Riverine and early ocean migration and mortality patterns of juvenile steelhead trout (<Emphasis Type="Italic">Oncorhynchus mykiss</Emphasis>) from the Cheakamus River,British Columbia

High mortality (65–73%) occurred in the first month of the smolt migration in a population of wild steelhead trout. We used acoustic telemetry to monitor the downstream, estuarine, and early ocean migration of tagged smolts and estimate their mortality rates. After entering the Strait of Georgia most smolts migrated north through Johnstone and Queen Charlotte Straits rather than south through the Strait of Juan de Fuca. Of 51 smolts tagged in 2004 (49 in 2005), 36–38 (41–42) survived to leave freshwater and 14–19 (13–14) survived to leave the Strait of Georgia system. Mortality rates in separate segments of the migration were correlated with segment distances. An additional component of mobile sampling showed that few smolts died during the migration through Howe Sound. Migration rates averaged 0.7–0.9 body lengths per second (BL s⁻¹) downstream and 1.0–2.6 BL s⁻¹ in ocean waters. Aggregated detection probabilities of 92–96% on lines of ocean receivers suggest that migration routes of small fishes can be quantified over several hundred kilometres, and survival rates can be estimated for even a modest number of tagged fish. Quantifying mortality patterns during the smolt migration could help to determine causes of low marine survival rates observed in recent years.     

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.     

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).     

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.     

The effects of water temperature on the migration of smolts of the Atlantic salmon Salmo salar L.

The dependence of the duration of downstream migration of smolts of Atlantic salmon (Salmo salar) on water temperature of the Lyl’ok River, a tributary of the Iokan’ga River (Kola Peninsula) is considered. It is shown that the duration of the migration of smolts depends on temperature conditions of the river in the second to third ten-day periods of July. An increase in the number of days with a daily average temperature over 20°C causes the deterioration of the physiological state of fish, the retardation of their linear growth, and a decrease in the intensity of migration, which results in a delayed stay in the river of smolts, shift of the dates of termination of migration, and an increase of its total duration. Nevertheless, smolts migrating from the river in August to September are ready to pass to life in the sea.     

Migration of wild and hatchery reared smolts of Atlantic salmon, Salmo salar L., through lakes

In 1982 and 1983 descending wild Atlantic salmon smolts, Salmo salar L., were caught in a fish-trap at the mouth of the River Imsa. Together with hatchery-reared smolts of the River Imsa strain they were tagged and released at three different sites of the Imsa-Lutsi watercourse: in the Imsa River 1 km above the trap, and in two lakes, 3 and 11 km upstream of the trap. The recapture-rate in the fish-trap decreased with increased migration distance. The hatchery-reared smolts migrated downstream faster than wild smolts. Lake-released smolts were considerably delayed in their downstream migration compared to the river released smolts. The results are discussed in relation to environmental and physiological parameters.     

Effects of training on functional variables of muscles in reared Atlantic salmon Salmo salar smolts: connection to downstream migration pattern

The relative amount of muscle contraction regulating dihydropyridine and ryanodine receptors in the swimming muscles of trained reared Atlantic salmon Salmo salar smolts was compared with those of untrained and wild smolts. After an optimized 2 week training period, i.e. swimming with a velocity of 1·5 body lengths per second for 6 h per day, the level of both receptors was significantly higher in the muscles of trained S. salar than in the untrained ones before they were released into the natural environment. This difference persisted after downstream migration in the river. The highest level of receptors was observed in wild S. salar. Swimming performance was also higher in trained fish compared to untrained ones. Furthermore, swimming performance was positively associated with the level of receptors in both red and white muscle types. Downstream migration after release into the wild was significantly slower in trained smolts than in untrained fish. This indicates that trained smolts were most probably swimming harder against the current in the river than untrained smolts. The possible advantages for a slower migration in the river are discussed. This study shows that the prerequisites for effective contraction of the swimming muscles are better met in trained S. salar compared to untrained fish, and the muscles of trained smolts more closely resemble those of wild smolts. The results also imply that the capacity of untrained, reared smolts to swim against the current is not equal to that of their trained or wild counterparts which affects the downstream migration pattern of S. salar smolts.     

