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

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

Lipid‐rich zooplankton subsidise the winter diet of benthivorous Arctic charr (Salvelinus alpinus) in a subarctic lake

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Habitat, diet and food assimilation of Arctic charr under the winter ice in two subarctic lakes

The Arctic charr Salvelinus alpinus populations of the subarctic lakes Takvatn and Fjellfrøsvatn, north Norway, concentrated in the littoral zones (0–15 m) of the lakes during the entire winter (December to May) despite very low temperatures (0·2 and 0·7° C). High prey availability, low predation and competition and comparatively better light under snow and ice in shallow compared with deep water are probable reasons. At ice break in June, all Arctic charr moved to the profundal zone for a brief period, probably in response to the sudden light increase and a profundal resource peak of chironomid pupae. In the summer, the Arctic charr are found in the pelagic, profundal and littoral zones of the lakes. These populations therefore perform regular habitat shifts between the littoral zone in the winter, the profundal zone at ice break and the whole lake in the summer and autumn. The fish fed continuously during winter despite the cold water and the poor light. Amphipods and chironomid larvae dominated the diet. Catch per unit effort, numbers of stomachs with food and food intake rates varied with the subarctic light cycle but were lowest after the winter solstice. The winter assimilation of energy was about equal to the standard metabolism in Takvatn but was higher in Fjellfrøsvatn. The assimilation increased in both lakes under the spring ice in May. The habitat choice, diet and energy assimilation indicate that the Arctic charr is well adapted to the extreme winter conditions of subarctic lakes.     

Winter ecology of Arctic charr (<Emphasis Type="Italic">Salvelinus alpinus</Emphasis>) and brown trout (<Emphasis Type="Italic">Salmo trutta</Emphasis>) in a subarctic lake,Norway

We studied habitat choice, diet, food consumption and somatic growth of Arctic charr (Salvelinus alpinus) and brown trout (Salmo trutta) during the ice-covered winter period of a subarctic lake in northern Norway. Both Arctic charr and brown trout predominantly used the littoral zone during winter time. Despite very cold winter conditions (water temperature <1°C) and poor light conditions, both fish species fed continuously during the ice-covered period, although at a much lower rate than during the summer season. No somatic growth could be detected during the ice-covered winter period and the condition factor of both species significantly declined, suggesting that the winter feeding rates were similar to or below the maintenance requirements. Also, the species richness and diversity of ingested prey largely decreased from summer to winter for both fish species. The winter diet of Arctic charr <20 cm was dominated by benthic insect larvae, chironomids in particular, and Gammarus lacustris, but zooplankton was also important in December. G. lacustris was the dominant prey of charr >20 cm. The winter diet of brown trout <20 cm was dominated by insect larvae, whereas large-sized trout mainly was piscivorous, feeding on juvenile Arctic charr. Piscivorous feeding behaviour of trout was in contrast rarely seen during the summer months when their encounter with potential fish prey was rare as the small-sized charr mainly inhabited the profundal. The study demonstrated large differences in the ecology and interactions of Arctic charr and brown trout between the winter and summer seasons.     

Selective predation on zooplankton by pelagic Arctic charr, Salvelinus alpinus, in six subarctic lakes

Selective predation by planktivore fish appears to be an important regulatory factor of zooplankton communities, potentially causing large changes in species composition and size distributions within populations. In this study, prey preferences and size-selective predation on zooplankton by Arctic charr were examined in six subarctic lakes with Arctic charr as the dominant pelagic fish species. Most of the lakes had a zooplankton community dominated by copepods (Cyclops scutifer and Eudiaptomus graciloides), but the pelagic charr evidently selected cladoceran species (Bythotrephes longimanus, Daphnia sp. and Bosmina sp.), likely because the copepods have a higher mobility and evasiveness than the cladocerans. Furthermore, a strong size selection was also revealed for both Bosmina sp. and Daphnia sp., as individual prey from Arctic charr stomachs were exclusively larger than individuals sampled in the environment. Additionally, visibility due to size, morphology and pigmentation (egg-carrying females) was also a major factor for the selection of zooplankton prey. In conclusion, Arctic charr was found to be highly selective on zooplankton both in respect to species composition and individual size of Bosmina sp. and Daphnia sp.     

