Audun H. Rikardsen

A critical life stage of the Atlantic salmon Salmo salar: behaviour and survival during the smolt and initial post‐smolt migration

The anadromous life cycle of Atlantic salmon Salmo salar involves long migrations to novel environments and challenging physiological transformations when moving between salt-free and salt-rich waters. In this article, (1) environmental factors affecting the migration behaviour and survival of smolts and post-smolts during the river, estuarine and early marine phases, (2) how behavioural patterns are linked to survival and (3) how anthropogenic factors affect migration and survival are synthesized and reviewed based on published literature. The timing of the smolt migration is important in determining marine survival. The timing varies among rivers, most likely as a consequence of local adaptations, to ensure sea entry during optimal periods. Smolts and post-smolts swim actively and fast during migration, but in areas with strong currents, their own movements may be overridden by current-induced transport. Progression rates during the early marine migration vary between 0.4 and 3.0 body lengths s(-1) relative to the ground. Reported mortality is 0.3-7.0% (median 2.3) km(-1) during downriver migration, 0.6-36% (median 6.0) km(-1) in estuaries and 0.3-3.4% (median 1.4) km(-1) in coastal areas. Estuaries and river mouths are the sites of the highest mortalities, with predation being a common cause. The mortality rates varied more among studies in estuaries than in rivers and marine areas, which probably reflects the huge variation among estuaries in their characteristics. Behaviour and survival during migration may also be affected by pollution, fish farming, sea lice Lepeophtheirus salmonis, hydropower development and other anthropogenic activities that may be directly lethal, delay migration or have indirect effects by inhibiting migration. Total mortality reported during early marine migration (up to 5-230 km from the river mouths) in the studies available to date varies between 8 and 71%. Hence, the early marine migration is a life stage with high mortalities, due to both natural and human influences. Factors affecting mortality during the smolt and post-smolt stages contribute to determine the abundance of spawner returns. With many S. salar populations in decline, increased mortality at these stages may considerably contribute to limit S. salar production, and the consequences of human-induced mortality at this stage may be severe. Development of management actions to increase survival and fitness at the smolt and post-smolt stages is crucial to re-establish or conserve wild populations.

Genetic versus Rearing-Environment Effects on Phenotype: Hatchery and Natural Rearing Effects on Hatchery- and Wild-Born Coho Salmon

With the current trends in climate and fisheries, well-designed mitigative strategies for conserving fish stocks may become increasingly necessary. The poor post-release survival of hatchery-reared Pacific salmon indicates that salmon enhancement programs require assessment. The objective of this study was to determine the relative roles that genotype and rearing environment play in the phenotypic expression of young salmon, including their survival, growth, physiology, swimming endurance, predator avoidance and migratory behaviour. Wild- and hatchery-born coho salmon adults (Oncorhynchus kisutch) returning to the Chehalis River in British Columbia, Canada, were crossed to create pure hatchery, pure wild, and hybrid offspring. A proportion of the progeny from each cross was reared in a traditional hatchery environment, whereas the remaining fry were reared naturally in a contained side channel. The resulting phenotypic differences between replicates, between rearing environments, and between cross types were compared. While there were few phenotypic differences noted between genetic groups reared in the same habitat, rearing environment played a significant role in smolt size, survival, swimming endurance, predator avoidance and migratory behaviour. The lack of any observed genetic differences between wild- and hatchery-born salmon may be due to the long-term mixing of these genotypes from hatchery introgression into wild populations, or conversely, due to strong selection in nature—capable of maintaining highly fit genotypes whether or not fish have experienced part of their life history under cultured conditions.

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.

Effects of Floy and Soft VIalpha Tags on Growth and Survival of Juvenile Arctic Char

Abstract The Floy FTF-69 fingerling tag and the new soft VIalpha (VI) tag can both be used on small fish (>100 mm and >150 mm, respectively) according to their manufacturers, but information on their use and effect on fish has not been available. In this study, these two tags were compared for one year on wild Arctic char Salvelinus alpinus, parr (lake dwelling), and smolts (100–229 mm) from two small lakes in northern Norway. Comparisons of growth rate, survival, and tag retention were analyzed. In general, VI-tagged smolts grew significantly better than Floy-tagged smolts in the sea. However, the difference was largest for the smallest fish (100–189 mm); for the larger fish (190–229 mm), there was less or no difference. Survival rates in the sea were also significantly higher (11–15%) for the smallest (<170 mm) VI-tagged smolts than for Floy-tagged smolts of the same size; larger VI-tagged smolts (≥170 mm) only survived at a slightly higher (1.5–4.0%) and nonsignificant rate than corresponding Floy-tagg...

