Smolt Year Research Articles

Time series covering up to four decades reveals major changes and drivers of marine growth and proportion of repeat spawners in an Atlantic salmon population.

Wild Atlantic salmon populations have declined in many regions and are affected by diverse natural and anthropogenic factors. To facilitate management guidelines, precise knowledge of mechanisms driving population changes in demographics and life history traits is needed.Our analyses were conducted on (a) age and growth data from scales of salmon caught by angling in the river Etneelva, Norway, covering smolt year classes from 1980 to 2018, (b) extensive sampling of the whole spawning run in the fish trap from 2013 onwards, and (c) time series of sea surface temperature, zooplankton biomass, and salmon lice infestation intensity.Marine growth during the first year at sea displayed a distinct stepwise decline across the four decades. Simultaneously, the population shifted from predominantly 1SW to 2SW salmon, and the proportion of repeat spawners increased from 3 to 7%. The latter observation is most evident in females and likely due to decreased marine exploitation. Female repeat spawners tended to be less catchable than males by anglers.Depending on the time period analyzed, marine growth rate during the first year at sea was both positively and negatively associated with sea surface temperature. Zooplankton biomass was positively associated with growth, while salmon lice infestation intensity was negatively associated with growth.Collectively, these results are likely to be linked with both changes in oceanic conditions and harvest regimes. Our conflicting results regarding the influence of sea surface temperature on marine growth are likely to be caused by long‐term increases in temperature, which may have triggered (or coincided with) ecosystem shifts creating generally poorer growth conditions over time, but within shorter datasets warmer years gave generally higher growth. We encourage management authorities to expand the use of permanently monitored reference rivers with complete trapping facilities, like the river Etneelva, generating valuable long‐term data for future analyses.

Growth variation of Atlantic salmon Salmo salar at sea affects their population-specific reproductive potential

Understanding the processes shaping the dynamics of anadromous fish populations is essential for their management and conservation. Yet, little is known about how variation in performance at sea affects their population dynamics. Here we show that variation in body growth at sea contributes to explaining variation in the reproductive potential for 2 Atlantic salmon Salmo salar populations, but to a varying extent. To this end, we assembled data collected during 50 yr for 2 Baltic salmon populations of hatchery origin, including annually released smolts, survival at sea estimates, size-specific growth at sea, annual length distributions of returning adult females and their reproductive potential. The regression models fitted to explain the reproductive potential of our 2 study populations improved when growth at sea was included as an explanatory variable, in addition to smolt year class abundance and estimates of their survival at sea. This link between body growth at sea and population-level reproductive potential suggests that growth at sea can be important to consider when resolving variation in recovery and dynamics among salmon populations sharing the same sea.

Increasing temperature associated with increasing grilse proportion and smaller grilse size of Atlantic salmon

Effects of temperature on Atlantic salmon (Salmo salar) were analysed using Carlin tag recovery data (1985–2014), and mixed‐stock catch data (smolt years from 2001 to 2012) in northern parts of the Baltic Sea. During warmer summers, the mean smolt length of the recaptured salmon tended to be smaller, and salmon were recaptured more frequently in feeding grounds closer to the home rivers in the Gulf of Bothnia, while colder summers were associated with more recaptures further south, in the Baltic Main Basin. Moreover, a warmer spring in the smolt year was associated with decreased weight of male grilses in mixed stock data. Further, warmer spring temperatures during the smolt year were associated with a higher proportion of one‐sea‐winter (1SW) males during the return migration in mixed stock data. These results suggest that the increasing global temperature may affect Atlantic salmon life history demographics.

Context-dependent impact of an ectoparasite on early marine growth in Atlantic salmon

The impact of sea lice on post-smolt growth (PSG) was studied by comparing growth patterns in the scales of recaptured adult salmon originating from hatchery-reared smolts that were untreated or were previously treated for sea lice infestation during the years 2003–2014. Growth in the returning hatchery-reared fish was also compared to the growth of wild fish from the same smolt year class. PSG was greater in treated versus untreated fish in the early part of the time series, which was a period of low marine survival. However, the differentiation in PSG disappeared in the later portion of the time series, which coincided with a shift from low to higher marine survival. This transition was demarcated by the year 2009. During this later period, PSG correlated with the marine survival of salmon, whereas in the early period, this correlation was not evident. Wild fish had faster PSG than untreated hatchery fish throughout the study period, but had slower PSG than treated hatchery fish during the early part of the time series. The effects of sea lice alone cannot explain the observed shift in marine survival occurring in 2009. However, it seems evident that the effect of antiparasitic treatment was contingent on poor marine survival, showing that the impacts of parasites on the PSG of Atlantic salmon are context dependent.

