Ribosomal DNA sequences indicate isolated populations of Ichthyophonus hoferi in geographic sympatry in the north-eastern Pacific Ocean

Infections of Ichthyophonus hoferi, a cosmopolitan parasite of marine fish, have recently been reported in rockfish, Sebastes spp., from the north‐eastern Pacific. Because I. hoferi also infects Pacific herring, Clupea pallasi Valenciennes, and salmonids in this region, we wanted to determine if Ichthyophonus parasites from rockfishes, Pacific herring and chinook salmon, Oncorhynchus tshawytscha (Walbaum), were the same. Small subunit ribosomal deoxyribonucleic acid sequence data revealed two haplotypes that were fixed among host species in geographic sympatry, one from rockfish and the other from both Pacific herring and salmon. These isolated populations of Ichthyophonus could be part of the same species that are ecologically separated because of host behaviours, or they could be distinct species that are host specific. Dietary patterns of the hosts indicate that ecological separation among hosts is possible, but the presence of distinct species may better explain the observed Ichthyophonus haplotype association with host species.     

Brandt's cormorant diet (1994–2012) indicates the importance of fall ocean conditions for northern anchovy in central California

Effective ecosystem‐based management requires a comprehensive understanding of the functional links in the system. In many marine systems, forage species constitute a critical link between primary production and upper trophic level marine predators. As top predators, seabirds can be indicators of the forage species they consume and the ocean processes that influence these populations. We analyzed the diet and breeding success for the years 1994, 2003, 2005, and 2007–2012 of the Brandt's cormorant (Phalacrocorax penicillatus), a piscivorous diving seabird, breeding in central California, to evaluate the extent to which cormorant diet composition relates to prey availability, and how diet composition relates to breeding success and ocean conditions. Cormorant diet was primarily composed of young‐of‐the‐year (YOY) northern anchovy (Engraulis mordax), YOY rockfish (Sebastes spp.), and several species of small flatfish (order Pleuronectiformes). YOY rockfish consumption was positively related to their abundance as measured in a late spring pelagic midwater trawl survey. Northern anchovy appeared to be the most important prey as its consumption was positively related to cormorant breeding success. More northern anchovy were consumed in years where warm‐water conditions prevailed in the fall season before cormorant breeding. Thus, warm ocean conditions in the fall appear to be an important contributing factor in producing a strong year‐class of northern anchovy in central California and consequently a strong‐year class of Brandt's cormorant on the Farallon Islands.     

Interannual variation in pelagic juvenile rockfish (Sebastes spp.) abundance – going with the flow

We report results from 28 yr of a midwater trawl survey of pelagic juvenile rockfish (Sebastes spp.) conducted off the central California coast. The fishery‐independent survey is designed to provide pre‐recruit indices of abundance for use in groundfish stock assessments. Standardized catch rate time series for 10 species were developed from delta‐generalized linear models that include main effects for year, station, and calendar date. Results show that interannual fluctuations of all 10 species are strongly coherent but highly variable, demonstrating both high‐ and low‐frequency components. A similarly coherent result is observed in the size composition of fish, with large fish associated with elevated catch rates. In contrast, spatial and seasonal patterns of abundance show greater species‐specific differences. A comparison of the shared common trend in pelagic juvenile rockfish abundance, derived from principal components analysis, with recruitments from five rockfish stock assessments shows that the time series are significantly correlated. An examination of oceanographic factors associated with year‐to‐year variability indicates that a signature of upwelled water at the time of the survey is only weakly related to abundance. Likewise, basin‐scale indices (the Multivariate El Niño‐Southern Oscillation Index, the Pacific Decadal Oscillation, the North Pacific Gyre Oscillation, and the Northern Oscillation Index) are poorly correlated with abundance. In contrast, sea level anomalies in the months preceding the survey are well correlated with reproductive success. In particular, equatorward anomalies in the alongshore flow field following the spawning season are associated with elevated survival and poleward anomalies with poor survival.     

