Habitat selection by juvenile Atlantic salmon: the interaction between physical habitat and abundance

The effect of physical river habitat variables on the distribution of juvenile Atlantic salmon Salmo salar L. in the Rivière de la Trinité, Québec, Canada, was examined using generalized additive modelling. A survey of Atlantic salmon fry and parr densities and habitat variables (flow velocity, water column depth and substratum size) was conducted in the summer months from 1984 to 1992. Clear patterns of habitat use existed: specific ranges of habitat variables were selected, with parr preferring greater velocities, depths and substratum sizes than fry. There was a large variation, however, in juvenile densities for given velocities, depths or substratum sizes, with this variation being greatest in optimal habitats. On examination of an individual year, interaction between the variables was found to explain some of the variation. On a year‐to‐year basis the juvenile Atlantic salmon population was found to exhibit an 'Ideal Free Distribution', which resulted in greatest variation in optimal habitats with year‐to‐year changes in population abundance.     

Temporal and spatial variation in growth of juvenile Atlantic salmon

Spatial and temporal variation in length‐at‐age and environmental factors affecting variation in growth rate of juvenile Atlantic salmon Salmo salar were studied using data from a long‐term study in the River Stjørdalselva, central Norway. Mean annual instantaneous growth rate among 1+ and 2+year juvenile Atlantic salmon varied between 0·59 and 1·50 g g⁻¹ year⁻¹ and mean instantaneous daily growth rate of young‐of‐the‐year (YOY) varied between 0·013 and 0·033 g g⁻¹ day⁻¹. Between year variation in growth was larger than the within year intra‐watercourse spatial variation. For YOY and 1+year Atlantic salmon, a major part of the observed between year variation in growth rates was explained by variation in mean daily water discharge and spring temperature. For 2+year juvenile Atlantic salmon, mean daily water discharge and cohort density were the only variables to significantly explain variation in growth rates. A large part of the within water‐course spatial variation could not be explained by temperature variations and juvenile Atlantic salmon in the uppermost areas of the river, experiencing the lowest ambient temperatures during the growth period, displayed the highest growth rates. Within the baselines set by temperature, biotic and abiotic factors connected to water flow regime and variation in food availability are suggested to be a major determinants of the temporal and spatial variation in juvenile Atlantic salmon growth rates.     

The feeding behaviour of juvenile Atlantic salmon in relation to turbulent flow

The feeding behaviour of juvenile Atlantic salmon Salmo salar in the Sainte‐Marguerite River, Quebec, Canada, varied with the characteristics of turbulent flow. Simulations indicated that juveniles would decrease their swimming costs during attacks by 19·8% in low and by 31·1% in high turbulent conditions by initiating movements in low‐speed flow events. The real swimming costs did not differ from the swimming costs estimated for a situation where fish initiate their movements at randomly selected flow velocities. The juvenile Atlantic salmon did not seem to prefer low‐speed flow events when initiating their movements. The proportion of time used for movements by fish decreased with an increase in the mean and the s . d . of the flow velocity.     

Temperature, light intensity and plasma melatonin levels in juvenile Atlantic salmon

Plasma melatonin synthesis in juvenile Atlantic salmon Salmo salar decreased with increasing light intensity. Melatonin profiles reflected accurately the photoperiod under which the salmon were maintained. Groups maintained at 12°C showed significantly higher ( P < 0·01) levels of dark phase plasma melatonin compared with the groups maintained at 4°C.     

An analysis of the distribution of juvenile Atlantic salmon (Salmo salar) in nature as a function of relatedness using microsatellites

The major objective of this study was to test the hypothesis that juvenile Atlantic salmon kin occupy adjacent territories in their natural habitat in order to profit from the benefits associated with kin-biased behaviours, as has been observed under controlled laboratory conditions. Microsatellites were used to establish the relatedness of salmon fry (in their first summer of life) and parr (in their second and third summer of life) captured in adjacent territories. We did not observe a relationship between the proximity and the relatedness of either parr of the same cohort or fry in their natural habitat. Although many pairs of fry were identified as being related when sampled immediately after emergence, most family groups did not occupy adjacent territories. The high dispersal potential in rivers, the low occupation rate of the habitat and the incidence of half-sibs in nature most probably reduce the opportunity and advantage of kin-biased behaviour, in contrast to laboratory studies conducted in artificial, high-density conditions.     

