Large predators and biogeochemical hotspots: brown bear (Ursus arctos) predation on salmon alters nitrogen cycling in riparian soils

Two important themes in ecology include the understanding of how interactions among species control ecosystem processes, and how habitats can be connected through transfers of nutrients and energy by mobile organisms. An impressive example of both is the large influx of nutrients and organic matter that anadromous salmon supply to inland aquatic and terrestrial ecosystems and the role of predation by brown bears (Ursus arctos) in transferring these marine-derived nutrients (MDN) from freshwater to riparian habitats. In spite of the recognition that salmon-bear interactions likely play an important role in controlling the flux of MDN from aquatic to riparian habitats, few studies have linked bear predation on salmon to processes such as nitrogen (N) or carbon (C) cycling. We combine landscape-level survey data and a replicated bear-exclosure experiment to test how bear foraging on salmon affects nitrous oxide (N₂O) flux, carbon dioxide (CO₂) flux, and nutrient concentrations of riparian soils. Our results show that bears feeding on salmon increased soil ammonium (NH₄ ⁺) concentrations three-fold and N₂O flux by 32-fold. Soil CO₂ flux, nitrate (NO₃ ⁻), and N transformation differences were negligible in areas where bears fed on salmon. Reference areas without concentrated bear activity showed no detectable change in soil N cycling after the arrival of salmon to streams. Exclosure experiments showed that bear effects on soil nutrient cycles were transient, and soil N processing returned to background conditions within 1 year after bears were removed from the system. These results suggest that recipient ecosystems do not show uniform responses to MDN inputs and highlight the importance of large mobile consumers in generating landscape heterogeneity in nutrient cycles.     

Stable isotope evidence indicates the incorporation into Japanese catchments of marine‐derived nutrients transported by spawning Pacific Salmon

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Effects of Salmon-Derived Nutrients and Habitat Characteristics on Population Densities of Stream-Resident Sculpins

Movement of nutrients across ecosystem boundaries can have important effects on food webs and population dynamics. An example from the North Pacific Rim is the connection between productive marine ecosystems and freshwaters driven by annual spawning migrations of Pacific salmon (Oncorhynchus spp). While a growing body of research has highlighted the importance of both pulsed nutrient subsidies and disturbance by spawning salmon, their effects on population densities of vertebrate consumers have rarely been tested, especially across streams spanning a wide range of natural variation in salmon densities and habitat characteristics. We studied resident freshwater prickly (Cottus asper), and coastrange sculpins (C. aleuticus) in coastal salmon spawning streams to test whether their population densities are affected by spawning densities of pink and chum salmon (O. gorbuscha and O. keta), as well as habitat characteristics. Coastrange sculpins occurred in the highest densities in streams with high densities of spawning pink and chum salmon. They also were more dense in streams with high pH, large watersheds, less area covered by pools, and lower gradients. In contrast, prickly sculpin densities were higher in streams with more large wood and pools, and less canopy cover, but their densities were not correlated with salmon. These results for coastrange sculpins provide evidence of a numerical population response by freshwater fish to increased availability of salmon subsidies in streams. These results demonstrate complex and context-dependent relationships between spawning Pacific salmon and coastal ecosystems and can inform an ecosystem-based approach to their management and conservation.     

Evaluating the consequences of salmon nutrients for riparian organisms: Linking condition metrics to stable isotopes

