An experimental test of the genetic component of the ontogenetic habitat shift in Arctic charr (Salvelinus alpinus)

Abstract— Fry of the Arctic charr, Salvelinus alpinus , were experimentally stocked into a small fish-free lake to test the hypothesis that the size-dependent habitat shift from the epibenthic to the pelagic habitat is genetically determined. The charr originated from a nearby lake inhabiting predatory brown trout Salmo trutta. The cohort of stocked charr was investigated for three years. The Arctic charr started to exploit the pelagic habitat in their first summer at a size of 7–9 cm in contrast to about 15 cm in the donor lake. In the next two summers, the pelagic fraction of the cohort increased. The main fraction lived in epibenthic areas, utilizing the same prey as pelagic charr. Water temperature moderated the habitat use of juveniles such that they avoided warm (>16°C) waters and resided in cool, deep areas. The result was consistent with the hypothesis of a tradeoff between feeding benefit and the predation risk producing spatial segregation of Arctic charr and demonstrated that the fish can facultatively respond to predation risk and adjust the size at which they migrate to the pelagic zone to feed on zooplankton.     

Brown trout (Salmo trutta L.) and Arctic charr (Salvelinus alpinus (L.)) as predators on three sympatric whitefish (Coregonus lavaretus (L.)) forms in the subarctic Lake Muddusjärvi

Abstract  – Brown trout ( Salmo trutta L.) and Arctic charr ( Salvelinus alpinus (L.)) use whitefish ( Coregonus lavaretus (L.)) as their main prey in the subarctic Lake Muddusjärvi. Brown trout dwelled in littoral and pelagic habitat, whereas Arctic charr lived only in epibenthic habitat. Both species shifted to whitefish predation at a length of 20–30 cm. At this size, brown trout fed on larger whitefish than Arctic charr. Whitefish occur in three sympatric forms, differing in their habitat, ecology and morphology. Both the predators preyed primarily upon the small-sized, densely rakered whitefish form (DR), which was the most numerous whitefish form in the lake. DR used both epibenthic and pelagic habitat, whereas two sparsely rakered whitefish forms dwelled (LSR and SSR) only in epibenthic habitat: LSR in littoral and SSR in profundal areas. Sparsely rakered whitefish forms had minor importance in predator diet.     

Effects of stocking of piscivorous brown trout, Salmo trutta L., on stunted Arctic charr, Salvelinus alpinus (L.)

To study the effects on a stunted freshwater population of Arctic charr, Salvelinus alpinus (L.), two groups of large (26–45 cm) individually tagged brown trout, Salmo trutta L., were released and recaptured with gillnets after 1, 7, 11 and 63 weeks. One group of trout was trained on a fish diet before release, and the other, reared on commercial dry pellets, served as a control. Specific growth rates in both groups were negative 1 week after release and approached zero after 63 weeks. Condition factor and internal fat content decreased during the experiment. Although only 11% of the trout stomachs examined contained fish prey, charr represented 79% of the total stomach weight content. Gillnet samples of charr before and 63 weeks after the release of trout indicated a decreasing population size of charr. Individual growth and mean length of charr increased after release of trout, especially for charr at age 4 years. After the release of trout, 35% of the charr were longer than 20 cm as compared with 6% before the release.     

Littoral spawning habitats of three southern Arctic charr (Salvelinus alpinus L.) populations

Abstract Arctic charr populations in southern latitudes are nonmigratory, with all life‐stages limited to freshwater lakes and in‐ or out‐flowing tributaries. Although many of these populations are reported to also spawn in lake littorals, little is known about the physical characteristics of putative spawning grounds. A total of 23 discrete spawning sites within three Irish lakes were located by fyke netting of spawning adults and snorkelling in littoral habitats. Spawning sites were found to be long, narrow strips running parallel to the shore at a maximum depth of 124 cm. Spawning sites were limited to areas of coarse mineral substrate with an adequate (c. 8 cm) depth of clean interstitial spaces. In individual lakes, combined areas of spawning sites made up 0.4–0.7% of available littoral. Egg densities varied considerably between sites (33–900·eggs m^?2) and were significantly correlated with gradient and width of spawning sites. No evidence of redd digging was found. The shallow, localised and restricted nature of spawning grounds makes such populations vulnerable to anthropogenically induced postoviposition changes in surface water level, eutrophication processes such as increased lake sedimentation and elevated nutrient status.     

