Stable isotope-based determinations of the average temperatures experienced by young-of-the-year Svalbard Arctic charr (Salvelinus alpinus (L.))

Mean ambient water temperatures experienced by individual young-of-the-year (YOY) Arctic charr, Salvelinus alpinus (L.), from a Svalbard lake were estimated using measurements of oxygen stable isotopes (δ¹⁸O) derived from fish otoliths. Otolith-derived water temperatures differed significantly from temperatures recorded at the outlet river of the Dieset watercourse but were consistent with temperatures previously recorded in shallow littoral areas of other Svalbard lakes where YOY charr are commonly found. This indicates that fixed-point monitoring does not necessarily represent the temperatures and thermal habitats used by individual fish. Otolith-derived water temperatures were also positively related to fish length-at-capture and otolith size, although much of the variation remained unexplained. Differences among individuals could be related to variability in food availability and food intake as well as variation in the initial fish size at hatching. Implications for subsequent investigations into how YOY charr respond and adapt to future climate change are discussed.     

Growth and mortality of Arctic charr and European whitefish reared at low temperatures

This comparative study explores how low temperatures affect the mortality and growth of first generation hatchery-reared progeny of subarctic populations of Arctic charr (Salvelinus alpinus L.) and European whitefish (Coregonus lavaretus L.). Replicate fish groups where held under simulated natural light regimes (70°N) at three constant temperatures (1, 3 and 6°C). The mortality of Arctic charr was low (≤1.4%) at all temperature treatments, whereas the mortality of whitefish increased with decreasing temperature from 6% at 6°C to 33% at 1°C. The Arctic charr exhibited higher growth rates than whitefish at all three temperature regimes. All groups of Arctic charr increased in weight, whereas whitefish held at 1°C did not gain weight throughout the experimental period of 133 days. Arctic charr exhibited a large intraspecific variability in growth leading to large variations in size-structure, whereas whitefish in contrast showed very homogenous growth and size-structure patterns; a dissimilarity probably related to species-specific differences in antagonistic behaviour. Evidently, Arctic charr are more cold water adapted than whitefish and are able to maintain growth at extremely low temperatures. Arctic charr thus appear to be the most suitable species for aquaculture at low water temperatures.     

Thermal habitat use and growth in young-of-the-year Arctic charr from proximal fluvial and lacustrine populations in Labrador,Canada

Predicted increases in temperature associated with climate change are expected to have consequences for fish, in particular for Arctic charr, Salvelinus alpinus, a cold-adapted fish species. Despite differences in predicted hydroecological responses to climate change in fluvial and lacustrine environments, little is known of whether fluvial and lacustrine Arctic charr populations may respond differently to increasing temperatures. In order to compare growth and thermal habitat use between habitat types, otolith-inferred average water temperatures estimated from whole otoliths and fork lengths at capture were measured for young-of-the-year (YOY) Arctic charr obtained from two proximal fluvial and lacustrine sites in Labrador, Canada. Otolith-inferred average experienced water temperatures were not significantly correlated with air temperatures at both sites, suggestive of behavioural thermoregulation by YOY. The majority of Kogluktokoluk Brook (fluvial) YOY were found using water temperatures consistent with laboratory determined preferred temperatures for juvenile Arctic charr, whereas most Tom's Pond (lacustrine) YOY were found using temperatures ranging between preferred temperatures and optimal temperatures for growth. There was no consistent difference between mean water temperatures used between YOY from the two sites. Otolith-inferred average experienced water temperatures were only correlated to fork lengths in Tom's Pond YOY. The lack of correlation in Kogluktokoluk Brook YOY is argued to reflect resource partitioning occurring as a result of territoriality known to occur among stream salmonids. The limited range of temperatures used by fluvial YOY in this study, particularly the lack of cooler temperatures, also suggests that fluvial YOY may face barriers to thermal refugia, and as a result may be particularly vulnerable to climate change.     

Influence of body weight and temperature on growth rates of Arctic charr, Salvelinus alpinus (L.)

