The origins and persistence of anadromy in brook charr

We synthesized the results of a coordinated study examining the spatial and temporal movements, genetic structure, and physiological characteristics of sympatric populations of resident and sea-run brook charr across eastern Canada. Our goal was to critically evaluate three working hypotheses that may explain anadromous behaviour in brook charr: (1) resident and anadromous forms have different phylogenic origins; (2) anadromy emerges from freshwater residents; and (3) freshwater residency emerges from anadromous individuals. Our synthesis is consistent with the conclusion that freshwater residency emerging from anadromous individuals is most probable. Overall, anadromy in brook charr is poorly developed and the tactic of sea-running may be described as partial, facultative anadromy if the species must be described in terms of anadromy. The sea-run tactic most probably results from the species’ propensity to move and disperse, the over-production of juveniles, an archetypical physiological ability to tolerate saline environments, and the persistence of critical habitats. When the spatial and temporal physical environments permit, the anadromous behaviour is expressed. The tactic creates an apparent fitness advantages related to growth, but it is not necessarily the only evolutionary stable strategy for a river. Most populations with sea-run forms are declining and successful conservation of sea-run forms will depend on managing harvests and more importantly, protection of the temporally and spatially complexity of critical habitats.     

The growth history of anadromous white-spotted charr in northern Japan: a comparison between river and sea life

Anadromous white-spotted charr Salvelinus leucomaenis in a natural river in northern Japan migrated to the sea mainly at age 3+ years. Sea-run charr grew faster than parr, particularly as first time migrants. Sea-run charr repeated seaward migration up to three times; the observed maximum age was 5+ years old. The likelihood ratio test identified significant differences in asymptotic size and individual growth variability between parr and sea-run charr. Sea-run charr had an asymptotic size ( L ^∞) twice as large as parr, while the growth parameter ( K ) was similar between sea-run and parr. Growth variability was higher for parr than for sea-run charr.     

Parasites as indicators of individual feeding specialization in Arctic charr during winter in northern Norway

The relationship between infection with the food-transmitted parasites Diphyllobothrium dendriticum, D. ditremum (Cestoda) and Cystidicola farionis (Nematoda) and prey selection was studied in individual Arctic charr from Lake Takvatn, northern Norway. There was no correlation between parasites transmitted throughout prey organisms from benthic habitats (amphipods) and pelagic habitats (copepods). A strong relationship between infection with a parasite species and the corresponding intermediate host from the stomach content of individual charr, indicated an individual feeding specialization. Independent of size, charr specialized on the intermediate hosts of all three parasite species. Some charr maintained this specialization on specific prey items throughout the winter period. These parasite species are considered to be useful indicators of past prey selection.     

Effect of female size on fecundity and egg size in white-spotted charr: comparison between sea-run and resident forms

The relationships between fecundity, egg size and female size of sea-run form were compared with resident form, using white-spotted charr, Salvelinus leucomaenis. Both fecundity and egg size increase with female size. However, the relationship between egg size and female size differed significantly between the resident and sea-run forms. Egg sizes of sea-run and resident were similar even though sea-run fish were much larger. ? 1998 The Fisheries Society of the British Isles     

Food borne parasites as indicators of trophic segregation between Arctic charr and brown trout

The habitat and diet choice and the infection (prevalence and abundance) of trophically transmitted parasites were compared in Arctic charr and brown trout living sympatrically in two lakes in northern Norway. Arctic charr were found in all main lake habitats, whereas the brown trout were almost exclusively found in the littoral zone. In both lakes the parasite fauna reflected the niche segregation between trout and charr. Surface insects were most common in the diet of trout, but transmit few parasites, and accordingly the brown trout had a relatively low diversity and abundance of parasites. Parasites transmitted by benthic prey such as Gammarus and insect larva, were common in both salmonid host species. Copepod transmitted parasites were much more common in Arctic charr, as brown trout did not include zooplankton in their diets. Parasite species that may use small fish as transport hosts, were far more abundant in piscivorous fish, especially brown trout. The seasonal dynamics in parasite infection were also consistent with the developments in the diet throughout the year. The study demonstrates that the structure of parasite communities of charr and the trout is highly dependent on shifts in habitat and diet of their hosts both on an annual base and through the ontogeny, in addition to the observed niche segregation between the two salmonid species.     