Density,length distribution,and diet of age-0 arctic charrSalvelinus alpinus in the surf zone of Thingvallavatn,Iceland

Synopsis Population densities of age-0 arctic chaff in the surf zone averaged 1.83 and 4.70 fish m^-2 in August 1984 and June 1985, respectively. Length variation of the littoral fish was low in early summer, increasing in late summer and autumn. Newly emerged charr, ∼ 20 mm long, appeared in the stony shallow water zone during both May and June. From length variation and variation in mouth position of the young charr, it is concluded that at least two of the four chaff morphs in the lake are present in the surf zone during spring and summer. In August, some of the larger age-0 charr had moved out from the surf zone, into the pelagic and the deeper epibenthic waters. The food of young littoral charr was dominated by large chironomid larvae (instar 3 and 4) and pupae Contribution from the Thingvallavatn project     

Habitat use and life history of brown trout (Salmo trutta) and Arctic charr (Salvelinus alpinus) in some low acidity lakes in central Norway

Habitat utilization and the life history of browntrout Salmo trutta and Arctic charr Salvelinus alpinus were investigated in fivesympatric populations and five allopatric brown troutpopulations in Høylandet catchment, a atmosphaericlow deposition area in Mid Norway. There was asignificant inverse correlation in abundance ofepibenthic Arctic charr and brown trout in theselakes, indicating that the latter species is dominant.The largest numbers of sympatric brown trout andArctic charr were caught in epibenthic habitat. In twolakes, brown trout to some extent also occurredpelagically, while pelagic individuals of Arctic charrwere found in all five lakes. The main food items forboth epibenthic and pelagic brown trout wereterrestrial surface insects and chironomid pupae.Zooplankton was the primary food item for Arctic charrin both habitats. Although the age distribution wasvery different in the populations, neither speciesseem to suffer from recruitment failure. There was nosignificant difference in survival rates betweensympatric populations of brown trout and Arctic charr.We found a significant inverse correlation betweenepibenthic catches of brown trout and the mean weightof 4+ fish, the most abundant age group. However, ifusing weight data for three-year-old fish, no suchrelationship was found for Arctic charr. Brown troutand Arctic charr reached asymptotic lengths of197–364 mm and 259–321 mm, respectively. Both speciestypically reached sexual maturity at age 2–3, and nomaturation-induced mortality was evident. We concludethat fish populations in Høylandet lakes areregulated throughout their lifes by inter- andintraspecific competition.     

Habitat utilization by sympatric arctic charr Salvelinus alpinus L. and brown trout Salmo trutta L. in Lake Atnsjø, south-east Norway

SUMMARY. 1. Habitat utilization, as well as inter- and intraspecific relations of different size groups of arctic charr (Salvelinus alpinus (L.)) and brown trout (Salmo trutta L.) in Lake Atnsjø, south-east Norway, were investigated by analysing food and spatial niches from monthly benthic and pelagic gillnet catches during June-October 1985.
2. Small individuals (150–230 mm) of both arctic charr and brown trout occurred in shallow benthic habitats. However, they were spatially segregated as arctic charr dominated at depths of 5–15 m and brown trout at depths of 0–5 m.
3. Larger (>230 mm) arctic charr and brown trout coexisted in the pelagic zone. Both species occurred mainly in the uppermost 2-3 m of the pelagic, except in August, when arctic charr occurred at high densities throughout the 0–12 m depth interval. On this occasion, arctic charr were segregated in depth according to size, with significantly larger fish in the top 6 m. This was probably due to increased intraspecific competition for food.
4. The two species differed in food choice in both habitats, Arctic charr fed almost exclusively on zooplankton, whereas brown trout had a more variable diet, consisting of surface insects, zooplankton. aquatic insects and fish.
5. The data suggest that the uppermost pelagic was the more favourable habitat for both species. Large individuals having high social position occupied this habitat, whereas small individuals lived in benthic habitat where they were less vulnerable to agonistic behaviour from larger individuals and less exposed to predators. The more aggressive and dominant brown trout occupied the more rewarding part of the benthic habitat.     

Only the small survive: monitoring long-term changes in the zooplankton community of an Alpine lake after fish introduction