The vgll3 Locus Controls Age at Maturity in Wild and Domesticated Atlantic Salmon (Salmo salar L.) Males.

Wild and domesticated Atlantic salmon males display large variation for sea age at sexual maturation, which varies between 1–5 years. Previous studies have uncovered a genetic predisposition for variation of age at maturity with moderate heritability, thus suggesting a polygenic or complex nature of this trait. The aim of this study was to identify associated genetic loci, genes and ultimately specific sequence variants conferring sea age at maturity in salmon. We performed a genome wide association study (GWAS) using a pool sequencing approach (20 individuals per river and phenotype) of male salmon returning to rivers as sexually mature either after one sea winter (2009) or three sea winters (2011) in six rivers in Norway. The study revealed one major selective sweep, which covered 76 significant SNPs in which 74 were found in a 370 kb region of chromosome 25. Genotyping other smolt year classes of wild and domesticated salmon confirmed this finding. Genotyping domesticated fish narrowed the haplotype region to four SNPs covering 2386 bp, containing the vgll3 gene, including two missense mutations explaining 33–36% phenotypic variation. A single locus was found to have a highly significant role in governing sea age at maturation in this species. The SNPs identified may be both used as markers to guide breeding for late maturity in salmon aquaculture and in monitoring programs of wild salmon. Interestingly, a SNP in proximity of the VGLL3 gene in humans (Homo sapiens), has previously been linked to age at puberty suggesting a conserved mechanism for timing of puberty in vertebrates.

Between-watershed movements of two anadromous salmonids in the Arctic

A long-term study in the River Halselva in the Arctic region of Norway demonstrated that movements between watersheds were considerably higher in anadromous brown trout (Salmo trutta) than anadromous Arctic char (Salvelinus alpinus). Fourteen smolt year classes of both species were captured in a fish trap, individually tagged, and thereafter registered each time they passed the trap during their migration between the sea and fresh water every summer. Annual mean survival (i.e., recovery rate) after the first entry to sea as smolts was estimated as 31.4% in Arctic char and 26.6% in brown trout. Most surviving Arctic char returned to the River Halselva to overwinter after the same summer that they migrated to sea as smolts. However, several brown trout overwintered one to four times in other watersheds, mainly the considerably larger River Altaelva, before most eventually returned to the River Halselva upon maturation. The substantial difference in movement rate between watersheds between Arctic char (2.2%) and brown trout (39.6%) is expected to be a consequence of local geographic conditions combined with different habitat preferences of the two species.

Postsmolt Growth and Thermal Regime Define the Marine Survival of Steelhead from the Keogh River, British Columbia

AbstractThe population of anadromous steelhead Oncorhynchus mykiss in the Keogh River has been studied intensively, in part because of its pattern of declining recruitment, which is largely attributed to poor marine survival. Climate variability has changed the productivity of salmonid species in all regions of the North Pacific, with areas alternately shifting between periods of enhanced and depressed productivity. The mechanisms governing marine survival and adult recruitment are central to contemporary resource management concerns but are also of concern with regard to the long‐term prospects of managing biodiversity. We provide evidence that postsmolt growth contributes to the pattern of marine survival of Keogh River steelhead over the period corresponding to smolt years 1977–1999. Size at ocean entry did not appear to have sufficient contrast to significantly affect survival. However, assessment of scale growth suggested that the fish's initial growth at sea is not as important as the sustained growth conditions during summer and fall of the postsmolt year. The return rate of steelhead was negatively correlated with sea surface temperature in the ocean domains that were assumed to provide postsmolt nursery habitat, suggesting that growth is directly affected by warming conditions or that ocean warming affects the food web upon which steelhead depend. Steelhead appear to respond to changing climate and growth regimes in a manner similar to that of their North Atlantic analog, the Atlantic Salmon Salmo salar. Comparative data show that eastern basin Atlantic Salmon populations are negatively affected by a thermal regime of increasing temperature during the postsmolt year, suggesting a relationship between postsmolt growth and survival.Received August 14, 2013; accepted October 17, 2013