Impact of naturally spawning captive-bred Atlantic salmon on wild populations: depressed recruitment and increased risk of climate-mediated extinction

The assessment report of the 4th International Panel on Climate Change confirms that global warming is strongly affecting biological systems and that 20–30% of species risk extinction from projected future increases in temperature. It is essential that any measures taken to conserve individual species and their constituent populations against climate-mediated declines are appropriate. The release of captive bred animals to augment wild populations is a widespread management strategy for many species but has proven controversial. Using a regression model based on a 37-year study of wild and sea ranched Atlantic salmon (Salmo salar) spawning together in the wild, we show that the escape of captive bred animals into the wild can substantially depress recruitment and more specifically disrupt the capacity of natural populations to adapt to higher winter water temperatures associated with climate variability. We speculate the mechanisms underlying this seasonal response and suggest that an explanation based on bio-energetic processes with physiological responses synchronized by photoperiod is plausible. Furthermore, we predict, by running the model forward using projected future climate scenarios, that these cultured fish substantially increase the risk of extinction for the studied population within 20 generations. In contrast, we show that positive outcomes to climate change are possible if captive bred animals are prevented from breeding in the wild. Rather than imposing an additional genetic load on wild populations by releasing maladapted captive bred animals, we propose that conservation efforts should focus on optimizing conditions for adaptation to occur by reducing exploitation and protecting critical habitats. Our findings are likely to hold true for most poikilothermic species where captive breeding programmes are used in population management.     

The influence of life-history strategy on lipid metabolism in overwintering juvenile Atlantic salmon

From November to May, the lipid mass in the viscera and carcass of juvenile Atlantic salmon Salmo salar that were undergoing smolt transformation prior to seaward migration ('early migrants') were significantly greater than those of their siblings that would delay migration for at least a further year. During winter (November-February), the depletion of lipid associated with the viscera was significantly greater in early migrants, whilst lipid depletion in the remaining carcass was greater in delayed migrants. Early migrants continued to deplete both lipid compartments in spring (February-May), whereas delayed migrants depleted visceral lipid but replenished carcass lipid over the same period. Fatty acid accumulation rates (a measure of storage lipid synthesis rates) were two to six times greater in visceral than in carcass lipid throughout the study, suggesting that lipid turnover is much more rapid in the viscera. There were no differences in fatty acid accumulation rates between migrant groups in November, despite the much lower food consumption rate of delayed migrants at that time, suggesting that these fish allocated a larger proportion of their nutritional resources to lipid synthesis. In the carcass lipid of early migrants, and in both the visceral and carcass lipid of delayed migrants, the fatty acid accumulation rate was negatively correlated with lipid mass. Fatty acid accumulation rates increased from November to February in both visceral and carcass lipid in the two migrant groups. The fatty acid accumulation rate in carcass lipid was significantly higher in delayed migrants than in early migrants in February, but not in May. These results support the hypothesis that life history strategies involving rapid growth will result in a relatively low allocation of resources to lipid reserves.     

Age-specific aggregation of wild juvenile Atlantic salmon Salmo salar at cool water sources during high temperature events

This study quantified the use of cool water sources by wild 0+, 1+ and 2+ year Atlantic salmon Salmo salar during high water temperatures ( i.e. >23° C) in summer 1995 and 2004. During these events, 0+ year Atlantic salmon did not aggregate or increase in abundance in cool water sites. Interestingly, 1+ and 2+ year Atlantic salmon numbers increased in cool water sites. In addition, these older juveniles formed numerous, discrete aggregations along the plume created by a tributary with aggregation locations being similar between years. Aged 2+ year fish aggregations were at the coolest sources whereas 1+ year aggregations were in locations cooler than the main river. Fish in aggregations on average used deeper sections (average depth: 380 mm) compared with the coolest available habitat in the thermal plume (average depth: 230 mm). Hence, during high temperature events, older juvenile Atlantic salmon moved to cooler water sites and then aggregated in deeper microhabitats.     

