Habitat use of Atlantic salmon Salmo salar parr in a dynamic winter environment: the influence of anchor‐ice dams

The effect of anchor‐ice dams on the physical habitat and behavioural responses of Atlantic salmon Salmo salar parr in a small, steep stream was investigated. Anchor‐ice dams formed periodically, leading to a dynamic winter environment as the study reach alternated between riffle and walk dominated habitat. Parr demonstrated large individual variation in habitat use, utilizing most of the wetted stream width, and were generally unaffected by diel changes in the mesohabitat composition. Furthermore, parr displayed high site fidelity in areas with low embedded substrata, and demonstrated few large movements between the three mesohabitat classes present: shallow riffle, walk and pool. Findings from this study question the importance of hydraulic variables such as water depth, flow velocity and dynamic ice formation as single habitat features for juvenile stream salmonids during winter and emphasize the importance of access to substratum cover.     

Mid-winter activity and movement of Atlantic salmon parr during ice formation events in a Norwegian regulated river

A telemetry study in a Norwegian regulated river was conducted through a 12-day period in mid-winter 2003. The objective was to study activity (defined as number of movement per hour) and movement (defined as distance moved per hour) during different ice formation events. Twenty-four Atlantic salmon (Salmo salar L.) parr were radio tagged and continuously monitored by both manually tracking (N = 24) and by fixed recording stations (N = 15). Detailed data on climate, flow and ice formation and its spatial distribution were collected and used in the analyses. Fish activity was not found to be affected by their size (L _F). There was a significant difference in activity between diel periods with highest activity during dusk (5–6 p.m.). Between high and low flow (mean ± SD, 21.1 m³ s⁻¹ ± 1.7 SD and 11.1 m³ s⁻¹ ± 1.7 SD, respectively) no significant difference in activity was found. During the experiment extensive anchor ice growth occurred mainly in the riffle part with thickness up to 50 cm. Juveniles tend to avoid riffle section during anchor ice formation and exploited ice covered areas, indicating critical and preferable habitats respectively. Further, a significant difference in movement was found between five selected ice events with highest mean movement during an anchor ice event and lowest mean movement during an ice break up with no anchor ice formation. No significant difference in activity or movement between parr exposed to frazil ice and parr not exposed were found.     

Is Atlantic salmon production limited by number of territories?

Thirty Atlantic salmon Salmo salar parr (mean 135 mm total length, L _T) in the River Alta, Norway, were radio‐tagged and tracked during a 11–13 day period. The parr stayed within defined home ranges with a 95% probability of localization within an average area of 1286 m²(241–3484 m²). On average, each parr had overlapping home ranges with 7·7 other parr, and the overlap between pairs of fish covered on average 24% of their home range areas. Mean length of the river stretch used was 90 m (22–383 m). On average, during 38%(16–77%) of the tracking surveys, the fish had moved >10 m since the previous survey. Mean total distance moved during the whole study was 402 m (208–862 m). The Atlantic salmon were most often recorded in riffles, but 27% of the parr alternated between riffles and pools. The extensive movements, flexible habitat utilization and relatively large home ranges, together with the fact that all the parr had home ranges partly overlapping with other radio‐tagged parr, indicate a flexible and variable behaviour and, thereby, a more complex social structure than a rigid defence of a fixed territory. Thus, Atlantic salmon production seems not to be limited by the highest potential number of territories for large parr in a given area.     

