Behavioural transition during the estuarine migration of wild Atlantic salmon (Salmo salar L.) smolt

Abstract –  Ultrasonic telemetry and hydrodynamic modelling were used to study the migratory behaviour of 54 wild Atlantic salmon ( Salmo salar ) smolt captured in freshwater during their downstream migration and tracked in 2 years through a shallow estuary system. A high-density, fixed array of receivers provided detailed spatial and temporal resolution of behaviour in the second year of study. Smolt migration in the river occurred mostly at night and downstream migration was slower during the day. In the estuary, smolt moved seaward on ebbing tides and landward on flooding tides. The effect of current velocity was greater during the night than during the day. We documented for the first time that current velocity and diurnal period only accounted for approximately one-third of the variation in smolt ground speeds in the estuary, indicating that smolt movements were far less passive than previously reported. Smolt energetic status had no effect on smolt swimming behaviour or migratory performance. With an increase in salinity, smolt seaward movements during flooding tides were more frequent, and overall seaward ground velocity increased. The increase in salinity experienced by the smolt during their migration through the leading edge of saltwater intrusion thus induced a behavioural transition from a more passive, fluvial migration to a more active- and seaward-oriented migration.     

Fish larvae distribution off Mejillones Peninsula (northern Chile) during a coastal upwelling event in Spring 1999: interactions with the cold upwelling plume

We examined the interaction between vertical and horizontal distribution of fish larvae off Mejillones Peninsula (23°S), northern Chile, under conditions of active coastal upwelling. An oceanographic survey covered spatial variability in temperature, chlorophyll- a (chl a ), dissolved oxygen, salinity and water density. Fish larvae were sampled during daytime and nighttime periods through two consecutive days in four stations: two inside and two outside of a well-developed upwelling plume, and at three depth strata: 0–20, 20–80 and 80–200 m. Eighteen taxa were analysed, of which the Myctophidae Diogenychthys atlanticus , Diogenichthys laternatus , and the anchovy Engraulis ringens , were most abundant. Our data showed little evidence for diel vertical migration and larvae were more abundant at depth (>80 m) under low temperature (∼12°C) and low chl a (∼2 mg m^–3), below the highly advective upper layer. The exploratory K -means analysis allowed the separation of data into two distinct habitats: upwelling and nonupwelling types. Most taxa were allocated in nonupwelling waters, i.e. outside the cold plume. However, short-term variations (<24 h) in the position of the upwelling plume influenced both horizontal and vertical occurrence as well as abundance of taxa, and caused variability in temperature, oxygen and chl a . These changes in oceanographic conditions, caused by upwelling circulation and the dynamics of the cold plume, may sharply modify the habitat of fish larvae and have an important role in survivorship and recruitment success.     

Movements of two strains of radio tagged Altlantic salmon, Salmo salar L., smolts through a reservoir

Smolt migration through a shallow and turbid hydro-reservoir in a major Danish river system was investigated using radiotelemetry. Hatchery-reared 1+-year-old Atlantic salmon, Salmo salar L., smolts of equal size from two different non-native strains were radio-tagged and followed during their downstream migration through the 12-km-long reservoir. A total of 50 salmon smolts, 25 of Swedish (Øtran River) and 25 of Irish (Burrishoole River) origin, were surgically implanted with miniature radiotransmitters. The tagged smolts were tracked daily over a 3-week period in May 1996. The Øtran smolts initiated migration first ( P  < 0.001), moved faster ( P  < 0.01), were delayed less when passing a culvert ( P  < 0.001) and were more successful in moving through the reservoir than the Burrishoole smolts. The observed differences in migratory behaviour are interpreted as evidence of a genetic component influencing smolt migration.     

