Pelagic marine feeding of Arctic charr and sea trout

In a fjord in northern Norway, both immature and maturing individuals (170–500 mm fork length) of anadromous Arctic charr Salvelinus alpinus and anadromous brown trout (sea trout) Salmo trutta were captured up to 5000 m offshore. Both species had a high feeding intensity and an almost exclusive piscivorous diet, with herring Clupea harengus as the dominant prey, an apparently opportunistic response to a high density of small‐sized herring in the pelagic zone.     

Breeding ecology of the White-headed Petrel Pterodroma lessoni on the Kerguelen Islands

The breeding season of the White-headed Petrel Pterodroma lessoni on Mayes Island off the Kerguelen archipelago was studied between December 1985 and February 1987. The birds returned to land in November, and the chicks hatched at the end of January and fledged in May. These incubation and fledging periods are among the longest known in the family Procellariidae. Successful birds normally bred every two years, which has not previously been reported for any member of this family. The breeding ecology is compared to that of other gadfly petrels from sub-antarctic and tropical waters, and the breeding frequency is discussed.     

Temporal variability in marine feeding of sympatric Arctic charr and sea trout

The marine feeding pattern of anadromous brown trout (sea trout) Salmo trutta and Arctic charr Salvelinus alpinus was studied during June to August in 1992–1993 and 2000–2004 in a fjord in northern Norway. In general sea trout fed proportionally more on fishes than on crustaceans and insects (81, 1 and 18% by mass, respectively) by comparison with Arctic charr (52, 25 and 22% by mass, respectively). Herring Clupea harengus dominated the total fish diet of both species, but the Arctic charr also fed significantly on gadoids and sandlance Ammodytes spp. While sea trout became virtually all piscivorous at fork lengths ( L _F) ≥250 mm, the Arctic charr was ≥400 mm L _F before shifting totally to a fish diet. Despite annual variation in diet and forage ratios, there was a clear shift in diet from 1992–1993 to 2000–2004. Sandlance and different crustaceans constituted most of the diet during the initial period with a shift towards gadoids and especially herring during the latter period. This shift seemed to be associated with a high abundance of herring larvae during the latter sampling period, indicating a preferential selection on herring when present, particularly by sea trout. Furthermore, an index indicated dietary overlap in years with intensive feeding on herring of both species, and usually differences in the trophic ecology during years feeding mostly on other prey species. In combination, it was hypothesized that the two species reflect the type of marine prey present within a fjord system over time, and therefore provide an index of variation in the production and biological diversity of their potential prey within fjords.     

Upper thermal limits for feeding and growth of 0+ Arctic charr

When Arctic charr Salvelinus alpinus from two diVerent stocks were fed live Neomysis integer , the upper thermal limits for feeding and growth were established in the range 21·5–21·8° C. These critical temperatures might have been underestimates, because fish tend to show increased sensitivity to handling at high experimental temperatures. In the second experiment, the proportion of feeding undisturbed charr from four stocks decreased initially as temperature was raised in steps from 18 to 22° C. At the lower temperatures, 18 and 20) C, almost all fish resumed feeding, but the recovery time was longer and more fish ceased to feed at 20) C than at 18° C. When the temperature was increased to 21° C, 50% of the fish ceased feeding permanently, and all fish ceased feeding within 2 days at 22° C. It is concluded that 0+ charr cease to feed and grow at c .21·5) C and that the critical temperatures for feeding and growth coincide.     

Relationships between lake ecology and morphological characters in Icelandic Arctic charr,Salvelinus alpinus

The common occurrence of parallel phenotypic patterns suggests that a strong relationship exists between ecological dynamics and micro‐evolution. Comparative studies from a large number of populations under varying sets of ecological drivers could contribute to a better understanding of this relationship. We used data on morphology of arctic charr (Salvelinus alpinus) and ecological factors from 35 Icelandic lakes to test the hypothesis that morphological patterns among monomorphic charr populations from different lakes are related to interlake variation in ecological characteristics. There is extensive phenotypic diversity among populations of Icelandic charr, and populations are easily distinguished based on overall body morphology. The results obtained in the present study showed that the morphological diversity of charr was related to large‐scale diversity in lake ecology. Variation in charr morphology was related to water origin (e.g. spring fed versus run‐off), bedrock age, and fish community structure. The present study shows how various ecological factors can shape the biological diversity that we observe. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 103 , 761–771.     

