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.     

Mitigating the effects of high-head dams on the Columbia River, USA: experience from the trenches

Worldwide, humans have tremendously altered freshwater ecosystems and arguably, construction of dams has had the greatest effect. Maintaining natural ecological processes and developing mitigation strategies that will maintain species while retaining dam benefits is challenging. In the Columbia River, USA, over the last 30 years more than US$7 billion has been spent on efforts to save historically large runs of salmon. These efforts have included improving passage conditions at dams through construction of efficient fish ladders for adult salmon, effective fish passage facilities for downstream migrating juvenile salmon, voluntarily spilling water to decrease the number of downstream migrants that pass through turbines, modifying dam operations to provide more constant flow and providing additional flow from storage reservoirs to create more natural flow through areas inundated by dams. Construction of hatcheries to offset losses in habitat for wild fish has also occurred. Further, for salmon from the Snake River, the largest tributary to the Columbia River, a large percent of juvenile salmon smolts are collected at upstream dams and transported in barges to the lower river to avoid passage through dams, turbines, and reservoirs. Experiences in the Columbia River suggest that the sum of all of these actions may keep salmon stocks from going extinct, but the technological fixes will not likely provide complete mitigation for altered freshwater ecosystems. Guest editors: R. L. Welcomme & G. Marmulla Hydropower, Flood Control and Water Abstraction: Implications for Fish and Fisheries     

Size-related differences in behaviour and spatial distribution of juvenile Atlantic salmon in a novel environment

Within a month of first feeding sibling juvenile salmon had developed not only large differences in size (up to fivefold in weight), but also significant differences in behaviour. Large fish remained in refuges (overhead cover) significantly longer after having been frightened. On leaving the shelters most small fish established individual territories on the bottom and. downstream of the shelters. The greater aggressiveness of small fish in a novel environment (from holding tank to experimental stream) was related to this apparent territoriality. The increased shelter residence time and delayed establishment of individual territories might be interpreted as greater caution in large fish, which are able to avoid risky situations due to their higher energy reserves. In contrast, smaller fish need to leave refuges more quickly and to establish individual territories in a novel environment.     

An Approach to Assessing Hydrological Influences on Feeding Opportunities of Juvenile Atlantic Salmon (<Emphasis Type="Italic">Salmo salar</Emphasis>): A Case Study of Two Contrasting years in a Small,Nursery Stream

This case study sought to examine how temporal variability in hydrological and hydraulic conditions might affect the feeding opportunities of juvenile Atlantic salmon in two hydrologically contrasting years of 2002 and 2003, which were characterised by high and low flows respectively. Firstly, measures of hydraulic influence were calculated to define what might be ecologically meaningful disturbance periods during high flows. Secondly, for identifying such periods, the parameter Critical Displacement Velocity (CDV) was derived from a river discharge time series as a first approximation of the amount of time in two hydrologically extreme years when fish foraging strategies for specific age classes of juvenile Atlantic salmon (Salmo salar) might be disrupted by flows. The CDV estimates the threshold velocity above which juvenile salmon are unable to hold station and it is dependent upon fish size and stream temperature. In the wet year 2002, the CDV was exceeded on 18% and 21% of days for 0+ and 1+ fish respectively. In 2003 these respective numbers fell to 6% and 15%. The data suggest that hydrological conditions during certain times of the year have the potential to affect foraging behaviour. This in turn might have implications for recruitment and growth rates for juvenile salmon in upland streams.     

Use of glacier river-fed estuary channels by juvenile Coho Salmon: transitional or rearing habitats?

Estuaries are among the most productive ecosystems in the world and provide important rearing environments for a variety of fish species. Though generally considered important transitional habitats for smolting salmon, little is known about the role that estuaries serve for rearing and the environmental conditions important for salmon. We illustrate how juvenile coho salmon Oncorhynchus kisutch use a glacial river-fed estuary based on examination of spatial and seasonal variability in patterns of abundance, fish size, age structure, condition, and local habitat use. Fish abundance was greater in deeper channels with cooler and less variable temperatures, and these habitats were consistently occupied throughout the season. Variability in channel depth and water temperature was negatively associated with fish abundance. Fish size was negatively related to site distance from the upper extent of the tidal influence, while fish condition did not relate to channel location within the estuary ecotone. Our work demonstrates the potential this glacially-fed estuary serves as both transitional and rearing habitat for juvenile coho salmon during smolt emigration to the ocean, and patterns of fish distribution within the estuary correspond to environmental conditions.     

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.     

