Parasites of freshwater resident and anadromous Arctic charr (Salvelinus alpinus) in Greenland

Twenty-one metazoan parasite species were found in Arctic charr at three coastal sites in west Greenland, including Neascus sp. and the nematode Pseudoterranova decipiens , new records for Arctic charr. Twelve species are new records for Arctic charr in Greenland. The quantitative composition of the parasite communities varied, but the freshwater species ( Diphyllobothrium ditremwn, Eubothriutn salvelini, Proteocephalus longicollis, Crepidostotnutn farianis ) were usually numerically dominant, even in anadromous fish. The absence from Greenland of Mysidacea, Amphipoda, Ephemeroptera and Odonata excludes parasites requiring such organisms as intermediate hosts, including the cestode Cyathocephalus truncatus , nematodes of the genus Cystidicola , and freshwater acanthocephalans.     

Phenotypic variation in growth trajectories in the Arctic charr Salvelinus alpinus

Animals with determinate growth have shown little variation in individual growth patterns, but similar analyses for animals with indeterminate growth have been lacking. We analysed the amount of phenotypic variation in growth patterns across ages among individuals of a hatchery-based population of Arctic charr, Salvelinus alpinus, Salmonidae, using the infinite-dimensional model and including the effects of group size structure. There was little phenotypic variation in growth trajectories: individuals that were small (in relation to the mean) early in life were among the smallest 2.5 years later. If the genetic variation reflects phenotypic variation, not much evolutionary change can be expected. Our results show that there are ecological conditions that determine the strong covariation of size across ages, most likely size-related dominance behaviour, which can mask the true variation of growth patterns. Thus, social interactions can have strong evolutionary effects on traits not directly involved in the behavioural interactions.     

Mitochondrial DNA analysis of Scottish populations of Arctic charr, Salvelinus alpinus (L.)

Mitochondrial DNA analysis of a total of 82 individuals from seven Scottish populations of Arctic charr ( Salvelinus alpinus L.) using restriction fragment length polymorphism and direct nucleotide sequencing of the cytochrome b gene revealed little variation within or between populations. One of 19 restriction enzymes showed a polymorphism that was present in three of the seven populations. Only one nucleotide substitution was detected in a 288 base pair segment of the cytochrome b gene examined in 26 fish. Approximately 4% of the mitochondrial genome was surveyed with these complementary procedures.     

Gyrodactylus salaris infecting allopatric Arctic charr Salvelinus alpinus fry: an experimental study of host survival

Artificially reared Arctic charr Salvelinus alpinus fry (post yolk‐sac stage) from Skibotnelva in northern Norway were exposed to infection with the monogenean ectoparasite Gyrodactylus salaris to determine if the parasite caused mortality in fry. About 95% of the fry became infected within the first week post‐exposure, and at the end of the experiment (77 days) all fry carried the parasite. Mortality rate was significantly higher in the groups of infected fry than in the uninfected control groups, and when the experiment was terminated c. 30% of the infected and 8% of the uninfected fish had died. Gyrodactylus salaris had a pronounced negative effect upon fry survival and caused an additive host mortality. Moreover, the concurrent temporal changes in parasite intensities and aggregation indicated that the parasite‐induced host mortality was density dependent.     

The development of regular seasonal habitat shifts in a landlocked Arctic charr, Salvelinus alpinus L., population

Parts of the Arctic charr population of the subarctic Lake Visjön in north-west Sweden migrate upstream during the spring to two small, recently eutrophied and very productive lakes. Large repeat migrants arrive first, followed by young first-time migrants. Charr in the small lakes grow more rapidly than those resident in L. Visjön. In early September mature fish leave the lakes, followed by immature fish later in September and in October. Overwintering and spawning takes place in L. Visjön. Migratory females attain maturity at age 4 years and resident females at age 6 years. The migrant fish return annually until they are 5–6 years old. This limit may be due to reduced relative growth benefits of the habitat shift for larger individuals. The rapid development of these regular habitat shifts could be explained by an internally fixed exploratory behaviour in these Arctic charr that makes the detection and utilization of distant feeding resources possible. Migrants will possess a considerably higher fitness, if survival rates for migratory and resident fish are equal.     

