Conservation and management of the Arctic charr: a forward view

Abstract –  In this study, we synthesised the views of 34 participants in a workshop to consider the status and future conservation and management of the Arctic charr ( Salvelinus alpinus L.). These are expressed as a series of resolutions deriving from a conference on Arctic charr conservation held in Perth, Scotland in June 2004. Agreed resolutions from the conference were as follows. (i) The unique diversity of Arctic charr should be recognised for the contribution it makes to biodiversity of northern aquatic communities. (ii) The recognition by the public, nongovernmental organisations and national governments of the importance of Arctic charr in northern ecosystems is an important step to the management and protection that should be pursued. (iii) The taxonomic status of highly variable charr populations requires to be reviewed. (iv) There should be a separate system, complementary to the taxonomic one, which systematically catalogues the biological diversity of S. alpinus . (v) The Arctic charr provides a unique natural resource to study the process of evolution and this requires the highest level of protection from anthropogenic effects. (vi) The status of conservation policy for Arctic charr needs to be urgently reviewed to take account of its unique position in the fauna.     

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.     

Ecological and genetic differentiation among the Arctic charr of Lake Aigneau, Northern Québec

Abstract –  Two modal size groups of sexually mature Arctic charr ( Salvelinus alpinus ) differing in shape and found at different depths in Lake Aigneau in the Canadian sub-Arctic are described and tested for genetic and ecological differentiation. Forms consisted of a small littoral resident, mean size 21.7 cm, and a large profundal resident, mean size 53.9 cm. Mitochondrial DNA analysis indicated that seven of eight haplotypes were diagnostic for either the littoral or profundal fish, with 66.6% of the variation being found within form groupings. Pairwise tests of microsatellite data indicated significant differences in nine of 12 loci and a significant difference between the forms across all tested loci. Molecular variation was partitioned to 84.1% within and 15.9% between forms and suggestive of either restricted interbreeding over time or different allopatric origins. Stable isotope signatures were also significantly different, with the profundal fish having higher δ¹³C and δ¹⁵N values than the littoral fish. Overlap and separation, respectively, in the range of form δ¹³C and δ¹⁵N signatures indicated that carbon was obtained from similar sources, but that forms fed at different trophic levels. Littoral fish relied on aquatic insects, predominantly chironomids. Profundal fish were largely piscivorous, including cannibalism. Predominantly empty stomachs and low per cent nitrogen muscle-tissue composition among profundal fish further indicated that the feeding activity was limited to the winter when ice-cover increases the density of available prey at depth. Results provide evidence of significant differences between the modal groups, with origins in both genetics and ecology.     

Summer habitat use and feeding of juvenile Arctic charr,Salvelinus alpinus,in the Canadian High Arctic

Abstract – Owing to limited knowledge of the habitat use and diet of juvenile Arctic charr from the High Arctic, particularly young‐of‐the‐year (YOY), we assembled data obtained from samples taken in and around Lake Hazen, Nunavut, Canada, to assess juvenile habitat use and feeding. Juvenile charr demonstrated a preference for stream environments, particularly those fed by warm upstream ponds. Charr occupying both stream and nearshore lake habitats were found to feed similarly, with chironomids occurring most frequently in diets. Some older stream‐dwelling charr preyed on smaller, younger Arctic charr. Preferred stream occupancy is likely mediated by physical barriers created mainly by water velocity, and by distance from the lake, lake‐ice dynamics, low water depth, and turbidity. Water velocities resulted in stream habitat segregation by size, with YOY mainly found in low‐velocity pools and back eddies adjacent to stream banks, but not in water velocities >0.1 m·s^?1. Greatest charr densities in streams were found in small, shallow, slow‐flowing side channels, which are highly susceptible to drought. Under predicted climate change scenarios, streams fed by small ponds will be susceptible to intermittent flow conditions, which could result in increased competition among juvenile charr for the remaining stream habitats. In addition, glacier‐fed streams are likely to experience increased flow conditions that will exacerbate physical barriers created by water velocity and further reduce the availability of preferred stream habitat.     

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.     

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     

Arctic charr in Britain and Ireland – 15 species or one?

