Polymorphic segregation in Arctic charr Salvelinus alpinus (L.) from Vatnshlidarvatn, a shallow Icelandic lake

We studied the salmonid fish Arctic charr, Salvelinus alpinus , in a small and shallow landlocked lake in NW Iceland. The lake is productive but die only fish present is Arctic charr. Despite the apparent absence of discrete benthic and limnetic habitats for fish, two forms of Arctic charr are found in the lake. They show subde differences in morphology related to swimming performance and manoeuvrability, but differences in life history such as growth, and age and size at sexual maturation are more pronounced. Both forms have benthic feeding habits with one form consuming greater number of species than the other. We suggest that the segregation of these forms is based on the evolution of a specialist from a local generalist and that this has been made possible by the absence of a common fish competitor in similar lakes, the threespined stickleback Gasterosteous aculeatus.     

Extant and extinct forms of arctic charr Salvelinus alpinus (L.) complex from the Leprindo lake system (Transbaikalia): Differentiation in life history, morphology, and genetics

Interconnected lakes Bol’shoe Leprindo and Maloe Leprindo in Transbaikalia hosted large (extinct) and dwarf charr forms. Rarely “small” individuals intermediate in size between these forms are caught. In order to assess morphological, ecological, and genetic differentiation of sympatric charr forms and parapatric charr populations we studied their meristic and morphometric characters, feeding, breeding, and growth; we also investigated variation at 8 microsatellite loci using DNA isolated both from contemporary and historic samples. Profound differences were found between large and dwarf charr in growth rate, feeding (piscivores and highly specialized zooplanktivores, respectively), spawning time, and morphology. Dwarf charr from the two lakes demonstrate minor differences in morphology and growth rate. “Small” individuals are morphologically similar with dwarf charr and spawn together with them, they are recruited from dwarf form in late ontogeny as the result of transition to piscivorous feeding and growth acceleration. Microsatellite analysis showed that: (1) large and dwarf charr forms display high degree of genetic differentiation and reproductive isolation; (2) dwarf charr from interconnected lakes belong to different isolted populations; (3) “small charr” are genetically identical with dwarfs. The degree of ecomorphological and genetic differentiation between large and dwarf forms places charr from Leprindo Lakes among the most strongly differentiated Arctic charr forms’ flocks known at the vast range of S. alpinus complex.     

Ecological polymorphism in Arctic charr

Arctic charr, Salvelinus alpinus (L.), commonly exhibits two coexisting morphotypes, dwarf and normal charr, which are characterized by differences in adult body size and colouration. We tested whether or not the morphotypic differences were genetically determined in rearing experiments with offspring of the two morphs and of their crosses. The experiments suggest that this ecological polymorphism in Arctic charr is largely environmentally determined. When reared under similar conditions, offspring of each of the two morphs differed little in size at the same age, and they had the same early developmental rate and maturation pattern. Moreover, the presence of parr marks along the flanks of the fish, one characteristic of dwarf charr, depended on body size and not on parental morph. Genetic differences between the morphs were suggested for growth rate during the second year of life, and for jaw morphology. Comparisons between charr life histories in captivity and in the wild suggest that ecological polymorphism in Arctic charr is chiefly a result of variation in growth conditions between different habitats.     

Selection on Arctic charr generated by competition from brown trout

We experimentally explored population‐ and individual‐level effects on Arctic charr (Salvelinus alpinus) resulting from resource competition with its common European competitor, the brown trout (Salmo trutta). At the population level, we compared performance of the two species in their natural sympatric state with that of Arctic charr in allopatry. At the individual level, we established selection gradients for morphological traits of Arctic charr in allopatric and in sympatric conditions. We found evidence for interspecific competition likely by interference at the population level when comparing differences in average performance between treatments. The growth and feeding rates did not differ significantly between allopatric and sympatric Arctic charr despite lower charr densities (substitutive design) in sympatric enclosures indicating that inter‐ and intraspecific competition are of similar strength. The two species showed distinct niche segregation in sympatry, and brown trout grew faster than Arctic charr. Arctic charr did not expand their niche in allopatry, indicating that the two species compete to a limited degree for the same resources and that interference may suppress the growth of charr in sympatric enclosures. At the individual level, however, we found directional selection in sympatric enclosures against individual Arctic charr with large head and long fins and against individuals feeding on zoobenthos rather than zooplankton indicating competition for common resources (possibly exploitative) between trout and these charr individuals. In allopatric enclosures these relations were not significant. Diets were correlated to the morphology supporting selection against the benthic‐feeding type, i.e. individuals with morphology and feeding behaviour most similar to their competitor, the benthic feeding brown trout. Thus, this study lends support to the hypothesis that Arctic charr have evolved in competition with brown trout, and through ecological character displacement adapted to their present niche.     

