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.     

Stream food web response to a salmon carcass analogue addition in two central Idaho, U.S.A. streams

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 (δ¹⁵N) 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 products, and is easily transported; however, salmon carcass analogues should not be viewed as a replacement for naturally spawning salmon and the important ecological processes they provide.     

Trophic ecology of Pacific salmon (<Emphasis Type="Italic">Oncorhynchus</Emphasis> spp.) in the ocean: a synthesis of stable isotope research

Increasing interest in the marine trophic dynamics of Pacific salmon has been motivated by the recognition of their sensitivity to changing climate and to the competitive effects of hatchery fish on wild stocks. It has become more common to use stable isotopes to supplement traditional diet studies of salmon in the ocean; however, there have been no integrated syntheses of these data to determine whether stable isotope analyses support the existing conventional wisdom of feeding strategies of the Pacific salmon. We performed a meta-analysis of stable isotope data to examine the extent of trophic partitioning among five species of Pacific salmon during their marine lives. Pink, sockeye, and chum salmon showed very high overlap in resource use and there was no consistent evidence for chum relying on alternative food webs dominated by gelatinous zooplankton. δ¹⁵N showed that Chinook and coho salmon fed at trophic levels higher than the other three species. In addition, these two species were distinctly enriched in ¹³C, suggesting more extensive use of coastal food webs compared to the more depleted (pelagic) signatures of pink, sockeye, and chum salmon. This paper presents the first synthesis of stable isotope work on Pacific salmon and provides δ¹⁵N and δ¹³C values applicable to research on the fate of the marine derived nutrients these organisms transport to freshwater and riparian ecosystems.     

Mobile scavengers create hotspots of freshwater productivity

Adjacent communities and ecosystems often differ in underlying productivity but are connected by flows of nutrients, energy, and matter. Pacific salmon ( Oncorhynchus spp.) transport substantial quantities of nutrients from marine ecosystems to coastal freshwater habitats when they return to spawn and die. Nutrients from their carcasses are initially concentrated in spawning streams and lakes, but are subsequently dispersed by abiotic (floods, hyporheic flow) and biotic processes (predators and scavengers). In southwest Alaska, mobile avian scavengers (gulls; Larus spp.) breed on small islands within salmon nursery lakes and consume large quantities of spawning salmon during the chick-rearing period. However the role of birds as vectors of salmon-derived nutrients remains unknown. We examined how gulls – by transporting salmon tissues to their chicks – create hotspots of biological productivity in the aquatic habitats surrounding their nesting colonies. We found that algal production was ∼10× higher at islands with high gull densities compared to islands without nesting gulls, but was concentrated within 40 m of island shorelines. Carbon stable isotopes (δ¹³C) confirmed that gulls enhance primary production in local benthic communities and demonstrated that this production was transferred up the food web to grazers (snails) and carnivores (blackfish). Nitrogen stable isotopes (δ¹⁵N) confirmed that salmon dominated the diet of gulls and that nutrients from gull guano were incorporated into algae and passed up the food web. By relocating and concentrating salmon-derived nutrients into new and distant locations, gulls alter and magnify production in local aquatic communities. We offer the first evidence that the avian community can move salmon-derived nutrients great distances, enriching otherwise isolated habitats.     

