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Ontogenetic dietary shifts are common in fish and often impact trophically transmitted parasite communities. How parasite species composition and relative abundances change among size classes, and at what rate these changes occur, is rarely examined. Hosts with a broad trophic niche are potentially exposed to a large variety of parasite species. The degree of ontogenetic changes in parasite species composition versus changes in parasite abundance should suggestively differ between thropically generalist and specialist host species. In the present study, we explore ontogenetic dietary shifts and their impact on species composition and relative abundance of intestinal parasites in two sympatric salmonid fish species, Arctic charr (Salvelinus alpinus) and brown trout (Salmo trutta) caught in the littoral habitat of a subarctic lake. Our results highlight a close interplay between ontogenetic dietary niche shifts and alterations in the acquisition of trophically transmitted parasites, leading to host-specific differences in the component community of parasites. Ontogenetic changes in the intestinal parasite community related to dietary niche shifts were distinct but less pronounced in Arctic charr than in brown trout due to a broader and more consistent dietary niche of the former and an ontogenetic shift toward piscivory in the latter. At the component community level, changes in parasite assemblages of both host species were driven by a faster increase in the heterogeneity of parasite relative abundance than in the compositional heterogeneity, a pattern that partly may be related to a rather species-poor parasite community of this subarctic study system. Separating compositional heterogeneity from heterogeneity in relative parasite abundance is important to understand how size-dependent variability shapes parasite communities of host populations., Graphical abstract Image 1, Highlights • Ontogenetic dietary niche shifts generated host-specific differences in the component community of parasites. • Ontogenetic changes in the intestinal parasite community were less pronounced in Arctic charr than in brown trout. • Heterogeneity of parasite relative abundance increased faster than the compositional heterogeneity.

Life history strategies and potential marine niche use of Arctic charr Salvelinus alpinus (n = 237, 84–652 mm, total body length, LT) were determined during the ice‐free season (2012) at three different watercourses in south‐western Greenland. All Arctic charr were collected from freshwater habitats. Based on stable isotopes of δ34S, the Arctic charr were categorized as either marine‐ or freshwater‐dependent feeders. The use of time‐integrated trophic tracers (stable isotopes of δ13C, δ15N, δ34S) suggested that several trophic groups of Arctic charr operate alongside within each fjord system. The groups suggested were one group that specialized in the marine habitat, in addition to two freshwater resident morphs (small‐sized resident and/or large‐growing cannibalistic individuals). Stomach contents consisted entirely of freshwater and terrestrial prey (i.e., insects), indicating that marine‐dependent feeders also fed in freshwater habitats after return from their marine migration. Growth and maturity patterns further supported variable life history strategies within each watercourse. The life history strategy patterns and marine trophic niche use were consistent across the watercourses along several hundred kilometres of coastline. This study represents the first ecological baseline for partially anadromous populations of Greenland Arctic charr. © 2020 The Authors. Journal of Fish Biology published by John Wiley & Sons Ltd on behalf of The Fisheries Society of the British Isles. This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited.

This is the peer reviewed version of the following article: Henriksen, E.H., Frainer, A., Knudsen, R., Kristoffersen, R., Kuris, A.M., Lafferty, K.D. & Amundsen, P-A. (2019). Fish culling reduces tapeworm burden in Arctic charr by increasing parasite mortality rather than by reducing density-dependent transmission. Journal of Applied Ecology, 56(6), 1482-1491, which has been published in final form at https://doi.org/10.1111/1365-2664.13369. This article may be used for non-commercial purposes in accordance with Wiley Terms and Conditions for Use of Self-Archived Versions. Two common Dibothriocephalus (formerly Diphyllobothrium) tapeworm species were significantly reduced by experimental culling of their fish host Arctic charr (Salvelinus alpinus) in a subarctic lake. Between 1984 and 1991, funnel traps were used to cull ~35 metric tons of Arctic charr, reducing charr density by ~80%. As charr densities decreased, tapeworm prevalence and then intensity also declined over the following three decades, with D. dendriticus (formerly dendriticum) responding faster than D. ditremus (formerly ditremum). The two main hypotheses for how culling a host can decrease parasitism are reductions in parasite transmission due to reduced host density and reductions in parasite survival through increases in host mortality rates. We found little evidence that charr density was the main driver for reduced parasite transmission. Instead, decreased survivorship in charr, initially, through fishing‐induced changes in charr age structure, and later through increased predation rates by brown trout, led to increased parasite mortality. Although brown trout, which increased significantly after fish culling, are also hosts, they are often too big for the final host birds to eat, thus becoming parasite sinks. Synthesis and applications. Fish populations with heavy parasite burdens constitute a management problem. Our results show how fish culling can indirectly reduce transmitted parasites through increased parasite mortality. Managing overcrowded fish populations by culling can produce two desirable outcomes: an increase in fish growth rates and reduced parasite burdens.

Accepted manuscript version. Final version published in Hydrobiologica, 840(1), 261-270, is avaialble at https://doi.org/10.1007/s10750-018-3865-8. Trophically transmitted parasites are known to impair fish growth in experimental studies, but this is not well documented in natural populations. For Arctic charr [Salvelinus alpinus (L.)], individual growth is positively correlated with food consumption. However, increased food consumption will increase the exposure to trophically transmitted parasites. Using a correlative approach, we explore the association between parasite abundance and the individual growth of Arctic charr from five lakes within the same watercourse. The studied parasite species differ in their life cycles and cost to the host. We predicted a positive association between parasite abundance and fish growth for parasites of low pathogenicity reflecting high consumption rates, and a negative association at higher parasite abundances for more costly parasites. We found no direct negative associations between parasite abundance and fish growth. The relationship between parasite abundance and growth was linearly positive for the low costly Crepidostomum sp. and concave for the more costly Eubothrium salvelini. In natural fish populations, the negative effects of parasites on fish growth might be outweighed by the energy assimilated from feeding on the intermediate host. However, experimental studies with varying food consumption regimes are needed to determine the mechanisms underlying our observations.