Evidence for genetic differences in the offspring of two sympatric morphs of Arctic charr

The sub-arctic Lake Fjellfrøsvatn, northern Norway, has two morphs of Arctic charr that are reproductively isolated because they spawn 5 months apart. The smaller morph (≤14 cm L_F ) is confined to the profundal zone of the lake and the larger morph is mainly littoral. Three hypotheses were tested: (i) the offspring of the profundal Arctic charr grow slower than the offspring of the littoral Arctic charr under identical conditions, thus indicating a genetic basis for the slow growth of the profundal Arctic charr in the wild; (ii) the wild phenotypes of the two morphs are morphometrically different and the differences are persistent in the offspring; (iii) the offspring of the two morphs have different behaviour traits under similar treatments. The first hypothesis was rejected; offspring of the profundal morph grew slightly better than offspring of the littoral morph at 10° C in the laboratory. The second and third hypotheses were supported by the data. Wild-caught fish of the two morphs differed in several morphometric characters and most of the differences persisted in the offspring. In the laboratory, offspring of the littoral morph were more active, more aggressive and more pelagic than offspring of the profundal morph and naive offspring of the profundal morph were more effective in eating live chironomid larvae than were offspring of the littoral morph. The data for morphometry and behaviour, but not growth, provide evidence for genetic differences between the two Arctic charr morphs of Fjellfrøsvatn.     

Genetic differentiation of Arctic charr <Emphasis Type="Italic">Salvelinus alpinus</Emphasis> complex from Transbaikalia revealed by microsatellite markers

Variation at eight microsatellite loci was studied in Arctic charr Salvelinus alpinus complex from five Transbaikalian mountain lakes. Samples from three lakes included two sympatric charr forms (dwarf and small) differing in trophic specialization, morphology and life cycle parameters. Sympatric forms were genetically closer to each other than to charr from other lakes which evidences their independent origin in each of these lakes as the result of sympatric speciation. In each lake, gene pools of sympatric forms were segregated to a different degree (estimates of F _ST varying from 0.030–0.184 and those of ρ _ST varying from 0.119–0.359). Hierarchical analysis of allelic frequencies variance (AMOVA) in Arctic charr from Lake Baikal, the Vitim, and the Olekma basins showed that variation among and within these basins accounted for 19.5% of the interpopulational variance each. In the AMOVA design, investigating differences among sympatric forms in three lakes these differences accounted for 7.1% of the total variance.     

Three sympatric forms of Arctic charr Salvelinus alpinus complex (Salmoniformes,Salmonidae) from Lake Kamkanda,Northern Transbaikalia

We studied morphology, size and age structure, growth, feeding, and variation at microsatellite loci of three forms of Arctic charr Salvelinus alpinus complex (dwarf, small, large) from mountain Lake Kamkanda in River Olekma basin, northern Transbaikalia. The forms differ in meristic and morphometric characters, external appearance and size. The small form distinctly differs from the dwarf and large forms in higher number and length of gill rakers. The forms differ in growth rate; however, differences in growth between the dwarf and the small forms are not as large as between sympatric dwarf and small charr from other Transbaikalian lakes. The large form is heterogeneous in growth rate. The small form matures one year earlier than the dwarf form and has a shorter life span. The dwarf form is a benthophage, the small form is a planktophage, and the large form is a predator. The dwarf form spawns in September, while the small form spawns in November-December, and there is no overlap in their spawning time. The three forms have clear genetic differences, which support their reproductive isolation. It is assumed that the three forms of Arctic charr originated within Lake Kamkanda on the basis of trophic polymorphism and spawning time displacement and attained a high degree of morphological and genetic divergence.     