Prey selectivity and diel feeding chronology of juvenile chinook (Oncorhynchus tshawytscha) and coho (O. kisutch) salmon in the Columbia River plume

We studied salmon feeding selectivity and diel feeding chronology in the Columbia River plume. Juvenile chinook and coho salmon were caught by trawling at 2–3 h intervals throughout a diel period on three consecutive days (21–23 June 2000) at stations located 14.8 and 37 km offshore from the mouth of the Columbia River. A total of 170 chinook salmon were caught at the inshore and 79 chinook and 98 coho salmon were caught at the offshore station. After each trawl, potential prey were sampled at different depths with 2–3 different types of nets (1‐m diameter ring net, bongo net, neuston net). Despite the variability in zooplankton abundance, feeding selectivity was surprisingly constant. Both salmon species fed selectively on larger and pigmented prey such as hyperiid amphipods, larval and juvenile fish, various crab megalopae, and euphausiids. Hyperiid amphipods were abundant in the salmon diets and we hypothesize that aggregations of gelatinous zooplankton may facilitate the capture of commensal hyperiid amphipods. Small copepods and calyptopis and furcilia stages of euphausiids dominated the prey field by numbers, but were virtually absent from salmon diet. Juvenile chinook salmon, with increasing body size, consumed a larger proportion of fish. Stomach fullness peaked during morning hours and reached a minimum at night, suggesting a predominantly diurnal feeding pattern. In general, both chinook and coho salmon appear to be selective, diurnal predators, preying mostly on large and heavily pigmented prey items, in a manner consistent with visually oriented, size‐selective predation.     

The vertical distribution of juvenile salmon (Oncorhynchus spp.) and associated fishes in the Columbia River plume

Simultaneous trawling at surface and at depth at one location off the Columbia River, Oregon, in June 2000 identified the depth distribution of juvenile salmonids and associated fishes. Juvenile salmon off the Columbia River were distributed primarily near the surface, within the upper 12 m. Highest densities of subyearling chinook salmon (Oncorhynchus tshawytscha) off the Columbia River were associated with high surface currents and decreasing tidal levels, with time of day possibly a co‐factor. Densities of yearling chinook salmon increased with higher turbidity. Pacific herring (Clupea pallasi) was the most abundant and commonly caught forage fish, with density increasing at night, probably related to diel vertical migration. Catches of juvenile salmonids were not associated with catches of forage fishes. Daytime surface trawling appears to be an appropriate method for assessing the distribution and abundance of juvenile salmonids in marine habitats.     

Relative importance of chinook salmon abundance on resident killer whale population growth and viability

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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.     

Copepods and salmon: characterizing the spatial distribution of juvenile salmon along the Washington and Oregon coast,USA

Yearling Chinook (Oncorhynchus tshawytscha) and coho salmon (Oncorhynchus kisutch) were sampled concurrently with physical variables (temperature, salinity, depth) and biological variables (chlorophyll a concentration and copepod abundance) along the Washington and Oregon coast in June 1998–2008. Copepod species were divided into four different groups based on their water‐type affinities: cold neritic, subarctic oceanic, warm neritic, and warm oceanic. Generalized linear mixed models were used to quantify the relationship between the abundance of these four different copepod groups and the abundance of juvenile salmon. The relationships between juvenile salmon and different copepod groups were further validated using regression analysis of annual mean juvenile salmon abundance versus the mean abundance of the copepod groups. Yearling Chinook salmon abundance was negatively correlated with warm oceanic copepods, warm neritic copepods, and bottom depth, and positively correlated with cold neritic copepods, subarctic copepods, and chlorophyll a concentration. The selected habitat variables explained 67% of the variation in yearling Chinook abundance. Yearling coho salmon abundance was negatively correlated with warm oceanic copepods, warm neritic copepods, and bottom depth, and positively correlated with temperature. The selected habitat variables explained 40% of the variation in yearling coho abundance. Results suggest that copepod communities can be used to characterize spatio‐temporal patterns of abundance of juvenile salmon, i.e., large‐scale interannual variations in ocean conditions (warm versus cold years) and inshore‐offshore (cross‐shelf) gradients in the abundance of juvenile salmon can be characterized by differences in the abundance of copepod species with various water mass affinities.     

