Diel migration pattern of Atlantic salmon (Salmo salar) and sea trout (Salmo trutta) smolts: an assessment of environmental cues

The timing of smolt migration is a key phenological trait with profound implications for individual survival during both river descent and the subsequent sea sojourn of anadromous fish. We studied relationships between the time of smolt migration, water temperature and light intensity for Atlantic salmon (Salmo salar) and sea trout (Salmo trutta). During 2006–2012, migrating smolts descending the southern Norway River Storelva were caught in a rotary screw trap located at the river mouth. The date of 50% cumulative smolt descent correlated significantly with the date when the river temperature exceeded 8°C for both Atlantic salmon and sea trout smolts. In 2010, smolts of both species were passive integrated transponder (PIT)‐tagged, and the diel timing of their migration was precisely documented. The degree of night migration decreased in both species as the river temperature rose, and at temperatures above 12–13°C, more smolts migrated during day than during night. A multinomial model was fitted for estimating temperature and species effects on probabilities of migration during night, daytime, dusk and dawn. Atlantic salmon smolts preferred migrating under lower light intensities than sea trout smolts during early, but not late spring when both species migrated during bright daylight. In accordance with the early‐season tendency to migrate at night, Atlantic salmon smolts migrated more during darker hours of the day than sea trout. In both species, smaller smolts migrated under dark conditions than during light conditions. Most of the findings on thermal, light and temporal effects on the observed smolt migration pattern can be explained as adaptations to predation avoidance.     

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.     

Stress-induced impairment of predator evasion and non-predator mortality in Pacific salmon

This study examined whether rapid recovery from stress-induced impairment in predator evasion, observed in previous studies on coho salmon, Oncorhynchus kisutch (Walbaum), was a general characteristic of different stocks of the same species and different species of Pacific salmon. We monitored stress-induced non-predator mortality, predator evasion and Cortisol concentrations of smolts of coho and spring Chinook, O. tshawytscha (Walbaum), after administration of standardized single and multiple handling stresses. Marked differences in the response to handling stress among stocks of coho and spring chinook smolts were evident, with recovery from impaired predator evasion occurring within 24 h of a 30-s handling stress for coho smolts and 24 h of a 1-min handling stress for spring chinook smolts. Differences in stress-induced non-predator mortality among stocks were also observed. The results point to the importance of screening hatchery salmonid stocks to assess differing capabilities of dealing with stress.     

A test of the cumulative effect of river weirs on downstream migration success,speed and mortality of Atlantic salmon (Salmo salar) smolts: An empirical study

This study investigated the cumulative impact of weirs on the downstream migration of wild Atlantic salmon (Salmo salar) smolts in the River Foyle, Northern Ireland. In spring of 2013 fish were released in two tributaries of similar length; one tributary (impacted) had seven low‐head weirs along the migration pathway and the other was devoid of such structures (un‐impacted). Salmon smolts fitted with acoustic transmitters were monitored via a passive acoustic telemetry array during downstream migration. In 2014 the study was repeated only in the impacted tributary. Overall freshwater survival rates were high (>94%). There was no significant difference in mortality, movement pattern, delay or travel speeds between rivers or between years at any phase of migration. Escapement of salmon smolts through Lough Foyle (a marine sea lough) to the open ocean was low, approximately 18% in each year. Escapement did not differ between impacted and un‐impacted rivers. This study showed no postpassage effects of weirs on mortality, migration speed or escapement of downstream migrating smolts. This suggests that the elevated mortality at low‐head obstacles described in other studies is not inevitable in all river systems. Migration through rivers with natural riffle‐pool migration may result in similar effects as those from low‐head weirs. Causes of apparent high mortality in the early part of marine migration in this study, are unknown; however similar studies have highlighted the impact of fish predators on smolts.     

