Long-term changes in the smolt size and age of Atlantic salmon, Salmo salar L., in a northern Baltic river related to parr density, growth opportunity and postsmolt survival

Abstract –  The size of wild Atlantic salmon ( Salmo salar L.) smolts in 1972–2004 was studied in relation to parr density, smolt age, growth opportunity and postsmolt survival in the Simojoki River. There was a significant negative regression between the annual mean smolt size and the density of wild >1 year parr in the previous autumn, but not between the annual mean smolt size and age. The density of reared parr released into the river or the growth opportunity, based on the day length and air temperature during the previous summer, did not affect the size of wild smolts. The data on postsmolt survival based on recaptures of Carlin-tagged smolts showed a significant positive relationship ( P  < 0.01) between the survival of postsmolts and the annual size of wild smolts. It is hypothesised that the increased density of wild >1 year parr could have contributed to the decreased smolt size since the 1990s, and the reduced size of wild smolts could be included among the factors resulting in their declined postsmolt survival in the Baltic Sea.     

Night stocking facilitates nocturnal migration of hatchery-reared Atlantic salmon, Salmo salar, smolts

Abstract  Hatchery-reared salmon, Salmo salar L., smolts are generally stocked during daylight hours, but the natural migration of smolts tends to occur at night. Recapture rates and timing of migration were compared between Atlantic salmon smolts stocked during the day and during the evening. Timing of release had no significant effect on the number of smolts recaptured, but had a strong effect on nocturnal behaviour. When stocked in the evening (but not during the day) hatchery-reared smolts moved almost exclusively during the night. This study suggests that timing the release to coincide with the natural time of smolt migration may provide valuable acclimatisation and facilitate nocturnal smolt passage.     

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

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

The source of autumn and spring downstream migrating juvenile Atlantic salmon in a small lowland river

Over 3 years, 32,444 age‐0 group Atlantic salmon (Salmo salar) parr tagged with passive integrated transponder tags throughout the River Frome catchment were assigned to one of three groups, nonmigrants, autumn migrants and spring smolts, depending on the detection and the timing of detection at downstream tag readers (situated 8.6 km above the tidal limit). We examined the effect of density at the time of tagging (n·m^?2), distance upstream from the tidal limit (km), fish length (mm), Fulton condition index, habitat type (divided into two types, main river and carrier), days after 1 September that each fish was tagged and year (replication) on the proportions of fish in each of the migration groups. Distance upstream from tidal limit was strongly negatively related to the proportion of autumn migrants and positively related to the proportion of spring smolts. Nonmigrants had a lower average body size than migrants, although there were no differences in the sizes of autumn migrants and spring smolts in September prior to migration. Fish density had no effect on migration strategy. A lower proportion of fish migrated as autumn migrants from the smaller carrier habitats than the main river channel. There is some evidence that those parr destined to become autumn migrants underwent a lower mortality rate during September before tagging than those destined to become spring migrants indicating possible physiological or behavioural differences between these two groups of fish at that time. More research into the factors responsible for initiating the autumn migration is required.     

Inter-stage survival of wild juvenile Atlantic salmon, Salmo salar L.

A biological model was developed to calculate annual survival between life stages of juvenile Atlantic salmon, Salmo salar L., in Catamaran Brook, a small stream basin (52 km²) in the Miramichi River catchment in New Brunswick, Canada. Seven years’ data (1990–1996) were used in the model. Input variables included: daily fish counts and measurements of parr (3–4 age classes), smolts, and adult salmon at a fish-counting fence near the stream mouth; biennial quantification of all habitat types along the watercourse; fish density estimated by electric fishing at 30 sites; and estimates of young-of-the-year emigration via stream drift. Continuous recording of stream discharge provided data to assist in interpretation of survival estimates. Annual survival for juvenile salmon in their first 3 years of life in the stream averaged between 31% and 34%. The greatest annual variation (CV = 0.699) occurred at the egg to 0+ (summer) stage with a low of 9.2% survival recorded for a winter with an atypical midwinter flood event; parr and pre-smolt survival were similarly affected. Survival from egg deposition (after correction for losses caused by predation and retention/non-fertilization) to smolt emigration was between 0.16% and 0.52%, which is low relative to estimates from many other studies. Survival of smolts to returning 1-sea-winter adults (grilse) averaged 8.5%. Potential errors in the computation of the model are discussed, e.g. inaccurate counts of spawning adults during high autumn stream flow. A possible explanation for the low egg to smolt survival was the environmental conditions experienced during various winters. Mean egg survival was 1.3 times higher (39.3%) and egg to smolt survival increased to 1.03% when the two winters characterized by extremely low discharge or midwinter freshets were excluded from the calculation. Density-dependent factors related to a beaver dam, which limited spawning distribution, may also have contributed to poor survival and increased fry emigration in one year. Environmental factors, particularly winter conditions, in streams such as Catamaran Brook may act as bottlenecks to natural production of Atlantic salmon.     

