Brown trout (Salmo trutta L.) and Arctic charr (Salvelinus alpinus (L.)) as predators on three sympatric whitefish (Coregonus lavaretus (L.)) forms in the subarctic Lake Muddusjärvi

Abstract  – Brown trout ( Salmo trutta L.) and Arctic charr ( Salvelinus alpinus (L.)) use whitefish ( Coregonus lavaretus (L.)) as their main prey in the subarctic Lake Muddusjärvi. Brown trout dwelled in littoral and pelagic habitat, whereas Arctic charr lived only in epibenthic habitat. Both species shifted to whitefish predation at a length of 20–30 cm. At this size, brown trout fed on larger whitefish than Arctic charr. Whitefish occur in three sympatric forms, differing in their habitat, ecology and morphology. Both the predators preyed primarily upon the small-sized, densely rakered whitefish form (DR), which was the most numerous whitefish form in the lake. DR used both epibenthic and pelagic habitat, whereas two sparsely rakered whitefish forms dwelled (LSR and SSR) only in epibenthic habitat: LSR in littoral and SSR in profundal areas. Sparsely rakered whitefish forms had minor importance in predator diet.     

Resource partitioning and spatial segregation in native and stocked brown trout, Salmo trutta L., and Arctic charr, Salvelinus alpinus (L.), in a hydroelectric reservoir

Abstract. Habitat use, food and spatial segregation in native and stocked brown trout, Salmo trutta L., and Arctic charr, Salvelinus alpinus (L.), were studied during summer 1989 and 1990 in the hydroelectric reservoir Lake Tunhovdfjorden. There was no difference in habitat use and feeding habits between wild and stocked brown trout. In epibenthic areas brown trout lived chiefly down to 2 Secchi disc units, whereas Arctic charr were most abundant between 1 and 4 Secchi disc units. In pelagic areas the catches were low for both species, and they were chiefly confined to surface waters down to 1 Secchi disc unit. The food segregation between brown trout and Arctic charr was almost complete. Both pelagic and epibenthic Arctic charr fed mainly on cladocerans ( Bosmina longispina and Daphnia galeata ), whereas surface insects of terrestrial origin and Arctic charr were the dominant food items for brown trout. Pelagic Arctic charr were significantly older, larger and more homogeneous in size than epibenthic charr. During calm weather schools of Arctic charr were observed cruising with the dorsal fin above the surface.     

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.     

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.     

Recaptures and resource use of native and non-native brown trout, Salmo trutta L., released in a Norwegian lake

Abstract Rate of recapture (gill netting), habitat use, and diet of three strains of stocked brown trout, Salmo trutta L., were compared with resident brown trout in a Norwegian lake. The strains originated from an alpine lake, from a boreal lake, and from the native brown trout population in the lake. Overall recapture rate was 5–8% for all strains. The low recapture rate could be due to the relatively small size at stocking; mean fish length varied between 13.1 and 14.5 cm with strain and stocking method. Two years after release, the frequency of the different strains decreased from about 12% in the first year to stabilize at about 1%. The alpine strain showed the highest overall recapture rate, whereas the native strain was recaptured at an intermediate rate. The overall recapture rate of scatter-planted brown trout was higher than that of spot-planted brown trout. Immediately after being stocked, introduced fish ate less and had a less-varied diet than resident trout; however, stocked fish adopted a natural diet within the first summer. The distribution of trout between the pelagic and the upper epibenthic habitat was similar for both the resident and the stocked brown trout. Results indicate that the habitat use of stocked brown trout is adaptive and becomes similar to that of indigenous fish.     

Food resource partitioning of Arctic charr, Salvelinus alpinus (L.) and three-spined stickleback, Gasterosteus aculeatus L., in the littoral zone of lake Takvatn in northern Norway

The food resource partitioning of Arctic charr (Salvelinus alpinus L.) and three-spined stickleback (Gasterosteus aculeatus L.) were investigated in the littoral zone of lake Takvatn in northern Norway in the ice-free period June–November. Charr and sticklebacks had different feeding habits. Sticklebacks ate several small benthic prey items that were never eaten by charr, and the sticklebacks' diet were dominated by the benthic microcrustaceans Chydoridae and Ostracoda, chironomid larvae and stickleback eggs. Small charr (<17 cm) consumed a wide spectrum of chironomid pupae, terrestrial insects and zooplankton. Intermediate (17–20 cm) and small charr had quite similar feeding habits, while large charr (>20 cm) frequently ate both benthos, pelagic and terrestrial food. The diet overlap between small charr and sticklebacks was never larger than 0.6 (Schoener's index). The segregation in feeding habits indicates that small charr and sticklebacks are segregated in microhabitat when they are both in the littoral zone.     

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.     

