Effects of stocking of piscivorous brown trout, Salmo trutta L., on stunted Arctic charr, Salvelinus alpinus (L.)

To study the effects on a stunted freshwater population of Arctic charr, Salvelinus alpinus (L.), two groups of large (26–45 cm) individually tagged brown trout, Salmo trutta L., were released and recaptured with gillnets after 1, 7, 11 and 63 weeks. One group of trout was trained on a fish diet before release, and the other, reared on commercial dry pellets, served as a control. Specific growth rates in both groups were negative 1 week after release and approached zero after 63 weeks. Condition factor and internal fat content decreased during the experiment. Although only 11% of the trout stomachs examined contained fish prey, charr represented 79% of the total stomach weight content. Gillnet samples of charr before and 63 weeks after the release of trout indicated a decreasing population size of charr. Individual growth and mean length of charr increased after release of trout, especially for charr at age 4 years. After the release of trout, 35% of the charr were longer than 20 cm as compared with 6% before the release.     

Habitat use, size and age structure in sympatric brown trout (Salmo trutta) and Arctic charr (Salvelinus alpinus) stocks: resistance of populations to change following harvest

Abstract– Habitat use and population dynamics in brown trout Salmo trutta and Arctic charr Salvelinus alpinus were studied in an oligotrophic lake over a period of 10 years. Previous studies showed that the species segregated by habitat during summer. While brown trout occupied the surface water down to a depth of 10 m, Arctic charr were found deeper with a maximum occurrence at depth 10–15 m. Following the removal of a large number of intermediate sized fish in 1988–89, habitat segregation between the species broke down and Arctic charr were found in upper waters, while brown trout descended to deeper waters. The following year, both species were most frequently found in surface waters at depths of 0–5 m. During the last four years, the species reestablished their original habitat segregation despite another removal experiment of intermediate-sized fish in 1992–1994. The removal of fish resulted in an increased proportion of large (≥ 25 cm) fish in both species. Furthermore, the charr stock responded by reduced abundance and increased size-at-age. The results revealed plasticity and strong resistance to harvest populations of brown trout and Arctic charr. This is probably due to internal mechanisms of intraspecific competition within each population, which result in differential mortality among size classes.     

Long-term variation in the population structure of Arctic charr, Salvelinus alpinus, and brown trout, Salmo trutta

The effects of induced water level fluctuations and introduction of the mysid Mysis relicta Lovén on population structure of brown trout, Salmo trutta L., and Arctic charr, Salvelinus alpinus (L.), were studied during 1953–1995 in Limingen hydroelectric reservoir, Norway. The main response was a marked reduction in catch‐per‐unit‐effort (CPUE) for trout and charr, probably caused by reduced recruitment following increased variation in water level. For both species, mean length decreased until 1967 and increased thereafter, whereas mean mass‐at‐length increased for the whole period. Both length and mass‐at‐length were negatively correlated with CPUE. The increases in mean length and mass‐at‐length were probably because of reduced competition following the reduced recruitment. Mysis relicta has become an important food item for charr but not for brown trout, but the increases in mean length and mass‐at‐length of charr started prior to the appearance of M. relicta in the charr diet.     

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.     

Food composition, habitat use and growth of stocked and native Arctic charr, Salvelinus alpinus, in Lake Muddusjärvi, Finland

Habitat use, growth and food composition of native and stocked Arctic charr, Salvelinus alpinus (L.), were studied in the subarctic Lake Muddusjärvi, northern Finland, to investigate reasons for poor stocking success. Samples were collected with pelagic and epibenthic gill nets. Stocked and native charr occurred in similar epibethic habitats, whereas pelagic habitat was avoided. Native charr grew fast after shifting to piscivory. Growth rate of stocked charr was slow because only a small proportion of stocked fish became piscivorous during the first year after stocking. During the first lake year, stocked charr divided into slow-growing planktivores and fast-growing piscivores. Piscivorous stocked and native charr consumed only whitefish, Coregonus lavaretus (L.), as their prey. Small-sized (<10 cm) whitefish were preferred when shifting to piscivory.     

