Stream geomorphology regulates the effects on periphyton of ecosystem engineering and nutrient enrichment by Pacific salmon

1. Pacific salmon (Oncorhynchus spp.) returning to streams deliver substantial quantities of nutrients (nitrogen and phosphorus) that may stimulate primary production. Salmon can also affect the phytobenthos negatively via physical disturbance during nest excavation, a process that may counteract the positive effects of salmon‐derived nutrients on benthic algae. The ability of salmon to disturb benthic habitats may be a function of substratum particle size, and therefore, the geomorphology of streams could determine the net effect of salmon on benthic communities. 2. Based on surveys of 17 streams in southwest Alaska before the salmon run and during peak salmon density, we identified size thresholds for the disturbance of substratum particles by salmon and classified particles as vulnerable (<60 mm B‐axis), invulnerable (>110 mm) or transitional (61–110 mm). At the scale of individual rocks, algal biomass on vulnerable substrata decreased at peak spawning (relative to values before the run) as a power function of salmon density; transitional and invulnerable substrata showed no quantifiable pattern. However, invulnerable substrata in streams with more than 0.11 salmon m^?2 showed net algal accrual, or relatively smaller declines in algal biomass, than vulnerable substrata, indicating that large rocks provide refuge for benthic algae from salmon disturbance. 3. We expected that streams with proportionally larger rocks would respond positively to salmon at the whole‐stream scale, after accounting for the relative abundance of rocks of different sizes within streams. Invulnerable rocks made up only 0–12% of the total substratum particle size distribution in salmon‐bearing streams, however, and algal accrual on invulnerable substrata did not outweigh the strong disturbance effects on the more spatially extensive vulnerable substrata. The change in whole‐stream benthic algal biomass among streams was negatively related to salmon density. 4. Stable isotopes of nitrogen (δ¹⁵N) were used to track nutrients from salmon into benthic biota. Periphyton δ¹⁵N on rocks of all size classes was higher at peak salmon spawning than before the salmon run, indicating the uptake of salmon‐derived nitrogen. Peak δ¹⁵N values were positively related to salmon abundance and followed a two‐isotope mixing relationship. The per cent of N from salmon in periphyton was also related to salmon density and was best explained by a saturating relationship. Spring δ¹⁵N was unrelated to salmon returns in the previous year, suggesting little annual carryover of salmon nutrients.     

Pacific salmon effects on stream ecosystems: a quantitative synthesis

Pacific salmon (Oncorhynchus spp.) disturb sediments and fertilize streams with marine-derived nutrients during their annual spawning runs, leading researchers to classify these fish as ecosystem engineers and providers of resource subsidies. While these processes strongly influence the structure and function of salmon streams, the magnitude of salmon influence varies widely across studies. Here, we use meta-analysis to evaluate potential sources of variability among studies in stream ecosystem responses to salmon. Results obtained from 37 publications that collectively included 79 streams revealed positive, but highly inconsistent, overall effects of salmon on dissolved nutrients, sediment biofilm, macroinvertebrates, resident fish, and isotopic enrichment. Variation in these response variables was commonly influenced by salmon biomass, stream discharge, sediment size, and whether studies used artificial carcass treatments or observed a natural spawning run. Dissolved nutrients were positively related to salmon biomass per unit discharge, and the slope of the relationship for natural runs was five to ten times higher than for carcass additions. Mean effects on ammonium and phosphorus were also greater for natural runs than carcass additions, an effect attributable to excretion by live salmon. In contrast, we observed larger positive effects on benthic macroinvertebrates for carcass additions than for natural runs, likely because disturbance by live salmon was absent. Furthermore, benthic macroinvertebrates and biofilm associated with small sediments (<32 mm) displayed a negative response to salmon while those associated with large sediments (>32 mm) showed a positive response. This comprehensive analysis is the first to quantitatively identify environmental and methodological variables that influence the observed effects of salmon. Identifying sources of variation in salmon–stream interactions is a critical step toward understanding why engineering and subsidy effects vary so dramatically over space and time, and toward developing management strategies that will preserve the ecological integrity of salmon streams. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Separating physical disturbance and nutrient enrichment caused by Pacific salmon in stream ecosystems

