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Salmonid populations are often regulated by territorial competition among juveniles for food and space. In the canonical view, salmonid territories are spaced horizontally across the river bottom in a two-dimensional mosaic. However, some juveniles instead feed in tight, three-dimensional (3-D) social groups. To investigate whether territoriality is possible within such groups, we applied a new concept—the momentary home range—to quantify the size, exclusivity, and temporal dynamics of 3-D space use by juvenile Chinook salmon ( Oncorhynchus tshawytscha) in the Chena River, AK. Individual strategies spanned a broad continuum of exclusivity and stationarity. However, some of the largest, most dominant fish in each group aggressively defended stationary, exclusive feeding spaces and thus were unambiguously territorial. Transient floaters entered and left the group quickly. A majority of fish were not aggressive but nevertheless occupied exclusive, stationary spaces that probably function as territories with regard to resource distribution and population regulation. The presence of territoriality within social groups, in a 3-D configuration, expands the known domain of this important behavior.

Anadromous Pacific salmon are semelparous, and resource subsidies from spawning adults (marine-derived nutrients, or MDN) benefit juvenile salmonids rearing in freshwater. However, it remains unclear how MDN assimilation relates to spawner abundance within a watershed. To address this, we examined seasonal, watershed-scale patterns of MDN assimilation in rearing coho (Oncorhynchus kisutch) and Chinook (O. tshawytscha) salmon and compared it with spawner biomass and landscape features in a western Alaska watershed with contrasting structural complexity in two sub-drainages. Adult salmon biomass density was estimated from escapement and spawner distribution data, and MDN assimilation in juvenile salmon was estimated via stable isotopes. In the North River, MDN assimilation was lowest in early summer, prior to annual spawning migrations, increased after spawning, and peaked in late winter. In the more complex mainstem Unalakleet River, MDN assimilation was higher but varied minimally from summer through fall before increasing in late fall and winter. Summer MDN assimilation, prior to salmon spawning, was primarily a function of habitat complexity, where MDN was highest in sloughs and the more complex mainstem river. After salmon spawned, fall MDN assimilation was a function of adult pink and Chinook salmon biomass as well as MDN assimilation that occurred prior to spawning (that is, summer MDN), but unrelated to total summer biomass (all salmon species biomass combined). Thus, MDN assimilation by juvenile salmon in the fall was a function of species-specific adult spawner abundance but seasonal patterns of MDN assimilation were masked in complex habitat where summer MDN assimilation remained high.

Stream fish rely on a mix of terrestrial and aquatic prey sources. While the importance of terrestrial invertebrates as a food source for stream fish is well documented, the role of aquatic insects that emerge from the stream as winged adult insects (aquatic winged adults) and return to the stream as prey is less understood. In this study we determined the proportion of total diet for stream-rearing juvenile Coho Salmon (Oncorhynchus kisutch) that is derived from terrestrial and aquatic winged adult invertebrates which enter the stream from riparian habitats and consider how those cross-ecosystem prey contributions vary based on riparian habitat type. Study reaches were identified in three streams within the Kenai River watershed of Alaska that were representative of habitats found throughout the region and riparian vegetation was classified into grass/sedge, shrub and tree types using LiDAR. Juvenile Coho Salmon stomach contents were sampled seasonally in study reaches over a two-year period and ingested invertebrates were identified by taxa, life stage and origin. Our results showed that aquatic winged adult prey contributions to juvenile salmon diet were significantly lower in the grass/sedge study reach, and cross-ecosystem invertebrate prey represented a significantly higher proportion of juvenile salmon diet in the tree study reach. Invertebrate prey in the grass/sedge reach were composed primarily of the larval life stage of aquatic winged adults. These results suggest that change in riparian vegetation from tree/shrub to grass/sedge along Kenai streams as projected by regional climate change models, or that results from anthropogenic modification, will likely lead to lower availability of cross-ecosystem prey for stream fish. Management of riparian buffers along streams to preserve or increase occurrence of trees and shrubs is likely to help mitigate impacts of those possible changes.

Conservation of Arctic fish species is challenging partly due to our limited ability to track fish through time and space, which constrains our understanding of life history diversity and lifelong habitat use. Broad Whitefish (Coregonus nasus) is an important subsistence species for Alaska’s Arctic Indigenous communities, yet little is known about life history diversity, migration patterns, and freshwater habitat use. Using laser ablation Sr isotope otolith microchemistry, we analyzed Colville River Broad Whitefish 87Sr/86Sr chronologies (n = 61) to reconstruct movements and habitat use across the lives of individual fish. We found evidence of at least six life history types, including three anadromous types, one semi-anadromous type, and two nonanadromous types. Anadromous life history types comprised a large proportion of individuals sampled (collectively, 59%) and most of these (59%) migrated to sea between ages 0–2 and spent varying durations at sea. The semi-anadromous life history type comprised 28% of samples and entered marine habitat as larvae. Nonanadromous life history types comprised the remainder (collectively, 13%). Otolith 87Sr/86Sr data from juvenile and adult freshwater stages suggest that habitat use changed in association with age, seasons, and life history strategies. This information on Broad Whitefish life histories and habitat use across time and space will help managers and conservation planners better understand the risks of anthropogenic impacts and help conserve this vital subsistence resource.

