Competition for shelter among over‐wintering signal crayfish and juvenile Atlantic salmon

Three separate effects on refuge use by signal crayfish Pacifastacus leniusculus and Atlantic salmon Salmo salar were examined: (1) the effect on Atlantic salmon of an addition of signal crayfish (doubling the total number of animals), (2) the effect on signal crayfish of an addition of Atlantic salmon and (3) intraspecific compared with interspecific competition, compared by holding total density of animals constant and varying the proportion of signal crayfish and Atlantic salmon in trials. Observations were made during winter, when both species are nocturnal. The proportion of Atlantic salmon sheltering was significantly lower in the presence than in the absence of signal crayfish when the interspecific treatment (Atlantic salmon plus signal crayfish) effected a doubling in density compared to the intraspecific treatment (Atlantic salmon alone). The proportion of signal crayfish sheltering was independent of the presence of Atlantic salmon. When total density was constant, the proportion of Atlantic salmon sheltering was significantly higher in intraspecific (52·8%) than interspecific trials (27·3%). Atlantic salmon out of shelter during the day in winter are believed to be very vulnerable to predators and the capacity for fish to share shelters with one another is known to be very low. Therefore, competition from crayfish for winter shelters may lead to detrimental effects on Atlantic salmon populations.     

Effect of density on competition between wild and hatchery‐reared Atlantic salmon for shelter in winter

The effect of varying the density of hatchery‐reared Atlantic salmon Salmo salar on the ability of single wild fish to occupy a shelter is assessed. Although there was strong density‐dependence on sheltering overall, the ability of wild Atlantic salmon parr to occupy a shelter was not affected by the presence of hatchery‐reared fish even when outnumbered by four to one. These findings illustrate a competitive asymmetry for shelter in favour of the wild fish at the densities tested.     

Shelter selection in juvenile Atlantic salmon,or why do salmon seek shelter in winter?

During winter, juvenile Atlantic salmon Salmo salar become nocturnal and seek refuge during the day in the stream bed gravel interstitial spaces. The function of this behaviour is unclear, but two major types of hypothesis have been proposed. One is that the fish are hiding from something (e.g. a predator) and the other is that the fish are seeking shelter from the water current. These hypotheses were tested by examining the selection of juvenile salmon for refuges that offered different degrees of concealment or shelter. The fish clearly preferred refuges that allowed them to hide (i.e. they were dark and opaque) but offered little shelter from the current. Therefore, it can be assumed that the primary function of this nocturnal behaviour during winter is most likely to hide from diurnal predators.     

Production of juvenile salmonids in small Norwegian streams is affected by agricultural land use

1. We estimated the biomass and production of juvenile anadromous brown trout (Salmo trutta) and Atlantic salmon (Salmo salar) (parr) in 12 streams in the Skagerrak area of Norway to identify controlling environmental factors, such as land‐use and water chemistry. 2. Production estimates correlated positively with fish density in early summer, but not with the size of the catchment. The summer biomass of age‐0 brown trout and Atlantic salmon was smaller than that of age‐1 and constituted 27.4 and 25.7%, respectively, of the total biomass of the two groups. 3. Mean production of brown trout from July to September varied between streams, but in most cases it was below 2 g 100 m^?2 day^?1. Yearly cohort production from age‐0 in July to age‐1 in July was 10 g m^?2 or less, with mean annual production of 1.32 g 100 m^?2 day^?1, equivalent to 4.8 g m^?2 year^?1. The corresponding annual cohort production of Atlantic salmon was 0.38 g 100 m^?2 day^?1 or 1.4 g m^?2 year^?1. Annual production to biomass ratio (P/B) for brown trout of the same cohort in the various streams was between 1.47 and 4.37; the overall mean (±SD) for all streams was 2.25 ± 0.94. Mean turnover rate of Atlantic salmon was 2.73 ± 0.24. 4. Production of 0+ brown trout during the summer correlated significantly with the percentage of agricultural land and forest/bogs in the catchment, with maxima at 20 and 75%, respectively. Age‐0 brown trout production also correlated with concentration of nitrogen and calcium in the water, with maxima at 2.4 and 14 mg L^?1, respectively. 5. The results support the hypothesis that brown trout parr production reflects the quality of their habitat, as indicated by the dome‐shaped relationship between percentage of agricultural land and the concentration of nitrogen and calcium in the water.     