Contemporary ocean warming and freshwater conditions are related to later sea age at maturity in Atlantic salmon spawning in Norwegian rivers

Atlantic salmon populations are reported to be declining throughout its range, raising major management concerns. Variation in adult fish abundance may be due to variation in survival, growth, and timing of life history decisions. Given the complex life history, utilizing highly divergent habitats, the reasons for declines may be multiple and difficult to disentangle. Using recreational angling data of two sea age groups, one-sea-winter (1SW) and two-sea-winter (2SW) fish originated from the same smolt year class, we show that sea age at maturity of the returns has increased in 59 Norwegian rivers over the cohorts 1991–2005. By means of linear mixed-effects models we found that the proportion of 1SW fish spawning in Norway has decreased concomitant with the increasing sea surface temperature experienced by the fish in autumn during their first year at sea. Furthermore, the decrease in the proportion of 1SW fish was influenced by freshwater conditions as measured by water discharge during summer months 1 year ahead of seaward migration. These results suggest that part of the variability in age at maturity can be explained by the large-scale changes occurring in the north-eastern Atlantic pelagic food web affecting postsmolt growth, and by differences in river conditions influencing presmolt growth rate and later upstream migration.

Evidence of changing migratory patterns of wild Atlantic salmon Salmo salar smolts in the River Bush, Northern Ireland, and possible associations with climate change

The migration patterns, timing and biological characteristics of wild Atlantic salmon Salmo salar smolts in the River Bush, Northern Ireland, were examined over the period 1978-2008. A distinct change in the timing of the smolt run was detected with progressively earlier emigration periods evident across the time series. The shift in run timing ranged from 3.6 to 4.8 days 10 years(-1) for a range of standard migratory audit points. The timing of smolt emigration has been linked to ambient river temperature patterns. Distinct seasonal patterns were evident for biological characteristics of River Bush smolts with mean age and fork length decreasing throughout the emigration period. Marine survival patterns in 1 sea winter River Bush S. salar were strongly influenced by the run timing of the preceding smolt year such that later emigrating cohorts demonstrated increased survival. Possible mechanisms for this relationship based on local climatic variation have been explored, including the effect of potential thermal mismatch between freshwater and marine environments.

Migratory timing, marine survival and growth of anadromous brown troutSalmo truttain the River Imsa, Norway

The aim of the paper was to study sea migration, growth and survival of brown trout Salmo trutta of the River Imsa, 1976-2005. The migratory S. trutta were individually tagged and fish leaving or entering the river were monitored daily in traps located near the river mouth. The mean annual duration of the sea sojourn was 6-9 months for first-time migrants moving to sea between January and June. It was 8-18 months for those migrating to sea between July and December. Veteran migrants stayed 12 months or less at sea and most returned to the river in August. Early ascending fish stayed the longest in fresh water because most returned to sea in April to May. The day number of 50% cumulative smolt descent correlated negatively with mean water temperature in February to March and the February North Atlantic Oscillation index (NAOI). Mean annual sea growth during the first 2 years after smolting was higher for S. trutta spending the winter at sea than those wintering in the River Imsa. First year's sea growth was lower for S. trutta descending in spring than autumn. For first-time migrants, it correlated negatively with the February NAOI of the smolt year. Sea survival was higher for spring than autumn descending first-time migratory S. trutta with a maximum in May (14.9%). Number of anadromous S. trutta returning to the river increased linearly with the size of the cohort moving to sea, with no evidence of density-dependent sea mortality. Sea survival of S. trutta smolts moving to sea between January and June correlated positively both with the annual number of Atlantic Salmo salar smolts, the specific growth rate at sea, and time of seaward migration in spring. This is the first study indicating how environmental factors at the time of seaward migration influence the sea survival of S. trutta.

