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.     

The effects of environmental factors on the migratory movement patterns of Sacramento River yearling late-fall run Chinook salmon (Oncorhynchus tshawytscha)

Understanding smolt migration dynamics is a critical step in the preservation and conservation of imperiled salmonids in California’s Sacramento River system. Late-fall run Chinook salmon yearling smolts were acoustically tagged and tracked during their outmigration through California’s Sacramento River and San Francisco Estuary during 2007–2009. Migration rates were 14.3 km·day^-1 (± 1.3 S.E.) to 23.5 km?day^-1 (± 3.6 S.E.), similar to rates published for other West Coast yearling Chinook salmon smolt emigrations. Region-specific movement rates were fastest through the upper river regions, and slowest in the Sacramento/San Joaquin River Delta. River travel times were recorded for smolts travelling through a series of ten monitor-delimited reaches. Using these, a smolt travel time model determined by two parameters (movement rate and rate of population spreading) was then used to determine the influence of different factors on the model’s fit, using model selection with Akaike’s Information Criterion. The model that allowed for both year and reach to be expressed additively for both travel time and population spreading rate estimates, while accounting for a “release” effect, was the best supported model. Finally, several models incorporated environmental data as a linear predictor of movement rates. The addition of the environmental variables, in order of importance, river width to depth ratio, river flow, water turbidity, river flow to mean river flow ratio, and water velocity all resulted in improved model fit. Water temperature did not improve model fit. These environmental associations are discussed and potential improvements on the travel time model are suggested.     

Synchrony in the downstream migration of smolts and upstream migration of adult Atlantic salmon in the subarctic River Utsjoki

Seasonal and diel migration timing of wild Atlantic salmon Salmo salar smolts and adults were investigated annually (2001–2004) in the subarctic River Utsjoki, a tributary of the large River Teno (70° N), using underwater video monitoring. Submerged video cameras provided information on the exact timing and intensity of both migrations in a natural river channel, without disturbing the fish. In contrast to the mainly nocturnal migration pattern described from temperate rivers, the River Utsjoki smolts and adults migrated throughout the day. Smolts migrated most intensively during hours of rising (0300–0900 hours) and high sun (0900–1500 hours), while adults favoured the period of low sun (2100–0300 hours). Smolt migrations started in June and lasted on average 42 days. Adults usually ascended the site 2–3 weeks before the first descending smolts were observed and the adult migrations extended to the end of August. Seasonal synchrony was observed between smolt and adult migrations in years of slowly warming water, whereas in a year of exceptionally warm early summer (2002), smolts migrated earlier than adults. Thus, water temperature seemed to be an important environmental factor triggering the smolt migration, while the migration of adults was probably more fixed to a certain season. Weak positive correlations between fish counts and water temperature were observed, indicating that increasing water temperature may have promoted both smolt and adult migrations. The influence of discharge was negligible, although increasing discharge late in the season may have activated the remaining individuals in both groups.     

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.     

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.     

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.     

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.     

Migratory timing, marine survival and growth of anadromous brown trout Salmo trutta in the River Imsa, Norway

The aim of the paper was to study sea migration, growth and survival of brown trout Salmo trutta of the River Imsa, 1976–2005. The migratory S. trutta were individually tagged and fish leaving or entering the river were monitored daily in traps located near the river mouth. The mean annual duration of the sea sojourn was 6–9 months for first-time migrants moving to sea between January and June. It was 8–18 months for those migrating to sea between July and December. Veteran migrants stayed 12 months or less at sea and most returned to the river in August. Early ascending fish stayed the longest in fresh water because most returned to sea in April to May. The day number of 50% cumulative smolt descent correlated negatively with mean water temperature in February to March and the February North Atlantic Oscillation index (NAOI). Mean annual sea growth during the first 2 years after smolting was higher for S. trutta spending the winter at sea than those wintering in the River Imsa. First year's sea growth was lower for S. trutta descending in spring than autumn. For first-time migrants, it correlated negatively with the February NAOI of the smolt year. Sea survival was higher for spring than autumn descending first-time migratory S. trutta with a maximum in May (14·9%). Number of anadromous S. trutta returning to the river increased linearly with the size of the cohort moving to sea, with no evidence of density-dependent sea mortality. Sea survival of S. trutta smolts moving to sea between January and June correlated positively both with the annual number of Atlantic Salmo salar smolts, the specific growth rate at sea, and time of seaward migration in spring. This is the first study indicating how environmental factors at the time of seaward migration influence the sea survival of S. trutta .     

