Changing salmon: An analysis of body mass,abundance, survival,and productivity trends across 45 years in Puget Sound

Pacific salmon and trout (Oncorhynchus spp., Salmonidae) of the Puget Sound region of Washington State, USA, have experienced recent and longer‐term (multidecadal) variability in abundance while supporting robust fisheries. As part of the post‐season salmon management process, population‐specific estimates of total adult abundance to Puget Sound (Strait of Juan de Fuca) for pink (O. gorbuscha), chum (O. keta), coho (O. kisutch), sockeye (O. nerka), and Chinook (O. tshawytscha) salmon and steelhead trout (O. mykiss) are calculated annually. We compiled annual estimates of body mass, abundance and survival of hatchery‐ and naturally produced salmon from 1970 to 2015 to compare spatial and temporal patterns across species. Average weights of adult salmon and steelhead returning to Puget Sound, with the exception of coho salmon, have decreased since the 1970s. Temporal trends in abundance, survival and productivity varied by species and origin (hatchery vs. naturally produced). Generally, abundance and survival rates of natural‐origin species decreased whereas those of hatchery‐produced species did not, which is in contrast with other studies' general conclusions of decreasing survival among Puget Sound salmonids. Species diversity has decreased in recent years, with salmonids that rely on a short freshwater rearing phase in the natural environment (hatchery‐produced fish and naturally produced pink and chum) representing >90% of total returns in most years. This new information reveals patterns of body size, abundance, survival and productivity across species, life history and rearing type over the past 45 years and, in doing so, demonstrates the strength in multidecadal, multifactor time series to critically evaluate salmonid species.     

Estimating thermal exposure of adult summer steelhead and fall Chinook salmon migrating in a warm impounded river

Rising river temperatures in western North America have increased the vulnerability of many Pacific salmon (Oncorhynchus spp.) populations to lethal and sublethal risks. There is a growing need to predict and manage such risks, especially for populations whose life history or geography increases the likelihood of warm‐water exposure. We estimated thermal exposure of adult summer steelhead (O. mykiss) and fall‐run Chinook salmon (O. tshawytscha) as they migrated through a warm (often > 20 °C), 157‐km reach of the impounded Snake River, Washington. Archival temperature loggers and radiotelemetry were used to reconstruct thermal histories for 50 steelhead and 21 salmon. Encountered temperature maxima were mostly inside dam fishways and ranged from 15.8 to 24.0 °C (mean = 19.6 °C) for steelhead and from 18.0 to 21.6 °C (19.9 °C) for salmon. Behavioural thermoregulation was evident for ~50% of steelhead and ~30% of salmon in one of three reservoirs. Degree days (DDs) calculated from archival tags ranged from 74 to 973 DDs (median = 130) for steelhead and from 56 to 220 DDs (133) for salmon. Models using river temperature data and fish migration times accurately estimated total DDs for both species except some steelhead with extended thermoregulation. In a predictive application, we estimated exposure for 10,104 steelhead and 9071 Chinook salmon with passive integrated transponder‐tag detections at dams and found considerable DD variability across individuals, species and years. This estimation method, combined with baseline thermal surveys and existing monitoring infrastructure, can help to address long‐standing questions about how warm‐water exposure affects Snake River salmon and steelhead phenology, bioenergetics, physiology, survival and reproductive success.     

Ecological processes influencing mortality of juvenile pink salmon (Oncorhynchus gorbuscha) in Prince William Sound,Alaska