Pacific pink salmon (Oncorhynchus gorbuscha) invasions, thought to originate from populations introduced and established in Russia, occurred along the Norwegian coast in 2017 and 2019. Despite several thousand pink salmon entering and establishing in northern Norwegian rivers, current understanding of the ecological effect of the species in northern Europe is limited. Scavengers feeding on pacific salmon carcasses are important vectors for the transport of marine derived energy and nutrients to terrestrial ecosystems in the Pacific Northwest, North America, where the salmon naturally occur. However the role of terrestrial and aquatic scavengers in the consumption and removal of pink salmon beyond the salmon’s native range is unknown. This study has identified terrestrial and sub-aquatic vertebrate scavengers on pink salmon carcasses in a sub-arctic river in northern Norway. Avian scavengers filmed by a camera placed near sites baited with pink salmon carcasses included the Eurasian magpie (Pica pica), hooded crow (Corvus cornix), common raven (Corvus corax), the European herring gull (Larus argentatus), redwing (Turdus iliacus) and goosander (Mergus merganser). However, the largest carcass weight was removed by red foxes (Vulpes vulpes). Carcasses entering Vesterelv River in 2019 were estimated to provide energy and nutrients to the river ecosystem an order of magnitude lower than in the Pacific Northwest. This study provides some of the first information in northern Europe on the mechanisms and quantification of energy and nutrient transfer from the ocean to riparian environments via introduced Pacific pink salmon. Results help to begin to determine the ecological effect of pink salmon and the development of appropriate management strategies.

Lakes can be important to stream dwelling fishes, yet how individuals exploit habitat heterogeneity across complex stream‐lake networks is poorly understood. Furthermore, despite growing awareness that intermittent streams are widely used by fish, studies documenting the use of seasonally accessible lakes remain scarce. We studied Arctic grayling (Thymallus arcticus) in a small seasonally flowing (June–October) stream‐lake network in Alaska using PIT telemetry. Overall, 70% of fish visited two lakes, 8% used a single lake, and 22% used only stream reaches. We identified five distinct behavioural patterns that differed in dominant macrohabitat used (deep lake, shallow lake or stream reaches), entry day into the network and mobility. Some juvenile fish spent the entire summer in a shallow seasonally frozen lake (average 71 days), whereas others demonstrated prospecting behaviour and only entered the stream channel briefly in September. Another group included adult and juvenile fish that were highly mobile, moving up to 27 km while in the 3‐km stream‐lake network, and used stream reaches extensively (average 59 days). Lentic and lotic habitats served differing roles for individuals, some fish occupied stream reaches as summer foraging habitat, and other individuals used them as migration corridors to access lakes. Our study emphasises the importance of considering stream‐lake connectivity in stream fish assessments, even to shallow seasonally frozen habitats not widely recognised as important. Furthermore, we demonstrate that individuals may use temporary aquatic habitats in complex and changing ways across ontogeny that are not captured by typical classifications of fish movement behaviour.

Pacific salmon Oncorhynchus spp. face serious challenges from climate and landscape change, particularly in the southern portion of their native range. Conversely, climate warming appears to be allowing salmon to expand northwards into the Arctic. Between these geographic extremes, in the Gulf of Alaska region, salmon are at historically high abundances but face an uncertain future due to rapid environmental change. We examined changes in climate, hydrology, land cover, salmon populations, and fisheries over the past 30–70 years in this region. We focused on the Kenai River, which supports world-famous fisheries but where Chinook Salmon O. tshawytscha populations have declined, raising concerns about their future resilience. The region is warming and experiencing drier summers and wetter autumns. The landscape is also changing, with melting glaciers, wetland loss, wildfires, and human development. This environmental transformation will likely harm some salmon populations while benefiting others. Lowland s...

Fisheries technician Mike Lunde prepares to check a floating gill net on the middle Colville River. Photo credit: Jason C. Leppi.In the cold waters of Alaska's Arctic swims a mysterious fish with l...

