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.     

Individual variation in distribution, activity and growth rate of Arctic charr kept in a three-tank system

The movements and distribution of groups of Arctic charr Salvelinus alpinus were examined in a rearing system that offered a choice between two different feeding tanks separated by a larger non-feeding tank. The passages of individual fish were monitored continuously during a period of 3 weeks using the PIT (passive integrated transponder)-tag technique. The primary aim was to examine if only some charr were occupying the feeding tanks, thereby excluding other individuals, and whether differences in visit activity explained within-group variation in individual growth. On average, about 35 of the 40 charr in each group shoaled in the large non-feeding tank leaving only five individuals in the feeding tanks. Charr that spent a long total time in one of the feeding tanks made frequent excursions to the other tanks resulting in a continuous exchange of individuals. Individual growth rates were correlated positively with visit activity rather than with the total time spent in the feeding tanks. Thus, individuals with low growth rate spent as much time in feeding tanks as charr with high growth rate. However, less successful fish tended to visit the feeding tanks at night when the feeders were switched off. Based on behavioural and growth results obtained in this experiment, the use of multitank systems in the cultivation of Arctic charr is discussed.     

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.     

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.     

Stable isotope analysis reveals ecological segregation in a bimodal size polymorphism in Arctic charr from Loch Tay, Scotland

The serendipitous discovery of a body-size dimorphism amongst the sexually mature Arctic charr Salvelinus alpinus of Loch Tay is described. Sexually mature Arctic charr, collected by gill netting on spawning areas, showed a clear and distinct bimodal size distribution with no overlap in fork length distributions. The upper (19–29 cm L _F) and lower modes (8–16 cm) were not solely the result of sex or age differences. Analysis of stable isotope ratios of C and N in muscle showed highly significant differences in mean δ¹³C and δ¹⁵N between populations, demonstrating a difference in trophic ecology between the two body-size morphs. Overlap in the range of δ¹³C and δ¹⁵N values for the two morphs, however, suggested that they occasionally shared a common diet. Data from other studies strongly indicated that the proximate and ultimate mechanisms that control body-size dimorphisms in Arctic charr differed between sites. Clear differences in trophic ecology in the Loch Tay Arctic charr suggested that the available feeding opportunity may differ for the two morphs. The most likely proximate mechanism resulting in this dimorphism is growth rate differences resulting from differences in food availability for the two subgroups occupying alternative foraging niches in Loch Tay.     

A stable isotope analysis of trophic polymorphism among Arctic charr from Loch Ericht, Scotland

Analysis of nitrogen stable isotope ratios (δ¹⁵N) in white muscle and liver shows temporal stability in the trophic levels at which the two sympatric morphs of Arctic charr Salvelinus alpinus from Loch Ericht feed. The results confirm an ontogenetic dietary switch to a more piscivorous diet, occurred in all individuals of one of the morphs, above a size threshold of 16·5 cm fork length, indicating that this occurs at a high frequency in this population and that once made, the return to feeding to a lower trophic level is infrequent.     

Growth, maturation and reproductive investment in Arctic charr

Size and rates of growth in a cohort of 1 + Arctic charr housed in standard conditions were tracked over 12 months (December to December) and mature and immature males and females compared retrospectively. In both sexes, maturing fish were larger than non-maturing ones. In males, this size differential was the result of differences in growth in winter and early spring, but not in the remainder of the study period. In females, size differentials resulted mainly from growth rate differences immediately prior to breeding. In females but not in males, gonadosomatic index was predicted by growth rates in the months leading up to maturation, and among the females that matured, faster growing fish produced more eggs. Lipid reserves in July were correlated negatively with growth during the previous 7 months and, in females only, lipid reserves were significantly lower in maturing fish than in non-maturing fish, indicating that mobilization of lipid energy reserves in maturing fish had commenced by this time. Variation in investment in gonadal tissue, measured as gonadosomatic index, was not explained by variation in July lipid reserves for either males or females. However, July lipid reserves were negatively correlated with egg number, so females investing more in ova exhibited greater depletion of lipid reserves. These results are discussed in the context of the relationship between body condition and the onset of maturation in salmonids, relative investment in reproduction and sexual differences in the cost of reproduction.     

Upper thermal limits for feeding and growth of 0+ Arctic charr

When Arctic charr Salvelinus alpinus from two diVerent stocks were fed live Neomysis integer , the upper thermal limits for feeding and growth were established in the range 21·5–21·8° C. These critical temperatures might have been underestimates, because fish tend to show increased sensitivity to handling at high experimental temperatures. In the second experiment, the proportion of feeding undisturbed charr from four stocks decreased initially as temperature was raised in steps from 18 to 22° C. At the lower temperatures, 18 and 20) C, almost all fish resumed feeding, but the recovery time was longer and more fish ceased to feed at 20) C than at 18° C. When the temperature was increased to 21° C, 50% of the fish ceased feeding permanently, and all fish ceased feeding within 2 days at 22° C. It is concluded that 0+ charr cease to feed and grow at c .21·5) C and that the critical temperatures for feeding and growth coincide.     

