The upper critical thermal limits for alevins of Arctic charr from a Norwegian lake north of the Arctic circle

Mean values ±95% CL of the upper incipient (T_IL) and ultimate (T_UL) lethal temperatures, determined at five acclimation temperatures ( T_A ), increased for T_IL from 19.2 ± 0.4° C ( T_A 0.5° C) to 21.0 ± 0.4° C ( T_A 20° C), and for T_UL from 22.6 ± 0.1° C ( T_A 0.5° C) to 26.6 ± 0.4° C ( T_A 20° C). Mean values were close to those obtained for Arctic charr alevins from Windermere (north-west England). These comparative data for alevins, and previous data for 0+ year parr, indicate negligible geographical variation in the thermal limits of Arctic charr.     

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 impact of stocking on the genetic integrity of Arctic charr (Salvelinus) populations from the Alpine region

There is a long tradition of artificially stocking lakes and rivers with fish in the hope to increase the quality and amount of fish that can be harvested. The animals used for stocking often originate in distant hatcheries or hatcheries that breed fish from remote regions. This stocking practice could have effects on the genetic integrity of resident populations. We have analysed here a case of the influence of stocking on Alpine populations of the Arctic charr (usually included into Salvelinus alpinus, but revised to Salvelinus umbla Kottelat 1997 ) within a unique geographical and historical setting. The Königssee in the Bavarian Alps (Berchtesgaden) was heavily stocked several times during the last century. However, a sample of the ancestral Königssee population still exists in the Grünsee, which lies close to Königssee, but 1000 m higher. To trace the influence of stocking in Königssee we have analysed more than 300 individuals from 10 lake populations, including the source populations of the fish used for stocking. From these we have sequenced a part of the mitochodrial control region and have typed them at six microsatellite loci. The differential distribution of haplotypes, as well as assignment tests, show that the influence of stocking on the genetic integrity of the Königssee population has been negligible. However, our data reveal that in another lake included in our study (Starnberger See), the ancestral population was apparently replaced completely by the populations used for stocking. The major difference between the lakes is the relative preservation of ecological integrity. Königssee was ecologically stable in the past, whereas Starnberger See was heavily polluted at one point, with charr approaching extinction. Interestingly, in a lake neighbouring Starnberger See, the Ammersee, which was also subject to strong pollution but not stocked, the ancestral population has recovered. Our data suggest that the practice of artificial stocking should be reconsidered, or at least monitored for effectiveness.     

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.     

Microsatellite and mitochondrial DNA assessment of population structure and stocking effects in Arctic charr Salvelinus alpinus (Teleostei: Salmonidae) from central Alpine lakes

Despite geographical isolation and widespread phenotypic polymorphism, previous population genetic studies of Arctic charr, Salvelinus alpinus , have detected low levels of intra- and interpopulation variation. In this study, two approaches were used to test the generality of low genetic diversity among 15 Arctic charr populations from three major drainages of the central Alpine region of Europe. First, a representative subsample of each drainage was screened by PCR–RFLP analysis of mtDNA using 31 restriction enzymes. All individuals but one shared an identical haplotype. In contrast, microsatellite DNA variation revealed high levels of genetic diversity within and among populations. The number of alleles per locus ranged from six to 49, resulting in an overall expected heterozygosity from 0.72 ± 0.09 to 0.87 ± 0.04 depending on the locus. Despite evidence for fish transfers among Alpine charr populations over centuries, genetic diversity was substantially structured, as revealed by hierarchical Φ statistics. Eighteen per cent of total genetic variance was apportioned to substructuring among Rhône, Rhine, and Danube river systems, whereas 19% was due to partitioning among populations within each drainage. Cluster analyses corroborated these results by drainage-specific grouping of nonstocked populations, but also revealed damaging effects of stocking practices in others. However, these results suggest that long-term stocking practices did not generally alter natural genetic partitioning, and stress the importance of considering genetic diversity of Arctic charr in the Alpine region for sound management. The results also refute the general view of Arctic charr being a genetically depauperate species and show the potential usefulness of microsatellite DNAs in addressing evolutionary and conservation issues in this species.     

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.     

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.     

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.     

A non-destructive morphometric technique for estimation of body and mesenteric lipid in Arctic charr: a case study of its application

Weight and eight linear measurements were made on Arctic charr from the domesticated Hammerfest strain from Norway and offspring of wild charr of a pelagic morph from Loch Rannoch, Scotland. Guts and mesenteries were removed from the Hammerfest charr only, and the amount of lipid in both carcass and mesenteries measured by Soxhlet extraction. Lipid was extracted from the whole body of the Loch Rannoch charr. Multiple regression analysis was used to derive morphometric predictors of total lipid for the Hammerfest charr and percentage body lipid for the Loch Rannoch charr, the regressions explaining 83 and 59% of variance respectively. For the Hammerfest charr, multiple regression also provided a reliable predictor of mesenteric fat, accounting for 65% of its variance. Hammerfest charr that exhibited high aggression rates had 53% more whole body lipid and 100% more mesenteric fat than those with low aggression rates, using direct measures of lipid levels. Indirect, morphometrically-derived measures of lipid levels gave almost identical results. It is concluded that morphometric techniques can provide estimates of both whole body and mesenteric lipid in studies requiring repeated measures on the same individuals.     