Displacement,velocity preference,and substrate use of three native California stream fishes in simulated pulsed flows

We studied whether juvenile fishes were able to maintain swimming speed and position during simulated river pulsed flows in a laboratory flume. We used a glass flume (15.24 × 0.6 m) with river-rock substrate to determine the longitudinal displacement, movement distances and frequencies, velocity selection, and substrate use of juvenile (SL range: 6.1 ± 0.2 cm) hardhead Mylopharodon conocephalus (n = 13), rainbow trout Oncorhynchus mykiss (n = 11), and Sacramento sucker Catostomus occidentalis (n = 12) during a 100-min flow pulse, as velocity changed from slow to medium, fast, medium, and slow. Fish were capable of maintaining swimming speed and position up to the maximum flume velocity of 0.46 m·s⁻¹, except for one hardhead that impinged on the rear fish screen. Fish swam faster in the flume during the medium and fast intervals than the slow intervals, but fish speeds were similar among the medium and faster intervals, when some fish took cover behind the rock substrate. In comparison with a Brett-type swim-tunnel, fish showed less increase in mean swimming speed as the flume velocity increased. Fish in the flume were able to use the rock substrate as hydraulic cover, decreasing the encountered water velocity, and, presumably, conserving energy.     

Ephemeral floodplain habitats provide best growth conditions for juvenile Chinook salmon in a California river

We reared juvenile Chinook salmon for two consecutive flood seasons within various habitats of the Cosumnes River and its floodplain to compare fish growth in river and floodplain habitats. Fish were placed in enclosures during times when wild salmon would naturally be rearing in floodplain habitats. We found significant differences in growth rates between salmon reared in floodplain and river enclosures. Salmon reared in seasonally inundated habitats with annual terrestrial vegetation experienced higher growth rates than those reared in a perennial pond on the floodplain. Growth of fish in the non-tidal river upstream of the floodplain varied with flow in the river. When flows were high, there was little growth and high mortality, but when the flows were low and clear, the fish grew rapidly. Fish displayed very poor growth in tidally influenced river habitat below the floodplain, a habitat type to which juveniles are commonly displaced during high flow events due to a lack of channel complexity in the main-stem river. Overall, ephemeral floodplain habitats supported higher growth rates for juvenile Chinook salmon than more permanent habitats in either the floodplain or river. Variable responses in both growth and mortality, however, indicate the importance of providing habitat complexity for juvenile salmon in floodplain reaches of streams, so fish can find optimal places for rearing under different flow conditions.     

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.     

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.     

Intra- and Interspecific Competition in Hatchery Landlocked Salmon and Brown Trout in Semi-Natural Streams

I studied inter- and intraspecific competition in two hatchery stocks: landlocked salmon with long-hatchery background and a heterogenic brown trout stock. These species are potential competitors in the natural environment when landlocked salmon is being restored to wild by stocking hatchery juveniles. Behavioural responses were studied in four indoor laboratory flumes (400 cm long and 37 cm wide) and habitat use in six semi-natural outdoor streams (26 m long and 1.5 m long). Video recordings were used to monitor fish behaviour and electrofishing for fish positioning in the outdoor channels. The study design included five treatments: two densities of brown trout and salmon in solitary and both species together. The results of the study demonstrated that juvenile brown trout changed their behaviour in laboratory streams in response to presence of the landlocked salmon and the density of the conspecifics also tended to alter the habitat use by brown trout in semi-natural streams. Landlocked salmon juveniles showed no response to treatments. I conclude that possible poor adaptive ability to conditions outside hatchery by the hatchery salmon together and more competitive brown trout stocks may limit the success of management action in restoring landlocked salmon back to their natural streams of stocking.     

The effects of turbidity and vegetation on the risk of juvenile salmonids, Oncorhynchus spp., to predation by adult cutthroat trout, O. clarkii

Synopsis During their seaward migration, juvenile salmonids encounter structural and visual cover which varies between and within watersheds. In this study, the effects of two types of cover (turbidity and artificial vegetation) on the predation mortality of juvenile salmonids exposed to fish piscivores was investigated in outdoor concrete ponds. During experiments, adult coastal cutthroat trout, Oncorhynchus clarkii clarkii, were allowed to feed on juvenile salmonid prey — chinook salmon, O. tshawytscha, chum salmon, O. keta, sockeye salmon, O. nerka, and cutthroat trout — in separate trials. Daily instantaneous per capita predation rate was determined for each turbidity and vegetation treatment, within each trial. Mean predation rates varied between 1% and 76% daily. In the presence of cover, mean daily predation rates were 10–75% lower than those in controls (no vegetation and clear water), depending on prey species. Predation rates were significantly lower in the presence of vegetation cover and did not covary with prey size or species. The effects of turbidity were generally not significant and were not additive with the effects of vegetation. However, turbidity appeared to significantly reduce the effectiveness of vegetation as cover for juvenile chinook and sockeye salmon. We suggest that these two forms of cover do not affect risk of predation by fish piscivores to juvenile salmonids via the same mechanism.     