1. Pacific salmon (Oncorhynchus spp.) deliver marine‐derived nutrients to the streams in which they spawn and die, and these resource subsidies can increase the abundance of stream biota. In strong contrast, physical disturbance from salmon spawning activity can reduce the abundance of benthic organisms. Previous experimental designs have not established the relative effects of these two contrasting processes on stream organisms during a salmon run. 2. We combined manipulative and observational field studies to assess the degree of nutrient enrichment, physical disturbance, and the net effect of salmon on the abundance of benthic periphyton. Related salmon‐mediated processes were also evaluated for benthic macroinvertebrates. Mesh exclosures (2 × 2 m plots) prevented salmon from disturbing areas of the stream channel, which were compared with areas to which salmon had access. Sampling was conducted both before and during the late‐summer spawning run of pink (O. gorbushca) and chum (O. keta) salmon. 3. Streamwater nitrogen and phosphorus concentrations increased sharply with the onset of the salmon run, and highly significant positive relationships were observed between the numbers of salmon present in the stream and these dissolved nutrients. Before the salmon run, periphyton biomass (as chlorophyll a) and total macroinvertebrate abundance were very similar between control and exclosure plots. During the salmon run, exclosures departed substantially from controls, suggesting significant disturbance imparted on benthic biota. 4. Comparing exclosures before and during the salmon run enabled us to estimate the effects of salmon in the absence of direct salmon disturbance. This ‘nutrient enrichment potential’ was significant for periphyton biomass, as was a related index for macroinvertebrate abundance (although enhanced invertebrate drift into exclosures during the salmon run could also have been important). Interestingly, however, the net effect of salmon, evaluated by comparing control plots before and during the salmon run, was relatively modest for both periphyton and macroinvertebrates, suggesting that nutrient enrichment effects were largely offset by disturbance. 5. Our results illustrate the importance of isolating the specific mechanisms via which organisms affect ecosystems, and indicate that the relative magnitude of salmon nutrient enrichment and benthic disturbance determines the net effect that these ecologically important fish have on stream ecosystems.     

Effects of spawning Pacific salmon on the isotopic composition of biota differ among southeast Alaska streams

1. Adult Pacific salmon (Oncorhynchus spp.) transport marine nutrients to fresh waters and disturb sediments during spawning. The relative importance of nutrient fertilisation and benthic disturbance by salmon spawners can be modulated by environmental conditions (e.g. biological, chemical and physical conditions in the catchment, including human land use). 2. To determine the importance of the environmental context in modifying the uptake and incorporation of salmon‐derived material into stream biota, we measured the nitrogen (δ¹⁵N) and carbon (δ¹³C) isotopic composition of benthic algae (i.e. epilithon) and juvenile coho salmon (Oncorhynchus kisutch) in seven streams across a timber‐harvest gradient (8–69% catchment area harvested), both before and during the salmon run. Conditional bootstrap modelling simulations were used to assess variability in the response of epilithon and juvenile coho salmon to spawning salmon. 3. In response to spawning salmon, epilithon exhibited enrichment in both δ¹⁵N (mean: 1.5‰) and δ¹³C (2.3‰). Juvenile coho were also enriched in both δ¹⁵N (0.7‰) and δ¹³C (1.4‰). Conditional bootstrap models indicate decreased variation in data as spatial replication increases, suggesting that the number of study sites can influence the results of Pacific salmon isotope studies. 4. Epilithon isotopic enrichment was predicted by environmental conditions, with δ¹⁵N enrichment predicted by stream temperature and timber harvest (R² = 0.87) and δ¹³C enrichment by discharge, sediment size, timber harvest and spawner density (R² = 0.96). Furthermore, we found evidence for a legacy effect of salmon spawners, with pre‐spawner δ¹⁵N and δ¹³C of both epilithon and juvenile coho predicted by salmon run size in the previous year. 5. Our results show that the degree of incorporation of salmon‐derived nitrogen and carbon differs among streams. Furthermore, the environmental context, including putative legacy effects of spawning salmon, can influence background isotopic concentrations and utilisation of salmon‐derived materials in southeast Alaska salmon streams. Future studies should consider the variation in isotopic composition of stream biota when deciding on the number of study sites and samples needed to generate meaningful results.     

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.     

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.     