Atlantic salmon Salmo salar L., brown trout Salmo trutta L. and Arctic charr Salvelinus alpinus (L.): a review of aspects of their life histories

Abstract – Among the species in the family Salmonidae, those represented by the genera Salmo, Salvelinus, and Oncorhynchus (subfamily Salmoninae) are the most studied. Here, various aspects of phenotypic and life‐history variation of Atlantic salmon Salmo salar L., brown trout Salmo trutta L., and Arctic charr Salvelinus alpinus (L.) are reviewed. While many strategies and tactics are commonly used by these species, there are also differences in their ecology and population dynamics that result in a variety of interesting and diverse topics that are challenging for future research. Atlantic salmon display considerable phenotypic plasticity and variability in life‐history characters ranging from fully freshwater resident forms, where females can mature at approximately 10 cm in length, to anadromous populations characterised by 3–5 sea‐winter (5SW) salmon. Even within simple 1SW populations, 20 or more spawning life‐history types can be identified. Juveniles in freshwater can use both fluvial and lacustrine habitats for rearing, and while most smolts migrate to sea during the spring, fall migrations occur in some populations. At sea, some salmon undertake extensive oceanic migrations while other populations stay within the geographical confines of areas such as the Baltic Sea. At the other extreme are those that reside in estuaries and return to freshwater to spawn after spending only a few months at sea. The review of information on the diversity of life‐history forms is related to conservation aspects associated with Atlantic salmon populations and current trends in abundance and survival. Brown trout is indigenous to Europe, North Africa and western Asia, but was introduced into at least 24 countries outside Europe and now has a world‐wide distribution. It exploits both fresh and salt waters for feeding and spawning (brackish), and populations are often partially migratory. One part of the population leaves and feeds elsewhere, while another part stays as residents. In large, complex systems, the species is polymorphic with different size morphs in the various parts of the habitat. Brown trout feed close to the surface and near shore, but large individuals may move far offshore. The species exhibits ontogenetic niche shifts partly related to size and partly to developmental rate. They switch when the amount of surplus energy available for growth becomes small with fast growers being younger and smaller fish than slow growers. Brown trout is an opportunistic carnivore, but individuals specialise at least temporarily on particular food items; insect larvae are important for the young in streams, while littoral epibenthos in lakes and fish are most important for large trout. The sexes differ in resource use and size. Females are more inclined than males to become migratory and feed in pelagic waters. Males exploit running water, near‐shore and surface waters more than females. Therefore, females feed more on zooplankton and exhibit a more uniform phenotype than males. The Arctic charr is the northernmost freshwater fish on earth, with a circumpolar distribution in the Holarctic that matches the last glaciation. Recent mtDNA studies indicate that there are five phylogeographic lineages (Atlantic, Arctic, Bering, Siberian and Acadian) that may be of Pleistocene origin. Phenotypic expression and ecology are more variable in charr than in most fish. Weights at maturation range from 3 g to 12 kg. Population differences in morphology and coloration are large and can have some genetic basis. Charr live in streams, at sea and in all habitats of oligotrophic lakes, including very deep areas. Ontogenetic habitat shifts between lacustrine habitats are common. The charr feed on all major prey types of streams, lakes and near‐shore marine habitats, but has high niche flexibility in competition. Cannibalism is expressed in several cases, and can be important for developing and maintaining bimodal size distributions. Anadromy is found in the northern part of its range and involves about 40, but sometimes more days in the sea. All charr overwinter in freshwater. Partial migration is common, but the degree of anadromy varies greatly among populations. The food at sea includes zooplankton and pelagic fish, but also epibenthos. Polymorphism and sympatric morphs are much studied. As a prominent fish of glaciated lakes, charr is an important species for studying ecological speciation by the combination of field studies and experiments, particularly in the fields of morphometric heterochrony and comparative behaviour.     

The influence of predation risk on habitat selection and food intake by Arctic charr, Salvelinus alpinus (L.)

Abstract— Large piscivorous fish are assumed to affect habitat selection and food intake of prey fish. To study the effects of cannibalistic Arctic charr, Salvelinus alpinus (L.), on smaller stunted charr, we sampled the prey fish in littoral and pelagic habitats using gill nets, before and shortly after the release of large charr in a small lake (0.52 km²). In the habitats where the risk of predation was highest, the catch per unit effort de creased from 13.3 to 4.8 fish per 100 m² of gillnet after release of pred ators. The large decrease in numbers of charr < 18 cm corresponded with the predicted vulnerable prey sizes, according to a model based on the size distribution of predators. The occurrence of planktivorous fish and weight-specific food intake decreased in the high risk habitat and remained unaffected in the low risk habitats. Changes in the food intake of prey fish could not be explained in terms of fish length, indicating that prey fish changed diet when the risk of predation was high.     

