Movements and habitat use by PIT-tagged Atlantic salmon parr in early winter: the influence of anchor ice

1. Movements and habitat use by Atlantic salmon parr in Catamaran Brook, New Brunswick, were studied using Passive Integrated Transponder technology. The fish were tagged in the summer of 1999, and a portable reading system was used to collect data on individual positions within a riffle‐pool sequence in the early winter of 1999. Two major freezing events occurred on November 11–12 (Ice 1) and November 18–19 (Ice 2) that generated significant accumulations of anchor ice in the riffle. 2. Individually tagged parr (fork length 8.4–12.6 cm, n = 15) were tracked from 8 to 24 November 1999. Over this period, emigration (40%) was higher from the pool than from the riffle. Of the nine parr that were consistently located, seven parr moved <5 m up‐ or downstream, and two parr moved more than 10 m (maximum 23 m). Parr moved significantly more by night than by day, and diel habitat shifts were more pronounced in the pool with some of the fish moving closer to the bank at night. 3. During Ice 2, there was relatively little movement by most of the parr in the riffle beneath anchor ice up to 10 cm in thickness. Water temperature was 0.16 °C above the freezing point beneath anchor ice, suggesting the existence of suitable habitats where salmon parr can avoid supercooling conditions and where they can have access to low velocity shelters. To our knowledge, these are the first data on habitat use by Atlantic salmon parr under anchor ice.     

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.     

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.     

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.     

Functions of restored wetlands for juvenile salmon in an industrialized estuary

The Duwamish estuary is an industrialized waterway located in Seattle, WA, USA. Despite a history of habitat loss, naturally produced juvenile Chinook salmon use the estuary. In addition to experiencing degraded habitat in the estuary, wild salmon growth may be affected by competition with more than three million hatchery fish released yearly into the river. Restoring habitat to benefit salmon in the Duwamish River is a priority for trustees of public resources, and a number of wetland restoration sites have been created there. We tested the function of restored sites in the Duwamish estuary for juvenile Chinook salmon by comparing fish densities from enclosure nets or beach seines at three paired restored/un-restored sites and by applying environmental and diet data to a bioenergetics model. We also examined temporal and diet overlap of wild juvenile Chinook salmon with other salmon species and with hatchery-reared Chinook salmon using non-metric multidimensional scaling (NMDS). At a brackish upstream site with a relatively large opening to the river, we found higher densities of juvenile Chinook salmon at the restored site. NMDS results indicated that juvenile Chinook salmon fed on different taxa at the restored sites than at the reference sites. However, modeled growth was similar at restored and reference sites. Co-occurring juvenile chum and Chinook salmon fed differently, with chum eating smaller prey, and Chinook salmon eating larger prey. Co-occurring hatchery and wild juvenile Chinook salmon had similar diets, indicating that they may compete for prey. However, modeled growth was positive and did not differ between hatchery and wild fish, suggesting that food was not limiting. Bioenergetics models indicated that overall juvenile Chinook salmon growth potential at the brackish water site was consistently higher than at more saline sites. Our results suggest that restoration sites in the Duwamish estuary that have larger access openings and are located in brackish water may have increased function over other configurations.     

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.     

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.     

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

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

Does life-history variability in salmonids affect habitat use by juveniles? A comparison among streams open and closed to anadromy

1. Migratory and resident forms of salmonids coexist in many river systems. Although such coexistence is widespread, little is known about its ecological basis and no studies have compared the habitat use of premigratory juveniles and residents. 2. We employed a comparative approach to explore the differential habitat use of juvenile anadromous and resident brook trout. This required the investigation of habitat use in streams closed to anadromy, containing only resident brook trout Salvelinus fontinalis ('resident-only' streams) and streams open to anadromy, containing coexisting Atlantic salmon Salmo salar and anadromous and resident brook trout ('migrant-resident' streams). 3. We demonstrate that fast habitats (riffles) are occupied more frequently in streams with migratory brook trout relative to riffle habitats of streams with only resident brook trout. In contrast, occupation of slow current velocities (pools) was observed in both migrant-resident and resident-only streams as both stream types contain resident brook trout. The net effect is a wider distribution of occupied habitats (pool and riffles) in migrant-resident streams relative to resident-only streams, resulting in few, if any, unused habitats. 4. These results are consistent with previously reported bioenergetic, morphological and stable isotope differences observed between anadromous and resident brook trout. 5. Our findings suggest that a link exists between juvenile habitat use, metabolic costs and life-history strategies.     