Parasites of freshwater resident and anadromous Arctic charr (Salvelinus alpinus) in Greenland

Twenty-one metazoan parasite species were found in Arctic charr at three coastal sites in west Greenland, including Neascus sp. and the nematode Pseudoterranova decipiens , new records for Arctic charr. Twelve species are new records for Arctic charr in Greenland. The quantitative composition of the parasite communities varied, but the freshwater species ( Diphyllobothrium ditremwn, Eubothriutn salvelini, Proteocephalus longicollis, Crepidostotnutn farianis ) were usually numerically dominant, even in anadromous fish. The absence from Greenland of Mysidacea, Amphipoda, Ephemeroptera and Odonata excludes parasites requiring such organisms as intermediate hosts, including the cestode Cyathocephalus truncatus , nematodes of the genus Cystidicola , and freshwater acanthocephalans.     

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.     

The relationships between anadromy, sex ratio and parr growth of Arctic charr in a lake in North Norway

Relationships between the degree of anadromy, sex ratio and parr growth of Arctic charr Salvelinus alpinus were investigated in Lake Storvatn in Hammerfest town, by estimating the densities of resident and anadromous charr in the lake, and by comparing the smolt characteristics of fish captured in the outlet river with those of fish of the same age in the lake. About two-thirds of the charr older than 5 years were anadromous. The sex ratio of smolts was approximately 1 : 1 and the frequency of mature male parr in the lake was low. As females tended to dominate the mature portion of the anadromous population, there was probably a relatively higher mortality among sea-run male smolts. The youngest smolts (3+ and 4+) were probably recruited from fast-growing parr in the littoral zone, while older smolts (>4+) may have been recruited from all habitats in the lake.     

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.     

Competitive interactions for foraging microhabitat among introduced brook charr, Salvelinus fontinalis, and native bull charr, S. confluentus, and westslope cutthroat trout, Oncorhynchus clarki lewisi, in a Montana stream

Competitive interactions for foraging microhabitat among introduced brook charr, Salvelinus fontinalis, and native bull charr, S. confluentus, and westslope cutthroat trout, Oncorhynchus clarki lewisi, were studied by species removal experiments in a tributary of the Flathead Lake and River system, northwestern Montana, focusing on brook charr influences on bull charr. When the three species were in sympatry, they interacted with each other, forming a size-structured, mixed-species dominance hierarchy in two stream pools. The influences of interference interactions were examined by measuring changes in five characteristics of foraging microhabitat and behavior, focal point height and velocity, cover use, and foraging rate and distance, after the successive removal of two species. Cutthroat trout removal resulted in increased foraging rates and distances, and decreased cover use for brook charr, but no changes for bull charr. After removal of brook charr from the two-species system, bull charr also increased foraging rates and distances and occupied more exposed positions. Moreover, total fish densities, which had initially decreased owing to the removal experiments, were partly compensated for by subsequent bull charr immigration, implying that competitive interactions with brook charr are an important factor in the mechanisms responsible for the regulation of bull charr densities, at least on a local scale.     

Genetic diversity and population structure of white-spotted charr, Salvelinus leucomaenis, in the Lake Biwa water system inferred from AFLP analysis

The genetic variations—and the time dependence of such variations—of natural populations of the white-spotted charr ,Salvelinus leucomaenis, in the Lake Biwa water system as well as those of a hatchery-reared population were inferred from AFLP. Upon the application of principal coordinate analysis using 118 polymorphic AFLP fragments based on the Jaccard similarity index, specimens of each of six natural local populations from the inlet rivers of Lake Biwa grouped roughly together, suggesting that each local population was genetically differentiated. The hatchery-reared population was shown to be closely related to the local population in the Seri River, suggesting that the Seri River population originated from hatchery-reared charr due to extensive stocking. Furthermore, specimens of the Yasu River grouped in a somewhat different position from the other natural populations, agreeing well with its geographic distance from the other populations. The nucleotide diversities of six natural populations (Harihata River, Ishida River, two reaches of the Takatoki River, Ane River, and Yasu River) in 2002 or 2003 were relatively low (π = 0.067–0.146%) compared with that of the Seri River (0.278%) and the hatchery-reared charr (0.316%). The nucleotide diversity in the five local populations (Ishida River, two reaches of the Takatoki River, Ane River, and Yasu River) remained at a low level from 1994 to 2002/2003, but only the nucleotide diversity in the Harihata River actually decreased. From 1994 to 2002/2003, the nucleotide diversity in the Seri River remained at a higher level among the natural populations from 1994 to 2002/2003; it was enhanced by the artificial release of hatchery-reared charr before 1994. In order to conserve the genetic diversity of the white-spotted charr in the Lake Biwa water system, it is necessary to prevent the stocking of hatchery-reared charr in reaches where hatchery-reared charr have not previously been stocked.     

