Interactions between riparian shading and food supply: a seasonal comparison of effects on time budgets, space use and growth in Atlantic salmon Salmo salar

This study examines seasonal (winter v. summer) differences in space-time budgets, food intake and growth of Atlantic salmon Salmo salar parr in a controlled, large-scale stream environment, to examine the direction and magnitude of shifts in behaviour patterns as influenced by the availability of overhead cover and food supply. Salmo salar parr tested in the presence of overhead cover were significantly more nocturnal and occupied more peripheral positions than those tested in the absence of overhead cover. This increase in nocturnal activity was driven primarily by increased activity at night, accompanied by a reduction in daytime activity during winter. The presence of overhead cover had no effect on rates of food intake or growth for a given food supply in a given season. Growth rates were significantly higher for fish subjected to a high food supply than those subjected to a low food supply. Food supply did not affect the extent to which S. salar parr were nocturnal. These results were consistent between winter and summer. The use of riparian shading as a management technique to mitigate the effects of warming allows the adoption of more risk-averse foraging behaviour and may be particularly beneficial in circumstances where it serves also to increase the availability of food.     

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.     

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.     

Shelter availability affects behaviour, size-dependent and mean growth of juvenile Atlantic salmon

1. Anthropogenic disturbances of the physical habitat and corresponding effects on fish performance are key issues in stream conservation and restoration. Reduced habitat complexity because of increased sediment loadings and canalization is of particular importance, but it is not clear to what extent fish populations are influenced directly by changes in the physical environment, or indirectly through changes in the biotic environment affecting the food availability. 2. Here, we test for the direct effect of habitat complexity on the performance (growth) of juvenile Atlantic salmon by manipulating shelter availability (interstitial spaces in the substrate) across 20 semi‐natural stream channels without altering the substrate composition, and stocking each channel with a common density of fish. A simple method for measuring salmonid shelters using flexible PVC tubes was developed and tested. Daytime sheltering behaviour and growth rates were compared across the channels differing in shelter availability. 3. Measured shelter availability was strongly negatively correlated with observed number of fish not finding shelters and mass loss rates of the fish (growth performance) increased with decreasing number of measured shelters. Number and mean depth of interstitial spaces explained up to 68% and 24% of the among‐channel variation in sheltering behaviour and growth performance, respectively. Furthermore, negative effects of shelter reduction increased with fish body size. Thus, changes in habitat structure may even influence the size selection gradients. 4. Shelter availability is an easily measured variable, possibly affecting the population demographics and long‐term evolutionary processes, and is therefore a key habitat factor to be considered in stream restoration and habitat classification.     

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.     

Feeding chronology of Atlantic salmon parr in subarctic rivers: consistency of nocturnal feeding

During the ice-free season in three subarctic rivers in northern Norway and Finland, Atlantic salmon parr Salmo salar exhibited significant diel fluctuations in stomach content weights in August and September, the largest weights being recorded consistently at 0300 hours. Accordingly, the feeding rate of the parr was highest during the night and lowest in daytime, whereas during the other periods of the ice-free season, no pronounced diel fluctuations in food intake occurred. The consistent nocturnal feeding periodicity in early autumn in subarctic rivers may be associated with benthic feeding and an increased availability of benthic prey at night.     

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.     

Allozyme heterozygosity, date of first feeding and life history strategy in Atlantic salmon

The relations between allozyme heterozygosity, relative date of first feeding and life history strategy in juvenile Atlantic salmon Salmo salar were examined using eggs obtained from a 400 family cross (20 male × 20 female adult Atlantic salmon). Multilocus heterozygosity, through its positive associations with the timing of first feeding and growth rate, was correlated with life history strategy in juvenile Atlantic salmon, albeit under genotype × environmental (temperature, food availability) regulation. Under hatchery conditions, a 10 day difference was observed in the relative date of first feeding between early and late first feeding Atlantic salmon. Early first feeding Atlantic salmon exhibited a significantly higher mean heterozygosity, grew faster at ambient water temperature (April to November) and a significantly higher proportion adopted the early freshwater maturation (age 0+ years, male fish) or early migrant (age 1+ years, mainly female fish) strategies compared to late first feeding Atlantic salmon. Elevated water temperatures over the winter (December to April, >10·5° C) provided additional growth opportunity allowing previously mature male parr (mainly early first feeders) and lower modal group parr (mainly late first feeders) to adopt the early migrant strategy by the following spring.     