Marine survival of reared Atlantic salmon in the Baltic Sea: The effect of smolt traits and annual factors

We examined the factors underlying the deteriorating marine survival of a hatchery stock of Atlantic salmon, Salmo salar. The stock originated from the salmon population of the Neva River, which flows into the Gulf of Finland in the Baltic Sea. During 1981–2005, about 134,000 Carlin-tagged Neva salmon smolts were released in the Gulf of Finland and the Bothnian Sea. We investigated the effects of fin damage, smolt length, sexual maturity (precocious maturation), smolt year, the time (days) between tagging and release and the release date on survival. The survival of the released smolts clearly decreased towards present. In earlier years, larger smolts tended to survive better than smaller ones, but the advantage of a large smolt size decreased in the later years of the study. No significant effect of dorsal fin damage on survival was detected. On average, immature smolts survived better than precocious males, and the proportion of precocious males increased over time. However, the proportion of precocious males among the tagged fish did not explain the decreasing trend in marine survival. Release year was the most important predictor of the long-term decreasing trend in marine survival. The optimal tagging time varies: in some years smolts tagged in the autumn survived better, while in other years spring taggings were more advantageous. For most years, a late release (in May or June) was more favourable than an early one (in April). As the releases were mainly carried out quite early in the spring in the last 13 years of the study, an unfavourable timing of release may partly explain the lower marine survival recorded during this period.

Twenty years of hydropower regulation in the River Alta: long-term changes in abundance of juvenile and adult Atlantic salmon

The River Alta, northern Norway (70°N), was regulated for hydropower in 1987. Densities of juveniles and catches of adult Atlantic salmon have been studied since 1980–1981 to examine the effects of regulation. The need to control environmental variables during electrofishing was emphasized, as flow variables explained up to 42% of the variation in estimated juvenile densities. The number of spawning redds was counted along the river from 1996 to 2005. The annual number of spawning redds was correlated with the catch of multi-sea-winter salmon (predominantly females). In the upper 7 km section, just downstream of the power station outlet, juvenile densities were reduced by 80% from pre-regulation levels to minimum levels in 1992–1996. This was followed by partial recovery during 1997–2005, although not entirely back to pre-regulation levels. In contrast, the general trend in the middle part of the river was a linear increase in juvenile densities during 1981–2005. Decreased juvenile densities in the upper section was subsequently followed by reduced catches of adult salmon in this part of the river. The relative catches of smolt year classes migrating to sea in the upper section was reduced by up to 75% from 1991 onwards. Spawning and recruitment in the upper section have increased in recent years, probably back to the introduction of catch-and-release angling and an increase in salmon runs. However, present day smolt production in the upper section is still reduced compared to the middle part of the river, 18 years after regulation. The decreased densities of juvenile salmon in the upper section were probably caused by several factors, of which stranding mortality due to sudden drops in the water level and increased winter mortality due to changed environmental conditions, especially reduced ice-cover, may be the most important. In conclusion, the regulation caused a considerable reduction of the salmon production in the upper 16% of salmon reaches, but did not affect the salmon population negatively further downstream. This study illustrates that apparently small environmental disturbances can cause large changes in Atlantic salmon abundance in high latitude populations.

Population structuring in Atlantic salmon (Salmo salar): evidence of genetic influence on the timing of smolt migration in sub‐catchment stocks

Abstract – We compared timing of smolt migration for two populations of naturally spawned Atlantic salmon, Salmo salar, originating from an upper catchment (River Tummel) and a lower catchment (River Almond) tributary of the River Tay, Scotland. Smolts from the upper tributary began migration earlier than those from the lower tributary. On both occasions when fry derived from River Isla (lower tributary) stock were transferred to a location in the upper catchment, smolt migration was later than for native fish. Similarly, when fry from Tummel and Isla stocks were stocked in a common, upper catchment location, Isla origin fish migrated at a later date, in both of the two following smolt years. These differences are indicative of a genetic basis for the timing of smolt migration and suggestive of local adaptation. Mounting evidence points towards local genetic adaptation for the timing of expression of behaviours associated with migration. These aspects of variation should be accommodated in management theory and practices.