Salmon‐derived nutrients drive diatom beta‐diversity patterns

1. Pacific salmon are a textbook example of migratory animals that transfer nutrients between ecosystems, but little is known about how salmon‐derived nutrients (SDN) affect the biodiversity of recipient freshwater ecosystems. We examined paleolimnological records from six Alaskan lakes to define how changes in SDN from sockeye salmon (Oncorhynchus nerka) influenced sedimentary diatom community structure and beta‐diversity among lakes and through time. 2. Using an isotopic mixing model, we showed that SDN loading could account for >80% of the lake total nitrogen budgets and strongly regulated diatom community composition. Spatial dissimilarity in diatom communities was positively related to differences in SDN among lakes (r² = 0.69, P < 0.01, n = 10). Likewise, temporal dissimilarity in diatom communities was positively related to differences in SDN in a sediment core with substantial variation in salmon spawner dynamics between 1700 and 1950 AD (r² = 0.34, P < 0.01, n = 19). Finally, beta‐diversity metrics quantifying temporal turnover within each lake’s sediment record were also positively related to the variance in SDN loading among lakes (r² = 0.88, P < 0.05, n = 5). Mean SDN was only negatively correlated to temporal diatom beta‐diversity. 3. Spatially subsidised systems often receive temporally variable resource inputs, and thus, it is not surprising that, unlike previous studies, we found that resource variability was the key driver of community composition and beta‐diversity. In habitats that receive strongly fluctuating external nutrient loads, environment heterogeneity may overweigh stochastic community processes. In addition, freshwater diatoms are characterised by great dispersal capabilities and short life cycles and therefore may be a more sensitive indicator for evaluating the role of resource variability than previously used model organisms. These results suggest that productivity–diversity relationship vary with the nature of nutrient loading and the life history of the community studied. 4. Overall, our study highlights that the transport of nutrients by sockeye salmon across ecosystem boundaries is a significant driver of algal community and biodiversity in nursery lakes, mainly through changing the magnitude of nutrient variation. As such, freshwater species diversity in regions like the U.S. Pacific Northwest may become impoverished where there have been long‐term declines in salmon populations and decreases in nutrient variability among lakes.     

Biotic disturbance and benthic community dynamics in salmon-bearing streams

1. Organisms can impact ecosystems via multiple pathways, often with positive and negative impacts on inhabitants. Understanding the context dependency of these types of impacts remains challenging. For example, organisms may perform different functions at different densities. 2. Anadromous salmon accumulate > 99% of their lifetime growth in marine ecosystems, and then return to spawn, often at high densities, in relatively confined freshwaters. While previous research has focused on how salmon nutrients can fertilize benthic communities, we examined how an ecosystem engineer, sockeye salmon Oncorhynchus nerka, influences seasonal dynamics of stream benthic communities through their nest-digging activities in south-western Alaska, USA. Benthic invertebrate and algal abundance were quantified every 7-14 days during the open water seasons of 10 streams in riffle and run habitats across multiple years, leading to 25 different stream-year combinations that spanned a large gradient of salmon density. 3. In streams with few or no salmon, benthic algal and insect biomass were fairly constant throughout the season. However, in streams with more than 0.1 salmon m(-2), algal and insect biomass decreased by an average of 75-85% during salmon spawning. Algal biomass recovered quickly following salmon disturbance, occasionally reaching pre-salmon biomass. In contrast, in streams with more than 0.1 salmon m(-2), aquatic insect populations did not recover to pre-salmon levels within the same season. We observed no positive impacts of salmon on algae or insects via fertilization from carcass nutrients. 4. Salmon, when their populations exceed thresholds in spawning density, are an important component of stream disturbance regimes and influence seasonal dynamics of benthic communities. Human activities that drive salmon densities below threshold densities, as has likely happened in many streams, will lead to altered seasonal dynamics of stream communities. Human activities that alter animal populations that are sources of biogenic disturbance can result in shifts in community dynamics.     

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.     

Carcass analogues provide marine subsidies for macroinvertebrates and juvenile Atlantic salmon in temperate oligotrophic streams

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Biogeochemical transformation of a nutrient subsidy: salmon, streams, and nitrification

Migratory animals can alter ecosystem function via the provision of nutrient subsidies. These subsidies are heterogeneous in space and time, which may create hot spots or hot moments in biogeochemical transformations, in turn altering the ecosystem effect of the subsidy by changing the form of the nutrients. Annual migrations of Pacific salmon (Oncorhynchus spp.) transport nutrients from the marine environment to their natal freshwater ecosystems. Salmon subsidies provide high quality nutrients (e.g., nitrogen, phosphorus, carbon) that may also be large in quantity where salmon migrations are near historic levels. We hypothesized that the nutrient subsidy provided via the excretion of ammonium (NH₄ ⁺) by live salmon would stimulate microbially mediated nitrification rates in stream sediments and increase streamwater nitrate (NO₃ ^?) concentrations. We quantified sediment nitrification in seven streams in Southeast Alaska before and during the salmon run in 2007 and 2008. Nitrification rates increased 3-fold from before to during the salmon run (mean ± SE = 0.07 ± 0.01 to 0.24 ± 0.02 mgN gAFDM^?1 d^?1, respectively). The variation in nitrification was explained by both streamwater and exchangeable NH₄ ⁺ concentrations (R ² = 0.50 and 0.71, respectively), which were low before salmon and increased relative to the size of the salmon run. To experimentally test the effect of salmon subsidies on nitrification rates, we staked senesced salmon carcasses on stream sediments for 3 weeks during the salmon run and then measured nitrification rates directly under the carcasses. Sediment nitrification was 2–5 times higher under the carcasses compared to nearby sediments without the direct carcass influence. Our results confirm that biogeochemical transformations alter the form of salmon-derived nitrogen, representing an overlooked aspect in the dynamics of this subsidy. Therefore, animal-derived nutrient subsidies are not passively retained or exported in recipient ecosystems, but also transformed, thereby influencing the form and incorporation of these nutrient subsidies.     