Muscle carotenoid concentrations in sexually maturing and immature Arctic charr, Salvelinus alpinus (L.)

Muscle carotenoid concentrations were measured in immature and maturing 2+ Arctic charr, Salvelinus alpinus (L.), sampled from April until ovulation started in November. In immature fish, muscle carotenoid concentration increased steadily and had reached about 10 mg kg^?1 by the end of the experiment. In mature fish of both sexes, pigment concentrations decreased during the autumn and muscle carotenoid concentrations were significantly lower than in the immature fish during the latter stages of the trial. No significant difference in muscle pigmentation was observed between sexes. Carotenoid concentrations were lowest in the cranial region of the fillet, but decreased in all parts of the fillet when the charr became mature. In females, the decrease seemed to be most marked in the caudal region. At the termination of the experiment, fillets accounted for about one-half of the carotenoid content in mature fish of both sexes. High concentrations of pigment were recorded in the skin and fins of mature males. In mature female charr, pigment was almost equally distributed among skin, fins and ovaries     

The effect of dietary inulin on aerobic bacteria associated with hindgut of Arctic charr (Salvelinus alpinus L.)

The primary aim of the present study was to evaluate the population level of adherent (autochthonous) aerobic and facultative anaerobic bacteria in the hindgut of healthy Arctic charr (Salvelinus alpinus L.) fed dextrin or inulin. This was assessed by the dilution plate technique, and visualized using both transmission and scanning electron microscopy. A population level of 4.8 × 10⁵ adherent bacteria per gram wet mass was found in the hindgut of fish fed a casein‐based diet supplemented with 15% dextrin. However, substituting dextrin with 15% inulin reduced the bacterial population level in the hindgut (3.56 × 10⁴). A total of 217 bacterial isolates were identified by key phenotypical and biochemical characteristics. In addition, 22 strains were also identified by partial sequencing of the 16S rRNA gene. The composition of bacteria colonizing the hindgut of Arctic charr fed dextrin was dominated by the genera Staphylococcus, Pseudomonas, Micrococcus, Psychrobacter glacincola and Streptococcus. However, bacteria colonizing the hindgut of fish fed inulin were dominated by Gram‐positive bacteria of the genera Staphylococcus, Streptococcus, Carnobacterium and Bacillus. While Carnobacterium divergens‐like strains were isolated from charr fed dextrin, Carnobacterium maltaromicus‐like strains were isolated from the hindgut of fish fed inulin. Electron microscopical analysis of hindgut regions confirmed traditional culture‐based microbial analysis as fewer bacterial cells were observed between microvilli and associated with the surfaces of enterocytes of fish fed inulin rather than dextrin.     

The influence of physical watercourse parameters on the degree of anadromy in different lake populations of Arctic charr (Salvelinus alpinus (L.)) in northern Norway

The influence of physical watercourse parameters on the degree of anadromy was studied in 15 lake populations of Arctic charr (Salvelinus alpinus (L.)) in northern Norway. The anadromy of the different populations was scored based on the following criteria; marine parasites. fish length and length at maturity. There were significant negative correlations between the anadromy scores and both the length of the outlet rivers of the different lakes and a migration barrier index for the different watercourses (combining migration distance and water velocity). It is hypothsized that these correlations were due to differences in energy expenditure for charr during upstream migration and/or differences in predation pressure on charr during river migration. Short rivers with relatively high water velocity seem to be the most preferable conditions for the anadromous life-history strategy. There were no correlations between the degree of anadromy and mean river slope or minimum summer water discharge.     

Influence of feeding frequency on growth and size dispersion in Arctic charr Salvelinus alpinus (L.)

Abstract Dominance hierarchy formation in groups of fish is thought to be a major reason for the development of size heterogeneity. The objective of the current experiment was to examine the influence of different feeding frequencies on growth and changes in size distribution in groups of Arctic charr Salvelinus alpinus (L.). The fish were fed equivalent excess rations continuously for 24 h a day, for 2 h twice a day (early morning and evening), for 2 h once a day in the morning or for 2 h once every other day also in the morning. Growth was measured by individual weighing of fish four times during the 130‐day experiment. Coefficient of variation in size within rearing groups was used to assess size dispersion and the possible development of hierarchies within the groups. Feeding frequency did not influence variation in size within groups, and there were no signs of hierarchy formation.     