Salinity tolerance of freshwater acclimated, small-sized Arctic charr, Salvelinus alpinus from northern Labrador

Arctic charr, Salvelinus alpinus , less than 150 mm in size were frequently captured at sea in northern Labrador in areas where salinities of 30‰ or higher had been recorded. These captures were inconsistent with many earlier reports for other areas that indicated Arctic charr less than 150 mm in size were not found at sea. A series of salinity challenge tests was carried out in the field, using wild Ikarut River charr, and in the laboratory, using cultured Fraser River charr, to understand more about the potential fate of these small fish. The results of challenge tests with small Arctic charr (< 120 mm) indicated that at intermediate salinities (10–20‰), these fish can readily survive. In laboratory tests with salinities at 30‰, survival was size dependent and would suggest that in natural situations, small charr would require periodic access to fresh or brackish water to stay alive. The influence of water temperature on salinity tolerance may be important when fish are exposed to temperatures that are below 0° C.     

The periodicity of primary increment formation in the otoliths of Arctic charr Salvelinus alpinus (L.)

The role of environmental factors in the regulation of sub-annual increment formation in fish otoliths appears to differ markedly between species. To examine the periodicity of primary increment formation in the otoliths of O + Arctic charr, Salvelinus alpinus (L.), and the effects of temperature, and photoperiod on their formation, fish were held under controlled environmental conditions. Primary growth increments were found in the otoliths of fish held at constant temperature (18° C) and at ambient temperature [fluctuating with a circadian and circannual rhythm (4–18° C)]. Consistent and significant disruptions in increment formation occurred however, in experimental groups subjected to rapid change from ambient photoperiod to a 6L: 6D photo-period for 96 h. Disruptions in increment formation were also observed immediately following transportation of fish between holding facilities and following disease treatment. The number of otolith increments formed in fish held on an ambient photoperiod regime, correlated closely with time elapsed in days since checkmark formation ( r = 0.989, P ≤0.001) in fish sampled sequentially over a period of 10 to 105 days. Thus we demonstrate that under conditions of ambient photoperiod, primary increments are formed daily.     

Seawater acclimation of diploid and triploid brook charr (Salvelinus fontinalis), diploid Arctic charr (Salvelinus alpinus), and their diploid and triploid hybrids

Seawater acclimation of diploid (FF) and triploid (F2F) brook charr Salvelinus fontinalis , diploid (AA) Arctic charr Salvelinus alpinus , and diploid (FA) and triploid (F2A) hybrids between female brook charr and male Arctic charr was investigated. Triploidization of brook charr and the hybrid did not have any effect on the acclimation. Seawater acclimation of the hybrid was achieved during the experimental period and was comparable to that observed in brook charr. Acclimation could not be ascertained in Arctic charr since the level of cortisol, a stress indicator, was still high at the end of the experiment. No relationship between either length or condition factors and plasma osmolality was observed. Elevated plasma cortisol concentrations in Arctic charr and in diploid or triploid hybrids, both in fresh water and sea water, indicate more favourable rearing conditions for brook charr.     

The effect of stocking density on the behaviour of Arctic charr (Salvelinus alpinus L.)

Recent studies have demonstrated that Arctic charr ( Salvelinus alpinus ) exhibit higher growth rates when held at high stocking densities. It has been argued that these increased growth rates are a result of decreased social interactions at the increased densities. The current study was designed to test this hypothesis by examining: (1) the frequency of agonistic interactions, (2) the proportion of time spent swimming and (3) growth rates among groups of young of the year Arctic charr held at different stocking densities. It was found that charr reared at higher densities exhibited significantly higher mean weights and lengths than those at the lower densities. The behavioural observations revealed that charr initiated significantly fewer agonistic interactions and spent a significantly greater amount of time shoaling at high densities than charr held at the medium or low densities. It is argued that these behavioural responses may serve to lower energy expenditures, resulting in the increased growth rates.     