Beaded streams are prominent across the Arctic Coastal Plain (ACP) of Alaska, yet prey flow and food web dynamics supporting fish inhabiting these streams are poorly understood. Arctic grayling (Thymallus arcticus) are a widely distributed upper-level consumer on the ACP and migrate into beaded streams to forage during the short 3-month open-water season. We investigated energy pathways and key prey resources that support grayling in a representative beaded stream, Crea Creek. We measured terrestrial invertebrates entering the stream from predominant riparian vegetation types, prey types supporting a range of fish size classes, and how riparian plants and fish size influenced foraging habits. We found that riparian plants influenced the quantity of terrestrial invertebrates entering Crea Creek; however, these differences were not reflected in fish diets. Prey type and size ingested varied with grayling size and season. Small grayling ( 15 cm FL) foraged most heavily on ninespine stickleback (Pungitius pungitius) throughout the summer, indicating that grayling can be insectivorous and piscivorous, depending on size. These findings underscore the potential importance of small streams in Arctic ecosystems as key summer foraging habitats for fish. Understanding trophic pathways supporting stream fishes in these systems will help interpret whether and how petroleum development and climate change may affect energy flow and stream productivity, terrestrial–aquatic linkages and fishes in Arctic ecosystems.

Despite their widespread presence in northern-latitude ecosystems, the ecological role of Ninespine Stickleback Pungitius pungitius is not well understood. Ninespine Stickleback can occupy both top and intermediate trophic levels in freshwater ecosystems, so their role in food webs as a predator on invertebrates and as a forage fish for upper level consumers probably is substantial. We introduced Ninespine Sticklebacks to fishless ponds to elucidate their potential effects as a predator on invertebrate communities in Arctic lentic freshwaters. We hypothesized that Ninespine Stickleback would affect freshwater invertebrate communities in a top-down manner. We predicted that the addition of Ninespine Sticklebacks to fishless ponds would: 1) reduce invertebrate taxonomic richness, 2) decrease overall invertebrate abundance, 3) reduce invertebrate biomass, and 4) decrease average invertebrate body size. We tested our hypothesis at 2 locations by adding Ninespine Stickleback to isolated ponds and compa...

Most studies of cross-ecosystem resource subsidies have demonstrated positive effects on recipient consumer populations, often with very large effect sizes. However, it is important to move beyond these initial addition–exclusion experiments to consider the quantitative consequences for populations across gradients in the rates and quality of resource inputs. In our introduction to this special issue, we describe at least four potential models that describe functional relationships between subsidy input rates and consumer responses, most of them asymptotic. Here we aim to advance our quantitative understanding of how subsidy inputs influence recipient consumers and their communities. In the papers following, fish were either the recipient consumers or the subsidy as carcasses of anadromous species. Advancing general, predictive models will enable us to further consider what other factors are potentially co-limiting (e.g., nutrients, other population interactions, physical habitat, etc.) and better integrate resource subsidies into consumer–resource, biophysical dynamics models.

Contributions of terrestrial-, freshwater-, and marine-derived prey resources to stream fishes vary over time and space, altering the energy pathways that regulate production. In this study, we determined large-scale use of these resources by juvenile Chinook and coho salmon (Oncorhynchus tshawytscha and Oncorhynchus kisutch, respectively) in the glacial Susitna River, Alaska. We resolved spatial and temporal trophic patterns among multiple macrohabitat types along a 97 km segment of the river corridor via stable isotope and stomach content analyses. Juvenile salmon were supported primarily by freshwater-derived resources and secondarily by marine and terrestrial sources. The relative contribution of marine-derived prey to rearing salmon was greatest in the fall within off-channel macrohabitats, whereas the contributions of terrestrial invertebrate prey were generally greatest during midsummer, across all macrohabitats. No longitudinal (upstream–downstream) diet pattern was discernable. These results highlight large-scale spatial and seasonal patterns of energy flow and the dynamic interplay of pulsed marine and terrestrial prey subsidies to juvenile Chinook and coho salmon in a large, complex, and relatively pristine glacial river.

The spread of invasive species in riparian forests has the potential to affect both terrestrial and aquatic organisms linked through cross-ecosystem resource subsidies. However, this potential had not been explored in regards to terrestrial prey subsidies for stream fishes. To address this, we examined the effects of an invasive riparian tree, European bird cherry (EBC, Prunus padus), spreading along urban Alaskan salmon streams, by collecting terrestrial invertebrates present on the foliage of riparian trees, their subsidies to streams, and their consumption by juvenile coho salmon (Oncorhynchus kisutch). Riparian EBC supported four to six times less terrestrial invertebrate biomass on its foliage and contributed two to three times lower subsidies relative to native deciduous trees. This reduction in terrestrial invertebrate biomass was consistent between two watersheds over 2 years. In spite of this reduction in terrestrial prey resource input, juvenile coho salmon consumed similar levels of terrestrial invertebrates in stream reaches bordered by EBC. Although we did not see ecological effects extending to stream salmonids, reduced terrestrial prey subsidies to streams are likely to have negative consequences as EBC continues to spread.