An experimental comparison of cannibalistic response in different Arctic charr (Salvelinus alpinus (L.)) stocks

Abstract– The cannibalistic response of four domesticated Arctic charr stocks were compared experimentally using a radiographic technique. Charr from Arctic Spitsbergen (80°N) consistently exhibited a much higher frequency of cannibalistic response than fish from three charr strains from mainland Norway (70°N). The results indicate that the extent of cannibalistic feeding in Arctic charr is population specific and support the hypothesis that charr from Arctic populations have stronger cannibalistic tendencies than do fish from more temperate regions.     

Production and predation rates in a cannibalistic Arctic char (Salvelinus alpinus L.) population

Abstract – Production of cannibalistic Arctic char ( Salvelinus alpinus (L.)) and their prey conspecifics were estimated in an allopatric population, situated in an alpine ultraoligotrophic lake in central Norway. Estimates of population abundance were obtained by mark recapture and successive removal experiments. Assuming stable age distribution, we estimated mean yearly biomass (±95% CL) of char in sampled size groups vulnerable to predation (60 to 150 mm) to 7.93 (5.11–14.30) kg · ha⁻¹. Similarly, mean yearly biomass (±95% CL) of cannibalistic char (> 250 mm) was estimated to 0.62 (0.50–1.06) kg · ha⁻¹. Annual production (±95% CL) of char in length groups 60 to 150 mm was estimated to be 4.31 (2.74–8.03) kg · ha⁻¹ · year⁻¹, and production (±95% CL) of cannibalistic char to 0.19 (0.15–0.33) kg · ha⁻¹ · year⁻¹. Depending on the food conversion factor (set to vary from 0.1 to 0.4), the cannibalistic char removed from 10% to 40% of the production of char in sampled size-classes vulnerable to predation yearly. The overall ecological efficiency in energy transfer between the prey and predator population was 4.4%.     

Amino acid composition of Arctic charr, Salvelinus alpinus (L.) and the prediction of dietary requirements for essential amino acids

Embryo somatic tissues, non‐somatic yolk‐sac materials, and whole, individual fingerlings (age 0+) of Arctic charr, Salvelinus alpinus (L.), as well as a commercial trout diet, were analysed for a wide spectrum of amino acids. Analytical material consisted of prefeeding swim‐up fry that were separated into discrete yolk sac and somatic embryo tissue samples. Amino acid concentrations in fry somatic tissue and whole fingerlings were generally very similar to each other, but were lower than those measured in yolk materials. Higher correlations were observed between the majority of specific amino acid concentrations in the trout diet when compared with fingerling data (r² = 0.91) and fry somatic tissue data (r² = 0.89), than when correlated with fry yolk sac material (r² = 0.76). These results indicate that the essential amino acid profiles of fry somatic tissue and whole fingerlings are closer to that of a commercial feed than they are to the endogenous profiles found in the embryonic yolk sac material itself. The dietary ratios of individual essential amino acids were also compared with the total essential amino acid concentrations (A/E ratios) in whole fingerling tissues, and these ratios could be used to accurately estimate the apparent essential amino acid requirements of Arctic charr. The rationale for using carcass amino acid composition data to estimate the dietary essential amino acid requirements of Arctic charr is discussed.     

Divergent, genetically based feeding behaviour of two sympatric Arctic charr, Salvelinus alpinus (L.), morphs

Abstract –  Arctic charr, Salvelinus alpinus (L.), is one of the several northern fishes that show resource polymorphisms in postglacial lakes. Two reproductively isolated morphs of Arctic charr coexist in distinct ecological niches in the subarctic lake Fjellfrøsvatn, North Norway. Offspring of the two morphs (termed littoral charr and profundal charr) were reared separately but under identical conditions. Their feeding behaviour was compared experimentally using different kinds of live prey. The fishes had no experience with natural prey before the experiments. The littoral morph were more effective in eating live plankton ( Daphnia ) and littoral benthos ( Gammarus ), and had a higher attack rate against pleuston (surface prey, Gerris ) compared with the profundal morph. The two morphs behaved in accordance with expectations from their in situ niche utilisation towards the three prey types. This indicates a case of incipient ecological speciation where divergence in resource utilisation in contrasting niches has evolved adaptations in feeding behaviour by natural selection.     