Genetic changes in an Atlantic salmon population resulting from escaped juvenile farm salmon

The study was undertaken on three adjacent rivers in NW Ireland, on one of which an Atlantic salmon Salmo salar freshwater juvenile rearing unit is situated. Two markers which distinguished farm and wild populations were used. An Ava II-B RFLP in the ND1 region of mtDNA was at a frequency of 0.58 in the farm strain but absent in the wild populations. Allele E at minisatellite locus Ssa- A45/2/l was at a frequency of 0.91 in farm samples, but at a maximum of 0.41 in the populations in the two rivers adjacent to the one with the juvenile rearing unit. The farm strain showed a significant reduction in mean heterozygosity (0.281 ± 0.057), over three minisatellite loci examined, compared to wild samples (0.532 ± 0.063). The occurrence of farm genotypes and the independent occurrence of mtDNA and minisatellite markers in several parr samples from the river indicated that escaped juvenile salmon completed their life cycle, bred and interbred with native fish, upon their return to the river. Escaped fish homed accurately, as adults, to the site of escape, i.e. the area adjacent to the hatchery outflow in the upstream part of the river. Breeding of males in the lower part of the river was also indicated but this could have been due to mature male parr which had moved downstream. The return of adults of farm origin to the river to breed was indicated by the presence of the Ava II-B haplotype in adults netted in the estuary.     

Chondroid bone and secondary cartilage contribute to apical dentary growth in juvenile Atlantic salmon

The microstructure and tissue composition of the dentary bone in Atlantic salmon salmo salar parr were examined using a variety of histological and whole‐mount techniques. Proximally, the dentary is composed of typical cellular lamellar bone with Sharpey's fibres extending dorsally, proximally and ventrally. Meckel's cartilage is located medially through the entire length of the dentary, and degrades distally resulting in a short transitional zone between hyaline cartilage and connective tissue. At the distal tip of the dentary, isogenic clusters of chondrocytes of periosteal origin were observed secreting small amounts of pericellular cartilage matrix within the bone matrix. These characteristics are highly indicative of secondary chondrogenesis, and suggest that the apical part of the dentary bone in Atlantic salmon does not grow via 'pure' intramembranous ossification, but rather via a modified mode of periosteal ossification involving secondary cartilage and chondroid bone. Furthermore, the unusual mode of gender‐related dentary growth (kype formation) in adult male Atlantic salmon could be the continuation of a general mode of salmonid apical dentary growth.     

Influence of growth and temperature on strontium deposition in the otoliths of Atlantic salmon

Sr : Ca ratios measured in the otoliths of sea‐caged Atlantic salmon Salmo salar showed distinct seasonal peaks that were unrelated to water temperature and somatic growth, suggesting other factors have a greater influence on strontium deposition. Atlantic salmon remained in the sea‐cages for two seasons, or equivalent to two‐seawinter fish in the wild and were subjected to seasonally varying temperatures. Water temperature appeared to be inversely related to the Sr : Ca ratios, but this relationship was statistically not significant. Furthermore, water temperature could not explain the distinct increase in the strontium during the second year. The intensity of the Sr peaks increased in the second season while average winter water temperatures were consistent between years. Additionally, strontium deposition in the otoliths was unrelated to somatic growth. Somatic growth, as evidenced by circuli spacings on the scales, was largely invariant and therefore could not explain the observed peaks. Though not explicitly measured, the data are consistent with the notion that strontium deposition is a function of maturity state in Atlantic salmon.     

Prey selection by experienced and naive juvenile Atlantic salmon

Both in foraging groups and in a sequential prey encounter context, learning had a visible effect on the pattern of selection for three live prey types ( Ecdyonurus larvae, Hydropsyche larvae, and Gammarus ) by juvenile Atlantic salmon Salmo salar . Compared to wild-caught fish, naive, hatchery-reared fish that had not been exposed to natural prey ate Hydropsyche larvae in a remarkably low proportion, and consumed a higher proportion of Gammarus. Ecdyonurus experienced a high and rather steady predation rate across the experience gradient, but after a short period of experience with live prey the consumption rate for Hydropsyche increased drastically, and that of Gammarus decreased, matching the selection pattern exhibited by wild fish. Individual fish offered prey in a sequential encounter context increased consumption rates of all the prey types as they gained experience, but the improvement was higher for the prey that were less consumed initially. Fish became more selective as they approached satiation, conforming to the prediction of optimal foraging theory that higher predator's energy requirements, as well as low food availability, result in reduced selectivity. The results also suggest that fish from distinct populations can differ in the degree of diet selectivity according to their energetic requirements for growth. The fast learning response of Atlantic salmon parr towards novel prey probably allows fish to maintain a high foraging efficiency when faced with frequent changes in the availability of different prey types.     