Amphipods are often key species in aquatic food webs due to their functional roles in the ecosystem and as intermediate hosts for trophically transmitted parasites. Amphipods can also host many parasite species, yet few studies address the entire parasite community of a gammarid population, precluding a more dynamic understanding of the food web. We set out to identify and quantify the parasite community of Gammarus lacustris to understand the contributions of the amphipod and its parasites to the Takvatn food web. We identified seven parasite taxa: a direct life cycle gregarine, Rotundula sp., and larval stages of two digenean trematode genera, two cestodes, one nematode, and one acanthocephalan. The larval parasites use either birds or fishes as final hosts. Bird parasites predominated, with trematode Plagiorchis sp. having the highest prevalence (69%) and mean abundance (2.7). Fish parasites were also common, including trematodes Crepidostomum spp., nematode Cystidicola farionis, and cestode Cyathocephalus truncatus (prevalences 13, 6, and 3%, respectively). Five parasites depend entirely on G. lacustris to complete their life cycle. At least 11.4% of the overall parasite diversity in the lake was dependent on G. lacustris, and 16% of the helminth diversity required or used the amphipod in their life cycles. These dependencies reveal that in addition to being a key prey item in subarctic lakes, G. lacustris is also an important host for maintaining parasite diversity in such ecosystems.

Changes in abiotic and biotic factors between seasons in subarctic lake systems are often profound, potentially affecting the community structure and population dynamics of parasites over the annual cycle. However, few winter studies exist and interactions between fish hosts and their parasites are typically confined to snapshot studies restricted to the summer season whereas host‐parasite dynamics during the ice‐covered period rarely have been explored. The present study addresses seasonal patterns in the infections of intestinal parasites and their association with the diet of sympatric living Arctic charr (Salvelinus alpinus) and brown trout (Salmo trutta) in Lake Takvatn, a subarctic lake in northern Norway. In total, 354 Arctic charr and 203 brown trout were sampled from the littoral habitat between June 2017 and May 2018. Six trophically transmitted intestinal parasite taxa were identified and quantified, and their seasonal variations were contrasted with dietary information from both stomachs and intestines of the fish. The winter period proved to be an important transmission window for parasites, with increased prevalence and intensity of amphipod‐transmitted parasites in Arctic charr and parasites transmitted through fish prey in brown trout. In Arctic charr, seasonal patterns in parasite infections resulted mainly from temporal changes in diet toward amphipods, whereas host body size and the utilization of fish prey were the main drivers in brown trout. The overall dynamics in the community structure of parasites chiefly mirrored the seasonal dietary shifts of their fish hosts., Seasonal variation in prevalence and intensity of trophically transmitted intestinal parasites hosted by Arctic charr (Salvelinus alpinus) and brown trout (Salmo trutta) in high‐latitude lakes is linked to dietary shifts occurring throughout the seasons. Winter proved to be an important transmission window, with increased prevalence and intensity of amphipod‐transmitted parasites in Arctic charr and parasites transmitted through piscivory in brown trout. In Arctic charr, seasonal patterns in intestinal parasite infections resulted mainly from temporal changes in diet towards amphipods, whereas host body size and the utilization of fish prey were the main drivers in brown trout.

The littoral benthivorous charrs Salvelinus malma with subterminal mouth distributed in Lake Kronotskoe basin are separated into three distinct morphs. These morphs significantly differ in head proportions, jaws' length, skull structure, and degree of gill rakers' reduction. The morphs spawn in distant lake tributaries at the spawning grounds of various types. Joint spawning has never been observed for these morphs in any type of spawning grounds. The differences in the external morphology and growth rate of the morphs become evident during the riverine life stage. The adult charrs inhabit the same sites of the littoral zone. No reliable difference is found in the diet among morphs, since all nosed charrs dominantly feed on amphipods with the exception of a few individuals that fed on chironomid larvae.

This is a post-peer-review, pre-copyedit version of an article published in Hydrobiologia. The final authenticated version is available online at https://doi.org/10.1007/s10750-019-3953-4. Niche diversification of polymorphic Arctic charr can be altered by multiple anthropogenic stressors. The opossum-shrimp (Mysis relicta) was introduced to compensate for reduced food resources for fish following hydropower operations in Lake Limingen, central Norway. Based on habitat use, stomach contents, stable isotopes (δ13C, δ15N) and trophically transmitted parasites, the zooplanktivorous upper water-column dwelling ‘normal’ morph was clearly trophically separated from two sympatric deep-water morphs (the ‘dwarf’ and the ‘grey’) that became more abundant with depth (> 30 m). Mysis dominated (50–60%) charr diets in deeper waters (> 30 m), irrespective of morph. Mysis and/or zooplankton prey groups caused high dietary overlap (> 54%) between the ‘dwarf’ morph and the two other ‘normal’ and ‘grey’ morphs. After excluding Mysis, the dietary overlap dropped to 34% between the two profundal morphs, as the ‘dwarf’ fed largely on deep-water zoobenthos (39%), while the ‘grey’ morph fed on fish (59%). The time-integrated trophic niche tracers (trophically transmitted parasites and stable isotopes) demonstrated only partial dietary segregation between the two deep-water morphs. The high importance of Mysis in Arctic charr diets may have reduced the ancestral niche segregation between the deep-water morphs and thereby increased their resource competition and potential risk of hybridization.

The first international symposium on Arctic charr (Salvelinus alpinus) was held in May, 1981, in Winnipeg, Manitoba, Canada, following shortly after the release of the seminal publication Charrs: Salmonid Fishes of the Genus Salvelinus (Balon, 1980), and sought to bring together scientists from nations in the circumpolar region containing Arctic charr or its close relatives in the S. alpinus complex. Since then, symposia have been held in Japan (1988), Norway (1994), Canada (2000), Iceland (2006), Scotland (2009) and Russia (2012). On the 14th June 2015, reflecting a buoyant research interest in Salvelinus, 112 participants from 13 countries gathered for the 8th International Charr Symposium in Tromso, Norway. Hosted by the UiT, The Arctic University of Norway, delegates included: 9 from Russia, 13 from USA, 15 from Canada, 9 from Japan, 8 from Iceland, 6 from Sweden, 3 from Scotland, 2 from Finland, 1 each from France, The Netherlands, Switzerland and Spain and 43 from Norway. In total, there were 79 oral presentations and 28 posters presented over the 4 days of meetings. Topical themes covered by the presentations and posters included anadromy, aquaculture, climate change, ecology, evolution, genetics, fisheries management, life history, parasitology and pollution impacts. Although the majority (two-thirds) of studies presented at the symposium focused on Arctic charr, there were many devoted to Dolly Varden charr (Salvelinus malma), Lake charr (Salvelinus namaycush), Brook charr (Salvelinus fontinalis) and Whitespotted charr (Salvelinus leucomaenis). The contributions covered two major geographic areas: northern Europe, and the Pacific basin drainages of the Asian and North American coasts. The vast area comprising the range of Salvelinus between these two more highly investigated regions was represented by only a few presentations from the Kola Peninsula, Siberia and Greenland, with the geographic spread of studies Guest editors: M. Power, R. Knudsen, C. E. Adams, M. J. Hansen, J. B. Dempson, M. Jobling & M. Ferguson / Advances in Charr Ecology and Evolution

