OnlineOpen: This article is available free online at www.blackwell-synergy.com SUMMARY 1. Pacific salmon and steelhead once contributed large amounts of marine-derived carbon, nitrogen and phosphorus to freshwater ecosystems in the Pacific Northwest of the United States of America (California, Oregon, Washington and Idaho). Declines in historically abundant anadromous salmonid populations represent a significant loss of returning nutrients across a large spatial scale. Recently, a manufactured salmon carcass analogue was developed and tested as a safe and effective method of delivering nutrients to freshwater and linked riparian ecosystems where marine-derived nutrients have been reduced or eliminated. 2. We compared four streams: two reference and two treatment streams using salmon carcass analogue(s) (SCA) as a treatment. Response variables measured included: surface streamwater chemistry; nutrient limitation status; carbon and nitrogen stable isotopes; periphyton chlorophyll a and ash-free dry mass (AFDM); macroinvertebrate density and biomass; and leaf litter decomposition rates. Within each stream, upstream reference and downstream treatment reaches were sampled 1 year before, during, and 1 year after the addition of SCA. 3. Periphyton chlorophyll a and AFDM and macroinvertebrate biomass were significantly higher in stream reaches treated with SCA. Enriched stable isotope (d 15 N) signatures were observed in periphyton and macroinvertebrate samples collected from treatment reaches in both treatment streams, indicating trophic transfer from SCA to consumers. Densities of Ephemerellidae, Elmidae and Brachycentridae were significantly higher in treatment reaches. Macroinvertebrate community composition and structure, as measured by taxonomic richness and diversity, did not appear to respond significantly to SCA treatment. Leaf breakdown rates were variable among treatment streams: significantly higher in one stream treatment reach but not the other. Salmon carcass analogue treatments had no detectable effect on measured water chemistry variables. 4. Our results suggest that SCA addition successfully increased periphyton and macroinvertebrate biomass with no detectable response in streamwater nutrient concentrations. Correspondingly, no change in nutrient limitation status was detected based on dissolved inorganic nitrogen to soluble reactive phosphorus ratios (DIN/SRP) and nutrient-diffusing substrata experiments. Salmon carcass analogues appear to increase freshwater productivity. 5. Salmon carcass analogues represent a pathogen-free nutrient enhancement tool that mimics natural trophic transfer pathways, can be manufactured using recycled fish

Anadromous fishes represent an important ecosystem linkage between marine and inland aquatic and terrestrial habitats. These fishes carry organic matter and marine‐derived nutrient (MDN) subsidies across a vast landscape, often with profound influences on recipient ecosystem food web structure and function. In the Columbia River basin, century‐long declines in the abundance of anadromous fish populations have focused attention on potential mitigation efforts to address MDN deficits. In this study, we evaluate components of the stream food web response (periphyton, macroinvertebrate, and fish) to pasteurized salmon carcass analog (SCA) treatments in 15 streams across the Columbia River basin. Periphyton standing crop, macroinvertebrate density, and salmonid fish growth rates and stomach fullness measures increased following the addition of SCA. We found no significant change in dissolved nutrient concentrations after treatment, suggesting that biological demand exceeded supply. Nitrogen stable isotope signatures confirmed trophic transfer from SCA to lower trophic levels but were noticeably weak in fish tissue samples despite our marked growth and stomach fullness measures. These data indicate that SCA has the potential to increase the productivity of nutrient‐limited freshwater ecosystems and may provide a nutrient mitigation tool in ecosystems where MDNs are severely limited or unavailable.

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Salmon provide an important resource subsidy and linkage between marine and land-based ecosystems. This flow of energy and nutrients is not unidirectional (i.e., upstream only); in addition to passive nutrient export via stream flow, juvenile emigrants actively export nutrients from freshwater environments. In some cases, nutrient export can exceed import. We evaluated nutrient fluxes in streams across central Idaho, USA, using Chinook salmon (Oncorhynchus tshawytscha) adult escapement and juvenile production data from 1998 to 2008. We found in the majority of stream-years evaluated, adults imported more nutrients than progeny exported; however, in 3% of the years, juveniles exported more nutrients than their parents imported. On average, juvenile emigrants exported 22% ± 3% of the nitrogen and 30% ± 4% of the phosphorus their parents imported. This relationship was density-dependent and nonlinear; during periods of low adult abundance, juveniles were larger and exported up to 194% and 268% of parental nitrogen and phosphorus inputs, respectively. We highlight minimum escapement thresholds that appear to (i) maintain consistently positive net nutrient flux and (ii) reduce the average proportional rate of export across study streams. Our results suggest a state shift occurs when adult spawner abundance falls below a threshold to a point where the probability of juvenile nutrient exports exceeding adult imports becomes increasingly likely.