Diel feeding rhythms and daily food consumption of juvenile Atlantic salmon in the River Alta, northern Norway

Atlantic salmon fry (0+) sampled from the River Alta exhibited only minor differences in stomach content weights and feeding rates throughout diel periods, but feeding rates were generally lowest at night. In contrast, salmon parr (1+ to 3+) had large diel fluctuations in stomach content weight, with the largest weights usually being recorded during the night and early morning. Accordingly, their feeding rates were highest at night. This nocturnal feeding pattern was consistent throughout all sampling occasions, and appeared to reflect a persistent feeding periodicity in the salmon parr. The daily food consumption rates of both fry and parr were highest during midsummer and decreased towards autumn.     

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.     

Seasonal and Diel Changes in Behaviour,Microhabitat use and Preferences by Young Pool-dwelling Atlantic Salmon,Salmo salar,and Brown Trout,Salmo trutta

There was a pronounced decline in activity of young pool-dwelling Atlantic salmon, Salmo salar, and brown trout, Salmo trutta, as the water temperatures dropped in the autumn and early winter, and the fish switched from a predominantly diurnal towards a nocturnal activity pattern. Such a switch in activity pattern has previously been observed in young brown trout, but the present study is the first documentation for juvenile Atlantic salmon under natural conditions. Juvenile fish fed actively even when water temperatures were below 0°C, although foraging behaviour at near-freezing temperatures was recorded exclusively during night surveys. This indicates that other proximate factors, in addition to water temperature, affect the activity of young salmon and trout in rivers. Trout kept feeding positions significantly higher above bottom than salmon in August and September, but both species reduced the height above bottom at the onset of winter, possibly due to reduced swimming performance and lowered food availability in the upper part of the water column.     

Infection of Atlantic salmon, Salmo salar L., by Gyrodactylus salaris, Malmberg 1957, in the River Lakselva, Misvær in northern Norway

Gyrodacrylus salaris was most probably introduced to the River Lakselva in 1975 through stocking of Atlantic salmon from an infected hatchery. The parasite population grew rapidly, and the parasite spread throughout the entire watercourse during the summer of 1976. This epidemic situation led to mortality among the young Atlantic salmon, and the density of salmon parr was heavily reduced from 1976 to 1977. The density of salmon parr has remained close to zero since then, while there are no apparent trends toward decrease or increase in the density of brown trout. In spite of the reduced density of young salmon, a new epidemic has developed each year among the few young 0+ and 1+ Atlantic salmon present in the river. Results from successive sampling during the summer of 1987, 1988 and 1989 indicate that most of the presmolt salmon are attacked during their first summer or autumn of life. The infection develops into an epidemic during the first autumn, winter or the next summer. The build-up of the parasite burden on the fish leads in turn to mortality. Norwegian Atlantic salmon probably have no resistance against G. salaris, since the parasite has recently been introduced to Norwegian rivers.     

Effects of body size on sympatric shelter use in over‐wintering juvenile salmonids

The study tests two hypotheses: (1) the degree of shelter dominance in Atlantic salmon Salmo salar and brown trout Salmo trutta increases progressively with increasing size differential between heterospecific fish in a pair and (2) shelter dominance, standardized to size differential, correlates with aggression. The results support the first but not the second hypothesis, suggesting that the fitness consequences of high growth performance during the summer are likely to become evident during winter. At this time of year, when mortality is high among both Atlantic salmon and brown trout, shelter dominance may increase the chances of survival.     

Summer temperature variation and implications for juvenile Atlantic salmon

Temperature is important to fish in determining their geographic distribution. For cool- and cold-water fish, thermal regimes are especially critical at the southern end of a species’ range. Although temperature is an easy variable to measure, biological interpretation is difficult. Thus, how to determine what temperatures are meaningful to fish in the field is a challenge. Herein, we used the Connecticut River as a model system and Atlantic salmon (Salmo salar) as a model species with which to assess the effects of summer temperatures on the density of age 0 parr. Specifically, we asked: (1) What are the spatial and temporal temperature patterns in the Connecticut River during summer? (2) What metrics might detect effects of high temperatures? and (3) How is temperature variability related to density of Atlantic salmon during their first summer? Although the most southern site was the warmest, some northern sites were also warm, and some southern sites were moderately cool. This suggests localized, within basin variation in temperature. Daily and hourly means showed extreme values not apparent in the seasonal means. We observed significant relationships between age 0 parr density and days at potentially stressful, warm temperatures (≥23°C). Based on these results, we propose that useful field reference points need to incorporate the synergistic effect of other stressors that fish encounter in the field as well as the complexity associated with cycling temperatures and thermal refuges. Understanding the effects of temperature may aid conservation efforts for Atlantic salmon in the Connecticut River and other North Atlantic systems.     

