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.     

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.     

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     

Optimal protein requirements of young Arctic charr (Salvelinus alpinus) fed practical diets

Young Arctic charr, Salvelinus alpinus (L.), mean weight 2.56 ± 0.02 g, were fed nine isoenergetic (?16.6 MJ digestible energy (DE) kg^?1) practical diets formulated to supply digestible crude protein (DCP) at 40g kg^?1 increments from 230 to 550g kg^?1, for 84 days. Mean weight gain (MWG) and specific growth rate (SGR) were determined every 14 days while carcass composition was determined at the start and end of the experiment. Growth responses attained the highest values in the fish fed the diet with 350 g kg^?1 DCP. Carcass moisture gain, protein gain and apparent net lipid accumulation increased as DCP levels increased to a maximum at 350 g kg^?1 after which there were no differences among treatments. Total carcass lipid and lipid gain decreased as dietary DCP increased up to 470 g kg^?1 with no differences thereafter. Apparent net protein accretion decreased with increasing DCP levels up to 350 g kg^?1 after which there were few differences among treaments. Protein requirements were estimated by fitting MWG and SGR data to broken line regression, quadratic and saturation kinetics models. Results from these analyses suggest that dietary DCP should be provided at between 340 and 392 g kg^?1 (equivalent to ?370 and 420g kg^?1 crude protein) for optimal growth of young Arctic charr reared in similar conditions.     

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.     

Evidence for genetic distinction among sympatric ecotypes of Arctic char (Salvelinus alpinus) in south‐western Alaskan lakes

Resource polymorphism may play an important role in the process of speciation. The Arctic char (Salvelinus alpinus) exhibits great phenotypic and genetic diversity across its range, making it an ideal species for studies of resource polymorphism and divergence. Here, we investigated genetic variation at 11 microsatellite loci among 287 Arctic char from five isolated yet proximate postglacial lakes in south‐western Alaska that were previously examined for resource polymorphism. Significant differences in pairwise F_ST were detected among all lakes (range from 0.05 to 0.28, all P < 0.02). In one lake (Lower Tazimina Lake), we found evidence for two genetic groups of char and for significant differences in the distribution of microsatellite variability among at least two of the three previously described body size morphotypes (‘large’‐, ‘medium’‐, and ‘small’‐bodied char; maximum F_ST = 0.09; differences in admixture proportions). We also found a significant association between genetic admixture proportions and gill raker counts among body size morphs (r = ?0.73, P < 0.001). Our data represent the first record of genetically distinct sympatric morphs of Arctic char in Alaska and provide further evidence that differences in morphology associated with feeding (gill rakers) and growth trajectories reflect niche diversification and promote genetic divergence in Holarctic populations of Arctic char.     

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.     

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.     

Otolith size differences during early life of dwarf and cannibal Arctic char (Salvelinus alpinus)

In Arctic and alpine lakes, Arctic char [Salvelinus alpinus (L.)] often form two distinct morphs: invertebrate feeders (‘dwarfs’) and piscivores (‘giant or cannibals’). Here, we test for early life history growth variation in dimorphic Arctic char as a proximate explanation for the observed life history variation between the two forms. Char were sampled in 11 alpine and Arctic Norwegian lakes. Dwarfs (defined as sexually mature char less than 15 cm long; N = 304) had a mean total length of 105 mm, whereas the typical cannibal (body length above 20 cm; N = 153) was 388 mm long. A positive correlation between egg size and otolith hatching ring were ascertained in a separate hatching experiment with brown trout (Salmo trutta) and it is assumed that this relationship also is valid for Arctic char, and otolith size was used as a proxy for length. Otolith hatching ring from Arctic char cannibals was larger (mean ± SD; 187 ± 24 μm) than those from dwarfs (mean ± SD; 164 ± 23 μm). There were only minor size differences between dwarfs and cannibals during the next three years, after which dwarfs usually matured. Two mutually nonexclusive, proximate explanations for the differentiation into separate morphs (dwarfs and cannibals) are therefore maternal effects and/or genetic based differentiation. The high catchability of large piscivorous char and low production in alpine and Arctic lake ecosystems may make these stocks particularly vulnerable to overexploitation.     

Effects of short‐term starvation periods on flesh quality in Arctic charr (Salvelinus alpinus) in different seasons

Possible effects of short‐term starvation on flesh quality in Arctic charr were studied in spring (March) and summer (August). Groups of juvenile Arctic charr (mean weight March 536 g ± 24; August 461 g ± 15 SEM) were starved for 1, 2 and 4 weeks (March) and 1, 2 and 3 (August). After each starvation period, the fish were slaughtered, and flesh samples collected in order to investigate quality and textural properties in the different experimental groups. Starvation had a positive effect on flesh quality giving firmer texture and lower gaping scores. Starved fish had lower cathepsin activity at slaughter, and a similar difference was seen one‐week post mortem. The results showed that the effect of starvation period was seasonally dependent. Starvation had a larger effect in summer, where a three‐week starvation resulted in firmer texture, whereas this was not seen during spring.     