When reared for a period of 6 months at a temperature of 10°C Arctic charr, Salvelinus alpinus , increased in weight from 18 g to approximately 135 g. Specific growth rates decreased as the fish increased in size and the relationship between size and growth rate could be described by the equation:
where G _w is specific growth rate and W is fish weight in grams. Temperature effects upon growth were examined using previously published data. Below the optimum growth temperature, the growth rate of a fish of given size could be predicted using the equation:
where T is the rearing temperature.
Rates of growth of Arctic charr were as high as those reported for other salmonid species reared under similar conditions. Preliminary results suggested that growth rates of charr may be lower in salt water than in fresh water.     

A morphometric method for estimation of total lipid level in live Arctic charr: a case study of its application on wild fish

Linear body measurements were made on wild Arctic charr Salvelinus alpinus parr (100–200 mm L_F) from two populations in northern Norway during spring (May), summer (June and July) and autumn (October). To reduce handling stress, a method was developed were dorsal and ventral body measurements could subsequently be taken easily from a single picture of the anaesthetized fish. Multiple regression analyses investigated the possible correlation between the body measurements and both total and percentage lipid content of the fish. For both populations and during all seasons, the regression equations gave better estimates for total lipid than for percentage lipid. The regression equations for total lipid accounted for between 67·7 and 89·5% of the variance in lipid content for the different seasons within the lakes. The pooled data within each lake accounted for 62·9 and 81·0% of the variance in total lipid, while the pooled data for both lakes accounted for 67·7% of the variance. In general, the condition factor alone did not give a satisfactory estimate of lipid content of the Arctic charr (r² = 0·003–0·521). Shrinkage on validation values was high (0·20–0·52). Employing a ridge regression method resulted in models with lower r² values and lower shrinkage values (indicating more stable models). Published equations used for hatchery‐reared Arctic charr could not be used on wild fish from the same population. The close correlations between actual and predicted lipid levels found in this study show that morphometric measurements can predict total lipid levels in wild Arctic charr with reasonable accuracy. The most accurate results were obtained when equations were derived from each data set. Therefore, the method has the potential within a single study to estimate lipid levels in live fish as long as some fish can be sacrificed to develop a unique regression equation for each population or experiment.     

Isolation and cross‐familial amplification of 41 microsatellites for the brook charr (Salvelinus fontinalis)

The brook charr (Salvelinus fontinalis; Osteichthyes: Salmonidae) is a phenotypically diverse fish species inhabiting much of North America. But relatively few genetic diagnostic resources are available for this fish species. We isolated 41 microsatellites from S. fontinalis polymorphic in one or more species of salmonid fish. Thirty‐seven were polymorphic in brook charr, 15 in the congener Arctic charr (Salvelinus alpinus) and 14 in the lake charr (Salvelinus namaycush). Polymorphism was also relatively high in Oncorhynchus, where 21 loci were polymorphic in rainbow trout (Oncorhynchus mykiss) and 16 in cutthroat trout (Oncorhynchus clarkii) but only seven and four microsatellite loci were polymorphic in the more distantly related lake whitefish (Coregonus clupeaformis) and Atlantic salmon (Salmo salar), respectively. One duplicated locus (Sfo228Lav) was polymorphic at both duplicates in S. fontinalis.     

The Arctic charr (<Emphasis Type="Italic">Salvelinus alpinus</Emphasis>) populations of Windermere,UK: population trends associated with eutrophication,climate change and increased abundance of roach (<Emphasis Type="Italic">Rutilus rutilus</Emphasis>)

The north and south basins of Windermere in the English Lake District, UK, support autumn- and spring-spawning populations of Arctic charr, Salvelinus alpinus, which have been studied since the 1930s. Continuous investigations of the population dynamics of Arctic charr at this lake have involved gill netting since 1939, collection of fishery catch-per-unit-effort data since 1966, and hydroacoustic surveys since 1990. Analysis of these and associated long-term data on the abiotic environment and other components of the fish communities revealed recently contrasting fortunes of the Arctic charr populations of the north and south basins, the latter of which has been significantly impacted by eutrophication while both basins have shown elevated water temperatures and increasing roach, Rutilus rutilus, populations. Despite the introduction of phosphate stripping in 1992 and some subsequent initial improvement, the hypolimnion of the south basin still remains significantly eutrophicated and the fishery catch-per-unit-effort in this basin is now at record low levels. In addition, the spatial distribution of roach has expanded to form significant components of the fish communities of inshore and offshore surface habitats, where this cyprinid may compete with Arctic charr for zooplanktonic prey. It is concluded that the Arctic charr populations of Windermere, particularly those of the south basin, currently face a number of significant environmental pressures and continued management action is required to ensure their survival.     