Size variability of juvenile Atlantic salmon: links to environmental conditions

Climate change models predict a 2 to 6° C increase in air temperature within the next 100 years in the Maritime Provinces of eastern Canada. Higher air temperatures are expected to contribute to increased water temperatures, alterations in stream flow conditions, and ultimately reductions in fish growth. Mean annual size-at-age of juvenile Atlantic salmon Salmo salar decreased in the Northwest Miramichi and Southwest Miramichi Rivers between 1971–1999. Lengths-at-age of juveniles were significantly correlated between the two rivers. For Atlantic salmon parr, stronger associations between inter-cohort fork length ( L _F) than intra-cohort L _F were observed, suggesting that environmental conditions in the current year of growth have the more significant effects on size of age 2 year parr than conditions encountered the previous year by age 1 year parr of the same cohort. Fork lengths of parr were significantly and negatively associated with spring air and water temperatures. In the Miramichi River, increases in air and water temperature as predicted from climate change models may adversely affect growth of juvenile Atlantic salmon parr, reducing the overall productivity of the Atlantic salmon populations in this region.     

Moon phase and nocturnal density of Atlantic salmon parr in the Sainte‐Marguerite River, Québec

Nocturnal underwater counts of Atlantic salmon Salmo salar parr were made on four consecutive occasions (two lunar cycles, samplings at both the full and new moon) at four 40 m long sites in the Sainte‐Marguerite River, Québec, Canada, between 30 June and 14 August 2003. Atlantic salmon parr counts did not differ significantly between moon phases. Cloud cover ranged from 0 to 100% during the study, and had no significant effect on parr counts. There were significantly more Atlantic salmon parr in the near shore than in the midstream areas. The findings of this study suggest that the sampling strategy of summer studies aimed at assessing population abundance or developing habitat quality models can be designed without taking moon phase or cloud cover into consideration, but it should account for the higher relative abundance of Atlantic salmon parr in the near shore areas as compared to areas closer to the middle of streams.     

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.     

Occurrence of Atlantic salmon parr in redds at spawning time

Several Atlantic salmon parr of different sizes were found alive in salmon redds under a 10–30 cm layer of stones at spawning time. Parr, both mature and immature, remain inactive in redds during daylight but show nocturnal activity.     

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.     

Basin‐scale phenology and effects of climate variability on global timing of initial seaward migration of Atlantic salmon (Salmo salar)

Migrations between different habitats are key events in the lives of many organisms. Such movements involve annually recurring travel over long distances usually triggered by seasonal changes in the environment. Often, the migration is associated with travel to or from reproduction areas to regions of growth. Young anadromous Atlantic salmon (Salmo salar) emigrate from freshwater nursery areas during spring and early summer to feed and grow in the North Atlantic Ocean. The transition from the freshwater (‘parr’) stage to the migratory stage where they descend streams and enter salt water (‘smolt’) is characterized by morphological, physiological and behavioural changes where the timing of this parr‐smolt transition is cued by photoperiod and water temperature. Environmental conditions in the freshwater habitat control the downstream migration and contribute to within‐ and among‐river variation in migratory timing. Moreover, the timing of the freshwater emigration has likely evolved to meet environmental conditions in the ocean as these affect growth and survival of the post‐smolts. Using generalized additive mixed‐effects modelling, we analysed spatio‐temporal variations in the dates of downstream smolt migration in 67 rivers throughout the North Atlantic during the last five decades and found that migrations were earlier in populations in the east than the west. After accounting for this spatial effect, the initiation of the downstream migration among rivers was positively associated with freshwater temperatures, up to about 10 °C and levelling off at higher values, and with sea‐surface temperatures. Earlier migration occurred when river discharge levels were low but increasing. On average, the initiation of the smolt seaward migration has occurred 2.5 days earlier per decade throughout the basin of the North Atlantic. This shift in phenology matches changes in air, river, and ocean temperatures, suggesting that Atlantic salmon emigration is responding to the current global climate changes.