Environmental change influences the life history of salmon Salmo salar in the North Atlantic Ocean

Annual mean total length (L_T) of wild one‐sea‐winter (1SW) Atlantic salmon Salmo salar of the Norwegian River Imsa decreased from 63 to 54 cm with a corresponding decrease in condition factor (K) for cohorts migrating to sea from 1976 to 2010. The reduction in L_T is associated with a 40% decline in mean individual mass, from 2 to 1·2 kg. Hatchery fish reared from parental fish of the same population exhibited similar changes from 1981 onwards. The decrease in L_T correlated negatively with near‐surface temperatures in the eastern Norwegian Sea, thought to be the main feeding area of the present stock. Furthermore, S. salar exhibited significant variations in the proportion of cohorts attaining maturity after only one winter in the ocean. The proportion of S. salar spawning as 1SW fish was lower both in the 1970s and after 2000 than in the 1980s and 1990s associated with a gradual decline in post‐smolt growth and smaller amounts of reserve energy in the fish. In wild S. salar, there was a positive association between post‐smolt growth and the sea survival back to the River Imsa for spawning. In addition, among smolt year‐classes, there were significant positive correlations between wild and hatchery S. salar in L_T, K and age at maturity. The present changes may be caused by ecosystem changes following the collapse and rebuilding of the pelagic fish abundance in the North Atlantic Ocean, a gradual decrease in zooplankton abundance and climate change with increasing surface temperature in the Norwegian Sea. Thus, the observed variation in the life‐history traits of S. salar appears primarily associated with major changes in the pelagic food web in the ocean.     

Development of seawater tolerance and subsequent downstream migration in wild and stocked young‐of‐the‐year derived Atlantic salmon Salmo salar smolts

This study investigated the development of hypo‐osmoregulatory capacity and timing of downstream migration in wild Atlantic salmon Salmo salar smolts from the River Stjørdalselva and stocked young‐of‐the‐year (YOY), derived S. salar smolts from the tributary River Dalåa. Both wild and stocked S. salar smolts developed seawater (SW) tolerance in early May, persisting through June, measured as their ability to regulate plasma osmolality and chloride following 24 h SW (salinity = 35) exposure. Although the majority of downstream migration among the stocked S. salar smolts occurred later than observed in their wild counterparts, the development of SW tolerance occurred concurrently. The wild S. salar from Stjørdalselva and stocked YOY smolts from the River Dalåa started to migrate on the same cumulative day‐degrees (D°). The study revealed no downstream migration before development of SW tolerance. This emphasizes the importance of incorporating physiological status when studying environmental triggers for downstream migration of S. salar smolts. Overall, these findings suggest that the onset of smolt migration in stocked S. salar smolts was within the smolt window from an osmoregulatory point of view.     

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

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

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

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

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.     

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

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

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.     

Long‐term effects of photoperiod,temperature and their interaction on growth,gill Na+, K+‐ATPase activity,seawater tolerance and plasma growth‐hormone levels in Atlantic salmon Salmo salar

This study was undertaken to examine the long‐term effects of photoperiod, temperature and their interaction on growth, gill Na⁺,K⁺‐ATPase (NKA) activity, seawater tolerance and plasma growth‐hormone levels in Atlantic salmon Salmo salar pre‐smolts and smolts. The fish (mean ± s.e . initial body mass = 15·9 ± 0·4 g) were reared on two photoperiods (continuous light, LL, and simulated natural photoperiod, LDN, 60° 25′ N) and two temperatures (8·3 and 12·7° C) from June to May of the following year. Mean body mass was affected by photoperiod, temperature and their interactions. Both temperature groups on LL developed peak levels in gill NKA activity from October to November, 4–5 months prior to the natural season for the parr–smolt transformation. Fish at 12° C showed peak levels in NKA activity 4–6 weeks before the fish at 8° C. Fish in all four experimental groups showed maximum NKA activity within a similar size range (113–162 g). The present findings further indicate that smoltification in S. salar is to some extent driven by size, and that S. salar will develop smolt characteristics, e.g. a marked increase in NKA activity, within a similar size range. Faster‐growing S. salar will, thus, reach this size threshold at a relatively younger age.     