The effect of lake morphometry on thermal habitat use and growth in Arctic charr populations: implications for understanding climate‐change impacts

Oxygen stable isotope temperature reconstruction methods were used to estimate mean experienced summer temperatures from growth zones within individual Arctic charr otoliths sampled from lakes with contrasting morphologies but proximate locations. For either lake, otolith‐estimated temperatures were not significantly related to back‐calculated growth. Fish in the smaller lake evidenced an increase in growth with age related to increasing use of cooler thermal habitats, with the use of thermal habitat possibly governed by predation risks. No relationships between age, growth or temperature were observed in the larger lake. Significant negative effects on back‐calculated growth were observed due to increasing air temperatures in the smaller and shallower lake, possibly owing to warmer surface and littoral waters and a limited amount of preferred cool‐water habitat. A similar relationship was not observed in the larger and deeper lake and indicated that resident Arctic charr were not as vulnerable to the impacts of temperature warming, possibly because of better behavioural thermoregulation opportunities in the cooler, deeper lake. Results provide evidence for differing climate‐influenced growth outcomes among proximately located populations, with outcomes likely to depend on the differences among habitats, including lake size and morphometry which may act to influence fish densities in available preferred thermal habitats.     

Summer habitat use and feeding of juvenile Arctic charr,Salvelinus alpinus,in the Canadian High Arctic

Abstract – Owing to limited knowledge of the habitat use and diet of juvenile Arctic charr from the High Arctic, particularly young‐of‐the‐year (YOY), we assembled data obtained from samples taken in and around Lake Hazen, Nunavut, Canada, to assess juvenile habitat use and feeding. Juvenile charr demonstrated a preference for stream environments, particularly those fed by warm upstream ponds. Charr occupying both stream and nearshore lake habitats were found to feed similarly, with chironomids occurring most frequently in diets. Some older stream‐dwelling charr preyed on smaller, younger Arctic charr. Preferred stream occupancy is likely mediated by physical barriers created mainly by water velocity, and by distance from the lake, lake‐ice dynamics, low water depth, and turbidity. Water velocities resulted in stream habitat segregation by size, with YOY mainly found in low‐velocity pools and back eddies adjacent to stream banks, but not in water velocities >0.1 m·s^?1. Greatest charr densities in streams were found in small, shallow, slow‐flowing side channels, which are highly susceptible to drought. Under predicted climate change scenarios, streams fed by small ponds will be susceptible to intermittent flow conditions, which could result in increased competition among juvenile charr for the remaining stream habitats. In addition, glacier‐fed streams are likely to experience increased flow conditions that will exacerbate physical barriers created by water velocity and further reduce the availability of preferred stream habitat.     

Trophic interactions between introduced lake trout (Salvelinus namaycush) and native Arctic charr (S. alpinus) in a large Fennoscandian subarctic lake

Introduced fishes may have major impacts on community structure and ecosystem function due to competitive and predatory interactions with native species. For example, introduced lake trout (Salvelinus namaycush) has been shown to replace native salmonids and induce major trophic cascades in some North American lakes, but few studies have investigated trophic interactions between lake trout and closely related native Arctic charr (S. alpinus) outside the natural distribution of the former species. We used stomach content and stable isotope analyses to investigate trophic interactions between introduced lake trout and native Arctic charr in large subarctic Lake Inarijärvi in northern Finland. Both salmonids had predominantly piscivorous diets at >280 mm total length and were mainly caught from the deep profundal zone. However, lake trout had a more generalist diet and showed higher reliance on littoral prey fish than Arctic charr, whose diet consisted mainly of pelagic planktivorous coregonids. According to length at age and condition data, lake trout showed slightly faster growth but lower condition than Arctic charr. The results indicate that introduced lake trout may to some extent compete with and prey upon native Arctic charr, but currently have only a minor if any impact on native fishes and food web structure in Inarijärvi. Future monitoring is essential to observe potential changes in trophic interactions between lake trout and Arctic charr in Inarijärvi, as well as in other European lakes where the two salmonids currently coexist.     

Thermal habitat use and juvenile growth of Svalbard Arctic charr: evidence from otolith stable oxygen isotope analyses

Abstract – Stable oxygen isotopes (δ¹⁸O) derived from otoliths were used to estimate mean annual water temperatures experienced by individual Svalbard Arctic charr, Salvelinus alpinus (L.), during their first four growth seasons. The analysed Arctic charr experienced a high variety of temperatures, indicating the use of different thermal habitats. A higher proportion of the juveniles experienced warmer temperatures during their first summer compared with later summers, suggesting the selective use of the shallowest littoral areas of the lake. Although the estimated temperatures were consistent with water temperatures found in High Arctic rivers and lakes during summer, they did not represent the annual variation in air temperature registered over the 20 years of otolith measurement. Furthermore, summer otolith increment width did not correlate with the experienced temperature. However, after the second year, otolith increment width was highly dependent on increment width during the previous summer. This study estimated mean summer water temperatures experienced by individual Arctic charr during the first four growth seasons providing additional evidence that stable oxygen isotope analysis can be used to provide insight into the thermal habitat use by juvenile Arctic charr.     

