Physiological preparedness and performance of Atlantic salmon Salmo salar smolts in relation to behavioural salinity preferences and thresholds

This study investigated the relationships between behavioural responses of Atlantic salmon Salmo salar smolts to saltwater (SW) exposure and physiological characteristics of smolts in laboratory experiments. It concurrently described the behaviour of acoustically tagged smolts with respect to SW and tidal cycles during estuary migration. Salmo salar smolts increased their use of SW relative to fresh water (FW) from April to June in laboratory experiments. Mean preference for SW never exceeded 50% of time in any group. Preference for SW increased throughout the course of smolt development. Maximum continuous time spent in SW was positively related to gill Na⁺, K⁺‐ATPase (NKA) activity and osmoregulatory performance in full‐strength SW (measured as change in gill NKA activity and plasma osmolality). Smolts decreased depth upon reaching areas of the Penobscot Estuary where SW was present, and all fish became more surface oriented during passage from head of tide to the ocean. Acoustically tagged, migrating smolts with low gill NKA activity moved faster in FW reaches of the estuary than those with higher gill NKA activity. There was no difference in movement rate through SW reaches of the estuary based on gill NKA activity. Migrating fish moved with tidal flow during the passage of the lower estuary based on the observed patterns in both vertical and horizontal movements. The results indicate that smolts select low‐salinity water during estuary migration and use tidal currents to minimize energetic investment in seaward migration. Seasonal changes in osmoregulatory ability highlight the importance of the timing of stocking and estuary arrival.     

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.     

Olfactory clues play a critical role in the estuarine migration of silver-phase American eels

Estuarine migration of anosmic and control silver-phase American eels was examined during the fall spawning migration. Ultrasonic telemetry was used to track seventeen control and eight anosmic silver eels through 32 km of the Penobscot Estuary, Maine, U.S.A.. Twelve of seventeen control eels migrated out of the estuary in 97 h (approximately 4 d) on average. Only two of eight anosmic eels migrated out of the estuary. On average these two anosmic eels migrated out of the estuary within 180 h (approximately 7.5 d) of release and the other six had not left within 9 d. Most control eels progressed rapidly to the mouth of the estuary within a few days. Anosmic eels spent more time in the estuary and demonstrated different behavior from control eels due to their lack of olfaction. Some control eels moved with the appropriate tide, the ebb tide for transport out of the estuary, within one tidal cycle of being released into tidal freshwater. However, anosmic eels either did not move with the appropriate tide or took significantly longer to do so. Olfaction was probably used for orientation by control eels sensing chemical clues (organic and inorganic), which change throughout the tidal cycle. Increased migration times and errors in orientation were likely related to the inability of anosmic eels to use selective tidal stream transport for movement out of the estuary. Chemical clues seem to be one of the most important environmental clues used to guide estuarine migration of silver eels. However, a hierarchy of sensory mechanisms and environmental clues are most likely used for estuarine orientation.     

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.     

Spatial variation in the environmental control of crab larval settlement in a micro-tidal austral estuary

Settlement of benthic marine invertebrates is determined by the interaction between physical factors and biological processes, in which the tide, wind, and predation can play key roles, especially for species that recruit within estuaries. This complexity promotes high variability in recruitment and limited predictability of the size of annual cohorts. This study describes the settlement patterns of megalopae of the commercially important crab Cancer edwardsii at three locations (one in the center and two at the mouth of the estuary) within the Valdivia River estuary (~39.9°S), over three consecutive years (2006–2008). At each location, 12 passive benthic collectors with a natural substratum were deployed for 48?h at 7-day intervals, over a lunar cycle. Half of the collectors were covered with mesh to exclude predators. The main findings were as follows: (1) circulation changes due to upwelling relaxation or onshore winds controlled crab settlement at sites within the mouth of the estuary, (2) at the internal estuarine site, settlement was dominated by tidal effects, and (3) the effect of predation on settlement was negligible at all scales. The results show that the predominant physical factor controlling the return of competent crab larvae to estuarine environments varies spatially within the estuary. The lack of tidal influence on settlement at the mouth of the estuary can be explained by the overwhelming influence of the intense upwelling fronts and the micro-tidal regime in the study area.     

Predation on hatchery-reared and wild smolts of Atlantic salmon, Salmo salar L., in the estuary of River Orkla, Norway

Predation on wild and hatchery-reared Atlantic salmon smolts was studied in the estuary of River Orkla. Cod and saithe congregating in the estuary were the most serious predators on smolts. There was no difference between the mortality rates of wild and hatchery-reared smolts. Predation by cod was estimated at 20%. No evidence was found to indicate selective predation on the smallest wild and hatchery-reared smolts.     

