The relationships between anadromy, sex ratio and parr growth of Arctic charr in a lake in North Norway

Relationships between the degree of anadromy, sex ratio and parr growth of Arctic charr Salvelinus alpinus were investigated in Lake Storvatn in Hammerfest town, by estimating the densities of resident and anadromous charr in the lake, and by comparing the smolt characteristics of fish captured in the outlet river with those of fish of the same age in the lake. About two-thirds of the charr older than 5 years were anadromous. The sex ratio of smolts was approximately 1 : 1 and the frequency of mature male parr in the lake was low. As females tended to dominate the mature portion of the anadromous population, there was probably a relatively higher mortality among sea-run male smolts. The youngest smolts (3+ and 4+) were probably recruited from fast-growing parr in the littoral zone, while older smolts (>4+) may have been recruited from all habitats in the lake.     

Relationships between Na+, K+-ATPase, sialic acid content and fluidity in rainbow trout density-separated erythrocytes

Smolt of anadromous Arctic charr Sahelinus alpinus (L.) migrating from Storvatn, northern Norway, had a similar feeding intensity as resident Arctic charr from the benthic- and pelagic zone in early summer in fresh water. While smolts fed mostly on surface insects, however, resident Arctic charr had a more broader diet consisting mostly of chironomids, Bosmina sp., benthic prey and surface insects. This indicates that Arctic charr smolts have adopted a surface oriented feeding behaviour prior to their sea migration.     

Individual growth and phase differentiation of lacustrine masu salmon, Oncorhynchus masou, under artificial rearing conditions

 In lacustrine masu salmon, Oncorhynchus masou, originated from anadromous fish and inhabiting an artificial lake (Shumarinai Lake), we examined the relationship between individual growth during the juvenile stage and phase differentiation under artificial rearing conditions. In females, the mean fork length of potential (subsequent) 1+ smolts and potential 1+ parr (1+ parr show fish that neither smolted nor matured until 1+ autumn) always differed after their first summer (0+ summer), the former being larger. In males, the juveniles that grew faster during their first spring became 0+ mature male parr. After their first summer, potential 1+ smolts were always larger in body length than potential 1+ mature parr and potential 1+ parr. These results were similar to previous reports of anadromous masu salmon, suggesting that the lacustrine masu salmon studied seems to have maintained the phase differentiation as in the original fish, likely because only 60 years have passed since the formation of the lacustrine population.     

Growth and development of a bimodal length-frequency distribution during smoking in a wild population of white-spotted charr in northern Japan

Smolt age and size, and individual growth during smoking were examined in a wild population of white-spotted charr, Salvelinus leucomaenis , in a small stream in northern Japan. Smolt size (110–210 mm F.L.) differed neither between sexes nor among age cohorts of 2–7 years. Back-calculation of riverine growth of smolts using otoliths showed a gradually increasing discrepancy in fork lengths of these age cohorts in both males and females, the faster-growing individuals smolting at an earlier age. A mark—recapture study on individually-identified fish revealed that the initial (spring-summer) length-frequency distribution of parr was unimodal, but became slightly skewed to the larger side in autumn, resulting in a bimodal distribution being evident by the following spring, with the upper modal group primarily composed of smolts. However, closer examination revealed that both male and female potential (subsequent) smolts were larger than the potential parr in the previous spring, the minimum lengths of the former being around 90–100 mm f.l. Specific growth rates of the potential smolts remained relatively high after summer through to the following spring, whereas those of the potential parr decreased after summer, despite having been similar between the two groups during the previous spring and summer. These observations revealed that the developmental triggering of smelting possibly occurs in or following the summer season.     

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

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

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.     

Adaptations to Stochastic Environmental Variations: The Effects of Seasonal Temperatures on the Migratory Window of Svalbard Arctic Charr