Survival of reared Atlantic salmon and sea trout in relation to marine conditions of smolt year in the Baltic Sea

The marine survival of hatchery-reared Atlantic salmon ( Salmo salar) and sea trout ( Salmo trutta) was examined in relation to marine conditions during post-smolt migration and in relation to stock traits. In 1970–2001, Carlin-tagged smolts were released in the Iijoki and Oulujoki rivers, the northern Baltic Sea. When both species were analysed together, the abundance of the three prey fish, herring ( Clupea harengus), smelt ( Osmerus eperlanus) and vendace ( Coregonus albula) correlated positively with the survival of salmonids. In addition, the increase in smolt size appeared to improve the survival rate. Sea surface temperature (SST) may have affected indirectly through the abundance of prey fish during the post-smolt migration of salmon and sea trout. The smelt and vendace showed a statistical effect on survival only when the temperature effects were not included in the models. In sea trout, an increasing smolt length was not significantly correlated with the survival in good herring recruitment years, but in poor years survival increased very rapidly with increasing smolt size. The recapture rates of the salmonids tended to decrease between the years 1970 and 2001. During the same time period, the June SST slightly decreased. The positive correlation between the annual summer SST and recapture rate of salmon may partly explain the decreasing trend in recapture rates. An increase in smolt size did not compensate for the decline in the recapture rate of either species.

Mortality patterns in infectious salmon anaemia virus outbreaks in New Brunswick, Canada

Mortality levels attributed to infectious salmon anaemia viral (ISAV) infections were examined at the net pen and site level in the 1996 smolt year class in three areas of New Brunswick, Canada. The year class in this region was the first known to have potential exposure to ISAV beginning at the time of seawater transfer. There was considerable variability in mortality patterns among net pen groups of fish. Net pen outbreak definitions were based on at least seven high mortality days in which there were at least 100 per 100 000 fish per day or >5% cumulative mortality for the study period. There were 106 net pen outbreaks in a study population consisting of 218 net pens. Although the number of new cases decreased as water temperature decreased, overall mortality levels at the study sites did not decrease noticeably. The median peak daily mortality rate during outbreaks was 492 per 100 000 fish per day, with 10% of cases experiencing >5200 mortalities per 100 000 fish per day. The median duration of outbreaks in net pens for which the fish were not slaughtered during the outbreak was 33 days and the median total loss in those outbreaks was 6600 per 100 000 fish.

Is juvenile salmon abundance related to subsequent and preceding catches? Perspectives from a long-term monitoring programme

AbstractThe abundance of wild Atlantic salmon in the River Teno system has been monitored since the 1970s by estimating salmon catches and juvenile salmon densities at permanent electrofishing sites. Analysis of the time-series has shown significant relationships between juvenile densities (0+ and 1+) and subsequent 1SW and 2SW catches. Corresponding significant relationships have been detected between 1SW and 2SW female salmon in the catches and subsequent fry and parr densities. Monitoring juvenile densities allows evaluation of spawning escapement 1 and 2 years earlier, confirming the stock status information provided by catch statistics. These relationships between juvenile abundance and catches suggest that the monitoring programme has included feasible and biologically relevant variables and proper methodologies. Increasing trends were detected in the numbers of 1SW and 2SW salmon in catches between 1977 and 2003. Similarly, fry abundance indicated long-term increasing trends at most sites. Significant relationships were detected between abundances of subsequent sea-age groups in catches (1SW vs. 2SW 1 year later, etc.), indicating that strong smolt year classes influence the abundance of several subsequent sea-age groups, and that such relationships permit forecasting future catches of multi-sea-winter salmon by 1SW salmon catches.

Survival of reared and wild Atlantic salmon smolts: size matters more in bad years

Abstract We compared the marine survival of Carlin-tagged wild and hatchery-reared Atlantic salmon smolts of the Simojoki river, northern Baltic Sea. All the reared and released smolts were the offspring of native spawners returning to the river. Reared smolts were adipose-fin-clipped and released from the hatchery several weeks before tagging. The wild and reared smolts were simultaneously caught and tagged at a smolt trap located at the Simojoki river mouth. The study was conducted in two years, 1991 and 1993, when post-smolt survival in the Baltic Sea was different. Tags were returned by fishermen and return rates were used to estimate the survival of the smolt groups. We applied generalized linear models with survival as response variable and the year, origin, and smolt size as explanatory variables. On average, wild smolts had a 4.5 times higher survival rate than reared fish of the same smolt size. The difference in observed tag recovery rates as such was only about twofold or less, as the larger size of the reared smolts compared with the wild ones compensated for their lower survival rate. The better survival of wild than reared smolts was more pronounced in the low-survival year (1993 smolt year class) than in the high-survival year (1991 smolt year class).

