Influence of sea temperature and initial marine feeding on survival of Atlantic salmon Salmo salar post-smolts from the Rivers Orkla and Hals, Norway

The abundance of returning adult Atlantic salmon Salmo salar, in the River Orkla in mid‐norway (1 sea‐winter, SW, fish) and River Hals in north Norway (1–3 SW fish), was tested against the early marine feeding and the seawater temperature experienced by their corresponding year classes of post‐smolts immediately after entry into the Trondheimsfjord (Orkla smolts, 22 years of data) and Altafjord (Hals smolts, 17 years of data). In both river–fjord systems, there was a significant positive correlation between the abundance of returning S. salar and the mean seawater temperature at the time of smolts descending to the sea. The number of 1SW fish reported caught in River Orkla was positively correlated to the proportion of fish larvae in the post‐smolt stomachs in Trondheimsfjord. The abundance of returning S.salar was, however, neither correlated to forage ratio (R_F) nor other prey groups in post‐smolt stomachs in the two fjord systems. In the Altafjord, the post‐smolts fed mainly on pelagic fish larva (70–98%) and had a stable R_F (0·009–0·023) over the 6 years analysed. In the Trondheimsfjord, however, there was a higher variation in R_F (0·003–0·036), and pelagic fish larvae were dominant prey in only two (50 and 91%) of the 8 years analysed. These 2 years also showed the highest return rates of S. salar in River Orkla. These results demonstrate that the thermal conditions experienced by post‐smolts during their early sea migration may be crucial for the subsequent return rate of adults after 1–3 years at sea. Pelagic marine fish larvae seem to be the preferred initial prey for S. salar post‐smolts. As the annual variation in abundance of fish larvae is related to seawater temperature, it is proposed that seawater temperature at sea entry and the subsequent abundance of returning adult S. salar may be indirectly linked through variation in annual availability of pelagic fish larvae or other suitable food items in the early post‐smolt phase.     

Migratory behaviour and survival rates of wild northern Atlantic salmon Salmo salar post‐smolts: effects of environmental factors

To study smolt behaviour and survival of a northern Atlantic salmon Salmo salar population during river descent, sea entry and fjord migration, 120 wild S. salar were tagged with acoustic tags and registered at four automatic listening station arrays in the mouth of the north Norwegian River Alta and throughout the Alta Fjord. An estimated 75% of the post‐smolts survived from the river mouth, through the estuary and the first 17 km of the fjord. Survival rates in the fjord varied with fork length (L_F), and ranged from 97·0 to 99·5% km^?1. On average, the post‐smolts spent 1·5 days (36 h, range 11–365 h) travelling from the river mouth to the last fjord array, 31 km from the river mouth. The migratory speed was slower (1·8 L_F s^?1) in the first 4 km after sea entry compared with the next 27 km (3·0 L_F s^?1). Post‐smolts entered the fjord more often during the high or ebbing tide (70%). There was no clear diurnal migration pattern within the river and fjord, but most of the post‐smolts entered the fjord at night (66%, 2000–0800 hours), despite the 24 h daylight at this latitude. The tidal cycle, wind‐induced currents and the smolts' own movements seemed to influence migratory speeds and routes in different parts of the fjord. A large variation in migration patterns, both in the river and fjord, might indicate that individuals in stochastic estuarine and marine environments are exposed to highly variable selection regimes, resulting in different responses to environmental factors on both temporal and spatial scales. Post‐smolts in the northern Alta Fjord had similar early marine survival rates to those observed previously in southern fjords; however, fjord residency in the north was shorter.     