Accepted manuscript version. Published version available in Effects of fish species composition on Diphyllobothrium spp. infections in subarctic brown trout - is three-spined stickleback a key species? (2016) 39(11), s.1313-1323. . Subarctic populations of brown trout (Salmo trutta) are often heavily infected with cestodes of the genus Diphyllobothrium, assumedly because of their piscivorous behavior. This study explores possible associations between availability of fish prey and Diphyllobothrium spp. infections in lacustrine trout populations. Trout in i) allopatry (group T); ii) sympatry with Arctic charr (Salvelinus alpinus) (group TC); and iii) sympatry with charr and three-spined stickleback (Gasterosteus aculeatus) (group TCS) were contrasted. Mean abundance and intensity of Diphyllobothrium spp. were higher in group TCS compared to group TC and group T. Prevalence however was similarly higher in group TCS and group TC compared to group T. Zero-altered negative binomial modelling identified lowest probability of infection in group T and similar probabilities of infection in group TC and group TCS, whereas the highest intensity was predicted in group TCS. Evidently, the most infected trout were from the group co-occurring with stickleback (TCS), assumedly due to a high availability of suitable prey fishes. In conclusion, our study demonstrates elevated Diphyllobothrium spp. infections in lacustrine trout populations where fish prey are available, and suggests that highly available and easily caught stickleback prey may play a key role in the transmission of Diphyllobothrium spp. parasite larvae.

Seasonal dietary shifts by Arctic charr (Salvelinus alpinus (L.)) are common in subarctic lakes, but less is known about how the shift to pelagic zooplankton feeding during the growing season affects stable isotope ratios and mercury concentrations. We sampled Arctic charr, zooplankton and benthic macroinvertebrates from oligotrophic Lake Galggojavri, northern Norway, in June, August and September. Stomach content, age, total length, weight, condition and sex were recorded for all individuals, and liver and muscle tissues were sampled for stable isotope (δ13C, δ15N) and total mercury analyses. We predicted that a dietary shift to zooplankton would lead to depleted 13C and increased mercury levels in charr. Arctic charr consumed benthic prey in June, but shifted to zooplankton feeding in August–September. Stable isotope mixing models revealed increased pelagic reliance towards September. Mercury content in liver increased from June to September, whereas muscle showed opposite trend. In stepwise multiple regression analyses, mercury content in muscle was explained by fish length, month and δ13C (R 2 = 0.46), whereas in liver month, δ13C and δ15N were the main explaining factors (R 2 = 0.69). Seasonal dietary shifts appeared to have an effect on total mercury content in charr, and thus sampling month should be considered when designing future monitoring programmes.

Arctic charr (Salvelinus alpinus) is generally considered an inferior piscivore compared to brown trout (Salmo trutta). However, we demonstrate that a recently described profundal spawning piscivore PP-morph of Arctic charr in a subarctic lake has evolved typical piscivore traits (i.e. large jaws, robust skulls), similar to the co-occurring brown trout but different from the sympatric littoral spawning omnivore LO-morph of Arctic charr. A few large-sized LO-morph, most large-sized PP-morph and trout constitute the piscivore guild, but they differed in prey fish selection. Of the fish-eating deep-water PP-morph, 52% had consumed charr, with a peak during the polar night period (68–80% in November–January). In contrast, the LO-morph and trout were mainly (~90%) inter-specific piscivores eating stickleback. Consequently, the PP-morph was the main charr predator and started to feed on fish at a smaller size and took larger prey relative to their body length than the trout. Stable isotope (δ13C, δ15N) ratios reflect the dietary specialisations in the upper-water (LO-morph, trout) and the deep-water (PP-morph) environments, besides the piscivorous behaviour of the PP-morph and trout. The existence of a shallow-water piscivore (trout) may explain the origin of this PP-morph, taking benefit of underutilised resources of small-sized Arctic charr in the deep-water habitat.

We contrast the genetic consequences of allopatric and sympatric divergence from the littoral spawning Arctic charr morph from Lake Fjellfrosvatn. The littoral spawning Arctic charr has sympatrically diverged into a natural profundal adapted morph and via a recent (1930) translocation of about 40 adult Arctic charr established a new allopatric population in a nearby lake (Takvatn). The sympatric morph-pair in Fjellfrosvatn was more genetically differentiated (F ST = 0.121), the derived profundal morph had higher genetic variation (H e = 0.740 ± 0.220; N PAR = 4.87), and had a higher proportion of linkage disequilibrium among loci, than the Takvatn charr derived in allopatry (F ST = 0.066; H e = 0.584 ± 0.193; N PAR = 0.29). The genetic differentiation in allopatry supports a scenario of rapid population expansion, despite genetic founder effects, whereas the genetic differentiation found in the sympatric morph-pair suggests that this divergence is older and supports a scenario of divergence under ongoing gene flow. We suggest that the differences between the two types of divergence may originate from temporal effects and differences in life histories between the two derived populations along with specific factors related to the initial stages of the divergence.

A population of small-sized Arctic charr living their entire life cycle in deep water was recently found in Lake Skogsfjordvatn, Northern Norway. Their phenotype and life history closely resembles the profundal charr morph in another subarctic Norwegian lake, Fjellfrosvatn. Fjellfrosvatn has one littoral omnivorous and one small profundal benthivorous (PB) charr morph coexisting with brown trout, whereas Skogsfjordvatn has migratory salmon, trout, charr and eel populations in addition to threespined sticklebacks, resident trout and three resident charr morphs of which one is a large piscivorous profundal morph. We predicted that the small profundal charr morph in Skogsfjordvatn would have a similar trophic niche to the PB-morph in Fjellfrosvatn and tested this by using stomach content and stable isotope (δ13C and δ15N) analyses. Profundal benthic invertebrates, mainly chironomid larvae, Pisidium sp. mussels and small crustaceans, dominated the diets, and the δ13C and δ15N values indicated almost identical long-term trophic niches of the PB-morphs in both lakes. The prediction was therefore supported. This seems to indicate that the small profundal Arctic charr morphs in these lakes have evolved in parallel, in nearly identical and stable habitat and dietary niches in spite of likely different ecological interactions within the contrasting fish communities.