The circumpolar Arctic charr, Salvelinus alpinus, is ideal for studying how environmental factors affect life history in fishes. Charr populations demonstrate a tremendous ecological plasticity and adaptations to harsh environments. Arctic charr is the only freshwater fish on Svalbard, including anadromous, resident, and landlocked stocks. Freshwater lake systems on Svalbard are characterized by very low water temperatures, long-term or even permanent ice cover, and low levels of nutrients. Food is thus limited and may lead to growth stagnation and early maturity in Arctic charr. The individual growth pattern may alternatively follow a sigmoid-shaped curve, caused by a shift to either cannibalistic or anadromous (migration to sea) behaviour. In lake systems that include migratory charr, the population may consist of a mixture of parr, postsmolt, and adult migratory individuals, as well as small-sized resident, large-sized resident (cannibals), and large formerly resident individuals transformed to anadromy. Our study in the Lake Dieset watercourse (79°N), Svalbard, demonstrates that the annual water flow in the outlet river is strongly correlated to air temperatures and provides a passage to the sea, allowing the charr access to the nutrient-rich seawater environment, during at most two months each year. During one of the years studied, the youngest and small-sized part of the sea-going stock was prevented from ascending the river and probably suffered mortality during winter. The migratory window of the Arctic charr in Lake Dieset is therefore highly variable among years and thus unpredictable. We hypothesize that in worst case scenarios (cold years, low water discharge), climatic variations may occasionally prevent charr from migrating upstream in Svalbard lake systems in late autumn, resulting in high mortality in the population.     

Smolting characters in anadromous and resident Arctic charr, Salvelinus alpinus (L.)

Morphological differences, haematocrit value and chloride cells were examined in downstream migrating Arctic charr, Salvelinus alpinus (L.) from the Hals River, North Norway, and resident charr from Lake Storvann. Fish were classified as visual parr, silvery parr or smolt based on the degree of silvering and the Occurrence of lateral parr marks. On average, 47% of downstream migrating Arctic charr were classified as visual molts, but only 14% of the resident charr. Charr longer than 20 cm fork length were mainly classified as visual molts. Morphometrical analyses of body size and shape revealed that most of the variations could be explained in terms of variations in fork length. Length adjusted ratios of post-anal distances were significantly higher in migrating charr than in resident charr. The number of developed chloride cells, the cell nucleus diameter in the basin of secondary gill lamellae and blood haematocrit values were significantly higher among downstream migrating charr than in resident charr. However, morphological differences between anadromous and resident Arctic charr were not necessarily synchronized with the development of chloride cells.     

Temperature preference of juvenile Arctic charr originating from different thermal environments

Temperature preference of juvenile (age 1+) Arctic charr (Salvelinus alpinus L.) originating from four arctic and sub-arctic populations (Svalbard and mainland northern Norway), representing a range of habitats with different temperature conditions, was studied by use of a shuttle-box system which allowed individual fish to control their environmental temperature. Based on the assumption that adaptations to long-lasting differences in thermal environments would affect temperature preference, we expected that Arctic charr from the high arctic Svalbard would prefer a lower temperature than the charr from two well-studied sub-arctic mainland lakes (i.e. one anadromous charr population from Storvatn, Hammerfest and two sympatric resident charr morphs from Fjellfrøsvatn, Målselv). There were, however, no significant differences in temperature preference among the four populations after 24 h exposure to the shuttle-box system, although the charr from the omnivore upper-water sympatric morph of Fjellfrøsvatn used significantly longer time to reach a stable thermal preferendum than the fish of the other populations. The average temperature preference at the end of the trials ranged between 10.9 and 11.6 °C among the populations. The lack of population differences suggests that temperature preference is not a polymorphic trait under strong selection in Arctic charr.     

Temperature requirements of Atlantic salmon Salmo salar, brown trout Salmo trutta and Arctic charr Salvelinus alpinus: predicting the effects of climate change

Atlantic salmon Salmo salar, brown trout Salmo trutta (including the anadromous form, sea trout) and Arctic charr Salvelinus alpinus (including anadromous fish) provide important commercial and sports fisheries in Western Europe. As water temperature increases as a result of climate change, quantitative information on the thermal requirements of these three species is essential so that potential problems can be anticipated by those responsible for the conservation and sustainable management of the fisheries and the maintenance of biodiversity in freshwater ecosystems. Part I compares the temperature limits for survival, feeding and growth. Salmo salar has the highest temperature tolerance, followed by S. trutta and finally S. alpinus. For all three species, the temperature tolerance for alevins is slightly lower than that for parr and smolts, and the eggs have the lowest tolerance; this being the most vulnerable life stage to any temperature increase, especially for eggs of S. alpinus in shallow water. There was little evidence to support local thermal adaptation, except in very cold rivers (mean annual temperature <6·5° C). Part II illustrates the importance of developing predictive models, using data from a long-term study (1967-2000) of a juvenile anadromous S. trutta population. Individual-based models predicted the emergence period for the fry. Mean values over 34 years revealed a large variation in the timing of emergence with c. 2 months between extreme values. The emergence time correlated significantly with the North Atlantic Oscillation Index, indicating that interannual variations in emergence were linked to more general changes in climate. Mean stream temperatures increased significantly in winter and spring at a rate of 0·37° C per decade, but not in summer and autumn, and led to an increase in the mean mass of pre-smolts. A growth model for S. trutta was validated by growth data from the long-term study and predicted growth under possible future conditions. Small increases (<2·5° C) in winter and spring would be beneficial for growth with 1 year-old smolts being more common. Water temperatures would have to increase by c. 4° C in winter and spring, and 3° C in summer and autumn before they had a marked negative effect on trout growth.     