Atlantic salmon smolts in the Irish Sea: First evidence of a northerly migration trajectory

Results from an acoustic telemetry study revealed for the first time a northerly migration route for Atlantic salmon (Salmo salar L.) smolts leaving the east coast of Ireland. Atlantic salmon smolts were tagged in spring 2019 in the Castletown and Boyne rivers. Three tagged smolts registered on disparate deep‐water offshore marine receivers as they travelled northwards out of the Irish Sea through the North Channel. One fish had migrated an estimated 250 km in a period of 32 days. The remaining two individuals were detected on receivers located off the Northern Ireland coast, further corroborating the northward migration of salmon smolts through the Irish Sea.     

Effects of release timing on migration behaviour and survival of hatchery‐reared Atlantic salmon smolts in a regulated river

In this study, 221 two‐year‐old hatchery‐reared salmon, Salmo salar, smolts were tagged with radio transmitters over a period of three consecutive years and released in the river in groups of 20–21 fish in various dates between late April and early June. Tagged smolts were tracked during their downstream migration in the lower 36‐km stretch of the regulated River Oulujoki, with the focus on the effects of release date, water temperature and river flow on migration behaviour and survival. The results indicate that release timing and river temperature have profound effects on the initiation of migration, swimming speed and survival of released S. salar smolts. Smolts released early in the spring in cold waters ceased migration after brief downstream movement and were vulnerable to predation, whereas the migration speed and survival rates increased markedly for smolts released later in the spring.     

Simulation of juvenile sockeye salmon (Oncorhynchus nerka) migrations in the Strait of Georgia, British Columbia

Previous research has documented two main migratory routes of juvenile sockeye salmon (Oncorhynchus nerka) through the Strait of Georgia, British Columbia, Canada, and large interannual variability in marine survival rates of the Chilko Lake stock. Simulation models were used to explore the influence of surface currents on the migratory route of juvenile sockeye salmon (smolts) through the Strait of Georgia. We used a model of downstream migration to generate daily numbers of Chilko Lake sockeye salmon smolts entering the Strait of Georgia, based on daily counts of smolts leaving the rearing lake. A numerical hydrodynamic model (driven by surface wind, tide, and Fraser River discharge) hindcasted surface currents in the Strait of Georgia on a 2 km × 2 km grid. A smolt migration model simulated fish moving through the Strait with different compass-oriented migratory behaviours (i.e. swimming speed and directional orientation) within the time-varying surface advection field. Results showed that surface currents within the Strait of Georgia can affect the migratory route of sockeye salmon smolts in spite of their large size (8 cm). Wind is the forcing mechanism primarily responsible for determining which migratory route would be used. Under prevailing wind conditions (i.e. toward the north-west), most sockeye salmon smolts would use the eastern migratory route; however, relatively brief south-eastward wind events (lasting about 2 days) would force most smolts into the western migratory route. Given the heterogeneity of food for salmon within the Strait, we hypothesize that wind-driven variability in the annual proportion of smolts that use the western and eastern migratory routes in the Strait of Georgia affects early marine survival rates of Fraser River sockeye salmon.     

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.     

Atlantic salmon living on the edge: Smolt behaviour and survival during seaward migration in River Minho

The Atlantic salmon (Salmo salar) population of the River Minho represents the southern natural distribution edge of the species. In line with the general trend for Atlantic salmon, this population has been declining over the years and is now at a critically low level. With river connectivity compromised by a large dam just 80 km upstream the River Minho's outlet, and an expected deterioration of climatic conditions, it is urgent to increase our knowledge of this population and identify survival bottlenecks that can be addressed. In this study, we used radio and acoustic telemetry to track Atlantic salmon smolts during their migration towards the sea and record both survival rates and possible causes of mortality. The recorded survival for the tagged migrating Atlantic salmon remained below 55% in the three studied years, indicating that the in‐river loss of smolts is likely a strong constraint to this population. From the smolts to which a likely cause of mortality could be attributed (34%), most appear to have been removed from the river (25%), with two confirmed events of bird predation and one of mammal predation. Interestingly, eight tags were recorded moving back upstream, likely indicating predation by larger fish. Increasing predator populations (e.g. cormorants, Phalacrocorax carbo) and invasive predators (e.g. American mink, Neovison vison) lead to elevated predation pressure on this already strained Atlantic salmon population, and further studies quantifying their impact in more detail could prove crucial for future management considerations.     