The effect of stocking with 0+ year age‐class Atlantic salmon Salmo salar fry: a case study from the River Bush,Northern Ireland

An enhancement programme based on stocking 0+ year age‐class Atlantic salmon Salmo salar, conducted in the River Bush, Northern Ireland, U.K. over the period 1996–2005, was reviewed with reference to the performance and biological characteristics of wild fish. Wild ova to 0+ year fry (summer) survival was c. 8% with subsequent wild 0+ year fry‐to‐smolt survival c. 9%. Stocked unfed 0+ year juveniles gave c. 1% survival to smolt whilst fed 0+ year S. salar stocked in late summer exhibited survival at c. 5%. Stocking with unfed and fed fry contributed to increased smolt production and helped attain local management objectives between 2001 and 2005. Significant differences in biological characteristics were observed between wild and stocked‐origin fish. Wild‐smolt cohorts were dominated by 2+ year age‐class fish on the River Bush whilst smolts originating from fed fry mostly comprised younger 1+ year individuals. The mean mass of 1+ year smolts derived from stocked fed fry was significantly lower than that of wild 1+ year smolts, although these differences were not evident between older age classes. Differences in run timing between wild smolts and smolts derived from stocked fry were also apparent with the stocked‐origin fish tending to run earlier than wild fish. Although the stocking exercise was useful in terms of maximizing freshwater production, concerns over the quality of stocked‐origin recruits and the long term consequences for productivity are highlighted.     

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.     

Retrospective growth analysis of Atlantic salmon Salmo salar and implications for abundance trends

Scale archives of Atlantic salmon Salmo salar from Maine, U.S.A., were examined to determine whether ocean conditions affected the long‐term trends in S. salar populations in the southern tier of the species' range in North America. To date, scale analyses of southern tier populations have been limited to hatchery fish; previous studies suggest that post‐smolt growth does not influence recruitment, with the exception that winter growth may play a role in stock maturation rate. A time series of scales from the Machias and Narraguagus Rivers spanning the years 1946 to 1999 was analysed. Image analysis was used to measure intercirculi spacing, which provided proxy variables of growth rate. Post‐smolt growth increment has increased since the early 1990s, as returns have decreased, suggesting that survival factors act on post‐smolts independent of growth. The data support the hypothesis of a decoupling between freshwater size and early marine growth. Growth during the second sea winter was independent of post‐smolt growth, suggesting that individuals are capable of significant compensatory growth. Southern tier North American stocks exhibit a similar pattern of independence between growth and survival as observed for northern tier North American stocks. These data support the inference that the recruitment of the North American and European subspecies is governed by fundamentally different mechanisms.     

Changes in scale circulus spacings of an endangered Atlantic salmon Salmo salar population: evidence of a shift in marine migration?

Post‐smolt scale circulus spacing patterns for two Atlantic salmon Salmo salar populations from the Southern Upland (SU) of Nova Scotia, Canada, were compared with spacings from two endangered populations from the inner Bay of Fundy (iBoF) Nova Scotia and New Brunswick, to determine if growth patterns differed among these populations, and if growth patterns had changed as the abundance of these populations declined. An analysis of numbers of marine circuli from scales of post‐smolts and one‐sea‐winter adults of known age indicated that circuli were deposited at a rate of about one circulus per week in summer and slowed to one every 2 weeks in winter. During the summer and the autumn, mean circulus spacing in the iBoF populations, known to have occupied the outer Bay of Fundy during these seasons, was lower than in the SU populations, which are known to migrate to the North Atlantic. Similar circulus spacing patterns within SU populations is suggestive of a common marine distribution for these populations. In contrast, a cluster analysis revealed that within the geographically intermediate Big Salmon River (iBoF), some individuals exhibited wider spacing patterns that resemble the distant migrating SU populations, while others exhibited narrower spacing similar to other iBoF S. salar. Within the Big Salmon River, the relative abundance of the wider and the narrower spacing patterns varied in the earlier years, but all fish sampled since 1999, exhibited wider spacings similar to distant migrating SU S. salar. The apparent disappearance of the narrower pattern, characteristic of localized migration and indicative of historical iBoF populations, suggests that local migration may not presently be a successful strategy for these populations.     

Water discharge affects Atlantic salmon Salmo salar smolt production: a 27 year study in the River Orkla,Norway

A model that explains 48% of the annual variation in Atlantic salmon Salmo salar smolt production in the River Orkla, Norway, has been established. This variation could be explained by egg deposition, minimum daily discharge during the previous winter and minimum weekly discharge during the summer 3 years before smolt migration. All coefficients in the model were positive, which indicates that more eggs and higher minimum discharge levels during the winter before smolt migration and the summer after hatching benefit smolt production. Hence, when the spawning target of the river is reached, the minimum levels of river discharge, in both winter and summer, are the main bottlenecks for the parr survival, and hence for smolt production. The River Orkla was developed for hydropower production in the early 1980s by the construction of four reservoirs upstream of the river stretch accessible to S. salar. Although no water has been removed from the catchment, the dynamics of water flow has been altered, mainly by increasing discharges during winter and reducing spring floods. In spite of the higher than natural winter discharges, minimum winter discharge is still a determinant of smolt production. Hence, in regulated rivers, the maintenance of discharges to ensure that they are as high as possible during dry periods is an important means of securing high S. salar smolt production.     

