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 .     

Morphological and physiological changes of female tropical eels, Anguilla celebesensis and Anguilla marmorata, in relation to downstream migration

Acoustic telemetry and microchemical analysis of otolith strontium-calcium ratios were used to evaluate how exotic brown trout Salmo trutta have responded to Japanese riverine environments of south-western Hokkaido by observing their migratory patterns. The existence of anadromous S. trutta was also verified. Most S. trutta caught in rivers for otolith analysis were freshwater residents (95·6%), whereas those caught in the sea were mainly smolts (91·3%), which had just migrated from rivers during spring. Anadromous S. trutta (n = 6) were captured in rivers and in the sea, confirming the existence of mature pre- and post-spawning fish. According to telemetry results, both mature and immature S. trutta used the river in winter, and their estimated sea-run timings showed individual differences. Through the combination of these two methods, migratory patterns on various spatio-temporal scales were observed. This first documentation of the presence of both male and female anadromous S. trutta in the same region within Japan indicated the risk of further colonization of exotic S. trutta via oceanic migration.     

Archival tags reveal that Arctic charr Salvelinus alpinus and brown trout Salmo trutta can use estuarine and marine waters during winter

By tagging north-Norwegian anadromous riverine Arctic charr Salvelinus alpinus and brown trout Salmo trutta with simple archival tags that measured ambient water temperature and relating the recordings to the temperature in the surrounding environments, it was demonstrated that 91% of the S. trutta and 80% of the S. alpinus utilized the estuarine and marine environment during the winter. There was large individual variation in migratory behaviour. Salvelinus alpinus on average entered the estuary on 12 January and the marine environment on 26 February, had continuous marine migrations lasting up to 55 days, and spent on average 40 days in the estuary and 25 days in the sea during the winter. The corresponding numbers for S. trutta were 15 December and 4 January for first entry in the estuary and sea, maximum 39 days in the marine environment and average number of days in the estuary and sea were 34 and 50. Most individuals of both species frequently shifted between the three habitats. These findings thus contradict previous studies conducted on lake-dwelling populations, which suggested that northern populations of both species solely overwinter in fresh water. The use of inexpensive temperature recording archival tags gave detailed continuous information on seasonal migrations between habitats with different thermal characteristics.     

Identification of juvenile natural hybrids between Atlantic salmon (Salmo solar L.) and trout(Salmo trutta L.)

The identification of two wild salmonid smolts, suspected to be hybrids between Atlantic salmon ( Salmo salar ) and trout ( S trutta ) on morphological grounds, was confirmed by electrophoretic separation of serum proteins. A third, trout-like, hybrid was identified in a sample of parr. Hybrids showed presence of both parent types of esterase, prealbumin and transferrin.     

Migrations of sea trout, Salmo trutta L., from the Vardnes river in northern Norway

During the periods 1956–1963 and 1967–1970 traps were operated to catch upstream- and downstream-migrating sea trout, Salmo trutta L. A total of 15 788 sea trout were tagged, using Carlin tags. The number of recaptures made in the traps was 4481, of which 1796 were recaptured more than once.
The distribution of the 2122 recaptures in the sea provides a picture of the sea-migration pattern. Of the sea recaptures, 52.8% were reported as within a distance of 3 km from the river mouth, compared to 0.7% more than 80 km away. All the different size-groups of sea trout were represented among both the long-distance and the short-distance migrants. The results of this study of sea trout migrations are discussed in relation to the published results for Atlantic salmon, Salmo salar L., and sea charr, Salvelinus alpinus (L.), from the same river.
The four highest values recorded for mean distance of daily travel away from the river were 20, 8, 8 and 6km day⁻¹ by smolts and 6, 6, 5 and 5km day⁻¹ by larger-sized sea trout.
Recaptures of tagged sea trout in rivers other than the Vardnes totalled 506, of which 306 had been tagged as smolts. The calculated minimum percentage of stray is 15.5%. The proportion of sea trout from the Vardnes river that actually spawn in other rivers is not known. No significant difference in length distribution was found between the sea trout caught in the Vardnes river and those caught in other rivers. An hypothesis concerning the selective advantages of straying by anadromous salmonids living in small rivers is discussed.     

Effect of water temperature and density of juvenile salmonids on growth of young-of-the-year Atlantic salmon Salmo salar

A von Bertalanffy growth model for young-of the-year Atlantic salmon Salmo salar in a small French coastal stream was fitted using water temperatures and densities of juvenile salmonids (S. salar and brown trout Salmo trutta) as covariates influencing daily growth rate. The Bayesian framework was used as a template to integrate prior information from external data sets. The relative influence of the covariates on parr growth was quantified and results showed that growth of S. salar juveniles depended on both water temperatures and densities, but that most of the spatiotemporal variability of growth resulted from local spatiotemporal variations of 0+ age salmonid (S. salar and S. trutta) densities. Further analysis revealed that the fluctuations in young-of-the-year salmonid densities are likely to dominate the effects of potential future warming of water temperature due to climate change. It is concluded that factors that could affect salmonid densities might well have a greater effect on S. salar population dynamics than factors influencing water temperatures.     