Relationships between fishes and habitat in rainforest streams in Sabah, Malaysia

Distinct fish assemblages were found at the mesohabitat scale in 14 streams in eastern Sabah, Malaysia. Sites were designated a priori as pool, run or riffle on the basis of physical habitat structure and properties. Principal components analysis of physical habitat data confirmed the validity of the a priori designation with a major axis of three correlated variables: water velocity, depth and substratum type. Canonical discriminant analysis on fish abundance and biomass data confirmed the existence of a specialized assemblage of fishes from riffle areas of all streams. Overall, pool and run assemblages were highly variable, dependent on stream size, but also variable between streams of the same size. Multiple regression of species richness, diversity, abundance and biomass data on principal components revealed significant but low correlations with measured habitat variables. Riffle habitats showed lower species richness and diversity but high abundance. The fish assemblage in riffles was dominated by balitorid species, specialized for fast-water conditions. Pool assemblages had the highest species diversity and were dominated by cyprinid species of a number of morphological and ecological guilds. Run assemblages were intermediate in assemblage characteristics between riffle and pool assemblages. Between-stream variation in assemblage composition was less than within-stream variation. Of 38 species collected, seven could be designated as riffle specialists, 18 as pool specialists and 13 as ubiquitous, although most of the latter showed size-specific habitat use with larger size classes found in slower, deeper water.     

Visual isolation furthers access to drift‐feeding positions for subordinate juvenile brown trout (Salmo trutta) in dominance hierarchies

Juvenile salmonids are visual predators that primarily feed on drifting invertebrates and compete for suitable feeding positions in swift water. We used an outdoor experimental stream to test the effect of visual isolation on agonistic interactions and habitat use by age‐1 brown trout ( Salmo trutta ) in riffle‐pool sections. We hypothesized that dominant fish defend suitable feeding positions in riffle and that visual obstruction between individuals enhance access to riffle for subordinates. Groups of juveniles, caught in the wild, were stocked in high and low visibility treatments. Visual isolation was manipulated by placing dark plastic ribbons or opaque Plexiglas boards onto the substrate of riffles. As expected, dominant fish held profitable positions in riffle and the proportion of fish in riffle significantly increased in presence of artificial structures. In high visibility treatment, the dominant fish despotically excluded subordinates from the riffle. In low visibility treatment, the proportion of fish that foraged on benthic prey in the pool and the number of major aggressive acts (chase, nip) decreased. Our results support the hypothesis that screening effect of physical structures in the water column loosen resource monopolization in dominance hierarchies of juvenile salmonids.     

Juvenile salmon in estuaries: comparisons between North American Atlantic and Pacific salmon populations

All anadromous fishes, including juvenile salmon, encounter estuarine habitats as they transition from riverine to marine environments. We compare the estuarine use between juvenile Atlantic salmon (Salmo salar) in the Penobscot River estuary and Pacific salmon (Oncorhynchus spp.) in the Columbia River estuary. Both estuaries have been degraded by anthropogenic activities. Atlantic and Pacific salmon populations in both basins rely heavily on hatchery inputs for persistence. Pacific salmon, as a group, represent a continuum of estuarine use, from species that move through rapidly to those that make extensive use of estuarine habitats. While Atlantic salmon estuarine use is predominantly similar to rapidly moving Pacific salmon, they can exhibit nearly the entire range of Pacific salmon estuarine use. Both slow and rapidly migrating Atlantic and Pacific salmon actively feed in estuarine environments, consuming insect and invertebrate prey. Interactions between juvenile salmon and estuarine fish communities are poorly understood in both estuaries, although they experience similar avian and marine mammal predators. Estuaries are clearly important for Atlantic and Pacific salmon, yet our understanding of this use is currently insufficient to make informed judgments about habitat quality or overall estuary health. This review of salmonid migration through and residency within estuaries identifies actions that could hasten restoration of both Atlantic and Pacific salmon populations.     

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.     

Return migration of Atlantic salmon in the River Tana: the role of environmental factors

Multi-sea-winter Atlantic salmon (75–115 cm fork length L _F, 2–4-winter fish) were radio-tagged in the Tanafjord (700 N), Norway, in 1992–1993, and 130 fish entered the large subarctic River Tana (Teno). They entered the fresh water at any time of the tidal cycle but more so during the high and ebbing tides. No diel rhythm was detected in river entry under polar day conditions. There were no differences in the change of flow between days when salmon moved and when they did not, but during active migration increasing discharge was associated with increased swimming activity of salmon, especially later in the summer. Increasing air temperature was also associated with enhanced migration activity. Low river flow was associated with increasing delay in salmon passing the first riffle area of the river, 35 km from the sea.     

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.     

