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.     

Association between environmental factors, smolt size and the survival of wild and reared Atlantic salmon from the Simojoki River in the Baltic Sea

The survival of Atlantic salmon Salmo salar in the Baltic Sea was examined in relation to smolt traits (length and origin) and annual environmental factors [sea surface temperature (SST) and seasonal North Atlantic Oscillation (NAO) index], and prey fish abundance (herring Clupea harengus and sprat Sprattus sprattus) in the main basin and the southern Gulf of Bothnia. The study was based on recapture data for Carlin‐tagged hatchery‐reared and wild smolts from the Simojoki, a river flowing into the northern Gulf of Bothnia. The survival of the wild and reared groups was analysed using an ANOVA model and a stepwise regression model, with the arcsin‐transformed proportion of recaptured fish as the response variable. The results demonstrated a combined influence of smolt traits and environmental factors on survival. For the reared Atlantic salmon released in 1986–1998 (28 groups), the increasing annual mean SST in July in the southern Gulf of Bothnia and increasing mean smolt size improved survival. If the SST in July was excluded from the model, the NAO index in May to July also had a positive effect on survival (P < 0·10). The log₁₀‐transformed abundance of 0+ year herring in the southern Gulf of Bothnia entered the model (P < 0·15) if the SST and NAO index were excluded. For the wild Atlantic salmon released in 1972–1993 (21 groups), only the increasing SST in July showed a significant association with improved survival (P = 0·004). Prey fish abundance in the main basin of the Baltic Sea had no influence on the survival of reared or wild smolt groups. The interaction between smolt size and the SST in July was not significant. The origin was a better, but not a significant, predictor of marine survival compared to the smolt size or the SST in July. The mean recapture rate of the wild groups was twice that of the reared groups in the whole data. The results suggest that cold summers in the Gulf of Bothnia reduce the survival of young Atlantic salmon in both wild and reared groups. The larger smolt size of the reared groups compared with the wild groups to some extent compensated for their lower ability to live in the wild.     

Stomach analyses of Baltic salmon from 1959–1962 and 1994–1997: possible relations between diet and yolk-sac-fry mortality (M74)

In recent years, Baltic Sea salmon Salmo salar have suffered high larval mortality (M74) which can be cured by thiamine treatment. Analyses of long term mortality records (1928–1998) from two salmon hatcheries suggest that before the 1970s M74 did not occur, or was less frequent. This indicates that varying M74 did not cause the long-term fluctuations of Baltic salmon catches in this period. The frequency of M74 has been correlated positively to the abundance of the salmon's primary prey, sprat Sprattus sprattus. Sprat, herring Clupea harengus and three-spined stickleback Gasterosteus aculeatus dominated the diet in both the periods of study, 1959–1962 and 1994–1997. The mean size of consumed sprat was significantly smaller in 1994–1997 compared with 1959–1962. Herring and, to a lesser extent, three-spined stickleback increased in the diet of salmon, while sprat appeared to constitute a smaller part of the diet in 1994–1997. The cause of M74 and the thiamine deficiency involved remains unknown, but is thought to be related to changes in thiamine or thiaminase content in forage fish, winter-feeding of salmon or general changes in the pelagic food web, caused by overfishing or eutrophication.     

Development of length–bimodality and smolting in wild stocks of Atlantic salmon, Salmo salar L., under different growth conditions

Growth dynamics of juvenile Atlantic salmon, Salmo salar L., from two sections of the Narcea River and one of the Esva River (Northern Spain) were examined in relation to the development of bimodality in their size–frequency distributions. Size–bimodality was clearer under intermediate growth (section A) than under relatively fast or slow growth. The proportion of fish entering the upper modal group increase with growth intensity. Composition of upper and lower modal groups became fixed prior to December, and at this time both groups separated on the 90–95 mm interval. Fish exhibiting smolt appearance in late March (larger than 130 mm) had already been upper group fish in December, while parr-like fish and those that remained in the river by May (potential 2-year-old smolts) had formed the lower modal group. Anadromous salmon catch in the Narcea River was mostly of previously 1-year-old smolts (97.6%), of which 94% were larger than 100 mm by their first winter. In the Esva River, slow growth of juveniles is consistent with a large proportion of 2-year-old smolts (47.9%) among anadromous salmon. Both juvenile samples and scale analysis of anadromous salmon indicate that 2-year-old smolts were larger than 1-year-olds. Early disappearance of the former (before March) is, at least, partially related to earlier migration of large fish, since sexual maturity of parr does not provide a complete explanation. The Narcea stock have a minimum length at smolting of about 130 mm and an optimum smolt size in the 155–175 mm interval. Mean smolt length did not vary although the winter length changed between years.     