Water chemistry and its effects on the physiology and survival of Atlantic salmon Salmo salar smolts

The physiological effects of episodic pH fluctuations on Atlantic salmon Salmo salar smolts in eastern Maine, U.S.A., were investigated. During this study, S. salar smolts were exposed to ambient stream-water chemistry conditions at nine sites in four catchments for 3 and 6 day intervals during the spring S. salar smolt migration period. Plasma chloride, plasma glucose, gill aluminium and gill Na(+)- and K(+)-ATPase levels in S. salar smolts were assessed in relation to ambient stream-water chemistry during this migration period. Changes in both plasma chloride and plasma glucose levels of S. salar smolts were strongly correlated with stream pH, and S. salar smolt mortality occurred in one study site with ambient stream pH between 5·6 and 5·8 during the study period. The findings from this study suggest that physiological effects on S. salar smolts are strongly correlated with stream pH and that in rivers and streams with low dissolved organic carbon (DOC) concentrations the threshold for physiological effects and mortality probably occurs at a higher pH and shorter exposure period than in rivers with higher DOC. Additionally, whenever an acidification event in which pH drops below 5·9 coincides with S. salar smolt migration in eastern Maine rivers, there is potential for a significant reduction in plasma ions of S. salar smolts.     

Genetic variation for tolerance to acidic water in salmonids

Water pH is an important factor affecting the general water quality as well as quality traits in fishes, and the magnitude of the effect varies among species. The massive and negative effect of acidification of rivers and lakes became evident during the 1960s and 1970s and caused the depletion of fish stocks in several countries in the northern hemisphere. Significant variation in tolerance to acidic water has been documented among salmonid species, and large genetic variation has been identified among strains of brown trout Salmo trutta, brook trout Salvelinus fontinalis and Atlantic salmon Salmo salar. For S. trutta, S. fontinalis and S. salar, there is considerable additive genetic variation in tolerance to acidic water, with heritabilities (h(2)) ranging from 0·09 to 0·27 for dead eyed-eggs (the period most sensitive to low pH). The main reasons for depletion of freshwater fish stocks are discussed.     

Pre-winter lipid stores in brown trout Salmo trutta along altitudinal and latitudinal gradients

Pre-winter lipid stores of brown trout Salmo trutta L. parr were compared along altitudinal (0-920 m a.s.l.) and latitudinal (58-71° N) gradients. There were increases in lipid content (size adjusted to common lipid-free dry mass of 2·0 g, corresponding to fresh mass of 10 g) with both increasing altitude and latitude. Mean size-adjusted lipid content for S. trutta in high altitude rivers was 60% higher than at low altitude (0·29 and 0·18 g, respectively). Mean size-adjusted lipid content for S. trutta in northern rivers was 30% higher compared to that in southern rivers (0·30 and 0·23 g, respectively). There was a marked between-river variation in mean lipid storage, probably reflecting different strategies or opportunities for the pre-winter acquisition of lipid both locally within rivers and between different populations. This study shows that temperature or winter length, not latitudinal covariates such as annual light regime, governs lipid storage patterns in juvenile salmonids.     

Wild adult hybrids of Salmo salar L. and Salmo trutta L.

Ovarian development was impaired in three adult Salmo salar L. × S. trutta L. hybrids identified among adult salmonids in Scottish fisheries. Species-specific variation at enzyme loci indicated that the fish were F1 hybrids and mitochondrial DNA analysis showed them to be the progeny of S. salar females.     

The effect of temperature on sperm motility and enzymatic activity in brown trout Salmo trutta, burbot Lota lota and grayling Thymallus thymallus

The effect of temperature on sperm motility was investigated in brown trout Salmo trutta, burbot Lota lota and grayling Thymallus thymallus using water and sperm motility prolonging saline solution (SMPS) for motility activation. The effect of temperature (4-20° C) on spermatozoal enzymes for energy supply [malate dehydrogenase (MDH), pyruvate kinase (PK), adenylate kinase (AK)], flagellar movement [Mg(2+) adenosine triposphatase (ATPase)] and oxidative defence [peroxidase (POX)] were measured in S. trutta and L. lota. Temperatures yielding the highest initial sperm motility rates and swimming velocities were 4-6° C for S. trutta [investigated range (IR) = 4-12° C] and L. lota (IR = 2-8° C) and 8-16° C (IR = 4-16° C) for T. thymallus. Motility variables were re-measured after 30 s in S. trutta, after 45 s in T. thymallus and after 60 s in L. lota in water and after 2 min in all investigated species in SMPS. Motility variables were increased by low temperatures and the results differed between water and SMPS. In S. trutta and L. lota, the temperature resulting in highest activities of MDH, PK, AK and ATPase was 4° C. POX had a very narrow temperature optimum at 20° C in both species. This may indicate that the temperature optimum of enzymes of energy supply and flagellar movement are closely related to motility. The present data show that the variables are affected by temperatures in an ecologically relevant range. Too low, as well as too high temperatures affected sperm motility, and the winter spawners (S. trutta and L. lota) have a narrower temperature optimum than the spring spawner T. thymallus.     