Differences in Lepeophtheirus salmonis abundance levels on Atlantic salmon farms in the Broughton Archipelago, British Columbia, Canada

Sea lice data collected from Atlantic salmon farms in the Broughton Archipelago between 2003 and 2005 were examined for inter-regional differences in mobile Lepeophtheirus salmonis (Krøyer) abundance using the generalized linear model procedure. Factors such as age of the salmon populations, location of farms and time of year had a significant effect on the abundance of the mobile stages of L. salmonis whereas water temperature and salinity did not. Separate evaluation of SLICE® treatment data found no significant difference in treatment frequency among the areas but did show that there were significantly lower numbers of farm treatments during the summer months when compared with other seasons. The role of environment and wild fish in influencing sea lice abundance on the farmed salmon is discussed. The findings suggest that effective management programmes for sea lice should not only be based on geographical location but should take into account other factors which could influence lice abundance levels.     

The effect of artificial light treatment and depth on the infestation of the sea louse Lepeophtheirus salmonis on Atlantic salmon (Salmo salar L.) culture

Two field studies were carried out with farmed Atlantic salmon (Salmo salarL.) in sea cages to examine various effects of artificial light (AL) and the vertical distribution of salmon on lice infestation.

The use of AL light caused an overall increase in lice infestation in both experiments. The first study showed that salmon held at 0–4 m depth in cages developed higher infestation than salmon held at greater depths (4–8 and 8–12 m) under both natural light (NL) and AL. In the second study, salmon maintained in 14-m deep sea cages that were exposed to AL with different light intensities. The AL treatments resulted directly in different diel and seasonal patterns of vertical distribution of the salmon and also different temporal patterns in lice infestations. So indirectly the infestation pattern appeared to be correlated with median day-time swimming depth of the salmon.     

Ocean distribution and habitat associations of yearling coho (Oncorhynchus kisutch) and Chinook (O. tshawytscha) salmon in the northern California Current

Yearling juvenile coho and Chinook salmon were sampled on 28 cruises in June and September 1981–85 and 1998–07 in continental shelf and oceanic waters off the Pacific Northwest. Oceanographic variables measured included temperature, salinity, water depth, and chlorophyll concentration (all cruises) and copepod biomass during the cruises from 1998–07. Juvenile salmonids were found almost exclusively in continental shelf waters, and showed a patchy distribution: half were collected in ～5% of the collections and none were collected in ～40% of the collections. Variance‐to‐mean ratios of the catches were high, also indicating patchy spatial distributions for both species. The salmon were most abundant in the vicinity of the Columbia River and the Washington coast in June; by September, both were less abundant, although still found mainly off Washington. In June, the geographic center‐of‐mass of the distribution for each species was located off Grays Harbor, WA, near the northern end of our sampling grid, but in September, it shifted southward and inshore. Coho salmon ranged further offshore than Chinook salmon: in June, the average median depth where they were caught was 85.6 and 55.0 m, respectively, and in September it was 65.5 and 43.7 m, respectively. Abundances of both species were significantly correlated with water depth (negatively), chlorophyll (positively) and copepod biomass (positively). Abundances of yearling Chinook salmon, but not of yearling coho salmon, were correlated with temperature (negatively). We discuss the potential role of coastal upwelling, submarine canyons and krill in determining the spatial distributions of the salmon.     

Length and sex-specific associations between spiny dogfish (Squalus acanthias) and hydrographic variables in the Bay of Fundy and Scotian Shelf

The associations between spiny dogfish ( Squalus acanthias ) and hydrographic variables (temperature, salinity and depth) were examined in the Bay of Fundy and Scotian Shelf from 1970 to 1998. Data were obtained from standard groundfish bottom trawl surveys. Dogfish sex affected habitat associations. Males were found to occupy bottom water of significantly higher salinities and depths than that occupied by females. Length also significantly affected habitat associations. Smaller dogfish occupied relatively deep, high salinity bottom water compared with larger dogfish. Overall, the occupied temperatures, salinities and depths were significantly different from those which were available. Dogfish occupied warmer temperatures along a narrow range (6.62–9.19°C) compared with those which were available (1.57–9.35°C). Occupied salinity (32.70–34.43 ppt) and occupied depth (88.62–184.66 m) were also distributed along a narrower range than available salinity (32.16–34.79 ppt) and available depth (55.00–218.10 m). Sex-specific, length-specific and overall environmental preference by dogfish may bias traditional `offshore' groundfish surveys while large scale changes in hydrographic parameters may alter dogfish distribution and their interactions with other marine fauna.     

