Joseph R. Benjamin

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.

Potential Effects of Changes in Temperature and Food Resources on Life History Trajectories of Juvenile Oncorhynchus mykiss

Abstract Increasing temperatures and changes in food resources owing to climate change may alter the growth and migratory behavior of organisms. This is particularly important for salmonid species like Oncorhynchus mykiss, where some individuals remain in freshwater to mature (nonanadromous Rainbow Trout) and others migrate to sea (anadromous Steelhead). Whether one strategy is adopted over the other may depend on the individuals growth and size. In this study, we explored (1) how water temperature in Beaver Creek, a tributary to the Methow River, Washington, may increase under four climate scenarios, (2) how these thermal changes may alter the life history trajectory followed by O. mykiss (i.e., when and if to smolt), and (3) how changes in food quality or quantity might interact with increasing temperatures. We combined bioenergetic and state-dependent life history models parameterized for O. mykiss in Beaver Creek to mimic baseline life history trajectories. Based on our simulations, when mean water t...

Measuring adult insect emergence from streams: the influence of trap placement and a comparison with benthic sampling

Abstract Increased need to quantify adult insects emerging from streams as a part of foodweb and ecosystem studies has placed new demands on techniques used to sample adults. The population sampled must be better understood to establish the scope of inferences that may be drawn from emergence data. We used data from 2 different studies to compare the structure of insect assemblages represented by benthic samples and emergence-trap samples and to compare adult insect assemblages collected in emergence traps placed at mid-channel vs streambank locations. Nonmetric multidimensional scaling (NMDS) ordination showed that some components of the benthic assemblage were underrepresented in the emergence-trap samples and others were underrepresented in benthic samples. These differences were mainly caused by taxa that emerged by crawling out on the stream bank (e.g., Plecoptera) or whose larvae reside in habitats, such as stream margins, that are underrepresented with traditional benthic sampling (e.g., Dixidae). The flux of insects into traps placed mid-channel did not differ significantly from the flux into traps placed along the stream bank. Taxa collected by mid-channel and streambank traps overlapped considerably, but midchannel traps tended to collect proportionally more Trichoptera, Ephemeroptera, and Diptera, whereas streambank traps collected proportionally more Plecoptera. Our results can be used to improve trapping designs for future assessments of aquatic insect emergence in studies of insect behavior and life histories and as part of foodweb and ecosystem research.

Do Nonnative Salmonines Exhibit Greater Density and Production than the Natives They Replace? A Comparison of Nonnative Brook Trout with Native Cutthroat Trout

Abstract The replacement of native species by nonnative species has been widely documented. In aquatic ecosystems, nonnative trout often replace native trout species to which they are closely related. For example, nonnative brook trout Salvelinus fontinalis have replaced native cutthroat trout Oncorhynchus clarkii throughout western North America. We hypothesized that when brook trout replace cutthroat trout, they exhibit greater density, biomass, and annual production than were achieved by the cutthroat trout before replacement. We tested this hypothesis in five paired streams in the Teton River drainage, Idaho, over a 4-year period. Streams were paired based on similar habitat characteristics and the presence of brook trout or cutthroat trout as the dominant species. Fish were surveyed annually by means of multipass depletion along representative stream reaches. Consistent with our hypothesis, brook trout exhibited 2.4 times greater density, 1.7 times greater biomass, and 2.5 times higher annual product...

Incorporating food web dynamics into ecological restoration: a modeling approach for river ecosystems

Restoration is frequently aimed at the recovery of target species, but also influences the larger food web in which these species participate. Effects of restoration on this broader network of organisms can influence target species both directly and indirectly via changes in energy flow through food webs. To help incorporate these complexities into river restoration planning, we constructed a model that links river food web dynamics to in-stream physical habitat and riparian vegetation conditions. We present an application of the model to the Methow River, Washington, USA, a location of on-going restoration aimed at recovering salmon. Three restoration strategies were simulated: riparian vegetation restoration, nutrient augmentation via salmon carcass addition, and side channel reconnection. We also added populations of nonnative aquatic snails and fish to the modeled food web to explore how changes in food web structure mediate responses to restoration. Simulations suggest that side channel reconnection may be a better strategy than carcass addition and vegetation planting for improving conditions for salmon in this river segment. However, modeled responses were strongly sensitive to changes in the structure of the food web. The addition of nonnative snails and fish modified pathways of energy through the food web, which negated restoration improvements. This finding illustrates that forecasting responses to restoration may require accounting for the structure of food webs, and that changes in this structure, as might be expected with the spread of invasive species, could compromise restoration outcomes. Unlike habitat-based approaches to restoration assessment that focus on the direct effects of physical habitat conditions on single species of interest, our approach dynamically links the success of target organisms to the success of competitors, predators, and prey. By elucidating the direct and indirect pathways by which restoration affects target species, dynamic food web models can improve restoration planning by fostering a deeper understanding of system connectedness and dynamics.

Thermal Regimes, Nonnative Trout, and Their Influences on Native Bull Trout in the Upper Klamath River Basin, Oregon

AbstractThe occurrence of fish species may be strongly influenced by a stream’s thermal regime (magnitude, frequency, variation, and timing). For instance, magnitude and frequency provide information about sublethal temperatures, variability in temperature can affect behavioral thermoregulation and bioenergetics, and timing of thermal events may cue life history events, such as spawning and migration. We explored the relationship between thermal regimes and the occurrences of native Bull Trout Salvelinus confluentus and nonnative Brook Trout Salvelinus fontinalis and Brown Trout Salmo trutta across 87 sites in the upper Klamath River basin, Oregon. Our objectives were to associate descriptors of the thermal regime with trout occurrence, predict the probability of Bull Trout occurrence, and estimate upper thermal tolerances of the trout species. We found that each species was associated with a different suite of thermal regime descriptors. Bull Trout were present at sites that were cooler, had fewer high-t...

Response of ecosystem metabolism to low densities of spawning Chinook Salmon

Marine derived nutrients delivered by large runs of returning salmon are thought to subsidize the in situ food resources that support juvenile salmon. In the Pacific Northwest, USA, salmon have declined to <10% of their historical abundance, with subsequent declines of marine derived nutrients once provided by large salmon runs. We explored whether low densities (<0.001 spawners/m2) of naturally spawning Chinook Salmon (Oncorhynchus tshawytscha) can affect ecosystem metabolism. We measured gross primary production (GPP) and ecosystem respiration (ER) continuously before, during, and after salmon spawning. We compared downstream reaches with low densities of spawning salmon to upstream reaches with fewer or no spawners in 3 mid-sized (4th-order) rivers in northern Washington. In addition, we measured chemical, physical, and biological factors that may be important in controlling rates of GPP and ER. We observed that low densities of spawning salmon can increase GPP by 46% during spawning, but values quickly return to those observed before spawning. No difference in ER was observed between up- and downstream reaches. Based on our results, salmon density, temperature, and the proximity to salmon redds were the most important factors controlling rates of GPP, whereas temperature was most important for ER. These results suggest that even at low spawning densities, salmon can stimulate basal resources that may propagate up the food web. Understanding how recipient ecosystems respond to low levels of marine derived nutrients may inform nutrient augmentation studies aimed at enhancing fish populations.