Sea trout Salmo trutta in the subarctic: home-bound but large variation in migratory behaviour between and within populations

Anadromous brown trout (sea trout), Salmo trutta, is currently in decline throughout its range, largely due to anthropogenic stressors in freshwater and marine habitats. Acoustic telmetry was utilized to study the marine migration of sea trout post-smolts from three populations in a relatively pristine subarctic fjord system. While at sea, the sea trout spent a substantial part of their time close to their natal river, preferred near shore over pelagic habitats and were strongly surface oriented. Despite a fidelity towards local areas, the sea trout utilized various parts of the fjord system, with maximum dispersion >30 km and total migration distance >300 km. Almost half of the sea trout (44%) migrated between river outlets, indicating that a metapopulation approach may be appropriate when managing neighbouring sea trout populations at high latitudes. Furthermore, the different populations displayed different migratory behaviours in terms of distance migrated, dispersion from origin and the likelihood of leaving their home area. This variation in migratory behaviour is likely influenced by spatiotemporal differences in habitat quality between sites, indicating that local habitat variations may promote population-specific behavioural responses even in relatively confined fjord systems.     

Time and size at seaward migration influence the sea survival of Salmo salar

Whether time of seaward migration of young Atlantic salmon Salmo salar influences their subsequent survival and growth was investigated in the River Imsa, south‐western Norway. Salmo salar were tagged when moving downstream through a trap near the outlet between 1976 and 2010 and recaptured on their adult return. Most descended as smolts in April and May, but some descended during the other months of the year. Annual variation in timing of the smolt migration was significantly correlated with variation in water temperature during spring. Mean total body length of the descending S. salar varied with month of seaward migration. The sea survival of S. salar emigrating from the River Imsa between January and May was 2·8 times higher than for those descending between June and December. The sea survival of the various cohorts decreased with increasing river temperature in April to May, prior to the smolt migration, and decreasing day number when the smolts moved to sea. The size of smolts descending the river between April and May did not affect the survival at sea as much as it affected the survival of migrants descending in any other month of the year. The majority of the downstream migrating S. salar were 2 years old, but proportionally, more 1 year olds moved downstream in the autumn than in the rest of the year. Mean duration between downstream migration of the young and the return migration of the grilse was shortest (12·7 months) for those descending in July and August and longest for those descending in October (21 months). Mean monthly specific growth rate was highest for those migrating downstream between May and July and lowest for those emigrating in September. Based on the present results, it was hypothesized that S. salar emigrating between April and August migrated directly out into the ocean, while those that emigrated between October and March stayed in the estuary until the subsequent spring.     

Marine depth use of sea trout Salmo trutta in fjord areas of central Norway

The vertical behaviour of 44 veteran sea trout Salmo trutta (275–580 mm) in different marine fjord habitats (estuary, pelagic, near shore with and without steep cliffs) was documented during May–February by acoustic telemetry. The swimming depth of S. trutta was influenced by habitat, time of day (day v. night), season, seawater temperature and the body length at the time of tagging. Mean swimming depth during May–September was 1·7 m (individual means ranged from 0·4 to 6·4 m). Hence, S. trutta were generally surface oriented, but performed dives down to 24 m. Mean swimming depth in May–September was deeper in the near‐shore habitats with or without steep cliffs (2·0 m and 2·5 m, respectively) than in the pelagic areas (1·2 m). May–September mean swimming depth in all habitats was slightly deeper during day (1·9 m) than at night (1·2 m), confirming that S. trutta conducted small‐scale diel vertical movements. During summer, S. trutta residing in near‐shore habitat progressively moved deeper over the period May (mean 1·1 m) to August (mean 4·0 m) and then reoccupied shallower areas (mean 2·3 m) during September. In winter (November and February), individuals residing in the innermost part of the fjords were found at similar average depths as they occupied during the summer (mean 1·3 m). The swimming depths of S. trutta coincide with the previously known surface orientation of salmon lice Lepeophtheirus salmonis. Combined with previous studies on horizontal use of S. trutta, this study illustrates how S. trutta utilize marine water bodies commonly influenced by anthropogenic factors such as aquaculture, harbours and marine constructions, marine renewable energy production or other human activity. This suggests that the marine behaviour of S. trutta and its susceptibility to coastal anthropogenic factors should be considered in marine planning processes.     