This study revealed between-lake genetic structuring between Coregonus lavaretus collected from the only two native populations of this species in Scotland, U.K. (Lochs Eck and Lomond) evidenced by the existence of private alleles (12 in Lomond and four in Eck) and significant genetic differentiation (FST = 0·056) across 10 microsatellite markers. Juvenile C. lavaretus originating from eggs collected from the two lakes and reared in a common-garden experiment showed clear phenotypic differences in trophic morphology (i.e. head and body shape) between these populations indicating that these characteristics were, at least partly, inherited. Microsatellite analysis of adults collected from different geographic regions within Loch Lomond revealed detectable and statistically significant but relatively weak genetic structuring (FST = 0·001–0·024) and evidence of private alleles related to the basin structure of the lake. Within-lake genetic divergence patterns suggest three possibilities for this observed pattern: (1) differential selection pressures causing divergence into separate gene pools, (2) a collapse of two formerly divergent gene pools and (3) a stable state maintained by balancing selection forces resulting from spatial variation in selection and lake heterogeneity. Small estimates of effective population sizes for the populations in both lakes suggest that the capacity of both populations to adapt to future environmental change may be limited.

Most ecosystems are exposed to multiple stressors acting in concert and their combined effects on parasite prevalence in freshwater, marine and terrestrial habitats are largely unknown. We investigated the relationships between farming intensity, water abstraction intensity and parasite prevalence in the mud snail Potamopyrgus antipodarum from 20 stream sites within the Manuherikia River catchment (New Zealand) by using generalized linear models and an information-theoretic model-selection approach. Three trematode taxa that use water birds as definitive hosts were found in the snail host. The average prevalence of all parasites infecting Potamopyrgus in the catchment was 5%. Microphallus sp. “lively”, the most common parasite, was most prevalent at high farming intensity and low water abstraction, besides showing an antagonistic interaction between the two agricultural stressors. These findings highlight the importance of considering multiple stressors and their potential interactions when studying host–parasite systems. Because snails often play key roles in aquatic communities, providing an important link between primary producers and higher trophic levels, and are a common intermediate host to a high diversity of trematode parasites, this host–parasite model system may represent a promising bioassessment tool for detecting anthropogenic disturbances in freshwater systems.

Parasite distribution patterns in lotic catchments are driven by the combined influences of unidirectional water flow and the mobility of the most mobile host. However, the importance of such drivers in catchments dominated by lentic habitats are poorly understood. We examined parasite populations of Arctic charr Salvelinus alpinus from a series of linear-connected lakes in northern Norway to assess the generality of lotic-derived catchment-scale parasite assemblage patterns. Our results demonstrated that the abundance of most parasite taxa increased from the upper to lower catchment. Allogenic taxa (piscivorous birds as final host) were present throughout the entire catchment, whereas their autogenic counterparts (charr as final hosts) demonstrated restricted distributions, thus supporting the theory that the mobility of the most mobile host determines taxa-specific parasite distribution patterns. Overall, catchment-wide parasite abundance and distribution patterns in this lentic-dominated system were in accordance with those reported for lotic systems. Additionally, our study highlighted that upper catchment regions may be inadequate reservoirs to facilitate recolonization of parasite communities in the event of downstream environmental perturbations.

Source at https://doi.org/10.1016/j.ijppaw.2018.01.007. Accepted manuscript version, licensed CC BY-NC-ND 4.0. The eye represents an immune privileged organ where parasites can escape host reactions. This study provides the first systematic evidence of the pathology associated with Diplostomum sp. infection in the eye retina of fish (i.e. Arctic charr). Histological sections showed that the trematodes caused mechanical disengagement between the retinal pigmentary epithelium and the neurosensory retina, with damaged cones and rods in the outer segment and epithelium reduced to a single layer of pigmentary cells. The metacercariae were “floating” in possibly fluid-filled vesicles together with several round cells, mostly located in the anterio-dorsal and anterio-ventral areas of the eye near the iris. The round cells may indicate internal retinal damage repair mechanisms, without connections to the general immune system. Metacercariae intestines contained pigmented cellular debris indicating that they feed on retinal epithelium. These retinal lesions may have similar vision effects as focal retinal detachment in vertebrates. Diplostomum metacercaria alters fish visual acuity but may in a lesser degree lead to a severe or total visual impairment because of repairing mechanisms. The pathology in the retina seems thereby to be dependent on fish size, age and dose.

Foraging trait specialization is important for polymorphic Arctic charr and whitefish, but visual capabilities of different morphs are unexplored. Photoreceptor complements and absorbance spectra of rod visual pigments were studied by microspectrophotometry in two sympatric Arctic charr morphs and three sympatric whitefish morphs from two subarctic lakes. Four spectral classes of photoreceptor cells, rods and three types of cones, were found in all morphs of both species. Arctic charr rods had a pure A1 pigment (rhodopsin) with wavelength of maximum absorbance λ max ≈ 511–512 nm and no significant differences either between littoral and profundal morphs or sampling times (January/August). Rods of littoral and pelagic whitefish had practically pure A2 pigment (porphyropsin), whereas profundal whitefish had chromophore mixtures with A2:A1 ≈ 0.8:0.2 in June, A1 decreasing to a smaller fraction in September. λ max values of littoral and pelagic whitefish rods were similar and did not change significantly with season (539.3 ± 0.3 nm/539.3 ± 1.1 nm and 538.4 ± 0.4/539.8 ± 0.3 nm in June/September) but differed from profundal whitefish (λ max = 531.5 ± 0.8/536.7 ± 1.0 nm). Differences between Arctic charr and whitefish morphs suggest importance of local light environment determining visual pigment composition.

Seasonal growth patterns have been investigated in juveniles of two reproductively isolated Arctic charr [Salvelinus alpinus (L.)] morphs, which coexist in distinct ecological niches (upper-water and deep-water environments) in a subarctic lake. Offspring (F1-generation) of each of the littoral spawning charr (LO) and the profundal spawning charr (PB) were reared individually over a period of 12 months under simulated natural photoperiod (at 69°N) and two different temperature regimes to mimic the profundal zone conditions with constantly low temperature (~4 °C) and the littoral zone conditions with seasonally cycling temperatures. All charr increased in weight during the experimental period. The LO-morph performed generally best at naturally fluctuating temperature, whereas the PB-morph grew best at a constantly low temperature (4 °C). The LO-morph showed a typical seasonal growth pattern found earlier for anadromous Arctic charr, with elevated specific growth rates (SGR) during June–October and profoundly reduced or negative SGR-values during November–February suggesting that the fish ceased feeding during this period. In contrast, the PB-morph maintained a steady growth rate, regardless of temperature regime, and showed less signs of reduced appetite during the polar night period and exhibited significantly higher SGR-values during late autumn and early winter compared to the LO-morph. This study suggests different heritable adaptations in somatic growth trajectories where the deep-water PB-morph do not show a down-regulation of appetite and voluntary fasting during winter conditions with typically low light intensity and low temperatures, and up-regulation of appetite with hyperphagia during summer, as characteristically found for the upper-water LO-morph and anadromous Arctic charr.