The Swedish Arctic charr breeding programme

Results and experiences from a selective breeding programme aiming at improving the performance of an Arctic charr strain in aquaculture are presented. The programme, which has been running since 1985, uses traditional quantitative genetic methods based on relatedness and trait measurements. Traits considered included growth, age at sexual maturity, flesh colour, fat content and other features. Estimates in the early phase of the programme showed promising heritabilities for most traits, e.g. growth heritabilities of 0.34–0.52, and suggested that improvement by selection was feasible. Several traits such as a high rate of early maturation and poor flesh colour that were considered problematic by the Arctic charr farming industry at the programme’s start no longer hamper farming. Considerable improvement of growth by selection, estimated at 8% per generation, has contributed to shortening the production cycle in commercial farming. Results from studies of genotype–environment interactions are presented and discussed. Poor survival of fertilised eggs is a major problem in Swedish Arctic charr farming as indicated by presented survival rates. Efforts have consistently been made to avoid inbreeding in the selected strain, and at the present seventh generation the accumulated increase in inbreeding is estimated from pedigree data to be only approximately 5%.     

Polymorphism and speciation in Arctic charr

The Arctic charr Salvelinus alpinus exhibits 1–4 sympatric morphs in postglacial lakes, of which one or two are epibenthic zoobenthos feeders, one is a limnetic planktivore and one is a piscivorous form. In addition, northern rivers support partly migratory populations with anadromous and freshwater resident fish. The morphs vary in their coloration, morphology, life history, behaviour and genetic characteristics. The morphs usually differentiate according to their ontogenetic stage at maturity, which parallels paedomorphism in amphibians. The young usually start as epibenthic zoobenthivores, but may become pelagic at a certain size according to the predation risk experienced at that time. From a length of > 20–25 cm, charr start to become piscivorous. The proportion of piscivorous fish increases with increasing body size. In partly anadromous populations, fish that mature before smolting become freshwater resident, the others anadromous. In some rivers, the morphs occupy separate niches (epibenthic and limnetic), from emergence onwards. The morphs exhibit different degrees of reproductive isolation that vary from a high degree of interbreeding to complete isolation. Usually, they spawn within morph (assortative mating), but alternative male mating behaviour (sneaking, fighting) may occur in stream‐spawning populations and at great depths in lakes. Morphologically specialized morphs appear to feed more effectively than intermediate forms, and selection according to feeding mode, site fidelity and associated assortative mating are prerequisites for the evolution of the different morphs. Charr morphs develop into stable feeding niches under conditions of intense intraspecific competition when there is little competition with other species. Sympatric morphs exhibit different degrees of speciation, but similar morphs in different systems are not individual species because of (1) their polyphyletic origin, (2) the supporting systems are often young, transient environments making the future situation for the populations uncertain, and (3) the genetic differentiation among morphs is low. Sympatric morphs may interbreed and produce fertile hybrids. Nevertheless, sympatric charr morphs should be managed as separate species. Changes in the natural conditions or human impacts to which the morphs are adapted will have a strong influence on the persistence and survival of each different morph.     