The effect of physiological and environmental conditions on smolt migration in Atlantic salmon Salmo salar

Hydropower development has negatively influenced Atlantic salmon (Salmo salar) populations. Compensatory hatchery‐rearing programmes exist, but released fish suffer from high mortality that may be related to the lack of experience from natural environments in hatchery‐reared smolts and their large body size and high energetic state. Here, we used acoustic telemetry to test how body size, energetic state, and the environmental conditions of the river affect migration in hatchery‐reared smolts. The study was conducted in three consecutive years between 2011 and 2013 in the lower part of the River Umeälven, Sweden. For individual fish, there was no effect of body size but the energetic state of the fish had a negative effect on sea entry. The most important factor affecting sea entry rate was the water discharge in the old river bed that differed among years. Smolts were more likely to enter the sea in years when the discharge was high or when the discharge increased substantially shortly after release. Hatchery‐reared fish had higher migration speed at a slower flowing section compared with a faster flowing section, which was likely a result of large hesitation to enter the rapid section. The increase in water discharge led to an increase in fish migration speed disproportional to the increase in water velocity. Our results highlight the importance of water discharge for the smolts during smolt migration, and we argue that concern should be given to migrating fish when managing regulated rivers.     

Estimation of infectious dose and viral shedding rates for infectious pancreatic necrosis virus in Atlantic salmon,Salmo salar L., post‐smolts

Infectious dose and shedding rates are important parameters to estimate in order to understand the transmission of infectious pancreatic necrosis virus (IPNV). Bath challenge of Atlantic salmon post‐smolts was selected as the route of experimental infection as this mimics a major natural route of exposure to IPNV infection. Doses ranging from 10² to 10^?4 50% end‐point tissue culture infectious dose (TCID₅₀) mL^?1 sea water were used to estimate the minimum infectious dose for a Scottish isolate of IPNV. The minimum dose required to induce infection in Atlantic salmon post‐smolts was <10^?1 TCID₅₀ mL^?1 by bath immersion (4 h at 10 °C). The peak shedding rate for IPNV following intraperitoneal challenge using post‐smolts was estimated to be 6.8 × 10³ TCID₅₀ h^?1 kg^?1 and occurred 11 days post‐challenge. This information may be incorporated into mathematical models to increase the understanding of the dispersal of IPNV from marine salmon sites.     

Stocking location and predation by marine fishes affect survival of hatchery‐reared Atlantic salmon smolts

Abstract Release strategies of hatchery‐reared Atlantic salmon, Salmo salar L., smolts were compared by studying survival and migration of smolts (n = 99) and their predators (Atlantic cod, Gadus morhua L., n = 8; and saithe, Pollachius virens (L.), n = 2) during the first 37 km of the marine migration using acoustic telemetry. Survivorship was higher in smolts released at the river mouth (30%) compared with smolts released in the river (12%). This was likely due to mortality or reduced migratory behaviour in fresh water. The marine mortality was 37% during the first 2 km after leaving the river (at least 25% mortality because of predation from marine fishes), and total marine mortality over 37 km was 68%. Detection‐depth data were useful for evaluating whether the tagged smolts were alive or predated; mortality during the first 2 km of outward migration would have been underestimated at 26% instead of 37% without the analysis of depth detection. Transmitters from consumed post‐smolts remained in predators for up to 47 days (average 29 days).     