Our collaborative work focused on understanding the system of mechanisms influencing the mortality of juvenile pink salmon (Oncorhynchus gorbuscha) in Prince William Sound, Alaska. Coordinated field studies, data analysis and numerical modelling projects were used to identify and explain the mechanisms and their roles in juvenile mortality. In particular, project studies addressed the identification of major fish and bird predators consuming juvenile salmon and the evaluation of three hypotheses linking these losses to (i) alternative prey for predators (prey‐switching hypothesis); (ii) salmon foraging behaviour (refuge‐dispersion hypothesis); and (iii) salmon size and growth (size‐refuge hypothesis). Two facultative planktivorous fishes, Pacific herring (Clupea pallasi) and walleye pollock (Theragra chalcogramma), probably consumed the most juvenile pink salmon each year, although other gadids were also important. Our prey‐switching hypothesis was supported by data indicating that herring and pollock switched to alternative nekton prey, including juvenile salmon, when the biomass of large copepods declined below about 0.2 g m^?3. Model simulations were consistent with these findings, but simulations suggested that a June pteropod bloom also sheltered juvenile salmon from predation. Our refuge‐dispersion hypothesis was supported by data indicating a five‐fold increase in predation losses of juvenile salmon when salmon dispersed from nearshore habitats as the biomass of large copepods declined. Our size‐refuge hypothesis was supported by data indicating that size‐ and growth‐dependent vulnerabilities of salmon to predators were a function of predator and prey sizes and the timing of predation events. Our model simulations offered support for the efficacy of representing ecological processes affecting juvenile fishes as systems of coupled evolution equations representing both spatial distribution and physiological status. Simulations wherein model dimensionality was limited through construction of composite trophic groups reproduced the dominant patterns in salmon survival data. In our study, these composite trophic groups were six key zooplankton taxonomic groups, two categories of adult pelagic fishes, and from six to 12 groups for tagged hatchery‐reared juvenile salmon. Model simulations also suggested the importance of salmon density and predator size as important factors modifying the predation process.     

Change in feeding ecology and trophic dynamics of Pacific salmon (Oncorhynchus spp.) in the central Gulf of Alaska in relation to climate events

The effects of climate events on the feeding ecology and trophic dynamics of Pacific salmon (Oncorhynchus spp.) in offshore waters of the central Gulf of Alaska were investigated during early summers (1994–2000), based on analyses of stomach contents, and carbon and nitrogen stable isotopes (δ¹³C and δ¹⁵N). Gonatid squids (mainly Berryteuthis anonychus) were the dominant prey of all salmon species except for chum salmon (O. keta). During the 1997 El Niño event and the 1999 La Niña event, squids decreased sharply in the diets of all Pacific salmon except coho salmon (O. kisutch) in the Subarctic Current, and chum salmon diets changed from gelatinous zooplankton (1995–97) to a more diverse array of zooplankton species. A δ¹³C and δ¹⁵N analysis indicated that all salmon species occupied the same branch of the food web in 1999–2000. We hypothesize that high‐seas salmon adapt to climate‐induced changes in their prey resources by switching their diets either within or between trophic levels. To understand the effects of climate change on Pacific salmon in the Gulf of Alaska, biological oceanographic research on B. anonychus and other important prey resources is needed.     

Assessing the diet of North American Atlantic salmon (Salmo salar L.) off the West Greenland coast using gut content and stable isotope analyses

Investigations on the marine feeding of Atlantic salmon (Salmo salar L.) in the Northwest Atlantic are limited compared with the Northeast Atlantic. Climate‐induced changes to food webs in Atlantic salmon feeding areas have been noted, alongside increased mortality despite a cessation of most marine fisheries. As forage efficiency may be hampering survival, it was important to address this knowledge gap. Atlantic salmon were sampled at three sites on the West Greenland coast (Sisimiut, Nuuk and Qaqortoq) between 2009 and 2011. Gut content and stable isotope analyses were combined to assess spatial and temporal differences in feeding. Capelin (Mallotus villosus) dominated the diet at Nuuk and Qaqortoq, whereas boreoatlantic armhook squid (Gonatus fabricii) was the dominant prey at Sisimiut. Hyperiid amphipods (Themisto spp.) and sand lance (Ammodytes spp.) were also important. Significant differences were found among sites for both gut contents and stable isotope analyses, with fewer differences evident temporally. Dietary differences were also evident across larger scales, with little overlap demonstrated with Northeast Atlantic diets and the emergence of boreoatlantic armhook squid as an important prey item over time. Atlantic salmon diets are frequently anchored on one or two prey items, on which they appear to specialize, but they will diversify to consume other available pelagic prey. Thus, Atlantic salmon are an opportunistic, generalist predator within the pelagic food web. The variability evident in diet suggests that the limited data available are insufficient to appropriately understand potential vulnerabilities that the species may have to ecosystem changes, and suggest further research is needed.     