Physiological responses to continuous swimming in wild salmon (Salmo salar L.) parr and smolt

Wild salmon (Salmo salar) parr and smolt were forced to swim against constant flow (50 cm.s^–1) for 8 hours. Physiological properties describing the hormonal status, the energy metabolism and the ionic and osmotic balance of fish were measured from the fish prior to and at the end of the swimming test.Plasma cortisol levels were elevated in response to enforced swimming; the response of the smolt was clearly greater than that of the parr. Plasma thyroxine concentration increased in the parr but stayed at the initial level in the smolts. The parr consumed much of their coelomic fat, but the glycogen stores stayed nearly constant. The smolts had very low fat stores, and the glycogen stores were depleted in the test. The ionic and osmotic balance of the parr was stable in the test, but in smolts, the plasma Cl^–1 and osmotic concentrations decreased and muscle moisture increased.The results indicate that downstream migration smolts have markedly lower physiological capacity for continuous swimming than parr.     

Comparison of physiological smolt status in descending and nondescending wild brown trout (Salmo trutta) in a Danish stream

Abstract –  Downstream movement of a wild population of brown trout was examined in a small Danish stream in relation to morphological and physiological smolt status from March to May. Downstream movement was monitored in a Wolf-type trap covering all possible passage routes in the stream. Trout caught in the trap were classified as parr, pre-smolt or smolt based on morphological criteria and compared with trout randomly caught by electrofishing upstream of the trap. Representative gill samples from trap-caught and electrofished trout were analysed for gill Na⁺, K⁺-ATPase activity and used as a measure of physiological smolt status. Only a few parr occurred in the trap. Few pre-smolts occurred in the trap evenly in March and early April. In late April, pre-smolt movement peaked. By comparison, the main downstream movement of smolts occurred in distinct peaks through late March and April. The majority of fish caught in the trap were judged as pre-smolts or smolts based on morphological criteria's and they were characterised by relatively high gill Na⁺, K⁺-ATPase activity compared with trout judged as parr. Trout caught by electrofishing upstream the trap, were classified as parr, pre-smolts and smolts early in the season (March). During and after the main smolt-run in April the distribution of the remaining trout in the brook became skewed in favour of pre-smolt and parr. The study suggests that smolting trout initiate downstream movement once having reached a certain physiological smolt condition (judged by increased gill Na⁺, K⁺-ATPase activity).     

The smolt run and postsmolt survival of Atlantic salmon, Salmo salar L., in relation to early summer water temperatures in the northern Baltic Sea

Abstract –  The timing of the smolt migration of Atlantic salmon, Salmo salar L., was investigated during 1972–2002 in the Simojoki, a river flowing into the northern Baltic Sea. The onset of the smolt run was positively correlated with the river water temperature; a rise in water temperature above 10 °C being the main proximate environmental triggering factor. There was also a weaker correlation between the decreasing river discharge in the spring and the onset of the smolt migration. The duration of the main run was shorter in the years when the onset of the smolt run was delayed. No differences were found in the onset timing or in the duration of the smolt run between wild smolts and semi-wild smolts released into the river as parr. A polynomial equation fitted to the annual data on the survival of Carlin-tagged wild smolts and the sea surface temperature (SST) in June off the river mouth appeared to follow a dome-shaped pattern. Survival was lower in cold early summers (SST <9 °C) than in those with an average SST (9–11.9 °C), and lower again, although not significantly, in warm early summers (SST ≥12 °C). Too low and probably also too a high water temperature in early summer could thus be one of the underlying reasons for the fluctuations observed in postsmolt survival in the Baltic Sea.     