State-dependent feed acquisition among two strains of hatchery-reared Arctic charr

The effects of genetic and environmental factors on aggression and feeding hierarchies were studied using X-radiography to measure food intake by hatchery-reared individuals of two strains (Hammerfest and Svalbard) of Arctic charr Salvelinus alpinus . A reduction in food rations and/or water current speed increased intraspecific aggression, and both factors led to increased interindividual variability in food intake, increasing the coefficient of variation (CV). Following a return to pre-manipulation conditions, CVs decreased to their original level. In control groups, CVs and share of group meals were stable throughout the experiment. The increase in CVs following manipulation was the result of a small number of dominant individuals obtaining a high share of the meal. Restriction in food ration affected share of meals, specific growth rates and the frequencies of non-feeding fish, while reductions in water current speed affected only share of meals. Feeding hierarchies were size-dependent in the control groups. In contrast, no relationships between body weight and feeding rank were evident in groups in which food ration or water current speed were reduced. A small, but consistent, difference was revealed in feeding hierarchy responses between the two strains.     

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.     

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.     

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.     

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.     

Cannibalism and protective behavior of eggs in Arctic charr (Salvelinus alpinus)

From video recordings of spawning events, we quantified protective and cannibalistic behavior of Arctic charr occurring immediately after spawning. The number of fish cannibalizing on stray eggs was examined regarding (a) whether more than one male shed milt during the spawning event, that is, whether sperm competition occurred, (b) whether the sperm competition included few or many males, that is, the intensity of sperm competition, and (c) the density of fish at the spawning site. Response behavior toward egg cannibalism was also examined among females and dominant males in order to determine any parental investment toward protecting the eggs after spawning. Cannibalistic behavior was seen in almost 50% of the spawnings, and the multiple spawning events showed the highest numbers of fish cannibalizing on eggs. Both the number of males releasing milt and the number of fish approaching the spawning site were positively correlated with egg cannibalism. Sperm competition was, however, not a prerequisite for egg cannibalism. Although we also observed partial filial cannibalism, protective behavior of eggs was seen both among dominant males and females, suggesting that charr actually conduct parental care.     

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.     

Heterozygote deficiencies in small lacustrine populations of brook charr Salvelinus Fontinalis Mitchill (Pisces,Salmonidae): a test of alternative hypotheses

Empirical studies of natural populations have commonly reported departures from Hardy-Weinberg expected proportions of heterozygote individuals. Recent advances in statistical population genetics now offer the potential to exploit individual multilocus genotypic information to test more rigorously for possible sources of heterozygote deficiencies. In a previous study in lacustrine brook charr (Salvelinus fontinalis), we reported stronger deficits in small than in large lakes. In the present paper, we propose a methodology for empirically testing alternative hypotheses to identify the cause of the deficits observed in three of the smallest lakes (85, 109 and 182 ha) analysed. First, as in several salmonid species, brook charr may exhibit a trophic polymorphism in north temperate lakes. If morphs are genetically divergent, indiscriminate sampling of both forms would result in less heterozygote individuals than expected in a randomly mating population (Wahlund effect). Using an individual-based method aiming at detecting cryptic population structure, we can reject this explanation as the sole source of deficits for all three lakes. Secondly, mating among relatives could also be frequent in small lakes and lead to heterozygote deficiencies. Significantly more fish than expected at random had low individual multilocus heterozygosity in two of the lakes, suggesting that inbred fish may have been present. Thirdly, sampling of genetically related fish would also lead to departures from Hardy-Weinberg proportions. In the same two lakes, the distribution of pairwise relatedness coefficients departed from its random expectation, suggesting that non-random sampling of kin may have occurred.     

Variations in juvenile growth, energy allocation and life-history strategies of two populations of Arctic charr in North Norway

Within the populations of Arctic charr Salvelinus alpinus in Storvatn and Rungavatn, Norway, fish that attained the largest size in the late parr stage migrated as 4+ smolts, fish that attained a medium size became 5+ smolts, and the smallest parr became lake residents. Within the last 2 years of the parr stage, those that became anadromous had a lower growth rate during the winter and early summer than those that became resident, and vice versa in late summer. Thus, anadromous fish grew faster in the parr stage, but in years prior to migration, their growth pattern seemed to differ from that of parr that became resident. Due to early size differences between parr that became anadromous or resident, it is suggested that some of the basis for the decision to smoltify or not in charr depends on the growth rate through the whole parr stage, but with significant modification by the local environment. Rungavatn parr had a significantly higher population density (5 ×), higher lipid content and mortality, an earlier maturation, a lower rate of growth, and only one-third the degree of anadromy than those from Storvatn. It is suggested that the two charr populations have evolved different life-history adaptations due to unequal growth, energy allocation and competition opportunities in the lakes.