Change in chum salmon (Oncorhynchus keta) stomach contents associated with fluctuation of pink salmon (O. gorbuscha) abundance in the central subarctic Pacific and Bering Sea

The abundance and stomach contents of salmonids (Oncorhynchus spp.) and the biomass of prey organisms were examined in the central subarctic Pacific and Bering Sea in the summer of 1991 and 1992. Salmonids were caught by surface longline using the same level of fishing effort. Chum (O. keta) and pink (O. gorbuscha) salmon were the predominant species, representing 44% and 36% sof the total catch (n = 1275) in 1991. In 1992, chum salmon composed 85% of the total catch (n = 603), but the catch of pink salmon decreased to 1% of the total catch due to the odd/even year fluctuation of Asian pink salmon abundance in the study area. It was found that chum salmon changed their dominant diet from gelatinous zooplankton (pteropods, appendicular-ians, jellyfishes, chaetognaths, polychaetes and unidentified materials) in 1991, when pink salmon were abundant, to a diet of crustaceans (euphausiids, cope-pods, amphipods, ostracods, mysids and decapods) in 1992, when pink salmon were less abundant. Local crustacean biomass (wet weight; mg m^-3) had significant negative correlation with the CPUE (catch number per 30 hachi) of pink salmon in 1991 (r = -0.586; P = 0.026) and that of chum salmon in 1992 (r =–0.616; P = 0.014). There may be a limitation in the available prey resource for production of salmonids.     

Use of juvenile salmon growth and temperature change indices to predict groundfish post age‐0 yr class strengths in the Gulf of Alaska and eastern Bering Sea

Juvenile marine growth (SW1) of salmon and a new temperature change (TC) index were evaluated as ecosystem indicators and predictors for the post age‐0 year class strength (YCS) of groundfish in the Gulf of Alaska (GOA) and eastern Bering Sea (EBS). Our hypothesis was that SW1, as measured on the scales of adult Pacific salmon (Oncorhynchus spp.), is a proxy for ocean productivity on the continental shelf, a rearing area for young salmon and groundfish. Less negative TC index values are the result of a cool late summer followed by a warm spring, conditions favorable for groundfish YCS. In the GOA, SW1 was a positive predictor of age‐1 pollock (Theragra chalcogramma), but not age‐2 sablefish (Anoplopoma fimbria) YCS, indicating that the growth of the Karluk River sockeye salmon that enter Shelikof Strait is a proxy for ocean conditions experienced by age‐0 pollock. Contrary to our hypotheses, the TC index was a negative predictor of GOA pollock YCS; and the SW1 a negative predictor of EBS pollock and cod YCS since the 1980s. Recent fisheries oceanography survey results provide insight into possible mechanisms to support the inverse SW1 and YCS relationship. For the EBS, the TC index was a significant positive predictor for pollock and cod YCS, supporting the hypothesis that a cool late summer followed by a warm spring maximizes the over‐wintering survival of pollock and cod (Gadus macrocephalus), especially since the 1980s. The TC and SW1 index showed value for the assessment of pollock and cod, but not sablefish.     

Interannual variability in the feeding and condition of subyearling Chinook salmon off Oregon and Washington in relation to fluctuating ocean conditions

Chinook salmon (Oncorhynchus tshawytscha) is one of several economically‐important species of salmon found in the Northeast Pacific Ocean. The first months at sea are believed to be the most critical for salmon survival, with the highest rate of mortality occurring during this period. In the present study, we examined interannual diet composition and body condition trends for late‐summer subyearling Chinook salmon caught off Oregon and Washington from 1998 to 2012. Interannual variability was observed in juvenile salmon diet composition by weight of prey consumed. Juvenile subyearling Chinook salmon were mainly piscivorous, with northern anchovy (Engraulis mordax) being especially important, making up half the diet by weight in some years. Annual diets clustered into two groups, primarily defined by their proportion of invertebrate prey (14% versus 39% on average). Diet composition was found to influence adult returns, with salmon from high‐invertebrate years returning in significantly larger numbers 2–3 yrs later. However, years that had high adult returns had overall lower stomach fullness and poorer body condition as juveniles, a counterintuitive result potentially driven by the enhanced survival of less fit individuals in better ocean conditions (top‐down effect). Ocean conditions in years with a higher percentage of invertebrates in salmon diets were significantly cooler from May to August, and bottom‐up processes may have led to a fall plankton community with a larger proportion of invertebrates. Our results suggest that the plankton community assemblage during this first fall may be critical in predicting adult returns of Chinook salmon in the Pacific Northwest.     