Winter ecology of Arctic charr (<Emphasis Type="Italic">Salvelinus alpinus</Emphasis>) and brown trout (<Emphasis Type="Italic">Salmo trutta</Emphasis>) in a subarctic lake,Norway

We studied habitat choice, diet, food consumption and somatic growth of Arctic charr (Salvelinus alpinus) and brown trout (Salmo trutta) during the ice-covered winter period of a subarctic lake in northern Norway. Both Arctic charr and brown trout predominantly used the littoral zone during winter time. Despite very cold winter conditions (water temperature <1°C) and poor light conditions, both fish species fed continuously during the ice-covered period, although at a much lower rate than during the summer season. No somatic growth could be detected during the ice-covered winter period and the condition factor of both species significantly declined, suggesting that the winter feeding rates were similar to or below the maintenance requirements. Also, the species richness and diversity of ingested prey largely decreased from summer to winter for both fish species. The winter diet of Arctic charr <20 cm was dominated by benthic insect larvae, chironomids in particular, and Gammarus lacustris, but zooplankton was also important in December. G. lacustris was the dominant prey of charr >20 cm. The winter diet of brown trout <20 cm was dominated by insect larvae, whereas large-sized trout mainly was piscivorous, feeding on juvenile Arctic charr. Piscivorous feeding behaviour of trout was in contrast rarely seen during the summer months when their encounter with potential fish prey was rare as the small-sized charr mainly inhabited the profundal. The study demonstrated large differences in the ecology and interactions of Arctic charr and brown trout between the winter and summer seasons.     

Evaluating catches of salmonids taken by gillnets

Data from 10 185 gillnetted brown trout ( Salmo trutta L.) and 5777 Arctic charr [ Salvelinus alpinus (L.)] were analysed. Relative to mesh size, the most efficiently caught fish length (the modal length) was somewhat larger with floated nets than with smaller nets placed on the bottom. There were no such consistent differences between the two species. Fish that exceeded the modal lengths were caught more efficiently than smaller fish. The relationship of most efficient mesh size to fish length and condition factor fitted excellently to a linear equation. Based on this equation, the selectivity curve for Arctic charr gillnetted in the Nesjø reservoir, the different efficiency of nets of 0.1 and 0.17mm twine and an equation expressing the increasing height of the selectivity curve with increasing mesh size, a model is presented to compare length frequencies and total numbers of gillnet catches taken with any combination of mesh sizes.     

Growth and Dietary Niche in Salvelinus Alpinus and Salvelinus Fontinalis as Revealed by Stable Isotope Analysis

Arctic charr, Salvelinus alpinus, and brook charr, Salvelinus fontinalis, inhabiting three lakes in the de la Trinité River and adjacent watershed, north shore, Gulf of St. Lawrence, were sampled in 1998. Arctic charr growth differed among lakes with the smallest fish coming from the largest lake. Arctic charr weight–length equation exponents were almost identical at all sample sites. Brook charr growth was also similar in all lakes. July stomach samples from Arctic charr consisted almost entirely of cladocerans in the largest lake, less so in the intermediate sized lake and were mostly aquatic insects in the smallest lake. Brook charr stomach contents were more varied and included fish. Carbon, nitrogen and sulphur stable isotope analyses (SIA) were used to provide a spatially and temporally integrated image of charr diets. SIA corroborated observed among-lake differences and similarities in species diets and suggested lake morphometry may influence measured results. The ¹⁵N signature in brook charr muscle increased with fork-length, as a result of a shift towards piscivory with size. The ¹⁵N signature in Arctic charr muscle tissue showed a significant negative correlation with fork-length in two of the studied lakes that appears related to dietary niche shifts. Results demonstrate the ability of SIA to detect dietary shifts otherwise unobservable from standard gut content analysis.     