Changes in the density and growth of brown trout (Salmo trutta) after intensive removal of sympatric Arctic charr (Salvelinus alpinus) in the sub‐Arctic lake Møkkelandsvatn,Norway

This study tests the basic hypothesis that the removal of charr, Salvelinus alpinus (L.), would cause an increase in both the growth and density of a sympatric trout population, Salmo trutta L. The charr population was characterised by slow‐growing individuals, with a high proportion of mature fish, that is typical for so‐called overpopulated populations. A total of 31,000 charr was removed from the lake in the period 1990–1992, and the density of younger trout (1+, 2+), but not older trout (3+, 4+), increased. The growth of older trout (3+, 4+) increased, but the evidence for similar growth increases of younger trout (1+, 2+) was limited. From 1989 to 1990, the proportion of trout increased from 30 to only 40% of the total catch, but from 1991 to 1994, it was significantly higher (60–80%) than that of charr. Total trout biomass increased to a maximum in 1992 and then decreased so that the biomass of 1994 was nearly similar to that of 1989, that is before the start of the charr removal. Back‐calculated lengths of trout from otoliths showed that 2+ and 3+ trout caught in the pelagic were growing consistently faster over previous years than those caught in the littoral, while this was not the case for the 4+ fish. Therefore, the hypothesis was partially supported; the growth rate of trout increased (age groups 1+ to 4+), while the density of juvenile trout (1+, 2+), but not the older trout (3+, 4+), increased after the removal of charr.     

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.     

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.     

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.     

Family × Environment interactions in the growth and survival of Arctic charr (Salvelinus alpinus) grown in brackish and fresh water

We investigated whether Fraser strain Arctic charr Salvelinus alpinus L. reared in fresh water (FW) differed in survival and growth from full siblings transferred to brackish water (BW) and whether family performance was consistent throughout 1 year of commercial growth. Fish were grown communally (n = 3600) in either FW or BW from May 2008 to May 2009. Mean family survival in BW was moderately correlated with mean family survival in FW (r = 0.342; P = 0.032). Fish which survived until the conclusion of the experiment were significantly heavier in body weight at the baseline assessment than their full‐sibs which died. Family by treatment interactions were detected for body weight, condition factor and specific growth across all measurement dates (May 2008, October 2008 and May 2009) and growth intervals. Despite the varied response of families to treatment, mean family body weight in FW was correlated phenotypically with BW body weight in October 2008 (r = 0.633, P < 0.0001) and marginally correlated in May 2009 (r = 0.289, P = 0.061). These data suggest that body weight in FW and body weight in BW should be analysed as separate, but correlated traits in Arctic charr breeding programs.     

The influence of predation risk on habitat selection and food intake by Arctic charr, Salvelinus alpinus (L.)

Abstract— Large piscivorous fish are assumed to affect habitat selection and food intake of prey fish. To study the effects of cannibalistic Arctic charr, Salvelinus alpinus (L.), on smaller stunted charr, we sampled the prey fish in littoral and pelagic habitats using gill nets, before and shortly after the release of large charr in a small lake (0.52 km²). In the habitats where the risk of predation was highest, the catch per unit effort de creased from 13.3 to 4.8 fish per 100 m² of gillnet after release of pred ators. The large decrease in numbers of charr < 18 cm corresponded with the predicted vulnerable prey sizes, according to a model based on the size distribution of predators. The occurrence of planktivorous fish and weight-specific food intake decreased in the high risk habitat and remained unaffected in the low risk habitats. Changes in the food intake of prey fish could not be explained in terms of fish length, indicating that prey fish changed diet when the risk of predation was high.     

Assessing the food web impacts of an anadromous Arctic charr introduction to a sub‐Arctic watershed using stable isotopes

Anadromous Arctic charr, Salvelinus alpinus (L.), was introduced to a sub‐Arctic river–lake system near the village of Kujjuuaq, Nunavik, and the stable isotope values and diets of key resident fish species were used to assess changes in feeding patterns. Stable isotope values for most species did not differ significantly between the pre‐ and post‐introduction periods, with observed shifts being within the bounds of expected natural variation. Lake chub, Couesius plumbeus (Agassiz), were the single species to show a difference between study periods, with a small but significant increase in δ¹⁵N. No significant post‐introduction changes were seen in lake trout, Salvelinus namaycush (Walbaum), omnivory or in any of the assessed quantitative food web metrics. Gut contents of major fish species similarly showed significant temporal overlap between the pre‐ and post‐introduction periods, and there was no significant change in species' weight–length relationships. The minor ecological impact was interpreted in relation to the availability of open niches exploitable by ecological generalists such as Arctic charr. The explanation accords with the known habitat and feeding flexibility of Arctic charr and the ecological immaturity of sub‐Arctic lakes known to have driven adaptive variation among Arctic charr. Findings suggest that anadromous Arctic charr may be introduced at moderate densities to other sub‐Arctic watersheds without major negative food web consequences for other resident fish species.     