Trophic polymorphism amongst Arctic charr from Loch Rannoch, Scotland

This paper describes the results of a multivariate and univariate morphometric analysis of three groups of Arctic charr Salvelinus alpinus from Loch Rannoch, Perthshire, Scotland, distinguished a priori on the basis of coloration (bright v. cryptic) and site of capture (from two locations 12 km apart). The analysis produced a clear and highly significant distinction, not only between brightly and cryptically coloured charr, but also between the two cryptically coloured groups, one of which was more robust in terms of several size-independent head measurements. Stomach analysis showed that the brightly coloured charr fed entirely on zooplankton, the less robust, cryptically coloured fish, fed on benthic macroinvertebrates, whereas a significant proportion of the diet of the more robust cryptically-coloured form consisted of other fish. Differences in length at age also distinguished the three forms, with the robust, piscivorous charr, which live longer and potentially reach a larger final size, attaining smaller sizes at a given age. These data clearly confirm the previous identification of a distinct planktivorous morph of Arctic charr in Loch Rannoch and extend the morph analysis by distinguishing two additional morphs, a benthivorous morph and a piscivorous morph that are morphologically and ecologically distinct. The results are discussed in the context of other systems in which sympatric morphs of Arctic charr have been described.     

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.     

Temperature preference of juvenile Arctic charr originating from different thermal environments

Temperature preference of juvenile (age 1+) Arctic charr (Salvelinus alpinus L.) originating from four arctic and sub-arctic populations (Svalbard and mainland northern Norway), representing a range of habitats with different temperature conditions, was studied by use of a shuttle-box system which allowed individual fish to control their environmental temperature. Based on the assumption that adaptations to long-lasting differences in thermal environments would affect temperature preference, we expected that Arctic charr from the high arctic Svalbard would prefer a lower temperature than the charr from two well-studied sub-arctic mainland lakes (i.e. one anadromous charr population from Storvatn, Hammerfest and two sympatric resident charr morphs from Fjellfrøsvatn, Målselv). There were, however, no significant differences in temperature preference among the four populations after 24 h exposure to the shuttle-box system, although the charr from the omnivore upper-water sympatric morph of Fjellfrøsvatn used significantly longer time to reach a stable thermal preferendum than the fish of the other populations. The average temperature preference at the end of the trials ranged between 10.9 and 11.6 °C among the populations. The lack of population differences suggests that temperature preference is not a polymorphic trait under strong selection in Arctic charr.     

Diet of Arctic charr (Salvelinus alpinus (L.)) and brown trout (Salmo trutta L.) in sympatry in two high altitude alpine lakes

Lake Pisses and Lake Labarre are two oligotrophic high altitude alpine lakes that have sympatric populations of Arctic charr and brown trout. These two lakes have similar morphometric, physical and chemical characteristics. The zooplanktonic and benthic fauna show little diversity. But the density of benthos (Chironomidae) and zooplankton is higher in Lake Pisses. The fish fauna of Lake Pisses is slightly more abundant than that of Lake Labarre, althought in both lakes fish density is low. A study of the diet of the two species revealed differences. In Lake Pisses, where the food supply is better, Arctic charr takes exclusively pelagic and benthic prey, whereas in Lake Labarre it also takes exogenous prey and thus comes into competition with trout. Length and body weight growth rates for Arctic charr are higher in Lake Pisses than in Lake Labarre. For trout, maximum length recorded was in Lake Pisses. The results show that the abundance of Chironomidae favours coexistence of the two species in Lake Pisses and confirm that, in the face of shortage of food, Arctic charr is better adapted than trout. This revised version was published online in July 2006 with corrections to the Cover Date.     

Habitat use and diet of sympatric Arctic charr (<Emphasis Type="Italic">Salvelinus alpinus</Emphasis>) and whitefish (<Emphasis Type="Italic">Coregonus lavaretus</Emphasis>) in five lakes in southern Norway: not only interspecific population dominance?

Stable coexistence of Arctic charr and whitefish does occur in a number of native lake fish communities in Scandinavia. Even so, whitefish introductions into Arctic charr lakes have resulted in serious decline and possibly local extinction of Arctic charr. In this article, we analyze the habitat use and diet of the two species in five Norwegian lakes differing in basin shape and environmental conditions. In two of the lakes, both species are native, and appear to live in a relatively stable coexistence. Here, whitefish mainly occupy the littoral and upper pelagic zone, while Arctic charr live in the deeper habitats. Diets are generally quite different in terms of the zooplankton species eaten. In the three other lakes, either whitefish or both species have been introduced. In the shallowest lake, habitat segregation is similar to that seen in the pristine lakes, although Arctic charr appears to be on the brink of extinction. In the remaining two lakes, however, Arctic charr dominates, and occurs in higher numbers than whitefish in all the habitats. Our observations indicate that coexistence of the two species in oligotrophic and relatively pristine lakes requires an extensive profundal zone to serve as a refugium for Arctic charr. If the littoral zone is rendered inaccessible or unprofitable for whitefish due to dominance of a third competitor or predator, or as a result of lake regulation, then Arctic charr may be the dominant species.     

