Effect of origin, sex and sea age of Atlantic salmon on their recapture rate after river ascent

The recapture rate of Atlantic salmon (Salmo salar L.) after river ascent was examined by the trapping and tagging of ascending spawners in the lower reaches of the Simojoki River, which flows into the northern Baltic Sea. In 1997 and 1998, altogether 825 Carlin‐tagged salmon were released to continue their upstream migration. Of these, 800 could be sexed and categorized as reared (91%) or wild (9%) salmon. In 1997, most of the ascending salmon were multi‐sea‐winter (MSW) fish, whereas in 1998 almost all were one‐sea‐winter (1SW) male grilse due to the late trapping season. About 10% of all tagged fish were recaptured, two‐thirds of which were caught in the river before their descent to the sea. There was no difference in the recapture rate between salmon of wild (8.5%) or reared (9.5%) origin, or between females (11.6%) and males (9.3%). Generalized linear models for data from 1997 showed that the recapture rate increased with length and age of females, but that the opposite was true for males. River fishing did not seem to remove proportionally more early ascending salmon than fish that ascended later.     

Divergence in smolt production from the stocking of 1-summer-old and 1-year-old Atlantic salmon parr in a northern Baltic river

One‐year‐old and 1‐summer‐old salmon parr have been stocked in the Simojoki, a northern Baltic river, to protect the natural salmon stock from extinction. This enhancement practice was studied during a 5‐year stocking period, when an average of 14.3 1‐summer‐old parr and 5.4 1‐year‐old parr was needed to produce one sea running smolt. The mean yield from stocking was 70 smolts per 1000 stocked 1‐summer‐old parr and 186 smolts per 1000 stocked 1‐year‐old parr. The mean cost of smolts produced by stocking 1‐year‐old and 1‐summer‐old parr was 1.2 and 2.6 times the cost of producing one 50 g smolt in the hatchery, respectively. There was a negative relationship between density of wild parr and length of smolts stocked as 1‐summer‐old parr in the river, but no relationship between wild parr density and length of smolts stocked as 1‐year‐old parr. This indicated that 1‐summer‐old parr were more susceptible than the older parr to competition with wild parr. Considering production costs and smolt yield, stocking of 1‐year‐old parr appeared to be more profitable than that of 1‐summer‐old parr in the enhancement of the endangered salmon stock.     

The survival of semi‐wild, wild and hatchery‐reared Atlantic salmon smolts of the Simojoki River in the Baltic Sea

The recapture rate and survival of hatchery‐reared Atlantic salmon Salmo salar stocked as 1 year‐old parr (semi‐wild) with that of hatchery‐reared Atlantic salmon stocked as 2 year‐old smolts and wild smolts of Atlantic salmon in the northern Baltic Sea were compared. This was done through tagging experiments carried out in 1986–1988 and 1992. The recapture rate of the semi‐wild groups varied from 1·0 to 13·1%, being similar in 3 tagging years and lower in 1 year than that of the wild groups (1·7–17·0%). The recapture rate of the semi‐wild groups was similar (in 2 years) or higher (in 2 years) than that of the hatchery‐reared groups stocked as smolts (1·3–6·3%). The survival of semi‐wild smolts during the sea migration was as high as that of wild Atlantic salmon of an equal size and two to three times higher than hatchery‐reared Atlantic salmon stocked as smolts. The survival rate was positively associated with smolt size. The suitability of hatchery‐reared parr and smolts in the management of reduced Atlantic salmon stocks is compared.     

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.     

Post-smolt migration of Atlantic salmon, Salmo salar L., from the Simojoki river to the Baltic Sea

The migration pattern of Atlantic salmon ( Salmo salar L.) post-smolts in the Baltic Sea was investigated based on tag recoveries of Carlin-tagged wild and hatchery-reared smolts released in the Simojoki river in 1972–2005. Exact date of sea entry was known only for the wild smolts. Tag recoveries of wild salmon in the estuary within 10 km from the river mouth were received on average 3.5 days (±2.0 SD) after release. Time required for emigration from the estuary was dependent on the sea surface temperature (SST) off the river ( R ² = 0.625, P = 0.004), being shorter in years with warmer than colder sea temperatures. Outside the estuary, the wild and hatchery-reared post-smolts migrated southwards along the eastern coast of the northern Gulf of Bothnia, the tag recoveries coinciding with the warm thermal zone in the SST occurring along the coastal area. After arriving in the southern Gulf of Bothnia in late summer the post-smolts mostly migrated near the western coast, reaching the Baltic main basin in late autumn. The relationships between the marine conditions and migration patterns are discussed.     