Signals of large scale climate drivers,hatchery enhancement,and marine factors in Yukon River Chinook salmon survival revealed with a Bayesian life history model

Understanding how species might respond to climate change involves disentangling the influence of co‐occurring environmental factors on population dynamics, and is especially problematic for migratory species like Pacific salmon that move between ecosystems. To date, debate surrounding the causes of recent declines in Yukon River Chinook salmon (Oncorhynchus tshawytscha) abundance has centered on whether factors in freshwater or marine environments control variation in survival, and how these populations at the northern extremity of the species range will respond to climate change. To estimate the effect of factors in marine and freshwater environments on Chinook salmon survival, we constructed a stage‐structured assessment model that incorporates the best available data, estimates incidental marine bycatch mortality in trawl fisheries, and uses Bayesian model selection methods to quantify support for alternative hypotheses. Models fitted to two index populations of Yukon River Chinook salmon indicate that processes in the nearshore and marine environments are the most important determinants of survival. Specifically, survival declines when ice leaves the Yukon River later in the spring, increases with wintertime temperature in the Bering Sea, and declines with the abundance of globally enhanced salmon species consistent with competition at sea. In addition, we found support for density‐dependent survival limitations in freshwater but not marine portions of the life cycle, increasing average survival with ocean age, and age‐specific selectivity of bycatch mortality in the Bering Sea. This study underscores the utility of flexible estimation models capable of fitting multiple data types and evaluating mortality from both natural and anthropogenic sources in multiple habitats. Overall, these analyses suggest that mortality at sea is the primary driver of population dynamics, yet under warming climate Chinook salmon populations at the northern extent of the species’ range may be expected to fare better than southern populations, but are influenced by foreign salmon production.     

Quantifying associations of large vertebrates with salmon in riparian areas of British Columbia streams by means of camera-traps,bait stations,and hair samples

The importance of transfers of marine derived nutrients (MDN) in salmon between stream ecosystems and terrestrial riparian systems is a subject of much recent research which has established that MDN subsidies support populations of many animal species, affect riparian zone productivity and may control the biodiversity of riparian plant communities. Numerous biophysical processes are involved in the transfer of MDN between aquatic and riparian zones of streams but, the largest proportion of MDN transport across the aquatic-terrestrial interface appears to depend on the actions of a predator–scavenger complex (PSC) that in British Columbia includes at least 23 species of mammals and birds. Consequently, the role of salmon in maintaining riparian ecosystem integrity depends not only on spatial and temporal variations in salmon abundance but also on variations in the composition and strength of linkages between salmon and other PSC species. Measuring these linkages with traditional methods using direct observation would be prohibitively expensive and time consuming. In the current paper laser activated camera systems, salmon carcass manipulation and deoxyribonucleic acid (DNA) analysis of hair samples were employed to (1) identify large vertebrate members of the PSC of riparian zone ecosystems and (2) quantify the strength of linkages of these PSC vertebrates to spatial and temporal changes in salmon abundance. Techniques developed and applied here to gain insights into the salmon PSC have several advantages over traditional survey methods. These include semi-automated capture of large volumes of data, identification of diurnally and nocturnally active PSC members, ready inclusion of experiments in field survey design, reduced impacts of unintentional human interference on survey results, and greatly reduced cost relative to direct observational studies. Results reported here confirm black bears (Ursus americanus), pine marten (Martes americana), cougar (Felis concolor), wolverine (Gulo luscus), eagles (Haliaeetus leucocephalus), ravens (Corvus corox), and vultures (Cathartes aura) as large vertebrate members of the regional salmon PSC. However, the composition and strength of linkages between salmon and these PSC members vary so much in space and time that most appear unsuitable as generally useful indicators of the potential magnitude of salmon-dependent, MDN-transport from aquatic to riparian-zone ecosystems. Black bears however, given their high relative abundance, broad geographic distribution, and strong linkages to salmon represent a superior candidate for further development in combination with salmon, as indicators of the maintenance of MDN transport potential in future assessments of riparian ecosystem integrity of salmon bearing streams.     

