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.     

Can research on the early marine life stage of juvenile chum salmon <Emphasis Type="Italic">Oncorhynchus keta</Emphasis> forecast returns of adult salmon? A case study from eastern Hokkaido,Japan

To examine the efficacy of juvenile salmon research as a tool for forecasting adult returns, the results from a study on the early marine life stage of juvenile chum salmon, conducted in the Nemuro Strait during 1999–2002 (i.e., 1998–2001 brood years), were compared with the return rates of adult salmon. Among the four brood years, the 2000 brood year (i.e., salmon migrating to the sea in 2001) was previously reported as showing higher abundance, higher growth rate and better somatic condition during the coastal residency period. Consequently, we expected it to have the highest return rate, under a hypothesis that juvenile survival in coastal residency regulates brood-year strength. Contrary to this expectation, the 2000 brood year had almost the lowest return rate. Alternatively, a statistical model in which sea surface temperature during the first year of marine life and size at release were utilized as explanatory variables reconstructed the actual variability in return rates more accurately than that based on the early marine life stage. Possible reasons for the discrepancy between the results of the juvenile salmon research and adult returns are discussed, and we suggest improvements for future research on juvenile salmon.     

Environmental risk factors associated with amoebic gill disease in cultured salmon, Salmo salar L., smolts in Ireland

A 2-year study was carried out on amoebic gill disease (AGD) involving monthly samples of 1+ Atlantic salmon, Salmo salar L., smolts, histological assessment of the gills and analysis of environmental data. Gill pathology was seen before amoebae could be detected microscopically. These changes in gill integrity were associated with marine environmental conditions, particularly elevated ammonium, nitrite and chlorophyll levels. The results suggest that the environmental changes predispose salmon to colonization by amoebae and ciliates. High densities of histophagous scuticociliates were observed in the gills during periods of advanced gill pathology. A number of different amoebae were observed in close association with gill pathology. Neoparamoeba was not seen in high densities, nor was it associated with gill pathology, indicating that Neoparamoeba may not be the primary agent of the AGD in Irish salmonid culture.     

Life stage‐specific,stochastic environmental effects overlay density dependence in an Atlantic salmon population

Atlantic salmon populations appear to fluctuate stochastically through time. It is suspected that both density‐dependent and density‐independent factors cause these fluctuations but the relative importance of each, and the life stages at which they operate, is not well known. In this study, a long‐term data set on Atlantic salmon migrants returning to the Foyle catchment, Ireland, was used to determine the role of density‐dependent and life stage‐specific environmental factors regulating population size. A Ricker density‐dependent model showed that spawning adult population size significantly predicted variation in the resultant filial generation; however, a large amount of variation (ca. 68%) remained unexplained. It was shown that environmental factors were significant in explaining some of the remaining variance and that these influences were linked to specific life stages. Three life stages—spawning and incubation, fry emergence and marine survival—were shown to have significant environmental effects that resulted in changes in the returning cohort strength. It is concluded that these life stage‐specific environmental effects are likely to contribute to the stochastic variation in population size resulting from the application of traditional stock–recruitment models. The identification and quantification of these effects should allow improved model accuracy.     

Time‐varying relationships between ocean conditions and sockeye salmon productivity

Environmental change is occurring at unprecedented rates in many marine ecosystems. Yet, environmental effects on fish populations are commonly assumed to be constant across time. In this study, I tested whether relationships between ocean conditions and productivity of North American sockeye salmon (Oncorhynchus nerka) stocks have changed over the past six decades. Specifically, I evaluated the evidence for non‐stationary relationships between three widely used ocean indices and productivity of 45 sockeye salmon stocks using hierarchical Bayesian models. The ocean indices investigated were the Pacific Decadal Oscillation (PDO), North Pacific Gyre Oscillation (NPGO), and sea surface temperature (SST). I found partial support for time‐varying salmon–ocean relationships. Non‐stationary relationships were strongest for the NPGO and weaker for the SST and PDO indices. Productivity–NPGO correlations tended to shift gradually over time with opposite trends for stocks in British Columbia (B.C.) and western Alaska; for B.C. stocks, the NPGO correlations shifted from significantly negative prior to 1980 to significantly positive after 1990, whereas for western Alaska stocks, the correlations shifted from positive to negative. Productivity–SST correlations showed declining trends for B.C. and Gulf of Alaska stocks, that is, correlations became more negative (B.C.) or less positive (Gulf of Alaska) over time. For the PDO, correlations weakened during the 1980s for western Alaska and B.C. stocks. Overall, these results provide evidence for time‐varying relationships between salmon productivity and environmental conditions over six decades, highlighting the need to recognize that historical responses of salmon populations to environmental change may not be indicative of future responses.     

