Migratory timing, marine survival and growth of anadromous brown trout Salmo trutta in the River Imsa, Norway

The aim of the paper was to study sea migration, growth and survival of brown trout Salmo trutta of the River Imsa, 1976–2005. The migratory S. trutta were individually tagged and fish leaving or entering the river were monitored daily in traps located near the river mouth. The mean annual duration of the sea sojourn was 6–9 months for first-time migrants moving to sea between January and June. It was 8–18 months for those migrating to sea between July and December. Veteran migrants stayed 12 months or less at sea and most returned to the river in August. Early ascending fish stayed the longest in fresh water because most returned to sea in April to May. The day number of 50% cumulative smolt descent correlated negatively with mean water temperature in February to March and the February North Atlantic Oscillation index (NAOI). Mean annual sea growth during the first 2 years after smolting was higher for S. trutta spending the winter at sea than those wintering in the River Imsa. First year's sea growth was lower for S. trutta descending in spring than autumn. For first-time migrants, it correlated negatively with the February NAOI of the smolt year. Sea survival was higher for spring than autumn descending first-time migratory S. trutta with a maximum in May (14·9%). Number of anadromous S. trutta returning to the river increased linearly with the size of the cohort moving to sea, with no evidence of density-dependent sea mortality. Sea survival of S. trutta smolts moving to sea between January and June correlated positively both with the annual number of Atlantic Salmo salar smolts, the specific growth rate at sea, and time of seaward migration in spring. This is the first study indicating how environmental factors at the time of seaward migration influence the sea survival of S. trutta .     

Differential response to water current in offspring of inlet-and outlet-spawning brown trout Salmo trutta

Two-year-old hatchery-reared progeny of inlet- and outlet spawning brown trout from Lake Tytifjorden were released at the mouth of the R. Imsa, south-western Norway. There were significant differences in migratory direction of juveniles between the two populations. After release, juvenile fish from the outlet river population moved against the current and ascended the R Imsa, while the inlet rivet fish tended to migrate with the water current to the sea. This differential response to water current in juveniles appears to be due to genetic differences between the populations, and parallels that found in their ancestors native environments.     

Effects of seawater-acclimatization and release sites on survival of hatchery–reared brown trout Salmo trutta

To test whether seawater–acclimatization of hatchery–reared anadromous and freshwater resident brown trout before release increased the survival of adults, smolts were retained 0, 2, 4 and 8 weeks in sea water before release. Total recapture rate increased for smolts retained 4 and 8 weeks in sea water before release relative to the controls. This trend was more pronounced for Freshwater resident than for anadromous stocks, Offspring of anadromous fish stayed longer at sea than offspring of freshwater resident fish. Recapture rates in fresh water were higher for brown trout released in the river than in the fjord in the R, Drammen area, but not in the R. Imsa. In both cases, most fish were recaptured in the sea. Moving into the R. Imsa (relative to other rivers) appeared higher for fish released at the mouth of the river (93%) than in the fjord (47%). Judged from the recapture rates, sea survival appeared to be the same whether released in the fjord or at the mouth of the river.     

Factors affecting river entry of adult Atlantic salmon in a small river

In this study, effects of stock origin, fish size, water flow and temperature on time of river ascent of adult Atlantic salmon Salmo salar were tested. Brood stocks were collected in eight Norwegian rivers situated between 59 and 69° N. The fish were reared to smolts, individually tagged and released in the River Imsa, south-west Norway (59° N). Adults from all stocks approached the Norwegian coast concurrently, but Atlantic salmon ≥70 cm in natural tip length entered coastal water slightly earlier during summer than smaller fish. Atlantic salmon <70 cm, however, ascended the river significantly earlier and at lower water flow and higher water temperature than larger fish. Although largest in size, the fish from the northern populations (62–69° N) ascended the River Imsa almost 1 month earlier than those from the south (59–60° N). They seemed less restricted by the environmental factors than the fish originating from the more southern rivers. There was no apparent trend among years in time of river ascent. Maximum ascent per day occurred at water discharges between 12·5 and 15 m³ s⁻¹ and at water temperatures between 10 and 12·5° C. There was a significant positive correlation between water flow and river ascent during the first part of the upstream run from July to September with best correlation for September, when multiple regression analysis indicated that water temperature had an additional positive effect. Stock origin, fish size and water discharge were important variables influencing the upstream migration of Atlantic salmon in small rivers.     