Wild arctic char Salvelinus alpinus and trout Salmo trutta: hosts and reservoir of the salmonid pathogen Spironucleus salmonicida (Diplomonadida; Hexamitidae)

Spironucleus salmonicida is a diplomonad flagellate known to cause systemic infections in farmed salmonids. In northern Norway, outbreaks of spironucleosis in farmed Atlantic salmon Salmo salar have been a recurring problem. Common to all these outbreaks was the origin of smolts: all came from the same farm. In the present study, wild Arctic char Salvelinus alpinus and brown trout Salmo trutta were sampled from the lakes used as a water source for the smolt supplier. In addition, smolt and three-spined sticklebacks Gasterosteus aculeatus were sampled from the smolt farm. Bile and intestinal contents from the sampled fish were examined by light microscopy and PCR. Spironucleus salmonicida was identified in both wild Arctic char and brown trout from the lakes used as water sources by the smolt farm, suggesting that the farmed fish were exposed to this pathogen before transfer to the sea. Spironucleus barkhanus and Spironucleus salmonis were also identified in the sampled fish. The present study also demonstrated that infections with multiple Spironucleus species are present in wild salmonids. No indications of disease related to diplomonad infections were observed in the wild fish, suggesting that wild salmonids are reservoir hosts of Spironucleus salmonicida.     

An experimental study of the multiple effects of brown trout Salmo trutta on the bioenergetics of two Arctic charr Salvelinus alpinus morphs

This study investigated the importance of competition with brown trout Salmo trutta as a driver of the morphological and behavioural divergence of two morphs of Arctic charr Salvelinus alpinus. The morphs originated from two lakes differing in absence or presence of the competitor. The bioenergetics and behaviour of S. alpinus were quantified in replicate experimental enclosures (mean volume: 150 m(3) ) stocked with 15 S. alpinus of one morph or the other and in the absence or presence of nine S. trutta. The presence of S. trutta decreased growth rate, affected food consumption and increased activity costs in S. alpinus, but provided little support for the hypothesis that competition with S. trutta is a major driver of the divergence of the two S. alpinus morphs. Both morphs responded similarly in terms of mean growth and consumption rates per enclosure, but the association between individual morphology and growth rate reversed between allopatric and sympatric enclosures. While the activity patterns of the two morphs were unaffected by the presence of S. trutta, their swimming speed and activity rate differed. Since the profound differences in the structure of the physical habitat of the source lakes provided a more likely explanation for the difference observed among these two morphs than interspecific competition, it is hypothesized that physical habitat may sometimes be a significant driving force of the phenotypic divergence.     

The functional relationship between peak spring floods and survival and growth of juvenile Atlantic Salmon (Salmo salar) and Brown Trout (Salmo trutta)

1. The effects of high spring floods on survival and growth of juvenile Atlantic Salmon, Salmo salar, and Brown Trout, Salmo trutta , are explored, using data from a long-term study in the River Saltdalselv, northern Norway. The flow regime in this river is typical for northern rivers.
2. There was considerable variation in year class strength of both species.
3. Mortality of Atlantic Salmon increased significantly in years with high discharge during the alevin stage as well as the first week after emergence. High discharge during the egg stage and more than 1 week after emergence seemed to be of minor importance. Water temperature at emergence was rather high (average 10·5 °C) and did not significantly affect year class strength.
4. Brown Trout emerged earlier than Atlantic Salmon at an average water temperature of 8·2 °C. Highest mortality was observed in years with low water temperatures at emergence as well as high discharge during the alevin stage.
5. For 1-year-old fish or older, the size of the spring peak flood did not influence mortality significantly.
6. Growth of Atlantic Salmon parr was diminished in years with a high peak spring flood. A similar effect on Brown Trout was not detected.     

Predation on Atlantic salmon and sea trout during their first days as postsmolts

Radio-tagged smolts of Atlantic salmon Salmo salar and sea trout Salmo trutta were predated heavily by sea birds after crossing the saline limit in the estuary of the River Skjern, Denmark. Most predation took place within the first 9 h after estuarine entry. The field data do not contradict the hypothesis of maladaptive anti-predatory behaviour.     