Piscivory and cannibalism in Arctic charr

Piscivory and cannibalism in Arctic charr, Salvelinus alpinus , were studied in three lakes in northern Norway: Guolasjavri, which contains only charr, Takvatn, where Arctic charr coexist with three-spined sticklebacks, Gasterosteus aculeatus and brown trout, Salmo trutta , and Stuorajavri, where whitefish, Coregonus lavarelun dominate a fish community containing six species. The prevalence of piscivory in the Arctic charr populations generally increased with increasing predator size. In all three lakes, many charr larger than 20 cm were piscivorous, but the extent of piscivory and cannibalism varied. The greatest prevalence of cannibalism was found in Guolasjavri, where 27% of charr greater than 20 cm in length had fed upon smaller conspeciflcs. In Takvatn, 5% of charr larger than 20 cm were cannibalistic, and an additional 9% had eaten three-spined sticklebacks. In Stuorajavri, up to 74% of the charr greater than 20 cm had eaten whitefish but cannibalism was not recorded. The possible role of cannibalism in population regulation within Arctic charr populations is considered.     

Ecological parasitology of polymorphic Arctic charr, Salvelinus alpinus (L.), in Thingvallavatn, Iceland

The helminth endoparasite fauna in four Arctic charr morphs, Salvelinus alpinus (L.), small benthivorous (SB), large benthivorous (LB), planktivorous (PL) and piscivorous (PI) charr, from Thingvallavatn, Iceland consisted of: Crepidostomum farionis (Trematoda: Allocreadiidae); Diplosttomum sp. (Trematoda: Diplostomatidae); Eubothrium salvelini; Diphyllobothrium dendriticum; D. ditremum (Cestoda: Pseudophyllidae); Proteocephalus longicollis (Cestoda: Proteocepha-lidae): and Philonema oncorhynchi (Nematoda: Filariidae). The morphs exhibited distinctive patterns in prevalences and parasite burdens (mean intensity and mean relative density of parasites). SB charr had high prevalence and parasite burden of the eye fluke Diplostomum sp. and none to very light infections of the other parasite species. LB charr had relatively high prevalence and parasite burden of the intestinal fluke C. farionis , whereas infections of the remaining parasite species were light to moderate. PL and PI charr had high prevalences and worm burdens of Diphyllobothrium spp. and P. longicollis . PL charr differed from PI charr in higher worm burden off P. longicollis and lighter burden of £. salvelini . Prevalences of P. oncorhynchi were high in PL and PI charr. Association of parasite intensities and age and length offish were investigated. The different infection patterns among the morphs agree well with their partitioning in food and habitat utilization, and confirm that there is a high degree of ecological segregation between the morphs. The results demonstrate the importance of ecological factors influencing transmission efficiency of parasites to the fish host.     

Sex chromosome polymorphisms in Arctic charr and their evolutionary origins

Current data on the Y-specific sex-determining region of salmonid fishes from genera Salvelinus, Salmo, and Oncorhynchus indicate variable polymorphisms in the homologous chromosomal locations of the sex-specific determining region. In the majority of the Atlantic lineage Arctic charr, including populations from the Fraser River, in Labrador Canada, as well as Swedish and Norwegian strains, the sex-determining locus maps to linkage group AC-4. Previously, sex-linked polymorphisms (i.e., variation in the associated sex-linked markers on AC-4) have been described in Arctic charr. Here, we report further evidence for intraspecific sex linkage group polymorphisms in Arctic charr (i.e., the detection of the SEX locus on either the AC-1 or AC-21 linkage group) and a possible conservation of a sex linkage arrangement in Icelandic Arctic charr and Atlantic salmon, involving sex-linked markers on the AC-1/21 homeologs and the European AS-1/6 homeologous linkage groups in Atlantic salmon. The evolutionary origins for the multiple sex-determining regions within the salmonid family are discussed. We also relate the variable sex-determining regions in salmonids to their ancestral proto-teleost karyotypic origins and compare these findings with what has been observed in other teleost species in general.     