The influence of precocious sexual maturation on survival to adulthood of river stocked Baltic salmon, Salmo salar, smolts

During three consequtive years, 1975–1977, Individually tagged Baltic salmon Salmo salar smolts of sexually immature male and female fish (n = 35027, mean size: 15.2 cm) and precocious males (n = 6518, mean size: 14.2 cm) were released into Umeälven (Ume river), northern Sweden. Rate of survival (% captured adults) based on 3714 recoveries was significantly higher (p < 0.01) for smolts from immature fish (10.2%) than those from smolts of early maturing males, i.e. precocious males (2.2%). corresponding to an average yield of 474 and 85 kg per KHX) smolts released, respectively. Gain in survival was on average 2.5% and 1.4% per cm increase in smolt size for immature smolts and smolts from precocious males, respectively. The poor survival among smolts of precocious males is suggested to he related to an interaction between sexual maturation and smolting linked to incompletely resorbed gonads leading to a non migratory behaviour. These non migratory males are then suggested to suffer heavily by predation in the river.
The two smolt categories had a similar growth pattern in sea. Smolts from precocious males did not mature early in sea indicating no relation to grisling, i.e. sexually maturing fish returning after first winter in sea. Adult weight of fish returning the fourth summer after release was related to smolt size (P < 0.05). Our Response Surface Model (RSA) predicted that large smolts (19.0 cm) had a higher specific growth rate over their life-span compared to small smolts (<15.0 cm), 0.86% d⁻¹ and 0.46% d⁻¹, respectively. Large smolts (19.0 cm) attained a size of 3.0 kg during their second winter in sea about six months earlier than small smolts (13.0 cm). The paper discusses alternative release strategies that can be employed if the ultimate goal of salmon stocking is maximizing yield.     

Does increased abundance of sea lice influence survival of wild Atlantic salmon post‐smolt?

A synthesis of results from two projects was assessed to analyse possible influence of sea lice Lepeophtheirus salmonis on marine Atlantic salmon Salmo salar survival. During the years 1992–2004, trawling for wild migrating post-smolts was performed in Trondheimsfjord, a fjord in which no Atlantic salmon aquaculture activity is permitted. Prevalence and intensity of sea lice infections on migrating wild post-smolts differed between years. A correlation analysis between 1 sea-winter (SW) Atlantic salmon catch statistics from the River Orkla (a Trondheimsfjord river) and sea lice infections on the migrating smolts in the Trondheimsfjord was not significant. Up to 2% reduction in adult returns due to sea-lice infection was expected. In addition, experimental releases from 1996 to 1998 with individually tagged groups of hatchery-reared Atlantic salmon smolts given protection against sea-lice infection was performed. Higher recaptures of adult Atlantic salmon from 1998 treated smolts compared to the control group may correspond to high abundance of sea lice found on the wild smolt, and may indicate influence on post-smolt mortality. These studies indicate that post-smolt mortality in Trondheimsfjord is marginally influenced by sea lice infection; however, the methods for assessing wild smolt mortality might be insufficient. Higher infections of sea lice farther out in the fjord may indicate more loss in Atlantic salmon returns in some years.     

Geographical differences in marine feeding of Atlantic salmon post‐smolts in Norwegian fjords

Stomach content analyses were conducted on Atlantic salmon Salmo salar post‐smolt (average size, 119–154 mm fork length, L _F) caught in eight large Norwegian fjord systems along a north–south geographical axis during 1998–2001. In general, post‐smolts from southern Norway showed low feeding intensity in the fjords, whereas extensive feeding was observed in fjords in the northern and middle parts of Norway. The marine diet mainly included different crustaceans and in particular marine pelagic fish larvae (sand-eels Ammodytes spp., herring Clupea harengus and gadoids), but with a substantial spatial and annual variation in prey diversity and feeding intensity. Insects were most frequently taken in the estuary, although fishes often made a large contribution in mass. In contrast, fishes, and to some extent various crustaceans (particularly Hyperiidae, Gammaridae, Euphausiacea and Copepoda) dominated the diet in the middle and outer parts of the fjords, where post‐smolts also fed more extensively than in the inner part. The results indicate that extensive feeding immediately after sea entrance may be more common for post‐smolts in the northern and middle parts of Norway, than in the southern fjords. The observed differences in post‐smolt feeding may be due to spatial and temporal differences in prey availability within and between the different types of fjord systems, and this might influence post‐smolt growth and survival.     