Stable isotope ratios (δ¹³C and δ¹⁵N) have been used extensively to trace nutrients from Pacific salmon, but salmon transfer more than carbon and nitrogen to stream ecosystems, such as phosphorus, minerals, proteins, and lipids. To examine the importance of these nutrients, metrics other than isotopes need to be considered, particularly when so few studies have made direct links between these nutrients and how they affect riparian organisms. Our study specifically examined δ¹³C and δ¹⁵N of riparian organisms from salmon and non‐salmon streams in Idaho, USA, at different distances from the streams, and examined whether the quality of riparian plants and the body condition of invertebrates varied with access to these nutrients. Overall, quality and condition metrics did not mirror stable isotope patterns. Most notably, all riparian organisms exhibited elevated δ¹⁵N in salmon streams, but also with proximity to both stream types suggesting that both salmon and landscape factors may affect δ¹⁵N. The amount of nitrogen incorporated from Pacific salmon was low for all organisms (<20%) and did not correlate with measures of quality or condition, probably due to elevated δ¹⁵N at salmon streams reflecting historical salmon runs instead of current contributions. Salmon runs in these Idaho streams have been declining, and associated riparian ecosystems have probably seen about a 90% reduction in salmon‐derived nitrogen since the 1950s. In addition, our results support those of other studies that have cautioned that inferences from natural abundance isotope data, particularly in conjunction with mixing models for salmon‐derived nutrient percentage estimates, may be confounded by biogeochemical transformations of nitrogen, physiological processes, and even historical legacies of nitrogen sources. Critically, studies should move beyond simply describing isotopic patterns to focusing on the consequences of salmon‐derived nutrients by quantifying the condition and fitness of organisms putatively using those resources.     

Variation in responses to spawning Pacific salmon among three south-eastern Alaska streams

1. Pacific salmon are thought to stimulate the productivity of the fresh waters in which they spawn by fertilising them with marine‐derived nutrients (MDN). We compared the influence of salmon spawners on surface streamwater chemistry and benthic biota among three south‐eastern Alaska streams. Within each stream, reaches up‐ and downstream of barriers to salmon migration were sampled during or soon after spawners entered the streams. 2. Within streams, concentrations of dissolved ammonium and soluble reactive phosphorus (SRP), abundance of epilithon (chlorophyll a and ash‐free dry mass) and biomass of chironomids were significantly higher in reaches with salmon spawners. In contrast, biomass of the mayflies Epeorus spp. and Rhithrogena spp. was significantly higher in reaches lacking spawners. 3. Among streams, significant differences were found in concentrations of dissolved ammonium, dissolved organic carbon, nitrate and SRP, abundance of epilithon, and the biomass of chironomids and Rhithrogena. These differences did not appear to reflect differences among streams in spawner density, nor the changes in water chemistry resulting from salmon spawners. 4. Our results suggest that the ‘enrichment’ effect of salmon spawners (e.g. increased streamwater nutrient concentrations) was balanced by other concurrent effects of spawners on streams (e.g. sediment disturbance). Furthermore, the collective effect of spawners on lotic ecosystems is likely to be constrained by conditions unique to individual streams, such as temperature, background water chemistry and light attenuation.     

Pacific Salmon,Nutrients, and the Dynamics of Freshwater and Riparian Ecosystems

Pacific salmon (Oncorhynchus spp.) accumulate substantial nutrients in their bodies as they grow to adulthood at sea. These nutrients are carried to predominantly oligotrophic lakes and streams, where they are released during and after spawning. Research over more than 3 decades has shown that the annual deposition of salmon-borne marine-derived nutrients (MD-nutrients) is important for the productivity of freshwater communities throughout the Pacific coastal region. However, the pathways and mechanisms for MD-nutrient transfer and accumulation in freshwater and riparian ecosystems remain virtually unexplored, consequently, there are many uncertainties in this area. This article addresses three related topics. First, we summarize recent advances in our understanding of the linkages among MD-nutrients, freshwater (including riparian) ecosystems, and community dynamics by addressing the importance of MD-nutrients to lakes and streams and by then reviewing large-scale and long-term processes in the atmosphere and ocean that govern variability in salmon populations. Second, we evaluate the validity of the discoveries and their implications for active ecosystem management, noting areas where extrapolation from these results still requires great caution. Finally, we outline five key research issues where additional discoveries could greatly augment our understanding of the processes shaping the structure and dynamics of salmon populations and the characteristics of their freshwater habitat and associated riparian zones. Collectively, the data suggest that the freshwater portion of the salmon production system is intimately linked to the ocean. Moreover, for the system to be sustainable, a holistic approach to management will be required. This holistic approach will need to treat climate cycles, salmon, riparian vegetation, predators, and MD-nutrient flowpaths and feedbacks as an integrated system. Received 3 July 2001; accepted 14 December 2001.     