Size-scaling of zooplankton foraging in Arctic charr

Prey capture rate (number of prey s⁻¹) and the mode of feeding of Arctic charr Salvelinus alpinus were studied by performing foraging experiments with two sizes (1·1 and 1·8 mm) of Daphnia longispina prey. Arctic charr were particulate feeders at all densities tested. Adjusted for the effect of prey density, the capture rate showed a hump-shaped relationship with Arctic charr size for both sizes of D. longispina . Estimated attack rates ( a ) also tended to show a hump-shaped relationship with fish size. The estimated size-scaling exponent of the attack rate function, however, was relatively small, implying small changes in attack rate over fish sizes. Simultaneous estimations of a and handling time were used in combination with published data on fish metabolism and dry mass rations of prey to estimate maintenance resource density of prey as a function of Arctic charr mass. Maintenance resource densities increased monotonically with Arctic charr size, and rapidly as optimum fish size relative to attack rate on prey was passed.     

Ecosystem size predicts eco‐morphological variability in a postglacial diversification

Identifying the processes by which new phenotypes and species emerge has been a long‐standing effort in evolutionary biology. Young adaptive radiations provide a model to study patterns of morphological and ecological diversification in environmental context. Here, we use the recent radiation (ca. 12k years old) of the freshwater fish Arctic charr (Salvelinus alpinus) to identify abiotic and biotic environmental factors associated with adaptive morphological variation. Arctic charr are exceptionally diverse, and in postglacial lakes there is strong evidence of repeated parallel evolution of similar morphologies associated with foraging. We measured head depth (a trait reflecting general eco‐morphology and foraging ecology) of 1,091 individuals across 30 lake populations to test whether fish morphological variation was associated with lake bathymetry and/or ecological parameters. Across populations, we found a significant relationship between the variation in head depth of the charr and abiotic environmental characteristics: positively with ecosystem size (i.e., lake volume, surface area, depth) and negatively with the amount of littoral zone. In addition, extremely robust‐headed phenotypes tended to be associated with larger and deeper lakes. We identified no influence of co‐existing biotic community on Arctic charr trophic morphology. This study evidences the role of the extrinsic environment as a facilitator of rapid eco‐morphological diversification.     

Effects of Partially Anadromous Arctic Charr (Salvelinus alpinus) Populations on Ecology of Coastal Arctic Lakes

Little research has been conducted on effects of iteroparous anadromous fishes on Arctic lakes. We investigated trophic ecology, fish growth, and food web structure in six lakes located in Nunavut, Canada; three lakes contained anadromous Arctic charr (Salvelinus alpinus) whereas three lakes did not contain Arctic charr. All lakes contained forage fishes and lake trout (Salvelinus namaycush; top predator). Isotope ratios (δ¹³C, δ¹⁵N) of fishes and invertebrates did not differ between lakes with and without anadromous Arctic charr; if anadromous Arctic charr deliver marine-derived nutrients and/or organic matter to freshwater lakes, these inputs could not be detected with δ¹³C and/or δ¹⁵N. Lake trout carbon (C):nitrogen (N) and condition were significantly higher in lakes with Arctic charr (C:N = 3.42, K = 1.1) than in lakes without Arctic charr (C:N = 3.17, K = 0.99), however, and ninespine stickleback (Pungitius pungitius) condition was significantly lower in lakes with Arctic charr (K = 0.58) than in lakes without Arctic charr (K = 0.64). Isotope data indicated that pre-smolt and resident Arctic charr may be prey for lake trout and compete with ninespine stickleback. Linear distance metrics applied to isotope data showed that food webs were more compact and isotopically redundant in lakes where Arctic charr were present. Despite this, lake trout populations in lakes with Arctic charr occupied a larger isotope space and showed greater inter-individual isotope differences. Anadromous Arctic charr appear to affect ecology and feeding of sympatric freshwater species, but effects are more subtle than those seen for semelparous anadromous species.     