Piscivory and cannibalism in Arctic charr

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

The functional significance of inherited differences in feeding morphology in a sympatric polymorphic population of Arctic charr

Artificially fertilised eggs from wild-caught Arctic charr parents of two sympatric morphs (benthivorous and planktivorous) from Loch Rannoch, Scotland were reared in the laboratory under identical conditions. During the subsequent 2 years, aspects of their trophic anatomy and feeding behaviour were compared. As previously described for wild-caught fish, charr derived from the benthivorous morph had an increasingly wider mouth gape for a given body length than those derived from the planktivorous morph. The functional significance of these differences in gape was tested by comparing the maximum size of prey that could be handled by each of the two morphs. In both forms, a larger gape enabled larger food particles to be eaten, but the elevation of the regression of maximum prey size on gape was higher in the benthivorous form, indicating the existence of additional morphological and/or behavioural differences influencing the size of prey consumed. When offered a choice between a typical benthic prey item and a typical pelagic food item, charr of benthivorous origin were more likely to feed on the former, whereas those of planktivorous origin were more likely to feed on the latter. Thus inherited differences in gape place constraints on foraging ability and are associated with inherited differences in dietary preference. We conclude that the functional significance of the foraging specialisations indicate a strong selection pressure for the evolution of the divergence and propose that heterochronic growth is the mechanism resulting in the divergence of tropic anatomy.     

Ecological polymorphism in Arctic charr

Arctic charr, Salvelinus alpinus (L.), commonly exhibits two coexisting morphotypes, dwarf and normal charr, which are characterized by differences in adult body size and colouration. We tested whether or not the morphotypic differences were genetically determined in rearing experiments with offspring of the two morphs and of their crosses. The experiments suggest that this ecological polymorphism in Arctic charr is largely environmentally determined. When reared under similar conditions, offspring of each of the two morphs differed little in size at the same age, and they had the same early developmental rate and maturation pattern. Moreover, the presence of parr marks along the flanks of the fish, one characteristic of dwarf charr, depended on body size and not on parental morph. Genetic differences between the morphs were suggested for growth rate during the second year of life, and for jaw morphology. Comparisons between charr life histories in captivity and in the wild suggest that ecological polymorphism in Arctic charr is chiefly a result of variation in growth conditions between different habitats.     

Epigenetic regulation of trophic morphology through feeding behaviour in Arctic charr, Salvelinus alpinus

Several models of speciation suggest that in species that are phenotypically plastic, selection can act on phenotypic variation that is environmentally induced in the earliest stages of divergence. One trait that could be subject to this process is foraging behaviour, where discrete foraging strategies are common. One species which is highly plastic in the expression of phenotype, the Arctic charr, Salvelinus alpinus (L.), is characterized by discrete variation in the anatomy of the head and mouthparts. These traits have been shown to have a functional significance, but the expression of which is thought to be at least partly phenotypically plastic. Here we test the hypothesis that foraging behaviour may regulate the anatomy of the head and mouthparts in Arctic charr. In a dyad experiment, size‐matched pairs of fish from a mixed family group were fed a diet of either Mysis (a hard‐bodied shrimp) or Chironomid larvae. Nine morphometric measures of head dimensions that describe wild trophic morphs were measured at the start of the experiment and 24 weeks later. Principal component scores of size‐corrected morphometric measures showed highly significant differences between fish exposed to the two diets. Univariate ANOVA analysis of the head morphometric variables showed that fish fed on Chironomids developed longer, wider jaws, longer heads and a larger eye for a given body length than did those fish fed upon Mysis. We conclude that foraging anatomy in Arctic charr is phenotypically plastic and that variation in foraging behaviour that results in feeding specialization in the wild could induce variation in head anatomy. This in turn could reinforce foraging specialization. Very rapid epigenetic divergence into distinct feeding morphs (as demonstrated here) would allow selection to act at more than one mode and thus could promote rapid evolutionary divergence, initially prior to genetic segregation, in species which are highly plastic. © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 78 , 43–49.     