Effects of juvenile size at release and early marine growth on adult return rates for Hokkaido chum salmon (Oncorhynchus keta) in relation to sea surface temperature

Using path analyses, we investigated relationships between size at release from hatcheries, the early marine growth of juveniles, and adult return rates for chum salmon from five river stocks of Hokkaido, Japan, in relation to sea surface temperature during ocean residence. Marine growth was estimated using scales collected from 11 760 adults of age 0.3 (1980–2004). The growth and survival of each stock appeared to have a different suite of regulatory processes. Interannual variability in return rates was mainly regulated by size at release in two stocks from the Sea of Okhotsk. A similar relationship was found in one stock from the Sea of Japan, but growth during coastal residency also affected their return rates. In two stocks from the Pacific coast of Hokkaido, variability in return rates was not related to size at release or to the coastal growth of juveniles, but with offshore growth in the Sea of Okhotsk, the nursery area for juveniles after leaving Japanese coastal waters. Whereas coastal growth tended to be negatively correlated with size at release in some stocks, offshore growth was positively associated with the August–November sea surface temperature in all stocks. This study confirmed that mortality of juvenile salmon occurred in two phases, during the coastal residency and the late period of the growing season, but the relative importance of both phases varied by stock and region, which probably regulated year‐class strength of Hokkaido chum salmon.     

Evidence for spatial coherence in time trends of marine life history traits of Atlantic salmon in the North Atlantic

A hierarchical Bayesian life cycle model is presented that considers spatial covariation of marine life history traits of Atlantic salmon (Salmo salar) populations in the North Atlantic. The model is based on a collective analysis of the dynamics of 13 stock units (SUs) from two continental stock groups (CSGs) in North America and Southern Europe in a single hierarchical model over the period 1971–2014. The model sets up a new assessment framework for Atlantic salmon stocks. It also provides a framework to investigate the drivers of changes in Atlantic salmon population dynamics including disentangling the effects of fisheries from those of environmental factors in a hierarchy of spatial scales. It is used to test the hypothesis of a strong spatial synchrony in marine life history dynamics of Atlantic salmon populations. The trends in two key parameters associated with the early marine phase of the life cycle are estimated: (i) the marine survival during the first summer–autumn spent at sea and (ii) the proportion of fish maturing after the first winter at sea. The results provide evidence of a decline in the marine survival together with an increase in the proportion of fish that mature after the first winter at sea, common to all SUs. Our results show an increased coherence in the covariations of trends in these two marine life history traits related to geographic proximity of SUs which support the hypothesis of a coherent response of geographically proximate Atlantic salmon populations that likely share similar migration routes.     

Running the trophic gauntlet: Empirical support for reduced foraging success in juvenile salmon in tidally mixed coastal waters

The productivity of Fraser River sockeye salmon has declined in recent years, with 2019 being the lowest return on record. The cause of the decline is still not fully understood; however, bottom‐up drivers and trophic interactions during the early marine migration are considered to be important contributing factors. McKinnell et al. (Fisheries Oceanography, 23, 2014 and 322) developed a “trophic gauntlet hypothesis,” proposing that low biological productivity leads to an energy deficit from poor foraging opportunities in migrating salmon. When combined with poor foraging conditions in typically productive waters elsewhere on the migration, low marine survival may result. Our study examined prey availability and stomach fullness of juvenile sockeye salmon along the 120 km stretch of the coastal migration through the Discovery Islands and Johnstone Strait to determine whether this section of the migration is indeed food limited. We observed low stomach fullness throughout tidally mixed waters, providing empirical support for the trophic gauntlet hypothesis. Zooplankton abundance was high in these regions so it appeared that unfavourably small prey size may have been the cause of low foraging success. We also observed foraging hotspots at both ends of the gauntlet, indicating that such areas may be key feeding grounds for migratory salmon.     

Entry of adult Atlantic salmon into a tributary of the Aberdeenshire Dee,Scotland

Upstream counts of adult salmon in a tributary to the River Dee during autumn and winter were examined over a 13‐year period using an optical fish counter. Statistical analysis indicated that salmon were mainly primed to enter the tributary at a particular time of year, peaking at early December. However, environmental factors also had a large influence. Entry of salmon was initiated by high flow rates in the main stem of the River Dee. Increases in water level in the tributary itself were not needed to stimulate the entry of salmon. Moreover, a change in flow from 1 day to the next did not result in a response from salmon. Rather, they reacted only after more than a day of flow increase. There was no consistent threshold level of flow that triggered tributary entry. The upstream passage of salmon was reduced at low temperatures (below 3°C). The numbers of salmon migrating upstream showed a gradual increase as the temperatures increased up to 11°C, and then sharply decreased at higher temperatures. Female salmon migrated earlier by about 2 weeks on average than males. Increasingly, counters and traps are being installed on spawning tributaries to examine the state of salmon stocks. Caution should be exercised, however, in using upstream entry data from individual tributaries to provide an overall assessment of salmon abundance, as local conditions in the tributaries may affect the salmon counts in particular years.     