Pelagic marine feeding of Arctic charr and sea trout

In a fjord in northern Norway, both immature and maturing individuals (170–500 mm fork length) of anadromous Arctic charr Salvelinus alpinus and anadromous brown trout (sea trout) Salmo trutta were captured up to 5000 m offshore. Both species had a high feeding intensity and an almost exclusive piscivorous diet, with herring Clupea harengus as the dominant prey, an apparently opportunistic response to a high density of small‐sized herring in the pelagic zone.     

Experimental and natural host specificity of Loma salmonae (Microsporidia)

The microsporidian Loma salmonae (Putz, Hoffman & Dunbar, 1965) Morrison & Sprague, 1981 has caused significant gill disease in Pacific salmon Oncorhynchus spp. Host specificity of the parasite was examined experimentally by per os challenge of selected salmonids and non-salmonids with infective chinook salmon O. tshawytscha gill material. Pink Oncorhynchus gorbuscha and chum salmon O. keta, brown Salmo trutta and brook trout Salvelinus fontinalis, and chinook salmon (controls) were positive, whereas Atlantic salmon Salmo salar and Arctic char Salvelinus alpinus were negative. In addition, no non-salmonids were susceptible to experimental exposure. Wild Pacific salmon species in British Columbia, Canada, were examined for L. salmonae during their freshwater life history stages (smolts, prespawning, spawning). All stages were infected, although infections in smolts were only detectable using a L. salmonae-specific PCR test. Many previous Loma spp. described from Oncorhychus spp. are likely L. salmonae based on host, parasite morphology, and site of infection.     

Thermal growth performance of juvenile brown trout Salmo trutta: no support for thermal adaptation hypotheses

Using thermal growth data from eight populations of anadromous and lake-feeding brown trout Salmo trutta , hypotheses of adaptation to local optima and countergradient variation in growth were tested. The adaptation to local optima hypothesis suggests that natural selection can shift optimal performance temperatures to match the prevailing temperature in a new or changed thermal niche. In contradiction, the countergradient variation hypothesis suggests that populations from hostile environments perform better than conspecifics from benign environments at all temperatures. In this study, growth capacity varied between populations but there was no significant correlation between any of the estimated thermal performance parameters ( e.g. lower and upper thermal growth limits, optimal temperature for growth and maximum growth capacity) and natural climatic conditions among populations. Hence, S. trutta growth response to temperature lends no support for either of the two suggested thermal adaptation hypotheses. Instead, growth capacity among populations tended to correlate positively with female size at maturity.     

Marine feeding of anadromous Salmo trutta during winter

Juvenile and adult anadromous trout Salmo trutta utilize the sea for feeding during the winter in the Skagerrak. This finding conflicts with the traditional view that anadromous trout overwinter in fresh water. Adults, just prior to spawning, were captured at sea in October to December, and spent fish were caught at sea from October to April, showing that the fish may leave the stream and move to sea just after spawning and spend the winter there. During mid‐winter (January to February), the feeding probability (chance of finding a fish with food in its stomach) increased markedly with increasing body length, with no similar effect during early and late winter (October to December and March to April). Among individuals with food in their stomach (72·5%), there was no evidence for variation in feeding intensity [stomach fullness = (mass of stomach content)(fish body mass)⁻¹] among early, mid‐, and late winter.     

Host specificity and geographical distribution of Eubothrium in European salmonid fish

The host specificity and distribution of Eubothrium crassum (Bloch, 1779) and Eubothrium salvelini (Schrank, 1790), morphologically fairly similar pseudophyllidean tapeworms parasitizing salmonid fish, were critically assessed on the basis of morphological and genetic evaluation of extensive material collected from different definitive hosts and geographical regions in Europe. Eubothrium crassum occurs in fish of the genera Salmo, i.e. salmon (S. salar - both freshwater and marine), sea trout (S. trutta trutta), brown trout (S. trutta fario), and lake trout (S. trutta lacustris), and also in Danubian salmon (Hucho hucho) and vendace (Coregonus albula). Eubothrium salvelini parasitizes Arctic char (Salvelinus alpinus) and brook trout (Salvelinus fontinalis) in Europe, and also whitefish (Coregonus wartmanni). Rainbow trout (Oncorhynchus mykiss), which is not a native European fish species, was found to be a suitable definitive host for both Eubothrium species, which may occur simultaneously in the same fish. Previous records of E. crassum in Arctic char and brook trout, and those of E. salvelini in fish of the genus Salmo were most probably misidentifications. Most studies of Eubothrium have involved salmonids from the northern part of Europe, with few records from southern and south-eastern Europe. This study also confirmed the reliability of the morphology of the apical disc for the discrimination of E. crassum and E. salvelini.