Hydropower-related mortality and behaviour of Atlantic salmon smolts in the River Sieg,a German tributary to the Rhine

We studied downstream migration of 256 radio-tagged Atlantic salmon smolts passing a low-head power station where technical facilities have been improved to allow safe migration via several bypass routes. Extra mortality was 7 and 17% (two years) in the power station reservoir, and a minimum of 10 and 13% at the power station compared to in a control stretch. The majority of the smolts followed the main flow at the power station, towards the turbines. Sloped bar racks with 10 mm bar spacing hindered smolts from entering the turbines, hence there was no turbine mortality. Smolts used surface openings in the racks, which directed them to a bypass route outside the turbines. The extra mortality in the reservoir and at the power station was related to physical injuries in bypass routes and to predation. The mortality risk in the reservoir and at the power station decreased with increasing migration speed. Migration speeds increased with water discharge. Migration was slower when the smolts passed the power station than on other stretches. This study shows that hydropower regulation caused elevated mortality and delays for downstream migrating fish, even with improved technical facilities to reduce mortality.     

Electrophysiological Study of Effects of Environmental Factors on Respiration and Cardiac Activity in the Fry of the Atlantic Salmon <Emphasis Type="Italic">Salmo salar</Emphasis>

Effects of temperature and level of illumination on cardiac and respiratory rhythms have been analyzed in the juvenile Atlantic salmon Salmo salar L. at different stages of ontogenesis. The study was carried out under field conditions, which allowed avoiding to a considerable degree the effect of artifacts due to chemical and physical properties of water and to fish transposition. The “handlinglp”-reaction of the fry cardiac and respiratory activities was examined. A statistically significant decrease of the respiration rate (RR) by the 30th min and of the heart rate (HR) by the 15th min of experiment was revealed. A more expressed reaction of the decrease of cardiac and respiratory activities after transfer to the working chamber was found in juvenile salmons at early stages of their development. A linear dependence of frequency characteristics of electrocardiograms and electropneumograms (ECG and EPG) was shown in parrs and smolts on temperature in the range from 4 to 24°C. According to reaction of the respiratory system of smolts there was identified a signaling significance of the illumination level in the range of 10–20 kLx, which is confirmed by data on fish behavior in nature. Study of the changes in EEG and EPG allowed suggesting the presence of correlation of frequency characteristics of cardiac and respiratory activities of juvenile salmons with the metabolic level. However, moments of fundamental organism rearrangement in smolts are to be identified, which complicates the use of RR and HR as the measure of the metabolic level at this stage of the Atlantic salmon development.__________Translated from Zhurnal Evolyutsionnoi Biokhimii i Fiziologii, Vol. 41, No. 2, 2005, pp. 139–145.Original Russian Text Copyright © 2005 by Bakhmet.     

Polymorphism of smolts of pink salmon <Emphasis Type="Italic">Oncorhynchs gorbuscha</Emphasis> in the Indera River (Kola Peninsula)

Pink salmon introduced into the White Sea started to exploit as spawning grounds middle and upper reaches of the river 20 years after its appearance in the Indera River. As a result of this, the migration pathway of smolts and late smolts appeared in addition to early smolts. The intraspecies polymorphism of smolts is confirmed by differences of early and late smolts by body length and weight, migration dates, food spectrum, and indices of stomach fullness. The food spectra of late juveniles of pink salmon coincide with those of parr of Atlantic salmon Salmo salar and of brown trout S. trutta. Greater abundance of late migrants of pink salmon may cause competition of these species for food.     

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.     

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.     