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.     

Migration of hatchery reared juvenile Atlantic salmon, Salmo salar L., smolts down a release ladder. 1. Environmental effects on migratory activity

The effects of light intensity, water temperature and river spate conditions on the rate of migration of hatchery-reared Atlantic salmon smolts down a release ladder were examined. Low light intensity and high day time water temperatures raised smolt migration rates: water temperature had littleor no effect at night. Thediel patternofdown-ladder movement wasdetermined by these two environmental stimuli. Smolts demonstrated a threshold response to both light and water temperature: a fall in light intensity, or increase in water temperature, below or above their respective thresholds elicited no further response. As fish progressed through the ladder their movement became increasingly nocturnal, and most entry into the estuary occurred at night. Heavy rainhll caused rapid migration of smolts through the ladder. As ladder discharge remained constant, smolts must have been responding to some other change in the water conditions. Smolts responded to spate conditions irrespective of water temperature and light intensity, and they responded to light intensity irrespective of water temperature. A hierarchy of environmental cues, responsible for triggering migratory behaviour on a day to day basis during the smolt run, of spate> light intensity > water temperature is therefore suggested.     

Riverine, estuarine and marine migratory behaviour and physiology of wild and hatchery-reared coho salmon Oncorhynchus kisutch (Walbaum) smolts descending the Campbell River, BC, Canada

Eighty coho salmon Oncorhynchus kisutch smolts (40 wild and 40 hatchery-reared) were surgically implanted with acoustic transmitters and released into the Quinsam River over 2 days. Differences in physiology, travel time and migratory behaviour were examined between wild and hatchery-reared fish. In addition, tagged and control fish of both wild and hatchery-reared stock were raised for 3 months following surgery to compare survival and tag retention. Detection ranges of the acoustic receivers were tested in the river, estuary and ocean in a variety of flow conditions and tide levels. Receivers were placed in the river, estuary and up to 50 km north and south from the river mouth in the marine environment. Wild smolts were significantly smaller by mass, fork length and condition factor than hatchery-reared smolts and exhibited significantly higher levels of sodium, potassium and chloride in their blood plasma than hatchery-reared smolts. The gill Na⁺K⁺-ATPase activity was also significantly higher in the wild coho smolts at the time of release. Ninety-eight per cent of wild and 80% of hatchery-reared fish survived to the estuary, 8 km downstream of the release site. No difference was found in migration speed, timing or survival between smolts released during daylight and those released after dark. Wild smolts, however, spent less time in the river and estuary, and as a result entered the ocean earlier than hatchery-reared smolts. Average marine swimming speeds for wild smolts were double those of their hatchery-reared counterparts. While hatchery smolts dispersed in both a northward and southward direction upon entering the marine environment, the majority of wild smolts travelled north from the Campbell River estuary. The wild coho salmon smolts were more physiologically fit and ready to enter sea water than the hatchery-reared smolts, and as a result had higher early survival rates and swimming speeds.     

Migration route selection of juvenile Chinook salmon at the Delta Cross Channel, and the role of water velocity and individual movement patterns

We examined movement tracks of ultrasonic-tagged juvenile Chinook salmon (Oncorhynchus tshawyscha) smolts at the juncture of two migratory pathways. This migratory juncture occurs where the Delta Cross Channel splits from the Sacramento River in California’s Sacramento–San Joaquin Delta. Smolt tracks were analyzed to compare the importance of river flow and individual parameters in migratory route selection. The two routes differ significantly in smolt survival probabilities (Perry et al. N Am J Fish Manag 30:142–156, 2010), thus a clearer understanding of the variables contributing to route selection will be valuable for management of this declining species. A comparison of the two migratory groups showed that fish remaining within the Sacramento River: 1) Encountered the migratory juncture when river water velocities were much higher than those in the Delta Cross Channel (p?<?0.0001), 2) showed more direct swimming paths (p?=?0.03) and 3) migrated at higher speeds (p?=?0.04). Logistic regression models showed that the ratio of mean water velocity between the two routes was a much stronger predictor of ultimate route selection than any other variable tested. However, parameters for both the lateral position of smolts within the river and smolt size added predictive power to the final model. Our results suggest that river flow remains the most important variable for predicting smolt migration route, but note that knowledge of individual smolt attributes and movement patterns can increase our predictive ability.     

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

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

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.     