Effects of the North Pacific Current on the productivity of 163 Pacific salmon stocks

Horizontal ocean transport can influence the dynamics of higher‐trophic‐level species in coastal ecosystems by altering either physical oceanographic conditions or the advection of food resources into coastal areas. In this study, we investigated whether variability in two North Pacific Current (NPC) indices was associated with changes in productivity of North American Pacific salmon stocks. Specifically, we used Bayesian hierarchical models to estimate the effects of the north‐south location of the NPC bifurcation (BI) and the NPC strength, indexed by the North Pacific Gyre Oscillation (NPGO), on the productivity of 163 pink, chum, and sockeye salmon stocks. We found that for salmon stocks located in Washington (WA) and British Columbia (BC), both the BI and NPGO had significant positive effects on productivity, indicating that a northward‐shifted bifurcation and a stronger NPC are associated with increased salmon productivity. For the WA and BC regions, the estimated NPGO effect was over two times larger than the BI effect for pink and chum salmon, whereas for sockeye salmon the BI effect was 2.4 times higher than the NPGO. In contrast to WA and BC stocks, we found weak effects of both horizontal ocean transport processes on the productivity of salmon stocks in Alaska. Our results indicated that horizontal transport pathways might strongly influence population dynamics of Pacific salmon in the southern part of their North American ranges, but not the northern part, suggesting that different environmental pathways may underlie changes in salmon productivity in northern and southern areas for the species under consideration.     

Density‐dependent effects of eastern Kamchatka pink salmon (Oncorhynchus gorbuscha) and Japanese chum salmon (O. keta) on age‐specific growth of western Alaska chum salmon

Hatcheries release >4.5 billion juvenile Pacific salmon (Oncorhynchus spp.) into the North Pacific Ocean annually, raising concerns about competition with wild salmon populations. We used retrospective scale analysis to investigate how the growth of chum salmon (O. keta) from western Alaska is affected by the abundance of chum salmon from Japanese hatcheries and wild pink salmon (O. gorbuscha) from the Russian Far East. Over nearly five decades, the growth of Kuskokwim River chum salmon was negatively correlated with the abundance of Japanese hatchery chum salmon after accounting for the effects of sex and spring/summer sea‐surface temperature in the Bering Sea. An effect of wild eastern Kamchatka pink salmon abundance on the growth of Kuskokwim River salmon was detectable but modest compared to the intraspecific competitive effect. A decrease in Japanese hatchery chum salmon releases in 2011–2013 was not associated with increased growth of Bering Sea chum salmon. However, the abundance of wild chum salmon from the Russian Far East increased during that time, possibly obscuring reduced competition with hatchery chum salmon. Our results support previous evidence that chum salmon are affected by intraspecific competition, and to a lesser extent interspecific competition, in the North Pacific, underscoring that the effects of salmon hatchery production transcend national boundaries.     

Ecosystem controls of juvenile pink salmon (Onchorynchus gorbuscha) and Pacific herring (Clupea pallasi) populations in Prince William Sound,Alaska

Five years of field, laboratory, and numerical modelling studies demonstrated ecosystem‐level mechanisms influencing the mortality of juvenile pink salmon and Pacific herring. Both species are prey for other fishes, seabirds, and marine mammals in Prince William Sound. We identified critical time‐space linkages between the juvenile stages of pink salmon and herring rearing in shallow‐water nursery areas and seasonally varying ocean state, the availability of appropriate zooplankton forage, and the kinds and numbers of predators. These relationships defined unique habitat dependencies for juveniles whose survivals were strongly linked to growth rates, energy reserves, and seasonal trophic sheltering from predators. We found that juvenile herring were subject to substantial starvation losses during a winter period of plankton diminishment, and that predation on juvenile pink salmon was closely linked to the availability of alternative prey for fish and bird predators. Our collaborative study further revealed that juvenile pink salmon and age‐0 herring exploit very different portions of the annual production cycle. Juvenile pink salmon targeted the cool‐water, early spring plankton bloom dominated by diatoms and large calanoid copepods, whereas young‐of‐the‐year juvenile herring were dependent on warmer conditions occurring later in the postbloom summer and fall when zooplankton was composed of smaller calanoids and a diversity of other taxa. The synopsis of our studies presented in this volume speaks to contemporary issues facing investigators of fish ecosystems, including juvenile fishes, and offers new insight into problems of bottom‐up and top‐down control. In aggregate, our results point to the importance of seeking mechanistic rather than correlative understandings of complex natural systems.     