The host transfer ability of Lepeophtheirus salmonis (Copepoda: Caligidae) from farmed Atlantic salmon, Salmo salar L.

The ability of mobile stages of Lepeophtheirus salmonis to transfer between salmon, Salmo salar L., hosts was examined by two methods. The first of these was the removal of certain stages on farmed fish, while the second comprised the placement of mobile lice on clean fish in the laboratory. In one-sea-winter farmed salmon, 63% of males and 52% of female lice transferred to new hosts over a period of 4 days. In laboratory trials on post-smolts, 61% of males and 69% of female lice transferred to new hosts over 3 days. In another experiment, free-swimming adult male lice were added to the tank, and 90% of these were found on fish after 24 h. Immigration of lice to fish in trial cages and the large proportion of lice transferring between fish in both laboratory and farm trials indicates that L. salmonis can transfer between hosts.     

Environmental conditions and physiological state influence estuarine movements of homing sockeye salmon

The reproductive migration of anadromous salmonids through estuarine waters is one of the most challenging stages of their life cycle, yet little is known about the environmental and physiological conditions that influence migratory behaviour. We captured, sampled tissues, tagged and released 365 sockeye salmon (Oncorhynchus nerka) homing through inner coastal waters towards the Fraser River, British Columbia, Canada. Biotelemetry was used to assess the behaviour of individual sockeye salmon approaching estuarine waters and at river entry, which were related to both fish physiological condition at release and to prevailing environmental conditions. Sockeye salmon tended to stay close to the shore, migrated during the day, and movements were related to tide. Sockeye salmon migration rate was linked to wind‐induced currents, salinity and an individual's physiological state, but these factors were specific to location and stock. We propose that wind‐induced currents exposed sockeye salmon entering the estuary to stronger olfactory cues associated with Fraser River water, which in turn resulted in faster migration rates presumably due to either an increased ability for olfactory navigation and/or advanced reproductive schedule through a neuroendocrine response to olfactory cues. However, once the migration had progressed further into more concentrated freshwater of the river plume, sockeye salmon presumably used wind‐induced currents to aid in movements towards the river, which may be associated with energy conservation. Results from this study improve our biological understanding of the movements of Fraser River sockeye salmon and are also broadly relevant to other anadromous salmonids homing in marine environments.     

Spatial correlation patterns in coastal environmental variables and survival rates of salmon in the north-east Pacific Ocean

We examined spatial correlations for three coastal variables [upwelling index, sea surface temperature (SST), and sea surface salinity (SSS)] that might affect juvenile salmon ( Oncorhynchus spp.) during their early marine life. Observed correlation patterns in environmental variables were compared with those in survival rates of pink ( O. gorbuscha ), chum ( O. keta ), and sockeye ( O. nerka ) salmon stocks to help identify appropriate variables to include in models of salmon productivity. Both the upwelling index and coastal SST were characterized by strong positive correlations at short distances, which declined slowly with distance in the winter months, but much more rapidly in the summer. The SSS had much weaker and more variable correlations at all distances throughout the year. The distance at which stations were no longer correlated (spatial decorrelation scale) was largest for the upwelling index (> 1000 km), intermediate for SST (400–800 km in summer), and shortest for SSS (< 400 km). Survival rate indices of salmon showed moderate positive correlations among adjacent stocks that decreased to zero at larger distances. Spatial decorrelation scales ranged from approximately 500 km for sockeye salmon to approximately 1000 km for chum salmon. We conclude that variability in the coastal marine environment during summer, as well as variability in salmon survival rates, are dominated by regional scale variability of several hundred to 1000 km. The correlation scale for SST in the summer most closely matched the observed correlation scales for survival rates of salmon, suggesting that regional-scale variations in coastal SST can help explain the observed regional-scale covariation in survival rates among salmon stocks.     

