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1. Streams and their adjacent riparian zones are closely linked by reciprocal flows of invertebrate prey. We review characteristics of these prey subsidies and their strong direct and indirect effects on consumers and recipient food webs. 2. Fluxes of terrestrial invertebrates to streams can provide up to half the annual energy budget for drift-feeding fishes such as salmonids, despite the fact that input occurs principally in summer. Inputs appear highest from closed-canopy riparian zones with deciduous vegetation and vary markedly with invertebrate phenology and weather. Two field experiments that manipulated this prey subsidy showed that it affected both foraging and local abundance of stream fishes. 3. Emergence of adult insects from streams can constitute a substantial export of benthic production to riparian consumers such as birds, bats, lizards, and spiders, and contributes 25-100% of the energy or carbon to such species. Emergence typically peaks in early summer in the temperate zone, but also provides a low-level flux from autumn to spring in ice-free streams. This flux varies with in-stream productivity, and declines exponentially with distance from the stream edge. Some predators aggregate near streams and forage on these prey during periods of peak emergence, whereas others rely on the lower subsidy from autumn through spring when terrestrial prey are scarce. Several field experiments that manipulated this subsidy showed that it affected the short-term behaviour, growth, and abundance of terrestrial consumers. 4. Reciprocal prey subsidies also have important indirect effects on both stream and riparian food webs. Theory predicts that allochthonous prey should increase density of subsidised predators, thereby increasing predation on in situ prey and causing a negative indirect effect via apparent competition. However, short-term experiments have produced either positive or negative indirect effects. These contrasting results may be due to characteristics of the subsidies and individual consumers, but could also result from differences in experimental designs. 5. New study approaches are needed to better determine the direct and indirect effects of reciprocal prey subsidies. Experiments coupled with comparative research will be required to measure their effects on individual consumer fitness and population demographics. Future work should investigate whether reciprocal prey fluxes stabilise linked stream-riparian ecosystems, explore how landscape context affects the magnitude and importance of subsidies, and determine how impacts of human disturbance can propagate between streams and riparian zones via these trophic linkages. 201 reciprocal connections is helping to define a more holistic perspective of catchments, and has the potential to shape new directions for ecology in general.

Habitat alteration and biotic invasions are the two leading causes of global environmental change and biodiversity loss. Recent innovative experiments have shown that habitat disturbance can have drastic effects that cascade to adjacent ecosystems by altering the flow of resource subsidies from donor systems. Likewise, exotic species invasions could alter subsidies and affect distant food webs, but very few studies have tested this experimentally. Here we report evidence from a large-scale field experiment in northern Japan that invasion of nonnative rainbow trout (Oncorhynchus mykiss) interrupted reciprocal flows of invertebrate prey that drove stream and adjacent riparian forest food webs. Rainbow trout usurped terrestrial prey that fell into the stream, causing native Dolly Varden charr (Salvelinus malma) to shift their foraging to insects that graze algae from the stream bottom. This indirectly increased algal biomass, but also decreased biomass of adult aquatic insects emerging from the stream to the forest. In turn, this led to a 65% reduction in the density of riparian-specialist spiders in the forest. Thus, species invasions can interrupt flows of resources between interconnected ecosystems and have effects that propagate across their boundaries, effects that may be difficult to anticipate without in-depth understanding of food web relationships.

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.

The distribution and abundance of bull trout (Salvelinus confluentus) spawning were affected by geomorphology and hyporheic groundwater - stream water exchange across multiple spatial scales in streams of the Swan River basin, northwestern Montana. Among spawning tributary streams, the abundance of bull trout redds increased with increased area of alluvial valley segments that were longitudinally confined by geomorphic knickpoints. Among all valley segment types, bull trout redds were primarily found in these bounded alluvial valley segments, which possessed complex patterns of hyporheic exchange and extensive upwelling zones. Bull trout used stream reaches for spawning that were strongly influenced by upwelling. However, within these selected reaches, bull trout redds were primarily located in transitional bedforms that possessed strong localized downwelling and high intragravel flow rates. The changing relationship of spawning habitat selection, in which bull trout selected upwelling zones at one spatial scale and downwelling zones at another spatial scale, emphasizes the importance of considering multiple spatial scales within a hierarchical geomorphic context when considering the ecology of this species or plans for bull trout conservation and restoration.

Abstract. 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 foodweb structure can affect energy flow and interaction strengths, and we show that these changes have consequences for food-web function and response to perturbations.

Physical characteristics of riverine habitats, such as large wood abundance, pool geometry and abundance, riparian vegetation cover, and surface flow conditions, have traditionally been thought to constrain fish production in these ecosystems. Conversely, the role of food resources (quantity and quality) in controlling fish production has received far less attention and consideration, though they can also be key productivity drivers. Traditional freshwater food web illustrations have typically conveyed the notion that most fish food is produced within the local aquatic habitat itself, but the concepts and model we synthesize in this article show that most fish food comes from external or very distant sources-including subsidies from marine systems borne from adult returns of anadromous fishes, from fishless headwater tributaries that transport prey to downstream fish, and from adjacent streamside vegetation and associated habitats. The model we propose further illustrates how key trophic pathways and food sources vary through time and space throughout watersheds. Insights into how food supplies affect fishes can help guide how we view riverine ecosystems, their structure and function, their interactions with marine and terrestrial systems, and how we manage natural resources, including fish, riparian habitats, and forests.

An experiment in >1000 river and riparian sites found spatial patterns and controls of carbon processing at the global scale.