Migratory behaviour and growth of hatchery-reared post-smolt Atlantic salmon Salmo salar

Individually lagged, 1+ and 2+ hatchery-reared smolts of Atlantic salmon were released in spring and early summer at the mouth of the R. Imsa, south-western Norway. The post-smolts moved mainly northwards in the sea with the coastal current. The estimated mean migratory speed (± s.d. ) of those captured in the sea along the Norwegian coast was 7.45 (± 6.26) km day ⁻¹; in the fjords it was 1.63 (± 2.33) km day⁻². Many of the post-smohs ascended rivers the same year as released; 37.3% of the total number recaptured were caught in R. Imsa, upstream from the site of release, and 5.8% were caught in other rivers throughout middle and southern parts of Norway. The fish recaptured in rivers was probably sexually mature and entered rivers to spawn. Mean specific growth rate for post-smolts caught in the sea was higher than for those caught in R. Imsa (P <0.001) but not for those caught in other rivers (P> 0.05). Post-smolts ascending R. Imsa were smaller at release than those ascending other rivers. However, there was no size difference at release between post-smolts captured in the sea and those recaptured in rivers other than the R. Imsa.     

Ecological Forms of Black Sea Brown Trout (<Emphasis Type="Italic">Salmo trutta labrax</Emphasis>) in the Mzymta River as Manifestation of Ontogenetic Plasticity

Populations of brown trout in the Mzymta River and its tributaries include anadromous (mainly female) and resident (mainly males) fish. Some resident males in the basin of the Mzymty River attain sexual maturity at the age 1+, and resident females mature at the age 2+ or 3+. The maximum age of resident fish is 4+ in the samples studied. Migrations of anadromous brown trout to the sea occur at the ages 1+, 2+, or 3+. Future spawners spend from 1 to 4 years at feeding grounds in the sea. Smolts of the population are characterized by performing not only spring but also autumn migrations to the sea. One smolt specimen has been detected upstream from the dam in the river where spawners of anadromous brown trout do not migrate; this means that the capability for sea migrations persists long in the population represented only by resident specimens of brown trout. The diversity of life cycles and ecological forms in populations of brown trout is not lower than in populations of brown trout in Northern and Western Europe. The comparison of the data obtained with published data makes it possible to come to the conclusion about the high plasticity of ontogenesis of Black Sea brown trout.     

Reproductive migration of brown trout in a small Norwegian river studied by telemetry

The movement of 34 large (39–73 cm standard length) brown trout Salmo trutta was monitored using radio telemetry for up to 74 days in Brumunda, a small Norwegian river (mean annual discharge 3·3 m³ s⁻¹) flowing into the large Lake Mjøsa. The maximum range of movement in the river was 20 km. No clear relationships existed between individual movement and water discharge, temperature and barometric pressure. Brown trout migrated at all levels of water discharge. At low discharge (<2 m³ s⁻¹) movements were nocturnal. A weir 5·3 km from the outlet restricted ascending brown trout at low ( c . 6° C), but not at high ( c . 8° C) water temperatures. Spawning occurred in September to October and tagged individuals spent 2–51 days at the spawning sites. Mean migration speed from tagging to when the fish reached the spawning area, and from when they left the spawning areas and reached the lake was 1·0 and 2·3 km day⁻¹, respectively. All tagged brown trout that survived spawning returned to the lake after spawning.     