Feed intake and growth of Arctic charr, Salvelinus alpinus (L.), fed diets supplemented with oxytetracycline and squid extract

This study examined the long-term effects of a potential feeding deterrent, oxytetracycline (OTC), and a putative feeding stimulant, squid extract, on feed intake, growth and dry matter (DM) digestibility in Arctic charr, Salvelinus alpinus (L.). Fish were fed one of four diets for 9 weeks: 1. commercial feed formulation (basic); 2. BM (basic plus 20 g kg⁻¹ OTC); 3. BMS (BM plus 10 g kg⁻¹ squid extract); 4. BS (basic plus 10 g kg⁻¹ squid extract). The feed intake of Arctic charr was not reduced by the addition of OTC to the basic feed, and addition of squid extract to the medicated feed did not enhance feed consumption. DM digestibility and growth tended to be reduced in groups of fish fed diets supplemented with the antibiotic (BM and BMS). This suggests that OTC may have interfered with nutrient digestion and absorption. OTC does not seem to be a feeding deterrent for Arctic charr, possibly indicating that the charr is a rather unselective feeder. Because the medicated feeds were readily accepted, there may be little to be gained by addition of feeding stimulants to these types of feeds for charr.     

Morphological variability among three geographically distinct Arctic charr (Salvelinus alpinus L.) populations reared in a common hatchery environment

Abstract –  The morphology of three lake-resident Arctic charr, Salvelinus alpinus , populations was studied at two life-history stages in a common-garden experiment. The fish of the same year class were reared under standard hatchery conditions, and 27 morphometric variables (a truss network) were measured from the sampled individuals. Most of the total variation was explained by the overall body robustness, dimensions of the head and caudal peduncle length. After controlling for a body size, significant heterogeneity in body shape was found among populations at both ages. Independent of age, the populations were morphologically highly distinct, although some integration of characters could be found as the fish reached sexual maturity. Sexual divergence accounted for a large part of the within-population shape variation, the mature males having more robust bodies, larger head dimensions and longer pectoral fins compared with the mature females/immature fish. Although the cultured fish may not be totally representative of their wild counterparts, it is reasonable to expect that the observed morphological differences have a genetic basis, presumably reflecting adaptation to local environmental conditions experienced by the charr in their original habitat.     

Effects of temperature and salinity on the reproductive success of Arctic charr, Salvelinus alpinus (L.): egg composition, milt characteristics and fry survival

The effects of temperature and salinity on the reproductive success of Arctic charr, Salvelinus alpinus (L.), were examined by holding broodstock under the following conditions from mid‐May until the end of September: fresh water at ambient temperature (NFW; 8–16 °C); salt water (25–30‰) at ambient temperature (NSW; 4–10 °C); fresh water cooled to saltwater temperature (CFW; 4–10 °C); or salt water heated to freshwater temperature (HSW; 8–16 °C). The relative fecundity of females was similar among groups (P > 0.05; 2685 ± 706 eggs), but females reared in NSW produced significantly larger eggs than those raised in NFW. The highest spermatozoa concentrations were found in milt from males reared in SW and the highest milt glucose concentration was from males kept under coldwater conditions (CFW, NSW). Eggs from NSW and HSW females contained more proteins than eggs produced by NFW females. Eggs from NSW females also contained 40% more lipids than was observed in the other groups, and total energy content was 27% higher in eggs from NSW females than in eggs from NFW females. When FW was cooled (CFW), females produced eggs with protein contents similar to those in NSW, but the lipid contents remained 30% lower. Finally, the best survival at the eyed stage and at hatch was observed in families produced by NSW broodstock. Intermediate values were observed in families from NFW or CFW while the highest mortality occurred in families from the HSW group. All these results suggest that, under the experimental conditions used in the present study, coastal seawater conditions offered the most favourable summer rearing conditions with respect to the reproductive success of Arctic charr.     