The zooplankton community of Alpine lake Seehornsee (1,779 m a.s.l.) was studied over a period of 13 years. In 1994, a typical high-altitude zooplankton community, consisting of two calanoid copepods (Mixodiaptomus laciniatus, Arctodiaptomus alpinus), one cladoceran (Daphnia rosea), and two rotifers (Keratella quadrata, Synchaeta pectinata) coexisted with infertile charr hybrids, which had been introduced in 1969 and again in 1974. When the aged fish were removed by intensive gill netting, they had fed predominantly on aquatic insects. After a fish-free period of 4 years, 2000 fertile juvenile Alpine charr (Salvelinus umbla) were stocked in 1998 and again in 1999. They preyed on benthic (chydorids, ostracods, cyclopoid copepods, chironomid larvae and pupae) and planktonic prey (diaptomid copepods, Daphnia). Between 2004 and 2006 charr successfully reproduced. Nine years after stocking of fertile charr, the two calanoids had virtually disappeared, and Daphnia rosea had notably declined in abundance. In concordance with the size efficiency hypothesis (Brooks and Dodson 1965), the newly appearing and smaller cladoceran Ceriodaphnia pulchella, together with the two resident, and two emerging species of rotifers (Polyarthra luminosa, Gastropus stylifer) dominated the zooplankton community.     

Comparison of growth, diet and food consumption of sea-run and lake-dwelling Arctic charr

Sea-run post-smolt Arctic charr Salvelinus alpinus , (15–26 cm) from Storvatn, northern Norway (70°39'48"N) had significantly higher average specific growth rates in two years (1·64 and 1·66) than the corresponding lake-dwelling charr (0·53 and 1·20). The post-smolts displayed fast compensatory growth in the first 2–3 weeks of their sea residency, but then almost stopped growing prior to their return to fresh water. Lake-dwelling charr grew more evenly during the same time period. Thus, the anadromous charr may return to the lake after only 5–6 weeks in the sea, because the potential to maintain a high growth rate in the sea is reduced. The marine diet consisted mainly of the two crustacean plankton species Calanus finmarchicus , and Thysanoëssa , sp. (88%), and less of fish (6%), insects (4%) and benthos (2%). The diet of lake-dwelling charr consisted mainly of insects (58%, mostly chironomid pupae) and zoobenthos (29%), and less of zooplankton (13%) during the same time period. Although post-smolts had the highest growth rates, they had significantly lower food consumption rates and higher frequencies of empty stomachs than the corresponding lake-dwelling fish. Possible explanations for this paradox are discussed in relation to stomach evacuation rates, water temperature, feeding behaviour and the energy content of the food in the two environments.     

Takvatn Through 20 Years: Long-term Effects of an Experimental Mass Removal of Arctic Charr, Salvelinus Alpinus, From a Subarctic Lake

Between 1984 and 1989, the experimental removal of 31 tons (666000 fish) of stunted Arctic charr, Salvelinus alpinus, from Takvatn in northern Norway, had strong effects on the populations of Arctic charr, brown trout, Salmo trutta, and three-spined sticklebacks, Gasterosteus aculeatus. The littoral catch per unit effort (CPUE) of charr had decreased by 90% in 1990 and then increased to about 50% of the initial level by 1994 while the pelagic CPUE had decreased to zero. Growth in both charr and trout greatly improved when the charr density had decreased, and large fish of both species appeared in the catches. These large fish became predators on small charr in the littoral zone. The incidence of trout increased from below 1% to 15% from 1988 to 1999 after a brief peak at 30% in 1992 and 1993. The charr population attained a bimodal size distribution and did not return to the stunted state during the 10 years following the intensive fishing period. The mass removal experiment showed that it is possible to change the structure of a charr population by intensive fishing. Predation on small charr from cannibals and large trout was probably essential for maintaining the new population structure. An increase in the growth of young charr from 1995 to 1997 was related to a high consumption of Daphnia and Eurycercus. Rapid changes in the growth of charr followed the density fluctuations in sticklebacks, which show large annual variations in this system; the rapid changes in charr growth were probably caused by variations in the competition intensity for cladoceran prey between young charr and sticklebacks. Twenty years of data has provided important information, but even more time is needed to follow the long-term trends in northern lakes such as Takvatn.     

Trophic differentiation of the nosed charr <Emphasis Type="Italic">Salvelinus schmidti</Emphasis> Viktorovsky, 1978 in Lake Kronotskoe (Kamchatka)

The nosed charr Salvelinus schmidti that inhabits the littoral zone of Lake Kronotskoe is divided into two groups according to food preferences and parasite fauna. Fish of the first group (G) predominantly feed on gammarids and are characterized by a high infestation with Cystidicola farionis, Cyathocephalus truncatus, and Crepidostomum spр. Fish of the second group (A) do not consume gammarids but feed mostly on chironomid larvae and pupae and on mollusks; the predominant parasites are Phyllodistomum umblae and Proteocephalus longicollis. The significant difference in the abundance of C. farionis, which remains in fish at least for 2 years, is indicative of a long-term and persistent trophic diversification between the charr groups. Moreover, significantly higher growth rates, a larger body size, and a longer lifespan are observed for fish of group G. The feeding strategy of the nosed charr is maintained throughout the life and does not depend on the sex of fish and their distribution within the littoral zone of the lake.     