On the interaction of turbulent flow and feeding behaviour of juvenile Atlantic Salmon (Salmo salar)

Anthropogenic activities can dramatically modify the riverine habitat of juvenile Atlantic salmon ( Salmo salar ). In the perspective of protecting and restoring the fluvial habitat, bioenergetic models are often used to estimate fish habitat quality. These models determine the habitat quality as the ratio between the energetic gains (food) and costs (growth, metabolism) of a fish. The energetic costs of swimming in a river are generally estimated using the average flow velocity without consideration of the effect of turbulence. Juvenile Atlantic salmon (JAS) live in rivers characterized by intense velocity fluctuations, which are often described as a succession of high‐ and low‐speed flow regions. These flow structures are likely to affect the JAS activity that consists of long periods of sit‐and‐wait at the top of a protuberant rock interrupted by short bursting motions to capture drifting food particles. To minimize the energetic costs, it is hypothesized that JAS use low‐speed flow regions to initiate and undertake their feeding motions.  To improve bioenergetic modelling, this study aimed at analyzing the relation between turbulent flow structures and the feeding behaviour of JAS in a natural gravel‐bed river. We filmed eight JAS during 30 min with a submersible video camera while simultaneously measuring velocity fluctuations close to the fish in the St. Marguerite River, Quebec, Canada. Our results show that the proportion of time used for feeding motions decreases with increasing turbulent intensity and mean flow velocity; and that JAS do not seem to prefer low‐speed flow regions to initiate their feeding motions.     

The performance advantage of a high resting metabolic rate in juvenile salmon is habitat dependent

1.?Basal levels of metabolism vary significantly among individuals in many taxa, but the effects of this on fitness are generally unknown. Resting metabolic rate (RMR) in juvenile salmon and trout is positively related to dominance status and ability to obtain a feeding territory, but it is not clear how this translates into performance in natural conditions. 2.?The relationships between RMR, dominance, territoriality and growth rates of yearling Atlantic salmon Salmo salar were examined in relation to predictability in food supply and habitat complexity, using replicate sections of a large-scale controlled semi-natural stream. 3.?Estimated RMR was a strong predictor of dominance, and under conditions of a predictable food supply in a structurally simple habitat, high estimated RMR fish obtained the best feeding territories and grew faster. 4.?When the spatial distribution of food was made less predictable, dominant (high estimated RMR) fish were still able to occupy the most profitable feeding locations by periodically moving location to track the changes in food availability, but RMR was no longer a predictor of growth rate. Moreover, when a less predictable food supply was combined with a visually more complex (and realistic) habitat, fish were unable to track changes in food availability, grew more slowly and exhibited greater site fidelity, and there were no relationships between estimated RMR and quality of occupied territory or growth rate. 5.?The relative benefit of RMR is thus context dependent, depending on both habitat complexity and the predictability of the food supply. Higher habitat complexity and lower food predictability decrease the performance advantages associated with a high RMR.     

Relative production of Atlantic salmon from fluvial and lacustrine habitats estimated from analyses of scale characteristics

Empirical and back-calculated growth of Atlantic salmon parr were compared between fish reared in fluvial and lacustrine habitats of Conne River, Newfoundland. Length at age was significantly higher for lacustrine parr. Various classification and maximum likelihood estimators indicated that 75% or more of the fish used lakes for rearing. Lacustrine use is another aspect of the inherent variability and plasticity of Atlantic salmon life-history traits. As most Newfoundland river systems include lakes, estimates of regional spawning targets and potential smolt production will need to take lake habitat into account.     

Effects of discharge and local density on the growth of juvenile Atlantic salmon Salmo salar

The study explored the combined effects of density, physical habitat and different discharge levels on the growth of juvenile Atlantic salmon Salmo salar in artificial streams, by manipulating flow during both summer and winter conditions. Growth was high during all four summer trials and increased linearly with discharge and mean velocity. Differences in fish densities (fish m^?3) due to differences in stream volume explained a similar proportion of the variation in mean growth among discharge treatments. Within streams, the fish aggregated in areas of larger sediment size, where shelters were probably abundant, while growth decreased with increasing densities. Fish appeared to favour the availability of shelter over maximization of growth. Mean growth was negative during all winter trials and did not vary among discharge treatments. These results suggest that increased fish densities are a major cause of reduced summer growth at low discharge, and that habitat‐mediated density differences explain the majority of the growth variation across habitat conditions both during summer and winter.     

Fish may fight rather than feed in a novel environment: metabolic rate and feeding motivation in juvenile Atlantic salmon

This study tested the hypothesis that juvenile Atlantic salmon Salmo salar with a high resting metabolic rate and probability of dominance will also have a correspondingly higher feeding motivation in a novel environment to offset their greater costs of maintenance. The opposite was found to be the case: Atlantic salmon with a high standard metabolic rate had a slightly but significantly lower feeding motivation. It is hypothesized that Atlantic salmon with higher maintenance costs opt to be more aggressive at the expense of the costly activities associated with feeding, since elevated aggression in a new habitat is a more successful strategy for acquiring a feeding territory (and hence sustaining food intake in the long-term) than a high feeding motivation alone.