Growth patterns of Labrador Sea Atlantic salmon postsmolts and the temporal scale of recruitment synchrony for North American salmon stocks

We measured postsmolt growth patterns from scales of Atlantic salmon postsmolts captured in the Labrador Sea and adult returns from stocks at the southern end of the range of salmon in North America. Because Labrador Sea postsmolts are believed to represent the juvenile nursery for the entire stock complex, the growth variation for southern stocks may yield insights on the time scale of stock mixing during the postsmolt year. Circuli spacing patterns were extracted from the scales of 1525 salmon for 3 smolt years. For 2 of the 3 years, growth trajectories for fish from the southern stocks intersected the trajectories for Labrador Sea postsmolts after 4-5 circuli pairs. The time that it takes to deposit a circulus during the spring and summer seasons suggests that distribution patterns for regional groups begin to overlap or stocks begin to experience similar environmental conditions by June or July of the postsmolt year, or 1 to 2 months after their migration to sea. In some years, however, it would appear that this mixing does not occur until fall. These data provide the first indication of the time scale of events during the postsmolt year that regional influences may be acting on stocks to produce synchronous recruitment.

The straying of Icelandic ranched Atlantic salmon, Salmo salar L.: release and recapture techniques

The straying rate of ranched Atlantic salmon, Salmo salar L., into rivers in Iceland was estimated on the basis of coded wire tag recoveries. Out of a total of 15 158 recaptured tagged salmon from the releases between 1987 and 1992, 189 fish (1.3%) strayed into 25 out of the 79 salmon rivers observed. Most of the strayers were found in neighbouring rivers to the ranching stations. There were no significant differences in straying rate between different age classes of salmon returning from the same smolt year class. Generally, there was a delay of 26-27 days in running time between strayers in rivers and salmon returning to the ranching station. Higher straying rates were observed for ranching stations using riverine traps than for stations using estuary traps. By close inspection of seven key rivers in Iceland and assuming a 50% exploitation rate (fishing effort) in the rivers, on average, 2.1% of the returning salmon in ranching were estimated to stray to native salmon rivers over the years. A limitation of using these numbers to estimate geneflow between ranching stocks and wild populations is discussed.

Relationships between smolt size, postsmolt growth and sea age at maturity in Atlantic salmon ranched in the Baltic Sea

Tagging data were used to examine the relationships between smolt size, post-smolt growth and sea age at first maturity for the short-migrating Neva strain of Atlantic salmon (Salmo salar L.) ranched in the Bothnian Sea and the Gulf of Finland. The results provided evidence that post-smolt growth was influenced by both relative and absolute smolt size. For both sea-areas, 2-year smolts of small relative size within a release group grew more rapidly in the sea than did smolts of higher relative, but equivalent absolute size. The negative influence of increasing relative smolt size on marine growth was, however, outweighed by the stronger positive influence of increasing absolute smolt size. A 160-mm increase in smolt size (140–300 mm) resulted in an overall growth advantage of about 1 year. In the Bothnian Sea, the predicted mean length after 1 year in the sea was 288 ± 25 mm for 140-mm smolts and 560 ± 16 mm for 300-mm smolts. Under the more favourable conditions of the Gulf of Finland, the respective mean lengths were 369 ± 15 mm and 613 ± 12 mm. The sea age at first maturity was inversely related to both freshwater and marine growth rates. For both sea areas, large smolts yielded proportionately more grilse than did small ones. Smolt years with good post-smolt growth rates yielded more grilse than did years with poor growth rates. The overall level of grilsing was higher in the Gulf of Finland than in the Bothnian Sea. These results suggest that the relationships between smolt size, post-smolt growth and age at first maturity in the sea are influenced by the environmental conditions of the respective sea area. A framework explaining the links between smolt size, marine growth, survival and sea age at maturity in Neva salmon is presented for the Gulf of Finland and the Bothnian Sea.