Behavioural responses of benthic and pelagic Arctic charr to substratum heterogeneity

Responses to environmental heterogeneity were studied in laboratory-reared offspring of two morphs of Arctic charr Salvelinus alpinus from Loch Rannoch, Scotland, one occupying the pelagic zone and feeding predominantly on zooplankton and the other being benthic in habit and feeding mainly on macroinvertebrates. When housed in groups in tanks with a black-and white striped base, benthic charr demonstrated a clear preference for dark areas, whereas pelagic fish positioned themselves at random with respect to substratum colour. In general, pelagic charr were much less aggressive than benthic charr. In pelagic fish, neither spacing nor aggression was affected by the visual heterogeneity of the substrate. In contrast, benthic charr swam closer together and fought more when housed over a uniform as opposed to a non-uniform substratum. The results are discussed in the context of habitat-specific visual requirements and of an interaction between visual complexity and territoriality previously described for Atlantic salmon Salmo salar .     

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.     

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.     

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.     

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.     

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.     

Arctic charr strain crosses: effects on growth and sexual maturity

Three Arctic charr strains and their crosses were evaluated with respect to growth and sexual maturation frequency. No significant heterosis was observed for growth. One out of five crosses was heavier, but not significantly so, than the best performing parental strain at ages 1 + and 2. The five other crosses did not perform better than the best performing parental strain at any age. Cross-breeding was not recommended as an effective method to improve growth performance in Arctic charr. Growth performance of reciprocal crosses differed significantly in all strain combinations: crosses resembled more the male parent strain than the female parent strain. ANOVA for male maturation frequency showed a highly significant effect of the male parent strain and a weakly significant effect of female parent strain. In the Ottsjö strain mature males were significantly larger than immature fish and females. Such size difference was also observed in the offspring when sires of the Ottsjö strain were crossed with dams of the faster growing Honavan strain or with dams of the Rensjö strain. When Ottsjö dams were crossed to sires of the other strains no significant size difference between mature males and other fish was found.     

The size structure of lacustrine Arctic charr (Pisces: Salmonidae) populations

The size structure of Arctic charr (Salvelinus alpinus) populations was analysed using published information; 44% of populations were bimodal, with both large (normal) and small (dwarf) morphs occurring within a cohort. The remaining populations were unimodal, consisting of normal-sized or of stunted adults. Bimodal populations increased in frequency with latitude and were characteristic of large, deep lakes with few fish species. The age and size at which bimodality developed, the size difference between the morphs, and the frequency of cannibalism in charr populations increased with latitude. A variety of genotypic and phenotypic explanations are examined. The evidence for specific differences between the morphs is unconvincing. Various competition and predation hypotheses fail to explain the occurrence of bimodality. Cannibalism does not cause bimodality despite being strongly associated with it since bimodality develops before charr become cannibalistic. Much of the variation in charr size structure is suggested to be a consequence of increased seasonality in food supply in more northerly environments, coupled with feeding size thresholds. The latter result in larger members of a cohort being able to maintain growth rates on seasonally abundant prey while smaller individuals which cannot catch these items form a second mode of more slowly growing fish. Bimodality is documented in a number of other, predominantly northern, fish species.     

Behavioural variation in juvenile Arctic charr in relation to body size

Young Arctic charr Salvelinus alpinus , derived from one male and one female only from Lake Ölvesvatn in northern Iceland (a stock that has been popular in Icelandic aquaculture), showed large variance in body size, primarily as a consequence of variable egg size. Shortly after the onset of exogenous feeding, large fish (0·11–0·14 g) were more active and fed mainly from the water surface. Small fish (0·06–0·09 g) moved less and made fewer foraging attempts. They spent most of the time on the bottom, and foraged equally on the bottom and surface. These findings can be used to improve growth and performance of juveniles in rearing tanks.     

Environmental effects on primary increment formation in the otoliths of newlyhatched Arctic charr

Otolith microincrements were investigated in Arctic charr Salvelinus alpinus , reared from hatching under various temperatures (1, 3, 5, 7° C) and feeding conditions (starved, fed every third day, fed daily). Larval charr otoliths were marked with oxytetracycline and alizarin complexone. Alizarin complexone was found to be 100 per cent successful in marking otoliths while oxytetracycline marks could be seen in <10 per cent of the otoliths viewed. Otolith microincrements were viewed by light and scanning electron microscopy to investigate the daily nature of increment deposition. Low temperatures (1 and 3° C) and starvation depressed daily increment formation. Increment deposition was found to be daily among the larvae reared at warmer temperatures (5 and 7° C) and fed at least every third day. Scanning electron microscopic analysis allowed us to confirm the results of light microscope increment counts from all temperatures except 1 ° C, where the number of increments enumerated were higher than the number obtained during light microscopy analyses. Increased feeding and warmer temperatures also resulted in increased increment width. The difference in increment number and width seems to be dependent upon fish growth rate which we have found to be affected by both temperature and feeding conditions.     

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.