Straying of hatchery salmon in Prince William Sound,Alaska

The straying of hatchery salmon may harm wild salmon populations through a variety of ecological and genetic mechanisms. Surveys of pink (Oncorhynchus gorbuscha), chum (O. keta) and sockeye (O. nerka) salmon in wild salmon spawning locations in Prince William Sound (PWS), Alaska since 1997 show a wide range of hatchery straying. The analysis of thermally marked otoliths collected from carcasses indicate that 0–98% of pink salmon, 0–63% of chum salmon and 0–93% of sockeye salmon in spawning areas are hatchery fish, producing an unknown number of hatchery-wild hybrids. Most spawning locations sampled (77%) had hatchery pink salmon from three or more hatcheries, and 51% had annual escapements consisting of more than 10% hatchery pink salmon during at least one of the years surveyed. An exponential decay model of the percentage of hatchery pink salmon strays with distance from hatcheries indicated that streams throughout PWS contain more than 10% hatchery pink salmon. The prevalence of hatchery pink salmon strays in streams increased throughout the spawning season, while the prevalence of hatchery chum salmon decreased. The level of hatchery salmon strays in many areas of PWS are beyond all proposed thresholds (2–10%), which confounds wild salmon escapement goals and may harm the productivity, genetic diversity and fitness of wild salmon in this region     

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.     

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

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Disturbance of freshwater habitats by anadromous salmon in Alaska

High densities of habitat modifiers can dramatically alter the structure of ecosystems. Whereas spawning sockeye salmon (Oncorhynchus nerka) dig nests that cover over 2 m² and are at least 20 cm deep, and can spawn at high densities, relatively little attention has been devoted to investigating the impacts of this disturbance. We hypothesized that this temporally and spatially predictable bioturbation has large impacts on the coastal aquatic habitats used by sockeye. We experimentally investigated the impacts of disturbance caused by spawning sockeye in two streams and two lakes in Alaska by excluding salmon from 2.25 m² plots where they traditionally spawn. We sampled exclusions and control plots before, during, and after spawning. During sockeye spawning, fine sediment accumulated in areas where sockeye were excluded from spawning. In addition, sockeye spawning significantly decreased algal biomass by 80% compared to exclusion plots. We found mixed effects of spawning on the invertebrate assemblage. Tricladida and Chironomidae densities increased by 3x in exclusion plots relative to control plots in one creek site. However, for most taxa and sites, invertebrate densities declined substantially as spawning progressed, regardless of experimental treatment. Habitat modification by spawning salmon alters both community organization and ecosystem processes.     

Environmental variability and the ecological effects of spawning Pacific salmon on stream biofilm

1. Variation in resource subsidies can create or reinforce heterogeneity in recipient ecosystems. Related activities of organisms delivering resource subsidies, such as ecosystem engineering by Pacific salmon spawners (Oncorhynchus spp.), also alter heterogeneity. We studied whether heterogeneity in stream environmental conditions and spawner abundances were reflected in the net ecological effects of salmon (i.e. enrichment by resource subsidies and disturbance by ecosystem engineering) on benthic biofilm. 2. We sampled seven Southeast Alaska streams over 3 years, both before and during the salmon run. In each stream and year, stream environmental characteristics and their influence on responses of benthic biofilm [mean and coefficient of variation of chlorophyll a (chl a), ash‐free dry mass (AFDM) and autotrophic index (AFDM:chl a)] to spawners were assessed. 3. Streams and periods before and during the salmon run were distinct based on their environmental characteristics. The responses of most biofilm metrics to spawners were stream‐ and year‐specific, suggesting that the ecological effect of spawners ranged from net enrichment to net disturbance depending on the stream or year studied. The environmental context, especially temperature, large wood, and sediment size, explained >50% of biofilm variability during the run, but <30% over the entire study, suggesting that salmon can alter environmental constraints. 4. Precision of biofilm estimates improved by increasing either the number of streams or the number of years sampled (i.e. spatial or temporal replication). However, combining data from different North Pacific Rim ecoregions inflated the confidence interval as compared with a single ecoregion, indicating the importance of regional environmental contexts for net salmon effects. 5. Our results suggest that biofilm responses to salmon can vary greatly, even within a single ecoregion, and that environmental conditions can modify net salmon effects. Consequently, generalisations about biofilm responses across the native range of salmon may be challenging.     