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

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

Effects of the eye fluke Diplostomum spathaceum on the energetics and feeding of Arctic charr Salvelinus alpinus

In the present study, effects of the trematode eye fluke Diplostomum spathaceum infection and parasite‐induced cataract on energetic costs and feeding of hatchery‐bred one‐summer‐old Lake Saimaa Arctic charr Salvelinus alpinus were examined. The energetic effects (measured as oxygen uptake) of acute eye fluke infection on S. alpinus were negligible, but in the chronic infections mean oxygen consumption of infected fish was significantly higher than that of controls. Feeding capability of S. alpinus was studied in aquarium experiments, and uninfected healthy‐eyed fish reacted to zooplankton more rapidly and caught zooplankton with better success compared to individuals with cataracts in the same school. The results suggested that to compensate energetic costs and impaired feeding capability caused by eye fluke, cataract‐bearing fish spend more time foraging, which may result in increased predation risk.     

The influence of previous feeding regime on the compensatory growth response of maturing and immature Arctic charr, Salvelinus alpinus

Alternating periods of food deprivation with those of unlimited provision of food depressed the growth of Arctic charr, Salvelinus alpinus , below that of controls. Fish that were deprived of food and then fed on alternate weeks (1:1) were larger than those that were exposed to periods of 1 5- or 3-week deprivation and feeding (1·5:1·5 or 3:3). On receiving excess food supplies following 24 weeks on the restricted feeding regimes the previously-restricted fish grew more rapidly than the controls. The greatest compensatory growth was displayed after the 3:3 regime, followed by the 1·5: 1·5 and then the 1:1 feeding regime. At the termination of the experiment there were no significant differences in body weight between fish fed according to the different regimes during the period that food restriction was imposed. Growth patterns of the immature males and females were similar, but mature males were significantly lighter than the immature fish by the end of the experiment. Both immature and maturing fish displayed a compensatory growth response on return to adequate feeding. Beginning food restriction in May did not influence the proportions of male fish ( c . 60%) which were mature in the autumn.     

The complete history of salmonid introductions in the Kerguelen Islands, Southern Ocean

Since the early 1950s, several species of salmonids have been introduced more or less successfully in the Kerguelen Islands, a 7,215 km² archipelago located in the Southern Ocean (49°S, 70°E) and previously devoid of any freshwater fish. The aim of this work was to establish a documented chronicle of these events from available archives, to better understand the causes of the colonization failure or success for the different species. The history that emerged from the analysis of the archives appeared much more complex than previously published. Stocks of various origins were used, and numerous attempts were made at different sites involving variable numbers of fish released at different life stages. Between 1951 and 1991, 22 importation attempts took place, involving about 2 million individuals. Of the 8 species introduced (Salmo trutta, S. salar, Oncorhynchus mykiss, O. tshawytscha, O. kisutch, Salvelinus namaycush, S. fontinalis and S. alpinus), only 3 failed to establish local populations (O. mykiss, O. tshawytscha and S. namaycush). Overall, 23 watersheds were stocked. At present, 45 watersheds are colonized by one or several species. S. trutta, S. fontinalis, S. alpinus and O. kisutch were capable of migrating toward new habitats. The brown trout (S. trutta) was the only species to colonize a large number of watersheds (32 in about 10 generations). Its success can be explained by the diversity of origins, the number and importance of introduction and transfer attempts, the diversity of release sites and the peculiarities of its life cycle.     

Size-scaling of zooplankton foraging in Arctic charr

Prey capture rate (number of prey s⁻¹) and the mode of feeding of Arctic charr Salvelinus alpinus were studied by performing foraging experiments with two sizes (1·1 and 1·8 mm) of Daphnia longispina prey. Arctic charr were particulate feeders at all densities tested. Adjusted for the effect of prey density, the capture rate showed a hump-shaped relationship with Arctic charr size for both sizes of D. longispina . Estimated attack rates ( a ) also tended to show a hump-shaped relationship with fish size. The estimated size-scaling exponent of the attack rate function, however, was relatively small, implying small changes in attack rate over fish sizes. Simultaneous estimations of a and handling time were used in combination with published data on fish metabolism and dry mass rations of prey to estimate maintenance resource density of prey as a function of Arctic charr mass. Maintenance resource densities increased monotonically with Arctic charr size, and rapidly as optimum fish size relative to attack rate on prey was passed.     