Diel food selection of pelagic Arctic charr, Salvelinus alpinus (L.), and brown trout, Salmo trutta L., in Lake Atnsjø, SE Norway

Patterns of diel food selection in pelagic Arctic charr, Salvelinus alpinus (L.) and brown trout, Salmo trutta L. were investigated in Lake Atnsjo, SE Norway, by gillnet sampling during July-September 1985. Arctic charr feed almost exclusively on zooplankton both day and night, while brown trout had a diurnal shift in diet. For this species zooplankton made up a considerable part of the diet in the daytime, while at night the diet consisted mainly of surface insect and chironomid pupae. Both species had a selective feeding mode on zooplankton during the day and night. Arctic charr had a higher gill raker number and a denser gill raker spacing compared with brown trout. Still, the differences in prey size between the two species were small. We argue that the observed differences in food selection between Arctic charr and brown trout can be explained by differing abilities to detect food items under low light conditions.     

Effects of Fish-farm Activity on the Limnetic Community Structure of Brown Trout, Salmo trutta, and Arctic Charr, Salvelinus alpinus

Establishment of four fish-farms during the period 1971 to 1994 in the oligotrophic lake Skogseidvatnet affected Arctic charr, Salvelinus alpinus, but not brown trout, Salmo trutta. From 1971 to 1987, an increase in mean individual size of Arctic charr was recorded, while the mean individual size of brown trout remained stable. Arctic charr were found to use deeper benthic areas than brown trout. Approximately 8% of the Arctic charr population (>26cm), were found to switch to waste food from fish-farms, resulting in a novel feeding habitat for the species. They were, however, found in gillnets distant from the fish farm cages, indicating high mobility. The habitat segregation between the two species can most likely be explained by selective differences and asymmetric competition with brown trout as the dominant species. Based on the present results, changes in the Arctic charr population may be due to increased food availability and due to a new habitat use as a waste food feeder. The reason for the brown trout population to have remained stable with respect to mean size, growth pattern and habitat use, may be due to a different diet choice than Arctic charr in this lake. Brown trout were found to feed mainly on terrestrial insects, while Arctic charr fed mainly on zooplankton and on waste food.     

Mitochondrial DNA diversity among four sympatric morphs of Arctic charr, Salvelinus alpinus L., from Thingvallavatn, Iceland

Restriction fragment length polymorphisms in mitochondrial DNA (mtDNA) were examined in progeny of four sympatric morphs of Arctic charr ( Salvelinus alpinus L.) from Thingvallavatn, Iceland. The mtDNA analysis with 46 hexanucleotide restriction enzymes indicates that the Thingvallavatn morphs are very closely related. Sequence divergence was less than 0.2% between any of the five clones detected. Although not significant, the topology of the mtDNA UPGMA dendogram was similar to that from a previous allozyme survey. Icelandic Arctic charr show greater affinity to charr from the British Isles than those from North America supporting their current taxonomic distinction.     

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.     

Stable isotope analysis reveals ecological segregation in a bimodal size polymorphism in Arctic charr from Loch Tay, Scotland

The serendipitous discovery of a body-size dimorphism amongst the sexually mature Arctic charr Salvelinus alpinus of Loch Tay is described. Sexually mature Arctic charr, collected by gill netting on spawning areas, showed a clear and distinct bimodal size distribution with no overlap in fork length distributions. The upper (19–29 cm L _F) and lower modes (8–16 cm) were not solely the result of sex or age differences. Analysis of stable isotope ratios of C and N in muscle showed highly significant differences in mean δ¹³C and δ¹⁵N between populations, demonstrating a difference in trophic ecology between the two body-size morphs. Overlap in the range of δ¹³C and δ¹⁵N values for the two morphs, however, suggested that they occasionally shared a common diet. Data from other studies strongly indicated that the proximate and ultimate mechanisms that control body-size dimorphisms in Arctic charr differed between sites. Clear differences in trophic ecology in the Loch Tay Arctic charr suggested that the available feeding opportunity may differ for the two morphs. The most likely proximate mechanism resulting in this dimorphism is growth rate differences resulting from differences in food availability for the two subgroups occupying alternative foraging niches in Loch Tay.     