Diel vertical migration of myctophid fishes (Family Myctophidae) in the transitional waters of the western North Pacific

The diel vertical migration patterns of adult myctophid fishes were determined in the transitional waters of the western North Pacific off Japan, using day–night sampling from 20 to 700 m depths with a commercial otter trawl in the summer of 1995. A total of 12 species belonging to 9 genera were collected. Four patterns were recognized in the diel vertical migration of 11 of the 12 species. (1) Migrants showing clear day–night habitat separation with peak abundance above 200 m at night: Symbolophorus californiensis, Tarletonbeania taylori, Notoscopelus japonicus, Diaphus theta, Ceratoscopelus warmingi, and Diaphus gigas. (2) Semi-migrants, in which part of the population often remains in the daytime habitat at night. The distribution depths of migratory and nonmigratory individuals do not overlap: Stenobrachius leucopsarus . (3) Passive-migrants, in which there is no separation of day–night habitats, but the upper limit of daytime distribution depth shifts to a shallower layer at night, probably as the fish follow migratory prey: Lampanyctus jordani . (4) Nonmigrants: Stenobrachius nannochir, Lampanyctus regalis (> 140 mm SL), and Protomyctophum thompsoni . The day–night habitat temperature ranges are also given for the 11 species. No remarkable east–west differences were seen in the vertical migration patterns compared with previous knowledge of eight of these species in the eastern Pacific. The diel migration patterns are newly described for three other species endemic to the western Pacific. The standing stock of myctophids in the study area was conservatively estimated at 18.5 ± 4.7 g m⁻² (avg. ± SD).     

Depth, temperature and salinity preferences of newly recruited penaeid shrimps in Kuwait waters

Associations between catches and depth, temperature and salinity data obtained from the 1994–97 preseason shrimp trawl surveys in Kuwait waters were analysed using cumulative distribution functions and overlap coefficients. No single environmental factor had a controlling influence on shrimp distribution. Shrimp associations with the joint distributions of two environmental variables were stronger and more consistent over time than those with any single environmental factor. All three shrimp species examined in this study were significantly associated with the joint distributions of depth and temperature, and depth and salinity, indicating depth as the most influential factor in structuring the shrimp distributions during the surveys. Metapenaeus affinis and Parapenaeopsis stylifera were distributed at similar ranges of depth (8–15 m), temperature (26–31°C) and salinity (35–41‰), and had very high overlap coefficients. These two species can be categorized as co-occurring species in the sampled area. Penaeus semisulcatus had the widest range of the three environmental variables investigated in this study, particularly depth (8–32 m) and temperature (24–32°C).     

How are the salmon lice (Lepeophtheirus salmonis Krøyer, 1837) in Atlantic salmon farming affected by different control efforts: A case study of an intensive production area with coordinated production cycles and changing delousing practices in 2013–2018

The aims of the present study were to describe the salmon lice (Lepeophtheirus salmonis Krøyer, 1837) situation in an intensive salmon production area in mid‐Norway and to consider implications of changing practices of how salmon lice infestation can be controlled. The results in this study suggest that there are steps that can be carried out to keep salmon lice under control even during years when the temperature facilitates a quick salmon lice development. The present work indicates that the use of cleaner fish can delay the time it takes adult female lice to reach 0.1 per salmon in the beginning of a production cycle. It suggests that the timing of cleaner fish deployment into salmon cages can influence its effectiveness in controlling salmon lice. It also gives caution to letting salmon lice develop unchecked, even at levels far below the current lice limit, because of the difficulties to control salmon lice when the external infection pressure is too high. This study took place during a rapid change in delousing methods, in an area with coordinated salmon production. Despite its exploratory nature, this study offers insights into the salmon lice fluctuations in relation to efforts aimed at controlling it.     

