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     

Ecological characteristics of ninespine stickleback Pungitius pungitius from southern Baffin Island,Canada

Abstract – Stable δ¹³C and δ¹⁵N isotopes, diet and parasites demonstrated that the prey consumed by ninespine stickleback Pungitius pungitius in a small lake on Baffin Island changed during the summer and also revealed intraspecific variation in their ecological niche. In July, there were differences in the diets of male and female ninespine stickleback as indicated by the stable isotopes, differences corroborated by the data on diet composition and the parasite fauna. Differences suggested that the sexes occupied different habitats during spawning. During July, females utilise the shallower littoral areas consuming zooplankton and benthic organisms, while males occupy deeper areas of the littoral zone feeding mainly on pelagic zooplankton. Parasite data support these observations as males had higher infections of copepod‐transmitted parasites than females. There appeared to be no segregation of resources between males and females in late August, although the diet of both male and female ninespine stickleback shifted towards more benthic organisms, compared with July. Differences in δ¹³C isotope, diet composition and infections of co‐occurring parasites demonstrated that sympatric ninespine stickleback and Arctic char Salvelinus alpinus captured in the littoral zone occupied separate niches. Ninespine stickleback preyed mainly on zooplankton and chironomids, while Arctic char consumed a greater variety of prey items, including zooplankton and larger‐sized prey such as insects and ninespine stickleback. The multifaceted approach improved our understanding of the trophic ecology of ninespine stickleback in southern Baffin Island and quantified resource use and dietary overlap with Arctic char.     

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.     

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.     

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.     

Lake-use by juvenile Atlantic salmon (Salmo salar L.) and other salmonids in northern Norway

Abstract– The utilization of lakes, and inlet and outlet streams by juvenile Atlantic salmon ( Salmo salar L.), brown trout ( Salmo trutta L.) and Arctic charr ( Salvelinus alpinus (L.), were investigated in 16 watercourses northern Norway, all known to inhabit salmon stocks. In lakes, fish were caught by small mesh size gill nets, while in rivers fish were caught electrofishing. In the shallow littoral (0-3 m depth) there were juvenile salmon in 15 of 19 investigated lakes, juvenile trout in 17 and juvenile charr in seven. Trout dominated significantly in numbers in the shallow littoral of seven lakes, while salmon and charr dominated in three lakes each. When trout and salmon were frequent in the shallow littoral, charr was usually not present in this habitat, but were found in the profundal zone in most of the lakes. Atlantic salmon parr utilized both shallow and deep lakes, and used both stones and macrophytic vegetation as shelter. The utilization of lakes by salmon parr seemed to be closely related to utilization of small inlet streams for spawning. In most inlet and outlet streams salmon dominated over trout in numbers, while charr were absent. This is the first documentation of lake-use by naturally occurring salmon parr in Scandinavia.     

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.     

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.     

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.     

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.     

Prey selection and growth in 0+ Eurasian perch Perca fluviatilis L. in littoral zones of seven temperate lakes

We studied the relationships between the planktonic food base and feeding patterns of juvenile mid-summer/early autumn Eurasian perch Perca fluviatilis L., a common predatory freshwater fish in large parts of Europe and Asia. The feeding of 0+ perch was studied during summer and autumn in littoral habitats of seven lakes with different environmental conditions – four Latvian (Auciema, Riebinu, Vārzgūnes, Laukezers) and three Estonian (Kaiavere, Prossa and Akste) lakes. Simultaneously, the abundance, biomass and structure of zooplankton communities were examined. We focused on the littoral areas because many studies in lakes suggest that littoral habitats are particularly important for 0+ fish growth and survival. We were interested in the question: can the diet and growth of 0+ perch be explained by zooplankton community structure? We also presumed that if the amount of zooplankton is low, more benthic invertebrates will be consumed by 0+ perch. Opposite to expectations, we found that zooplankton always counted for over 90% of diet biomass in perch. There were also clear correlations between the zooplankton biomass in a given lake, the zooplankton biomass in 0+ perch stomachs, and the fish growth rate. The study also suggested that nutrient enrichment can positively impact the 0+ perch feeding conditions in lakes.     

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.     

Gill development in sympatric morphs of Arctic charr from Loch Awe,Scotland: A hidden physiological cost of macrobenthos feeding?