Determination of body shape variation in Irish hatchery‐reared and wild Atlantic salmon

Discriminant function analysis was used to distinguish morphologically between samples of parr, smolts and adult Atlantic salmon Salmo salar from several hatchery and river systems in Ireland. The effect of habitat shift was investigated in Atlantic salmon parr. Parr grown from the eyed‐egg stage with a non‐sibling group in a hatchery environment, came to resemble the mean body shape of their host hatchery Atlantic salmon stock more closely than that of a full sibling group grown at their natal hatchery. Wild Atlantic salmon smolts differed in shape from hatchery‐reared smolts. This difference was less pronounced, but still statistically significant when wild adults were compared with hatchery‐reared adults captured in the coastal drift‐net fishery after a year spent at sea. Rearing conditions had a significant impact on the production and growth of fish body shape. This in turn may have affected adaptability and survivorship of ranched Atlantic salmon in the marine environment.     

The effect of sexual maturation on the spatial distribution of Baltic salmon

This study examines the migratory habits of tagged mature and immature Atlantic salmon parr Salmo salar released from the Norrfors hatchery on the River Umeälven, Sweden from 1975 to 1977 and 1988 to 1990. Tags were recaptured in the Baltic Main Basin, the Gulf of Bothnia, and in the Umeälven. Ninety-three per cent of previously mature males were recovered in the Umeälven compared with 23% of the previously immature smolts during the calendar year of release. In the second year after release (grilse year), the proportion of early maturing males recovered in the Umeälven was significantly greater than the proportion of previously immature smolts recovered in the Umeälven. Likewise, the proportion of previously mature males recovered in the Main Basin was significantly less than the proportion of previously immature smolts recovered in the Main Basin in the second calendar year after release. Previously mature males rematured after fewer years at sea, on average, than the previously immature smolts. Following the second calendar year after release, the proportions of previously mature males and previously immature smolts were not significantly different throughout the Baltic Sea.     

The importance of stocking age in the enhancement of River Kymijoki salmon (Salmo salar)

Hatchery‐reared Atlantic salmon parr and smolts are regularly released into the River Kymijoki, Southern Finland, to enhance the local river and coastal fisheries. In 1988–1994, a series of micro‐ and Carlin‐tagging experiments were carried out to examine the influence of stocking age on the stocking results. Five age groups were compared: 1‐year and 2‐year‐old smolts, and 1‐summer, 1‐year and 2‐summer‐old parr. The aim was to determine the most profitable way of producing salmon for fishing in the River Kymijoki. Stocking age had a strong influence on the stocking results, measured as the proportion of adult recoveries. The most favourable results were obtained using 2‐year‐old smolts (survival index 100), followed by 1‐year‐old smolts (52), 2‐summer‐old parr (51), 1‐year‐old parr (37) and 1‐summer‐old parr (24). Available data on rearing costs suggest that 2‐year‐old smolts were also the most economically profitable choice, followed by 1‐year‐old parr, 1‐year‐old smolts and 1‐summer‐old parr. The most expensive way of producing salmon was by stocking 2‐summer‐old parr.     

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.     

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.     

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.     

Genetic implications of hatchery rearing in Atlantic salmon: effects of rearing environment on genetic composition

In an experiment to investigate genetic consequences of hatchery rearing in salmon, allozyme variation at five polymorphic loci was examined in Atlantic salmon of known initial genetic composition, which were reared throughout freshwater life in the hatchery or stocked into the wild as swim-up fry. The genetic composition of the juveniles in the hatchery remained homogeneous from fertilization up to stocking, and from stocking to 2+ in the wild, however, those remaining at the hatchery developed genetic differences among smolting and nonsmolting 1+ parr. These differences were attributed to conditions leading to early smolting at 1+ among the hatchery fish, with 1+ smolts diverging from the gene pool from which they were derived, whereas those stocked into the wild did not smolt until a year later and retained the original genetic composition. The results are discussed in relation to hatchery rearing of salmon and implications for the use of reared fish in stocking and enhancement programmes.     