Lipid dynamics in anadromous Arctic charr, Salvelinus alpinus (L.): seasonal variations in lipid storage depots and lipid class composition

Changes in lipid class composition of tissues of anadromous Arctic charr, Salvelinus alpinus, were studied in relation to seasonal patterns of lipid deposition and lipid mobilisation. Charr of a north Norwegian population were collected in late May when fish were migrating towards the sea, and in mid-July when the fish re-entered fresh water. Some maturing fish captured in mid-July were held in captivity until late September to assess lipid dynamics associated with maturation. The carcass (head, fins, skeleton and skin) accounted for approx. 50% of the total body lipid content when the fish re-entered fresh water from the sea. Triacylglycerols (TAG) comprised 75-80% of the carcass lipid in ascending charr, confirming that the carcass is a major lipid depot. The skeletal muscle also represented a major lipid store: muscle of ascending charr contained 35-40% of the body lipids, with TAG being dominant (approx. 75% of muscle lipid). The gut (gastrointestinal tract and visceral lipid) and liver of ascending charr each held only 4-5%; of the body lipid, TAG being the dominant lipid class in both of these depots (Gut TAG: 60-65%; Liver TAG: 50-55%). Body lipid decreased 30-40% during maturation: lipids were depleted from all depots, but mobilisation of from the carcass and muscle was quantitatively the most important. In mature females, the ovaries held almost 25% of the body lipid, with the proportion of TAG exceeding that of polar lipids (PL)(TAG: 62%; PL 12.4%). In the mature males, the testes accounted for less than 3% of the total lipids. TAG tended to dominate the testis lipids (TAG: 29.5%), but there were also quite high proportions of free fatty acids, cholesterol and polar lipids (FFA: 23.5%; C: 15%; PL 9.7%). Females lost approx. 80% of their body lipid during spawning and overwintering, the majority of this being TAG. Males also lost large amounts of TAG during the winter, but in contrast to the females lipid depletion amounted to only 50-55% of that present at the time of ascent into fresh water.     

Food composition, habitat use and growth of stocked and native Arctic charr, Salvelinus alpinus, in Lake Muddusjärvi, Finland

Habitat use, growth and food composition of native and stocked Arctic charr, Salvelinus alpinus (L.), were studied in the subarctic Lake Muddusjärvi, northern Finland, to investigate reasons for poor stocking success. Samples were collected with pelagic and epibenthic gill nets. Stocked and native charr occurred in similar epibethic habitats, whereas pelagic habitat was avoided. Native charr grew fast after shifting to piscivory. Growth rate of stocked charr was slow because only a small proportion of stocked fish became piscivorous during the first year after stocking. During the first lake year, stocked charr divided into slow-growing planktivores and fast-growing piscivores. Piscivorous stocked and native charr consumed only whitefish, Coregonus lavaretus (L.), as their prey. Small-sized (<10 cm) whitefish were preferred when shifting to piscivory.     

Food resource partitioning of Arctic charr, Salvelinus alpinus (L.) and three-spined stickleback, Gasterosteus aculeatus L., in the littoral zone of lake Takvatn in northern Norway

The food resource partitioning of Arctic charr (Salvelinus alpinus L.) and three-spined stickleback (Gasterosteus aculeatus L.) were investigated in the littoral zone of lake Takvatn in northern Norway in the ice-free period June–November. Charr and sticklebacks had different feeding habits. Sticklebacks ate several small benthic prey items that were never eaten by charr, and the sticklebacks' diet were dominated by the benthic microcrustaceans Chydoridae and Ostracoda, chironomid larvae and stickleback eggs. Small charr (<17 cm) consumed a wide spectrum of chironomid pupae, terrestrial insects and zooplankton. Intermediate (17–20 cm) and small charr had quite similar feeding habits, while large charr (>20 cm) frequently ate both benthos, pelagic and terrestrial food. The diet overlap between small charr and sticklebacks was never larger than 0.6 (Schoener's index). The segregation in feeding habits indicates that small charr and sticklebacks are segregated in microhabitat when they are both in the littoral zone.     

Long-term retention of coded wire tags in juvenile Arctic charr Salvelinus alpinus

Abstract  A hatchery experiment was organised to find out if the high loss rate of coded wire tags, noticed in connection with an Arctic charr Salvelinus alpinus (L.) restocking programme, was size-dependent and also to test head moulds for future taggings. Six moulds made by Northwest Marine Technology Inc. and five moulds made at the Finnish Game and Fisheries Research Institute were used. A total of 100 charr were tagged with standard-length coded wire tags using each mould. One-summer old charr at tagging were held for 534 days and two-summer old charr for 320 days. The tag loss rate varied between 0% and 54% and was negatively related to the size of the charr at tagging. The high loss rate was connected with the poor alignment of some moulds, such that a large proportion of the tags were located in the nasal cavity instead of the nasal cartilage. Testing of the moulds and careful grading of the fish for tagging is emphasised, especially if Arctic charr smaller than 20 g in weight are to be tagged.