Published version available at http://dx.doi.org/10.3354/dao02893 Parasite−host systems show markedly variable patterns in isotopic fractionation: parasites can be either depleted or enriched in 15N and 13C as compared to their hosts. However, it remains unknown whether isotopic fractionation patterns are similar in comparable parasite−host systems from markedly different ecosystems. Results of this study show that large-sized Schistocephalus spp. endoparasites are consistently depleted in 15N (by on average −2.13 to −2.20‰) as compared to their nine-spined stickleback Pungitius pungitius and three-spined stickleback Gasterosteus aculeatus hosts. The differences between parasites and host for both δ15N and δ13C were consistent in both study systems despite marked biogeographical differences between the study localities. Although the stable isotope values in general were strongly correlated between the hosts and their parasites, Schistocephalus specimens occupying the same nine-spined stickleback host showed sometimes substantial individual variation in δ13C. This might be due to selective use of different carbon sources, or different metabolic or feeding rates. Further studies on selective feeding, physiology and metabolism of parasites are needed to better understand the role of parasites in the structure and functioning of aquatic food webs.

To identify trematode diversity and life-cycles in the sub-Arctic Lake Takvatn, Norway, we characterised 120 trematode isolates from mollusc first intermediate hosts, metacercariae from second intermediate host fishes and invertebrates, and adults from fish and invertebrate definitive hosts, using molecular techniques. Phylogenies based on nuclear and/or mtDNA revealed high species richness (24 species or species-level genetic lineages) and uncovered trematode diversity (16 putative new species) from five families typical in lake ecosystems (Allocreadiidae, Diplostomidae, Plagiorchiidae, Schistosomatidae and Strigeidae). Sampling potential invertebrate hosts allowed matching of sequence data for different stages, thus achieving molecular elucidation of trematode life-cycles and exploration of host-parasite interactions. Phylogenetic analyses also helped identify three major mollusc intermediate hosts (Radix balthica, Pisidium casertanum and Sphaerium sp.) in the lake. Our findings increase the known trematode diversity at the sub-Arctic Lake Takvatn, showing that digenean diversity is high in this otherwise depauperate sub-Arctic freshwater ecosystem and indicating that sub-Arctic and Arctic ecosystems may be characterised by unique trematode assemblages.

Source at https://doi.org/10.1002/ece3.3224 . A study of body and head development in three sympatric reproductively isolated Arctic charr (Salvelinus alpinus (L.)) morphs from a subarctic lake (Skogsfjordvatn, northern Norway) revealed allometric trajectories that resulted in morphological differences. The three morphs were ecologically assigned to a littoral omnivore, a profundal benthivore and a profundal piscivore, and this was confirmed by genetic analyses (microsatellites). Principal component analysis was used to identify the variables responsible for most of the morphological variation of the body and head shape. The littoral omnivore and the profundal piscivore morph had convergent allometric trajectories for the most important head shape variables, developing bigger mouths and relatively smaller eyes with increasing head size. The two profundal morphs shared common trajectories for the variables explaining most of the body and head shape variation, namely head size relative to body size, placement of the dorsal and pelvic fins, eye size and mouth size. In contrast, the littoral omnivore and the profundal benthivore morphs were not on common allometric trajectories for any of the examined variables. The findings suggest that different selective pressures could have been working on traits related to their trophic niche such as habitat and diet utilization of the three morphs, with the two profundal morphs experiencing almost identical environmental conditions.

Arctic charr and European whitefish are considered to be strong competitors in lakes, with the latter usually being the superior species. However, high niche plasticity and lake morphometry may suggestively facilitate resource partitioning and coexistence between charr and whitefish. Here, we explore the trophic niche utilization (diet and habitat use) of charr and whitefish co-occurring with brown trout in the deep and oligotrophic Lake Fyresvatnet, southern Norway (59˚05’N, 8˚10’E). Using CPUE, stomach contents and stable isotope analyses, a distinct resource partitioning was revealed between brown trout and the other two species. Brown trout typically occupied the littoral zone, feeding on benthic invertebrates, surface insects and small-sized whitefish. In contrast, charr and whitefish were predominantly zooplanktivorous, but diverged somewhat in habitat utilization as charr shifted seasonally between the profundal and the littoral zone, whereas whitefish were found in the upper water layers (littoral and pelagic habitats). Accordingly, the stable isotope values of carbon (δ13C) reflected a pelagic orientated prey resource use for both charr and whitefish, whereas brown trout had elevated carbon and nitrogen (δ15N) signatures that reflected their benthivore and piscivore diet, respectively. The findings suggest that charr may not rely upon the profundal zone as a feeding habitat but as a refuge area, and may coexist with whitefish if a third competitive and predatory species like brown trout co-occur in the lake. The study indicates that a general high habitat plasticity of Arctic charr may be essential in the presently observed coexistence with a competitively superior fish species like whitefish, and that a third fish species like brown trout may facilitate this particular fish community structure. peerReviewed

Temperature preference of juvenile (age 1+) Arctic charr (Salvelinus alpinus L.) originating from four arctic and sub-arctic populations (Svalbard and mainland northern Norway), representing a range of habitats with different temperature conditions, was studied by use of a shuttle-box system which allowed individual fish to control their environmental temperature. Based on the assumption that adaptations to long-lasting differences in thermal environments would affect temperature preference, we expected that Arctic charr from the high arctic Svalbard would prefer a lower temperature than the charr from two well-studied sub-arctic mainland lakes (i.e. one anadromous charr population from Storvatn, Hammerfest and two sympatric resident charr morphs from Fjellfrosvatn, Malselv). There were, however, no significant differences in temperature preference among the four populations after 24 h exposure to the shuttle-box system, although the charr from the omnivore upper-water sympatric morph of Fjellfrosvatn used significantly longer time to reach a stable thermal preferendum than the fish of the other populations. The average temperature preference at the end of the trials ranged between 10.9 and 11.6 °C among the populations. The lack of population differences suggests that temperature preference is not a polymorphic trait under strong selection in Arctic charr.

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.