Impacts of marine-derived nutrients on stream ecosystem functioning

Energy and nutrient subsidies transported across ecosystem boundaries are increasingly appreciated as key drivers of consumer-resource dynamics. As purveyors of pulsed marine-derived nutrients (MDN), spawning salmon are one such cross-ecosystem subsidy to freshwaters connected to the north Pacific. We examined how salmon carcasses influenced detrital processing in an oligotrophic stream. Experimental manipulations of MDN inputs revealed that salmon carcasses indirectly reduced detrital processing in streams through temporarily decoupling the detrital resource-consumer relationship, in which detrital consumers shifted their diet to the high-nutrient resource, i.e. salmon carcasses. The average decomposition rate of alder leaves with salmon carcass addition was significantly lower than that without the carcass, which was associated with lower abundance and biomass of detritivorous Trichoptera on the carcass-treated leaves. There were generally larger in size Trichopteran detritivores on the carcasses than on leaves. These results imply that cross-boundary MDN subsidies indirectly retard the ecosystem processing of leaf litter within the short term, but may enhance those food-limited detritivorous consumers. Because unproductive freshwaters in the Pacific northwest are highly dependent upon the organic matter inputs from surrounding forests, this novel finding has implications for determining conservation and management strategies of salmon-related aquatic ecosystems, in terms of salmon habitat protection and fisheries exploitation.     

Alaskan brown bears (Ursus arctos) aggregate and display fidelity to foraging neighborhoods while preying on Pacific salmon along small streams

The interaction between brown bears (Ursus arctos) and Pacific salmon (Oncorhynchus spp.) is important to the population dynamics of both species and a celebrated example of consumer‐mediated nutrient transport. Yet, much of the site‐specific information we have about the bears in this relationship comes from observations at a few highly visible but unrepresentative locations and a small number of radio‐telemetry studies. Consequently, our understanding of brown bear abundance and behavior at more cryptic locations where they commonly feed on salmon, including small spawning streams, remains limited. We employed a noninvasive genetic approach (barbed wire hair snares) over four summers (2012–2015) to document patterns of brown bear abundance and movement among six spawning streams for sockeye salmon, O. nerka, in southwestern Alaska. The streams were grouped into two trios on opposite sides of Lake Aleknagik. Thus, we predicted that most bears would forage within only one trio during the spawning season because of the energetic costs associated with swimming between them or traveling around the lake and show fidelity to particular trios across years because of the benefits of familiarity with local salmon dynamics and stream characteristics. Huggins closed‐capture models based on encounter histories from genotyped hair samples revealed that as many as 41 individuals visited single streams during the annual 6‐week sampling season. Bears also moved freely among trios of streams but rarely moved between these putative foraging neighborhoods, either during or between years. By implication, even small salmon spawning streams can serve as important resources for brown bears, and consistent use of stream neighborhoods by certain bears may play an important role in spatially structuring coastal bear populations. Our findings also underscore the efficacy of noninvasive hair snagging and genetic analysis for examining bear abundance and movements at relatively fine spatial and temporal scales.     

Chemosensory responses of juvenile Coho salmon, Oncorhynchus kisutch, Dolly Varden, Salvelinus malma, and sculpins (Cottus spp.) to eggs and other tissues from adult Pacific salmon