Laboratory and field investigations of salmon lice [Lepeophtheirus salmonis (Krøyer)] infestation on Atlantic salmon (Salmo salar L.) post-smolts

Hatchery‐reared 1‐year‐old Atlantic salmon post‐smolts (Salmo salar L.), artificially infected with salmon lice [Lepeophtheirus salmonis (Krøyer)] copepodids, were found to suffer from primary alterations (increased cortisol levels) at early lice stages. Secondary alterations, such as osmotic stress (increased chloride levels), first occurred after the preadult stages of the lice appeared. Fish with the highest salmon lice infections died throughout the experiment. Seven years of field investigation of Trondheimsfjorden showed that Atlantic salmon post‐smolts descending coastal waters can become heavily infected with salmon lice. The migrating post‐smolts were only infected with the chalimus stages, showing that the fish had only recently left the rivers. The infection level, however, varied considerably between the years, and, in 1998, the infection was higher than previous years. The experimental results have been combined with the field data to appraise the consequences of the infection.     

Reservoir entrapment and dam passage mortality of juvenile Chinook salmon in the Middle Fork Willamette River

Abstract – High‐head dams in Oregon’s Willamette River basin inhibit seaward migration and present significant mortality risks to ESA‐listed juvenile Chinook salmon (Oncorhynchus tshawytscha). Over 7 years, we passively collected 13,365 downstream‐migrating juvenile salmon in rivers above and below Willamette dams. Most salmon emigrated from upstream sites in February–June, but passed dams in November–February when reservoirs were drawn down near annual lows, and access to deep‐water passage routes improved. Samples collected above reservoirs were dominated by subyearlings, whereas below‐dam samples were a phenotypically diverse mix of subyearling, yearling and older salmon. The life history data indicated that Willamette reservoirs seasonally entrap many salmon and some sea‐ready smolts probably residualise. Annual dam‐passage mortality estimates were 8–59% (mean = 26%). Individual salmon mortality risk increased significantly with body length and varied with reservoir elevation and discharge. Operational changes that allow timely volitional emigration and development of less hazardous passage routes would benefit this threatened population.     

Migration pathways,speed and mortality of Atlantic salmon (Salmo salar) smolts in a Scottish river and the near‐shore coastal marine environment

Long‐distance migration of Atlantic salmon (Salmo salar) is known to result in high levels of mortality. For a species experiencing global population decline, it is thus vital to better understand migration behaviour, both in the river and marine stages. Atlantic salmon smolts (n = 50) were tracked using acoustic telemetry in the River Deveron, Scotland, and adjacent coastal area. Higher rates of mortality were observed in the river (0.77% per km) than the early marine stage of migration (0.0% per km). Mortality likely resulted from predation. Higher swim speeds were recorded in the early marine stage compared with the river (marine = 7.37 ± 28.20 km/day; river = 5.03 ± 1.73 km/day [mean ± SD]), a potential predator avoidance behaviour. The majority of smolts leaving the river did so in darkness and on a flooding tide. Overall river and marine migration success were linked to nights of lower lunar brightness. Marine migration speed decreased with increasing environmental noise levels, a finding with implications for fisheries management. The migration pathway in the early marine environment did not follow obvious geographical features, such as the coastline. Thus, we suggest that early marine environment pathways are more influenced by complex water currents. These findings highlight factors that influence smolt migration survival and behaviour, areas on which future research should focus.     

Riverine and fjord migration of wild and hatchery‐reared Atlantic salmon smolts

Migration timing, speed, survival and effects of environmental parameters on migration, between wild and hatchery produced Atlantic salmon, Salmo salar L., smolts in the River Lærdalselva were studied. Hatchery‐reared (n = 40) and wild pre‐smolts (n = 40) were tagged with acoustic tags, and an array of receivers along the migration route was deployed. In all, 77 and 85% of the fish from the two groups, respectively, were recorded as migrating smolts, that is, predation rate and/or numbers of fish opting to remain in the river were low. Hatchery‐reared smolts showed a migration pattern, speed and migration route similar to wild smolts, even though the time period between river release and onset of migration was relatively short. Both groups of smolt showed high migration speed through both the river and the fjord compared with other studies.     