In polymorphic populations morphs usually diverge in morphology, ecology and life history, which is most likely driven by adaptations to different environments or resources. Sympatric morphs may develop differences in several life history traits to be able to maximize fitness in alternative niches and habitats. Here, the contrasting life history traits of three sympatric Arctic charr (Salvelinus alpinus (L.)) morphs in a deep and oligotrophic lake in sub-arctic Norway are addressed. The charr morphs differ in spawning habitat and trophic niche. One is a littoral spawning morph that feeds on benthic invertebrates and zooplankton in the littoral and pelagic zones (referred to as the LO-morph), and two other are profundal spawning morphs that either utilize profundal soft bottom benthos as food resource (the PB-morph) or are piscivorous (the PP-morph). The LO-morph typically had intermediate life-history traits relative to the two profundal morphs that had highly contrasting life history traits, especially in growth and age and size of maturity. The PB-morph matured at a young age (∼3 years) and at a small body size (∼8.5 cm), thereby increasing their fitness by investing in reproduction early in life, which results in a short generation time and decreased probability of being predated before first reproduction. The PP-morph on the other hand, matured at an old age (∼9.2 years) and a large body size (∼26 cm), thereby increasing their fitness by investing in somatic growth to enhance initial fecundity, and also to reach a large body size profitable for piscivory. The different trade-off regime between the PP- and PB-morphs seems to be caused by adaptation to alternative trophic niches, and appears to be an important factor for the co-occurrence of the two sister-morphs in the profundal zone.

Two unconnected time-integrated tracers of niche use provided similar conclusions about individual foraging behaviour and niche adaptations (functional traits in head shape) within a subarctic lake population of Arctic charr (Salvelinus alpinus). Zooplanktivorous individuals mainly from the pelagic zone were characterised by having low δ13C values, high infections of the parasites transmitted by pelagic copepods (especially Diphyllobothrium spp.) and slender heads with long snouts. In contrast, fish individuals that had consumed benthic prey in the littoral zone were enriched in δ13C and had high abundances of parasites transmitted by Gammarus lacustris (Cyathocephalus truncatus and Cystidicola farionis) and a robust head shape. There were strong positive correlations between individual δ13C values and the abundance of the two parasite species transmitted by Gammarus, but a negative correlation between δ13C and the infection of copepod-borne parasites. The close relationships between diet variation (foraging behaviour), the time-integrated ecological tracers (SI and parasites) and functional trophic morphology (niche adaptations) evidently reflect long-term temporally stable niche use of each individual predator. The two independent time-integrated tracers both gave valuable information of specialised trophic behaviour at the individual level, which is an important basis for studies related to ecological (e.g., resource partitioning) and evolutionary (e.g., polymorphism) topics within a population.

SUMMARY 1. Many species are expanding their distribution towards higher latitudes and altitudes in response to climate change. These range shifts are expected to change fish community structure and alter food-web dynamics in subarctic lakes. However, the impacts of invading species on native fish and invertebrate prey communities remain understudied. 2. The trophic ecology of invasive species determines the likelihood of direct resource competition with native taxa. In Northern Europe, perch (Perca fluviatilis), a trophic generalist, and ruffe (Gymnocephalus cernuus), a benthic specialist, are expanding their distribution ranges northwards, colonising lakes inhabited by a native generalist, whitefish (Coregonus lavaretus). We predicted that increased fish diversity and density would deplete the invertebrate community and increase resource competition between native and invasive species. 3. To assess the degree of resource competition between native and invasive species, we compared (i) fish and invertebrate community structure; (ii) diet and stable carbon and nitrogen ratios of whitefish, ruffe and perch and (iii) growth, condition and relative population size of whitefish in two noninvaded lakes with two lakes containing one and two lakes containing both invasive species. Each lake was sampled on a single occasion between August and September. 4. Benthic macroinvertebrate density and community structure were unaffected by increased consumer diversity, while top-down control of pelagic zooplankton density and size was evident in lakes with increased fish diversity. 5. Differences in diet and stable isotope ratios were evident between all whitefish populations, although these were not directly related to the presence of invasive species. Specialised adaptations of invasive species may confer a competitive advantage in invaded lakes; ruffe dominated the profundal niche, while perch displayed an ontogenetic shift to piscivory, reducing niche overlap with native whitefish. 6. Growth rate and population density of whitefish were largely independent of fish community structure and were governed by local variations in lake productivity. However, there was a sign of lowered condition of whitefish in invaded systems. Shallow and more productive lakes with higher food availability supported populations of native and invasive species. 7. Our findings indicate that trophic specialisations of invasive species play a key role in determining their impacts on the systems they invade. This study focussed on early stages of invasion, and the outcome of species interactions may change following the establishment of new species. In addition, these impacts will not be uniform across the invaded landscape as lake-specific variations in

In the study of mechanisms structuring fish communities, mixed competition–predation interactions where large predators feed on prey fish versus those in which small predators compete with prey fish for a shared prey have been the focus of substantial research. We used a long-term data set from a system inhabited by brown trout (Salmo trutta) (predator) and Arctic char (Salvelinus alpinus) (prey) to evaluate whether mixed interspecific interactions were present in this system as suggested in other studies focusing on this species pair. We found no evidence for a negative interspecific density dependence in individual performance in either Arctic char or brown trout. In contrast, a negative intraspecific density dependence was present, especially in Arctic char. Furthermore, large brown trout condition showed a positive response to encounter rate with Arctic char (related to the density of small Arctic char). The most parsimonious interaction module to explain the Arctic char – brown trout interaction patterns in the studied system does therefore not need to include interspecific competition. We suggest that size-structured mixed competition–predation interactions in different systems are realized as being either mainly structured through interspecific predation or by competition depending on species life history characteristics and environmental conditions.

Three genetically discrete morphs of Arctic charr in Loch Rannoch, Scotland originated from a recent divergence within the lake (in situ) (piscivore and benthivore morphs) and from secondary contact of two older lineages (ex situ; a planktivore–piscivore/ benthivore divergence). To test if the expression of traits with strong functional roles was linked to the age of the divergence, fin and gill anatomy, and dentition were quantified and compared across morphs. Five additional working hypotheses suggesting a rank order of trait expression amongst morphs were also tested. The planktivorous morph had more rays in the dorsal and pectoral fins, longer gill rakers (but not more) as well as a smaller gill cavity than the other two morphs. The piscivorous morph had more palatine teeth and longer teeth on the mandible, pre-maxillary and glossohyal bones, and a larger buccal cavity. These differences indicate a differential response to selection in these functional anatomical features most likely related to morph foraging specialisms. Notably, between-morph divergences in the expression of these traits were not simply linked to the length of divergence between morphs and have arisen equally quickly in the recent (in situ) divergence as they have in older, ex situ divergences.