Trends in coho marine survival in relation to the regime concept

There was a synchronous and significant decrease in marine survival of coho salmon in the Strait of Georgia, Puget Sound, and off the coast from California to Washington after 1989. This large-scale, synchronous change indicates that trends in coho marine survivals were linked over the southern area of their distribution in the north-east Pacific, and that these linkages were associated with a common event. Indicators of large-scale climate change (the Aleutian Low Pressure Index) and of recent regional climate change (the April flows from the Fraser River) also changed abruptly about the same time. The synchrony of trends in marine survival of aggregates of coho stocks from three distinct marine areas and trends in climate indices implies that climate/ocean changes can have profound impacts on the population dynamics of coho salmon. The trend towards low marine survival may persist as long as the trends in the climate indicators do not change.     

Spatial variability in ocean‐mediated growth potential is linked to Chinook salmon survival

Early ocean survival of Chinook salmon, Oncorhynchus tshawytscha, varies greatly inter‐annually and may be the period during which later spawning abundance and fishery recruitment are set. Therefore, identifying environmental drivers related to early survival may inform better models for management and sustainability of salmon in a variable environment. With this in mind, our main objectives were to (a) identify regions of high temporal variability in growth potential over a 23‐year time series, (b) determine whether the spatial distribution of growth potential was correlated with observed oceanographic conditions, and (c) determine whether these spatial patterns in growth potential could be used to estimate juvenile salmon survival. We applied this method to the fall run of the Central Valley Chinook salmon population, focusing on the spring and summer period after emigration into central California coastal waters. For the period from 1988 to 2010, juvenile salmon growth potential on the central California continental shelf was described by three spatial patterns. These three patterns were most correlated with upwelling, detrended sea level anomalies, and the strength of onshore/offshore currents, respectively. Using the annual strength of these three patterns, as well as the overall growth potential throughout central California coastal waters, in a generalized linear model we explained 82% of the variation in juvenile salmon survival estimates. We attributed the relationship between growth potential and survival to variability in environmental conditions experienced by juvenile salmon during their first year at sea, as well as potential shifts in predation pressure following out‐migration into coastal waters.     

Energy dynamics of subyearling Chinook salmon reveal the importance of piscivory to short‐term growth during early marine residence

Variation in prey quantity and quality can influence growth and survival of marine predators, including anadromous fish that migrate from freshwater systems. The objective of this study was to examine the energy dynamics of subyearling Chinook salmon (Oncorhynchus tshawytscha) following freshwater emigration. To address this objective, a population of Chinook salmon and their marine prey were repeatedly sampled from June to September over 2 years in coastal waters off Oregon and Washington. Subyearlings from the same population were also reared under laboratory conditions. Using a bioenergetics model evaluated in the laboratory, we found that growth rate variability in the field was associated more with differences in northern anchovy (Engraulis mordax) consumption and less with variation in diet energy density or ocean temperature. Highest growth rates (2.43–3.22% body weight/day) occurred in months when anchovy biomass peaked, and the timing of peak anchovy biomass varied by year. Our results support a general pattern among subyearling Chinook salmon occurring from Alaska to California that feeding rates contribute most to growth rate variability during early marine residence, although dominant prey types can differ seasonally, annually, or by ecosystem. In the northern California Current, faster growth appears to be associated with the availability of age‐0 anchovy. Identifying factors that influence the seasonal development of the prey field and regulate prey quantity and quality will improve understanding of salmon growth and survival during early marine residence.     

Behavioural ecology at sea of Atlantic salmon (Salmo salar L.) kelts from a Newfoundland (Canada) river