Holarctic phylogeography of Arctic charr (Salvelinus alpinus L.) inferred from mitochondrial DNA sequences

This study evaluated mitochondrial DNA (mtDNA) sequence variation in a 552-bp fragment of the control region of Arctic charr (Salvelinus alpinus) by analyzing 159 individuals from 83 populations throughout the entire range of the complex. A total of 89 (16.1%) nucleotide positions were polymorphic, and these defined 63 haplotypes. Phylogenetic analyses supported the monophyly of the complex and assigned the observed haplotypes to five geographic regions that may be associated with different glacial refugia. Most notably, a formerly defined major evolutionary lineage (S. a. erythrinus) ranging from North America across the Arctic archipelago to the Eurasian continent has now been partitioned into the Arctic group and the newly identified Siberian group. The Beringian group, formed entirely by specimens assigned to S. malma (Dolly Varden), encompassed the area formerly assigned to S. a. taranetzi. The latter, due to a unique haplotype, became the basal member of the Arctic group. Overall, the S. alpinus complex reflects divergent evolutionary groups coupled with shallow intergroup differentiation, also indicated by an analysis of molecular variance that attributed 73.7% (P < 0.001) of the total genetic variance among groups. Time estimates, based on sequence divergence, suggest a separation of the major phylogeographic groups during early to mid-Pleistocene. In contrast, colonization of most of today's range started relatively recently, most likely late Pleistocene during the last retreat of ice sheets some 10,000-20,000 years ago. This time scale obviously is too shallow for detecting significant variation on a smaller scale using mtDNA markers. However, other studies using nuclear microsatellite DNA variation strongly suggested ongoing evolution within groups by revealing strong population-genetic substructuring and restricted gene flow among populations. Thus, Arctic charr could serve as a model organism to investigate the linkage between historical and contemporary components of phylogeographic structuring in fish, and, with a global perspective of the distribution of genetic variation as a framework, meaningful comparisons of charr studies at a smaller geographic scale will now be possible.     

Diel food selection of pelagic Arctic charr, Salvelinus alpinus (L.), and brown trout, Salmo trutta L., in Lake Atnsjø, SE Norway

Patterns of diel food selection in pelagic Arctic charr, Salvelinus alpinus (L.) and brown trout, Salmo trutta L. were investigated in Lake Atnsjo, SE Norway, by gillnet sampling during July-September 1985. Arctic charr feed almost exclusively on zooplankton both day and night, while brown trout had a diurnal shift in diet. For this species zooplankton made up a considerable part of the diet in the daytime, while at night the diet consisted mainly of surface insect and chironomid pupae. Both species had a selective feeding mode on zooplankton during the day and night. Arctic charr had a higher gill raker number and a denser gill raker spacing compared with brown trout. Still, the differences in prey size between the two species were small. We argue that the observed differences in food selection between Arctic charr and brown trout can be explained by differing abilities to detect food items under low light conditions.     

Population genetic structure of Arctic charr, Salvelinus alpinus from northwest Europe on large and small spatial scales