Brown bears selectively kill salmon with higher energy content but only in habitats that facilitate choice

Pacific salmon return to spawn in thousands of streams across the Pacific Rim, from large rivers to tiny headwater streams. Once on the spawning grounds, salmon undergo dramatic biochemical changes as they metabolize stored lipid and protein reserves; at stream entrance, they will contain up to 85% more lipid and 40% more protein than at their senescent death a week or two later. Foraging brown and black bears that congregate at spawning streams thus encounter salmon that vary dramatically in their energy content and thus energetic reward. We hypothesized that bears would selectively kill salmon that are highest in energy content (fewest number of days on the spawning grounds) when they pursue salmon at small shallow streams where little effort is necessary to capture salmon, i.e. habitats that facilitate choice. In contrast, bears in environments where foraging is difficult (deeper, more complex streams) should be less selective and should capture salmon that are most available. We tested these ideas by examining predation rates on fish of different in-stream ages (i.e. energy content) at three different streams that varied in physical habitat attributes. At a very shallow, simple stream, bears preferentially killed salmon that had spent the fewest days in the stream. At two streams where deeper water and woody debris provided refuges for salmon, predation rates increased with in-stream age. At the shallowest streams encounter rates and capture success are likely equal among the high- and low-energy salmon and thus predation rates reflect active choice by bears. In contrast capture success probably increases on the older salmon at the larger streams (due to a loss of vigor), and thus 'preference' for these fish increases due to decreasing effort necessary to capture them.     

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.     

Balancing competing life‐stage requirements in salmon habitat rehabilitation: between a rock and a hard place

Gravel augmentation is often applied to rivers and streams to rehabilitate salmonid spawning and incubation habitat. However, the effect of gravel size on salmon spawning utilization and embryo survival during incubation is not well understood. We conducted an experiment on a regulated and previously mined Northern California salmonid‐bearing stream in which different sized gravel (small, medium, and large) patches were placed into the stream's degraded spawning reach. We documented Oncorhynchus tshawytscha (Chinook salmon) spawning activity within the three gravel sizes for two seasons. In addition, we deployed Chinook salmon embryos into each gravel size patch and allowed them to incubate until estimated emergence time. Although all experimental gravel sizes were predicted to be within the spawning population's mobilization capabilities, model results indicated the probability of salmon building redds decreased as substrate size increased. Conversely, embryo survival increased as gravel size increased. A possible mechanism of disparate Chinook salmon embryo survival is provided by an observed decrease in embryo survival correlating with greater presence of embryo predators (leeches), which are associated with smaller gravel. Our results indicate a parent‐offspring conflict in optimal spawning gravel size for Chinook salmon, and suggest that an intermediate gravel size would maximize overall reproductive success across both spawning and incubation life stages.     

Diet partitioning in sympatric Atlantic salmon and brown trout in streams with contrasting riparian vegetation

Prey intake by Atlantic salmon Salmo salar and brown trout Salmo trutta was measured across different riparian vegetation types: grassland, open canopy deciduous and closed canopy deciduous, in upland streams in County Mayo, Western Ireland. Fishes were collected by electrofishing while invertebrates were sampled from the benthos using a Surber sampler and drifting invertebrates collected in drift traps. Aquatic invertebrates dominated prey numbers in the diets of 0+ year Atlantic salmon and brown trout and 1+ year Atlantic salmon, whereas terrestrial invertebrates were of greater importance for diets of 1+ and 2+ year brown trout. Terrestrial prey biomass was generally greater than aquatic prey for 1+ and 2+ year brown trout across seasons and riparian types. Prey intake was greatest in spring and summer and least in autumn apart from 2+ year brown trout that sustained feeding into autumn. Total prey numbers captured tended to be greater for all age classes in streams with deciduous riparian canopy. Atlantic salmon consumed more aquatic prey and brown trout more terrestrial prey with an ontogenetic increase in prey species richness and diversity. Atlantic salmon and brown trout diets were most similar in summer. Terrestrial invertebrates provided an important energy subsidy particularly for brown trout. In grassland streams, each fish age class was strongly associated with aquatic, mainly benthic invertebrates. In streams with deciduous riparian canopy cover, diet composition partitioned between conspecifics with older brown trout associated with surface drifting terrestrial invertebrates and older Atlantic salmon associated with aquatic invertebrates with a high drift propensity in the water column and 0+ year fish feeding on benthic aquatic invertebrates. Deciduous riparian canopy cover may therefore facilitate vertical partitioning of feeding position within the water column between sympatric Atlantic salmon and brown trout. Implications for riparian management are discussed.     

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.     