Predicting the effects of whale population recovery on Northeast Pacific food webs and fisheries: an ecosystem modelling approach

The recovery of whale populations from historical depletion may have the potential to noticeably affect Northeast Pacific ecosystems and fisheries. Surplus production models based on whaling catch records were used to reconstruct the historical abundances of five large whale species in the waters surrounding Haida Gwaii, British Columbia. The results suggest that the local abundances of all five species were vastly higher before the onset of modern whaling. A comparison of ecosystem models representing the states of the local marine food web before and after full whale recovery indicates that abundant whales could consume large proportions of the annual production of their principal prey, ranging up to 87% for Pacific herring (Clupea pallasii) and 72% for piscivorous rockfish (Sebastes spp.). Dynamic modelling of the food web effects of whale recovery, including simulations of simultaneous top‐down and bottom‐up forcing and a Monte Carlo sensitivity analysis, revealed noticeable (～6–12%) top‐down effects on Pacific herring biomass owing to increased predation by humpback and fin whales. However, these effects cannot explain the magnitude of recent declines in local herring biomass. The dynamic modelling results also suggest that top‐down effects of whale recovery could result in reduced biomasses of large rockfish as a result of predation by sperm whales, as well as potential cascading effects on many demersal fish groups. These findings have numerous practical implications for ecosystem‐based fisheries management and whale conservation strategies in Northeast Pacific waters.     

Relationship between zooplankton abundance and the early marine life history of juvenile chum salmon <Emphasis Type="Italic">Oncorhynchus keta</Emphasis> in eastern Hokkaido,Japan

Interannual variations in abundance, timing of outmigration from rivers, growth rate and condition of juvenile chum salmon (Oncorhynchus keta) were studied in the Nemuro Strait (eastern Hokkaido, Japan) during 1999–2002 to establish a possible relationship to zooplankton abundance. The otolith microstructure of juveniles was examined each year in late June to determine their time and size at sea entry (i.e., outmigration), and to estimate the early marine growth rates. Salmon outmigration peaked in mid- or late May, which coincided, in three of the four study years, with the peak release of juveniles into rivers within the study area. Abundance, growth rate and condition of fish were higher in 2001, when—compared to other years—smaller fish experienced higher growth rates, coinciding with greater zooplankton abundance for that year. Our results suggest that high zooplankton abundance positively influenced juvenile chum salmon growth and the condition of the fish during their early marine life despite their small size at sea entry.     

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.     

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.     

Diving back in time: Extending historical baselines for yelloweye rockfish with Indigenous knowledge

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Food habits and marine survival of juvenile Chinook and coho salmon from marine waters of Southeast Alaska

Little is known about the food habits of juvenile Chinook (Oncorhynchus tshawytscha) and coho (Oncorhynchus kisutch) salmon in marine environments of Alaska, or whether their diets may have contributed to extremely high marine survival rates for coho salmon from Southeast Alaska and much more modest survival rates for Southeast Alaskan Chinook salmon. To address these issues, we documented the spatial and temporal variability of diets of both species collected from marine waters of Southeast Alaska during summers of 1997–2000. Food habits were similar: major prey items of both species included fishes, crab larvae, hyperiid amphipods, insects, and euphausiids. Multivariate analyses of diet composition indicated that the most distinct groups were formed at the smallest spatial and temporal scales (the haul), although groups also formed at larger scales, such as by month or habitat type. Our expectations for how food habits would influence survival were only partially supported. As predicted, Southeast Alaskan coho salmon had more prey in their stomachs overall [1.8% of body weight (BW)] and proportionally far fewer empty stomachs (0.7%) than either Alaskan Chinook (1.4% BW, 5.1% empty) or coho salmon from other regions. However, contrary to our expectations, coho salmon diets contained surprisingly few fish (49% by weight). Apparently, Alaskan coho salmon achieved extremely high marine survival rates despite a diet consisting largely of small, less energetically‐efficient crustacean prey. Our results suggest that diet quantity (how much is eaten) rather than diet quality (what is eaten) is important to marine survival.     