Growth hormone-induced effects on mortality, energy status and growth: a field study on Brown Trout (Salmo trutta)

1. Growth hormone (GH) treatment increases the growth rate and competitive ability of salmonids under laboratory conditions. Since fast growth should increase fitness, why is endogenous secretion of GH not higher in wild fish? To address this question, three hypotheses were suggested. H₁: high GH levels reduce antipredator responses and may therefore increase mortality from predation. H₂: high GH levels reduce long-term (e.g. over winter) survival by reducing allocation to critical energy reserves. H₃: GH is not beneficial for growth under natural conditions.
2. To test these hypotheses, the performance of GH-treated juvenile Brown Trout ( Salmo trutta ) and control (placebo) trout was compared in an enclosed stream section subjected to natural predation. Four experiments were conducted during winter, spring, summer and autumn, respectively.
3. Mortality rates were not significantly different between GH-treated and control trout in any of the four experiments so H₁ was not supported. Energy reserves were generally lower in GH-treated fish, which is consistent with H₂, whereas growth rates in mass were higher in GH-treated fish than in controls so H₃ was not supported. This suggests that GH promotes growth at the expense of investment in maintenance.
4. Judging from growth and mortality rates, the fitness of GH-treated and control trout appeared similar. Thus, escaped GH-manipulated fish may compete successfully with wild fish.
5. Hatchery-raised trout with higher initial condition index suffered higher mortality rates than more slender fish. This novel finding may be explained by reduced escape ability related to body morphology, reduced behavioural responses towards predators by high-condition trout, or predator preferences for high-condition fish.     

Experimental sea ranching of brook trout, Salvelinus fontinalis Mitchill

An experiment to induce anadromy in a population of wild brook trout, Salvelinus fontinalis , was conducted near Sept-Iles, Quebec, in 1978–1979. Brook trout were captured from the Matamek River, tagged and transported to the Matamek River estuary during late spring and early summer, and allowed free movement between an impassable waterfall 0.7 km upstream and the sea. Fish were recaptured in autumn as they returned to fresh water. Over two years, 34.0% of the released fish were recaptured. Best returns were in the 2+ and 3+ age classes with 38.0 and 62.1% recaptured, respectively. Straying of transplanted fish appeared to be <1%. All age classes included sea run brook trout (sea trout) but the largest percentages of sea trout occurred in older fish. Growth was better in sea trout than in fish which did not develop anadromy, presumably a function of an increased food supply at sea. Severe tagging effects stunted growth and probably suppressed anadromy, especially among younger fish. Sexual characteristics of recaptured fish indicated suppressed maturation of gonads in sea trout compared to fish remaining in fresh water and there was a shift to a larger percentage of females in the sea trout. Comparisons between our results and data on other anadromous Salvelinus species underscore the potential for sea-ranching of trout and char as a moderate effort, high yield aquaculture technique.     

Time and size at seaward migration influence the sea survival of Salmo salar

Whether time of seaward migration of young Atlantic salmon Salmo salar influences their subsequent survival and growth was investigated in the River Imsa, south‐western Norway. Salmo salar were tagged when moving downstream through a trap near the outlet between 1976 and 2010 and recaptured on their adult return. Most descended as smolts in April and May, but some descended during the other months of the year. Annual variation in timing of the smolt migration was significantly correlated with variation in water temperature during spring. Mean total body length of the descending S. salar varied with month of seaward migration. The sea survival of S. salar emigrating from the River Imsa between January and May was 2·8 times higher than for those descending between June and December. The sea survival of the various cohorts decreased with increasing river temperature in April to May, prior to the smolt migration, and decreasing day number when the smolts moved to sea. The size of smolts descending the river between April and May did not affect the survival at sea as much as it affected the survival of migrants descending in any other month of the year. The majority of the downstream migrating S. salar were 2 years old, but proportionally, more 1 year olds moved downstream in the autumn than in the rest of the year. Mean duration between downstream migration of the young and the return migration of the grilse was shortest (12·7 months) for those descending in July and August and longest for those descending in October (21 months). Mean monthly specific growth rate was highest for those migrating downstream between May and July and lowest for those emigrating in September. Based on the present results, it was hypothesized that S. salar emigrating between April and August migrated directly out into the ocean, while those that emigrated between October and March stayed in the estuary until the subsequent spring.     