Resource partitioning between lake-dwelling Atlantic salmon (Salmo salar L.) parr, brown trout (Salmo trutta L.) and Arctic charr (Salvelinus alpinus (L.))

Abstract – Resource partitioning between Atlantic salmon parr, brown trout and Arctic charr was studied throughout the ice-free season in a north Norwegian lake. Juvenile salmon and trout (≤160 mm) utilized the littoral zone and juvenile charr the profundal, while adult trout and charr (>160 mm) were found in both. Juvenile salmon and trout had a similar diet, although trichopteran larvae were more important for the trout and chironomid pupae and three-spined sticklebacks for the salmon parr. Small salmon and trout parr (≤120 mm) had a higher diet overlap than larger parr (121–160 mm). The feeding habits of adult trout were similar to that of juvenile trout, but the former took larger prey items. At the population level, both salmon and trout were generalistic feeders with a broad diet, but at the individual level, both species had specialized on a single or a few prey categories. Juvenile charr were segregated from salmon and trout in both habitat and food utilization; they had a narrow diet consisting of chironomids and zooplankton, possibly reflecting their confinement to the profundal habitat which have a low diversity of potential prey. Larger charr also took zoobenthos and sticklebacks in the littoral zone. ^Note     

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.     

Littoral spawning habitats of three southern Arctic charr (Salvelinus alpinus L.) populations

Abstract Arctic charr populations in southern latitudes are nonmigratory, with all life‐stages limited to freshwater lakes and in‐ or out‐flowing tributaries. Although many of these populations are reported to also spawn in lake littorals, little is known about the physical characteristics of putative spawning grounds. A total of 23 discrete spawning sites within three Irish lakes were located by fyke netting of spawning adults and snorkelling in littoral habitats. Spawning sites were found to be long, narrow strips running parallel to the shore at a maximum depth of 124 cm. Spawning sites were limited to areas of coarse mineral substrate with an adequate (c. 8 cm) depth of clean interstitial spaces. In individual lakes, combined areas of spawning sites made up 0.4–0.7% of available littoral. Egg densities varied considerably between sites (33–900·eggs m^?2) and were significantly correlated with gradient and width of spawning sites. No evidence of redd digging was found. The shallow, localised and restricted nature of spawning grounds makes such populations vulnerable to anthropogenically induced postoviposition changes in surface water level, eutrophication processes such as increased lake sedimentation and elevated nutrient status.     

Consequence of habitat segregation to growth rate of two sparsely rakered whitefish (Coregonus lavaretus (L.)) forms in a subarctic lake

Abstract  – The aim of this study was to explain the growth differences of the two sympatric sparsely rakered whitefish ( Coregonus lavaretus (L.)) forms in the subarctic Lake Muddusjärvi by comparing their habitat and food selection to available benthic food resources. The average number of gillrakers was 22.9 for large sparsely rakered whitefish (LSR) and 16.8 for small sparsely rakered whitefish (SSR). LSR dwelled primarily in the littoral zone at depths < 10 m, whereas SSR used mostly profundal habitats in depths > 10 m. LSR and SSR consumed mainly benthic macroinvertebrates, their abundance, biomass and diversity was highest in the littoral, decreasing towards the profundal. As the individual size of fish increased, LSR was able to shift to larger food items, which were available in the littoral. Possibility of SSR to ontogenetic food shift was scarce in the profundal, where availability of larger benthic macroinvertebrates was low. Because of abundant food resources in the littoral, growth rate of LSR was faster than that of SSR.     