Density-dependent refuge use among over-wintering wild Atlantic salmon juveniles

Sheltering behaviour of wild juvenile Atlantic salmon in an indoor stream was found to be density dependent; the proportion of fish sheltering decreased significantly with increasing population density. The mean number of fish occupying refugia was ≤1.5 fish per refuge even at very high densities (potentially 5 fish per shelter). These results suggest that shelter availability has potentially important consequences for the carrying capacity for natural populations of salmon in streams.     

Rearing conditions influence refuge use among over-wintering Atlantic salmon juveniles

At low population densities, both hatchery- and wild-reared Atlantic salmon Salmo salar , which were progeny of a common genetic stock, occupied sub-gravel shelters alone. However, at high population densities, hatchery-reared fish occurred within shelters in significantly higher numbers than wild-reared fish. There was a significant effect of time after introduction to the experimental arenas on the proportion of hatchery-reared fish sheltering. These results show that rearing environment influences the behaviour of Atlantic salmon parr during winter and may also affect the carrying capacity of their environment.     

Migration of hatchery‐reared Atlantic salmon and wild anadromous brown trout post‐smolts in a Norwegian fjord system

Hatchery‐reared Atlantic salmon Salmo salar ( n  = 25) and wild anadromous brown trout (sea trout) Salmo trutta ( n  = 15) smolts were tagged with coded acoustic transmitters and released at the mouth of the River Eira on the west coast of Norway. Data logging receivers recorded the fish during their outward migration at 9, 32, 48 and 77 km from the release site. Seventeen Atlantic salmon (68%) and eight sea trout (53%) were recorded after release. Mean migratory speeds between different receiver sites ranged from 0·49 to 1·82 body lengths (total length) per second (bl s⁻¹) for Atlantic salmon and 0·11–2·60 bl s⁻¹ for sea trout. Atlantic salmon were recorded 9, 48 and 77 km from the river mouth on average 28, 65 and 83 h after release, respectively. Sea trout were recorded 9 km from the release site 438 h after release. Only four (23%) sea trout were detected in the outer part of the fjord system, while the rest of the fish seemed to stay in the inner fjord system. The Atlantic salmon stayed for a longer time in the inner part than in the outer parts of the fjord system, but distinct from sea trout, migrated through the whole fjord system into the ocean.     

Long‐term effects of stock transfers: synergistic introgression of allochthonous genomes in salmonids

The genus Salmo was employed as a model to study introgression of genes between species due to secondary contacts. Seven microsatellite loci, the LDH‐C1* locus and the 5S ribosomal DNA were studied. Results showed the mutually enhanced introgression of allochthonous genomes into southern European salmonids. This phenomenon appears to go beyond a simple consequence of the altered behaviour of domestic individuals. Invasions of autochthonous genomes by allochthonous genes would be enhanced by human activities such as stock transfers, which would simultaneously promote allochthonous and allospecific (from other species) introgressions in a synergistic process in Atlantic salmon Salmo salar and brown trout Salmo trutta. As a minor result, the data do not support the value of the microsatellite locus SsaD486 as a species‐specific marker.     

Seasonal changes in the body composition of young riverine Atlantic salmon and brown trout

The body composition of protein and fat in Atlantic salmon Salmo salar and brown trout Salmo trutta before and after winter was investigated in a temperate, small river, normally ice covered from the middle of November until the end of March. Fat, protein and specific energy declined greatly in winter but were replenished rapidly in spring. Rates of decline were slower for the smallest fish, which also had the lowest specific content of fat, protein, and energy, while the differences in absolute amounts were greatest for the largest fish. The mean specific fat content was reduced by 45–70% during winter, relative to the pre-winter period (September). Mean daily reductions in specific enegy of the larger size groups of brown trout (3·7 × 10⁻³ kJ g⁻¹ day⁻¹) were almost half of the corresponding values for the largest Atlantic salmon (6·3 × 10⁻³ kJ g⁻¹ day⁻¹) during winter. A minor reduction in protein content was found during winter, with mean reductions of 6–10% in comparison to those in September. During spring the fat content was replenished rapidly, particularly for the smallest salmon fry (a threefold increase from April to June). Fat content in the larger salmon and trout increased by about 1·8 times. Based on estimated metabolic rates, digested energy during wintertime may contribute about two-thirds of the brown trout fry's energy demand. For Atlantic salmon, the corresponding value is about 50%. The winter period put considerable stress on the young salmonids living in lotic environments, in particular for the smallest fry with the lowest energy content before winter and the largest losses during winter. This should make the fry more vulnerable to adverse abiotic and biotic factors.     