The development of the respiratory surfaces was compared in two sympatric, lacustrine morphs of Arctic charr. A macrobenthic invertebrate feeding specialist, that forages in the littoral benthic zone, had a gill cavity that was 54% larger in volume and had 31% greater respiratory surface area than that of a zooplankton feeding morph that forages in the pelagic zone. The large respiratory surface area in the benthic‐feeding form was the result of longer gill arches, more and longer gill filaments and more numerous secondary lamellae. The difference in gill cavity volume and filament length appears to be the result of a larger head, but not body size, in the benthic‐feeding form. This suggests that differences in these characteristics may have arisen as a by‐product of the expression of larger head size commonly described in macrobenthos foraging specialist charr. The other differences, particularly the more numerous secondary lamellae and the length of the gill arches, were not the result of head size differences between morphs, and thus, these are most likely an adaptation to greater respiratory requirements. Benthic‐feeding fish may have a greater respiratory capacity to allow them to forage in areas with lower levels of dissolved oxygen and/or engage in a more active lifestyle compared to the pelagic‐feeding form. In any event, the strikingly larger respiratory surface is likely to impose an additional ionoregulatory stress on the benthic‐feeding and thus may represent a hidden physiological cost of specialisation for foraging on benthic prey.     

Feeding ecology and habitat of the threespine stickleback, Gasterosteus aculeatus microcephalus, in a remnant population of northwestern Baja California, México

Abstract – The feeding ecology and habitat of the threespine stickleback ( Gasterosteus aculeatus microcephalus Girard, 1854) was studied from November 1996 to May 1997 in a remnant population of northwestern Baja California, México. The analysis of the stomach content of 179 individuals (25 to 56 mm standard length [SL]) showed a diet dominated by cyclopoid copepods (43.8%) and chironomid larvae (39.1%). Diet in relation to size and sex of the fish was dominated by copepods in autumn and winter and by chironomid larvae during spring. Diet overlap (Schoener's index) was significant (≥60%) between fish size-classes in January, March and April and between sexes for most sampling months. The average size of prey consumed was independent of fish mouth size. The feeding strategy of the threespine stickleback shifted from opportunist in winter to specialist in spring.     

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.     

Growth and diet of anadromous Arctic charr after their return to freshwater

Abstract – The post smolt of anadromous Arctic charr ( Salvelinus alpinus ) from Lake Storvatn, northern Norway (70°39'48"), continued to feed and grow after their return to freshwater in late summer, but in late autumn (October) both their growth rate and feeding intensity ceased along with the drop in water temperature. In contrast, adult, sexually mature anadromous charr did not appear to feed during their freshwater residency. The stomach fullness of post smolts were similar to that of resident Arctic charr during early autumn, and their diets were similar, being dominated by pelagic prey such as surface insects, Holopedium sp., Bosmina sp. and chironomid pupae. Later in the autumn, the resident charr also included some more benthic prey, in particular gastropods, whereas the post smolts continued to feed predominantly on zooplankton and surface insects, but with a lower stomach fullness than the resident fish. This suggests that after arrival in freshwater, the post smolts continue their pelagic feeding behaviour displayed during the sea-migration.     

Feed intake,growth and osmoregulation in Arctic charr,Salvelinus alpinus (L.), transferred from freshwater to saltwater at 8°C during summer and winter

The purpose of the current study was to examine seasonal changes in seawater tolerance and growth performance of anadromous Arctic charr (Salvelinus alpinus L.) held at the same temperature (8°C) during winter and summer. Charr (20–27 cm), previously reared in freshwater under natural photoperiod, were transferred either directly (DT) from freshwater to seawater (35 ppt), from freshwater to brackish water (20 ppt), or were gradually adapted (GT) to seawater over a period of 10 days. Control fish were held in freshwater. Feed intake and osmoregulatory ability were then monitored on three occasions during the following 59 days. Two experiments were carried out, one during winter (December–January) and the other during summer (June–July). In both experiments fish mortality was low. Plasma osmolalities recorded in fish transferred to seawater were within normal ranges, but osmolalities on day 10, were significantly lower in summer (313 mOsm/kg (DT), 328 mOsm/kg (GT)) than in winter (323 mOsm/kg (DT), 352 mOsm/kg (GT)). In winter, feed intake and growth rates were high in fish kept in fresh and brackish water, but charr transferred directly to seawater ate little and lost weight. Fish that were gradually adapted to seawater occupied an intermediate position. During summer the observed differences in feed intake were small and all fish had relatively high growth rates. These results suggest that Arctic charr display seasonal changes in feed intake and growth performance that parallel seasonal changes in hypoosmoregulatory capacity. The ability to survive and hypoosmoregulate in full strength seawater does not, however, seem to be a particularly good indicator of successful seawater adaptation with respect to the ability to display high rates of feed intake and growth. During winter, a gradual transfer to seawater appeared to lead to improved feeding and growth compared to direct transfer.     