The carcasses of semelparous Pacific salmon (Oncorhynchus spp.) provide nutrients that enter aquatic ecosystems by various pathways, including direct consumption of tissue by fishes. Salmonids and other species frequently eat eggs and other tissues from dead salmon but the roles of vision and olfaction are unclear, as is the relative attraction to different tissues. Accordingly, we conducted a series of in situ experiments using minnow traps in two natural streams in Alaska to test the relative roles of chemosensory and visual cues in attraction of fishes to eggs from adult Pacific salmon, and then compared catch rates of traps baited with eggs, muscle, liver, and testis. Experiments indicated that chemical traces were necessary and sufficient to attract juvenile Dolly Varden (Salvelinus malma), coho salmon (Oncorhynchus kisutch), and sculpins (Cottus spp.) into traps. Combining both sites, 70 salmonids and 19 sculpins were trapped using visual and chemical cues, and 53 and 21, respectively, for traps with only chemical cues. Traps with only the sight of eggs caught no salmonids and only 5 sculpins, comparable to empty control traps. In addition, eggs were markedly more attractive than the other tissues, trapping 68?% of the salmonids and 69?% of the sculpins, compared to 14?% and 15?% for muscle tissue, 12?% and 11?% for liver, and 6?% and 5?% for the testis. Visual cues undoubtedly play a role in egg consumption in streams, but these experiments indicated a very important role of chemical traces in attracting fish to the vicinity of the eggs, and selective attraction of eggs over other salmon tissues.     

Forest succession and prey availability influence the strength and scale of terrestrial‐aquatic linkages in a headwater salamander system

1. Trophic linkages between terrestrial and aquatic ecosystems are common and sensitive to disruption. However, there is little information on what causes variation in the strength and spatial scale of these linkages. 2. In the highly aquatic adults of the headwater salamander Gyrinophilus porphyriticus (family Plethodontidae), use of terrestrial prey decreases along a gradient from early‐ to late‐successional riparian forests. To understand the cause of this relationship, we tested the predictions that (i) terrestrial prey abundance is lower in late‐successional forests, and (ii) G. porphyriticus adults cannot move as far from the stream to forage in late‐successional forests, thus limiting access to terrestrial prey. 3. We established 100‐m long study reaches on six headwater streams in the Hubbard Brook Experimental Forest, New Hampshire. Three reaches were in early‐successional forests and three were in late‐successional forests. We conducted pitfall trapping for invertebrate prey in June and July of 2005, with three traps at 0, 2, 5 and 10 m from the stream at each reach. In June, July and August of 2004 and 2005, nighttime salamander surveys were conducted at each reach along ten, 10‐m long by 2.5‐m wide transects perpendicular to the stream. 4. Abundance of terrestrial prey was consistently lower in late‐successional forests, suggesting that consumption of terrestrial prey by G. porphyriticus is affected by prey abundance. Contrary to our prediction, G. porphyriticus adults moved farther from the stream in late‐successional forests, suggesting that habitat conditions in late‐successional forests do not limit movement away from the stream, and that lower abundances of terrestrial prey in these forests may cause salamanders to move farther from streams. 5. Our results provide novel insight on the extent of terrestrial habitat use by G. porphyriticus. More broadly, these results indicate that major habitat gradients, such as forest succession, can affect the strength and scale of terrestrial‐aquatic linkages. Application of this insight to the design of vegetation buffers along headwater streams would have widespread benefits to freshwater ecosystems.     

Barriers to salmon migration impact body condition,offspring size,and life history variation in an avian consumer

The impacts of barriers, such as dams, and associated stream effects on the migration of salmon have been well documented, but there is much less information on consequences for terrestrial vertebrates in these freshwater systems. Salmon themselves provide food resources to higher trophic levels and deposit rich marine‐derived nutrients (MDN) that benefit the base of freshwater food webs. We studied a higher‐order terrestrial consumer of aquatic prey, the American dipper Cinclus mexicanus, in a riparian food web, to test the hypothesis that barriers to salmon migration negatively impact body condition, and ultimately life history. We predicted that, compared to dippers breeding above barriers, those in areas with salmon would receive MDN and thus be in better condition, have greater breeding effort (frequency of multiple brood attempts), produce larger offspring in better condition, be more likely to maintain year‐round territories, and have higher annual survival. Using stable isotopes as signatures of MDN, we found that ¹³C and ¹⁵N were more enriched in dippers below barriers, confirming access to MDN. Female dippers below barriers had greater mass corrected for body size, were more likely to attempt multiple broods within a season and produced larger female offspring. Furthermore, dippers below barriers were more likely to maintain year‐round territories and had higher annual survival. Our results demonstrate how river barriers such as dams influence vital rates and life histories of higher‐order consumers.     