Early marine survival and movements of escaped Atlantic salmon Salmo salar L. juveniles from a land‐based smolt farm during autumn

The aim of this study was to examine early marine survival and movements of simulated escaped Atlantic salmon Salmo salar L. pre‐smolt and smolt from a commercial smolt farm during autumn. One‐third of the pre‐smolt most likely died in the immediate vicinity of the release location, whereas the corresponding mortality for smolts was lower (8.5%) during the 5‐week study period. The surviving pre‐smolt left the farm area after 2–3 days, predominantly along the shore. In contrast, most of the surviving smolts left the farm area during the first day and 54% seemed to move away from the shore and adopt a more pelagic movement pattern than pre‐smolt. The number of surviving fish recorded in the fjord decreased throughout the study period, possibly due to a combination of fish migrating out of the fjord or undetected mortality. Compared with existing knowledge on migration of released farmed smolts during spring, our results indicate less directional and slower movement rates during autumn. Only two of the tagged fish were detected upstream in the rivers following release. A rapid dispersion of escapees indicates that the potential for recapturing escapees is limited unless recapture efforts are initiated immediately after escape. Hence, there is a need for development of technology that detects and prevents escapees to enter the sea.     

Development of an accurate model to predict the phenology of Atlantic salmon smolt spring migration

1. Changes in migration timing, resulting from the alteration in river continuity or the effect of climate change, can have major consequences on the population dynamics of diadromous fish. Forecasting the phenology of fish migration is thus critically important to implement management actions aimed at protecting fish during their migration.
2. In this study, an 11‐year monitoring survey of Atlantic salmon smolts (Salmo salar) from the Ourthe River, Belgium, was analysed within a European Special Area of Conservation to improve the understanding of environment‐induced spring migration. A logistic model was fitted to forecast smolt migration and to calculate phenological indicators for management, i.e. the onset, end, and duration of migration, while accounting for the influence of photoperiod, water temperature, and hydrological conditions.
3. The results indicated that the photo‐thermal units accumulated by smolts above a 7°C temperature threshold was a relevant proxy to reflect the synergistic effect between temperature and photoperiod on smolt migration. After integrating the effect of river flow pulses, the model accurately explained the inter‐annual changes in migration timing (R² = 0.95). The model predictions provide decisive management information to identify sensitive periods during which mitigation measures (e.g. hydropower turbine shutdown, river discharge management) should be conducted to promote smolt survival.
4. The model was used to predict phenological characteristics under future scenarios of climate change. The results suggest a joint effect of hydrological alterations and water warming. Temperature increases of 1–4°C were associated with earlier initiation of migration, 6–51 days earlier, and spring flood events greatly influenced the duration of the migration period. Accordingly, the combined effects of human‐induced modifications of the hydrological regimes and increasing temperatures could result in a mismatch between the smolt and favourable survival conditions in the marine environment.

     

The effects of stocking Atlantic salmon, Salmo salar, in a Norwegian regulated river

Abstract  Stocking is undertaken in the River Suldalslågen, western Norway, to compensate for an estimated annual loss of 20 000 Atlantic salmon smolts, Salmo salar L., caused by regulating the river for hydropower production. The annual contribution to angling catches from stocked hatchery fish varied from 7 to 334 kg, or <15% of the total number caught. Between 160 000 and 250 000 one-summer old fish were stocked, but only between 6 and 10 (<0.005%) were recaptured as adults in the river. Recaptured stocked fish never exceeded 0.03% by number, despite smolts dominating the stocking material in recent years. It is not certain whether the slight increase in catches comes in addition to or at the expense of natural reproduction. In most years more adults were used as parent stock than were caught as offspring. The lack of positive response to stocking is possibly due to lesser age, smaller size and later migration of hatchery smolts, and that seawater tolerance of hatchery smolts is poorly developed, all factors increasing mortality at sea.     

Epidemiology of Gyrodactylus salaris (Monogenea) in the River Tornionjoki, a Baltic wild salmon river

The occurrence of Gyrodactylus salaris in the River Tornionjoki was investigated in 2000-2004. Infection of salmon parr, Salmo salar, was common in the uppermost reach of the river system but decreased downstream and was rare in the lowermost reach. This pattern was consistent across the study period regardless of varying water temperatures. The oldest age groups of parr were more often infected than younger ones throughout the river system, irrespective of their origin (wild or stocked). Parasite-free hatchery-reared 1-year-old parr became infected during their first summer in the wild. Downmigrating salmon smolts had a high prevalence of infection, but their role in the distribution of infection seemed unimportant. On grayling, Thymallus thymallus, we observed only the grayling-specific clade of Gyrodactylus. We found no indication of grayling participating in the epidemiology of infection on salmon. The salmon parr and smolt population in the Tornionjoki has been at its height during the late 1990s and 2000s. Our results indicate that G. salaris infection in this Baltic river has no devastating effects on the salmon population as it has had in salmon rivers flowing into the North Atlantic and White Sea.     