Integrative research is increasingly a priority within the scientific community and is a central goal for the evolving field of sustainability science. While it is conceptually attractive, its successful implementation has been challenging and recent work suggests that the move towards interdisciplinarity and transdisciplinarity in sustainability science is being only partially realized. To address this from the perspective of social-ecological systems (SES) research, we examine the process of conducting a science of integration within the Southcentral Alaska Test Case (SCTC) of Alaska-EPSCoR as a test-bed for this approach. The SCTC is part of a large, 5 year, interdisciplinary study investigating changing environments and adaptations to those changes in Alaska. In this paper, we review progress toward a science of integration and present our efforts to confront the practical issues of applying proposed integration frameworks. We: (1) define our integration framework; (2) describe the collaborative processes, including the co-development of science through stakeholder engagement and partnerships; and (3) illustrate potential products of integrative, social-ecological systems research. The approaches we use can also be applied outside of this particular framework. We highlight challenges and propose improvements for integration in sustainability science by addressing the need for common frameworks and improved contextual understanding. These insights may be useful for capacity-building for interdisciplinary projects that address complex real-world social and environmental problems.

Freshwater growth of juvenile sockeye salmon (Oncorhynchus nerka) depends upon the quality and quantity of prey and interactions with potential competitors in the foraging environment. To a large extent, knowledge about the ecology of lake-rearing juvenile sockeye salmon has emerged from studies of commercially important runs returning to deep nursery lakes, yet information from shallow nursery lakes (mean depth ≤ 10 m) is limited. We examined seasonal and ontogenetic variation in diets of juvenile sockeye salmon (N = 219, 30–85 mm) and an abundant potential competitor, threespine stickleback (Gasterosteus aculeatus; N = 198, 42–67 mm), to understand their foraging ecology and potential trophic interactions in a shallow Alaska lake. This study revealed that adult insects made up 74% of all sockeye salmon diets by weight and were present in 98% of all stomachs in Afognak Lake during the summer of 2013. Diets varied temporally for all fishes, but small sockeye salmon (

Summary Surface water connectivity can influence the richness and composition of fish assemblages, particularly in harsh environments where colonisation factors and access to seasonal refugia are required for species persistence. Studies regarding influence of connectivity on Arctic fish distributions are limited and are rarely applied to whole assemblage patterns. To increase our understanding of how surface water connectivity and related hydrologic variables influence assemblage patterns, we investigated species richness and composition of Arctic lake fishes over a large region, 8500 km2, of the central Arctic Coastal Plain, Alaska. We collected fish presence/non-detection data from 102 lakes and used a hierarchical multispecies occupancy framework to derive species richness and inform species composition patterns. Our mean estimate of regional richness was 12.3 (SD 0.5) species. Presence of a permanent channel connection was an overriding factor affecting species richness (mean 3.6, 95% CI 3.1–4.9), presumably driving lake colonisation potential. In lakes without a permanent channel connection, data suggest richness (mean 2.0, 95% CI 1.7–3.3) increased with the availability of in-lake winter refugia and with the potential of ephemeral connections during spring floods. Fish species functional traits and environmental faunal filters contributed to patterns of richness and assemblage composition. Composition corresponded with richness in a coherent manner, where each successive level of richness contained several discrete assemblages that showed similar responses to the environment. Lakes with permanent channel connections contained both widespread and restricted species, while the species-poor lakes that lacked a connection contained mainly widespread species. This work provides useful baseline information on the processes that drive the relations between patch connectivity and fish species richness and assemblage composition. The environmental processes that organise fish assemblages in Arctic lakes are likely to change in a warming climate.

The need for theories that address food web assembly and complexity over multiple spatial scales are critical to understanding their stability and persistence. In a meta-food web – an integrated network of local food webs – spatial heterogeneity in physical processes may have profound effects on food web function and energy flow. In the Arctic, surface water connectivity plays a vital role in determining fish assemblage composition, and potentially, food web structure. We examined lentic food web complexity associated with heterogeneity in surface water connectivity among Arctic lakes at the local scale, by contrasting lakes over a stream-lake connectivity gradient, and at the regional scale, by contrasting two locations with different surface water conditions (i.e., wet and dry) on the Arctic Coastal Plain of Alaska. Among lakes and across locations, increased hydrologic connectivity between streams and lakes increased the number of fish species and increased the complexity of the food web. The interaction of the region's hydrologic connectivity, local stream-lake connections, and the trophic niches of relevant fish species produced integrated, complex meta-food webs. Fully understanding mechanisms that support meta-food web stability are crucial when assessing future changes to Arctic stream-lake networks and the function and persistence of aquatic food webs.