Oxygen consumption, ammonia excretion and protein use in response to thermal changes in juvenile Atlantic salmon Salmo salar

Experiments were designed to examine the effects of various temperature challenges on oxygen consumption and ammonia excretion rates and protein utilization in juvenile Atlantic salmon Salmo salar . Fish acclimated to 15° C were acutely and abruptly exposed to either 20 or 25° C for a period of 3 h. To simulate a more environmentally relevant temperature challenge, a third group of fish was exposed to a gradual increase in temperature from 15 to 20° C over a period of 3 h ( c. 1·7° C h⁻¹). Oxygen consumption and ammonia excretion rates were monitored before, during and after the temperature shift. From the ammonia excretion and oxygen consumption rates, protein utilization rates were calculated. Acute temperature changes (15–20° C or 15–25° C) caused large and immediate increases in the oxygen consumption rates. When the temperature was gradually changed ( i.e. 1·7° C h⁻¹), however, the rates of oxygen consumption and ammonia excretion were only marginally altered. When fish were exposed to warmer temperatures ( i.e. 15–20° C or 15–25° C) protein use generally remained at pre-exposure (15° C) levels. A rapid transfer back to 15° C (20–15° C or 25–15° C) generally increased protein use in S. salar . These results indicate that both the magnitude and the rate of temperature change are important in describing the physiological response in juvenile salmonids.     

Winter behaviour of juvenile Atlantic salmon Salmo salar L. in experimental stream channels: effect of substratum size and full ice cover on spatial distribution and activity pattern

Activity and choice of areas offering different cover (substratum or surface ice) for juvenile Atlantic salmon Salmo salar were studied in experimental stream channels during winter. Channels were completely ice covered between December and March. During this period, the ice thickness increased from 50 to 300 mm after which 50% of the ice was experimentally removed and followed by c. 2·5-fold increase in discharge to simulate the effects of spring flood. Large substrata provided preferred habitats but areas with small substratum sizes were also used when full surface ice provided above-stream cover and the stream discharge was relatively low. The fish remained nocturnal throughout the study but the level of day activity significantly increased as the surface ice became thicker. Maximum movement distance during a 24 h period and homing-at-dawn behaviour remained at a constant level throughout the main winter, but significantly changed during the simulated spring flood (mean ± s . e . maximum extent of movements within 24 h increased from 1·1 ± 0·1 to 3·0 ± 0·5 m; homing behaviour decreased from the highest level of 89·3 to 34·6% during spring flood). Overwinter survival was high (92·9%). Relative mass increase during the study ranged from –8·3 to 28·5%, and 84% of the juvenile Atlantic salmon gained mass. The highest rates of mass increase were associated with frequent movements between areas of different substratum size. The results indicate that during winter: (1) Atlantic salmon parr preferred large substratum cover compared with surface ice cover at the fish densities studied here, (2) juvenile Atlantic salmon were predominantly nocturnal but diurnal activity increased as surface ice became thicker and (3) increase in water discharge during spring altered the behaviour of juvenile Atlantic salmon and may have caused additional habitat shifts.     

Diel activity pattern of juvenile Atlantic salmon (Salmo salar) in early and late winter

Radiotelemetry was used to investigate the diel activity pattern of juvenile Atlantic salmon (Salmo salar) in early and late winter. Fish were active throughout the diel cycle. However, there was significantly less daytime than nighttime movement and movement declined significantly with increasing fork length. Maximizing winter growth rate, through an overall increase in foraging activity, may reduce the risk of starvation in smaller fish. The results of the present study provide evidence that the activity patterns of juvenile salmonids are quite complex and support the suggestion that individual variation in activity patterns are, at least, partially driven by body size.     

Time series covering up to four decades reveals major changes and drivers of marine growth and proportion of repeat spawners in an Atlantic salmon population