To examine the population genetic structure of lake-resident Arctic charr, Salvelinus alpinus from northwest Europe on multiple spatial scales, 2367 individuals from 43 lakes located in three geographical regions (Iceland, the British Isles and Scandinavia) were genotyped at six microsatellite loci. On a large scale, data provided little evidence to support clustering of populations according to geographical region. Hierarchical analysis of molecular variance indicated that, although statistically significant, only 2.17% of the variance in allelic frequencies was partitioned at the among-region level. Within regions, high levels of genetic differentiation were typically found between lakes regardless of the geographical distance separating them. These results are consistent with the hypothesis of rapid postglacial recolonization of all of northwest Europe from a single charr lineage, with subsequent restriction of gene flow. On a smaller scale, there was evidence for close genetic relationships among lakes from within common drainage basins in Scotland. Thus, interlake genetic structure reflects localized patterns of recent (or contemporary) gene flow superimposed onto a larger scale structure that is largely a result of historical processes. There was also evidence for widespread genetic structuring at the within-lake level, with sympatric populations detected in 10 lakes, and multilocus heterozygote deficits found in 23 lakes. This evidence of the Wahlund effect was found in all lakes known to contain discrete phenotypic morphs, as well as many others, suggesting that morphs may often represent separate breeding populations, and also that the phenomenon of polymorphism in this species may be more widespread than is currently realized.     

Diel and seasonal locomotor activity patterns in Arctic charr, Salvelinm alpinus (L.)

One-year-old Arctic charr, Sulvelinus alpinus (L.), of the Hornavan strain were tested from February 1985 to January 1986 in an attempt to get an increased understanding of the annual rheotactic behaviour as well as the die1 and seasonal locomotor activity pattern. An annular stream tank equipped with photocells was used to measure the direction of swimming movements as well as the number of passings. From February to late May the locomotor activity was low but increased in July and peaked in September. After November the locomotor activity was again at low winter levels. During the activity peak from July to November the majority ofall movements was directed against the current while no preference for direction was noted during the rest of the year. The high level of swimming movements directed against the current in late summer and autumn may be related to an innate habitat change. From February until June, the charr exhibited a bimodal diurnal activity pattern. In July activity was evenly spread over the whole 24- hour period and in August and September activity was again mainly diurnal with a bimodal pattern. In October and November the activity was mainly nocturnal and in December and January activity was concentrated in the short light period. Both annual and die1 activity are discussed in relation to earlier findings in general locomotor activity in Arctic charr and other salmonids.     

The effects of kinship on the growth of juvenile Arctic charr

The effect of kinship on growth and growth variability was studied by rearing young-of-the-year Arctic charr in full-sibling, mixed-sibling or non-kin groups for 11 weeks. Both weight and length were found to be significantly greater among full-sibling v. mixed or non-kin groups. Also, variance in weight and length of the individuals within groups was found to be significantly lower in full-sibling v. mixed or non-kin groups. These data suggest that relatedness of group members has a significant effect on both the growth rate and variation in growth of juvenile Arctic charr.     

The genetics of sympatric Arctic charr [Salvelinus alpinus (L.)] populations from Loch Rannoch, Scotland

Two morphologically and ecologically distinct forms of Arctic charr, Salvelinus alpinus , are found in Loch Rannoch, Scotland. The differences in morphology are adaptations to different modes of life, one being pelagic, the other benthic. Both forms have been the subjects of extensive genetic studies including cytogenetics, nuclear and mitochondrial DNA analysis, and protein electrophoresis. Significant differences between the two forms are revealed by some techniques but not others and provide evidence for the reproductive isolation of these two morphs. The findings are discussed in relation to the derivation of sympatry and the phylogenetics of Arctic charr.     

Seasonal patterns of growth, lipid deposition and lipid depletion in anadromous Arctic charr