Migration of hatchery reared juvenile Atlantic salmon, Salmo salar L., smolts down a release ladder. 3. Reactions on exiting the ladder

Twenty-eight hatchery-reared Atlantic salmon ( Salmo salar L.) smolts were tracked as they left a release ladder and moved through the estuary of the Lussa River and out to sea. Smolt passage through the estuary was fastest at high water and slowest at low water; the reverse of the situation observed for passive objects floating in the surface water layer. At low tide the water channel was effectively a freshwater stream and the smolts behaved in it accordingly, attempting to maintain station. At high water smolts moved downstream with the current close to the water surface. The degree of variation in individual speed through the estuary was the main difference observed between immature S2, stripped mature male S2, unstripped mature male S2 and S1 smolts. Smolts moved directly out of the estuary with no apparent check in their progress. Sea contact times in the vicinity of the river mouth, for both passive objects and smolts, were longest around high water and shortest around low tide. In the sea no differences were observed in the behaviour of the four different smolt types. While passive objects showed no tendency to move off in a particular direction, smolts showed a significant inclination to head off on a southerly bearing.     

Migratory behaviour of post-smolt Atlantic salmon during initial stages of seaward migration

The movements of 24 hatchery-reared Atlantic salmon Salmo salar smolts, with miniature acoustic transmitters (pingers) implanted surgically, were determined after release in the coastal waters of Passamaquoddy Bay (mean tide range 6 m), New Brunswick, Canada, to describe the first stages of seaward migration. Automated pinger detection at fixed sites, and pinger location and tracking by boat were used. Post-smolts left the release area rapidly, and the majority moved to open waters of the bay within several tidal cycles. Initially, post-smolts moved with a seaward orientation on ebb tides and held positions on flood tides. Their movements into open waters were diurnal, and the timing corresponded with the state of the tide during which they moved through a narrow channel. Post-smolts moved preferentially through this passageway with the aid of the tidal stream. Successful movements out through the channel occurred during ebb tides and any movements back in were during flood tides. Ground speed of fish moving through the channel was 4·2 body lengths s⁻¹ and faster than the tidal stream velocities in the channel. The relative velocity of fish swimming through the channel was 2 body lengths s⁻¹. Post-smolt movement was indicative of active, directed swimming with a reliance on ebb-tide transport for migration through a coastal area with strong tidal currents. Some post-smolts moved seaward directly with no apparent period of acclimation for the transfer to the marine environment, whereas others delayed their departure. These differences in behaviour were probably related to asynchrony in smolting when fish were released.     

The seaward movement of Atlantic salmon smolts in the Usk estuary, Wales, as inferred from power station catches

The greatest numbers of Atlantic salmon smolts were caught at Uskmouth power station during the day on the flood tide and the least on an ebbing night tide. Catch rates during the day on an ebb tide and at night on a flood tide represented an intermediate state. The results, together with tracking data, suggest that salmon smolts prefer to migrate seaward through the lower Usk estuary during the night Oncc an ebbing tide.     

Survival and behavior of juvenile steelhead trout (Oncorhynchus mykiss) in two estuaries in Oregon, USA

Anadromous salmonids are viewed as a prized commodity and cultural symbol throughout the Pacific coast of North America. Unfortunately, several native salmonid populations are threatened or at risk of extinction. Despite this, little is known about the behavior and survival of these fish as the juveniles transition from freshwater to the ocean. Our primary objectives were to estimate survival of juvenile steelhead migrating between trapping sites and the ocean and evaluate whether survival in the estuary varies temporally (within a year) or spatially (within and between estuaries) within the same distinct population segment. We also evaluated whether flow or fork length were correlated with survival and collected information on variables that have been demonstrated to affect smolt survival in other studies to lend insight regarding differences in survival estimates between basins. We compared run timing, migration rate, survival, condition factor, age composition and time of residence in the estuary for steelhead outmigrants from each basin and measured parasite loads in outmigrating steelhead to evaluate potential differences in parasite density and parasite community between basins. In 2009, we implanted acoustic transmitters in 139 wild steelhead smolts in two small rivers on the Oregon Coast. In general, only 40–50 % of the wild steelhead smolts tagged at upstream smolt traps were detected entering the ocean. The majority of mortality occurred in the lower estuary near the ocean. Wild steelhead smolts typically spent less than 1 day in the estuary in both basins. Using similar data from previous studies in the Nehalem and Alsea basins, we showed that survival appears to be negatively correlated with flow in most releases, and in 2009 fork length was not correlated with survival. Our observations provide baseline information on factors that could influence smolt survival through the estuary as well as smolt to adult survival in these basins, and emphasize the importance of monitoring smolt survival in the estuary.     