Growth, production and bioenergetics of brown trout in upland streams with contrasting riparian vegetation

1. A series of laboratory-based equations on trout growth and bioenergetics developed by J.M. Elliott were applied to data collected for brown trout ( Salmo trutta L.) under field conditions in Co. Mayo, Western Ireland. Fish were collected by electrofishing eight upland streams with contrasting riparian vegetation; grassland, open canopy and closed canopy deciduous.
2. Stream temperatures, one of the main influencing factors on fish growth and energetics, did not differ significantly between riparian types.
3. Observed growth rates were lower than the predicted maximum growth rates and were not influenced by riparian vegetation type. Growth ranged between 0.66% day⁻¹ for 0 + trout to 0.08% day⁻¹ for 2 + trout.
4. Production estimates showed no clear difference between riparian vegetation types over the growing season.
5. Fish densities and biomass tended to be greater in closed canopy streams particularly in summer.
6. Actual ration sizes calculated for trout were similar to the ration required for maintenance metabolism and were only 45–63% of the maximum potential rations. Although there was an ontogenetic increase in ration size with increasing fish age, the proportion of ration available for growth (i.e. the difference between actual and maintenance rations) did not differ between age classes but was greatest in summer. 1+ and 2+ trout show greatest ration available for growth in grassland streams.
7. Trout growth did not differ between riparian vegetation types but did vary seasonally with greatest attainment in summer. Growth was limited in the present study possibly due to combined effects of reduced prey available to fish and low stream temperatures reducing metabolic requirements. In such food limited systems, terrestrial invertebrate energy subsidies could have significant benefits to brown trout growth, production and bioenergetics.     

Evidence for an autumn downstream migration of Atlantic salmon Salmo salar (Linnaeus) and brown trout Salmo trutta (Linnaeus) parr to the Baltic Sea

In the eastern Baltic rivers, anadromous salmonid parr are known to smoltify and migrate to the sea from March until June, depending on latitude, climate and hydrological conditions. In this study, we present the first records of autumn descent of brown trout Salmo trutta and Atlantic salmon Salmo salar from the Baltic Sea Basin. Otolith microchemistry analyses revealed that these individuals hatched in freshwater and had migrated to the brackish water shortly prior to capture. The fish were collected in 2006, 2008, 2009 and 2013 from Eru Bay (surface salinity 4.5–6.5 ‰), Gulf of Finland. This relatively wide temporal range of observations indicates that the autumn descent of anadromous salmonids is not a random event. These results imply that autumn descent needs more consideration in the context of the effective stock management, assessment and restoration of Baltic salmonid populations and their habitats.     

Competitive habitat displacement of brown trout by Siberian sculpin: the role of size and density

Habitat competition in brown trout Salmo trutta and Siberian sculpin Cottus poecilopus was investigated by varying density, fish size, and species composition in stream channels providing areas of different substratum particle sizes. In allopatry, both small (52 ± 4 mm L _T) and large (86 ± 6 mm L _T) brown trout exhibited strong preference for the intermediate (8–11 cm diameter) and large (17–21 cm) gravel substrata. There was a tendency for more brown trout to occupy finer (2–4 cm) substrata with increasing density, in particular for large brown trout. Also, more small brown trout were observed on finer substrata when tested with large brown trout, suggesting interspecific competition for restricted space. Both small (56 ± 6 mm L _T) and large (88 ± 10 mm L _T) Siberian sculpin preferred the large gravel in all tests, and did not change their substratum preferences much with increasing densities, suggesting higher tolerance for 'crowding'. The large Siberian sculpin preferred the coarser substratum, and the largest individuals were consistently found on it. In sympatry with large Siberian sculpin, habitat displacement of brown trout occurred, indicative of interspecific competition. A higher proportion of small and large brown trout occupied the finer substrata than in allopatry. Habitat selection by large Siberian sculpin appeared to be unaffected by species composition and density. Small Siberian sculpin were displaced to finer substrata when tested with large Siberian sculpin, suggesting intraspecific competition. The results indicate that Siberian sculpin are potential habitat competitors for young brown trout.     