Periodicity in Atlantic salmon Salmo salar L. smolt migration

Behavioural and physiological mechanisms postulated for the control of downstream migration of Atlantic salmon smolts are reviewed briefly, and some new evidence is presented for their refusal to undergo sustained swimming. Although these mechanisms imply passive displacement as the primary means of emigration, it is likely that active components must also exist as the rates of travel of smolts through loch systems are only slightly slower than those recorded for river systems. The timing of these movements within 24 h periods is reviewed and it is shown that the predominantly nocturnal emigration pattern is evident on occasions in alevin, fry and parr stages also. Thus at migration the diel periodicity probably represents a seasonal locomotor rhythm which, under changed behavioural and physiological circumstances, results in downstream displacement.     

Diel movements of out-migrating Chinook salmon (Oncorhynchus tshawytscha) and steelhead trout (Oncorhynchus mykiss) smolts in the Sacramento/San Joaquin watershed

We used ultrasonic telemetry to describe the movement patterns of late-fall run Chinook salmon (Oncorhynchus tshawytscha) and steelhead trout (O. mykiss) smolts during their entire emigration down California’s Sacramento River, through the San Francisco Bay Estuary and into the Pacific Ocean. Yearling hatchery smolts were tagged via intracoelomic surgical implantation with coded ultrasonic tags. They were then released at four upriver locations in the Sacramento River during the winters of 2007 through 2010. Late-fall run Chinook salmon smolts exhibited a nocturnal pattern of migration after release in the upper river. This is likely because individuals remain within a confined area during the day, while they become active at night and migrate downstream. The ratio between night and day detections of Chinook salmon smolts decreased with distance traveled downriver. There was a significant preference for nocturnal migration in every reach of the river except the Estuary. In contrast, steelhead smolts, which reside upriver longer following release, exhibited a less pronounced diel pattern during their entire migration. In the middle river, Delta, and Estuary, steelhead exhibited a significant preference for daytime travel. In the ocean Chinook salmon preferred to travel at night, yet steelhead were detected on the monitors equally during the night and day. These data show that closely related Oncorhynchus species, with the same ontogenetic pattern of out-migrating as yearlings, vary in migration tactic.     

Behavior of early life intervals of Klamath River green sturgeon, <Emphasis Type="Italic">Acipenser</Emphasis><Emphasis Type="Italic">medirostris</Emphasis>, with a note on body color

We studied ontogenetic behavior, migration, and wintering behavior of young Klamath River green sturgeon, Acipenser medirostris, in the laboratory to provide insight into likely behavior of wild sturgeon. Hatchling free embryos preferred cover but were poor swimmers and could not move farther than a few centimeters to cover. The poor swimming ability and cover preference of hatchlings suggests evolution for habitat selection of females to place eggs in habitat with cover for eggs (and hatchlings), and for egg characteristics (large, dense, and weakly adhesive) to cause rapid sinking into cover without drifting. A day or so after fish developed into larvae (first life interval feeding exogenously), day-12 larvae initiated a 12-day downstream nocturnal migration. A totally nocturnal migration is unlike other Acipenser migrants yet studied. Migrant larvae had a dark-colored body typical of other Acipenser species that migrate as larvae. Tail color was a dark black (black-tail phenotype) only during the early larva period, suggesting a morphological adaptation for migration, foraging, or both. Post-migrant larvae and early juveniles to day 84 foraged diurnally with a nocturnal activity peak. Day 110–181 juveniles moved downstream at night until water temperature decreased to about 8°C, indicating wild juveniles migrate downstream to wintering habitat. Habitat preference of month 9–10 wintering juveniles suggests wild juveniles are in deep pools with low light and some rock structure. Wintering juveniles were only active at night. Initiation and cessation of daily activity was at dusk and dawn during illumination changes of <1.0lx. This sensitivity to illumination has not been found before in sturgeons. During the first 10months of life, nocturnal activity of early life intervals is a dominant feature of migration, foraging, and wintering.     