Influence of the marine abundance of pink (Oncorhynchus gorbuscha) and sockeye salmon (O. nerka) on growth of Ozernaya River sockeye

The length and weight of Russian sockeye ( Oncorhynchus nerka ) returning to the Ozernaya River (Kamchatka) was substantially reduced in years when the ocean abundances of Kamchatkan pink ( O. gorbuscha ) and sockeye salmon were high. We found that the density-dependent reduction in sockeye growth on a per-capita basis was greater for sockeye than for pink salmon. However, the overall effect of pink salmon abundance on sockeye growth was greater because of the higher numerical abundance of pink salmon. The strongest statistical relationships were found for sockeye from separate age groups; pooled data combining all age classes were statistically insignificant. We estimate that, if pink salmon were absent, the most strongly affected age group of sockeye salmon (2.1 males) would weigh twice as much at maturity than if pink salmon populations from eastern and western Kamchatka were both simultaneously at peak observed abundances. Trophic competition in the ocean between pink and sockeye salmon can therefore have a significant influence on the productivity of sockeye populations for the most strongly affected age groups. These effects are large enough that they should be explicitly considered in the management of salmon populations.     

Early ocean distribution of juvenile Chinook salmon in an upwelling ecosystem

Extreme variability in abundance of California salmon populations is often ascribed to ocean conditions, yet relatively little is known about their marine life history. To investigate which ocean conditions influence their distribution and abundance, we surveyed juvenile Chinook salmon (Oncorhynchus tshawytscha) within the California Current (central California [37°30′N) to Newport, Oregon (44°00′N]) for a 2‐week period over three summers (2010–2012). At each station, we measured chlorophyll‐a as an indicator of primary productivity, acoustic‐based metrics of zooplankton density as an indicator of potential prey availability and physical characteristics such as bottom depth, temperature and salinity. We also measured fork lengths and collected genetic samples from each salmon that was caught. Genetic stock identification revealed that the majority of juvenile salmon were from the Central Valley and the Klamath Basin (91–98%). We constructed generalized logistic‐linear negative binomial hurdle models and chose the best model(s) using Akaike's Information Criterion (AIC) to determine which covariates influenced the salmon presence and, at locations where salmon were present, determined the variables that influenced their abundance. The probability of salmon presence was highest in shallower waters with a high chlorophyll‐a concentration and close to an individual's natal river. Catch abundance was primarily influenced by year, mean fork length and proximity to natal rivers. At the scale of sampling stations, presence and abundance were not related to acoustic indices of zooplankton density. In the weeks to months after ocean entry, California's juvenile Chinook salmon population appears to be primarily constrained to coastal waters near natal river outlets.     

Foraging behaviour of juvenile pink salmon (Oncorhynchus gorbuscha) and size‐dependent predation risk

Two hypotheses related to effects of juvenile pink salmon (Oncorhynchus gorbuscha) foraging behaviour and size on their predation risk were evaluated using field data collected in Prince William Sound, Alaska 1 995–97. My results supported the hypothesis that low macrozooplankton density leads to dispersion of juvenile salmon from shallow nearshore habitats and greater predation risk, but zooplankton type was an important factor. When the biomass of large copepods (primarily Neocalanus spp.) declined, salmon dispersed from shallow nearshore habitats, and mean daily individual predator consumption of salmon increased by a factor of 5. A concomitant five‐fold increase in the probability of occurrence of salmon in predator stomachs supported the notion that increased predation on salmon was caused by a greater overlap between predator and prey when salmon dispersed offshore, not an increase in the number of salmon consumed per feeding bout. The results also generally supported the hypothesis that the timing of predation events modifies the nature of size‐dependent predation losses of salmon to different predator groups (small and large planktivores and piscivores). Size‐dependent vulnerabilities of salmon to predators were a function of both predator and prey sizes. When simulated predation was shifted from May to June, the vulnerability of salmon became more dependent on their growth than initial size. But, the size‐ and growth‐dependent vulnerabilities of salmon differed more among predator groups than between May and June, suggesting that changes in the composition of predator fields could more strongly affect the nature of size‐dependent predation than changes in the timing of predation losses.     