Pacific hake (Merluccius productus Ayres, 1855) hydrolysates as feed attractants for juvenile Chinook salmon (Oncorhynchus tshawytscha Walbaum, 1792)

Soybean meal (SBM) inclusion in salmonid diets can lower feed cost, but dramatically reduces growth and feed utilization, and increases mortality in juvenile chinook salmon Oncorhynchus tshawytscha, due to diminished diet palatability and/or other adverse physiological effects exerted by antinutritional factors in SBM. The objective of this study was to investigate whether commercial Antarctic krill meal Euphausia superba or hydrolysates enzymatically produced from Pacific hake Merluccius productus could reverse the negative palatability effects of SBM inclusion in juvenile chinook salmon diets. Diets without SBM or with SBM and no added feed attractant were used as positive and negative control diets respectively. Incorporation of 2% krill meal or Alcalase^®‐produced hydrolysates into SBM‐containing diets (20% of dry matter by isonitrogenous replacement of fishmeal) significantly (P < 0.05) increased feed intake, feed utilization, fish weight gain and thermal growth coefficient during a 5‐week trial. Nevertheless, the negative effects on fish performance incurred by dietary inclusion of 20% SBM could not be fully reversed, indicating that most of those effects were likely unrelated to palatability. This study demonstrates the potential for using Pacific hake hydrolysates as a dietary feed attractant for salmonid diets, and supports the need for further research to optimize its application for ideal fish performance.     

Comparing abundance‐based and tag‐based estimates of coho salmon marine survival

Obtaining reliable estimates of marine survival is essential for understanding anadromous salmon population dynamics. Two common approaches to estimating marine survival are (a) dividing abundance of returning adult salmon abundance by abundance of smolts from the same cohort, or (b) tagging a portion of the migrating smolts and estimating the return rate of tagged adults. This study compared these two approaches to estimating marine survival for coho salmon, Oncorhynchus kisutch (Walbaum), across multiple years in three California streams. Abundance‐based survival estimates were higher than tag‐based estimates; average estimates for the two techniques differed from 1.5‐fold to 7.4‐fold across streams. One likely cause for these divergent estimates is migration of juveniles from natal habitat before smolt trapping begins, resulting in an underestimate of smolt abundance and an overestimate of marine survival rate for the abundance‐based method. Estimates of marine survival obtained from abundance estimates and tag returns are not directly comparable.     

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.     

Linking habitat characteristics with juvenile density to quantify Salmo salar and Salmo trutta smolt production in the river Sävarån,Sweden

Abstract Habitat mapping along 85 km of river was related to juvenile (15 years of electric fishing) and smolt (3 years of screw‐trapping) abundance data to estimate salmon, Salmo salar L., and sea trout, Salmo trutta L., smolt production in the River Sävarån, northern Sweden. Spawning site selection by radio‐tagged salmon (n = 12) and sea trout (n = 4) was also assessed. Fifty‐one hectares of potential spawning and nursery habitat was found in the main stem river, representing 25% of the total river area. These areas were estimated to yield 1300–7580 salmon and 630–3540 sea trout smolts based on juvenile densities, equating with 3 years of screw‐trap data (2990–5080 salmon and 680–2520 trout smolts, respectively). A hypothetical maximum production of about 19 900 salmon smolts was predicted for the river at a density of 40, 0+ salmon 100 m^?2. Tracking adults during the spawning period identified optimal and potential reproductive areas.     

The GABA a agonist muscimol enhances locomotor activity, but does not alter the behavioural effects of CRH in juvenile spring chinook salmon (Oncorhynchus tshawytscha)†

The present study investigated the effects of manipulating the GABAergic system on locomotor activity in juvenile spring chinook salmon, Oncorhynchus tshawytscha. In addition, we evaluated whether the GABAergic system is important for mediating the behavioural effects of corticotropin-releasing hormone (CRH). An intracerebroventricular (ICV) injection of the GABA_a agonist muscimol caused an acute and dose dependent increase in locomotor activity in juvenile spring chinook salmon. ICV injections of the GABA_a antagonist bicuculline prevented the increase in activity when administered concurrently with muscimol. The GABA_b agonist baclofen had no effect on locomotor activity in this study. Furthermore, we found no evidence that the locomotor response to exogenous CRH was altered by the concurrent administration of muscimol or bicuculline. These results provide evidence to support the hypothesis that endogenous GABA within the central nervous system is involved in the control of locomotor activity in fish. The data also suggest that there is no interaction between the GABAergic system and CRH with regards to the control of locomotor activity in this species.