Temporal characteristics of downstream migration of smolts of the European river lamprey <Emphasis Type="Italic">Lampetra fluviatilis</Emphasis> in the Chernaya River

Data on the seasonal and diurnal dynamics of downstream migration of smolts of the European river lamprey Lampetra fluviatilis are obtained. On the basis of published data and surveys in nature, it is shown that in the largest part of the area the downstream migration of smolts occurs mainly in spring, in the period of high water in rivers caused by snow melting. The smolts undertake downstream migration at night, under minimal illumination. It has been determined experimentally that the smolts are characterized by negative photopreferendum (>90% of smolts preferred illumination <0.4 lx).     

Effects of downstream migration on myosin heavy chain expression and dihydropyridine receptor density in farmed smolt of Atlantic salmon

The expression of myosin heavy chain (MHC) subunits and dihydropyridine receptors (DHPRs) from red and white tail muscles of cultured smolts of Atlantic salmon Salmo salar was analysed from samples taken: (1) before the fish were transferred to the river and (2) after the migration distance of c . 50 km. The relative work load of migrating fish, estimated on the basis of swimming speed of Atlantic salmon and discharge rate of the River Simojoki, Finland, was maximally 1700 times higher than that of fish in culture. The data show that after five migration days the density of DHPRs in the muscles of the smolts was increased already. Furthermore, the results indicated a transition from the fast‐to‐slow MHC isoform. Transfer of fish to a natural environment and downstream migration thus had a significant effect on the expression of muscle proteins.     

Influence of sea temperature and initial marine feeding on survival of Atlantic salmon Salmo salar post-smolts from the Rivers Orkla and Hals, Norway

The abundance of returning adult Atlantic salmon Salmo salar, in the River Orkla in mid‐norway (1 sea‐winter, SW, fish) and River Hals in north Norway (1–3 SW fish), was tested against the early marine feeding and the seawater temperature experienced by their corresponding year classes of post‐smolts immediately after entry into the Trondheimsfjord (Orkla smolts, 22 years of data) and Altafjord (Hals smolts, 17 years of data). In both river–fjord systems, there was a significant positive correlation between the abundance of returning S. salar and the mean seawater temperature at the time of smolts descending to the sea. The number of 1SW fish reported caught in River Orkla was positively correlated to the proportion of fish larvae in the post‐smolt stomachs in Trondheimsfjord. The abundance of returning S.salar was, however, neither correlated to forage ratio (R_F) nor other prey groups in post‐smolt stomachs in the two fjord systems. In the Altafjord, the post‐smolts fed mainly on pelagic fish larva (70–98%) and had a stable R_F (0·009–0·023) over the 6 years analysed. In the Trondheimsfjord, however, there was a higher variation in R_F (0·003–0·036), and pelagic fish larvae were dominant prey in only two (50 and 91%) of the 8 years analysed. These 2 years also showed the highest return rates of S. salar in River Orkla. These results demonstrate that the thermal conditions experienced by post‐smolts during their early sea migration may be crucial for the subsequent return rate of adults after 1–3 years at sea. Pelagic marine fish larvae seem to be the preferred initial prey for S. salar post‐smolts. As the annual variation in abundance of fish larvae is related to seawater temperature, it is proposed that seawater temperature at sea entry and the subsequent abundance of returning adult S. salar may be indirectly linked through variation in annual availability of pelagic fish larvae or other suitable food items in the early post‐smolt phase.     

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.     

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.     

Migration of farmed adult Atlantic salmon with and without olfactory sense, released on the Norwegian coast

The dispersal and migration of farmed Atlantic salmon, Salmo salar , allowed to escape during the summer was studied. Three groups of 4–year–old fish of the River Imsa stock were released in coastal waters off south-western Norway: one group, with functional olfactory organs, was released at a fish farm 4 km away from the R. Imsa; two other groups, one with transected olfactory nerves and the other with functional olfactory organs, were released in the sea 90 km from the R. Imsa. To compare them with the migration pattern of reared, large smolts of the Imsa stock, a group of 3 + smolts was released in the R. Imsa.
Adults of salmon released as 3–year–old smolts homed with high precision to the R. Imsa. Four– year–olds released in the sea were recaptured in the fjord and in the coastal current, the majority north of the places of release. Immatures migrated to feeding areas in the North Atlantic. Matures seemed to enter rivers at random when ready to spawn. There was no difference in migration pattern between anosmics and controls. The olfactory sense was not mandatory for entering fresh water. The results indicate that the homing behaviour of Atlantic salmon is not a direct consequence of a single imprinting of the smolts, and that there is not a direct genetic link for return to a particular river. The present results support the sequential imprinting hypothesis proposed by Harden Jones (1968).