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

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

Lake size and fish diversity determine resource use and trophic position of a top predator in high‐latitude lakes

Prey preference of top predators and energy flow across habitat boundaries are of fundamental importance for structure and function of aquatic and terrestrial ecosystems, as they may have strong effects on production, species diversity, and food‐web stability. In lakes, littoral and pelagic food‐web compartments are typically coupled and controlled by generalist fish top predators. However, the extent and determinants of such coupling remains a topical area of ecological research and is largely unknown in oligotrophic high‐latitude lakes. We analyzed food‐web structure and resource use by a generalist top predator, the Arctic charr Salvelinus alpinus (L.), in 17 oligotrophic subarctic lakes covering a marked gradient in size (0.5–1084 km²) and fish species richness (2–13 species). We expected top predators to shift from littoral to pelagic energy sources with increasing lake size, as the availability of pelagic prey resources and the competition for littoral prey are both likely to be higher in large lakes with multispecies fish communities. We also expected top predators to occupy a higher trophic position in lakes with greater fish species richness due to potential substitution of intermediate consumers (prey fish) and increased piscivory by top predators. Based on stable carbon and nitrogen isotope analyses, the mean reliance of Arctic charr on littoral energy sources showed a significant negative relationship with lake surface area, whereas the mean trophic position of Arctic charr, reflecting the lake food‐chain length, increased with fish species richness. These results were supported by stomach contents data demonstrating a shift of Arctic charr from an invertebrate‐dominated diet to piscivory on pelagic fish. Our study highlights that, because they determine the main energy source (littoral vs. pelagic) and the trophic position of generalist top predators, ecosystem size and fish diversity are particularly important factors influencing function and structure of food webs in high‐latitude lakes.     

Diet, gastric evacuation rates and food consumption in a stunted population of Arctic charr, Salvelinus alpinus L., in Takvatn, northern Norway

Diet and food consumption of a stunted population of Arctic charr in Takvatn, northern Norway, was studied throughout the ice-free season. Chironomid pupae dominated the diet in June, July and August, while zooplankton and three-spined sticklebacks dominated in September and October. Estimates of gastric evacuation rates were obtained from laboratory experiments with wild-caught, acclimated charr, and these estimates were used for in situ estimation of food consumption from weights of stomach contents. The daily food consumption was greatest in July, after which it decreased markedly towards autumn. The estimated food intake rates were low, especially in September and October. The food supply appears to be restricted, which is probably the main reason why the charr in Takvaln exhibit stunted growth.     

Sticklebacks from Greenland

Greenland freshwater sticklebacks Gasterosteus aculeatus were low-plated. Length distribution of 1 + and 2+ fish peaked at 33 and 46 mm, in early July. Pre-reproductive fish including 1 + fish were infected with Schistocephalus solidus . The diet consisted of chironomid larvae and pupae. Infected individuals ate smaller prey than non-infected ones, on one occasion where competition for food was likely to occur. Infected fish had lower stomach fullness, and the parasite compressed the stomach in heavily infected fish. The other fish species was charr, and the stickleback seemed to occupy a wide range of habitats.     

Life history characteristics and distribution of Bythotrephes longimanus Leydig (Crustacea,Onychopoda) in the Biesbosch reservoirs

The diel and horizontal distributions of Bythotrephes longimanus Leydig, Leptodora kindti (Focke) and Anomopoda (cladocerans), as well as several life history parameters of each developmental stage of B. longimanus were studied in the non-stratified Biesbosch reservoirs. Anomopoda avoided the surface layer during the day only in the pelagic zone. In the surface layer near the shore, however, they were very abundant. Because densities in the pelagic zone during the day were very low, and high during the night, the population undoubtedly moved from the littoral zone or from the sediments to the pelagic. Accumulation of zooplankton at the littoral site is either the result of random dispersal or wind-induced movements. Bythotrephes' diel distribution pattern is possibly related to that of their prey, the anomopods. Leptodora mostly exhibited a normal diel vertical migration with a small amplitude. Wind most likely influenced the horizontal distribution of the zooplankton. Bythotrephes body length increased with higher temperatures, whereas spine length was constant throughout the year. Bythotrephes can already be fertile in the first developmental stage, indicating that a rapid adaptation to a favourable environment is possible. The number of parthenogenetically produced eggs per ovigerous female was higher at the start of the growing season and constant throughout the rest of the year. Relatively few resting eggs and males were found in autumn.     