Genetic assessment of Atlantic salmon Salmo salar and sea trout Salmo trutta stocking in a Baltic Sea river

Microsatellite DNA variation was used to assess the outcome of stocking Atlantic salmon Salmo salar and migratory trout Salmo trutta in River Sävarå, N Sweden. No information on pre‐stocking genetic composition of S. salar and S. trutta in River Sävarå was available. In 2 year‐classes of S. salar smolt, microsatellite data indicated that post‐stocking genetic composition differed markedly (F_ST= 0·048) from the main donor strain, Byskeälven S. salar, and from other Gulf of Bothnia S. salar stocks (F_ST 0·047 and 0·132). The STRUCTURE programme failed to detect any substructuring within Sävarå salmon. It was concluded that only minor introgression estimated to a proportion of 0·11 (95% CI 0·07–0·16) has occurred in S. salar. Salmo trutta showed overall low differentiation among populations with maximum F_ST of 0·03 making analysis more cumbersome than in S. salar. Still, the SävaråS. trutta deviated significantly from potential donor populations, and STRUCTURE software supported that majority of trout in Sävarå formed a distinct genetic population. Admixture was more extensive in S. trutta and estimated to 0·17 (95% CI 0·10–0·25).     

Migration of Atlantic Salmon, Salmo salar, Smolt through the Estuary Area of River Ellidaar in Iceland

Synopsis We tagged both wild and hatchery Atlantic salmon, Salmo salar, smolts from River Ellidaar (64 ° 08′ N, 21 ° 50′ W) with ultrasonic tags. We caught the wild smolts in a smolt trap and selected the largest individuals from the run. We implanted the transmitters in the abdominal cavity of the fish and then released them in River Ellidaar close to the estuary. We used four ultrasonic receivers; one in the river’s estuary, one outside the estuary and two further away on both sides of an island in the estuary zone. The receivers recorded all transmitters within a 600 m radius. The hatchery smolts were larger than the wild smolts. Some of the smolts were lost on the way through the estuary. Only 4 out of 9 wild smolts and 14 of 17 hatchery smolts were recorded all the way through. The tags and the tagging likely affected the survival of the smolts especially the smaller fish. There were no differences in the smolt migration between the 2 years of study and no differences in the migration behavior between the wild and the hatchery smolts. After being released the smolts stayed on average for 10 h in the river then migrated into the estuary were they stayed for 54 h on average. Then they migrated straight through the area to the sea at approximately 0.2 fish lengths per second. We recorded large differences in the migration.     

Changes in smolt traits of Atlantic salmon (Salmo salar Linnaeus, 1758) and linkages to parr density and water temperature

Smolt traits (length, age) and timing of smolt migration of wild Atlantic salmon, Salmo salar L., were investigated in the Simojoki River, northern Baltic Sea. The aim was to determine whether they responded to changes in parr length, parr density and temperature from 2000 to 2014. Annual electrofishing surveys and smolt numbers determined parr densities by springtime trapping in the river mouth. During the smolt trapping period captured parr and smolts were aged from scales. Water temperature was measured daily. Mean length decreased from 137 mm (TL) to 129 mm among 2‐year‐old smolts, and from 150 mm to 139 mm among 3‐year‐olds. Median date of the smolt migration was 10 days earlier, from early June to late May during the study period, linked to the rise in air temperature in May at the nearby Kemi‐Tornio airport. However, the median day temperature and the mean daily water temperatures during the second (Q₂) and third (Q₃) migration quartiles did not change. This implied that migration began when a suitable water temperature was reached, independent of the date.     

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

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

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

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

Does size matter? A test of size‐specific mortality in Atlantic salmon Salmo salar smolts tagged with acoustic transmitters

Mortality rates of wild Atlantic salmon Salmo salar smolts implanted with acoustic transmitters were assessed to determine if mortality was size dependent. The routinely accepted, but widely debated, ‘2% transmitter mass: body mass’ rule in biotelemetry was tested by extending the transmitter burden up to 12·7% of body mass in small [mean fork length (L_F) 138·3 mm, range 115–168 mm] downstream migrating S. salar smolts. Over the short timescale of emigration (range 11·9–44·5 days) through the lower river and estuary, mortality was not related to S. salar size, nor was a relationship found between mortality probability and transmitter mass: body mass or transmitter length: L_F ratios. This study provides further evidence that smolt migration studies can deviate from the ‘2% rule’ of thumb, to more appropriate study‐specific measures, which enables the use of fishes representative of the body size in natural populations without undue effects.