Habitat utilization of juvenile Atlantic salmon (Salmo salar L.), brown trout (Salmo trutta L.) and Arctic charr (Salvelinus alpinus (L.)) in two lakes in northern Norway

Abstract— Habitat utilization of juvenile Atlantic salmon, brown trout and Arctic charr was investigated in two lakes in northern Norway during the icefree season. Both the vertical distribution and the distribution among different habitat types were studied by gillnetting with small mesh sized gillnets (8-15 mm) in different habitats. Salmon and trout were predominantly caught in the littoral and sublittoral zones (0-6 m depth). Access to shelter seemed to be the most important factor determining the horizontal distribution of small salmon and trout. Most of these fish were caught in stony or vegetated habitats, while few salmon and trout were caught on sandy locations or in the pelagic zone. In one of the lakes, there were significantly higher catch rates of salmon than of trout in the stony littoral (0-3 m), while in the other lake there were no significant differences in spatial distribution between these two species. Charr were primarily found in the profundal, sublittoral or pelagic zones of the lakes.     

The influence of physical watercourse parameters on the degree of anadromy in different lake populations of Arctic charr (Salvelinus alpinus (L.)) in northern Norway

The influence of physical watercourse parameters on the degree of anadromy was studied in 15 lake populations of Arctic charr (Salvelinus alpinus (L.)) in northern Norway. The anadromy of the different populations was scored based on the following criteria; marine parasites. fish length and length at maturity. There were significant negative correlations between the anadromy scores and both the length of the outlet rivers of the different lakes and a migration barrier index for the different watercourses (combining migration distance and water velocity). It is hypothsized that these correlations were due to differences in energy expenditure for charr during upstream migration and/or differences in predation pressure on charr during river migration. Short rivers with relatively high water velocity seem to be the most preferable conditions for the anadromous life-history strategy. There were no correlations between the degree of anadromy and mean river slope or minimum summer water discharge.     

Divergent, genetically based feeding behaviour of two sympatric Arctic charr, Salvelinus alpinus (L.), morphs

Abstract –  Arctic charr, Salvelinus alpinus (L.), is one of the several northern fishes that show resource polymorphisms in postglacial lakes. Two reproductively isolated morphs of Arctic charr coexist in distinct ecological niches in the subarctic lake Fjellfrøsvatn, North Norway. Offspring of the two morphs (termed littoral charr and profundal charr) were reared separately but under identical conditions. Their feeding behaviour was compared experimentally using different kinds of live prey. The fishes had no experience with natural prey before the experiments. The littoral morph were more effective in eating live plankton ( Daphnia ) and littoral benthos ( Gammarus ), and had a higher attack rate against pleuston (surface prey, Gerris ) compared with the profundal morph. The two morphs behaved in accordance with expectations from their in situ niche utilisation towards the three prey types. This indicates a case of incipient ecological speciation where divergence in resource utilisation in contrasting niches has evolved adaptations in feeding behaviour by natural selection.     

An experimental test of the genetic component of the ontogenetic habitat shift in Arctic charr (Salvelinus alpinus)

Abstract— Fry of the Arctic charr, Salvelinus alpinus , were experimentally stocked into a small fish-free lake to test the hypothesis that the size-dependent habitat shift from the epibenthic to the pelagic habitat is genetically determined. The charr originated from a nearby lake inhabiting predatory brown trout Salmo trutta. The cohort of stocked charr was investigated for three years. The Arctic charr started to exploit the pelagic habitat in their first summer at a size of 7–9 cm in contrast to about 15 cm in the donor lake. In the next two summers, the pelagic fraction of the cohort increased. The main fraction lived in epibenthic areas, utilizing the same prey as pelagic charr. Water temperature moderated the habitat use of juveniles such that they avoided warm (>16°C) waters and resided in cool, deep areas. The result was consistent with the hypothesis of a tradeoff between feeding benefit and the predation risk producing spatial segregation of Arctic charr and demonstrated that the fish can facultatively respond to predation risk and adjust the size at which they migrate to the pelagic zone to feed on zooplankton.     

Planktivory and diet-overlap of densely rakered whitefish (Coregonus lavaretus (L.)) in a subarctic lake

Abstract –  Three sympatric whitefish ( Coregonus lavaretus (L.)) forms, one being pelagic and two benthic, segregate available habitat and food resources in subarctic Lake Muddusjärvi, northern Finland. Zooplankton availability in the lake, food composition, diet-overlap and growth of densely rakered (DR) whitefish were examined during June to September to explore the reasons for the small individual size of the pelagic form. DR whitefish used zooplankton as main food item and prey selection followed zooplankton species density proportions in the lake. Zooplankton density and water temperature was highest in July–August. The average lengths of Bosmina spp., Daphnia spp., Calanoida and Cyclopoida in DR whitefish stomach were higher than in zooplankton sample during June–September, except Calanoida in June. Diet-overlap between DR whitefish age groups was high at all months indicating intercohort resource competition. DR whitefish reached sexual maturity at 3 years of age and at the length of 12 cm, after which somatic growth almost ceased. Reason for the small average size and slow growth of DR whitefish were connected to high diet-overlap between age groups and early sexual maturation.     