Interactive effects of life history and season on size‐dependent growth in juvenile Atlantic salmon

Size‐dependent growth (SDG) is an important process in structuring populations as well as determining life history outcomes. Despite its importance, there have been few investigations from observational studies focusing on the interaction between life history decisions and SDG. In this study, we used data on individually tagged Atlantic salmon from both the laboratory and the field to investigate differences in SDG among two life history groups, parr and smolts. In the laboratory, we found little evidence of SDG in parr but seasonally dependent SDG in the smolt group. Smolts showed at strong compensatory response over the winter months just prior to the smolt transformation window. In the field, we found little evidence of SDG early in ontogeny (i.e., age 0+ fall and winter). There was some evidence of depensatory growth (positive SDG) during the age 1+ spring among both life history groups that may reflect random habitat variation or the monopolisation of resources. After the age 1+ spring, we found that smolts were more likely to show a compensatory effect (negative SDG) than parr. This effect was strongest, as they approached the smolt window in the spring of their age 2+ year. These results suggest (i) SDG is common in Atlantic salmon; however, the form and extent of life history depends on (ii) season and (iii) life history. For individuals that adopt a smolt life history, trade‐offs between freshwater survival and sea survival may lead to a convergent growth pattern, as they approach the smolt migration window.     

Effects of smolt size and timing of migration on recovery rate of wild masu salmon <Emphasis Type="Italic">Oncorhynchus masou</Emphasis>

Little is known about the survival rate of wild masu salmon Oncorhynchus masou. To examine the effects of smolt length and migration timing on the recovery rate of wild masu salmon, we reanalyzed past tagging and recovery data (1993–1994). The tagging study was conducted in the Shokanbetsu River, northern Japan; 863 wild masu salmon smolts were captured, tagged, and released in a downstream site, and a total of 19 fish were recovered in coastal fisheries and in the natal river the following year. The data were analyzed by a logistic regression analysis with recapture as a response variable and tagging date and smolt length as explanatory variables; the tagging date had a significant effect on the recapture rate, whereas the effect of smolt size was not significant. Despite the small number of recaptures, this study indicates that migration timing is a factor affecting the marine survival of wild masu salmon smolts, although this conclusion has been repeatedly documented for other species of Pacific salmon Oncorhynchus spp.     

Nomads no more: early juvenile coho salmon migrants contribute to the adult return

The downstream movement of coho salmon fry and parr in the fall, as distinct from the spring migration of smolts, has been well documented across the range of the species. In many cases, these fish overwinter in freshwater, but they sometimes enter marine waters. It has long been assumed that these latter fish did not survive to return as adults and were ‘surplus’ to the stream's carrying capacity. From 2004 to 2010, we passive integrated transponder tagged 25,981 juvenile coho salmon in three streams in Washington State to determine their movement, survival and the contribution of various juvenile life histories to the adult escapement. We detected 86 returning adults, of which 32 originated from fall/winter migrants. Half of these fall/winter migrants spent ~1 year in the marine environment, while the other half spent ~2 years. In addition, the median return date for fall/winter migrants was 16 days later than spring migrants. Our results indicated that traditional methods of spring‐only smolt enumeration may underestimate juvenile survival and total smolt production, and also overestimate spring smolt‐to‐adult return (SAR). These are important considerations for coho salmon life cycle models that assume juvenile coho salmon have a fixed life history or use traditional parr‐to‐smolt and SAR rates.     

Exposure to seawater stimulates lipid mobilization from depot tissues of juvenile coho (Oncorhynchus kisutch) and chinook (O. tshawytscha) salmon

Tissue lipid content and lipolytic enzyme activity was determined in selected tissues of coho salmon,Oncorhynchus kisutch, at various developmental stages (freshwater parr, freshwater smolt, seawater smolt, and seawater stunt) and in tissues of coho salmon and chinook salmon,O. tshawytscha, exposed to seawater periodically during smoltification. Among developmental groups, total lipid concentration of liver and dark muscle was highest in freshwater (FW) parr. Lipid concentration in both liver and dark muscle was significantly lower in FW smolts, seawater (SW) smolts and SW stunts; no difference was observed among these groups. Alterations in lipid composition were reflected in depot triacylglycerol lipase activity. FW smolts, SW smolts and SW stunts displayed significantly higher lipase activity than FW parr in each of the tissues examined (live, dark muscle and mesenteric fat). Early in smoltification (March, April), exposure to seawater results in enhanced lipid depletion from liver, dark muscle and mesenteric fat, both 30 and 60 days after exposure, compared to FW controls. This depletion was accompanied by increased liver (March and April) dark muscle (March) and mesenteric fat (March) lipase activity. Later in smoltification (May), salinity-induced alterations in lipid metabolism were not observed. These results indicate that exposure to seawater stimulates lipid depletion in juvenile salmon and that the depletion can be explained, in part, by increased depot lipase activity. Furthermore, these data confirm that metabolic dysfunction is associated with stunting.     