Carlin tag recoveries as an indicator of predation on salmon smolts by goosanders and red-breasted mergansers

Between 1984 and 1990 a total 221 Carlin tags used to mark salmon Salmo salar smolts in the River North Esk, NE Scotland, were recovered from the stomachs of goosanders Mergus merganser and red-breasted mergansers M. serrator . Both Carlin-tagging and adipose-clipping affected the predation of salmon smolts by sawbill ducks. The mean (± S.D.) sizes of tagged smolts taken by both species were similar (117 ± 3 mm) and significantly smaller than the mean sizes of smolts in the river, possibly due to a reduction in the swimming performance of small smolts bearing tags. Large adipose-clipped smolts (±mean smolt size) were predated significantly more than unclipped smolts, but no such difference was observed for small smolts (相似文献   

Marine post-smolt growth and age at maturity of Atlantic salmon

The annual variation in sea-age of maturation for a hatchery dependent stock of Atlantic salmon was compared to variation in post-smolt growth as evidenced by circuli spacing patterns. The proportion of returns of 1-seawinter (1 SW) and 2 SW salmon and the fraction of the smolt year class or cohort that maturated as 1 SW fish, were compared to seasonal growth indices determined from circuli spacing on the scales of smolt class survivors returning as 1 SW and 2 SW spawners. Using image processing techniques, we extracted inter-circuli distances from scales from 2244 recaptured fish. Spacing data for the first year at sea were collected and then expressed as seasonal growth indices for the spring period, when post-smolts first enter the ocean; the summer, when growth appears maximal; and winter, when growth appears to be at a minimum. In general, circuli spacings were wider for 1 SW than for the 2 SW returns of the same smolt cohort. The 1 SW fraction was significantly and positively correlated with late summer growth, suggesting that growth during this season is pivotal in determining the proportion of a smolt class that matures early.     

Do Norwegian Atlantic salmon feed in the northern Barents Sea? Tag recoveries from 70 to 78° N

Three tagged Atlantic salmon Salmo salar were recaptured as subadults or adults (1·4–3 kg) between 70·5 and 78° N in the western Barents Sea, two of which originated from the Alta Fjord region in northern Norway and one from the Drammen River, south-eastern Norway. An additional tag was recovered from the stomach of a Greenland halibut Reinhardtius hippoglossoides captured south-west of Bear Island at >600 m depth; this tag was from a smolt released in the River Alta 1 month earlier. These are the northernmost tag recoveries reported for Atlantic salmon, and indicate that Norwegian Atlantic salmon, especially the fish from northern populations, may use the northern Barents Sea as a feeding area during part of their life cycle.     

The physiological effects of salmon lice infection on post-smolt of Atlantic salmon

The physiological effects of salmon lice infections on post-smolt of Atlantic salmon were examined by experimentally infecting hatchery reared post-smolts with infective copepodids. Even at high infection intensities, ranging from 30–250 lice per fish, early chalimus stages did not have severe, physiological effects on the fish. There was a sudden increase in fish mortality after the appearance of preadult I stages. Infected fish were then suffering due to lesions and osmoregulatory failure. Plasma chloride level increased significantly and total protein, albumin and haematocrit decreased significantly in infected compared to uninfected fish. All infected fish became moribund before adult lice appeared. Infection intensities above 30 salmon lice larvae per fish thus appear to cause death of Atlantic salmon post-smolt soon after the lice reach their pre-adult stage.     

Variation in the post-smolt growth pattern of wild one sea-winter salmon (Salmo salar L.), and its linkage to surface warming in the eastern North Atlantic Ocean

Variation in circulus spacing on the scales of wild Atlantic salmon is indicative of changes in body length growth rate. We analyzed scale circulus spacing during the post-smolt growth period for adult one sea-winter salmon (n = 1947) returning to Scotland over the period 1993–2011. The growth pattern of the scales was subjectively and visually categorized according to the occurrence and zonal sequence of three intercirculus spacing criteria (“Slow”, “Fast” and “Check” zones). We applied hierarchical time-series cluster analysis to the empirical circulus spacing data, followed by post hoc analysis of significant changes in growth patterns within the 20 identified clusters. Temporal changes in growth pattern frequencies showed significant correlation with sea surface temperature anomalies during the early months of the post-smolt growth season and throughout the Norwegian Sea. Since the turn of the millennium, we observed (a) a marked decrease in the occurrence of continuous Fast growth; (b) increased frequencies of fish showing an extended period of initially Slow growth; and (c) the occurrence of obvious growth Checks or hiatuses. These changes in post-smolt growth pattern were manifest also in decreases in the mean body length attained by the ocean midwinter, as sea surface temperatures have risen.     