In Arctic ecosystems, freshwater fish migrate seasonally between productive shallow water habitats that freeze in winter and deep overwinter refuge in rivers and lakes. How these movements relate to seasonal hydrology is not well understood. We used passive integrated transponder tags and stream wide antennae to track 1035 Arctic grayling in Crea Creek, a seasonally flowing beaded stream on the Arctic Coastal Plain, Alaska. Migration of juvenile and adult fish into Crea Creek peaked in June immediately after ice break-up in the stream. Fish that entered the stream during periods of high flow and cold stream temperature traveled farther upstream than those entering during periods of lower flow and warmer temperature. We used generalized linear models to relate migration of adult and juvenile fish out of Crea Creek to hydrology. Most adults migrated in late June – early July, and there was best support (Akaike weight = 0.46; w i ) for a model indicating that the rate of migration increased with decreasing discharge. Juvenile migration occurred in two peaks; the early peak consisted of larger juveniles and coincided with adult migration, while the later peak occurred shortly before freeze-up in September and included smaller juveniles. A model that included discharge, minimum stream temperature, year, season, and mean size of potential migrants was most strongly supported (w i = 0.86). Juvenile migration rate increased sharply as daily minimum stream temperature decreased, suggesting fish respond to impending freeze-up. We found fish movements to be intimately tied to the strong seasonality of discharge and temperature, and demonstrate the importance of small stream connectivity for migratory Arctic grayling during the entire open-water period. The ongoing and anticipated effects of climate change and petroleum development on Arctic hydrology (e.g. reduced stream connectivity, earlier peak flows, increased evapotranspiration) have important implications for Arctic freshwater ecosystems.

Yukon River Chinook salmon (Oncorhynchus tshawytscha) populations are declining for unknown reasons, creating hardship for thousands of stakeholders in subsistence and commercial fisheries. An informed response to this crisis requires understanding the major sources of variation in Chinook salmon productivity. However, simple stock–recruitment models leave much of the variation in this system’s productivity unexplained. We tested adding environmental predictors to stock–recruitment models for two Yukon drainage spawning streams in interior Alaska — the Chena and Salcha rivers. Low productivity was strongly associated with high stream discharge during the summer of freshwater residency for young-of-the-year Chinook salmon. This association was more consistent with the hypothesis that sustained high discharge negatively affects foraging conditions than with acute mortality during floods. Productivity may have also been reduced in years when incubating eggs experienced major floods or cold summers and falls. These freshwater effects — especially density dependence and high discharge — helped explain population declines in both rivers. They are plausible as contributors to the decline of Chinook salmon throughout the Yukon River drainage.

We critically evaluate some of the key ecological assumptions underpinning the use of nutrient replacement as a means of recovering salmon populations and a range of other organisms thought to be linked to productive salmon runs. These assumptions include: (1) nutrient mitigation mimics the ecological roles of salmon, (2) mitigation is needed to replace salmon-derived nutrients and stimulate primary and invertebrate production in streams, and (3) food resources in rearing habitats limit populations of salmon and resident fishes. First, we call into question assumption one because an array of evidence points to the multi-faceted role played by spawning salmon, including disturbance via redd-building, nutrient recycling by live fish, and consumption by terrestrial consumers. Second, we show that assumption two may require qualification based upon a more complete understanding of nutrient cycling and productivity in streams. Third, we evaluate the empirical evidence supporting food limitation of fish populations and conclude it has been only weakly tested. On the basis of this assessment, we urge caution in the application of nutrient mitigation as a management tool. Although applications of nutrients and other materials intended to mitigate for lost or diminished runs of Pacific salmon may trigger ecological responses within treated ecosystems, contributions of these activities toward actual mitigation may be limited.