Effect of rearing temperatures on the growth and maturation of Arctic charr (Salvelinus alpinus) during juvenile and on‐growing periods

The effect of rearing temperature on the growth and maturation of Arctic charr (Salvelinus alpinus) was investigated. Arctic charr juveniles were reared for 6 months (phase I, October–April, size range 20–500 g) at constant temperatures of 9, 12 and 15 °C and according to two temperature‐step groups (T_step) i.e. fish transferred from 15 to 12 °C or from 12 to 9 °C. All the previous treatments were then reared either at 7 °C or at 12 °C for an additional 4 months (phase II, size range 300–1000 g) and then slaughtered in August 2008. The overall growth rate was the highest at a constant temperature of 15 °C for the first 6 months of the trial, with the fish in this group being 44% and 78% heavier than the fish reared at a constant temperature of 12 or 9 °C respectively. Arctic charr showed a negative response in terms of the growth rate when transferred from higher to lower temperatures, especially for groups previously reared at 15 °C. There was a trend for higher gonadosomatic index values at the end of the experiment for groups of fish that were exposed to higher rearing temperatures during the juvenile phase i.e. 4.18% (±0.79) and 7.29% (±0.89), for the temperature groups of 12 and 15 °C, respectively, compared with 2.49% (±0.74) for the 9 °C group. Our results suggest that for the production of fish >1000 g, moderate or low temperatures (here 9 °C) should be applied during the juvenile phase in order to reduce the negative effects arising from maturation. Farmers with access to heat sources should accordingly choose more moderate rearing temperatures during the juvenile stage, especially if the fish is to be moved down in the temperature regime during the on‐growing period.     

Temperature–growth patterns of individually tagged anadromous Arctic charr Salvelinus alpinus in Ungava and Labrador,Canada

Individual measurements of annual, or within‐season growth were determined from tag‐recaptured Arctic charr and examined in relation to summer sea surface temperatures and within‐season capture timing in the Ungava and Labrador regions of Eastern Canada. Differences between two years of growth (2010–2011) were significant for Ungava Bay Arctic charr, with growth being higher in the warmer year. Growth of Labrador Arctic charr did not vary significantly among years (1982–1985). Regional comparisons demonstrated that Ungava Arctic charr had significantly higher annual growth rates and experienced warmer temperatures than Labrador Arctic charr. The higher annual growth of Ungava Bay Arctic charr was attributed to the high sea surface temperatures experienced in 2010–2011 and the localised differences in nearshore productivity as compared to Labrador. Within‐season growth rates of Labrador Arctic charr peaked in June, declined towards August and were negatively correlated with the length of time spent at sea and mean experienced sea surface temperatures. A quadratic model relating growth rate to temperature best explained the pattern of within‐season growth. Collectively, results suggest that increases in water temperature may have profound consequences for Arctic charr growth in the Canadian sub‐Arctic, depending on the responses of local marine productivity to those same temperature increases.     

Apparent digestibility of macro‐nutrients and phosphorus in plant‐derived ingredients for Atlantic salmon,Salmo salar and Arctic charr,Salvelinus alpinus

Alternative protein sources for aquafeeds need to be indentified in order to increase the efficiency of production. Many studies have examined terrestrial plant meals/protein concentrates as alternatives. Recently the focus has turned to aquatic protists and plants as well as by‐products from other industries, such as breweries. Atlantic salmon, Salmo salar, and Arctic charr, Salvelinus alpinus, were fed diets containing canola meal, soybean meal, corn gluten meal, soy protein concentrate, barley protein concentrate, and solar dried algae included at 30% of the test diet. Barley protein concentrate had the highest apparent protein digestibility values for both species (96.3% for Atlantic salmon and 85.1% for Arctic charr), followed by corn gluten meal. Algae had the highest organic matter digestibility value for arctic charr (80.1%) while corn gluten meal had the highest organic matter digestibility value for Atlantic salmon (88.4%). Algae had a high energy apparent digestibility coefficient (82.4 salmon, 82.7 charr) along with corn gluten meal (78.5 salmon, 82.7 charr) for both species. In general, Atlantic salmon had higher apparent digestibility coefficients compared to Arctic charr for most of the tested ingredients. Both corn gluten and barley protein concentrate appear good candidates as alternative protein sources with both species.