Static habitat partitioning and dynamic selection by sympatric young Atlantic salmon and brown trout in south-west England streams

Direct underwater observation of micro‐habitat use by 1838 young Atlantic salmon Salmo salar [mean L_T 7·9 ± 3.1(s.d.) cm, range 3·19] and 1227 brown trout Salmo trutta (L_T 10·9 ± 5·0 cm, range 3·56) showed both species were selective in habitat use, with differences between species and fish size. Atlantic salmon and brown trout selected relatively narrow ranges for the two micro‐habitat variables snout water velocity and height above bottom, but with differences between size‐classes. The smaller fishes <7 cm held positions in slower water closer to the bottom. On a larger scale, the Atlantic salmon more often used shallower stream areas, compared with brown trout. The larger parr preferred the deeper stream areas. Atlantic salmon used higher and slightly more variable mean water velocities than brown trout. Substrata used by the two species were similar. Finer substrata, although variable, were selected at the snout position, and differences were pronounced between size‐classes. On a meso‐habitat scale, brown trout were more frequently observed in slow pool‐glide habitats, while young Atlantic salmon favoured the faster high‐gradient meso‐habitats. Small juveniles <7 cm of both species were observed most frequently in riffle‐chute habitats. Atlantic salmon and brown trout segregated with respect to use of habitat, but considerable niche overlap between species indicated competitive interactions. In particular, for small fishes <7 cm of the two species, there was almost complete niche overlap for use of water depth, while they segregated with respect to water velocity. Habitat suitability indices developed for both species for mean water velocity and water depth, tended to have their optimum at lower values compared with previous studies in larger streams, with Atlantic salmon parr in the small streams occupying the same habitat as favoured by brown trout in larger streams. The data indicate both species may be flexible in their habitat selection depending on habitat availability. Species‐specific habitat overlap between streams may be complete. However, between‐species habitat partitioning remains similar.     

Movement and habitat shift responses of juvenile Atlantic Salmon (Salmo salar) to annually permanent stream flooding

Reservoir formation in a river system changes a lotic environment to more lacustrine conditions, with impacts throughout the ecosystem. In this study, a river reach containing typical salmonid riffle/run habitat was flooded to create a large, deep pool from June to September in each of 3 years. We test the hypothesis that juvenile Atlantic salmon (Salmo salar) with their preference for run/riffle habitats will respond to the transformation to a lentic environment by moving into adjacent lotic environments. Movements of juvenile Atlantic salmon were monitored using a combination of biotelemetry (radio- and passive integrated transponder-tagging) and electrofishing. Results showed that no tracked fish moved away from the created pool habitat. Mass-specific growth rates showed the created pool habitat resulted in net growth of juveniles. The results confirm that fish may not immediately (i.e., at least for an approximate 2 months) respond to rapid, large-scale habitat alterations by moving to find similar habitat conditions outside the altered habitat. This is most probably related to plasticity of behavior and habitat use, and no change in biological conditions to a point that would negatively impact fish growth and survival, for example food availability, competition, or predation. The results also support the hypothesis that the relative importance of physical habitat variables is not universal among streams and populations, therefore limiting the value of applying standard habitat suitability criteria and use.     

A synopsis of ‘hydropeaking’ studies on the response of juvenile Atlantic salmon to experimental flow alteration

In insular Newfoundland, Canada, studies were conducted from 1999 to 2003 on the effects of ‘simulated’ hydropeaking power generation on juvenile Atlantic salmon (Salmo salar). In 1999, Atlantic salmon parr were released into an experimental reach below a hydroelectric facility and flow was manipulated over a range of discharge (1.0–4.2 m³ s⁻¹) during a series of ‘experiments’ simulating hydropeaking in both summer and fall. Fish were implanted with radio transmitters, manually tracked, precisely located (±1 m), habitat selection evaluated, and movement response determined. Experiments were continued in 2002 and 2003 to contrast response of salmon between summer and winter, the magnitude of flow changes were greater (0.7–5.2 m³ s⁻¹) and changes were made more rapidly (instantly). As discharge was increased, velocity and depth use by parr increased, and fish adapted behaviourally by increased contact with the substrate. Salmon parr also exhibited two distinct movement patterns in the summer and fall of 1999 studies; high site fidelity or considerable movement during trials. Salmon were more mobile during both static and dynamic flow conditions and throughout the diel cycle in the summer of 2002 experiment, and 2 fish were stranded and died, the only time this happened in the four series of experiments. Within each experiment generally there were no differences between movements at static high and low flows for day and night movements, with one exception, and night time movements were always greatest, again with one exception. During dynamic flow changes, within each experiment, distances moved during down ramping and up ramping were not significantly different except in the summer of 1999. Overall, comparing between experiments for up and down ramping events, distances moved in the summer of 2002 were statistically higher than for all other experiments. Not surprisingly, the home ranges of fish in the summer of 2002 were also the greatest while the smallest home ranges were in the winter of 2003. Results suggest hydropeaking regimes may be energetically costly potentially affecting over-winter survival which is related to energy reserves obtained during summer. Collectively these studies provide comprehensive information on the response of juvenile Atlantic salmon parr to hydropeaking, on both diel and seasonal scales, and will assist hydro producers and regulators design and operate hydropeaking regimes to minimize ecological impact. Guest editors: R. L. Welcomme & G. Marmulla Hydropower, Flood Control and Water Abstraction: Implications for Fish and Fisheries     