Body size and the adoption of a migratory tactic in brook charr

The presence of a conditional strategy based on size attained before migration and a sex-ratio at migration biased towards females were explored in a population of brook charr Salvelinus fontinalis in the Sainte-Marguerite River, Quebec Province, Canada, where anadromous and resident forms live sympatrically. Seaward migration in the system occurred at 1 and 2 years old. Comparisons between backcalculated size-at-age of migrant and resident fish revealed that smaller fish at age 1 year delay migration to the following year and that bigger fish either migrated at age 1 year or remained resident for the rest of their life-cycle. Slow growth was associated with migration later in life (age 2 years) at a bigger size, which is consistent with the hypothesis of a threshold size for migration. No difference in size at age 1 year between migrant and resident fish suggests that other factors, such as growth efficiency and the presence of heritability of the tactics, are involved. Overall the sex ratio was equal for migrant and resident fish, while an age-specific bias was found: more males migrated at age 1 year and more females at age 2 years. These differences suggest that different tactics are adopted by different sexes.     

The Effect of Carlin Tags on Survival and Growth of Anadromous Arctic Charr, Salvelinus Alpinus

Fish tagging methods are important tools in the research and management of salmonids; the Carlin tag is widely used, especially in anadromous salmonids. However, this tagging method may interfere with fish behaviour and increase post-smolt mortality. To estimate Carlin-tag-inflicted mortality in Arctic charr, we used jet injection of alcian blue (colour tags) as a control. Recapture rates were higher for the colour-tagged fish than for the Carlin-tagged fish for both one- and two-year-old smolts. For one-year-old smolts in 1992, the recapture rate was 18.9% for colour-tagged fish and 10.0% for Carlin-tagged ones. For two-year-old smolts in the same year, 34.0% of colour- and 28.6% of Carlin-tagged fish were recaptured; in 1993, recapture rates were 17.2% and 6.3% for colour- and Carlin-tagged fish, respectively. Colour-tagged one-year-old Arctic charr also grew faster than their Carlin-tagged counterparts. No differences in growth rates were found between tagging methods in two-year-olds. These results imply that for hatchery-reared Arctic charr, survival rates using Carlin tagging may be half to one third that of colour tagging. Growth was also higher among colour-tagged fish, indicating that Carlin tags alter the behaviour of fish in the sea.     

PCB impairs smoltification and seawater performance in anadromous Arctic charr (Salvelinus alpinus)

The impacts of polychlorinated biphenyl (PCB) exposure on smoltification and subsequent seawater performance were investigated in hatchery-reared, anadromous Arctic charr (Salvelinus alpinus). The fish were subjected to a 2-month summer seawater residence, after which they were orally dosed with 0 (Control, C), 1 (Low Dose, LD) or 100 mg Aroclor 1254 kg(-1) body mass (High Dose, HD) in November. They were then held in fresh water, without being fed (to mimic their natural overwintering in freshwater), until they had smolted in June the next year. The smolts were then transferred to seawater and fed to mimic their summer feeding residence in seawater, followed by a period without food in freshwater from August until maturation in October. Compared with C and LD charr, the HD charr had either a transient or a permanent reduction in plasma growth hormone, insulin-like growth factor-1, and thyroxin and triiodothyronine titers during the period of smoltification. These hormonal alterations in the HD charr corresponded with impaired hyposmoregulatory ability in May and June, as well as reduced growth rate and survival after transference to seawater. Consequently, fewer fish in the HD group matured in October compared to the other two treatments. The HD fish had a liver PCB concentration ranging between 14 and 42 mg kg(-1) wet mass, whereas there were similar, and very low, liver PCB concentrations in LD and C fish throughout the smolting period. Our findings suggest that PCB might compromise mechanisms important for fitness in a fish species living in an extreme environment.     