Prior residence, territory quality and life-history strategies in juvenile Atlantic salmon (Salmo salar L.)

Three groups of juvenile salmon were introduced sequentially into an artificial stream to investigate the effects of prior residence on behaviour and territory choice. Almost half of the first group obtained and defended distinct territories, the other half being constrained to an area approximately the size of one large territory. All of the fish in the subsequent groups, bar one, were also constrained to the same site. Since the fish were of similar size, prior residence alone seemed to influence which individuals obtained territories. However, within the first group, the fish that obtained territories were larger and more aggressive. The territorial fish did not appear to choose the most profitable territories, although they had the greatest opportunity to do so. Since juvenile salmon emerge from their gravel nests fairly synchronously, a time constraint on site sampling is hypothesised: there may be a risk in taking time to sample sites, since these same sites may become occupied with conspecifics. However, fish with territories fed at faster rates than non-territorial fish, possibly because of reduced competition for prey items. Consequently, fish from the first group (containing most of the territorial fish) grew faster than the other two groups. Moreover, most of the territory holders, but only one of the non-territorial fish, reached the threshold size that increases their probability of smolting the following year. This suggests that ability to obtain a defensible territory, primarily through prior residence, influences the age at which juvenile salmon can migrate to sea.     

Development of length–bimodality and smolting in wild stocks of Atlantic salmon, Salmo salar L., under different growth conditions

Growth dynamics of juvenile Atlantic salmon, Salmo salar L., from two sections of the Narcea River and one of the Esva River (Northern Spain) were examined in relation to the development of bimodality in their size–frequency distributions. Size–bimodality was clearer under intermediate growth (section A) than under relatively fast or slow growth. The proportion of fish entering the upper modal group increase with growth intensity. Composition of upper and lower modal groups became fixed prior to December, and at this time both groups separated on the 90–95 mm interval. Fish exhibiting smolt appearance in late March (larger than 130 mm) had already been upper group fish in December, while parr-like fish and those that remained in the river by May (potential 2-year-old smolts) had formed the lower modal group. Anadromous salmon catch in the Narcea River was mostly of previously 1-year-old smolts (97.6%), of which 94% were larger than 100 mm by their first winter. In the Esva River, slow growth of juveniles is consistent with a large proportion of 2-year-old smolts (47.9%) among anadromous salmon. Both juvenile samples and scale analysis of anadromous salmon indicate that 2-year-old smolts were larger than 1-year-olds. Early disappearance of the former (before March) is, at least, partially related to earlier migration of large fish, since sexual maturity of parr does not provide a complete explanation. The Narcea stock have a minimum length at smolting of about 130 mm and an optimum smolt size in the 155–175 mm interval. Mean smolt length did not vary although the winter length changed between years.     

Comparison between PIT and radio telemetry to evaluate winter habitat use and activity patterns of juvenile Atlantic salmon and brown trout

Winter habitat use and activity patterns of juvenile Atlantic salmon and brown trout were analysed in a comparative study between Passive Integrated Transponder (PIT) technology, radio telemetry and underwater observation by snorkelling. Two study periods were conducted in Stoney River, Newfoundland, Canada. During Study period I, 49 juvenile Atlantic salmon (fork length: 11.0–18.0 cm) and 7 brown trout (11.0–17.3 cm) were tagged with PIT tags and/or radio transmitters in late winter of 2004. During Study period II, 18 juvenile Atlantic salmon (fork length: 12.0–18.4 cm) and 23 brown trout (10.9–20.8 cm) were tagged and tracked twice a day at 10:00 h and 22:00 h on five consecutive days in late winter of 2005. From the 56 fish released during Study period I, on average 19.6 ± 6.0% of the PIT tagged fish and 99.3 ± 2.2% of the radio tagged fish were relocated during any given survey. Over the Study period II, 39% of fish emigrated from the study site. PIT technology had an efficiency of 39.2 ± 14.1% to detect the remaining fish. In contrast, radio telemetry relocated on average 96.9 ± 6.5% of the tagged fish whereas by snorkelling on average only 4.1 ± 5.6% of the tagged fish were observed. PIT telemetry may however be more efficient in smaller, less heterogeneous streams. The advantage of PIT technology over radio telemetry is clearly that it is relatively less costly permitting higher numbers of individuals to be tagged and there is no limit in the operational life of the transponder. In winter, juvenile salmonids preferred low flow velocity and no preferences were observed for any specific water depth over the range of available water depths. Fish selected preferentially boulder habitat over other substrates in the environment. Habitat utilisation did not differ between day and night. The use of winter preference indices may be important for future habitat modelling.     