Archival tags reveal that Arctic charr Salvelinus alpinus and brown trout Salmo trutta can use estuarine and marine waters during winter

By tagging north-Norwegian anadromous riverine Arctic charr Salvelinus alpinus and brown trout Salmo trutta with simple archival tags that measured ambient water temperature and relating the recordings to the temperature in the surrounding environments, it was demonstrated that 91% of the S. trutta and 80% of the S. alpinus utilized the estuarine and marine environment during the winter. There was large individual variation in migratory behaviour. Salvelinus alpinus on average entered the estuary on 12 January and the marine environment on 26 February, had continuous marine migrations lasting up to 55 days, and spent on average 40 days in the estuary and 25 days in the sea during the winter. The corresponding numbers for S. trutta were 15 December and 4 January for first entry in the estuary and sea, maximum 39 days in the marine environment and average number of days in the estuary and sea were 34 and 50. Most individuals of both species frequently shifted between the three habitats. These findings thus contradict previous studies conducted on lake-dwelling populations, which suggested that northern populations of both species solely overwinter in fresh water. The use of inexpensive temperature recording archival tags gave detailed continuous information on seasonal migrations between habitats with different thermal characteristics.

Survival, Migration Speed and Swimming Depth of Atlantic Salmon Kelts During Sea Entry and Fjord Migration

In contrast to most species of Pacific salmon (Oncorhynchus spp.), Atlantic salmon (Salmo salar L.) is an iteroparous species such that it may survive and return to spawn repeatedly. Little information exists on these survivors (kelts) even though they might contribute significantly to salmon production when returning as repeat spawners. In order to estimate survival, timing of migration, swimming progression and swimming depth of Atlantic salmon kelts during sea entry and fjord migration, 60 individuals were captured, tagged with acoustic transmitters (of which 20 with depth sensors) and released in the River Alta, Northern Norway. In addition, 172 kelts were also tagged with external Carlin tags to obtain reliable recapture rates in the fisheries. Nearly all (95%) kelts tagged with acoustic transmitters were recorded during their outward migration at four transects of acoustic receivers deployed across the river mouth and Alta Fjord. Most of the kelts migrated through the 30 km long fjord in only 1–2 days (mean time 33 h, range 7–138 h) and generally stayed close to the surface during the fjord migration (individual mean depth of 2 m, range of individual means 0–15 m). Ninety-two percent (55/60) were recorded at the outermost transect 30 km from the river mouth, indicating a high minimum survival rate. The high survival rate and fast progression of kelts throughout the fjord indicate that sea-entry and early sea migration is not a critical phase for Atlantic salmon kelts, despite their weakened condition after spawning and overwintering in the river. Even though kelt migration overlapped partly with the fishing season both in the river and the fjord, and the kelts migrated in shallow waters exposed to several types of fishing gear, reported recapture rates were small (3%, 6 of all 232 tagged kelts).

Early marine migration of European silver eel (Anguilla anguilla) in Northern Norway

To study the migratory behaviour in wild northern European silver eel Anguilla anguilla during sea entry and early marine migration, 32 individuals were tagged with acoustic transmitters and registered at four automatic listening station arrays from the mouth of the north Norwegian River Alta and throughout the Alta Fjord. The A. anguilla entered the fjord during all parts of the tidal cycle and did not seem to utilize the outgoing tidal currents. They migrated mainly during the night, in both the river mouth and the fjord. On average, they spent 2·7 days travelling from the river mouth to the outermost array, 31 km from the river mouth, corresponding to an average migratory speed of 0·5 km h(-1) . The A. anguilla generally migrated in the central part of the fjord and in the uppermost 10-25% of the water column, but with frequent dives to greater depths. Already 4 km after sea entry, A. anguilla were observed diving deeper than 130 m within 20-30 min periods. Hence, this study demonstrated that A. anguilla may perform an active diving behaviour during the early marine migration. The study took place in a pristine area with a minimum of anthropogenic interventions and by individuals from a population still uninfected by the introduced parasite Anguillicoloides crassus. The results may therefore be used as a baseline for future studies of the A. anguilla early marine migration.