Arctic charr in sympatry with burbot: ecological and evolutionary consequences

The trophic niche and parasite infection of Arctic charr (Salvelinus alpinus) were explored in two lakes with sympatric burbot (Lota lota) and two lakes without burbot in subarctic Norway. The CPUE of burbot and charr were similar in one lake, but burbot had a low population density in the other. Burbot were benthivorous in both lakes. Other co-occurring species like brown trout (Salmo trutta), Atlantic salmon parr (Salmo salar), grayling (Thymallus thymallus) and minnow (Phoxinus phoxinus) were also benthivores. At high densities, benthivorous burbot forced the whole Arctic charr population to utilise mainly the limnetic trophic niche. In contrast, at low burbot density or without burbot present, Arctic charr were primarily benthivorous in the littoral zone. Thus, a clear interactive segregation in diet was observed between Arctic charr and burbot at high burbot densities. There was also a high predation pressure from burbot on young Arctic charr along the benthic zones. The extensive use of zooplankton as prey caused a high parasite infection pressure of copepod transmitted Diphyllobothrium spp. larvae, with the potential for high negative impact on the Arctic charr population. As the benthivore trophic niche was occupied by burbot, the ecological opportunities for polymorphism with benthivorous ecotypes or morphs of Arctic charr were probably prevented. Therefore, the sympatry with burbot seems to have large ecological and evolutionary consequences for this Arctic charr population compared with neighbouring lakes where burbot is absent.     

Salinity tolerance of freshwater acclimated, small-sized Arctic charr, Salvelinus alpinus from northern Labrador

Arctic charr, Salvelinus alpinus , less than 150 mm in size were frequently captured at sea in northern Labrador in areas where salinities of 30‰ or higher had been recorded. These captures were inconsistent with many earlier reports for other areas that indicated Arctic charr less than 150 mm in size were not found at sea. A series of salinity challenge tests was carried out in the field, using wild Ikarut River charr, and in the laboratory, using cultured Fraser River charr, to understand more about the potential fate of these small fish. The results of challenge tests with small Arctic charr (< 120 mm) indicated that at intermediate salinities (10–20‰), these fish can readily survive. In laboratory tests with salinities at 30‰, survival was size dependent and would suggest that in natural situations, small charr would require periodic access to fresh or brackish water to stay alive. The influence of water temperature on salinity tolerance may be important when fish are exposed to temperatures that are below 0° C.     

Thermal adaptation of Arctic charr: experimental studies of growth in eleven charr populations from Sweden, Norway and Britain

1. Experimental growth data for Arctic charr (Salvelinus alpinus L.), all fed on excess rations, from 11 European watercourses between 54 and 70°N were analysed and fitted to a new general growth model for fish. The model was validated by comparing its predictions with the growth rate of charr in the wild. 2. Growth performance varied among populations, mainly because of variation in the maximum growth potential, whereas the thermal response curves were similar. The estimated lower and upper temperatures for growth varied between ?1.7 to 5.3 and 20.8–23.2 °C, respectively, while maximum growth occurred between 14.4 and 17.2 °C. 3. There was no geographical or climatic trend in growth performance among populations and therefore no indication of thermal adaptation. The growth potential of charr from different populations correlated positively with fish body length at maturity and maximum weight in the wild. Charr from populations including large piscivorous fish had higher growth rates under standardised conditions than those from populations feeding on zoobenthos or zooplankton. Therefore, the adaptive variation in growth potential was related to life‐history characteristics and diet, rather than to thermal conditions.     

Adaptations to Stochastic Environmental Variations: The Effects of Seasonal Temperatures on the Migratory Window of Svalbard Arctic Charr

The circumpolar Arctic charr, Salvelinus alpinus, is ideal for studying how environmental factors affect life history in fishes. Charr populations demonstrate a tremendous ecological plasticity and adaptations to harsh environments. Arctic charr is the only freshwater fish on Svalbard, including anadromous, resident, and landlocked stocks. Freshwater lake systems on Svalbard are characterized by very low water temperatures, long-term or even permanent ice cover, and low levels of nutrients. Food is thus limited and may lead to growth stagnation and early maturity in Arctic charr. The individual growth pattern may alternatively follow a sigmoid-shaped curve, caused by a shift to either cannibalistic or anadromous (migration to sea) behaviour. In lake systems that include migratory charr, the population may consist of a mixture of parr, postsmolt, and adult migratory individuals, as well as small-sized resident, large-sized resident (cannibals), and large formerly resident individuals transformed to anadromy. Our study in the Lake Dieset watercourse (79°N), Svalbard, demonstrates that the annual water flow in the outlet river is strongly correlated to air temperatures and provides a passage to the sea, allowing the charr access to the nutrient-rich seawater environment, during at most two months each year. During one of the years studied, the youngest and small-sized part of the sea-going stock was prevented from ascending the river and probably suffered mortality during winter. The migratory window of the Arctic charr in Lake Dieset is therefore highly variable among years and thus unpredictable. We hypothesize that in worst case scenarios (cold years, low water discharge), climatic variations may occasionally prevent charr from migrating upstream in Svalbard lake systems in late autumn, resulting in high mortality in the population.     