Age, sex ratio and timing of the catch of kelts and ascending Atlantic salmon in the subarctic River Teno

By 15 June, 82% of the catch of Atlantic salmon Salmo salar kelts had been taken from the middle part of River Teno, northern Scandinavia. The median date of capture was 4 June for males and 8 June for females. Salmon of 1–4 sea–winters (SW) of both sexes survived spawning to return to sea as kelts. Among males, 1 SW kelts were caught earliest in the spring and 3 SW latest, but among females 4 SW were earliest, then 3 SW and finally 1 and 2 SW. There were 17 river and sea–age combinations among the kelts compared with 23 among the ascending salmon. The smolt age distribution and the mean smolt age differed significantly only between female 2 SW ascending salmon (3·97 years) and kelts (4·14 years). The proportion of 1 SW females was higher and that of 3 SW males lower among kelts than among ascending salmon. The proportion of males among 1 SW ascending salmon was 80% but among kelts only 57%. Similarly, the proportion of males among 3 SW fish was 21% for ascending salmon but only 7% for kelts. Hence overwinter mortality was higher among males. Male and female kelts of 1 and female kelts of 2 SWhad a greater mean length than ascending salmon in corresponding groups indicating a better survival of larger fish within an age group. Grilse ascend rivers after most kelts have left, but the main catch of ascending 2–3 SW salmon takes place concurrently with kelts leaving the river, inadvertently targeting kelts in the fishery.     

Feeding of wild and hatchery reared Atlantic salmon (<Emphasis Type="Italic">Salmo salar</Emphasis> L.) smolts during downstream migration

In general, hatchery salmonid smolts experience higher mortality during migration than wild smolts, which is suggested to be due to domestication effects and that hatchery fish lack experience of the natural environment. However, possible differences in feeding during smolt migration between hatchery and wild smolts have rarely been addressed. We compared the number of feeding smolts and stomach fullness among wild Atlantic salmon smolts, hatchery-reared smolts released as 1-year-old parr, and hatchery-reared smolts released as 2-year-old smolts during their descent to sea in River Tornionjoki. In addition, estimations of prey selection among the smolt groups were conducted. A high proportion of wild smolts and smolts stocked as parr actively fed during the smolt migration. A lower proportion of smolts stocked as smolts was feeding and their stomach fullness were much reduced in comparison with the two other groups. The study also indicated that the feeding of migrating smolts is selective rather than opportunistic. In conclusion, this study suggests that stocked 2-year-old smolts may enter sea with an inferior foraging behaviour and it is a possibility that this may contribute to the observed low post-smolt survival in the Baltic Sea.     

The effect of temperature and somatic growth on otolith growth: the discrepancy between two clupeid species from a similar environment

Otolith growth rates of the early life stages of herring Clupea harengus ( n = 472) and smelt Osmerus eperlanus ( n = 348) collected in the Vistula Lagoon (Baltic Sea) during 1997–1999 were analysed. The larvae and early juveniles were not only collected in the same geographical area they were also of the same size (range 15–43 mm standard length, L _S), similar ages and were collected during the same seasons (May to July). Although the two clupeid species experienced very similar environmental conditions, there were significant discrepancies in the analysed relationships. The otolith growth of larval and juvenile smelt was very strongly related to somatic growth while temperature had a minor effect. In herring, the effect of somatic growth, although clearly visible and statistically highly significant, was of less importance than temperature. Furthermore, variation in the otolith size and L _S relationship was affected by temperature and somatic growth in both species, but the variance of otolith size at L _S was higher for herring than for smelt. Although growth backcalculation from otoliths can presently be recommended as an appropriate method for use with both smelt and herring (despite possibly lower precision and accuracy with the latter), other methods referring directly to short-term increment width changes ( e.g. marginal increment analysis) are recommended for smelt but not for herring.     

The effect of prey quality and ice conditions on the nutritional status of Baltic gray seals of different age groups

We analyzed a long-term data set of the body condition of Baltic gray seals (Halichoerus grypus) over time and investigated how average subcutaneous blubber thickness of different age groups of seals corresponds to environmental factors. Blubber thickness of pups declined until 2010. The decreasing weight of 5–6-year-old herring (Clupea harengus), the main prey fish for Baltic gray seals, explained well the decline. In the Gulf of Finland, the blubber thickness of pups declined also in recent years (2011–2015) with declining number of days with permanent ice cover. In other regions, the blubber thickness of pups increased during recent years with increasing weight of herring. The blubber thickness of sub-adults in Baltic Proper and that of hunted adult females in the Bothnian Bay also increased during recent years, and the weight of age 6+ or 7-year-old herring best explained the increase. The blubber thickness of all age groups of seals was thinnest in the Bothnian Bay where also herring weight was lowest. There was a negative correlation between blubber thickness of seals and herring catch size (an index of herring abundance) suggesting that herring quality, not the quantity, is important for the nutritional status of Baltic gray seals. Nutritional status of gray seals may thus reveal changes in the marine food web which affect herring quality. Marine food web, in turn, may be affected, e.g., by climate change. The warming climate also has an impact on ice cover and thus body condition of seal pups.     