Idiosyncratic responses of Pacific salmon species to land cover,fragmentation, and scale

Salmon are critical to the ecology and livelihood of the Pacific Northwest, and are declining throughout much of their range. While much of their life cycle occurs in open ocean, freshwater conditions also contribute to population trends. Because stream habitats are connected to uplands by water flow, salmon can be influenced by the characteristics of terrestrial systems. We analyzed the relationships between the population trends of Pacific salmon (1953–2006) and land cover, fragmentation, and forest age derived from remotely‐sensed, landscape level datasets. Analyses included 425 populations of all native salmon species in 156 watersheds on Vancouver Island, British Columbia, Canada. Vancouver Island salmon escapements exhibited general patterns of decline, which may be largely controlled by broad‐scale marine conditions. The spatial variation in these population trends was related to landscape variables at watershed and riparian scales with regression trees. Results were found to be species specific, but characteristics indicating a legacy of historic and current forest management (such as fragmented forests and non‐forested or early‐successional forest cover) generally had negative effects, driven by a small subset of highly fragmented watersheds. Chum and coho had strong negative relationships with fragmentation, pink had a strong positive relationship with wetland abundance, and Chinook and sockeye were most closely related to geomorphology. There was no ‘single best’ scale of analysis. Salmon trends were generally more closely related to variables estimated over the entire watershed, however, the relative importances of watershed and riparian level predictors varied by both variable and species. Efforts to restore salmon habitat will be complicated by marine and freshwater processes, terrestrial conditions throughout watersheds, and the idiosyncratic requirements of each species.     

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.     

Watershed‐scale climate influences productivity of Chinook salmon populations across southcentral Alaska

The ecosystems supporting Pacific salmon (Oncorhynchus spp.) are changing rapidly as a result of climate change and habitat alteration. Understanding how—and how consistently—salmon populations respond to changes at regional and watershed scales has major implications for fisheries management and habitat conservation. Chinook salmon (O. tshawytscha) populations across Alaska have declined over the past decade, resulting in fisheries closures and prolonged impacts to local communities. These declines are associated with large‐scale climate drivers, but uncertainty remains about the role of local conditions (e.g., precipitation, streamflow, and stream temperature) that vary among the watersheds where salmon spawn and rear. We estimated the effects of these and other environmental indicators on the productivity of 15 Chinook salmon populations in the Cook Inlet basin, southcentral Alaska, using a hierarchical Bayesian stock‐recruitment model. Salmon spawning during 2003–2007 produced 57% fewer recruits than the previous long‐term average, leading to declines in adult returns beginning in 2008. These declines were explained in part by density dependence, with reduced population productivity following years of high spawning abundance. Across all populations, productivity declined with increased precipitation during the fall spawning and early incubation period and increased with above‐average precipitation during juvenile rearing. Above‐average stream temperatures during spawning and rearing had variable effects, with negative relationships in many warmer streams and positive relationships in some colder streams. Productivity was also associated with regional indices of streamflow and ocean conditions, with high variability among populations. The cumulative effects of adverse conditions in freshwater, including high spawning abundance, heavy fall rains, and hot, dry summers may have contributed to the recent population declines across the region. Identifying both coherent and differential responses to environmental change underscores the importance of targeted, watershed‐specific monitoring and conservation efforts for maintaining resilient salmon runs in a warming world.     