Resource use of native and stocked brown trout Salmo trutta L., in a subarctic lake

Habitat use, food composition and growth of stocked and native brown trout, Salmo trutta L., were studied in the subarctic Lake Muddusjärvi in northern Finland. Stocked brown trout and native brown trout preferred littoral and pelagic areas. Trout were stocked in October. In June stocked trout fed primarily on invertebrates while native fish were piscivorous. From July onwards the composition of the diet of both stocked and native trout was similar and consisted almost entirely of small‐sized whitefish. Brown trout were already piscivorous at a length of about 20 cm. The mean length of prey consumed was about 12 cm. Mean length‐at‐age was similar from the second year in the lake despite of the larger size of stocked fish during the first year in the lake.     

Changes in the population density of pelagic salmonids in relation to changes in lake enrichment in Windermere (northwest England)

From July 1989 to December 1994, an echo sounder provided monthly estimates, usually for both day and night, of pelagic salmonid densities in the North and South Basins of Windermere, the largest natural lake in England. Sampling was along contiguous transects, three in the North Basin and five in the South Basin. Records for Arctic charr (Salvelinus alpinus) could not be separated from those for brown trout (Salmo trutta), but previous sampling by gill-nets and anglers showed that charr formed over 90% of this mixed population in the North Basin and about 60–75% in the South Basin. Associated with the increasing eutrophication of the lake, there has been a decline in anglers' catches of charr and, since 1984, an increase in brown trout taken in the pelagic zone of the South Basin. The echo-sounder data showed that pelagic salmonid density in the North Basin was about two to five times that in the more eutrophic South Basin in 1989, 1990 and 1991. Since the start, in April 1992, of the reduction of phosphorus discharged from sewage works, this ratio has decreased, especially at night when the highest densities were recorded. This improvement was chiefly due to a significant (P<0.001) increase in the density of small fish (length <20 cm), in both the upper (depth <20 m) and deeper (depth >20 m) water layers. Although a similar improvement has still to be shown in the upper water layer by larger fish above the size limit for removal by angling (20 cm), there has been a significant increase (P<0.01) in the density of these fish in the deeper water layer of the South Basin. The increased density of small fish suggests that the stock available to charr anglers (fish >20 cm at water depths <20 m) should increase in the next few years, especially in the South Basin. It is therefore important to continue the monitoring program and thus ensure that there is advance warning of any marked changes in charr stocks.     

Trophic ecology of brown trout (Salmo trutta L.) in subarctic lakes

In subarctic lake systems, fish species like brown trout are often important predators, and their niche performance is a key characteristic for understanding trophic interactions and food web functioning at upper trophic levels. Here, we studied summer habitat use and stomach contents of brown trout under both allopatric and sympatric conditions in six subarctic lakes to reveal its trophic role, and population‐ and individual‐level niche plasticity. In allopatry, brown trout mainly used the littoral habitat, but also less commonly used the pelagic zone. In sympatry with stickleback, there was always a considerable habitat overlap between the two species. In contrast, sympatric populations of brown trout and Arctic charr generally revealed a distinct habitat segregation. In the sympatric systems, in general, there was a distinct resource partitioning between the trout and charr, whereas the observed diet overlap between trout and stickleback was much larger. Trout modified their individual dietary specialisation between the littoral and pelagic zone, always being lower in the pelagic. Piscivorous behaviour of trout was only found in sympatric systems, possibly contributing to a competitive advantage of trout over charr and stickleback. Hence, the trophic level of trout was strongly related to the fish community composition, with a higher trophic level in sympatric systems where piscivorous behaviour was frequent. These changes in the trophic level of trout linked with the observed food resource partitioning might be an important mechanism in the ecosystem functioning of subarctic lakes to allow coexistence among sympatric‐living fish species.     

