -1 . Using Kdrop we scaled k600 to the remainder of the Colorado River in Grand Canyon. At the scale corresponding to the segment length where 80% of the O2 exchanged with the atmosphere (mean length ¼ 26.1 km), k600 varied 4.5-fold between 56 and 272 cm h -1 with a mean of 113 cm h -1 . Gas transfer velocity for the Colorado River was higher than those from other aquatic ecosystems because of large rapids. Our approach of scalingk600 based onKdrop allows comparing gas transfer velocity across rivers with spatially heterogeneous morphology.

Flow management and fish density regulate salmonid recruitment and adult size in tailwaters across western North America

Rainbow and brown trout have been intentionally introduced into tailwaters downriver of dams globally and provide billions of dollars in economic benefits. At the same time, recruitment and maximum length of trout populations in tailwaters often fluctuate erratically, which negatively affects the value of fisheries. Large recruitment events may increase dispersal downriver where other fish species may be a priority (e.g., endangered species). There is an urgent need to understand the drivers of trout population dynamics in tailwaters, in particular the role of flow management. Here, we evaluate how flow, fish density, and other physical factors of the river influence recruitment and mean adult length in tailwaters across western North America, using data from 29 dams spanning 1-19 years. Rainbow trout recruitment was negatively correlated with high annual, summer, and spring flow and dam latitude, and positively correlated with high winter flow, subadult brown trout catch, and reservoir storage capacity. Brown trout recruitment was negatively correlated with high water velocity and daily fluctuations in flow (i.e., hydropeaking) and positively correlated with adult rainbow trout catch. Among these many drivers, rainbow trout recruitment was primarily correlated with high winter flow combined with low spring flow, whereas brown trout recruitment was most related to high water velocity. The mean lengths of adult rainbow and brown trout were influenced by similar flow and catch metrics. Length in both species was positively correlated with high annual flow but declined in tailwaters with high daily fluctuations in flow, high catch rates of conspecifics, and when large cohorts recruited to adult size. Whereas brown trout did not respond to the proportion of water allocated between seasons, rainbow trout length increased in rivers that released more water during winter than in spring. Rainbow trout length was primarily related to high catch rates of conspecifics, whereas brown trout length was mainly related to large cohorts recruiting to the adult size class. Species-specific responses to flow management are likely attributable to differences in seasonal timing of key life history events such as spawning, egg hatching, and fry emergence.

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.

High Diet Overlap between Native Small-Bodied Fishes and Nonnative Fathead Minnow in the Colorado River, Grand Canyon, Arizona

AbstractRiver 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 Schoeners 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 a...

Building a better sticky trap: description of an easy-to-use trap and pole mount for quantifying the abundance of adult aquatic insects

Abstract: Insect emergence is a fundamental process in freshwaters. It is a critical life-history stage for aquatic insects and provides an important prey resource for terrestrial and aquatic consumers. Sticky traps are increasingly being used to sample these insects. The most common design consists of an acetate sheet coated with a nondrying adhesive that is attached to a wire frame or cylinder. These traps must be prepared at the deployment site, a process that can be time consuming and difficult given the vagaries of field conditions. Our goals were to develop a sturdy, low-cost sticky trap that could be prepared in advance, rapidly deployed and recovered in the field, and used to estimate the flight direction of insects. We used 150-mm Petri dishes with lids. The dishes can be coated cleanly and consistently with Tangle-Trap® adhesive. Deploying traps is simple and requires only a pole set near the body of water being sampled. Four dishes can be attached to the pole using Velcro and aligned in 4 different directions to enable quantification of insect flight direction. After sampling, Petri dishes can be taped closed, packed in boxes, and stored indefinitely. Petri traps are comparable in price to standard acetate sheet traps at ∼US

Pulsed flows, tributary inputs and food‐web structure in a highly regulated river

0.50/directional deployment, but they require more space for storage than acetate sheet traps. However, a major benefit of Petri traps is that field deployment times are those of acetate traps. Our study demonstrated that large Petri dishes are an ideal platform for sampling postemergent adult aquatic insects, particularly when the study design involves estimating flight direction and when rapid deployment and recovery of traps is critical.