SSU rRNA gene sequence reveals two genotypes of Spironucleus barkhanus (Diplomonadida) from farmed and wild Arctic charr Salvelinus alpinus

Spironucleus barkhanus isolated from the blood of Arctic charr Salvelinus alpinus from a marine fish farm were genetically compared with S. barkhanus isolated from the gall bladder of wild Arctic charr. The wild Arctic charr were caught in the lake used as the water source for the hatchery from which the farmed fish originated. Sequencing of the small subunit ribosomal RNA gene (SSU rDNA) from these 2 populations showed that the isolates obtained from farmed and wild Arctic charr were only 92.7 % similar. Based on the sequence differences between these isolates, it is concluded that the parasites isolated from the farmed fish have not been transmitted from wild Arctic charr in the hatchery's fresh water source. It is therefore most likely that the farmed fish were infected by S. barkhanus after they were transferred to seawater. S. barkhanus isolated from diseased farmed Arctic charr were 99.7% similar to the isolates obtained from diseased farmed Chinook (Canada) and Atlantic salmon (Norway). The high degree of sequence similarity between S. barkhanus from farmed Arctic charr, Chinook and Atlantic salmon indicates that systemic spironucleosis may be caused by specific strains/variants of this parasite. The genetic differences between the isolates of farmed and wild fish are of such magnitude that their conspecificity should be questioned.     

Vaccination influences growth of Arctic charr

There is limited knowledge about the effects of oil-based vaccines on the growth of Arctic charr Salvelinus alpinus, in particular at different rearing temperatures. One-year-old Arctic charr were immunized intraperitoneally at 2.9 degrees C with a metabolizable oil-adjuvanted, bivalent vaccine containing killed typical and atypical Aeromonas salmonicida bacteria. After vaccination the non-vaccinated (controls) and vaccinated individually marked fish were held for 20 d at 10.0 degrees C and then for 7 wk at 10.3, 14.1 or 18.1 degrees C. During the first 20 d at 10.0 degrees C the growth rate (G) was higher for non-vaccinated than vaccinated fish. Thereafter vaccinated charr had higher G than control fish at 10.3 and 14.1 degrees C. In contrast, at 18.1 degrees C there was no difference in G and therefore no compensation of earlier growth suppression in vaccinated fish was observed at that temperature. The study indicates that vaccination has no ultimate negative effects on the growth of Arctic charr at temperatures ranging from 10.3 to 14.1 degrees C.     

Lake size and fish diversity determine resource use and trophic position of a top predator in high‐latitude lakes

Prey preference of top predators and energy flow across habitat boundaries are of fundamental importance for structure and function of aquatic and terrestrial ecosystems, as they may have strong effects on production, species diversity, and food‐web stability. In lakes, littoral and pelagic food‐web compartments are typically coupled and controlled by generalist fish top predators. However, the extent and determinants of such coupling remains a topical area of ecological research and is largely unknown in oligotrophic high‐latitude lakes. We analyzed food‐web structure and resource use by a generalist top predator, the Arctic charr Salvelinus alpinus (L.), in 17 oligotrophic subarctic lakes covering a marked gradient in size (0.5–1084 km²) and fish species richness (2–13 species). We expected top predators to shift from littoral to pelagic energy sources with increasing lake size, as the availability of pelagic prey resources and the competition for littoral prey are both likely to be higher in large lakes with multispecies fish communities. We also expected top predators to occupy a higher trophic position in lakes with greater fish species richness due to potential substitution of intermediate consumers (prey fish) and increased piscivory by top predators. Based on stable carbon and nitrogen isotope analyses, the mean reliance of Arctic charr on littoral energy sources showed a significant negative relationship with lake surface area, whereas the mean trophic position of Arctic charr, reflecting the lake food‐chain length, increased with fish species richness. These results were supported by stomach contents data demonstrating a shift of Arctic charr from an invertebrate‐dominated diet to piscivory on pelagic fish. Our study highlights that, because they determine the main energy source (littoral vs. pelagic) and the trophic position of generalist top predators, ecosystem size and fish diversity are particularly important factors influencing function and structure of food webs in high‐latitude lakes.