Brown trout Salmo trutta released to support recreational fishing in a Norwegian fjord

One- and two-year-old hatchery reared juveniles of seven freshwater resident and anadromous populations of Scandinavian brown trout were released in the outer and inner part of the Oslofjord and in the River Akerselva, flowing through the city of Oslo. Recapture rates were highest (mean 20.3%) for river released fish and lowest for those released in the outer (16.8%) and inner (12.1%) fjord. In general, recapture rate increased with fish size at release (r=0.76). When released in fresh water, most of the recaptures were from fresh water and when released in the fjord, most recaptures were from the fjord. In general, freshwater resident stocks showed a higher degree of freshwater residency than anadromous stocks. However, mean migratory distance was longer for freshwater resident than anadromous fish. Trout moved longer distances at sea when released in the outer than in the inner fjord. Specific growth rate and size at recapture varied among release sites and stocks; they were highest for fish released in the outer fjord and lowest for river-released trout. There was no consistent difference in sea growth between freshwater resident and anadromous stocks. Estimated total yield was highest for fish released in the outer fjord, whereas there was no significant difference in yields between trout released in the river and the inner Oslofjord.     

Choice of feeding station in Atlantic salmon, Salmo salar, parr: effects of predation risk, season and life history strategy

The current speed at which underyearling salmon parr held feeding stations was examined from late summer to early winter in laboratory flume tanks that offered a choice between (a) areas with high water flow, high food availability but high predation risk and (b) areas with low flow, little food but shelter from predators. In August, those fish that would become smolts aged 1 + (and which by late winter formed the upper modal group, UMG, of the bimodal size distribution) adopted positions in faster currents than did the fish which would take a further year to reach the smolt stage (the lower modal group, LMG). However, the chosen current speed of UMG fish decreased through the period of study, so that by December all fish were found in areas of low flow, and hence little food. Both date and water temperature had independent effects on the chosen current speed of UMG fish.
The effect of predation risk was investigated using a model trout. A brief sight of this predator caused 47% of fish that had been in the main, exposed currents to move to slacker, sheltered areas; they took 1 h, on average, to return to their previous position. The fish that remained in position upon seeing the predator reduced their rate of tail beating, presumably increasing crypticity. Eventual UMG parr were less likely than were LMG fish to move away upon seeing a predator. The fish moved to faster currents than normal 2–3 h after seeing the predator, possibly compensating for the earlier reduction in feeding rate.     

Polymorphism and speciation in Arctic charr

The Arctic charr Salvelinus alpinus exhibits 1–4 sympatric morphs in postglacial lakes, of which one or two are epibenthic zoobenthos feeders, one is a limnetic planktivore and one is a piscivorous form. In addition, northern rivers support partly migratory populations with anadromous and freshwater resident fish. The morphs vary in their coloration, morphology, life history, behaviour and genetic characteristics. The morphs usually differentiate according to their ontogenetic stage at maturity, which parallels paedomorphism in amphibians. The young usually start as epibenthic zoobenthivores, but may become pelagic at a certain size according to the predation risk experienced at that time. From a length of > 20–25 cm, charr start to become piscivorous. The proportion of piscivorous fish increases with increasing body size. In partly anadromous populations, fish that mature before smolting become freshwater resident, the others anadromous. In some rivers, the morphs occupy separate niches (epibenthic and limnetic), from emergence onwards. The morphs exhibit different degrees of reproductive isolation that vary from a high degree of interbreeding to complete isolation. Usually, they spawn within morph (assortative mating), but alternative male mating behaviour (sneaking, fighting) may occur in stream‐spawning populations and at great depths in lakes. Morphologically specialized morphs appear to feed more effectively than intermediate forms, and selection according to feeding mode, site fidelity and associated assortative mating are prerequisites for the evolution of the different morphs. Charr morphs develop into stable feeding niches under conditions of intense intraspecific competition when there is little competition with other species. Sympatric morphs exhibit different degrees of speciation, but similar morphs in different systems are not individual species because of (1) their polyphyletic origin, (2) the supporting systems are often young, transient environments making the future situation for the populations uncertain, and (3) the genetic differentiation among morphs is low. Sympatric morphs may interbreed and produce fertile hybrids. Nevertheless, sympatric charr morphs should be managed as separate species. Changes in the natural conditions or human impacts to which the morphs are adapted will have a strong influence on the persistence and survival of each different morph.     