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

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

Mismatch between fishway operation and timing of fish movements: a risk for cascading effects in partial migration systems

Habitat fragmentation is a growing problem worldwide. Particularly in river systems, numerous dams and weirs hamper the movement of a wide variety of species. With the aim to preserve connectivity for fish, many barriers in river systems are equipped with fishways (also called fish passages or fish ladders). However, few fishways provide full connectivity. Here we hypothesized that restricted seasonal opening times of fishways can importantly reduce their effectiveness by interfering with the timing of fish migration, for both spring‐ and autumn‐spawning species. We empirically tested our hypothesis, and discuss the possible eco‐evolutionary consequences of affected migration timing. We analyzed movements of two salmonid fishes, spring‐spawning European grayling (Thymallus thymallus) and autumn‐spawning brown trout (Salmo trutta), in Norway's two largest river systems. We compared their timing of upstream passage through four fishways collected over 28 years with the timing of fish movements in unfragmented river sections as monitored by radiotelemetry. Confirming our hypothesis, late opening of fishways delayed the migration of European grayling in spring, and early closure of fishways blocked migration for brown trout on their way to spawning locations during late autumn. We show in a theoretical framework how restricted opening times of fishways can induce shifts from migratory to resident behavior in potamodromous partial migration systems, and propose that this can induce density‐dependent effects among fish accumulating in lower regions of rivers. Hence, fragmentation may not only directly affect the migratory individuals in the population, but may also have effects that cascade downstream and alter circumstances for resident fish. Fishway functionality is inadequate if there is a mismatch between natural fish movements and fishway opening times in the same river system, with ecological and possibly evolutionary consequences for fish populations.     

Population dynamics and production of Jesogammarus annandalei, an endemic amphipod, in Lake Biwa, Japan

SUMMARY 1. Population dynamics and production of Jesogammarus annandalei , an endemic amphipod in Lake Biwa, were examined from April 1997 to June 1998. The life cycle of this species was 1 year with the new generation beginning in early autumn. They preferred low temperature (<12 °C) and their spatial distribution varied seasonally and accordingly.
2. In deep water, the abundance of J. annandalei ranged from 200 to 63 000 m⁻² and decreased towards summer and the biomass (0.01∼3.6 g C m⁻²) was on average comparable that of zooplankton. The density was much higher than that recorded by a study conducted 35 years ago.
3. Individual growth rate of this amphipod was high in winter and spring but decreased in summer. Annual production of J. annandalei (6.2 g C m⁻² year⁻¹) was only 2% of primary production but was at the higher end of the range reported for amphipods in oligo- and mesotrophic lakes.
4. These results are consistent with the view that Lake Biwa is becoming more eutrophic, with a consequent decrease in the abundance of predatory fish in the profundal zone.     

The duration of sea and freshwater residence of the sea trout,Salmo trutta,from the Vardnes River in northern Norway

Synopsis The sea trout,Salmo trutta, from the Vardnes River generally stays in freshwater during winter. The results from 11 years of tagging experiments showed that the mean annual duration of the feeding migration in the sea was 68 ± 21 days (yearly means ranging between 54 and 88 days). A pronounced variation was found with size, sex and time of migration of the fish and between the different years. The males stayed for a shorter time in the sea (mean 66 days) than the females (mean 69 days) and they usually descended later and ascended earlier. Those fish that descended first stayed for the longest period in the sea. The mean annual duration of stay in the sea for first and second time migrants to the sea was positively correlated with the mean sea temperature in May (r² > 0.5). A positive correlation was also found between the annual duration of stay in the sea for the sea trout above 33 cm and the mean temperature in the river in July (r² > 0. 6). Prolonged mean yearly sea sojourn was generally correlated with higher sea temperatures. Water temperature-salinity effects are discussed.     