Rhythmic aspects of estuarine migration of hatchery-reared Atlantic salmon (Salmo salar) smolts

Seasonal, diel and tidal rhythmic activity of hatchery-reared Atlantic salmon ( Salmo salar ) smolts migrating through a large estuary was studied by ultrasonic tracking of 46 individuals during two seasons. Prior to 10 May each year most smolts were inactive and remained near shore in shallow water. After 10 May nearly all smolts moved away from the release point into swift water and made rapid seaward progress. Initiation of migration each year occurred when river and hatchery water temperatures rose above 9°C. Migration in the estuary was largely passive drift, and as a result there were distinct tidal rhythms of ground ('swimming') speed and net seaward travel. There were no diel rhythms in ground speed or net seaward travel; smolts drifted seaward on the tides during both day and night. Smolts may be slightly deeper during day than night.     

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.     

The turbidity maximum in the Tamar estuary

Many estuaries of medium to high tidal range exhibit an accumulation of fine cohesive material in their upper reaches in the region of the limit of saline intrusion. Much, or all, of this material is suspended each tidal cycle and the entire region undergoes a seasonal variation which appears to depend on fluvial input. Two factors which are throught to influence the formation and maintenance of turbidity maxima are the differing magnitudes of the bed shear stress (τ₀) on flood and ebb tides and the large vertical density gradient which developes on the ebb tide. Crucial to the importance of the first factor is that τ₀ exceeds a critical value, at which erosion occurs, for a greater period on the flood than on the ebb. The effect of the density gradient is that upward propagation of bed generated eddies is inhibited and the sediment is not transported into the upper part of the flow where it will be most effectively transported. It is not clear which, if either, of these mechanisms is dominant. Data consisting of vertical profiles of velocity, salinity and suspended solids were collected at four stations in the Tamar estuary during a high range tidal cycle. One station, at which the depth mean salinity (S _d ) varied from 0.0 to ∼ 12.0‰, was occupied permanently. The other stations were occupied such that data were collected asS _d varied in the range 0.0 to ∼ 4.0‰. In this way each station was occupied for a period of time on the ebb and flood tide. Observations show that during the early ebb, when the flow is relatively deep and slow, stratification persist untilS _d ∼ 0.0‰ and that no significant transport occurs while the flow is saline but that there is a rapid increase in suspended solids concentrations after this time. During the later ebb the shallower faster flow allowed the density gradient to be erode and significant transport was observed atS _d ∼ 5.0‰. On the flood tide the flow in the low salinity region is well-mixed troughout. Computation of the fluxes and total transport per unit breath of estuary show that on the ebb tide the quantity of solid material being transported by the low salinity (0–3‰) region remains nearly constant as this region of the flow is advebted seaward. On the flood tide, however, as the same region is advected landward the quantity of material being transported increases. It is concluded that in the Tamar estuary the early ebb tide stratification contributed to the formation and maintenance of a turbidity maximum which is strongly associated with the low salinity region of the flow. It is also speculated that the differences in the ebb and flood tide transport are caused by differences in the availability of mobile material on the bed at different stages of the tidal cycle.     

Estuarine survival and migratory behaviour of Atlantic salmon Salmo salar smolts

To estimate mortality rates, assess the spatio‐temporal dynamics of natural mortality and examine migratory behaviour during the fresh to saltwater transition, 185 wild Atlantic salmon Salmo salar smolts were implanted with coded acoustic transmitters. Seaward migration of tagged S. salar from four river systems in an area of Nova Scotia, Canada known as the Southern Upland was monitored using fixed receivers and active telemetry over 3 years. Cumulative survival through the river, inner estuary, outer estuary and bay habitats averaged 59·6% (range = 39·4–73·5%). When standardized to distance travelled, survival rates followed two patterns: (1) constant rates of survival independent of habitat or (2) low survival most frequently associated with inner estuary habitats. In rivers where survival was independent of habitat, residency periods were also independent of habitat, post‐smolts exhibited few upstream movements, took a more direct route to the ocean and reached the ocean rapidly. Alternatively, in rivers where survival was habitat specific, residency was also habitat specific with overall increased residency, more frequent upstream movements and delayed arrival to the open ocean. The sudden disappearance of most (75–100%) smolts and post‐smolts assumed dead during the course of this study warrants further examination into the role of avian predators as a mortality vector.     