Age effects on diel activity patterns of juvenile Atlantic salmon: parr are more nocturnal than young‐of‐the‐year

In the summer Atlantic salmon Salmo salar parr were predominantly active at night, while young‐of‐the‐year were equally active during day and at night. These findings are consistent with the prediction that larger animals should be less willing to risk predation for a certain increase in size than smaller animals.     

Patterns of density‐dependent growth in juvenile stream‐dwelling salmonids

Literature data for 19 populations from 16 different studies of six species of stream‐dwelling salmonids were analysed to test the hypothesis that density‐dependent growth is stronger at low rather than at high population density. Fifteen of 19 populations showed evidence of a significant decrease in growth rate with increasing density. In 11 of these 15 populations, the pattern of density‐dependent growth was better described by a negative power curve than by a linear regression ( i.e . Akaike Information Criterion, AIC _linear −  AIC _power > 2), whereas only one population was better described by a linear regression than by a negative power curve; three populations were adequately described by both models ( AIC  < 2). In 10 of the 11 populations that were best described by a negative power curve, most of the decrease in growth rate occurred at population densities <1 fish m⁻², when space limitation is unlikely. This analysis provides broad support for the hypothesis that density‐dependent growth in stream salmonids occurs primarily at low population densities, probably due to exploitative competition.     

The cost of aggregation: juvenile salmon avoid sharing winter refuges with siblings

Kin selection advantages are usually accrued by individualsthat associate with close relatives. But aggregation may alsobe costly, by increasing the risk of predation or resource competition,for example. As a result, individuals should increase theirinclusive fitness by trading the costs and benefits of kin associationand aggregation. Studies of kin selection to date have focusedon situations where there is ample opportunity for kin-biasedbehavior and therefore for the formation of kin groups. Herewe used juvenile Atlantic salmon to test an alternative strategy:that under conditions where the potential for kin-biased behavioris negligible, individuals should, when aggregating, avoid ratherthan associate with kin to avoid imposing the costs of aggregationupon close relatives. By testing salmon during winter, whenjuveniles shelter inactively in streambed refuges, we testedwhether individuals associate with or avoid their siblings ata time when the opportunity for kin-directed behaviors is restricted.Our results provide the first evidence of kin avoidance in nonreproductiveanimals studied under semi-natural conditions.     

Influence of size, winter duration and density on sexual maturation of Atlantic salmon (Salmo salar) juveniles in Little Codroy River (southwest Newfoundland)

Factors affecting sexual maturation of Atlantic salmon ( Salmo salar L.) juveniles in fresh water were analysed under wild conditions in Little Codroy River (southwest Newfoundland). Large size after the first year of life and short winters favoured maturation of age 1 + male parr. Incidence of maturing 1+ males was greater where high densities of parr (>0.2 individuals m ²; fish >6cm fork length) were recorded in the second season of growth. These results were discussed according to current knowledge on early maturation in Atlantic salmon.     

Diel activity patterns of juvenile Atlantic salmon in rivers with summer water temperature near the temperature‐dependent suppression of diurnal activity

Direct day and night underwater observations of juvenile Atlantic salmon Salmo salar during summer and autumn showed a duality in response to temperature between 7 and 11° C for young‐of‐the‐year (YOY) Atlantic salmon. They were predominantly diurnal in early summer and nocturnal in late summer although water temperatures were similar. Post‐YOY Atlantic salmon did not show a strong response to temperature as they were mostly nocturnal during the study period. It is suggested that the difference in activity patterns between YOY and post‐YOY Atlantic salmon can be explained by size‐dependent trade‐off between growth and predation risk.     

Determination of body shape variation in Irish hatchery‐reared and wild Atlantic salmon

Discriminant function analysis was used to distinguish morphologically between samples of parr, smolts and adult Atlantic salmon Salmo salar from several hatchery and river systems in Ireland. The effect of habitat shift was investigated in Atlantic salmon parr. Parr grown from the eyed‐egg stage with a non‐sibling group in a hatchery environment, came to resemble the mean body shape of their host hatchery Atlantic salmon stock more closely than that of a full sibling group grown at their natal hatchery. Wild Atlantic salmon smolts differed in shape from hatchery‐reared smolts. This difference was less pronounced, but still statistically significant when wild adults were compared with hatchery‐reared adults captured in the coastal drift‐net fishery after a year spent at sea. Rearing conditions had a significant impact on the production and growth of fish body shape. This in turn may have affected adaptability and survivorship of ranched Atlantic salmon in the marine environment.     