Influence of maturity status on the annual cycles of feeding and growth in Arctic charr reared at constant temperature

Temporal changes in feeding and growth of immature (1 +), maturing male (1 +), and mature male and female (3+) Arctic charr were studied by monitoring feed intake and growth of individually-marked fish for over 1 year. The fish were held at a constant temperature (4°C) under conditions of liberal feed supply. Feed intake and growth remained low in the period December-April, with only 20–60% of the fish having fed on each occasion that feed intake was monitored. Feed intake and growth increased markedly during the late spring and summer, but the increases were delayed by approximately 1–2 months in the immature fish compared with their maturing counterparts. Maturing fish decreased feeding during the late summer and by September there had been an almost complete cessation of feeding. This was reflected in changes in fish body weight and condition, both of which declined from August-September onwards. By contrast, the immature fish peaked in weight and condition in September, and weight loss during the autumn tended to be less rapid than observed for the maturing fish. The fish appeared to reduce feeding once a'threshold condition' of 1–4—1–5 had been attained. It is suggested that the fish may become anorexic once there has been replenishment of the energy reserves required for overwintering and, in the case of maturing individuals, for the completion of gonadal growth and Spawning.     

Vaccination influences growth of Arctic charr

There is limited knowledge about the effects of oil-based vaccines on the growth of Arctic charr Salvelinus alpinus, in particular at different rearing temperatures. One-year-old Arctic charr were immunized intraperitoneally at 2.9 degrees C with a metabolizable oil-adjuvanted, bivalent vaccine containing killed typical and atypical Aeromonas salmonicida bacteria. After vaccination the non-vaccinated (controls) and vaccinated individually marked fish were held for 20 d at 10.0 degrees C and then for 7 wk at 10.3, 14.1 or 18.1 degrees C. During the first 20 d at 10.0 degrees C the growth rate (G) was higher for non-vaccinated than vaccinated fish. Thereafter vaccinated charr had higher G than control fish at 10.3 and 14.1 degrees C. In contrast, at 18.1 degrees C there was no difference in G and therefore no compensation of earlier growth suppression in vaccinated fish was observed at that temperature. The study indicates that vaccination has no ultimate negative effects on the growth of Arctic charr at temperatures ranging from 10.3 to 14.1 degrees C.     

Genetic differences and environments of mussel populations in Kerguelen Islands

Summary Samples of subantarctic mussels, Mytilus desolationis, from 12 locations in the Kerguelen Islands (South Indian Ocean) were analysed electrophoretically at 5 polymorphic loci to study correlations between genotype frequencies and environmental factors. Thirty-one out 48 pairs (population-locus) were not in Hardy-Weinberg equilibrium proportions, and 41 out 60 showed heterozygote deficiency. Spatial heterogeneity analysed with correspondence analysis was observed at 3 loci (Lap2, Pgd and Pgm). The genetic differentiation of M. desolationis could be explained through both geographical origin of populations and adaptation to environmental conditions. Within each geographical group, the first parameter influencing spatial structure is salinity. Additional differentiation is related to a factor linked to maximal size and to a lesser extent to shore exposure. The higher genetic differentiation occurring within the Kerguelen populations than among continental populations could correspond to incomplete panmixia due to a reduced gene flow versus constant external immigration in continental populations.     

Upstream migratory behaviour in landlocked Arctic charr

Synopsis Extensive upstream migration of landlocked Arctic charr during spring floods was recorded in several tributaries of an oligotrophic lake in north-west Sweden. Migration was confined to a period of about two weeks and residence in most creeks was of short duration. Only fish migrating to two small productive lakes remained in the new habitat over the summer. Repeated annual migrations were only recorded in the creek leading to these lakes and no straying was observed among repeat migrants. Water temperatures provided the primary cues for initiation and direction of migration, although an ability to detect productive habitats by odour was indicated. Creek size, feeding opportunities during migration and conspeeific odour were subordinate guiding factors.