Water temperatures influence the marine area use of Salvelinus alpinus and Salmo trutta

The migratory behaviour and spatial area use of sympatric Arctic charr Salvelinus alpinus and brown trout Salmo trutta were investigated during their marine feeding migration. The likelihood of finding individuals of both species in the inner or outer fjord areas was dependent on water temperature in the inner area (especially for S. alpinus), the temperature difference between the inner and outer areas (especially for S. trutta) and fish fork length (both species). The strongest predictor was the water temperature in the inner area, and particularly S. alpinus left this area and moved to the outer areas with increasing temperatures in the inner area. At 8° C in the inner area, the likelihood of finding S. alpinus in the outer areas was >50%. This predictor had a smaller effect on S. trutta, and the likelihood of finding S. trutta in the outer areas only started to increase at around 14° C. The relationships between temperature and area use did not correspond to the species' optimal growth temperatures, but to their previously documented temperature preferences. Individuals of both species used mainly the littoral fjord areas, and to a lesser extent the pelagic areas. In conclusion, temperature differences between the inner and outer marine areas probably resulted in the segregated area use between the species, because water temperatures or factors influenced by temperature affected their migratory behaviour and habitat use differently. The results indicate that increased marine temperatures with global warming may lead to increased spatial overlap between S. trutta and S. alpinus, which again may lead to increased interspecific competition during their marine phase, and with S. alpinus probably being the more negatively affected.     

Residency time, migration route and survival of Atlantic salmon Salmo salar smolts in a Canadian fjord

Atlantic salmon Salmo salar smolts (n = 181) from two rivers were surgically implanted with acoustic transmitters and released to determine migration route, residency time and survival in a 50 km long estuarine fjord located on the south coast of Newfoundland, Canada. Data obtained from automated receivers placed throughout the Bay d'Espoir fjord indicated that migrating smolts used different routes to reach the outer areas of the fjord. The duration of time that smolts spent in the immediate estuary zone also differed between the two localities (7 and 17 days) although the total time smolts were resident in the fjord was similar and extensive (40 days). Many smolts were resident for periods of 4-8 weeks moving back and forth in the outer part of the fjord where maximum water depths range from 300 to 700 m. Survival in the estuary zone was greater for smolts with prolonged residency in estuarine habitat. Overall smolt survival to the fjord exit was moderately high (54-85%), indicating that the initial phase of migration did not coincide with a period of unusually high mortality.     

Factors influencing return rate and marine residence duration in sea trout populations in Central Norway

Brown trout (Salmo trutta) display extensive plasticity in marine migratory behaviours, with marine migrations considered to be an adaptive strategy which enables sea trout to maximize growth and reproductive potential. However, marine migrations are not without associated costs, including threats posed by ever-increasing salmon lice (Lepeophtheirus salmonis) infestations. In the present study, we used passive integrated transponder technology to characterize variability in sea trout migration behaviour amongst three catchments situated in a region of intensive salmon farming in central Norway. Specifically, we investigate how lice infestation, out-migration date and body size alter sea trout return rate and marine residence duration during the first out-migration to sea from each catchment. Distinct catchment-specific differences in sea trout out-migration size and the number of cohorts were observed, but larger body size did not guarantee the successful return of migrating trout. The marine residence duration of individuals that successfully returned to freshwater was positively correlated with lice infestation risk, suggesting for these individuals the lethal infestation threshold had not been reached. Our results also suggest that sea trout populations from lotic-dominated catchments are potentially at greater risk from size-related threats to their survival encountered during their marine migrations than sea trout from lentic-dominated catchments. The variability in sea trout migratory behaviour amongst catchments observed here emphasizes the challenges fisheries managers face when deciding the best actions to take to protect the anadromous portion of brown trout populations.     

Does reduced feeding prior to release improve the marine migration of hatchery brown trout Salmo trutta smolts?