Interhabitat and instream movements of large Atlantic salmon parr in a Newfoundland watershed in winter

Radio‐telemetry was used to investigate movement of large, mainly mature male (80%) Atlantic salmon Salmo salar parr in Stoney River, Newfoundland during early winter (November; water temperature 6·0 ± 0·1° C) and mid‐winter (January to February; 0·8 ± 0·0° C). Site fidelity of parr in early winter was low. Parr moved between fluvial and lacustrine habitats and were active throughout the diel cycle. Parr caught in fluvial habitats in mid‐winter were smaller and younger than parr caught in early winter. Site fidelity of parr in mid‐winter was greater than in early winter. Parr in mid‐winter moved between fluvial and adjacent small lacustrine habitats, but avoided a larger pond inhabited by large piscivorous fishes. Instream movement rates in mid‐winter were lower than in early winter and occurred primarily during hours of darkness (dawn, dusk and night). Fluvial habitats were relatively stable and ice‐free throughout the study periods. These results suggested that large Atlantic salmon parr utilize a variety of habitats and remain active throughout the winter, even under stable environmental conditions.     

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.     

Behavioural strategies for species co‐existence during environmental extremes: competition between trout and salmon parr during dewatering

Competition between species of animals can be predicted to be intense under extreme environmental conditions during which niche overlap increases. Fluvial aquatic systems fluctuate naturally across a broad range of time‐scales. Land management activities impose further extreme fluctuations, particularly when water is abstracted for irrigation during summer. This study focused on the interaction between Atlantic salmon and brown trout parr during acute dewatering events. Brown trout are known to compete strongly for pool habitat whereas salmon can coexist by using riffle areas during normal flows. It is not known, however, how competition between the species affects their behavioural responses to extreme low flows when riffle areas decrease. Replicated groups of salmon were held in allopatry and sympatry with trout in sections of a large indoor stream, each of which was landscaped into riffles and pools. Space use and behaviours of the fish were recorded by direct visual observations and a network of Passive Integrated Transponder detectors. Here we report the response of the fish to dewatering and consider the system as a model for natural and forced responses of communities to environmental extremes.     

Swimming performance of various freshwater Newfoundland salmonids relative to habitat selection and fishway design

Swimming ability of wild brook trout Salvelinus fontinalis , brown trout Salmo trutta , anadromous Atlantic salmon Salmo salar , and landlocked Atlantic salmon was examined using fixed and increasing velocity tests. Although brook trout and salmon parr were collected from the same site, brook trout were found generally in slow-moving pools whereas salmon were more common in faster riffle areas. Salmon parr could hold station indefinitely in currents in which brook trout could only maintain themselves briefly. Therefore, selection of fast-water areas by salmon parr may impose a velocity barrier to sympatric juvenile brook trout, reducing competition between the species. Performance comparisons also indicate that anadromous Atlantic salmon possess slightly greater sustained ability than landlocked salmon, possibly due to altered selective pressure associated with their different life histories. Finally, fishways and culverts in Newfoundland can now be designed using models generated from performance data collected from native salmonid species.     