Migration of Atlantic salmon smolts and post-smolts in the Rivière Saint-Jean, QC north shore from riverine to marine ecosystems

The characteristics of the Atlantic salmon (Salmo salar) smolt run of the Rivière Saint-Jean, Quebec, Canada, in 2009 and 2010 were determined using acoustic telemetry. Tagged smolts were tracked from freshwater release sites for 17 km, then through the estuary and for their entry into the Gulf of St. Lawrence. The smolt migration began in both years at water temperatures of 10 °C, but lasted twice as long in the cooler year. The smolts in 2009 crossed the river to ocean boundary faster than the smolts of 2010, despite being similar in size. Smolt speed over ground increased from the river to the estuary to the marine environment. Smolts migrated both during the day and the night, but most movements began near or just after sunset, with increasingly nocturnal movements in the ocean. Smolts crossed the estuary during ebbing tides and moved faster during the night than the day. The tide cycle also influenced estuarine smolt travelling rates, but only secondarily to the diel cycle.     

Tidal and diel timing of river entry by adult Atlantic salmon returning to the Aberdeenshire Dee, Scotland

The timing of salmon migration from the estuary of the Aberdeenshire Dee into the river in relation to tidal phase and time of day was studied by combined acoustic and radio-tracking of individual fish and by analysing records of untagged fish from a resistivity fish counter 0.8 km upstream from the tidal limit. Up-estuary movements that led to river entry were predominantly nocturnal and tended to occur during the ebb tide. Penetration into the non-tidal reaches of the river also tended to occur at night, but the timing of salmon movements was no longer significantly associated with tidal phase. The tracking data suggested that the reduction in the strength of the association between salmon migration and tidal phase resulted from variability in rates of progress from the estuary to the river. This variability may have been random, or related to changes in migratory behaviour during entry to fresh water. A simulation of upstream progress by groups of salmon illustrated the decay of the relationship between salmon movements and tidal phase and the advance of the average tidal phase of observed salmon movements with increasing distance of the observation point from where movements were initiated. The magnitude of these effects depended on the average rate of upstream progress and variability in the rate of progress, being greatest when upstream progress was slow and variable. These results highlight a limitation of point observations of migration with regard to identifying environmental stimuli for migration and quantifying their effects.     

Temporal and spatial patterns in the abundance of wintering Red-breasted Mergansers Mergus serrator in an estuary

Tidal, temporal and spatial aspects in the abundance of wintering Red-breasted Mergansers in the Ythan estuary, northeast Scotland, were investigated. The number and location of birds were examined in relation to four variables: tidal cycle and tidal range, and month and time of day. A significant increase in the numbers of this piscivorous bird occurred as the winter progressed. Numbers were highest in early morning and then decreased significantly during the day. Spring tides were favoured over neap tides and a significant increase in numbers was recorded between low tide and high tide. The lower part of the estuary was preferred over the upper part with no detectable shift in the preference occurring in relation to any of the four variables.     

Movement of adult Atlantic salmon in the Usk estuary, Wales

A total of 56 salmon was tagged in the Usk estuary using combined acoustic and radio tags. Those fish migrating within the estuary oscillated with the tide over c . 10 km, being towards the seaward end at low water and moving upstream on the flood tide. Fish migrating through the estuary moved upstream on the flood tide and stemmed displacement downstream during the ebb. These findings, together with information on the hydrodynamics of the estuary, indicate that the fish utilize tidal currents to migrate passively in their preferred direction.     

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.     

The influence of tidal cycles and freshwater inflow on the distribution and movement of an estuarine resident fish Acanthopagrus butcheri

Movement patterns and habitat utilization by black bream Acanthopagrus butcheri (Sparidae), an estuarine resident species, were investigated using acoustic telemetry in a small estuary on the east coast of Tasmania, Australia. Thirty‐four adult A. butcheri were tracked for periods of up to 187 days between August 2005 and January 2006. Although able to tolerate a wide range of salinities, the fish spent most of the time within the upper and middle regions of the estuary, where brackish conditions dominated. The species exhibited extensive movements linked to tidal cycles, with small‐scale upstream movements during incoming tides and downstream movements during out going tides. The extent of these movements was positively correlated with the tidal height difference between consecutive tidal peaks and troughs. Freshwater inflows and resultant changes in salinity also significantly influenced distribution and movement patterns. Fish moved downstream during the periods of heavy inflows, returning upstream as salinities increased to c. >10. During the peak of spawning period (November to December) fish moved into the upper region of the estuary, where they aggregated to spawn. Periodic increases in freshwater discharge, however, resulted in fish leaving the spawning grounds and moving downstream. Towards the end of the spawning season (January), the fish became more dispersed throughout the entire estuarine system.