Assessing seasonal changes in stock composition of Atlantic salmon catches in the Baltic Sea with genetic stock identification

The feasibility of using genetic stock identification to analyse seasonal changes in stock compositions of Atlantic salmon catches in the Baltic Sea was examined. The analysis employed seven variable allozyme loci from most of the potentially contributing stocks (16) from Finland and Sweden. Catch samples were collected from Finnish salmon fisheries in the eastern Bothnian Sea during the 1992 fishing season. Simulation studies were used to evaluate the feasibility of identifying Baltic salmon stocks with allozyme data. Special attention was paid to analysing the wild production of salmon stocks. Clear seasonal differences in stock composition were found. The estimates were compared with smolt production and Carlin-tag data. The proportions of the Neva and Oulujoki river stocks could be estimated as individual stocks, whereas the contributions of the remaining stocks were estimated as four composite stock groups. One of the groups consisted of wild stocks from the rivers Kalixälven and Simojoki. Identification of this group, which could be used as an index of wild production in the catches, requires catch sample sizes >300 salmon if <15% error is required.     

Sea growth, smolt age and age at sexual maturation in Atlantic salmon

Relationships between growth at sea, smolt size and age at sexual maturation of Atlantic salmon Salmo salar were tested. The fish were offspring of brood stocks sampled in eight Norwegian rivers at latitudes between 59° and 70° N, hatchery reared and released at smolting at the mouth of the River Imsa (59° N). Smolt size influenced the subsequent growth rate of Atlantic salmon. The larger the fish were at release, the slower the yearly length increment at sea. Mean sea age at sexual maturity, measured as proportion of the returning adults attaining sexual maturity at sea age 2 years, was significantly correlated with mean growth rate during the first year at sea and mean smolt size ( r ²= 0·74, P < 0·001). Fish attaining maturity at a relatively high sea age were more fast growing during their first year at sea than those maturing at a younger age. The results indicate that high sea age at sexual maturation is a population-specific characteristic and associated with high early growth rate at sea.     

A size-dependent migration strategy in Atlantic salmon smolts: Small smolts favour nocturnal migration

Migration theory states that migration behavioural strategies should be optimised to maximise fitness. Many studies have shown that in downstream migrating Atlantic salmon Salmo salar L. smolts, mortality from predation is high and negatively size dependent. The most common predators are birds and piscivorous fish that are mainly daylight feeders. Given the high mortality during this stage we should expect to observe smolts to follow predator avoidance strategies that may be affected by body size. We tested the hypothesis that small smolts have a higher tendency to exhibit predator avoidance strategies (i.e. nocturnal versus diurnal migration) than larger smolts. The number and size of out-migrating/downstream-migrating wild Atlantic salmon smolts was recorded as they passed through a glass-sided channel during April-May, 1996–1999. In all years, the mean size of nocturnal migrating smolts was significantly lower than the mean size of diurnal migrating smolts. Analysis of the size of smolts, during early and late stages of the migration period showed size-dependent nocturnal migration behaviour up to the end of April. After this, no such size dependent migration pattern was observed. However, small smolts (<100 mm) were absent during this period. We suggest that nocturnal migration is an adaptive behaviour that small Atlantic salmon smolts have to avoid predation by large daylight feeding visual piscivorous predators (e.g. pike Esox Lucius L. and fish eating birds).     

Interannual and regional variations in body length, weight and energy content of age-0 Pacific herring from Prince William Sound, Alaska

Age-0 Pacific herring were surveyed in October of 4 years in a large northern Gulf of Alaska estuary, to determine the range of variations in length, weight and whole body energy content (WBEC). These parameters reflect their preparedness for surviving their first winter's fast. During the surveys there were distinct regional and interannual variations in all three parameters for individual groups of herring in Prince William Sound. Likewise, with each collection there was typically a large range of size and WBEC values. The average standard length was (±S.D.) 80±13 mm (range=40–118), the mean whole body wet weight was 5·7±3·0 g (range=0·7–29·2) and the average WBEC of all age-0 herring captured, regardless of year or site (n=1471), was 5·4± 1·0 kJ g⁻¹ wet weight (range=2·4–9·4). The large range of WBEC and size indicates that age-0 herring at different capture sites were not all equally prepared for surviving their first winter.