Diets and food consumption rates of pelagic fish in Lake Malawi, Africa

• 1 Studies of diel feeding periodicity, rates of food consumption and diet composition of pelagic fish were undertaken to resolve the dispute regarding the existence of a vacant niche for a pelagic zooplanktivore in Lake Malawi.
• 2 Six species of zooplanktivore were abundant in the offshore pelagic zone, these were: Diplotaxodon limnothrissa, D.‘bigeye’, Rhamphochromis longiceps and Copadichromis quadrimaculatus (all haplochromine cichlids), Synodontis njassae (Mochokidae) and Engraulicypris sardella (Cyprinidae). The main piscivores were cichlids of the genus Rhamphochromis.
• 3 All cichlids were daytime feeders; some showed peaks in feeding activity at dawn and dusk that were related to vertical migration patterns of their prey. Synodontis njassae was able to feed throughout the day, but fed most actively at night.
• 4 Food consumption: biomass ratios (Q/B yr^-1) calculated from diel stomach contents data were 5.87–7.13 for D. limnothrissa, 12.79 for D.‘bigeye’, 4.20–24.7 for R. longiceps and 6.45 for S. njassae. Annual Q/B ratios for other species, which ranged from 4.74 to 9.28, were obtained from an empirical model relating food consumption to fish morphology. Published estimates were used for E. sardella and Opsaridium microcephalum (Cyprinidae). An estimate of total prey consumption by the pelagic fish population was obtained from Q/B ratios and fish biomass estimates from acoustic surveys covering the entire offshore pelagic zone of the lake (24 000 km²).
• 5 Diplotaxodon limnothrissa, R. longiceps and C. quadrimaculatus had a broad range of diets, with the ability to switch from crustacean zooplankton to larvae and pupae of Chaoborus edulis (Diptera) or larval and juvenile E. sardella. Diplotaxodon‘bigeye’ and S. njassae fed almost exclusively on fourth-instar larvae and pupae of C. edulis. Engraulicypris sardella fed mainly on crustacean zooplankton. The main prey of the large piscivorous Rhamphochromis species were Diplotaxodon spp. and E. sardella.
• 6 The fish community consumed only 3% of estimated crustacean zooplankton production directly. Predation pressure was intense (> 80% of estimated production consumed by predators) on late instars of the zooplanktivorous C. edulis larvae and on E. sardella larvae. Rhamphochomis spp. consumed 22–43% of estimated production of the zooplankton-eating fishes.
• 7 Although many members of the fish community do not feed directly on crustacean zooplankton, so that the food-chain supporting fish production involves an extra trophic level, it is considered that the endemic fish community is well adapted to feeding on the low-density prey in the pelagic zone, and that there is no vacant feeding niche.

     

The Relevance of Individual Size to Management of Arctic Charr, Salvelinus Alpinus, Populations

Arctic charr, Salvelinus alpinus, tend to form stunted populations presumably due to competition for limited resources. In this context a long-term intensive fishery programme aiming at reducing charr density, and thereby increasing growth and harvestable sizes, was initiated in the early 1980s in a Norwegian alpine lake. Here we present long-term data on catch statistics and changes in mean weights of charr caught with gill nets on spawning grounds, as well as changes in mean weight of juvenile charr caught with funnel traps during the 1990s. Furthermore, we present results from shorter-term studies on growth, size- and age-distribution, and size-related habitat and resource utilisation of Arctic charr and brown trout, Salmo trutta, in this lake. Mean weight (± SD) of charr caught on the spawning grounds increased significantly from 129.8g (±11.9) in the years 1982–1990 to 213.1g (±37.8) in the years 1996–1999, whereas catch per unit of effort decreased significantly. Mean weight of juveniles caught in funnel traps increased significantly from 21.2g (±6.9) in 1993 to 41.9g (±14.8) in 1999. Apparently the increase in weight of spawning charr coincided with the onset of trap fishing for juvenile charr. Compared to generally shallow dwelling trout, charr grew rapidly and were generally found in deeper areas of the lake. Charr went through a distinct size-related niche shift from mainly consuming small zooplankton in the pelagic to consuming large benthic prey in shallower waters. Resource and habitat utilisation in different size-groups of charr and trout are discussed with respect to possible competitive and predatory intra- and interspecific interactions, and with regard to management of charr populations.     

Biogenic methane contributes to the food web of a large,shallow lake

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