Daily feeding rhythms in Atlantic salmon II: size-related variation in feeding patterns of post-smolts under constant environmental conditions

Previous studies of feeding rhythms in post-smolt Atlantic salmon Salmo salar have produced variable results. Therefore, the present study was performed to clarify the source of these differences and to examine size-related variation in daily feeding patterns. A small group of hatchery reared post-smolts was kept in an indoor seawater tank under conditions of constant photoperiod and light intensity, and negligible diurnal variation in water temperature. Feeding responses were monitored by video camera throughout the day. The resultant feeding pattern for the population mirrored an earlier study, with the highest levels of feeding activity in the early morning and late evening. Differences between this and other studies may be due to variation in climate, season or life-history stage. Daily feeding patterns were size-dependent. The largest fish fed at a relatively constant level through the day, medium-sized fish fed in a pattern similar to that described above for the population and small fish fed most at first and last light. These differences were due probably to the effect of competitive interactions upon fish of different sizes.     

Microhabitat use by brown trout, Salmo trutta L. and Atlantic salmon, S. salar L., in a stream: a comparative study of underwater and river bank observations

Two methods, visual observation from the river bank and visual observation underwater by diving, were compared for microhabitat studies in young brown trout and Atlantic salmon in a stream. A wide range of habitat conditions were surveyed. Each method yielded different results with respect to microhabitat use. River bank observations missed small fish under surface turbulence and in deeper waters. Underwater observations missed small fish in shallow areas.     

Predator avoidance behaviour in wild and hatchery‐reared brown trout: the role of experience and domestication

Juvenile brown trout Salmo trutta from natural populations reacted to the presence of piscivorous brown trout by increasing the use of refuges. In contrast, second‐generation hatchery fish and the offspring of wild fish raised under hatchery conditions were insensitive to predation risk. The diel pattern of activity also differed between wild and hatchery brown trout. Second‐generation hatchery fish were predominantly active during daytime regardless of risk levels. Wild fish, however, showed a shift towards nocturnal activity in the presence of predators. These findings emphasize the potential role of domestication in weakening behavioural defences. They support the idea that the behavioural divergence between wild and domesticated individuals can arise from a process of direct or indirect selection on reduced responsiveness to predation risk, or as a lack of previous experience with predators.     

Return migration of one‐sea‐winter Atlantic salmon in the River Tana

Fifty‐three one‐sea‐winter Atlantic salmon Salmo salar (45–63 cm L _T) were radio‐tagged in the Tana fjord, Barents Sea, in 1995. Thirty‐seven fish (70%) entered the freshwater zone of the River Tana in an average of 3 days after release in the fjord. The migration speeds in the lowest river section below the first riffle area were significantly higher than in the subsequent river section below the second riffle area. Similarly, the observed time spent in the first riffle area was significantly lower than in the next riffle area. The majority of Atlantic salmon entered the river during the hours of high tide and the subsequent ebb tide. In addition, most river entries were recorded around midnight. No effects of river flow on the river entry or migration speed were detected, but the migration speed of Atlantic salmon in both river sections examined was greater at lower temperatures. Twenty‐eight fish (72%) were recaptured in the river, 71% of them with weirs and gillnets, and 29% by rod and line. Over half of the Atlantic salmon (54%) were recaptured within 3 weeks following river entry, and within the first 100 km of the river (56%). The results are discussed in relation to earlier studies on multi‐sea‐winter Atlantic salmon in the River Tana.     

Predicting microhabitat selection in juvenile Atlantic salmon Salmo salar by the use of logistic regression and classification trees

1. We compared the capacity of logistic regression (LR) and classification tree (CT) models to predict microhabitat use and the summer distribution of juvenile Atlantic salmon, Salmo salar, in two reaches of a small stream in eastern Quebec. 2. The models predicted the presence or absence of salmon at a location on the basis of habitat features (depth, current velocity, presence of instream and overhead cover, substratum particle size, and distance to stream bank) measured at that location. Models were validated by means of crossover field tests evaluating the performance of models developed for one reach (calibration trials) when applied to the other reach (validation trials). Model performance was evaluated with regard to accuracy, generality and ease of use and interpretation. Prediction maps based on habitat features were also built to compare the observed position of fish with those predicted by LR and CT models. 3. The spatial distribution of active fish differed markedly from that of resting fish, apparently as a result of the selection for water greater than about 30 cm depth by active fish and for the presence of rocky cover by resting fish. 4. All models made accurate predictions, validated by crossover trials. For both LR and CT models, the prediction maps reflected well the actual fish distributions. However, CT models were easier to build and interpret than LR models. CT models also had less variable performance and a smaller decline in predictive capability in crossover trials (for fish at rest), suggesting that they may be more transferable than LR models.