We synthesized an expert review of climate change implications for hydroecological and terrestrial ecological systems in the northern coastal temperate rainforest of North America. Our synthesis is based on an analysis of projected temperature, precipitation, and snowfall stratified by eight biogeoclimatic provinces and three vegetation zones. Five IPCC CMIP5 global climate models (GCMs) and two representative concentration pathways (RCPs) are the basis for projections of mean annual temperature increasing from a current average (1961–1990) of 3.2 °C to 4.9–6.9 °C (5 GCM range; RCP4.5 scenario) or 6.4–8.7 °C (RCP8.5), mean annual precipitation increasing from 3130 mm to 3210–3400 mm (3–9 % increase) or 3320–3690 mm (6–18 % increase), and total precipitation as snow decreasing from 1200 mm to 940–720 mm (22–40 % decrease) or 720–500 mm (40–58 % decrease) by the 2080s (2071–2100; 30-year normal period). These projected changes are anticipated to result in a cascade of ecosystem-level effects including: increased frequency of flooding and rain-on-snow events; an elevated snowline and reduced snowpack; changes in the timing and magnitude of stream flow, freshwater thermal regimes, and riverine nutrient exports; shrinking alpine habitats; altitudinal and latitudinal expansion of lowland and subalpine forest types; shifts in suitable habitat boundaries for vegetation and wildlife communities; adverse effects on species with rare ecological niches or limited dispersibility; and shifts in anadromous salmon distribution and productivity. Our collaborative synthesis of potential impacts highlights the coupling of social and ecological systems that characterize the region as well as a number of major information gaps to help guide assessments of future conditions and adaptive capacity.

Freshwater fishes utilising seasonally available habitats within annual migratory circuits time movements out of such habitats with changing hydrology, although individual attributes of fish may also mediate the behavioural response to environmental conditions. We tagged juvenile Arctic grayling in a seasonally flowing stream on the Arctic Coastal Plain in Alaska and recorded migration timing towards overwintering habitat. We examined the relationship between individual migration date, and fork length (FL) and body condition index (BCI) for fish tagged in June, July and August in three separate models. Larger fish migrated earlier; however, only the August model suggested a significant relationship with BCI. In this model, 42% of variability in migration timing was explained by FL and BCI, and fish in better condition were predicted to migrate earlier than those in poor condition. Here, the majority (33%) of variability was captured by FL with an additional 9% attributable to BCI. We also noted strong seasonal trends in BCI reflecting overwinter mass loss and subsequent growth within the study area. These results are interpreted in the context of size and energetic state-specific risks of overwinter starvation and mortality (which can be very high in the Arctic), which may influence individuals at greater risk to extend summer foraging in a risky, yet prey rich, habitat. Our research provides further evidence that heterogeneity among individuals within a population can influence migratory behaviour and identifies potential risks to late season migrants in Arctic beaded stream habitats influenced by climate change and petroleum development.

European bird cherry (Prunus padus) (EBC) is an invasive ornamental tree that is spreading rapidly in riparian forests of urban Alaska. To determine how the spread of EBC affects leaf litter processing by aquatic invertebrate shredders, we conducted complementary leaf pack experiments in two streams located in Anchorage, Alaska. The first experiment contrasted invasive EBC with three native tree species—thin-leaf alder (Alnus tenuifolia), paper birch (Betula neoalaskana), and black cottonwood (Populus trichocarpa)—in one reach of Chester Creek; finding that EBC leaf litter broke down significantly faster than birch and cottonwood, but at a similar rate to alder. The second experiment contrasted EBC with alder in four reaches of Campbell and Chester creeks; finding that while EBC leaf litter broke down significantly faster than alder in Chester Creek, EBC broke down at a similar rate to alder in Campbell Creek. Although EBC sometimes supported fewer shredders by both count and mass, shredder communities did not differ significantly between EBC and native plants. Collectively, these data suggest that invasive EBC is not currently exhibiting strong negative impacts on leaf litter processing in these streams, but could if it continues to spread and further displaces native species over time.

Thaw ponds on the Arctic Coastal Plain of Alaska are productive ecosystems, providing habitat and food resources for many fish and bird species. Permafrost in this region creates unique pond morphologies: deep troughs, shallow low-centred polygons (LCPs) and larger coalescent ponds. By monitoring seasonal trends in pond volume and chemistry, we evaluated whether pond morphology and size affect water temperature and desiccation, and nitrogen (N) and phosphorus (P) fluxes. Evaporation was the largest early-summer water flux in all pond types. LCPs dried quickly and displayed high early-summer nutrient concentrations and losses. Troughs consistently received solute-rich subsurface inflows, which accounted for 12 to 42 per cent of their volume and may explain higher P in the troughs. N to P ratios increased and ammonium concentrations decreased with pond volume, suggesting that P and inorganic N availability may limit ecosystem productivity in older, larger ponds. Arctic summer temperatures will likely increase in the future, which may accelerate mid-summer desiccation. Given their morphology, troughs may remain wet, become warmer and derive greater nutrient loads from their thawing banks. Overall, seasonal- to decadal-scale warming may increase ecosystem productivity in troughs relative to other Arctic Coastal Plain ponds. Copyright © 2014 John Wiley & Sons, Ltd.