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.     

Sensitivity of survival to migration routes used by juvenile Chinook salmon to negotiate the Sacramento-San Joaquin River Delta

Populations of juvenile salmon emigrating from natal rivers to the ocean must often traverse different migratory pathways that may influence survival. In regulated rivers, migration routes may consist of a network of channels such as in the Sacramento-San Joaquin River Delta, or of different passage structures at hydroelectric dams (e.g., turbines or spillways). To increase overall survival, management actions in such systems often focus on altering the migration routing of fish to divert them away from low-survival routes and towards high-survival routes. Here, we use a 3-year data set of route-specific survival and movement of juvenile Chinook salmon in the Sacramento-San Joaquin Delta to quantify the sensitivity of survival to changes in migration routing at two major river junctions in the Sacramento River. Our analysis revealed that changes in overall survival in response to migration routing at one river junction depended not only differences in survival among alternative routes, but also on migration routing at the other river junction. Diverting fish away from a low-survival route at the downstream river junction increased population survival by less than expected, given the difference in survival among routes, because part of the population used an alternative migration route at the upstream river junction. We also show that management actions that influence only migration routing will likely increase survival by less than actions that alter both migration routing and route-specific survival. Our analysis provides an analytical framework to help fisheries managers quantify the suite of management actions likely to maximize increases in population level survival.     

Seaward migrating subyearling chinook salmon avoid overhead cover

Approximately three‐quarters of migrating autumn (fall) chinook salmon Oncorhynchus tshawytscha smolts avoided a covered channel and selected an uncovered channel when presented with a choice in an experimental flume. Rejection of overhead cover occurred prior to, but was rare post‐channel entrance. Smolts may selectively reject riparian cover as an adaptive behavioural response to minimize predation risk and enhance rates of migration. The findings have implications for fish bypass design and riparian habitat and culvert restoration.     

Comprehensive passage (COMPASS) model: a model of downstream migration and survival of juvenile salmonids through a hydropower system

Migratory fish populations are impacted worldwide by river impoundments. Efforts to restore populations will benefit from a clear understanding of survival and migration process over a wide-range of river conditions. We developed a model that estimates travel time and survival of migrating juvenile salmonids (Oncorhynchus spp.) through the impounded Snake and Columbia rivers in the northwestern United States. The model allows users to examine the effects of river management scenarios, such as manipulations of river flow and spill, on salmonid survival. It has four major components: dam passage and survival, reservoir survival, fish travel time, and hydrological processes. The probability that fish pass through specific routes at a dam and route-specific survival probabilities were based on hydroacoustic, radio telemetry, PIT tag, and acoustic tag data. We related reservoir mortality rate (per day and per km) to river flow, water temperature, and percentage of fish passing through spillways and then fit the relationships to PIT-tag survival data. We related fish migration rate to water velocity, percentage of fish passing through spillways, and date in the season. We applied the model to two threatened “Evolutionarily Significant Units” (as defined under the US Endangered Species Act): Snake River spring/summer Chinook salmon (O. tshawytscha Walbaum) and Snake River steelhead (O. mykiss Walbaum). A sensitivity analysis demonstrated that for both species survival through the hydropower system was responsive to water temperature, river flow, and spill proportion. The two species, however, exhibited different patterns in their response. Such information is crucial for managers to effectively restore migratory fish populations in regulated rivers. Guest editors: R. L. Welcomme & G. Marmulla Hydropower, Flood Control and Water Abstraction: Implications for Fish and Fisheries     

Juvenile growth and aggression in diploid and triploid Chinook salmon Oncorhynchus tshawytscha (Walbaum)

Multilocus heterozygosity, aggressive and feeding behaviour, plasma cortisol levels and growth rate were evaluated among three groups of juvenile Chinook salmon Oncorhynchus tshawytscha : diploid, triploid and mixed groups of diploid and triploid fish. There was no difference between diploid and triploid fish in measurements of heterozygosity calculated using seven microsatellite loci, and these measurements did not correlate with performance measurements including feeding rate and growth rate. Aggression trials that examined small groups of fish revealed that after 4 days together in tanks, triploid fish were significantly less aggressive during feeding than diploid fish or fish in mixed groups. At the end of the trials, however, plasma cortisol levels did not differ among the three groups. Thirty-day growth trials in duplicate tanks of 60 fish revealed no difference in growth rate among diploid, triploid and mixed groups, but plasma cortisol levels were significantly lower in triploid fish than in either diploid fish or the mixed fish. Overall, independent of the above differences in aggressive behaviour and cortisol levels, these results suggest similar performance in diploid and triploid Chinook salmon, and thus provide support for the viability of triploid Chinook salmon culture in commercial aquaculture.