A comparison of Floy and soft VIalpha tags on hatchery Arctic charr, with emphasis on tag retention, growth and survival

The Roy PIP-69 ‘fingerling’ tag and the soft VIalpha tag are designed to be used on small salmonid fishes (> 100 and > 150 mm, respectively). The two tags were compared for 160 days on hatchery-reared Arctic charr, Salvelinus alpinus, (170–209 mm). Tag retention and effects on growth rate and survival were analysed. VIalpha-tagged and untagged control fish had similar growth rates and grew significantly faster than Roy-tagged fish during the experiment. Tag retention was significantly higher for Floy tags (94%) than for Vlalpha tags (78%). Most VIalpha tag loss (11%) took place within the first 10 days of tagging while there were no Floy tags lost during this period. Survival rates were about equal for all three groups (96–98%), and thus the tags did not seem to affect the survival of charr under hatchery conditions.

Prey availability and juvenile Atlantic salmon feeding during winter in a regulated subarctic river subject to loss of ice cover

Increased mortality of juvenile Atlantic salmon (Salmo salar L.), related to lowered levels of stored energy following the loss of ice cover during winter, has been observed after hydropower development in the subarctic River Alta, northern Norway. Drift samples were compared to examine if drift densities, and thus drift prey availabilities for juvenile salmon, were lower in the ice-free than the ice-covered area. In addition, juvenile salmon stomach contents were compared to benthos and drift in the ice-free area to examine salmon winter feeding habitat. Zooplankton, originating from the reservoir, dominated drift at the ice-free site but had lower densities at the downstream ice-covered site. Excluding zooplankton, Chironomidae comprised most of the remaining drift at both the ice-free and ice-covered site, followed by Ephemeroptera, Plecoptera and Simuliidae. No Trichoptera were found in the drift samples. There was no consistent diel periodicity in drift. Benthos was dominated by Chironomidae, followed by Ephemeroptera, Plecoptera and Trichoptera. Other invertebrates occurred in low numbers. Juvenile salmon demonstrated size-selective feeding and fed mainly on Ephemeroptera, followed by Trichoptera and Plecoptera. No zooplankton and few Chironomidae were found in the stomach samples. Stomach content was more similar to benthos than to drift, indicating a larger extent of benthic than drift feeding. No evidence was found for the hypothesis that lack of ice cover reduced the invertebrate drift or caused diel periodicity in the drift. Differences in drift between areas with and without ice could not account for the observed differences in mortality of juvenile salmon during the winter in these areas.

Pop-up satellite archival tag effects on the diving behaviour, growth and survival of adult Atlantic salmon Salmo salar at sea

The effects of large, externally attached pop-up satellite archival tags (PSATs) were compared with those of small implanted data storage tags (DSTs) on adult Atlantic salmon Salmo salar during their ocean migration in regards to depth utilization, diving depth, diving rate, diving speed and temperatures experienced. Additionally the return rate and growth of individuals tagged with PSATs was compared with those of small acoustic tags and DSTs. Overall, the depth distribution of individuals tagged with PSATs was similar to that of those tagged with DSTs, reflecting the pelagic nature of S. salar at sea. Individuals tagged with PSATs, however, dived less frequently and to shallower depths, and dived and surfaced at slower velocities. Sea surface temperatures experienced by individuals tagged with PSATs were similar to those experienced by those tagged with DSTs for the same time of year, suggesting that there were no large differences in the ocean migration. Return rates did not depend on whether individuals were tagged with PSATs or not, indicating that survival at sea was not impacted by PSATs in comparison to small internal tags. Individuals tagged with PSATs, however, had a smaller increase in body mass than those tagged with acoustic tags or DSTs. It was concluded that PSATs are suitable for use in researching large-scale migratory behaviour of adult S. salar at sea, but that some effects on their behaviour from tagging must be expected. Effects of PSATs may be largest in the short term when S. salar are swimming in bursts at high speeds. Even though individuals tagged with PSATs performed deep and frequent dives, the results of this study suggest that untagged individuals would perform even deeper and more frequent dives than tagged individuals.