Diverse foraging opportunities drive the functional response of local and landscape-scale bear predation on Pacific salmon

The relationship between prey abundance and predation is often examined in single habitat units or populations, but predators may occupy landscapes with diverse habitats and foraging opportunities. The vulnerability of prey within populations may depend on habitat features that hinder predation, and increased density of conspecifics in both the immediate vicinity and the broader landscape. We evaluated the relative effects of physical habitat, local, and neighborhood prey density on predation by brown bears on sockeye salmon in a suite of 27 streams using hierarchical Bayesian functional response models. Stream depth and width were inversely related to the maximum proportion of salmon killed, but not the asymptotic limit on total number killed. Interannual variation in predation was density dependent; the number of salmon killed increased with fish density in each stream towards an asymptote. Seven streams in two geographical groups with ≥23 years of data in common were then analyzed for neighborhood density effects. In most (12 of 18) cases predation in a stream was reduced by increasing salmon abundance in neighboring streams. The uncertainty in the estimates for these neighborhood effects may have resulted from interactions between salmon abundance and habitat that influenced foraging by bears, and from bear behavior (e.g., competitive exclusion) and abundance. Taken together, the results indicated that predator–prey interactions depend on density at multiple spatial scales, and on habitat features of the surrounding landscape. Explicit consideration of this context dependency should lead to improved understanding of the ecological impacts of predation across ecosystems and taxa.     

Implications of water column ammonium uptake and regeneration for the nitrogen budget in temperate, eutrophic Missisquoi Bay, Lake Champlain (Canada/USA)

Invasion by exotic trees into riparian areas has the potential to impact aquatic systems. We examined the effects of the exotic Salix fragilis (crack willow) on the structure and functioning of small streams in northern Patagonian Andes via a field survey of benthic invertebrates and leaf litter and an in situ experiment. We compared leaf decomposition of the native Ochetophila trinervis (chacay) and S. fragilis in reaches dominated by native vegetation versus reaches dominated by crack willow. We hypothesized that S. fragilis affects the quality of leaf litter entering the streams, changing the aquatic biota composition and litter decomposition. Our study showed that crack willow leaves decomposed slower than chacay, likely related to leaf properties (i.e., leaf toughness). Benthic leaf litter mass was similar between the two riparian vegetation types, though in stream reaches dominated by crack willow, leaves of this species represented 82% of the total leaf litter. Benthic invertebrate abundance and diversity were similar between reaches but species composition differed. Our study found little evidence for strong impacts of crack willow on those small streams. Further studies on other aspects of ecosystem functioning, such as primary production, would enhance our understanding of the impacts of crack willow on Patagonian streams.     

Effects of habitat features on size-biased predation on salmon by bears

Predators can drive trait divergence among populations of prey by imposing differential selection on prey traits. Habitat characteristics can mediate predator selectivity by providing refuge for prey. We quantified the effects of stream characteristics on biases in the sizes of spawning salmon caught by bears (Ursus arctos and U. americanus) on the central coast of British Columbia, Canada by measuring size-biased predation on spawning chum (Oncorhynchus keta) and pink (O. gorbuscha) salmon in 12 streams with varying habitat characteristics. We tested the hypotheses that bears would catch larger than average salmon (size-biased predation) and that this bias toward larger fish would be higher in streams that provide less protection to spawning salmon from predation (e.g., less pools, wood, undercut banks). We then we tested for how such size biases in turn translate into differences among populations in the sizes of the fish. Bears caught larger-than-average salmon as the spawning season progressed and as predicted, this was most pronounced in streams with fewer refugia for the fish (i.e., wood and undercut banks). Salmon were marginally smaller in streams with more pronounced size-biased predation but this predictor was less reliable than physical characteristics of streams, with larger fish in wider, deeper streams. These results support the hypothesis that selective forces imposed by predators can be mediated by habitat characteristics, with potential consequences for physical traits of prey.     