Abstract –  From both a modern and a historical perspective there is little doubt that the Arctic charr, Salvelinus alpinus , in Britain and Ireland (as well as elsewhere) is a 'difficult' species. Historically 15 separate species have been recognised from populations in Britain and Ireland and there have been recent attempts to reassert these specific names. Here we review the evidence for the status of these 'species'. We conclude that the evidence for these 15 being afforded full species status is poor. However, both historical and contemporary data show that Salvelinus alpinus (Linnaeus 1758) in Britain and Ireland exhibits levels of variability in form that is much greater than in other species. We argue that a comprehensive genotypic and phenotypic survey of charr populations in Britain and Ireland is required to determine the full extent of variability and the status of populations with a view of providing suitable protection.     

Thermal habitat use and juvenile growth of Svalbard Arctic charr: evidence from otolith stable oxygen isotope analyses

Abstract – Stable oxygen isotopes (δ¹⁸O) derived from otoliths were used to estimate mean annual water temperatures experienced by individual Svalbard Arctic charr, Salvelinus alpinus (L.), during their first four growth seasons. The analysed Arctic charr experienced a high variety of temperatures, indicating the use of different thermal habitats. A higher proportion of the juveniles experienced warmer temperatures during their first summer compared with later summers, suggesting the selective use of the shallowest littoral areas of the lake. Although the estimated temperatures were consistent with water temperatures found in High Arctic rivers and lakes during summer, they did not represent the annual variation in air temperature registered over the 20 years of otolith measurement. Furthermore, summer otolith increment width did not correlate with the experienced temperature. However, after the second year, otolith increment width was highly dependent on increment width during the previous summer. This study estimated mean summer water temperatures experienced by individual Arctic charr during the first four growth seasons providing additional evidence that stable oxygen isotope analysis can be used to provide insight into the thermal habitat use by juvenile Arctic charr.     

Genetic divergence among broodstocks of Arctic charr Salvelinus alpinus in eastern Canada derived from the same founding populations

We examined the amount and distribution of molecular variation at microsatellite loci in 21 broodstocks of Arctic charr Salvelinus alpinus derived from the Fraser River, Labrador and the Nauyuk Lake, Nunavut, Canada. Our goal was to assess the amount of genetic diversity and differentiation as broodstocks are subdivided and propagated in different hatcheries and grow‐out facilities. We observed significant heterogeneity across pairs of ancestral and descendant broodstocks in the mean numbers of alleles at microsatellite loci. We detected a significant decrease in the observed heterozygosity between ancestors and descendants but the amount of decrease did not depend on either the degree of removal from the wild (number of sequential transfers) or the strain (Fraser vs. Nauyuk). Based on allele frequency distributions, there was little genetic evidence of bottlenecks during the creation of subsequent broodstock populations after the initial founding events. All broodstock samples were significantly differentiated from each other but those within the same strain were more similar to each other than to broodstocks from different strains. Broodstocks from the Nauyuk Lake broodstocks showed greater differentiation from each other than did Fraser River broodstocks, which could be attributed to differences in the number of founders.     

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.     

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.     

Antipredator behaviour of naïve Arctic charr young in the presence of predator odours and conspecific alarm cues

Abstract –  We investigated how Arctic charr young respond behaviourally to olfactory cues from skin-damaged conspecifics, charr-fed pikeperch, and the combination of food-deprived pikeperch and skin-damaged conspecific cues in a two-channel Y-maze fluviarium test arena. Significant antipredator responses were found to skin-damaged conspecifics in three behavioural traits, to charr-fed pikeperch in two traits and to the combination of pikeperch and skin-damaged conspecific cues in all six behaviours investigated. The combination of predator and conspecific cues significantly increased spatial avoidance and cautiousness to approach the source of scent compared with odours from charr-fed predators. The results suggest that damage-released alarm cues exist in charr, and they strengthen the antipredator responses in conspecifics. As the charr used different behaviours and levels of response in the presence of different cues, they seem to have sophisticated skills to distinguish between different odour combinations and be able to adjust their behaviour according to the current predation risk level.     

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.     

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.     

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.     

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.     