The aim of this study was to test the hypothesis that hatchery brown trout Salmo trutta smolts, with 50% reduced or no feeding over the last 5 months before release, were more likely to migrate to the sea than individuals with standard feeding ratios. The juvenile fish were divided into three groups 176 days before release: (A) with no feeding, (B) with 50% and (C) with 100% feeding. To study their seaward migration, 40 fish from each feeding group were tagged with acoustic transmitters and tracked by automatic listening stations in the River Nidelva, Trondheim, Norway, its estuary and in the nearest marine environment. At the time of release, mean condition factor was significantly lower in group A and the fish from groups A and B had higher levels of Na+, K+‐ATPase. Significantly more fish from group A migrated to the sea, but the rate of downstream progression from release to the estuary did not differ between the three groups. In conclusion, the S. trutta smolts with no access to food in the last 176 day before release were more likely to migrate to the sea. Fish from all three feeding groups, however, appeared to smoltify and had the same rate of downstream progression to the estuary. This indicates that differences in migratory behaviour between individuals from the three feeding groups begin from the time when the fish reach saline waters. It is suggested that feeding in hatcheries has to be greatly reduced (by 50% or more) over several months to have a pronounced effect on the migratory behaviour in S. trutta.     

Migration behaviour of twaite shad Alosa fallax assessed by otolith Sr:Ca and Ba:Ca profiles

Individual migration behaviour during the juvenile and adult life phase of the anadromous twaite shad Alosa fallax in the Elbe estuary was examined using otolith Sr:Ca and Ba:Ca profiles. Between hatching and the end of the first year of life, juveniles showed two migration patterns. Pattern one exhibited a single downstream migration from fresh water to the sea with no return into fresh water. In contrast, pattern two showed a first migration into the sea, then a return into fresh water and, finally, a second downstream migration into marine water. This first report of migration plasticity for A. fallax points to different exposure times to estuarine threats depending on the migration strategy. In adults, high Sr:Ca and low Ba:Ca in the majority of individuals confirmed prior reports of a primarily marine habitat use. Patterns reflecting spawning migrations were rarely observed on otoliths, possibly due to the short duration of visits to fresh water.     

Estuarine survival and migratory behaviour of Atlantic salmon Salmo salar smolts

To estimate mortality rates, assess the spatio‐temporal dynamics of natural mortality and examine migratory behaviour during the fresh to saltwater transition, 185 wild Atlantic salmon Salmo salar smolts were implanted with coded acoustic transmitters. Seaward migration of tagged S. salar from four river systems in an area of Nova Scotia, Canada known as the Southern Upland was monitored using fixed receivers and active telemetry over 3 years. Cumulative survival through the river, inner estuary, outer estuary and bay habitats averaged 59·6% (range = 39·4–73·5%). When standardized to distance travelled, survival rates followed two patterns: (1) constant rates of survival independent of habitat or (2) low survival most frequently associated with inner estuary habitats. In rivers where survival was independent of habitat, residency periods were also independent of habitat, post‐smolts exhibited few upstream movements, took a more direct route to the ocean and reached the ocean rapidly. Alternatively, in rivers where survival was habitat specific, residency was also habitat specific with overall increased residency, more frequent upstream movements and delayed arrival to the open ocean. The sudden disappearance of most (75–100%) smolts and post‐smolts assumed dead during the course of this study warrants further examination into the role of avian predators as a mortality vector.     

Trophically transmitted parasites in wild Atlantic salmon post‐smolts from Norwegian fjords

The community structure of trophically transmitted intestinal helminths of Atlantic salmon Salmo salar post‐smolts was highly variable among four fjords in Norway. There were no severely pathogenic parasite species. Post‐smolts from the southernmost Trondheimsfjord had a higher diversity of freshwater parasite species compared to the three northern fjords (Tanafjord, Altafjord and Malangen). In contrast, the highest diversity and proportion of marine species was found in the three northern fjords. Post‐smolts were generally more infected with marine parasites in the outer rather than inner parts of all of the fjords. The prevalence of the acanthocephalan Echinorynchus gadi (range: 13–42%) and marine trematodes (range: 14–47%) was higher in post‐smolts in outer zones of the northern fjords than in fish from Trondheimsfjord (0 and 6%, respectively). The within‐fjord variability and north‐south geographical gradient in parasite infection patterns reflected differences in marine feeding of the post‐smolts on potential intermediate hosts such as amphipods ( E. gadi ) and fish larvae (trematodes), which were higher in the northern fjords (range: 27–28 and 67–85%, respectively) than in Trondheimsfjord (5 and 19%, respectively). High intensities of marine parasites suggest that some post‐smolts from northern fjords may have a prolonged fjord‐feeding compared to those from Trondheimsfjord. Parasites of both freshwater and marine origin appear to be suitable as bio‐indicators of feeding and migratory pattern of Atlantic salmon post‐smolts and preadults during their seaward migration.     