Asynchrony in population dynamics of sockeye salmon in southwest Alaska

Ecologists have examined the synchronization of population dynamics across space as a means to understand how populations respond to climate variation. However, response diversity may reflect important variation among local population dynamics driven by population‐specific responses to regional environmental change. We used long‐term data on sockeye salmon Oncorhynchus nerka from pristine watersheds of southwestern Alaska to show that populations spawning in close proximity (<40 km) to one another have a limited degree of synchrony in their dynamics, even after accounting for density‐dependent processes. In fact, the dynamics of local populations of stream‐spawning sockeye salmon were no more coherent than those of stocks at a much coarser resolution across this region of Alaska. We examined four hypotheses to explain the observed patterns of asynchrony among stream‐spawning populations, and found that populations spawning in dissimilar habitats, and using different nursery lakes were less synchronized in their productivity. Similarity in the age structure of spawning adults was less correlated with synchrony in productivity. These results emphasize the importance of maintaining diverse spawning and rearing habitat for the conservation of Pacific salmon, and should guide conservation planning for Pacific salmon populations in regions where natural dynamics have been altered by habitat loss, hatchery practices, and over‐fishing.     

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.     

Effects of Pacific salmon spawning and carcass availability on the caddisfly Ecclisomyia conspersa (Trichoptera: Limnephilidae)

1. The effects of spawning coho (Oncorhynchus kisutch) and chum salmon (Oncorhynchus keta) on the limnephilid caddisfly Ecclisomyia conspersa were evaluated by experimentally excluding salmon from the upper 14‐m stretch of a spawning channel by a wire‐meshed fence. Density, and development and growth rates, of larvae upstream of the fence (without salmon) were compared with those downstream (with salmon). 2. Larval density in the stretch with salmon declined during spawning, but increased again after spawning subsided and the carcasses of dead fish became available. In the stretch with salmon, larval density on salmon carcasses was seven to 37 times greater than on the adjacent channel substratum. The rate of larval development in the stretch with salmon was greater than that in the stretch without salmon. Two months after carcasses became available, 98% of larvae sampled from the stretch with salmon were in the fifth instar, compared to only 23% from the stretch without salmon. Body weight of E. conspersa in the stretches with and without salmon increased by an average of 3.04 and 2.38 mg, respectively, over a 6‐month period. 3. ¹⁵N values of larvae from the stretch with salmon increased following the arrival of the fish, suggesting that the larvae were feeding on salmon‐derived material, such as eggs and carcasses, which contain a high proportion of the heavier stable isotope. In contrast, ¹⁵N values of larvae from the stretch without salmon remained relatively constant throughout the experiment. The availability of salmon carcasses as a high‐quality food source late in larval development may increase survival and fecundity of E. conspersa. 4. These substantial differences were consistent with the view that they were due to the experimental exclusion of salmon and salmon carcasses from the upstream stretch, though the study was un‐replicated and thus precludes ascribing causation more definitely.     

Varying effects of anadromous sockeye salmon on the trophic ecology of two species of resident salmonids in southwest Alaska

1. Anadromous salmon transport marine‐derived nutrients and carbon to freshwater and riparian ecosystems upon their return to natal spawning systems. The ecological implications of these subsidies on the trophic ecology of resident fish remain poorly understood despite broad recognition of their potential importance. 2. We studied the within‐year changes in the ration size, composition and stable isotope signature of the diets of two resident salmonids (rainbow trout, Oncorhynchus mykiss; Arctic grayling, Thymallus arcticus) before and after the arrival of sockeye salmon (Oncorhynchus nerka) to their spawning grounds in the Bristol Bay region of southwest Alaska. 3. Ration size and energy intake increased by 480–620% for both species after salmon arrived. However, the cause of the increases differed between species such that rainbow trout switched to consuming salmon eggs, salmon flesh and blowflies that colonized salmon carcasses, whereas grayling primarily ate more benthic invertebrates that were presumably made available because of physical disturbances by spawning salmon. 4. We also observed an increase in the δ¹⁵N of rainbow trout diets post‐salmon, but not for grayling. This presumably led to the observed increase in the δ¹⁵N of rainbow trout with increasing body mass, but not for grayling. 5. Using a bioenergetics model, we predicted that salmon‐derived resources contributed a large majority of the energy necessary for growth in this resident fish community. Furthermore, the bioenergetics model also showed how seasonal changes in diet affected the stable isotope ratios of both species. These results expand upon a growing body of literature that highlights the different pathways whereby anadromous salmon influence coastal ecosystems, particularly resident fish.