Drift-feeding fish are challenged to discriminate between prey and similar-sized particles of debris, which are ubiquitous even in clear-water streams. Spending time and energy pursuing debris mistaken as prey could affect fish growth and the fitness potential of different foraging strategies. Our goal was to determine the extent to which debris influences drift-feeding fish in clear water under low-flow conditions when the distracting effect of debris should be at a minimum. We used high-definition video to measure the reactions of drift-feeding juvenile Chinook salmon (Oncorhynchus tshawytscha) to natural debris and prey in situ in the Chena River, Alaska. Among all potential food items fish pursued, 52 % were captured and quickly expelled from the mouth, 39 % were visually inspected but not captured, and only 9 % were ingested. Foraging attempt rate was only moderately correlated with ingestion rate (Kendall's τ=0.55), raising concerns about the common use of foraging attempts as a presumed index of foraging success. The total time fish spent handling debris increased linearly with foraging attempt rate and ranged between 4 and 25 % of total foraging time among observed groups. Our results help motivate a revised theo- retical view of drift feeding that emphasizes prey detection and discrimination, incorporating ideas from signal detec- tion theory and the study of visual attention in cognitive ecology. We discuss how these ideas could lead to better explanations and predictions of the spatial behavior, prey selection, and energy intake of drift-feeding fish.

Predation on returning runs of adult salmon (Oncorhynchus spp.) can have a large influence on their spawning success. At McNeil River State Game Sanctuary (MRSGS), Alaska, brown bears (Ursus arctos) congregate in high numbers annually along the lower McNeil River to prey upon returning adult chum salmon (O. keta). Low chum salmon escapements into McNeil River since the late 1990s have been proposed as a potential factor contributing to concurrent declines in bear numbers. The objective of this study was to determine the extent of bear predation on chum salmon in McNeil River, especially on pre-spawning fish, and use those data to adjust the escapement goal for the river. In 2005 and 2006, 105 chum salmon were radiotagged at the river mouth and tracked to determine cause and location of death. Below the falls, predators consumed 99% of tagged fish, killing 59% of them before they spawned. Subsequently, the escapement goal was nearly doubled to account for this pre-spawning mortality and to ensure ...

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.

Artificial additions of nutrients of differing forms such as salmon carcasses and analog pellets (i.e. pasteurized fishmeal) have been proposed as a means of stimulating aquatic productivity and enhancing populations of anadromous and resident fishes. Nutrient mitigation to enhance fish production in stream ecosystems assumes that the central pathway by which effects occur is bottom‐up, through aquatic primary and secondary production, with little consideration of reciprocal aquatic‐terrestrial pathways. The net outcome (i.e. bottom‐up vs. top‐down) of adding salmon‐derived materials to streams depend on whether or not these subsidies indirectly intensify predation on in situ prey via increases in a shared predator or alleviate such predation pressure. We conducted a 3‐year experiment across nine tributaries of the N. Fork Boise River, Idaho, USA, consisting of 500‐m stream reaches treated with salmon carcasses (n = 3), salmon carcass analog (n = 3), and untreated control reaches (n = 3). We observed 2–8 fold increases in streambed biofilms in the 2–6 weeks following additions of both salmon subsidy treatments in years 1 and 2 and a 1.5‐fold increase in standing crop biomass of aquatic invertebrates to carcass additions in the second year of our experiment. The consumption of benthic invertebrates by stream fishes increased 110–140% and 44–66% in carcass and analog streams in the same time frame, which may have masked invertebrate standing crop responses in years 3 and 4. Resident trout directly consumed 10.0–24.0 g·m−2·yr−1 of salmon carcass and

We examined how marine-derived nutrients (MDN), in the form of spawning Pacific salmon, influenced the nutritional status and δ15N of stream-dwelling fishes. We sampled juvenile coho salmon ( Oncorhynchus kisutch ) and Dolly Varden ( Salvelinus malma ) during spring and fall from 11 south-central Alaskan streams that ranged widely in spawning salmon biomass (0.1–4.7 kg·m–2). Growth rate (as indexed by RNA–DNA ratios), energy density, and δ15N enrichment in spring-sampled fishes increased with spawner biomass, indicating the persistence of spawner effects more than 6 months after salmon spawning. Point estimates suggest that spawner effects on nutrition were substantially greater for coho salmon than Dolly Varden (268% and 175% greater for growth and energy, respectively), indicating that both species benefitted physiologically, but that juvenile coho salmon accrued more benefits than Dolly Varden. Although the data were less conclusive for fall- than spring-sampled fish, they do suggest spawner effects were also generally positive during fall, soon after salmon spawned. In a follow-up analysis where growth rate and energy density were modeled as a function of δ15N enrichment, results suggested that both increased with MDN assimilation, especially in juvenile coho salmon. Our results support the importance of salmon runs to the nutritional ecology of stream-dwelling fishes.