Marine and freshwater regime changes impact a community of migratory Pacific salmonids in decline

Marine and freshwater ecosystems are increasingly at risk of large and cascading changes from multiple human activities (termed “regime shifts”), which can impact population productivity, resilience, and ecosystem structure. Pacific salmon exhibit persistent and large fluctuations in their population dynamics driven by combinations of intrinsic (e.g., density dependence) and extrinsic factors (e.g., ecosystem changes, species interactions). In recent years, many Pacific salmon have declined due to regime shifts but clear understanding of the processes driving these changes remains elusive. Here, we unpacked the role of density dependence, ecosystem trends, and stochasticity on productivity regimes for a community of five anadromous Pacific salmonids (Steelhead, Coho Salmon, Pink Salmon, Dolly Varden, and Coastal Cutthroat Trout) across a rich 40-year time-series. We used a Bayesian multivariate state-space model to examine whether productivity shifts had similarly occurred across the community and explored marine or freshwater changes associated with those shifts. Overall, we identified three productivity regimes: an early regime (1976–1990), a compensatory regime (1991–2009), and a declining regime (since 2010) where large declines were observed for Steelhead, Dolly Varden, and Cutthroat Trout, intermediate declines in Coho and no change in Pink Salmon. These regime changes were associated with multiple cumulative effects across the salmon life cycle. For example, increased seal densities and ocean competition were associated with lower adult marine survival in Steelhead. Watershed logging also intensified over the past 40 years and was associated with (all else equal) ≥97% declines in freshwater productivity for Steelhead, Cutthroat, and Coho. For Steelhead, marine and freshwater dynamics played approximately equal roles in explaining trends in total productivity. Collectively, these changing environments limited juvenile production and lowered future adult returns. These results reveal how changes in freshwater and marine environments can jointly shape population dynamics among ecological communities, like Pacific salmon, with cascading consequences to their resilience.     

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.     

Comparative estuarine and marine migration ecology of Atlantic salmon and steelhead: blue highways and open plains

This synthesis focuses on the estuarine and ocean ecology of Atlantic salmon (Salmo salar) and steelhead (Oncorhynchus mykiss) across their southern ranges in North America. General life history and ecology share many common traits including iteroparity, duration of freshwater (0–3 years) and marine (2–5 years) rearing, ocean emigration at relatively large sizes and strong surface orientation compared to other salmonids. Despite parallels in life history and anthropogenic pressures, several differences emerged for these species. First, steelhead have greater life history diversity and a broader geographic distribution. Generally, estuary habitats serve as short-term migration corridors for both species. However, some steelhead populations used lagoon habitat in south-coast watersheds. While both species are epipelagic, Atlantic salmon exhibit more vertical migration. Atlantic salmon tend to follow migratory highways—relatively narrow bands along the coastal shelf, then crossing the Atlantic to feed inshore and in fjords of West Greenland. Conversely, steelhead exit the coastal shelf quickly, dispersing across the Pacific, and rarely use coastal environments. Despite inhabiting rivers in warm dry Mediterranean climates, the extended range and stability of southern steelhead distribution is likely buffered by cool upwelled waters of the California Current. Whereas Atlantic salmon populations are restricted by warmer Northwest Atlantic circulation patterns lacking cool upwelling with greater susceptibility to warming associated with climate change. Determining the rate of marine habitat changes in the Atlantic and Pacific Oceans is important to the conservation of these species, including subtleties of temporal and spatial habitat use, and adaptability to ocean ecosystems under climate change.     

Sea to sky: impacts of residual salmon-derived nutrients on estuarine breeding bird communities

Pacific salmon (Oncorhynchus spp.) returning to streams around the North Pacific Rim provide a nutrient subsidy to these ecosystems. While many species of animals feed directly on salmon carcasses each autumn, salmon-derived nutrients can also be stored in coastal habitats throughout the year. The effects of this storage legacy on vertebrates in other seasons are not well understood, especially in estuaries, which can receive a large portion of post-spawning salmon nutrients. We examine the effects of residual salmon-derived nutrients, forest habitats and landscape features on summer breeding birds in estuary forests. We compared models containing environmental variables and combined chum (Oncorhynchus keta) and pink (Oncorhynchus gorbuscha) salmon biomass to test predictions concerning bird density and diversity. We discovered that total bird, insectivore, golden-crowned kinglet and Pacific wren densities and Shannon's diversity in the summer were strongly predicted by salmon biomass in the autumn. For most metrics, this relationship approaches an asymptote beyond 40 000 kg of salmon biomass. Foliage height diversity, watershed catchment area and estuary area were also important predictors of avian communities. Our study suggests that the legacy of salmon nutrients influences breeding bird density and diversity in estuaries that vary across a wide gradient of spawning salmon biomass.