Warm oceans exacerbate Chinook salmon bycatch in the Pacific hake fishery driven by thermal and diel depth-use behaviours

Fisheries bycatch impacts marine species globally and understanding the underlying ecological and behavioural mechanisms could improve bycatch mitigation and forecasts in novel conditions. Oceans are rapidly warming causing shifts in marine species distributions with unknown, but likely, bycatch consequences. We examined whether thermal and diel depth-use behaviours influenced bycatch of a keystone species (Chinook salmon; Oncorhynchus tshawytscha, Salmonidae) in the largest fishery on the US West Coast (Pacific hake; Merluccius productus, Merlucciidae) with annual consequences in a warming ocean. We used Generalized Additive Models with 20 years of data including 54,509 hauls from the at-sea hake fishery spanning Oregon and Washington coasts including genetic information for five salmon populations. Our results demonstrate that Chinook salmon bycatch rates increased in warm ocean years explained by salmon depth-use behaviours. Chinook salmon typically occupy shallower water column depths compared to hake. However, salmon moved deeper when sea surface temperatures (SSTs) were warm and at night, which increased overlap with hake and exacerbated bycatch rates. We show that night fishing reductions (a voluntary bycatch mitigation strategy) are effective in reducing salmon bycatch in cool SSTs by limiting fishing effort when diel vertical movements bring salmon deeper but becomes less effective in warm SSTs as salmon seek deeper thermal refugia during the day. Thermal and diel behaviours were more pronounced in southern compared with northern salmon populations. We provide mechanistic support that climate change may intensify Chinook salmon bycatch in the hake fishery and demonstrate how an inferential approach can inform bycatch management in a changing world.     

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.     

Variability of preferred environmental conditions for Atlantic bluefin tuna (Thunnus thynnus) larvae in the Gulf of Mexico during 1993–2011

The Gulf of Mexico (GOM) is the primary spawning ground for western Atlantic bluefin tuna (Thunnus thynnus). In this work, information reported by previous studies about the preferred environmental conditions for the occurrence of bluefin tuna larvae in the GOM is integrated into a dimensionless index, the BFT_Index. This index is used to evaluate the spatial and temporal variability of areas with favorable environmental conditions for larvae within the GOM during 1993–2011. The main findings of this work are that: (i) the proposed index successfully captures the spatial and temporal variability in the in situ occurrence of bluefin tuna larvae; (ii) areas with favorable environmental conditions for larvae in the GOM exhibit year‐to‐year spatial and temporal variability linked with mesoscale ocean features and sea surface temperature; and (iii) comparison of the BFT_Index‐derived variability with recruitment of age‐0 fish estimated from recent stock assessment indicates that changes in environmental conditions may drive a relevant component (~58%) of the recruitment variability. The comparison with the recruitment dataset further revealed the existence of key regions linked with recruitment in the central/northern GOM, and that the Loop Current may function as a trap for larvae, possibly leading to low survival rates. Above (below) average conditions for occurrence of larvae in the GOM during spring were observed in 2000, 2001, 2002, 2006–2008, and 2011 (1994, 1996, 1998, 1999, 2003 and 2010). Results reported here have potential applications to assessment of bluefin tuna.     

Differences in sea migration between wild and reared Atlantic salmon (Salmo salar L.) in the Baltic Sea

The effect of origin, smolt size and year of release on the sea migration pattern of Atlantic salmon (Salmo salar L.) in the Baltic Sea was examined by tagging experiments conducted in 1991–1993 on wild and reared smolts of the Simojoki river salmon stock. The tag recovery data analysed by log-linear models revealed significant differences in both spatial and temporal sea migrations between the wild and reared salmon; the variation was attributed to the year of release and to the origin of the fish. Grilse accounted for the majority of reared returners (76%) but for a smaller proportion (46%) of the wild fish. The effect of smolt size could be studied only in the smolt groups tagged in 1991. Wild fish were more frequently (71%) caught in the Baltic Main Basin than were reared fish (51%) during their second sea year, and the size variation between wild and reared smolts did not explain the recovery site. No such differences in spatial distribution were found during the third sea year. The tagging place (hatchery/trap) of the reared fish did not affect their later sea migration. The differences in sea migration patterns suggest that the wild salmon are more vulnerable to the intensive salmon fishery in the Baltic Main Basin than are reared fish.