1. Wild Atlantic salmon populations have declined in many regions and are affected by diverse natural and anthropogenic factors. To facilitate management guidelines, precise knowledge of mechanisms driving population changes in demographics and life history traits is needed.
2. Our analyses were conducted on (a) age and growth data from scales of salmon caught by angling in the river Etneelva, Norway, covering smolt year classes from 1980 to 2018, (b) extensive sampling of the whole spawning run in the fish trap from 2013 onwards, and (c) time series of sea surface temperature, zooplankton biomass, and salmon lice infestation intensity.
3. Marine growth during the first year at sea displayed a distinct stepwise decline across the four decades. Simultaneously, the population shifted from predominantly 1SW to 2SW salmon, and the proportion of repeat spawners increased from 3 to 7%. The latter observation is most evident in females and likely due to decreased marine exploitation. Female repeat spawners tended to be less catchable than males by anglers.
4. Depending on the time period analyzed, marine growth rate during the first year at sea was both positively and negatively associated with sea surface temperature. Zooplankton biomass was positively associated with growth, while salmon lice infestation intensity was negatively associated with growth.
5. Collectively, these results are likely to be linked with both changes in oceanic conditions and harvest regimes. Our conflicting results regarding the influence of sea surface temperature on marine growth are likely to be caused by long‐term increases in temperature, which may have triggered (or coincided with) ecosystem shifts creating generally poorer growth conditions over time, but within shorter datasets warmer years gave generally higher growth. We encourage management authorities to expand the use of permanently monitored reference rivers with complete trapping facilities, like the river Etneelva, generating valuable long‐term data for future analyses.

     

Monthly indices of the post-smolt growth of Atlantic salmon from the Drammen River, Norway

Indices of the monthly post-smolt growth of three maturity groups of Atlantic salmon Salmo salar from the Drammen River in Norway were examined. Scale samples were collected from 1983 to 2003, providing a retrospective growth history for 2854 individuals. Overall post-smolt growth of all three maturity groups decreased throughout the study period. When the post-smolt year was segmented into 8 monthly indices, there was a significant decline in growth during the fourth and fifth month, typically months of maximum growth. The pre-fishery abundance of one and two sea-winter fish from Norway also showed a decline throughout the study period and was positively correlated to the post-smolt growth of fish returning to the Drammen River. Freshwater growth was negatively correlated to the pre-fishery abundance, thus indicating that factors affecting post-smolt growth during the fourth and fifth months at sea may be responsible for declining return numbers in Norway.     

Behavioural influences on life-history variation in juvenile Atlantic salmon,Salmo salar

Synopsis Juvenile Atlantic salmon emigrate from the river in a given year as a consequence of a physiological decision influencing appetite and growth during the previous summer. The direction of the decision depends on developmental performance exceeding a genetically determined threshold at that time, and that performance is governed by environmental opportunity. The animal's foraging efficiency (ability to avoid predators, and to compete) determines how well that opportunity is used. Those fish which maintained relatively high growth after July preferred higher stream flows, and were more likely to hide than to flee from a predator. Predator vigilance reduced intake, and ability to discriminate edible particles. Early competitive ability depended on fierceness rather than size, and ultimate large size was a consequence of dominant status. Initial status among high ranking individuals (but not among low ones) predicted the likelihood of maintaining growth in late summer. Size by July was the better predictor for low ranked fish. The influence of high status on life-history variation depends on how much it suppresses the growth of those lower in the hierarchy. Invited review     

The effects of prior residence on behavior and growth rates in juvenile Atlantic salmon (Salmo salar)

It is well documented that prior residence confers advantagesin territorial disputes, but its impact on other aspects ofbehavior and fitness is less understood. We tested how priorresidence influences the subsequent feeding behavior and growthperformance of dispersing Atlantic salmon fry (Salmo salar)using experimental manipulations of residence in a seminaturalstream tank. In replicated trials, groups of seven "primary"fish were released into the stream tank 3 days ahead of seven"secondary" fish. Standardized behavioral observations were madeon each fish over the following 14 days, after which all fishwere removed and measured. Primaries and secondaries were initiallythe same size and body condition and exhibited the same degreeof site fidelity. However, primaries darted higher into thewater column to intercept prey items, fed at a higher rate,and subsequently grew faster. Larger fish (in terms of body length)tended to be more dominant, and dominants grew faster than subordinates.However, there was no difference in dominance between primaries andsecondaries. These results suggest that the well-documentedadvantage of early-emerging salmon fry over late-emerging frycannot be completely attributed to intrinsic differences andthat the advantage is partly mediated via a prior residenceeffect. Furthermore, prior residents gain foraging advantageswithout necessarily becoming more dominant.     

Recombinant genotypes in backcrosses of male Atlantic salmon × brown trout hybrids to female Atlantic salmon

When male hybrids of Atlantic salmon × brown trout were backcrossed to female Atlantic salmon, approximately 1% of diploid progeny hatched. These were shown to exhibit recombinant genotypes when examined electrophoreticalty at five enzyme loci. This is the first confirmation of genie recombination in backcrosses of these species. Triploidization greatly increases the proportion of backcross progeny which hatch.