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.     

Effect of hatchery environment on cranial morphology and developmental stability of Atlantic salmon (Salmo salar L.) from North‐West Russia

The study addresses the effect of hatchery rearing on morphological variation and developmental stability of Atlantic salmon parr from North‐West Russia. Totally, we collected nine samples. Four wild samples were collected from each of the rivers Kola, Umba, Keret’ and Shuia. Five samples of hatchery‐reared parr were the first‐generation progeny of wild adults from these rivers reared separately at the four hatcheries (one hatchery was represented by two samples). Ten meristic and 48 morphometric cranial characters were analysed. We studied the morphological divergence between wild and hatchery fishes of the same river of origin. To analyze developmental stability we used fluctuating asymmetry (random deviations from perfect bilateral symmetry). It was found that hatchery‐reared parr significantly differ from wild parr in both meristic characters and the shape of cranial bones. Different hatcheries caused similar effect on morphological variation in all populations. Fluctuating asymmetry in morphometric characters was significantly higher in hatchery fish than in wild from the Shuia River, indicating a higher level of developmental instability. However, wild parr from the Keret’ River had significantly higher fluctuating asymmetry than cultivated parr of the same origin, possible due to a high infection pressure of the parasite Gyrodactylus salaris Malmberg which has led to significant decline of the wild salmon population in this river, or from genetic changes caused by cultivation. The obtained results indicate a notable effect of hatchery environment on Atlantic salmon’s phenotype.     

Temporal variation in abundance, return rate and life histories of previously spawned Atlantic salmon in a large subarctic river

The 30 year time series analyses revealed large temporal variation in the return rates and a recent increase in abundance of previous spawning Atlantic salmon Salmo salar in the River Teno, northern Scandinavia. The mean proportion of repeat spawners was 7 and 4% in the total Atlantic salmon catch and 9 and 22% in multi‐sea‐winter (MSW) catch component for females and males, respectively. Previous spawners constituted on the average 7% of the catch in mass but up to 20%(31 t) and 30%(19 t) in 2003 and in 2004, respectively. In 1975–2000, the proportion of previous spawners varied between 1 and 6%(3–12% of MSW Atlantic salmon), whereas in 2001–2004, they accounted for 8–21%(16–35% of MSW Atlantic salmon) of the total Atlantic salmon catch. The number of previous spawners in the catch correlated significantly with the preceding numbers of respective 1–3 sea‐winter (SW) maiden Atlantic salmon 2 years earlier. The recent increase in the numbers of 1S1 and 2S1 (1 or 2 years at sea followed by first spawning and 1 year reconditioning period at sea) alternate spawning Atlantic salmon was a consequence of higher numbers of maiden 1SW and 2SW Atlantic salmon in the catches and increased sea temperatures. Similarly, the return rate of 1SW Atlantic salmon to second spawning has improved in recent years. Most previous spawners ascended and were captured early in the fishing season. The smolt and sea‐age combinations of repeat spawners comprised 68 age groups contributing with the annual mean of 15 age groups to the great diversity of the River Teno Atlantic salmon population complex.     

Physiological stress responses to confinement in diploid and triploid Atlantic salmon

Confinement of juvenile Atlantic salmon Salmo salar from four different populations (all–female diploids, all–female triploids, mixed sex diploids and mixed sex triploids) either before (FW parr) or after (SW smolts) transfer to sea elevated plasma cortisol and plasma lactate from control levels irrespective of ploidy status. Both before and after confinement, plasma cortisol levels in SW smolts (6–174 ng ml^–1) were higher than those in FW parr (4–58 ng ml^–1), which possibly reflected the physiological challenge of acclimation to SW. Mixed sex populations of SW smolts had higher cortisol levels than all–female populations. The duration of confinement (1, 3 or 6 h) affected the magnitude of the plasma cortisol and lactate responses in SW smolts. Plasma cortisol levels in diploid and triploid SW smolts subjected to 2 h of confinement decreased to pre–stress levels within 6 h post–confinement. Plasma lactate levels were not significantly different from pre–stress levels 48 h after confinement. As no difference exists in primary and secondary stress responses of Atlantic salmon of differing ploidy status, it is unlikely that differences in mortality rates between diploid and triploid populations under commercial conditions can be attributed to differences in their physiological responses to periods of stress lasting up to 6 h.     