Assessing the food web impacts of an anadromous Arctic charr introduction to a sub‐Arctic watershed using stable isotopes

Anadromous Arctic charr, Salvelinus alpinus (L.), was introduced to a sub‐Arctic river–lake system near the village of Kujjuuaq, Nunavik, and the stable isotope values and diets of key resident fish species were used to assess changes in feeding patterns. Stable isotope values for most species did not differ significantly between the pre‐ and post‐introduction periods, with observed shifts being within the bounds of expected natural variation. Lake chub, Couesius plumbeus (Agassiz), were the single species to show a difference between study periods, with a small but significant increase in δ¹⁵N. No significant post‐introduction changes were seen in lake trout, Salvelinus namaycush (Walbaum), omnivory or in any of the assessed quantitative food web metrics. Gut contents of major fish species similarly showed significant temporal overlap between the pre‐ and post‐introduction periods, and there was no significant change in species' weight–length relationships. The minor ecological impact was interpreted in relation to the availability of open niches exploitable by ecological generalists such as Arctic charr. The explanation accords with the known habitat and feeding flexibility of Arctic charr and the ecological immaturity of sub‐Arctic lakes known to have driven adaptive variation among Arctic charr. Findings suggest that anadromous Arctic charr may be introduced at moderate densities to other sub‐Arctic watersheds without major negative food web consequences for other resident fish species.     

Dietary plasticity of Arctic charr (Salvelinus alpinus) facilitates coexistence with competitively superior European whitefish (Coregonus lavaretus)

Abstract – Habitat use and diet of Arctic charr (Salvelinus alpinus) coexisting with European whitefish (Coregonus lavaretus) and grayling (Thymallus thymallus) were studied in one deep and two relatively shallow subarctic lakes in northern Norway. Stomach content and stable isotope analyses revealed clear and temporally stable resource partitioning between the species in all three lakes. Arctic charr had a wide and flexible trophic niche and was the only piscivorous species. In contrast, whitefish and grayling had remarkably stable planktivorous and benthivorous niches, respectively. In the deepest lake, Arctic charr together with grayling mainly utilised littoral benthos, while piscivory was more prevalent in Arctic charr in the two shallower lakes. In one of the shallow lakes, whitefish was apparently relegated to the inferior profundal niche because of dominance of the littoral by grayling. Our results suggest that Arctic charr may not necessarily need an extensive profundal zone as a refuge, but can coexist with whitefish if a third competing fish species like grayling occurs in the littoral habitat or if profitable small prey fish are available. The study demonstrates that strong dietary plasticity of Arctic charr is instrumental in the observed coexistence with the commonly competitively superior whitefish.     

Habitat and food choice of Arctic charr in Linnévatn on Spitsbergen, Svalbard: the first year-round investigation in a High Arctic lake

Abstract –  Habitat and diet of Arctic charr Salvelinus alpinus (L.) were studied by monthly sampling from late autumn to early summer in Linnévatn, Svalbard (78°3'N, 13°50'E). This is the first year-round study of a population of charr in the High Arctic, with samples being taken every 5–7 weeks. The ice cover lasted for more than 9 months, from mid-October to late July, with the greatest thickness in mid-May. Although most charr occupied the littoral zone during winter, the highest densities in April and October were found in the deeper areas (20 m) of the lake. The fish fed at all times of the year, but the number of stomachs with food and the stomach-filling indices were lowest during the darkest part of the season. The diet of smaller charr (<15 cm) varied strongly with season, showing a dominance of zooplankton in late autumn and chironomids in winter (larvae) and summer (pupae). The food choice was in accordance with the density of food items available. Larger fish (≥15 cm) were mostly cannibalistic during the entire year.     

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

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

The marine temperature and depth preferences of Arctic charr (Salvelinus alpinus) and sea trout (Salmo trutta), as recorded by data storage tags

Eleven Arctic charr (Salvelinus alpinus) (370–512 mm) and eight sea trout (Salmo trutta) (370–585 mm in length) were tagged externally or internally with depth‐ and temperature‐measuring data‐storage tags (DST) before they were released into the sea in the Alta Fjord in north Norway in June 2002. All sea trout were recaptured after they spent 1–40 days at sea, while all Arctic charr were recaptured after 0.5–33 days at sea. On average, trout preferred water about 0.6 m deeper and 1.3°C warmer than Arctic charr. Arctic charr spent >50% of their time between 0 and 1 m depth, while trout spent >50% of their time between 1 and 2 m depth. Both species spent >90% of their time in water no deeper than 3 m from the water surface. However, sea trout dove more frequently and to greater depths (max. 28 m) than Arctic charr (max. 16 m), and these deep dives were most frequently performed at the end of the sea migration. Arctic charr demonstrated a diel diving pattern, staying on average about 0.5 m deeper between 08:00 hours and about 15:00 hours than during the rest of the 24 h, even though there was continuous daylight during the experiments. When comparing data obtained from the DSTs with temperature measurements within the fjord system, the two species were observed to select different feeding areas during their sea migration, the sea trout choosing the inner and warmer parts of the fjord, in contrast to the Arctic charr that preferred the outer, colder parts of the fjord. The observed differences in migration behaviour between the two species are discussed in relation to species preferences for prey and habitat selection, and their optimal temperatures for growth.