Changes in demand feeding behaviour in Arctic charr, Salvelinus alpinus L., caused by differences in dietary energy content and reward level

The objective of this study was to determine whether Arctic charr, Salvelinus alpinus L., are able to adjust their demand feeding behaviour in accordance to differences in dietary energy content (experiment 1) and reward level (amount of food received in response to one trigger actuation) (experiment 2). Fish (initial size 215 g in experiment 1 and 88 g in experiment 2) were reared in 0.8-m³ indoor tanks at commercial stocking densities and fed using demand feeders for 57 and 180 days, respectively. Demand feeding activity did not differ significantly between groups of charr fed diets with a low (19.8 MJ kg^-1) gross energy content and those given high-energy (22.0 MJ kg^-1) feed. As a result, fish offered the high-energy diets grew significantly faster. The results show that charr held under culture conditions are unable to adjust their demand feeding activity based on the energy content of the food. On the contrary, Arctic charr are able to adjust their demand feeding activity to either low (0.33 g), medium (0.87 g) or high (1.52 g) rewards, and thereby, regulate their food supply to fit their needs. However, it took about 90 days before charr in the low-reward treatment released a daily food ration as high as that released in the high-reward groups. Consequently, there was a significant positive relationship between the size of the reward and final weights. To avoid any depression of initial growth rates, the optimal size of the reward should be 0.1 g per kg fish and trigger actuation.     

Trophic pathways supporting Arctic grayling in a small stream on the Arctic Coastal Plain,Alaska

Beaded streams are prominent across the Arctic Coastal Plain (ACP) of Alaska, yet prey flow and food web dynamics supporting fish inhabiting these streams are poorly understood. Arctic grayling (Thymallus arcticus) are a widely distributed upper‐level consumer on the ACP and migrate into beaded streams to forage during the short 3‐month open‐water season. We investigated energy pathways and key prey resources that support grayling in a representative beaded stream, Crea Creek. We measured terrestrial invertebrates entering the stream from predominant riparian vegetation types, prey types supporting a range of fish size classes, and how riparian plants and fish size influenced foraging habits. We found that riparian plants influenced the quantity of terrestrial invertebrates entering Crea Creek; however, these differences were not reflected in fish diets. Prey type and size ingested varied with grayling size and season. Small grayling (<15 cm fork length (FL)) consumed mostly aquatic invertebrates early in the summer, and terrestrial invertebrates later in summer, while larger fish (>15 cm FL) foraged most heavily on ninespine stickleback (Pungitius pungitius) throughout the summer, indicating that grayling can be insectivorous and piscivorous, depending on size. These findings underscore the potential importance of small streams in Arctic ecosystems as key summer foraging habitats for fish. Understanding trophic pathways supporting stream fishes in these systems will help interpret whether and how petroleum development and climate change may affect energy flow and stream productivity, terrestrial–aquatic linkages and fishes in Arctic ecosystems.     

Feeding strategy of the non‐native rainbow trout,Oncorhynchus mykiss,in low‐order Patagonian streams

The dietary composition and feeding strategy of rainbow trout, Oncorhynchus mykiss (Walbaum), in two low‐order Patagonian streams were studied. Benthic macroinvertebrate availability was estimated in both riffles and pools. Fish stomach contents were examined to determine prey richness and diversity, prey electivity, food‐niche width, and the feeding strategy employed by trout throughout the year. Availability of benthos varied seasonally with Ephemeroptera, Trichoptera, Plecoptera and Diptera species dominating. Rainbow trout diet was composed mainly of benthic macroinvertebrates, followed by terrestrial insects, fish, algae and plants. Different trout size classes segregated the use of food resources to reduce predation pressure. Elected prey included organisms displaying no to high mobility. A high feeding plasticity allows trout to buffer changes in food availability by switching from a specialised to a generalised feeding behaviour. Consequently, trout may exploit abundant but temporary food resources opportunistically, which would explain their marked expansion in Patagonian environments.