Estimating salmon escapement is one of the fundamental steps in managing salmon populations. The area-under-the-curve (AUC) method is commonly used to convert periodic aerial survey counts into annual salmon escapement indices. The AUC requires obtaining accurate estimates of stream life (SL) for target species. Traditional methods for estimating SL (e.g., mark–recapture) are not feasible for many populations. Our objective in this study was to determine the average SL of chum salmon Oncorhynchus keta in the McNeil River, Alaska, through radiotelemetry. During the 2005 and 2006 runs, 155 chum salmon were fitted with mortality-indicating radio tags as they entered the McNeil River and tracked until they died. A combination of remote data loggers, aerial surveys, and foot surveys were used to determine the location of fish and provide an estimate of time of death. Higher predation resulted in tagged fish below McNeil Falls having a significantly shorter SL (12.6 d) than those above (21.9 d). The st...

Wildfire and debris flows are important physical and ecological drivers in headwater streams of western North America. Past research has primarily examined short-term effects of these disturbances; less is known about longer-term impacts. We investigated wildfire effects on the invertebrate prey base for drift-feeding rainbow trout (Oncorhynchus mykiss, Walbaum) in Idaho headwater streams a decade after wildfire. Three stream types with different disturbance histories were examined: 1) unburned, 2) burned, and 3) burned followed by debris flows that reset channel morphology and riparian vegetation. The quantity of macroinvertebrate drift (biomass density) was more variable within than among disturbance categories. Average body weight and taxonomic richness of drift were significantly related to water temperature and influenced by disturbance history. During the autumn sampling period, the amount of terrestrial insects in rainbow trout diets varied with disturbance history and the amount of overhead canopy along the stream banks. Results indicate that there are detectable changes to macroinvertebrate drift and trout diet a decade after wildfire, and that these responses are better correlated with specific characteristics of the stream (water temperature, canopy cover) than with broad disturbance classes.

Land use activities often directly and indirectly limit the capacity of freshwater habitats to produce fish. Consequently, habitat creation and enhancement actions are often undertaken to increase the quantity and quality of resources available to aquatic communities within these impaired systems, with the intent to increase fish production. The objectives of this study were to (1) determine whether aquatic community colonization and development could be accelerated through additions of woody debris bundles and marine-derived nutrients (via salmon carcass analog pellets) and (2) measure how aquatic communities (biofilm, invertebrates, and fish) respond to these additions after the creation of off-channel (alcove) fish habitat in a stream in south-central Alaska. Biofilm, invertebrates, and juvenile coho salmon Oncorhynchus kisutch were sampled in four treatments (control, wood, analog, and analog plus wood). Biofilm chlorophyll-a concentrations were 4–10 times higher in analog-enriched treatments...

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

The energetic benefits enjoyed by consumers in streams with salmon runs depend on how those benefits are accrued. Adult Pacific salmon Oncorhynchus spp. deliver significant amounts of nutrients (i.e., nitrogen and phosphorus) and carbon to streams when they spawn and die; these nutrient additions can have demonstrable effects on primary production in streams. Consumption of carcass tissues or eggs provides for direct energy subsidies to consumers and may have significant effects on their condition. In this study, comparisons of juvenile coho salmon O. kisutch and aquatic insects exposed to terrestrial and marine energy sources demonstrated that direct consumption of marine-derived lipids had a significant effect on the lipid reserves of consumers. Direct consumption of marine-derived tissues was verified through fatty acid analysis. Selected aquatic insects and juvenile coho salmon were reared for 6 weeks in experimental streams supplied with terrestrial or marine energy sources. Chironomid midge...

Inorganic nutrient amendments to streams are viewed as possible restoration strategies for re-establishing nutrients and stream productivity throughout the western coast of North America, where salmon runs and associated marine-derived nutrient subsidies have declined. In a mesocosm experiment, we examined the short-term (6 weeks) comparative effects of artificial nutrient pellets and salmon carcasses, alone (low and high amounts) and in combination, on stream food webs. Response variables included dissolved nutrient concentrations, biofilm ash-free dry mass (AFDM) and chlorophyll-a levels, macroinvertebrate density, growth and body condition of juvenile coho salmon Oncorhynchus kisutch, and whole-body lipid content of invertebrates and juvenile coho salmon. Most of the response variables were significantly influenced by carcass treatment; the only response variable significantly influenced by fertilizer pellet treatment was soluble reactive phosphorus (SRP) concentration. Ammonium-nitrogen conce...