Fish abundance and habitat relationships in forest and grassland streams, northern Hokkaido, Japan

The relationships between habitat variables and population densities of masu salmon (Oncorhynchus masou), rosyface dace (Leuciscus ezoe), Siberian stone loach (Noemacheilus barbatulus) and wrinklehead sculpin (Cottus nozawae) were examined by data collected at 55 reaches in forest and grassland streams in northern Hokkaido, Japan. Regression analysis suggested that salmon and dace densities were affected by water temperature (negative for salmon, positive for dace) and structural habitat factors (woody debris for salmon, pools for dace). Salmon density was higher in forest reaches than in grassland reaches, whereas dace density was higher in grassland reaches, suggesting that the removal of riparian forest had raised water temperature and allowed upstream invasions by dace. In contrast to salmon and dace, neither the density of loach nor sculpin differed between the forest and grassland reaches. For their densities, a negative effect of each on the other was most important, suggesting a strong effect of interspecific competition between loach and sculpin on their distributions. However, regression models also suggested that substrate heterogeneity mediated the outcome of their interspecific competition. On the basis of the results, a scenario is predicted for a fish-assemblage change with a typical land-development process in Hokkaido, and the importance of leaving or restoring riparian buffer for conservation and restoration of stream habitat is emphasized.     

Do non-native Platanus hybrida riparian plantations affect leaf litter decomposition in streams?

In forest headwater streams where the riparian canopy limits autochthonous primary production, leaf litter decomposition is a key process controlling nutrient and carbon cycling. Any alteration of the riparian vegetation may influence litter decomposition and detrital food webs. We evaluated the effect of non-native Platanus hybrida riparian plantations on leaf litter decomposition in Mediterranean streams. The experiment was conducted in six headwater streams; three lined by native riparian vegetation and three crossing P. hybrida plantations. We have characterized the processing rates of alder leaves and the assemblages of shredder macroinvertebrates and fungi. Litter decomposition was significantly faster in the P. hybrida than in the reference streams. Although the dissolved inorganic nitrogen concentration was higher in P. hybrida, no significant effect was observed in decomposition rates. Differences in decomposition rates reflected the macroinvertebrate and shredder colonization in alder litter, with higher abundance and richness in the P. hybrida streams. However, aquatic hyphomycete sporulation rate was higher in reference streams, suggesting that the variation in decomposition rates is a direct consequence of shredder abundance. Our findings support part of the substrate quality-matrix quality (SMI) hypothesis, which expects that high-quality litter will show increased decomposition rates in a low-quality litter matrix.     

Quantitative Links Between Pacific Salmon and Stream Periphyton

Species’ impacts on primary production can have strong ecological consequences. In freshwater ecosystems, Pacific salmon (Oncorhynchus spp.) may influence stream periphyton through substrate disturbance during spawning and nutrient subsidies from senescent adults. The shape of relationships between the abundance of spawning salmon and stream periphyton, as well as interactions with environmental variables, are incompletely understood and may differ across the geographic range of salmon. We examined these relationships across 24 sockeye salmon (Oncorhynchus nerka) spawning streams in north-central British Columbia, Canada. The influence of salmon abundance and environmental variables (temperature, light, dissolved nutrients, water velocity, watershed size, and invertebrate grazer abundance) on post-spawning periphyton abundance and nitrogen stable isotope signatures, which can indicate the uptake of salmon nitrogen, was evaluated using linear regression models and Akaike Information Criterion. Periphyton nitrogen stable isotope signatures were best described by a positive log-linear relationship with an upstream salmon abundance metric that includes salmon from earlier years. This suggests the presence of a nutrient legacy. In contrast, periphyton abundance was negatively related to the spawning-year salmon density, which likely results from substrate disturbance during spawning, and positively related to dissolved soluble reactive phosphorus prior to spawning, which may indicate phosphorus limitation in the streams. These results suggest that enrichment from salmon nutrients does not always translate into elevated periphyton abundance. This underscores the need to directly assess the outcome of salmon impacts on streams rather than extrapolating from stable isotope evidence for the incorporation of salmon nutrients into food webs.     