Population structure of the masu salmon <Emphasis Type="Italic">Oncorhynchus masou</Emphasis> from the Kol River (Western Kamchatka) and geographic variation in the species area

According to materials from original investigations in Kamchatka as well as published data, the population structure and geographical variation of the masu salmon Oncorhynchus masou are considered in the species area. The intraspecies structure, dates of their run to rivers, dates of spawning, traits of spawning, size-age and sex composition, fecundity, biological traits of parr, downstream migration of smolts are investigated. The relationship is shown between the manifestations of geographical variation of masu salmon and the factor of temperature. It is assumed that the global warming of climate would contribute to the advancement of this species to northern areas and to the increase in its abundance.     

Basin‐scale phenology and effects of climate variability on global timing of initial seaward migration of Atlantic salmon (Salmo salar)

Migrations between different habitats are key events in the lives of many organisms. Such movements involve annually recurring travel over long distances usually triggered by seasonal changes in the environment. Often, the migration is associated with travel to or from reproduction areas to regions of growth. Young anadromous Atlantic salmon (Salmo salar) emigrate from freshwater nursery areas during spring and early summer to feed and grow in the North Atlantic Ocean. The transition from the freshwater (‘parr’) stage to the migratory stage where they descend streams and enter salt water (‘smolt’) is characterized by morphological, physiological and behavioural changes where the timing of this parr‐smolt transition is cued by photoperiod and water temperature. Environmental conditions in the freshwater habitat control the downstream migration and contribute to within‐ and among‐river variation in migratory timing. Moreover, the timing of the freshwater emigration has likely evolved to meet environmental conditions in the ocean as these affect growth and survival of the post‐smolts. Using generalized additive mixed‐effects modelling, we analysed spatio‐temporal variations in the dates of downstream smolt migration in 67 rivers throughout the North Atlantic during the last five decades and found that migrations were earlier in populations in the east than the west. After accounting for this spatial effect, the initiation of the downstream migration among rivers was positively associated with freshwater temperatures, up to about 10 °C and levelling off at higher values, and with sea‐surface temperatures. Earlier migration occurred when river discharge levels were low but increasing. On average, the initiation of the smolt seaward migration has occurred 2.5 days earlier per decade throughout the basin of the North Atlantic. This shift in phenology matches changes in air, river, and ocean temperatures, suggesting that Atlantic salmon emigration is responding to the current global climate changes.     

Growth,survival, and tag retention of steelhead trout (<Emphasis Type="Italic">O. mykiss</Emphasis>) surgically implanted with dummy acoustic tags

Recent advances in micro-electronics make the study of the migration of even small marine animals (>12 cm) over many 1000s of kilometres a serious possibility. Important assumptions in long-term studies are that rates of tag loss caused by mortality or tag shedding are low, and that the tagging procedure does not have an unacceptable negative effect on the animal. This paper reports results from a study to examine the retention of relatively large (24 × 8 mm) surgically-implanted dummy acoustic tags over a 7-month period in steelhead pre-smolts (O. mykiss), and the effects of implantation on growth and survival. Although there was some influence on growth to week 12, survival was high for animals > 13 cm FL. In the following 16-week period, growth of surgically implanted pre-smolts was the same as the control population and there was little tag loss from mortality or shedding. Currently available acoustic tags can be implanted in salmonid fish ≥12 cm FL, although combined losses from mortality and tag shedding were 33–40% for animals in the 12 and 13 cm FL size classes. By 14 cm FL, combined rates of tag loss (mortality plus tag shedding) for surgically implanted tags dropped to <15% and growth following surgery was close to that of the controls. Our results suggest that studies of ocean migration and survival over periods of many months are now feasible even for animals as small as salmon smolts. Surgically implanted salmon smolts are therefore good candidates for freshwater and coastal ocean-tracking studies on relatively long time scales (months). On such time scales, even relatively small salmon smolts may move thousands of kilometers in the ocean.     

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.