Distribution and structure of the population of sea lice, Lepeophtheirus salmonis Krøyer, on Atlantic salmon, Salmo salar L., under typical rearing conditions

Abstract. Samples of Atlantic salmon, Salmo salar L., held in sea cages and tanks with running water were examined for presence of sea lice, Lepeophtheirus salmonis. The number of lice, their sex stage (only post-chalimus stages) and location on the fish body were recorded. A new method has been used to express parasitic intensity in fish of different size. Effects of different factors on the degree and character of infestation (structure of the lice population), such as fish size, habitat and chemical treatment, have been studied. Different lice categories have shown different susceptibility to delousing with dichlorvos. Second pre-adults are most susceptible, and males generally more than females. Running water has been found to'flush out'the lice from the fish body. Pre-adults are flushed out easier than adults, and males generally easier than females. Adult males show high tolerance to water of lowered salinity and great mobility manifested in occasional transfer between hosts. The higher activity level of adult males may partly explain a drop in proportion of males after delousing and in running water. Frequency of mating has been found to increase with parasitic intensity.     

The effect of temperature and salinity on the settlement and survival of copepodids of Lepeophtheirus salmonis (Krøyer, 1837) on Atlantic salmon, Salmo salar L.

The effects of temperature and salinity on the settlement, subsequent survival and development of the copepodids of Lepeophtheirus salmonis on Atlantic salmon were investigated experimentally. There was a significantly greater settlement and survival of copepodids at 10 days post-infection (dpi) at 12 °C compared with at 7 °C at a constant salinity of 34‰. Development of L. salmonis was also more rapid at 12 °C. Settlement was significantly greater at a salinity of 34‰ than at 24‰. In one experiment, survival at 10 dpi was significantly greater at 34‰; however, a second experiment found that there was no significant difference between the two saline levels. This may have been because of a rise in water temperature for 2 dpi, which appears to have overridden the effect of low salinity. Development of L. salmonis was more rapid at 34‰. Copepodids settled on all of the external surfaces of the salmon, although the proportion on different surfaces varied between experiments. The gills, particularly at low temperatures, the body surface, and the pectoral and dorsal fins were especially favoured.     

Factors associated with severity of naturally occurring piscirickettsiosis in netpen- and tank-reared juvenile Atlantic salmon at a research aquarium in western Canada

The opportunistic examination of factors associated with an outbreak of piscirickettsiosis (SRS) is described in Atlantic salmon Salmo salar post-smolts held in an open netpen or in tanks supplied with raw sea water at a research aquarium in western Canada. During the outbreak, seawater temperature was significantly higher and salinity significantly lower in the netpen compared with the tanks. Mortality in the netpen began approximately 3 weeks prior to that in the tanks, and cumulative mortality in the netpen (34%) was significantly higher than in the tanks (12%). Piscirickettsia salmonis was confirmed by qPCR in tissues from moribund and dead fish and from colonies grown on enriched blood agar medium. Neither P. salmonis nor SRS were observed in salmon held concurrently in UV-irradiated sea water. The elevated mortality was curtailed by treatment with oxytetracycline. These observations further indicate warmer, less saline and periodically hypoxic seawater are risk factors for SRS. UV irradiation of sea water is shown to be a tool for SRS management in fish-holding facilities.     

Environmental preferences of longlining for yellowfin tuna (Thunnus albacares) in the tropical high seas of the Indian Ocean

A survey of yellowfin tuna, Thunnus albacares , fishing ground was carried out on board of the Chinese longliners from September 15 to December 12, 2005 in the tropical high seas of the Indian Ocean. The depth at which each yellowfin tuna was hooked was estimated using a stepwise regression analysis of theoretical hook depth and observed average hook depth measured using a temperature depth recorder. Water temperature, salinity, chlorophyll  a , dissolved oxygen and thermocline, which are important variables influencing yellowfin tuna habitats, were measured in the survey. Catch rates of yellowfin tuna were then analyzed with respect to depth, temperature, salinity, chlorophyll  a , dissolved oxygen and thermocline. We suggest that the optimum ranges of swimming depth, water temperature, chlorophyll  a and dissolved oxygen concentration for yellowfin tuna are 100.0–179.9 m, 15.0–17.9°C, 0.090–0.099  μ g L⁻¹, 2.50–2.99 mg L⁻¹, respectively; that salinity has less influence on the vertical distribution of adult yellowfin tuna; and that yellowfin tuna are mainly distributed within the thermocline in the high seas of the Indian Ocean. Our results match the yellowfin tuna's vulnerability to deep longline fishing gear well.     