Riverine and near coastal migration performance of hatchery brown trout Salmo trutta

To study migration performance and return rates of hatchery brown trout Salmo trutta smolts the first 5 months after release, 50 fish in each year (fork length, L_F, 158–288 mm) were in two subsequent years tagged with acoustic transmitters and recorded by automatic listening stations in the River Nidelva (central Norway), its estuary and in the marine environment. More than half of the smolts became anadromous migrants (52% in 2011 and 70% in 2012). The fish spent longer time in the estuary than in the marine environment and the results suggest that migratory behaviour of S. trutta smolts is not only restricted to be resident or anadrome–lacustrine, but that there is also an intermediary strategy of estuarine feeding. There were no differences in L_F or mass between groups of smolts with different migration patterns. Return rates from the sea within the first 5 months after release were in both years 16%. Median progression rate in the river was 0·090 L_F s^?1 but decreased significantly as the smolts entered the estuary (0·015 L_F s^?1). The long residential time in the estuary may increase the risk of negative effects of anthropogenic activities in estuaries, such as harbours and industrial development, and special attention should be given to evaluate effects of such activities.     

Net ground speed of downstream migrating radio-tagged Atlantic salmon (Salmo salar L.) and brown trout (Salmo trutta L.) smolts in relation to environmental factors

The downstream migration of Atlantic salmon (Salmo salarL.) and sea trout smolt (S. trutta L.) was investigated using radio telemetry in the spring of 1999 and 2000. Forty wild sea trout smolts, 20 F1 sea trout smolts, 20 hatchery salmon smolts and 20 salmon smolts from river stockings were radio tagged and released in the Danish River Lilleaa. The downstream migration of the different groups of fish was monitored by manual tracking and by three automatic listening stations. The downstream migration of radio tagged smolts of both species occurred concurrently with their untagged counterparts. The diel migration pattern of the radio tagged smolts was predominantly nocturnal in both species. Wild sea trout smolt migrated significantly faster than both the F1 trout and the introduced salmon. There was no correlation between net ground speed, gill Na⁺,K⁺-ATPase activity or fish length in any of the different groups. The migration speed of wild sea trout smolts was positively correlated with water discharge in both years. In F1 sea trout smolts, migration speed was positively correlated with temperature in 1999. The migration speed of salmon smolts did not correlate to any of the investigated parameters.     

The post‐spawning movements of migratory brown trout Salmo trutta L.

The post spawning behaviour of sea trout Salmo trutta was studied over a 2 year period in the river and estuary of the River Fowey, south‐west England. Forty‐five sea trout kelts were trapped immediately after spawning in December and intraperitoneally tagged with miniature acoustic transmitters. The subsequent emigration into coastal waters was monitored using acoustic receivers deployed throughout the river catchment. The levels of gill Na⁺K⁺ATPase activity in sea trout kelts sampled at the same time as the tagged fish were within the range of 2·5 to 4·5 μmol Pi per mg protein per h indicating that the post‐spawning fish were not physiologically adapted to salt water. The tagged kelts were resident in fresh water between 4 and 70 days before entering the estuary. Sixty two per cent of the tagged kelts subsequently migrated successfully into coastal waters, with a higher success rate for male fish (75%) than females (58%). There was a significant size related difference in the run‐timing of the kelts with the larger fish moving more quickly into coastal waters after spawning than smaller fish. Seaward migration within fresh water was predominantly nocturnal and generally occurred in conjunction with increasing river discharge and rising water temperature. Migration through the estuary continued to be predominantly nocturnal and occurred during an ebbing tide. Residency within the estuary varied amongst individuals although it was invariably short, with most fish moving out into coastal waters within one to two tidal cycles. Five tagged kelts returned from the coastal zone and re‐entered fresh water during April and June. Marine residence time varied between 89 and 145 days (mean 118 days) and the minimum estimated marine survival was c. 18%. One of these sea trout was subsequently recaptured after successfully spawning in the vicinity where it had been previously tagged demonstrating a degree of spawning site fidelity.     