Summer stream habitat partitioning by sympatric Arctic charr, Atlantic salmon and brown trout in two sub-arctic rivers

Summer habitat use by sympatric Arctic charr Salvelinus alpinus, young Atlantic salmon Salmo salar and brown trout Salmo trutta was studied by two methods, direct underwater observation and electrofishing, across a range of habitats in two sub-arctic rivers. More Arctic charr and fewer Atlantic salmon parr were observed by electrofishing in comparison to direct underwater observation, perhaps suggesting a more cryptic behaviour by Arctic charr. The three species segregated in habitat use. Arctic charr, as found by direct underwater observation, most frequently used slow (mean ±s .d . water velocity 7·2 ± 16·6 cm s⁻¹) or often stillwater and deep habitats (mean ±s .d . depth 170·1 ± 72·1 cm). The most frequently used mesohabitat type was a pool. Young Atlantic salmon favoured the faster flowing areas (mean ±s .d . water velocity 44·0 ± 16·8 cm s⁻¹ and depth 57·1 ± 19·0 cm), while brown trout occupied intermediate habitats (mean ±s .d . water velocity 33·1 ± 18·6 cm s⁻¹ and depth 50·2 ± 18·0 cm). Niche overlap was considerable. The Arctic charr observed were on average larger (total length) than Atlantic salmon and brown trout (mean ±s .d . 21·9 ± 8·0, 10·2 ± 3·1 and 13·4 ± 4·5 cm). Similar habitat segregation between Atlantic salmon and brown trout was found by electrofishing, but more fishes were observed in shallower habitats. Electrofishing suggested that Arctic charr occupied habitats similar to brown trout. These results, however, are biased because electrofishing was inefficient in the slow-deep habitat favoured by Arctic charr. Habitat use changed between day and night in a similar way for all three species. At night, fishes held positions closer to the bottom than in the day and were more often observed in shallower stream areas mostly with lower water velocities and finer substrata. The observed habitat segregation is probably the result of interference competition, but the influence of innate selective differences needs more study.     

Juvenile Anadromous Atlantic Salmon of Three Lakes of Newfoundland

Anadromous Atlantic salmon parr (Salmo salar) were captured from three Newfoundland lakes. Yearling and 2 year parr were captured most often in shallow water <2 m deep around lake perimeters and were not abundant at greater depths. The 3+ and 4+ parr age groups were rarely found inshore (<2 m) but were captured in deeper offshore areas. Though the parr population in one deep lake was too small to be estimated by mark-recapture, estimates of the yearlings plus 2+ age groups in two shallower lakes were 55 and 63 parr ha⁻¹. respectively. No under yearlings were found in lake habitats. Parr density in the three study lakes varied inversely with mean lake depth. Lacustrine parr growth rate was greater than or equal to that of stream dwelling parr. It is concluded that some shallow lakes of Newfoundland provide major rearing areas for juvenile anadromous Atlantic salmon.     

The evolution of cooperation and advanced social behaviour

Competition between species of animals can be predicted to be intense under extreme environmental conditions during which niche overlap increases. Fluvial aquatic systems fluctuate naturally across a broad range of time‐scales. Land management activities impose further extreme fluctuations, particularly when water is abstracted for irrigation during summer. This study focused on the interaction between Atlantic salmon and brown trout parr during acute dewatering events. Brown trout are known to compete strongly for pool habitat whereas salmon can coexist by using riffle areas during normal flows. It is not known, however, how competition between the species affects their behavioural responses to extreme low flows when riffle areas decrease. Replicated groups of salmon were held in allopatry and sympatry with trout in sections of a large indoor stream, each of which was landscaped into riffles and pools. Space use and behaviours of the fish were recorded by direct visual observations and a network of Passive Integrated Transponder detectors. Here we report the response of the fish to dewatering and consider the system as a model for natural and forced responses of communities to environmental extremes.     