Sea trout in North Argyll Sea lochs, population, distribution and movements

Sea trout were sampled by shore seining in the sea lochs of the west coast of Scotland between 1970 and 1974, This area is of special interest because of the indentation of the coastline and the varying degree of freshwater and marine influence in the sea lochs. The majority of fish caught were in their first year after smolt migration. These post-smolts were caught mainly in May and June, after which the number of trout present was low until fish at the 'whirling' stage appeared in the catches at the end of August, continuing through to the following spring. An additional recruitment of mainly unsilvered young trout from the rivers to the sea lochs was found in the autumn. Age and sex composition of both spring and autumn recruits, and of mature fish, were investigated and compared.
A total of 3228 sea trout were tagged, with 311 recaptures. These recaptures supported the evidence from smolt trapping and beach seine catches that, in the first post-migration year, the smolts migrate from the rivers from late March to early May, they then migrate from the sea lochs in May and June and return in late summer and autumn.     

Body composition and energy allocation in life-history stages of brown trout

Energy contents of immature parr and smolts, and mature resident and anadromous brown trout Salmo trutta sampled from a small stream in southern Norway were estimated from lipid, protein and carbohydrate concentrations. In immatures the lipid concentrations were highest in parr in the autumn. Mean lipid concentrations increased significantly with age in parr sampled in autumn (1·3% in age 0+ to 3·4% in age 3+), whereas they did not in smolts. The lipid concentrations of parr in spring were not significantly different from those of similarly aged smolts. By contrast, the relative water content (%) decreased with age in parr in the autumn and increased with age in smolts, mean values being slightly higher in smolts (78%) than in parr (77%). Protein and carbohydrate concentrations did not vary with age in the immature fish, mean protein concentrations being 18·0, 17·5 and 16·8% in parr in the autumn and spring, and in smolts, respectively. In residents, the concentrations of lipids were lower and of water higher, in age group 1 than in older fish, whereas there was no significant variation with age amongst anadromous trout. The energy concentration of 2+ smolts (349 kJ 100 g^-1) was similar to that of 0+ parr in the autumn. Mean somatic energy density in autumn was 1·1 times higher in freshwater residents than in parr at age 1+ (407 and 387 kJ 100 g^-1) and marginally different at age 2+ (462 and 426 kJ 100 g^-1, respectively). The energy contents per unit mass of residents were 1·3–1·6 times that of similar aged smolts. Mean somatic energy density of anadromous trout (504 kJ 100 g^-1) was higher than that of residents (455 kJ 100 g^-1). Somatic energy, lipid and protein concentrations were correlated highly with water contents of all life stages, age and sex groups.     

Performance of juvenile brown trout exposed to fluctuating water level and temperature

Individual daily food intake, mass‐specific growth rate and growth efficiency in groups of juvenile brown trout Salmo trutta were compared in tank experiments with three water level regimes (fluctuating, stable high and low water levels) and two temperature regimes (fluctuating between 10 and 14° C and constant 14° C) to simulate events during hydropeaking in regulated rivers. Fish exposed to high stable water level showed higher food intake and growth rate, and higher or similar growth efficiency than fish exposed to fluctuating or stable low water level. Both groups of slow‐growing and fast‐growing individuals fed less and grew slower at stable low and fluctuating water level than at stable high water level. Furthermore, growth and growth efficiency were lower in brown trout exposed to stable low water level and fluctuating temperature, particularly for groups of fish with slow growth. Temperature did not have any effect at high water level. For groups of fast‐growing fish, there was no difference in growth efficiency between treatments. It is concluded that fluctuating water level and temperature have a potentially detrimental effect on growth in juvenile brown trout and effects are more severe in slow‐ than fast‐growing fish.