Keystone Interactions: Salmon and Bear in Riparian Forests of Alaska

The term “keystone species” is used to describe organisms that exert a disproportionately important influence on the ecosystems in which they live. Analogous concepts such as “keystone mutualism” and “mobile links” illustrate how, in many cases, the interactions of two or more species produce an effect greater than that of any one species individually. Because of their role in transporting nutrients from the ocean to river and riparian ecosystems, Pacific salmon (Oncorhynchus spp.) and brown bear (Ursus arctos) have been described as keystone species and mobile links, although few data are available to quantify the importance of this interaction relative to other nutrient vectors. Application of a mass balance model to data from a southwestern Alaskan stream suggests that nitrogen (N) influx to the riparian forest is significantly increased in the presence of both salmon and bear, but not by either species individually. The interactions of salmon and bear may provide up to 24% of riparian N budgets, but this percentage varies in time and space according to variations in salmon escapement, channel morphology and watershed vegetation characteristics, suggesting interdependence and functional redundancy among N sources. These findings illustrate the complexity of interspecific interactions, the importance of linkages across ecosystem boundaries and the necessity of examining the processes and interactions that shape ecological communities, rather than their specific component parts.     

Influence of marine-derived nutrients from spawning salmon on aquatic insect communities in southeast Alaskan streams

The objective of this study was to evaluate the influence of the nutrient transfer system between anadromous salmon and aquatic insect communities across multiple, natural stream systems. Between 2000 and 2002, we sampled seven streams in southeast Alaska, seasonally. Of the seven study streams, four received large annual salmon runs (high-run streams), and three were no-run streams. All the streams selected had a natural waterfall barrier to salmon, providing an upstream control reach for each study stream. Insect density, biomass, richness, diversity and functional feeding groups were analyzed before, during and after the fall salmon run in each stream section (i.e. above and below the barrier) of the seven study streams between 2001 and 2002. Results showed that diversity and richness were similar across stream sections and run size within each period, except for during the run when both were significantly lower in downstream sections of high-run streams. Functional feeding group patterns showed higher abundance and biomass of collector–gatherers and shredders during the post spawning, carcass decomposition period. High-run streams had upstream sections with greater abundance and biomass of mayflies (dominated by Baetidae, Heptageniidae and Ephemerellidae) during the run, and downstream sections with greater abundance and biomass of dipterans (dominated by Chironomidae). This study suggests that the often published positive relationship between MDN and stream insect abundance and biomass may only exist for certain taxa, primarily chironomid midges.     

Salmon-derived nitrogen delivery and storage within a gravel bed: Sediment and water interactions

Post-spawning salmon carcasses are broadly recognized as a source of organic matter- and marine-derived nutrients (MDN) in Pacific salmon streams, but MDN delivery and retention processes are not well understood. Recent studies emphasize the interaction of inorganic particulate matter and salmon organic matter, through flocculation, as a delivery mechanism for MDN to the streambed. This study builds upon previous flocculation studies to look at nitrogen delivery and storage within the gravel bed of a re-circulating flume. Findings indicate that nitrogen storage in surface and interstitial water is lower than sediment-associated nitrogen. Flocculation of salmon organic matter and inorganic sediment is presented as a delivery mechanism in spawning and post-spawning periods that helps to maintain ecological productivity within Pacific salmon streams. Based on these findings it is recommended that salmon enhancement activities should include leaving post-spawn carcasses in-stream and that fertilization programs should consider flocculation processes to increase nutrient delivery to the streambed.