Impact of Beaver Dams on Abundance and Distribution of Anadromous Salmonids in Two Lowland Streams in Lithuania

European beaver dams impeded movements of anadromous salmonids as it was established by fishing survey, fish tagging and redd counts in two lowland streams in Lithuania. Significant differences in abundancies of other litophilic fish species and evenness of representation by species in the community were detected upstream and downstream of the beaver dams. Sea trout parr marked with RFID tags passed through several successive beaver dams in upstream direction, but no tagged fish were detected above the uppermost dam. Increase in abundances of salmonid parr in the stream between the beaver dams and decrease below the dams were recorded in November, at the time of spawning of Atlantic salmon and sea trout, but no significant changes were detected in the sections upstream of the dams. After construction of several additional beaver dams in the downstream sections of the studied streams, abundance of Atlantic salmon parr downstream of the dams decreased considerably in comparison with that estimated before construction.     

Metabolic recovery in Atlantic salmon fry and parr following forced activity

Atlantic salmon Salmo salar fry and parr were subjected to 5 min of forced activity and the subsequent changes in oxygen consumption and ammonia excretion rates were evaluated over a 24 h period. In a second experiment, individual Atlantic salmon fry and parr were freeze‐clamped in liquid nitrogen, before, immediately following a 5 min activity period, or after periods of recovery up to 2 h. Samples were analysed for whole body phosphocreatine (PCr), ATP and lactate. Five minutes of forced activity resulted in significant increases in both oxygen consumption and ammonia excretion rates. Changes in the oxygen consumption rates were greater in the parr compared with the fry. In contrast, the post‐exercise ammonia excretion rates were nearly twice as high for the fry compared with the parr. Exercise also caused a marked decrease in PCr levels (c. 47 and 65% in fry and parr, respectively), no change in ATP levels and a significant increase in lactate levels in Atlantic salmon fry and parr. Recovery of PCr occurred quickly (between 15 and 30 min) in fry and parr. Although the post‐activity levels of lactate were lower in fry (c. 3 μmol g^?1) compared with parr (c. 14 μmol g^?1), lactate levels returned to control levels within 60 min in fry, but it took >2 h for this metabolite to recover in parr. Compared with parr, these findings show that Atlantic salmon fry possess a reduced anaerobic capacity, and these results are consistent with the theoretical and experimental evidence that smaller fish support burst swimming through aerobic processes.     

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.     

Habitat-specific estimates of competition in stream salmonids: A field test of the isodar model of habitat selection

Summary The population densities of sympatric Atlantic salmon,Salmo salar and brook charr,Salvelinus fontinalis, were measured in riffle and pool stream habitats to test whether non-linear isodars, a multispecific model of habitat selection based on ideal distribution assumptions, could (1) predict the distribution of densities between habitats and (2) reproduce the processes postulated to underlie spatial segregation and species interactions in previous laboratory and field studies. The model provided a good fit to observed density patterns and indicated that habitat suitability declined non-linearly with increased heterospecific competitor densities. Competitive effects in riffles appeared to be due to exploitative resource use, with salmon always emerging as the superior competitor. No evidence was found for interference competition in riffles. In contrast, interspecific competition in pools seemed to occur through exploitation and interference. The specific identity of the superior competitor in pools depended on the density of both species; pools provided the charr with refuge from competition with the salmon, presumably through the adoption by the charr of density-dependent behaviours, such as schooling and group foraging, that mitigated the negative impact of the salmon. Charr were displaced from the riffles toward the pools as the total salmon density increased. The isodar analysis, based on limited density data, successfully reproduced the processes suggested to underlie spatial segregation in previous field and laboratory studies and provided new insights into how changes in competitor densities modify habitat suitability in this system.     

Visiting the hyporheic zone: young Atlantic salmon move through the substratum

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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.     

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.