Seasonal patterns of growth, lipid deposition and lipid depletion were studied in anadromous Arctic charr from a north Norwegian population. Samples were collected in late May when fish were migrating between fresh water and the sea, and in mid-July when the fish re-entered fresh water. A sample of maturing fish captured in mid-July was held in captivity until late September to assess lipid mobilization linked to the final stages of maturation. The fish increased substantially in weight during their 40–50-day summer residence in sea water (immatures from c . 300 to 600 g; maturing fish from c . 500 to 800 g), and body lipid stores were increased approximately fivefold. The carcass (head, skeleton and skin) was the major lipid depot, accounting for c . 50% of the total lipid content when the fish re-entered fresh water from the sea. The muscle of the ascending charr contained 35–40% of the body lipids, whereas the gut and liver each held 4–5%. Body lipid decreased 30–40% during the period between the re-entry of the fish to fresh water and spawning; although lipids were depleted from all depots the carcass and muscle were quantitatively the most important. By the time of spawning, the gonads of the females held almost 25% of the body lipid, whereas in the males the gonads accounted for <3% of the total lipids. Females lost c . 80% of their body lipid during spawning and overwintering, and, consequently, the lipid depots were severely depleted by the early spring. By contrast, lipid depletion in the males amounted to 50–55% of total lipid in the same period. These data suggest that the combination of spawning and overwintering in fresh water imposes a greater load on the females than on the males. It may be that the severity of the depletion prevents females from spawning in successive years.     

Effects of food quality on maturation rate in Arctic charr, Salvelinus alpinus (L.)

The effects of food quality on maturation rate were followed in progeny from wild Arctic charr, caught in Lake Fattjaure, northern Sweden. The fish were reared at five different food quality levels. In the first summer, the fry were given feed of three qualities: diluted with cellulose in three proportions (0, 15 and 25%). During the second winter the impact of changed food quality level was studied by transferring half the fish from the high (control) to the low food quality level and vice versa.
Maturation rate was lower in males reared at the high and improved food quality levels than in males reared at the medium, low and reduced food quality levels. The maturation rate in females was similar at all levels, though the rate tended to increase at the reduced food quality level.
Fish reared at the high and medium levels had similar growth rates, whereas fish at the low food quality level experienced slower growth. Reduced food quality did not arrest the growth offish, whereas improved food quality enhanced their growth.     

Piscivory by brown trout Salmo trutta L. and Arctic charr Salvelinus alpinus (L.) in Norwegian lakes

Size and frequency of occurrence of prey of brown trout Salmo trutta L. and Arctic charr Salvelinus alpinus (L.) were recorded in 13 Norwegian lakes during 1973–1990. Piscivores usually comprised less than 5% of the total population. Arctic charr were less piscivorous than brown trout. Trout and charr became piscivorous at 13 and 16 cm length, respectively. These size thresholds were similar to those of other facultative piscivorous freshwater fish species. When present, three-spined sticklebacks, Gasterosteus aculeatus (L.), were preferred by all length groups of piscivorous brown trout and Arctic charr. Length of prey increased with increasing predator length, and the mean body length of prey was about 33 and 25% of predator length for trout and charr, respectively. Yearlings of charr were not recorded as prey.     

The influence of environmental manipulations on inter– and intra–individual variation in food acquisition and growth performance of Arctic charr, Salvelinus alpinus

Repeated measurements of food intake made on juvenile Arctic charr, Salvelinus alpinus , held under different rearing conditions enabled examination of the effects of environmental manipulations on both intra– and inter–individual variations in food intake to be made. This permitted the assessment of the influences of differential food acquisition on individual growth rates and biomass gain. When charr were held in isolation individual fish showed relatively little day–to–day variability in food intake and inter–individual differences in intake were small ('base–fine' values). All fish exhibited positive rates of growth and the overall range was narrow. Nevertheless, there was a highly significant positive correlation between food intake and growth, indicating that those individuals that consumed the greatest quantities of food were also those that had the highest rates of weight gain. The rearing of charr in groups led to increases in both intra– and inter–individual variations in food intake to levels considerably above 'base–line'. This increased variability in food intake was reflected in rates of weight gain being more variable amongst the charr reared in groups, with fish that lost weight often being recorded. Manipulation of the rearing environment had marked influences upon intra–individual variability in food intake, inter–individual differences in food acquisition and rates of weight gain. High stocking densities and exposure of the fish to moderate water currents were most effective in reducing levels of variability to approach those observed under 'base–line' conditions.