In 1991, Snake River sockeye salmon Oncorhynchus nerka were listed as endangered. The Sawtooth Valley Project was initiated to conserve and rebuild sockeye salmon populations that historically spawned and reared in five Sawtooth Valley lakes designated as critical habitat in central Idaho. We evaluated smolt survival of sockeye salmon that were stocked as parr into Redfish, Pettit, and Alturas lakes. Smolt travel time, residuals of Salmon River discharge and smolt travel time, and specific growth rate of parr explained 58% of the variation in smolt survival in a multiple‐regression model. Smolt survival was inversely related to smolt travel time, and travel time was negatively correlated with mean May discharge in the Salmon River at Salmon, Idaho. We were particularly interested in the relationships between smolt survival and parr size at release, smolt size at migration, and parr growth rate. Smolt survival from nursery lakes to Lower Granite Dam was negatively correlated with mean parr size at release and mean smolt size at emigration. Smaller parr and smolts survived better. Smolt survival to the dam was correlated with parr growth rates in the three lakes combined; the relationship was nonlinear. Smolt survival increased as specific growth rate increased up to 0.06% per day, but further increases in growth rate were associated with reductions in survival. Absolute growth rates of parr were negatively related to parr weight at release. Smaller individuals grew faster than larger individuals, apparently as a result of size‐dependent metabolic demand and prey availability. The relationships between smolt survival and parr size, smolt size, and parr growth rate differed among lakes. These data suggest that successful migrants must at least maintain weight during the winter preceding migration and that the stocking of smaller parr with lower metabolic demand may be preferable to stocking larger parr when forage is limited.Received November 18, 2010; accepted June 1, 2011

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Dramatic declines of Pacific salmon (Oncorhynchus spp.) populations have decreased delivery of marine-derived material to Pacific Northwest streams where juvenile salmon reside. Managers use artificial nutrient additions to increase juvenile salmon growth and survival and typically assume nutrient-driven increases in biofilm production are an important pathway by which nutrients become available to higher trophic levels. To evaluate how biofilms respond to additions of salmon carcass analog, a pasteurized, processed form of nutrient mitigation materials, we quantified biofilm nutrient limitation, benthic and whole-stream metabolism, and biofilm standing crops before and following experimental additions in tributaries of the Salmon River, Idaho, USA. Biofilm nutrient limitation did not change and standing crop did not increase in response to analog additions at two different levels (low, 30 g·m −2 ; or high, 150 g·m −2 ) within 1 month of addition. In contrast, whole-stream and benthic primary productivity and respiration increased in a high-analog treated segment, but did not increase in a low-analog treated segment. Together, our results suggest that metabolism may be a more appropriate tool for assessing the ecosystem effects of nutrient additions than biofilm standing crop or nutrient limitation, which are constrained by a variety of abiotic and biotic factors like hydrology and grazing.Résumé : D'importantes diminutions des populations de saumons du Pacifique (Oncorhynchus spp.) ont entraîné une réduction des apports de matières d'origine marine dans les cours d'eau du Pacific Northwest où résident des saumons juvéniles. Les gestionnaires utilisent l'ajout de nutriments artificiels pour accroître la croissance et la survie des saumons juvéniles, présumant généralement que l'augmentation de la production de biofilm induite par les nutriments constitue une importante voie de transfert de nutriments vers des niveaux trophiques plus élevés. Afin d'évaluer la réaction des biofilms à l'ajout d'analogues de carcasse de saumon, une forme transformée et pasteurisée de matière utilisée pour compenser le manque de nutriments, nous avons quantifié la limitation des nutriments de biofilms, le métabolisme benthique et à l'échelle du cours d'eau et la biomasse de biofilm disponible avant et après des ajouts expérimentaux dans des affluents de la rivière Salmon (Idaho, États-Unis). La limitation des nutriments de biofilms n'a pas changé et la biomasse de biofilm disponible n'a pas augmenté en réaction à des ajouts d'analogue à deux taux différents (faible, 30 g·m −2 ; et fort, 150 g·m −2 ) durant le mois suivant ces ajouts. En revanche, la productivité primaire benthique et à l'échelle du cours d'eau et la respiration ont augmenté dans un tronçon ayant reçu un fort ajout d'analogue, mais n'ont pas augmenté dans un tronçon ayant reçu un faible ajout d'analogue. Nos résultats donnent à penser que le métabolisme pourrait être un outil mieux adapté pour évaluer les effets écosystémiques d'ajouts de nutriments que la biomasse de ...