This is the peer reviewed version of the following article: Kuhn, J.A., Knudsen, R., Kristoffersen, R. & Amundsen, P.A. (2017). A simplified method to estimate Diphyllobothrium spp. infection in salmonids. Journal of Fish Diseases, 40(7), 863-871., which has been published in final form at https://doi.org/10.1111/jfd.12566 . This article may be used for non-commercial purposes in accordance with Wiley Terms and Conditions for Self-Archiving. Some fish parasites constitute severe management problems as they may cause mortality of their fish host or are important zoonoses of humans. Parasite assessments are therefore critical to keep track of infections. If conventional sampling techniques can be simplified, parasite assessments might be easier to obtain, less time-consuming and more extensive. In this study, we compare the assessed number of Diphyllobothrium spp. cysts (CYST) with the counted number of Diphyllobothrium spp. plerocercoid larvae recovered using a conventional digestive technique (LARV). The aim was to determine the potential of using CYST as a simplified methodology for assessing Diphyllobothrium spp. infection in salmonids. In total, 365 brown trout and 424 Arctic charr were sampled from nine lakes in subarctic Norway. Strong correlation, significant linear relationship and large amount of explained variation were found between log10CYST and log10LARV in both fish species. The method had a slight, but not significant tendency to work better in charr compared to trout. In addition, absolute difference between CYST and LARV increased at parasite intensities >100 indicating that the method has reduced functionality when estimating parasite intensity in heavily infected salmonid populations. However, overall, using this simplified and less time-consuming methodology, a good indication of Diphyllobothrium spp. intensity, abundance and prevalence was obtained. We suggest that this method provides a sound proxy of the Diphyllobothrium spp. burden and have the potential to be used in parasite assessment during fish monitoring and fisheries management surveys, particularly if the time and resources for detailed parasite studies are not available.

The monogenean parasite Gyrodactylus salaris poses serious threats to many Atlantic salmon populations and presents many conservation and management questions/foci and challenges. It is therefore critical to identify potential vectors for infection. To test whether hybrids of native Atlantic salmon (Salmo salar) × brown trout (Salmo trutta) are suitable as reservoir hosts for G. salaris during winter, infected hybrid parr were released into a natural subarctic brook in the autumn. Six months later, 23.9% of the pit-tagged fish were recaptured. During the experimental period, the hybrids had a sixfold increase in mean intensity of G. salaris, while the prevalence decreased from 81% to 35%. There was high interindividual hybrid variability in susceptibility to infections. The maximum infrapopulation growth rate (0.018 day−1) of G. salaris throughout the winter was comparable to earlier laboratory experiments at similar temperatures. The results confirm that infrapopulations of G. salaris may reproduce on a hybrid population for several generations at low water temperatures (~1 °C). Wild salmon–trout hybrids are undoubtedly susceptible to G. salaris and represent an important reservoir host for the parasite independent of other co-occurring susceptible hosts. Consequently, these hybrids may pose a serious risk for G. salaris transmission to nearby, uninfected rivers by migratory individuals.

SUMMARY 1. Generalist fish species are recognised as important couplers of benthic and pelagic food-web compartments in lakes. However, interspecific niche segregation and individual specialisation may limit the potential for generalistic feeding behaviour. 2. We studied summer habitat use, stomach contents and stable isotopic compositions of the generalist feeder Arctic charr coexisting with its common resource competitor brown trout in five subarctic lakes in northern Norway to reveal population-level and individual-level niche plasticity. 3. Charr and trout showed partial niche segregation in all five lakes. Charr used all habitat types and a wide variety of invertebrate prey including zooplankton, whereas trout fed mainly on insects in the littoral zone. Hence, charr showed a higher potential to promote habitat and food-web coupling compared to littoral-dwelling trout. 4. The level of niche segregation between charr and trout and between pelagic-caught and littoralcaught charr depended on the prevailing patterns of interspecific and intraspecific resource competition. The two fish species had partially overlapping trophic niches in one lake where charr numerically dominated the fish community, whereas the most segregated niches occurred in lakes where trout were more abundant. 5. In general, pelagic-caught charr had substantially narrower dietary and isotopic niches and relied less on littoral carbon sources compared to littoral-caught conspecifics that included generalist as well as specialised benthivorous and planktivorous individuals. Despite the partially specialised planktivorous niche and thus reduced potential of pelagic-dwelling charr to promote benthic–pelagic coupling, the isotopic compositions of both charr subpopulations suggested a significant reliance on both littoral and pelagic carbon sources in all five study lakes. 6. Our study demonstrates that both interspecific niche segregation between and individual trophic specialisation within generalist fish species can constrain food-web coupling and alter energy mobilisation to top consumers in subarctic lakes. Nevertheless, pelagic and littoral habitats and food-web compartments may still be highly integrated due to the potentially plastic foraging behaviour of top consumers.

– 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.

There is now strong evidence that foraging niche specialisation plays a critical role in the very early stages of resource driven speciation. Here we test critical elements of models defining this process using a known polymorphic population of Arctic charr from subarctic Norway. We test the long-term stability of niche specialisation amongst foraging predators and discuss the possibility that contrasting foraging specialists are exposed to differing selection regimes. Inter-individual foraging niche stability was measured by combining two time-integrated ecological tracers of the foraging niche (each individual’s δ13C and δ15N stable isotope (SI) signatures and their food borne parasite fauna) with a short-term measure of foraging niche use (stomach contents composition). Three dietary subgroups of predators were identified, including zooplankton, gammarid and benthivore specialists foragers. Zooplanktivorous specialists had muscle low in δ 13C, a high abundance of parasites transmitted from pelagic copepods, a smaller head, longer snout and a more slender body-form than gammaridivorous specialist individuals which had muscle more enriched in δ 13C and high abundance of parasites transmitted from benthic Gammarus. Benthivorous individuals were intermediate between the other two foraging groups according to muscle SI-signals (δ13C) and loadings of parasites transmitted from both copepods and Gammarus. The close relationship between subgroups identified by stomach contents, time-integrated tracers of niche use (SI and parasites) and functional trophic morphology (niche adaptations) demonstrate a long-term temporally stable niche use of each individual predator. Differential habitat use and contrasting parasite communities and loadings, show differential exposure to different suites of selection pressures for different foraging specialists. Results also show that individual specialisation in trophic behaviour and thus exposure to different suites of selection pressures are stable over time, and thus provide a platform for disruptive selection to operate within this sympatric system.