Passing a seawater challenge test is not indicative of hatchery‐reared Atlantic salmon Salmo salar smolts performing as well at sea as their naturally produced conspecifics

Despite satisfactory reactions to seawater challenge tests indicative of appropriate physiological state, hatchery‐reared Atlantic salmon Salmo salar smolts stocked in the Eira River in Norway between 2001 and 2011 performed less well at sea in terms of growth, age at maturity and survival than smolts of natural origin. The mean rates of return to the river for hatchery‐reared and naturally produced S. salar were 0·98 and 2·35%. In the Eira River, c. 50 000 hatchery‐reared S. salar smolts of local origin were stocked annually to compensate for reduced natural smolt production following regulation for hydroelectric purposes, while a mean of 17 262 smolts were produced naturally in the river. This study demonstrates that, although captive S. salar perform well in seawater challenge tests, hatchery‐reared smolts are not necessarily as adaptable to marine life as their naturally produced counterparts. These findings suggest that production of hatchery‐reared smolts more similar to naturally produced individuals in morphology, physiology and behaviour will be necessary to improve success of hatchery releases. Where possible, supplementary or alternative measures, including habitat restoration, could be implemented to ensure the long‐term viability of wild stocks.     

Environmental cues and extended estuarine residence in seaward migrating eels (Anguilla australis)

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Migration of farmed adult Atlantic salmon with and without olfactory sense, released on the Norwegian coast

The dispersal and migration of farmed Atlantic salmon, Salmo salar , allowed to escape during the summer was studied. Three groups of 4–year–old fish of the River Imsa stock were released in coastal waters off south-western Norway: one group, with functional olfactory organs, was released at a fish farm 4 km away from the R. Imsa; two other groups, one with transected olfactory nerves and the other with functional olfactory organs, were released in the sea 90 km from the R. Imsa. To compare them with the migration pattern of reared, large smolts of the Imsa stock, a group of 3 + smolts was released in the R. Imsa.
Adults of salmon released as 3–year–old smolts homed with high precision to the R. Imsa. Four– year–olds released in the sea were recaptured in the fjord and in the coastal current, the majority north of the places of release. Immatures migrated to feeding areas in the North Atlantic. Matures seemed to enter rivers at random when ready to spawn. There was no difference in migration pattern between anosmics and controls. The olfactory sense was not mandatory for entering fresh water. The results indicate that the homing behaviour of Atlantic salmon is not a direct consequence of a single imprinting of the smolts, and that there is not a direct genetic link for return to a particular river. The present results support the sequential imprinting hypothesis proposed by Harden Jones (1968).     

Keeping close to the river,shore and surface: the first marine migration of brown trout (Salmo trutta) and Arctic charr (Salvelinus alpinus) post-smolts

Acoustic telemetry was utilized to track 49 brown trout (Salmo trutta) and 37 Arctic charr (Salvelinus alpinus) first-time migrants of wild origin [post-smolts; mean L_F (fork length): 169 and 172 mm] in a large fjord in northern Norway. The S. trutta were registered at sea for more than twice the time of the S. alpinus (medians of 54 and 22 days, respectively). Both species were mostly detected near river mouths (>80% of detections) and almost exclusively spent their time (>95%) within the interior 18 km of the fjord. They were surface oriented, with most detections at <1 m depth and S. trutta deeper on average (median mean depths of 0.7 and 0.5 m, respectively). This study concludes that post-smolts of both species stay closer to the surface and to river mouths than larger veteran migrants. This study emphasizes the importance of river mouths and upper water layers for the survival of both species during their first marine migration.