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.     

The second population of Arctic charr Salvelinus alpinus complex (Salmoniformes, Salmonidae) in the Lake Baikal basin, the highest mountain charr population in Russia

We report the finding of the second population of Arctic charr in Lake Baikal basin, in a nameless lake in the Barguzin mountain range in the outlet of the Svetlaya River. The lake is situated at an altitude of 1766 m above sea level; this is the highest lake inhabited by this species in the territory of Russia. Two abundant charr forms, dwarf and small, were recorded in the lake; data on their ecology, morphology, age composition, growth, and microsatellite variation are presented. Both forms feed mainly on zooplankton, though neither has pronounced morphological traits characteristic of specialized plankton-feeders. Dwarf and small charrs are rather close in meristic characters and in the length of gill rakers, but differ substantially in allele frequencies and allele diversity at the microsatellite loci, which is indicative of a high degree of reproductive isolation between them. The discovered population is the only one among Transbaikalian charr populations, that, due to its remoteness, has not been affected by man. For that reason, it can be considered as an etalon of an undisturbed natural charr population. The lake and its surroundings should receive official protected status.     

How fish-helminth associations arise: an example from Arctic charr in Loch Rannoch

Three sympatric morphs of Arctic charr Salvelinus alpinus occur in Loch Rannoch, Scotland, and are identified by their differing head morphology and diet. These are small-headed benthic, large-headed benthic and pelagic morphs. Six species of endoparasitic helminth were found in the fish, but the morphs had different patterns of infection. Overall infections in pelagic charr were heavier than in large-headed benthics, which were in turn heavier than in small-headed benthics, even though benthic charr live longer than pelagics. Pelagic fish had high prevalences and intensities of pseudophyllidean tapeworms, the intermediate hosts of which are copepods. The prevalence and intensity of metacercariae of Diplostomum sp. (the intermediate hosts of which are snails) were high in the benthic morphs. The results are discussed in terms of the effects of ecological factors on transmission of helminth parasites to their hosts and the evolution of host-parasite associations.     

Migration, growth and survival of wild and hatchery-reared anadromous Arctic charr (Salvelinus alpinus) in Finnmark, Northern Norway

Two groups of anadromous Arctic charr ( Salvelinus alpinus ) (size 200–350 mm) reared in heated water (6–12° C) under simulated natural photoperiod were individually tagged and released in spring 1988. The fish were released at two sites, in the estuary of the River Halselva and in the fjord, 2 km from the river mouth. Growth, timing of migration and survival of these hatchery-reared fish was compared to that of wild anadromous charr of the same size over a 4-year period. The hatchery-reared charr had poorer growth than the wild fish during their first year in sea water. They also resided longer in the sea and had a slightly lower survival than wild fish. During the second year, hatchery-reared charr displayed good growth, and after the third sea-season the fish were ready for slaughter at a size of approximately 800g. The results suggest that the successful development of Arctic charr ranching will be dependent upon production and release strategies that lead to improved migratory and feeding behaviour of the fish during their first season at sea.     

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.     

Trophic polymorphism in brook charr revealed by diet, parasites and morphometrics

Stomach contents, parasite assemblages and morphometrics were compared in brook charr Salvelinus fontinalis from the littoral and pelagic zone of two adjacent lakes on the Canadian Shield. In lac Baie des Onze Îles, fish from the littoral zone had greater abundance of benthic prey in their stomach and were more heavily infected by parasites that use intermediate hosts associated with the littoral zone than fish captured in the pelagic zone. Littoral and pelagic brook charr from this lake also differed in regard to body shape and fin length, with each group being anatomically adapted to exploit their respective habitats. The highly significant correlation between morphometric and parasite canonical scores supports the hypothesis of functional diversification of individuals within lac Baie des Onze Îles. While fish from littoral and pelagic zones of lac Caribou did not differ in terms of diet, parasite assemblages or morphometrics, they were different to fish from lac Baie des Onze Îles in that they were less frequently infected with parasites that use gastropods as intermediate hosts, and had shorter pectoral fins. The inter-lake comparisons suggested that parasite assemblages and morphometrics of brook charr reflected the dominance of the limnetic and littoral habitats in lacs Caribou and lac Baie des Onze Îles, respectively.