Effects of diets on the growth performance and shell pigmentation of Pacific abalone

This study investigated the effects of natural and artificial diets on growth performance and shell pigmentation of cultured abalone. A 7‐month feeding trial was conducted on 12 000 Pacific abalones (Haliotis discus hannai) with four diets including two extruder‐processed test diets (with & without addition of oleopaprika carotenoids), Pacific dulse (Palmaria mollis) and a combination of Pacific dulse and a test diet. The results showed that the two test diets resulted in higher survival of abalone, but with a lower growth rates when compared with Pacific dulse alone (P < 0.05). The combination diet achieved the highest survival and growth rates. The Pacific dulse resulted in abalone with dark‐brown shells, which are preferred by Asian markets. The test diets led to 52% and 55% of the animals with pink coloured shells and the supplementation of oleopaprika did not affect shell pigmentation. The aqueous acidic extracts from both dark‐brown and pink shells showed blue colour; HPLC‐MS chromatography revealed that the pigments in the extracts were consistent with a biliverdin and a cysteine‐biliverdin. These results are valuable for the development of abalone feed and the control of abalone shell colouration.     

Biogeography of pelagic food webs in the North Pacific

The tufted puffin (Fratercula cirrhata) is a generalist seabird that breeds throughout the North Pacific and eats more than 75 different prey species. Using puffins as samplers, we characterized the geographic variability in pelagic food webs across the subarctic North Pacific from the composition of ~10,000 tufted puffin meals (~56,000 prey items) collected at 35 colonies in the Gulf of Alaska (GoA) and Aleutian Archipelago. Cluster analysis of diet species composition suggested three distinct forage fish communities: (i) in the northern GoA, multiple age‐classes of coastal and shelf residents such as capelin, sand lance and herring dominated the food web, (ii) in the western GoA to eastern Aleutians, the shelf community was dominated by transient age‐0 walleye pollock, and (iii) in the western Aleutians, shelf‐edge and mesopelagic forage species such as squid, lanternfish, and Atka mackerel were prevalent. Geographic patterns of abundance of capelin and sand lance in tufted puffin diets were corroborated by independent research fisheries and diets of piscivorous fish, indicating that puffin diets reflect the local abundance of forage species, not just selection of favored species. Generalized additive models showed that habitat characteristics predict, in a non‐linear fashion, forage species distribution and abundance across two large marine ecosystems. We conclude that major biogeographic patterns in forage fish distribution follow gradients in key habitat features, and puffin diets reflect those patterns.     

Freshwater habitat associations between pink (Oncorhynchus gorbuscha), chum (O. keta) and Chinook salmon (O. tshawytscha) in a watershed dominated by sockeye salmon (O. nerka) abundance

To understand the interplay between habitat use and contemporary anadromous Pacific salmon, Oncorhynchus spp., distributions we explored the habitat associations of three species, pink (O. gorbuscha), chum (O. keta) and Chinook salmon (O. tshawytscha) in streams of the Wood River system of Bristol Bay, Alaska, where sockeye salmon (O. nerka) are numerically dominant. We developed models to investigate the occurrence of nondominant salmon in relation to habitat characteristics and sockeye salmon density, using four decades of salmon presence and abundance data. The frequency of occurrence and abundance of nondominant species increased with watershed drainage area and stream depth and decreased with sockeye salmon density. The range of occurrence varied from nonexistent to perennial for the other species in sockeye‐dominated streams. Increasing watershed area resulted in larger stream habitat area and deeper habitats, allowing for the sympatric occurrence and persistence of all salmon species. The relationships between habitat and the presence of these Pacific salmon help define their requirements but also remind us that the patterns of presence and absence, within the overall ranges of salmon species, have yet to be fully understood.     