For migratory fishes such as Atlantic salmon, temporal variation in the utilization of thermal and depth habitats in the marine environment is likely to occur at both broad and fine scales, reflecting both ecological and metabolic factors. To test this, we implanted 26 Atlantic salmon kelts (i.e., post‐spawners) descending Campbellton River, Newfoundland, with data storage tags that recorded date/time, internal and external temperatures, pressure, and light. The salmon that returned to Campbellton River after an average of 62 days at sea were consecutive spawning kelt. Detailed data were recovered from eight of these fish and revealed distinct patterns in their utilization of thermal and depth habitats. Water temperatures experienced over the period at sea showed a wide range for all fish (from <0 to nearly 20°C); however, there were two clear frequency modes; one at 6–7°C and the other at 11–12°C. The cooler mode was indicative of daytime profiles and the warmer mode of night‐time profiles. This corresponded with the depth profiles, which indicated that salmon dove more frequently below 5 m (mean ± SD = 23 ± 0.9 dives per day) and spent less time (approximately 18%) near the sea surface (<1 m) during the day than during the night (4 ± 0.4 dives per night; approximately 45% time <1 m). The diurnal pattern may be indicative of a reliance on vision for feeding at depth and the short duration of dives (<10 min) may be a strategy that allows salmon a metabolic advantage (e.g., swimming capacity) over their prey living constantly at depth in cool waters.     

Production of Atlantic salmon (Salmo salar) in fresh water and at sea at hgh and low densities

A basic management objective is to identify the production capacity of Atlantic salmon in both freshwater and marine environments. Production in the biological sense is the total elaboration of fish biomass over time, and it is a function of fish abundance, growth and survival. In this article, we consider the factors that affect production at low and high stock levels. Using data from New Brunswick (Miramichi and Restigouche Rivers) and Newfoundland (Western Arm Brook), we consider freshwater and marine production of salmon at low and high stock levels. For all life stages, freshwater and marine production are related to initial biomass of the year-class: thus, it is important to maintain adequate egg deposition. Production at high stock levels is stock-dependent and mortality is extremely variable, particularly in the marine environment.     

Spatial correlation patterns in coastal environmental variables and survival rates of salmon in the north-east Pacific Ocean

We examined spatial correlations for three coastal variables [upwelling index, sea surface temperature (SST), and sea surface salinity (SSS)] that might affect juvenile salmon ( Oncorhynchus spp.) during their early marine life. Observed correlation patterns in environmental variables were compared with those in survival rates of pink ( O. gorbuscha ), chum ( O. keta ), and sockeye ( O. nerka ) salmon stocks to help identify appropriate variables to include in models of salmon productivity. Both the upwelling index and coastal SST were characterized by strong positive correlations at short distances, which declined slowly with distance in the winter months, but much more rapidly in the summer. The SSS had much weaker and more variable correlations at all distances throughout the year. The distance at which stations were no longer correlated (spatial decorrelation scale) was largest for the upwelling index (> 1000 km), intermediate for SST (400–800 km in summer), and shortest for SSS (< 400 km). Survival rate indices of salmon showed moderate positive correlations among adjacent stocks that decreased to zero at larger distances. Spatial decorrelation scales ranged from approximately 500 km for sockeye salmon to approximately 1000 km for chum salmon. We conclude that variability in the coastal marine environment during summer, as well as variability in salmon survival rates, are dominated by regional scale variability of several hundred to 1000 km. The correlation scale for SST in the summer most closely matched the observed correlation scales for survival rates of salmon, suggesting that regional-scale variations in coastal SST can help explain the observed regional-scale covariation in survival rates among salmon stocks.     

An investigation of the biological basis of return variability for sockeye salmon (Oncorhynchus nerka) from Great Central and Sproat lakes,Vancouver Island

We tested whether variations in stock characteristics (spawner and smolt abundance) and biotic conditions (prey variability, predation, competition) during the early marine period explained variations in the return of sockeye salmon (Oncorhynchus nerka) to Great Central and Sproat lakes, adjacent lakes on the west coast of Vancouver Island. There are two freshwater age groups in each lake; fish spend 1 or 2 yrs in freshwater after hatching. We tested the influences of stock and biotic factors on the return of each of the two age groups from each of the two lakes. Results of regression analyses showed that prey biomass variability best explained the variation in return for all lake‐age groups. Euphausiid (Thysanoessa spinifera) and cladoceran (Evadne) prey biomass variability explained between 0.75 and 0.95 (adjusted R²) of the variation in return. There appear to be instances of a mismatch between the seasonality of prey productivity and the apparent critical period of feeding for juvenile sockeye.     

