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.     

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.     

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.     

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.     

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     

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.     

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.     

Determining the consistency of thermal habitat segregation within and among Arctic charr morphotypes in Gander Lake,Newfoundland

Abstract – Otolith carbon and oxygen isotope data obtained from distinct genetic and ecological groups of lacustrine Arctic charr, Salvelinus alpinus L., from Gander Lake, Newfoundland, were used to examine hypotheses regarding the consistency of differential habitat use among the groups. Results indicated thermal habitat separation by group, with small ‘pale’ individuals consistently remaining in cooler profundal habitats and larger ‘dark’ individuals more frequently occupying warmer upper water column habitats. Theoretical measures of resource separation and competition indicated lower thermal habitat overlap among the forms and greater within‐form competition. Depth at capture data indicated more varied short‐term use of available lake habitats by ‘dark’ form Arctic charr, possibly as a result of cannibalistic foraging on profundal ‘pale’ form fish. Nominal capture depth data only partially explained observed variation in the mean temperature of occupied thermal habitat, suggesting that capture depth can only be used as a rough index of thermal habitat use. Provided that sufficient thermal gradients exist in the environments being studied, otolith oxygen isotopes provide a useful means of establishing the significance of niche differentiation among individuals.     

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.     

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.     

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.     

Learning about danger: chemical alarm cues and local risk assessment in prey fishes

An individual's behaviour patterns can be conceptualized as a series of threat‐sensitive trade‐offs between ambient predation pressure and a suite of fitness‐related activities, such as resource defence, foraging and mating. Individuals that can reliably assess local predation risk could increase their fitness potential by exhibiting predator avoidance behaviours only at appropriate times. However, such learned risk assessment requires reliable information regarding current predation risks. A diverse range of prey fishes are known to possess chemical alarm cues, which when detected by conspecifics and some heterospecifics, elicit a variety of overt and covert responses. These chemical cues, either alone or as a part of a predator's dietary odour, provide reliable information regarding local predation risk. In this review, I describe recent works examining the role of chemosensory information in: (i) acquired predator recognition, (ii) predator inspection behaviour and (iii) the use of conspecific and heterospecific cues as social information sources.     

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.     

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.     

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.     

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.     

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.