Smolting characters in anadromous and resident Arctic charr, Salvelinus alpinus (L.)

Morphological differences, haematocrit value and chloride cells were examined in downstream migrating Arctic charr, Salvelinus alpinus (L.) from the Hals River, North Norway, and resident charr from Lake Storvann. Fish were classified as visual parr, silvery parr or smolt based on the degree of silvering and the Occurrence of lateral parr marks. On average, 47% of downstream migrating Arctic charr were classified as visual molts, but only 14% of the resident charr. Charr longer than 20 cm fork length were mainly classified as visual molts. Morphometrical analyses of body size and shape revealed that most of the variations could be explained in terms of variations in fork length. Length adjusted ratios of post-anal distances were significantly higher in migrating charr than in resident charr. The number of developed chloride cells, the cell nucleus diameter in the basin of secondary gill lamellae and blood haematocrit values were significantly higher among downstream migrating charr than in resident charr. However, morphological differences between anadromous and resident Arctic charr were not necessarily synchronized with the development of chloride cells.     

Late ontogeny growth acceleration and size form transformations in Transbaikalian Arctic charr, <Emphasis Type="Italic">Salvelinus alpinus</Emphasis> complex: evidence from fin ray cross section growth layers

In some polymorphic populations of Arctic charr in Transbaikalia, an individual can transform from a smaller to larger size form during their lifetime as a result of accelerated growth that follows a period of slow growth and reproduction as a small size form. Alternating periods of slow and fast growth are reflected in growth layer patterns visible in fin ray cross sections. Stained microtome fin ray cross sections were used to reveal the incidence of transformations from one form to another. Data were collected from 14 northern Transbaikalian lakes containing two or three sympatric Arctic charr forms (‘dwarf’, ‘small’, and ‘large’) exhibiting varying levels of morphological separation. Individuals recruited from the dwarf or small form were found in varying proportions among the small and/or large form in 12 lakes. Small or large form charr that grew without noticeable acceleration to the adult size typical of the form or experienced accelerated growth as juveniles prior to maturation were also observed. There were no transformations between sympatric forms that differed in the length and number of gill rakers and in some other meristic characters. Results indicate that in the region under study, transformations of sympatric Arctic charr size forms are a widespread, but not a ubiquitous, phenomenon. Such transformations reflect the plasticity of the developmental channels of the forms. In the course of intra-lacustrine form divergence and genetic differentiation, the frequency of the observed transformations decreases to zero.     

Microsatellite and mitochondrial DNA assessment of population structure and stocking effects in Arctic charr Salvelinus alpinus (Teleostei: Salmonidae) from central Alpine lakes

Despite geographical isolation and widespread phenotypic polymorphism, previous population genetic studies of Arctic charr, Salvelinus alpinus , have detected low levels of intra- and interpopulation variation. In this study, two approaches were used to test the generality of low genetic diversity among 15 Arctic charr populations from three major drainages of the central Alpine region of Europe. First, a representative subsample of each drainage was screened by PCR–RFLP analysis of mtDNA using 31 restriction enzymes. All individuals but one shared an identical haplotype. In contrast, microsatellite DNA variation revealed high levels of genetic diversity within and among populations. The number of alleles per locus ranged from six to 49, resulting in an overall expected heterozygosity from 0.72 ± 0.09 to 0.87 ± 0.04 depending on the locus. Despite evidence for fish transfers among Alpine charr populations over centuries, genetic diversity was substantially structured, as revealed by hierarchical Φ statistics. Eighteen per cent of total genetic variance was apportioned to substructuring among Rhône, Rhine, and Danube river systems, whereas 19% was due to partitioning among populations within each drainage. Cluster analyses corroborated these results by drainage-specific grouping of nonstocked populations, but also revealed damaging effects of stocking practices in others. However, these results suggest that long-term stocking practices did not generally alter natural genetic partitioning, and stress the importance of considering genetic diversity of Arctic charr in the Alpine region for sound management. The results also refute the general view of Arctic charr being a genetically depauperate species and show the potential usefulness of microsatellite DNAs in addressing evolutionary and conservation issues in this species.