Homing experiments with parr, smolt and residents of anadromous Arctic char Salvelinus alpinus and brown trout Salmo trutta: transplantation between neighbouring river systems

Abstract –  Olfaction is decisive for the homeward migration of anadromous salmonids. Two different olfactory hypotheses for explaining how this mechanism works have been proposed (the imprinting and the pheromone hypothesis), and they differ with regard to the origin of the odours that define home. The pheromone hypothesis assumes that fish respond to strain-specific pheromones and that no imprinting is necessary. The imprinting hypothesis assumes that juvenile fish become imprinted to odours of abiotic or biotic origin in natural rivers; the imprinting may be a single event or may occur sequentially during downstream migration as smolts. The two hypotheses were challenged by reciprocal transplantation of parr, residents and smolt of Arctic char ( Salvelinus alpinus ) and brown trout ( Salmo trutta ) between the Salangen and the Løksebotten rivers. Both rivers empty into the Salangen Fjord, Norway, about 10 km apart. Salangen Arctic char released in the Løksebotten River, and Løksebotten brown trout released in the Salangen River (parr, resident and smolt) initiated spontaneous homeward migration without preceding contact with home stream water. This indicates that site imprinting (single or sequential) could not have been involved. In the sea, the released fish apparently integrated into the migratory system of relatives from their river of origin. Recapture rates did not differ for fish allowed to receive single or sequential imprinting.     

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     

Char ecology. Natal homing in sympatric populations of anadromous Arctic char Salvelinus alpinus (L.): roles of pheromone recognition

Abstract –  This study revealed that progeny of sympatric Salangen Arctic char, omitted from any kind of imprinting to home water, returned with high precision to the home river and parental locality upon releases in their native marine environments. Fish did not select river randomly, and the homing precision was the same for progeny whether fertilized in home stream or in foreign water. The progeny studied originate from the lake Øvervatnet and the Salangen River population and were hatched and reared at a hatchery at Voss in southern Norway. Experimental material comprised freshwater resident and smolt released from parr directly. The freshwater resident is partly anadromous exhibiting about 74% smolt when released directly into the sea. Char at all stages (resident, transformed resident, and smolt released from parr directly) returned in significant numbers to the home river system and the site of the specific parental population.     

The effect of dietary inulin on aerobic bacteria associated with hindgut of Arctic charr (Salvelinus alpinus L.)

The primary aim of the present study was to evaluate the population level of adherent (autochthonous) aerobic and facultative anaerobic bacteria in the hindgut of healthy Arctic charr (Salvelinus alpinus L.) fed dextrin or inulin. This was assessed by the dilution plate technique, and visualized using both transmission and scanning electron microscopy. A population level of 4.8 × 10⁵ adherent bacteria per gram wet mass was found in the hindgut of fish fed a casein‐based diet supplemented with 15% dextrin. However, substituting dextrin with 15% inulin reduced the bacterial population level in the hindgut (3.56 × 10⁴). A total of 217 bacterial isolates were identified by key phenotypical and biochemical characteristics. In addition, 22 strains were also identified by partial sequencing of the 16S rRNA gene. The composition of bacteria colonizing the hindgut of Arctic charr fed dextrin was dominated by the genera Staphylococcus, Pseudomonas, Micrococcus, Psychrobacter glacincola and Streptococcus. However, bacteria colonizing the hindgut of fish fed inulin were dominated by Gram‐positive bacteria of the genera Staphylococcus, Streptococcus, Carnobacterium and Bacillus. While Carnobacterium divergens‐like strains were isolated from charr fed dextrin, Carnobacterium maltaromicus‐like strains were isolated from the hindgut of fish fed inulin. Electron microscopical analysis of hindgut regions confirmed traditional culture‐based microbial analysis as fewer bacterial cells were observed between microvilli and associated with the surfaces of enterocytes of fish fed inulin rather than dextrin.