Sea Trout of the North Esk

The sea trout population of the River North Esk was sampled by means of a stationary trap situated on Kinnaber Mill Lade which drains its water from the main river. Commencing in 1971 the number of trout taken in the trap was noted daily, and from 1976 all stages of trout caught in the trap were tagged. The peak of the sea trout smolt migration was usually in May or June, and over the period of study exhibited a tendency to occur later each season. Two-year-old smolts comprised the largest age class in the smolt samples with 3-year-old fish being the other substantial age class. Samples of smolts taken for sex determination were found to be predominantly female. In 1979 and 1980 the sea trout smolt production of the North Esk was estimated from a mark-recapture experiment to be 51,969 and 78,208 respectively. A proportion of finnock (post-smolts) returned to fresh water in the year of their smolt migration and were taken moving upstream into the trap from July onwards, with a peak of movement occurring in the autumn. The main run of adult sea trout into the trap was found to take place at any time between July and October. The adult sea trout sampled were predominantly maiden spawners of ages 2.1+ and 3.1+ years. Both finnock and adult sea trout samples exhibited a similar freshwater age composition to that of the smolts.     

Influence of environmental and biological factors on the overwinter growth rate of Atlantic salmon Salmo salar parr in a UK chalk stream

Smolt lengths are increasingly recognised as an important determinant of salmonid marine survival rates. Overwintering growth rates could thus strongly influence adult return rates. In Atlantic salmon Salmo salar, most overwintering studies focus on rivers in harsh climates with minimal growth, yet in more southerly rivers, overwintering growth rates can be relatively high. Here, the factors influencing annual overwinter growth rates were tested for juvenile S. salar in a temperate chalk stream in southern England over 13 years, where over 10,000 salmon parr were tagged annually in autumn and a proportion recaptured the following spring during smolt emigration. Winters of higher and more variable water temperatures, with longer periods of high flows, showed increased overwintering growth rates. Faster growth rates were recorded from sites further upstream and that had lower parr densities; smaller individuals also grew more than expected for their initial size. These results suggest that a range of factors influences overwintering salmonid growth rates and can be used to inform management decisions to maximise the quality of emigrating smolts.     

Density of brown trout, Salmo trutta L., and Atlantic salmon, Salmo salar L., parr after point and scatter stocking of fry

Atlantic salmon, Salmo salar L., and brown trout, Salmo trutta L., fry were point and scatter stocked in the early part of June at densities of 63–263 fry 100 m⁻² per species in the River Viantienjoki, a small river in northern Finland, and their population densities were assessed in late summer. Both species were always stocked together in similar quantities. Point stocking was used in the first 2 years and scatter stocking in the following 2 years. In point stocking, there was no correlation between the distance from the stocking sites (maximum = 250 m) and parr density in census sites ( r = −0.013 and 0.019 for brown trout and Atlantic salmon, respectively). The stocking density of fry did not influence parr density in August by either method or by species. Stocking density explained only from 11% to 23% of the parr survival depending on the species or stocking method. The mean densities of Atlantic salmon and brown trout parr did not differ significantly from each other at any fishing site ( P > 0.05). Both point and scatter stocking appear to be suitable methods for use in small rivers. The parr densities depend more on the other factors (e.g. habitat quality) than the stocking method, and the choice between methods could be based on the time and labour available.     

The Irish sea trout enhancement programme: an assessment of the parr stocking programme into the Burrishoole catchment

The success of the sea trout, Salmo trutta L., enhancement programme carried out in the Burrishoole catchment in the west of Ireland from 1993 until 1998 is reviewed in terms of the number of fish stocked, the number of fish which migrated to sea and the number which returned from sea during this period. The success of the programme is also evaluated in terms of the cost of producing parr for stocking and the subsequent value of returning post-smolts (0+ sea age) and rod caught fish. A total of 49 235 tagged fish were released into Lough Feeagh, between 1993 and 1998. Of these fish, 14 788 were microtagged and the remainder, 34 447 had either elastomer or alphanumeric visible implant (VI) tags. Tag loss rates at release varied from 3 to 5.7%. Over the course of the programme a maximum of 7801 fish migrated through the traps towards sea, of which 581, or 7.4%, returned. In any 1 yr a maximum 13.1% of fish returned from sea. The average cost of producing a parr for stocking out was €1.15, which translated into average values of €8.16 per smolt and €169 per returning post-smolt.