Individual growth and smoltification of juvenile masu salmon, Oncorhynchus masou Brevoort, under rearing conditions

In a reared population of individually marked juvenile masu salmon, individual growth was monitored from the first autumn in 1983 to the following spring. The potential smolts were not significantly greater in mean fork length and body weight than the potential parr in late August of the first year, but they then grew faster until March of the second year. As a result, the potential smolts formed the upper mode of the bimodal length distribution after February. Especially in autumn (October and November) the specific growth rates of potential smolts were significantly greater than those of parr, and the bimodality in growth rate distribution was more distinct for males than females. These suggest that there are two groups having different growth rates in autumn of the first year and that sufficient growth in this period may play an important role in smoltification in the following spring.     

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.     

Survival and migration patterns of naturally and hatchery-reared Atlantic salmon (Salmo salar) smolts in a Lake Ontario tributary using acoustic telemetry

1. Atlantic salmon (Salmo salar) smolts are often stocked into rivers to supplement natural reproduction, yet hatchery-reared fish have lower survival compared to wild conspecifics. However, few studies have assessed riverine migratory performance and survival differences in hatchery and wild smolts, or more specifically naturally reared smolts (hatchery fish released earlier as parr), particularly in rivers with weirs which may further reduce survival.
2. Using acoustic telemetry, including a subset of fish with novel transmitters that identify predation events, we assessed survival and migration patterns of hatchery- (2017: n = 32; 2018: n = 30) and naturally reared Atlantic salmon smolts (2017: n = 8; 2018: n = 30) in a Lake Ontario tributary with two weirs to better understand their ecology and assess the influence of environmental parameters on migration.
3. Naturally reared smolts were 13.9 times more likely to survive than hatchery-reared smolts and mark–recapture models indicated that weirs did not reduce survival for either group. Survival per km was lowest at the release site, indicating pre-migration mortality, and specifically high stocking-related mortality of hatchery-reared smolts. Speed and times of day fish migrated (i.e. migratory performance) did not vary by rearing group, suggesting that the high mortality of hatchery-reared smolts may be due to other factors related to hatchery and stocking operations. Overall mean (± SD) migration speed for smolts was 0.70 ± 0.39 km/hr and movements occurred significantly more frequently at night (18:00–06:00).
4. Smolts were detected in Lake Ontario after they left the river; however, the array in Lake Ontario was not conducive to providing much detail regarding movement patterns. There was no predation of the two predation tags detected in Lake Ontario, indicating that movements were made by smolts and not predators.
5. With ongoing restoration efforts of Atlantic salmon in Lake Ontario, it was important to understand the smolt migration patterns and success of the stocked fish. Our findings of similar migratory performance yet different relative survival of hatchery- and naturally reared smolts help inform management with regards to stocking strategies that could improve Atlantic salmon reintroduction success.

     

Fitness reduction and potential extinction of wild populations of Atlantic salmon, Salmo salar, as a result of interactions with escaped farm salmon

The high level of escapes from Atlantic salmon farms, up to two million fishes per year in the North Atlantic, has raised concern about the potential impact on wild populations. We report on a two-generation experiment examining the estimated lifetime successes, relative to wild natives, of farm, F(1) and F(2) hybrids and BC(1) backcrosses to wild and farm salmon. Offspring of farm and "hybrids" (i.e. all F(1), F(2) and BC(1) groups) showed reduced survival compared with wild salmon but grew faster as juveniles and displaced wild parr, which as a group were significantly smaller. Where suitable habitat for these emigrant parr is absent, this competition would result in reduced wild smolt production. In the experimental conditions, where emigrants survived downstream, the relative estimated lifetime success ranged from 2% (farm) to 89% (BC(1) wild) of that of wild salmon, indicating additive genetic variation for survival. Wild salmon primarily returned to fresh water after one sea winter (1SW) but farm and 'hybrids' produced proportionately more 2SW salmon. However, lower overall survival means that this would result in reduced recruitment despite increased 2SW fecundity. We thus demonstrate that interaction of farm with wild salmon results in lowered fitness, with repeated escapes causing cumulative fitness depression and potentially an extinction vortex in vulnerable populations.