Habitat and food choice of Arctic charr in Linnévatn on Spitsbergen, Svalbard: the first year-round investigation in a High Arctic lake

Abstract –  Habitat and diet of Arctic charr Salvelinus alpinus (L.) were studied by monthly sampling from late autumn to early summer in Linnévatn, Svalbard (78°3'N, 13°50'E). This is the first year-round study of a population of charr in the High Arctic, with samples being taken every 5–7 weeks. The ice cover lasted for more than 9 months, from mid-October to late July, with the greatest thickness in mid-May. Although most charr occupied the littoral zone during winter, the highest densities in April and October were found in the deeper areas (20 m) of the lake. The fish fed at all times of the year, but the number of stomachs with food and the stomach-filling indices were lowest during the darkest part of the season. The diet of smaller charr (<15 cm) varied strongly with season, showing a dominance of zooplankton in late autumn and chironomids in winter (larvae) and summer (pupae). The food choice was in accordance with the density of food items available. Larger fish (≥15 cm) were mostly cannibalistic during the entire year.     

Evidence for the recent extinction of two Arctic charr Salvelinus alpinus (L.) populations in the West of Ireland

1. This paper highlights the recent extinctions of two native Arctic charr populations in the West of Ireland, one population from the Lough Conn catchment, and a second from the Lough Corrib catchment, respectively. 2. The distribution of charr populations in the two catchments is reviewed. The current distribution information is drawn from recent fish stock surveys (Lough Corrib in 1996, and Lough Conn in 1997 and 1998). Previous records of charr in the literature and unpublished data are described to place the current distribution of charr within these catchments into a historical perspective. 3. The importance of the charr population in Lough Mask (Lough Corrib catchment) is emphasized, as it is one of the last extant examples of charr from a large Irish alkaline lake. 4. Impacts on charr populations in both catchments are discussed and conservation measures proposed. Copyright © 2001 John Wiley & Sons, Ltd.     

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.     

The digestion and selective absorption of dietary fatty acids in Arctic charr, Salvelinus alpinus

To compare the rates of digestion and absorption of individual fatty acids, Arctic charr, Salvelinus alpinus (L.), were fed isoenergetic diets containing 40 g kg^?1 coconut oil, and various combinations of 10 g kg^?1 of polyunsaturated fatty acids (PUFA) (18:2n-6 or 18:3n-3) and monounsaturated fatty acids (MONO) (20:1n-9 or 22:1n-9) in the form of free fatty acids (FFA) or triacylglycerol (TAG). The average lipid digestibility for all diets measured by use of the chromic oxide method in the pyloric caeca area, midgut and hindgut were 72%, 83% and 88%, respectively, showing that lipid digestion and absorption occur mainly in the pyloric caeca area, but also extend throughout the intestinal tract. Analyses of digesta present in the intestinal segments suggest the predominance of non-specific lipolytic activity producing primarily FFA and glycerol from dietary TAG. Comparisons of the fatty acid composition of the lipid classes in the digesta showed that the utilization of dietary TAG was dependent both on the rate of release of the individual fatty acids from TAG, and their subsequent rate of absorption. When supplied as either FFA or TAG, the levels of PUFA (18:2n-6 or 18:3n-3) in the digesta were very low, indicating almost complete utilization. Both MONO used (20:1n-9 or 22:1n-9) were absorbed less efficiently than PUFA, but the rate of release from TAG seemed to be rate limiting only for 22:1n-9, which accumulated in the digesta. The rates of absorption of 20:1n-9 and 22:1n-9 when fed as FFA were the same. Comparisons of the levels of fatty acids in the dietary coconut oil TAG with those of the digesta lipids showed that 12:0 was a good substrate for intestinal lipase and was quickly absorbed. The lipolysis of 14:0 and 16:0 was intermediate while the longer-chain 18:0 appeared very resistant to digestion and was a major component of TAG, diacylglycerols and monoacylglycerols present in particularly the hindgut digesta. The absorption of 18:0 also appeared to be very low. The results suggest that PUFA are released very rapidly from dietary TAG by intestinal lipases in Arctic charr, and are specifically absorbed compared with long-chain saturated and monounsaturated fatty acids. The rate of lipolysis may be the rate-limiting step in the digestion of very long chain monounsaturated fatty acids such as 22:1n-9, while both the rate of lipolysis and absorption may be rate limiting for long-chain saturated fatty acids such as 18:0.