Mortality surveys as a tool for studying marine anaemia in seapen-reared Chinook salmon, Oncorhynchus tshawytscha (Walbaum)

Because of their rapid autolysis and post mortem contamination, the examination of dead seapen salmon has been considered of limited value for investigating disease in farmed salmon. Marine anaemia is a recently described plasmacytoid leukemia of farmed salmon that is diagnosed primarily by histological findings. The objectives of this study were to determine if marine anaemia could be reliably diagnosed in dead fish under commercial conditions and to identify the factors that affect the quality of data that are derived from mortality surveys. Surface moribund fish and dead salmon were recovered from four adjacent pens on 31 farm visits to commercial salmon farms over a 6-month period. Mortality surveys provided six times as many fish for gross pathological diagnosis and three times as many fish for reliable histological diagnosis of marine anaemia than did surveys of surface moribund salmon. The interval between recovery of dead fish from a pen most strongly influenced the quality of information derived from the mortality surveys. Mortality surveys proved to be more sensitive to finding the disease than did surface morbidity surveys. The results demonstrated that mortality surveys can provide reliable information, which can generate new insights not only into the impact and epidemiology of marine anaemia, but also for other fatal diseases of seapen salmon.     

Oceanic and atmospheric extremes motivate a new hypothesis for variable marine survival of Fraser River sockeye salmon

In spite of a relatively optimistic pre‐season forecast, the total return of adult sockeye salmon (Oncorhynchus nerka) to the Fraser River (British Columbia, Canada) in 2009 was the lowest recorded since quantitative records began in the late 1940s. A plausible mechanism is proposed that links a sequence of extreme oceanic and climatic events to poor marine survival. It began with record‐setting snow packs in the coastal mountain range during the winter of 2007 that led to the development of unprecedented oceanographic conditions in the spring of 2007 from Queen Charlotte Strait in central British Columbia to Southeast Alaska. When combined with equally extreme atmospheric anomalies in the region in the spring of 2007, with a winter wind regime persisting through July, a coastal surface ocean with characteristics that are known to be associated with lower marine survival was established. Most of the sockeye salmon that were expected to return to the Fraser River as adults in 2009 passed through this atypical ocean as juveniles on their migration to the open ocean in 2007. A trophic gauntlet hypothesis is proposed as a new paradigm to describe the oceanic environment faced by sockeye salmon after they emigrate northward from the Strait of Georgia. The hypothesis identifies a new type of high nutrient low chlorophyll region that can explain how oceanographic extremes at critical locations along the migration route beyond the Strait of Georgia can reduce marine survival in some years.     

Influence of climate and abundance on migration timing of adult Atlantic salmon (Salmo salar) among rivers in Newfoundland and Labrador

Evidence mounts for the influence of climate variability on temporal trends in the phenology of many organisms including various species of fish. Accordingly, we examined variation in adult Atlantic salmon Salmo salar run timing in thirteen Newfoundland and Labrador rivers where returns were monitored at fishways or fish‐counting fences. Run timing varied significantly among rivers with the median date of return differing by up to 5 weeks. Duration of runs was generally short with most adults returning over a period of three to 5 weeks. A mixed model analysis incorporating a first‐order autoregressive error structure was used to generalise changes in run timing among all monitored rivers. Results indicated that the median date of return has advanced by almost 12 days over a 35‐year interval from 1978 to 2012, while several individual rivers have advanced by almost 21 days. The influence of climate on median timing was evident when the simultaneous effects of both climate and salmon abundance were controlled. We found earlier runs associated with overall warmer climate conditions on the Newfoundland and Labrador Shelf. Results contrast with those from the north‐east Atlantic where Atlantic salmon are returning later in some rivers coincident with warming climate conditions.     

Retrospective analysis of Atlantic salmon (Salmo salar) marine growth and condition in the northwest Atlantic based on tag‐recovery data

The life history of North American Atlantic salmon (Salmo salar) is characterized by extensive round‐trip migrations between freshwater rearing habitats and marine feeding grounds off the coasts of Canada and Greenland. Growth is rapid during the marine migration, and growth rate and condition factor may be indicators of salmon health during this period. Growth data were evaluated from a tag‐recovery program conducted from 1969 to 1991 using hatchery‐reared Atlantic salmon smolts released in the Penobscot River, Maine, U.S.A. Information from recaptures of 3167 salmon that were at large in the marine environment for 1 month to 3 yr was analyzed. Length–weight measurements coupled with time‐at‐large data were used to estimate von Bertalanffy and allometric growth parameters specific to the marine phase. Variations in growth and condition factor in relation to smolt age, release date, and temperature conditions in the northwest Atlantic were also examined. The von Bertalanffy k parameter declined with ordinal release date, indicating faster growth rates during the first year of smolts released earlier in the spring. The 2‐yr‐old smolts had a larger k than 1‐yr‐old smolts, although 1‐yr‐old smolts grew to a larger asymptotic size. Sea surface temperature had variable effects on growth parameters and condition factor, with temperature at the beginning of the migration and in overwintering habitat during the first year at sea having the greatest influence on length–weight relationships. Determining the mechanisms that influence growth of individuals during the marine phase will help elucidate the factors responsible for historic growth trends, establishing a baseline for current research.