Population structure, ecological segregation and reproduction in non-anadromous Arctic charr, Salvelinus alpinus (L), in four unexploited lakes in the Canadian high Arctic

Land-locked populations of Arctic charr in four lakes on Northern Ellesmere Island (80° N) were found to consist of two distinct sizes: 'dwarf' and 'normal' charr. Both groups attained sexual maturity but differed in appearance and habitat. The smaller fish, occupying the more marginal habitats, retained their parr-markings; the larger group had the general characteristics of smolts, being more silvery and without parr-marks. In their juvenile stages, the charr destined to attain the larger group were indistinguishable from members of the smaller group. Although fish in the larger group were capable of cannibalism, this was rarely observed. In general, the fish in the larger group were older than the smaller ones but great variation in size at a given age resulted in certain age classes containing representatives of both groups. The population structure varied considerably between lakes; a high proportion of 'normal' charr correlated well with a high growth rate in the first few years of development. It is postulated that the two groups live in dynamic equilibrium where the advantages of progenesis (retention of juvenile characters by adults) in the smaller type are traded against the larger proportion of the energy resources available to the larger type. The concept of heterochrony in an ecological setting is introduced.     

Life histories of Atlantic salmon altered by winter temperature and summer rearing in fresh- or sea-water

Increased growth during winter increased the incidence of age 1+ Salmo salar smolts in spring. High condition factor in spring and good growth in summer was associated with a high incidence of sexually mature males in autumn. In two experiments, groups (n=160–237 per group) of individually identified parr, either ungraded (lower and upper modal groups: LMG, UMG) or size-graded (LMG only), were reared at either 10, 6 or 3 °C overwinter (Nov to May). At 10 °C, up to 51% of parr originally in the LMG became smolts in spring at age 1+. In contrast, at either 6 or 3 °C (control), < 6% of LMG parr became smolts. The probability of being recruited into the UMG overwinter was positively related to initial body size, and was increased by size-grading. Smolt recruitment was two-fold higher among females compared to males; a proportion of males by age 0+ had already opted to mature at age 1+ rather than smolt at age 1+. In contrast, smolting at age 1+ was not inhibited in males previously mature at age 0+. During summer (May to Nov), all experimental groups were reared at ambient temperature, each subdivided between fresh water (max 21 °C) or seawater (max 15 °C). Good growth in seawater of winter recruits into the UMG confirmed they had completed smolting. Mortality in seawater among parr was 41–83%, and among smolts was 10–22%. Specific growth rate during summer was inversely related to winter rearing temperature. The incidence of sexual maturity in autumn at age 1+ among male parr was positively related to winter rearing temperature, fork length and condition factor in May, but there was large variation among individuals with respect to body size and maturity. Summer rearing in seawater reduced growth and the incidence of maturation. Parr and post-smolt maturity was 84–99% and 100% in fresh water respectively, 21–58% and 0% in seawater.     

Genetic architecture of growth and early life‐history transitions in anadromous and derived freshwater populations of steelhead

Heritabilities of growth, precocious maturation and smolting were measured in 75 families of juvenile steelhead or rainbow trout Oncorhynchus mykiss , progeny of within and between line matings (crosses) of wild, anadromous steelhead and wild, resident (lake) rainbow trout originally derived from the same anadromous stock 70 years earlier. The tagged yearling progeny were combined by line in common freshwater rearing containers and graded into three categories: mature, smolt or rearing (undifferentiated) at age 2 years. Heritabilities of precocious male maturity, smolting and growth were moderate to high, and the genetic correlation between growth and smolting was low. Smolting and precocious male maturity were highly variable among families within lines and significantly different between lines. Each of the four lines produced significant numbers of smolts at age two. Smolting and maturation were negatively genetically correlated, which may explain the persistence of smolting in the lake population despite strong selection against lake smolts; balancing selection on male maturation age may help to maintain variation for smolting. The high heritability of smolting, coupled with the inability of smolts that leave the lake to return to it indicates that the genetic potential for smolting can lie dormant or be maintained through a dynamic interaction between smolting and early maturation for decades despite complete selection against the phenotype. The results have significant implications for the preservation of threatened anadromous stocks in fresh water and the inclusion of resident fish of formerly anadromous populations, currently trapped behind long‐standing barriers to migration, as one component of the same population.     

Age composition of masu salmon smolts in northern Japan

Smolts of anadromous masu salmon Oncorhynchus masou aged 3+ years were found in a northern Japanese river. This is the first recording of 3+ year smolts in Japan. These fish appeared to originate in the cold upper river where 2+ year parr were found during autumn.