A synthesis of the life history and ecology of juvenile Pacific herring in Prince William Sound,Alaska

Physical and biological variables affecting juvenile Pacific herring (Clupea pallasi) in Prince William Sound (PWS) from 1995 to 1998 were investigated as part of a multifaceted study of recruitment, the Sound Ecosystem Assessment (SEA) program. Though more herring larvae were retained in eastern PWS bays, ages‐0 and ‐1 herring used bays throughout PWS as nursery areas. Water transported into PWS from the Gulf of Alaska (GOA) contributed oceanic prey species to neritic habitats. Consequently, variations in local food availability resulted in different diets and growth rates of herring among bays. Summer food availability and possible interspecific competition for food in nursery areas affected the autumn nutritional status and juvenile whole body energy content (WBEC), which differed among bays. The WBEC of age‐0 herring in autumn was related to over‐winter survival. The limited amount of food consumption in winter was not sufficient to meet metabolic needs. The smallest age‐0 fish were most at risk of starvation in winter. Autumn WBEC of herring and winter water temperature were used to model over‐winter mortality of age‐0 herring. Differences in feeding and energetics among nursery areas indicated that habitat quality and age‐0 survival were varied among areas and years. These conditions were measured by temperature, zooplankton abundance, size of juvenile herring, diet energy, energy source (GOA vs. neritic zooplankton), WBEC, and within‐bay competition.     

Competitive release modifies the impacts of hydrologic alteration for a partially migratory stream predator

Understanding ecological effects of altered stream flows is an essential objective. In a comparative field study of juvenile steelhead trout (Oncorhynchus mykiss) that compared natural‐ and reduced‐flow sites, we used mark–recapture modelling and information theory to quantify spatially and temporally explicit patterns of density, specific growth rate, survival and outmigration; and test predictions for biotic and abiotic drivers. Densities were lower in water withdrawal treatments, resulting in lower intraspecific competition and, higher specific growth rate and survival. We observed yearly differences in density and intraspecific competition, with a negative relationship between density and specific growth rate over a wide range of densities, but reductions in survival only at the highest densities. Moreover, individual variability within sites was important. At high density (sites and years), survival related negatively to body size. In contrast, when overall density was lower, specific growth rate was negatively related to body size. Lastly, individuals were more likely to outmigrate when they had larger body size, lower survival or reared in habitats with reduced flows, and these patterns appeared mediated by the intensity of intraspecific competition. Our results underscore the harsh bioenergetic conditions induced by higher temperatures and densities during summer baseflow (relative to other seasons), particularly for larger fish demanding more resources, and suggest a density‐dependent mechanism for why this period is important for regulating salmonid populations. We found that a complex combination of natural (e.g., density) and anthropogenic (e.g., withdrawal) factors affected juvenile salmon populations and life history expression in the face of altered flows.     

Introduced northern pike predation on salmonids in southcentral Alaska

Northern pike (Esox lucius) are opportunistic predators that can switch to alternative prey species after preferred prey have declined. This trophic adaptability allows invasive pike to have negative effects on aquatic food webs. In Southcentral Alaska, invasive pike are a substantial concern because they have spread to important spawning and rearing habitat for salmonids and are hypothesised to be responsible for recent salmonid declines. We described the relative importance of salmonids and other prey species to pike diets in the Deshka River and Alexander Creek in Southcentral Alaska. Salmonids were once abundant in both rivers, but they are now rare in Alexander Creek. In the Deshka River, we found that juvenile Chinook salmon (Oncorhynchus tshawytscha) and coho salmon (O. kisutch) dominated pike diets and that small pike consumed more of these salmonids than large pike. In Alexander Creek, pike diets reflected the distribution of spawning salmonids, which decrease with distance upstream. Although salmonids dominated pike diets in the lowest reach of the stream, Arctic lamprey (Lampetra camtschatica) and slimy sculpin (Cottus cognatus) dominated pike diets in the middle and upper reaches. In both rivers, pike density did not influence diet and pike consumed smaller prey items than predicted by their gape‐width. Our data suggest that (1) juvenile salmonids are a dominant prey item for pike, (2) small pike are the primary consumers of juvenile salmonids and (3) pike consume other native fish species when juvenile salmonids are less abundant. Implications of this trophic adaptability are that invasive pike can continue to increase while driving multiple species to low abundance.