Diel vertical distribution of postflexion larval Citharichthys spp. and Sebastes spp. off central California

The diel and spatial distributions of postflexion larval Citharichthys spp. and Sebastes spp. at 67 stations off central California were examined during March 1994 with a 10 m² MOCNESS during 24 h sampling operations. At each station, a CTD profiler with an attached PAR sensor was deployed to measure temperature, salinity, σ_t, dynamic height, and ambient light levels at depth. Catches of C. sordidus, C. stigmaeus, unidentified Sebastes spp., and S. jordani were larger at night than during the day and crepuscular periods, probably because of net avoidance during the day and crepuscular periods, and (for Citharichthys spp.) vertical migration into the upper water column at night. C. sordidus and C. stigmaeus migrated vertically through the pycnocline on a diurnal cycle, while Sebastes spp. apparently remained in the mixed layer above the pycnocline at all times. No definitive patterns were observed between horizontal contours of temperature, salinity, and σ_t at depth and catches of any larvae. This was attributed to the relatively homogeneous spatial pattern in the water column above the pycnocline, where the majority of the fish occurred.     

Migration motivation of cultured Atlantic salmon,Salmo salar L., smolts in relation to size,time of release and acclimatization period

Within the framework of a national sea-ranching programme in Norway a field study of marine release of smolts was conducted in Selsto Bay, west Norway. In March/early April 42 000 cultured Atlantic salmon, Salmo salar L., 10–20cm in body length were divided into two size groups consisting of fish smaller and larger than approximately 14–15 cm and reared in eight tanks, four of each size category. The eight groups were transferred to net pens located in a small marine bay where artificial estuarine salinity gradients had been established. Prior to release from net pens between 13 May and 3 June the fish were exposed to an acclimatization period of either 3 or 7 days. The migratory behaviour of the released smolts was studied by underwater video recordings and divers observations. Fast-migrating schools followed the seabed at 8–10m depth, leaving the bay. Non-migrating fish formed large groups close to the seabed (4–7 m depth), or smaller groups close to surface, in the inner part of the bay. Migratory motivation was higher for the large-size groups and increased during the release period for both size categories. Increasing the acclimatization period from 3 to 7 days may have stimulated the migration of small fish early in the release period but the effect of time of release seemed to be more important for the rest of the experiment. During the release period only small smolts were found in the stomachs of gill-netted predators in the bay. The data indicated that smolt size and time of release strongly influenced the migratory motivation of cultured smolts and that the release strategy may affect predation in the release area.     

Temporal and spatial variations in lice numbers on salmon farms in the Hardanger fjord 2004–06

The long and narrow Hardanger fjord in western Norway has a high density of salmon farms and has had severe salmon lice, Lepeophtheirus salmonis, problems. In the years 2004–06, salmon lice numbers were recorded in selected salmon farms in the fjord as part of a larger research project. Most farm sites participated in a strategic control programme and were deloused between November and January in each year. The aim of the programme was to achieve a mean abundance of <0.3 adult female lice at this time and to minimize the infection pressure on wild smolts in the spring. Dedicated teams carried out detailed counting of lice on farmed fish in April–September each year. Temperature conditions were fairly similar throughout the fjord and amongst years, but wide variations in salinities were observed. The two innermost zones, B and C, had the lowest lice mean abundances, whereas the outermost zones, D and E, consistently had more lice. General linear model analyses showed that differences in adult female lice abundance between the zones were associated with differing levels of salinity and emamectin benzoate treatments strategically administered. Mean fish weight was significantly positively correlated with mean abundance of adult female lice.