This comparative study explores how low temperatures affect the mortality and growth of first generation hatchery-reared progeny of subarctic populations of Arctic charr (Salvelinus alpinus L.) and European whitefish (Coregonus lavaretus L.). Replicate fish groups where held under simulated natural light regimes (70°N) at three constant temperatures (1, 3 and 6°C). The mortality of Arctic charr was low (≤1.4%) at all temperature treatments, whereas the mortality of whitefish increased with decreasing temperature from 6% at 6°C to 33% at 1°C. The Arctic charr exhibited higher growth rates than whitefish at all three temperature regimes. All groups of Arctic charr increased in weight, whereas whitefish held at 1°C did not gain weight throughout the experimental period of 133 days. Arctic charr exhibited a large intraspecific variability in growth leading to large variations in size-structure, whereas whitefish in contrast showed very homogenous growth and size-structure patterns; a dissimilarity probably related to species-specific differences in antagonistic behaviour. Evidently, Arctic charr are more cold water adapted than whitefish and are able to maintain growth at extremely low temperatures. Arctic charr thus appear to be the most suitable species for aquaculture at low water temperatures.

The trophic niche and parasite infection of Arctic charr (Salvelinus alpinus) were explored in two lakes with sympatric burbot (Lota lota) and two lakes without burbot in subarctic Norway. The CPUE of burbot and charr were similar in one lake, but burbot had a low population density in the other. Burbot were benthivorous in both lakes. Other co-occurring species like brown trout (Salmo trutta), Atlantic salmon parr (Salmo salar), grayling (Thymallus thymallus) and minnow (Phoxinus phoxinus) were also benthivores. At high densities, benthivorous burbot forced the whole Arctic charr population to utilise mainly the limnetic trophic niche. In contrast, at low burbot density or without burbot present, Arctic charr were primarily benthivorous in the littoral zone. Thus, a clear interactive segregation in diet was observed between Arctic charr and burbot at high burbot densities. There was also a high predation pressure from burbot on young Arctic charr along the benthic zones. The extensive use of zooplankton as prey caused a high parasite infection pressure of copepod transmitted Diphyllobothrium spp. larvae, with the potential for high negative impact on the Arctic charr population. As the benthivore trophic niche was occupied by burbot, the ecological opportunities for polymorphism with benthivorous ecotypes or morphs of Arctic charr were probably prevented. Therefore, the sympatry with burbot seems to have large ecological and evolutionary consequences for this Arctic charr population compared with neighbouring lakes where burbot is absent.

Arctic charr and European whitefish are common species in northern Europe, but have a parapatric distribution pattern and rarely coexist in substantial densities within the same lake. Whitefish is considered to be the strongest competitor, usually causing competitive relegation or exclusion of co-occurring charr. Some exceptions do however occur, and the present study addresses niche utilization and resource partitioning of an Arctic charr population coexisting with whitefish and grayling in a subarctic lake. There was a distinct resource partitioning with respect to habitat and diet utilization of the three species. Grayling were exclusively caught in shallow near-shore areas, feeding chiefly on surface insects and Trichoptera larvae. Whitefish dominated in all lake habitats, feeding predominantly on small-sized planktonic and benthic crustaceans. Arctic charr were mainly found along the benthic profile, feeding predominantly on insects and snails and showing a low dietary overlap with whitefish which appeared to have monopolised the zooplankton resources. The highly restricted trophic niche of Arctic charr suggests a severe competitive impact due to the presence of whitefish, but the coexistence of the two species may be facilitated by the presence of grayling.

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 20 cm. The winter diet of brown trout

There has been an increased focus on Salvelinus alpinus as a potential long-term host to Gyrodactylus salaris and, here, both susceptibility to G. salaris and ability to sustain a parasite population seasonally, was tested using fry and parr of S. alpinus from the River Skibotnelva, northern Norway. Fry were highly susceptible. Gyrodactylus salaris survived on allopatric S. alpinus parr during the 5 month-long winter when water temperatures were c. 1 degrees C. Salvelinus alpinus fry also maintained a pulse of G. salaris infection for over 155 days from early May until autumn. Gyrodactylus salaris are thus able to reproduce and survive on S. alpinus for long periods and at low water temperatures. In spring, newly hatched fry of S. alpinus may serve as an important host to maintain a G. salaris metapopulation within a river system. The results suggest that S. alpinus are adequate long-term hosts of G. salaris independent of the presence of the co-occurring highly susceptible S. salar.

Artificially reared Arctic charr Salvelinus alpinus fry (post yolk-sac stage) from Skibotnelva in northern Norway were exposed to infection with the monogenean ectoparasite Gyrodactylus salaris to determine if the parasite caused mortality in fry. About 95% of the fry became infected within the first week post-exposure, and at the end of the experiment (77 days) all fry carried the parasite. Mortality rate was significantly higher in the groups of infected fry than in the uninfected control groups, and when the experiment was terminated c. 30% of the infected and 8% of the uninfected fish had died. Gyrodactylus salaris had a pronounced negative effect upon fry survival and caused an additive host mortality. Moreover, the concurrent temporal changes in parasite intensities and aggregation indicated that the parasite-induced host mortality was density dependent.

In the past 40 years, there has been increasing acceptance that variation in levels of gene expression represents a major source of evolutionary novelty. Gene expression divergence is therefore likely to be involved in the emergence of incipient species, namely, in a context of adaptive radiation. In the lake whitefish species complex (Coregonus clupeaformis), previous microarray experiments have led to the identification of candidate genes potentially implicated in the parallel evolution of the limnetic dwarf lake whitefish, which is highly distinct from the benthic normal lake whitefish in life history, morphology, metabolism, and behavior, and yet diverged from it only approximately 15,000 years before present. The aim of the present study was to address transcriptional divergence for six candidate genes among lake whitefish and European whitefish (Coregonus lavaretus) species pairs, as well as lake cisco (Coregonus artedi) and vendace (Coregonus albula). The main goal was to test the hypothesis that parallel phenotypic adaptation toward the use of the limnetic niche in coregonine fishes is accompanied by parallelism in candidate gene transcription as measured by quantitative real-time polymerase chain reaction. Results obtained for three candidate genes, whereby parallelism in expression was observed across all whitefish species pairs, provide strong support for the hypothesis that divergent natural selection plays an important role in the adaptive radiation of whitefish species. However, this parallelism in expression did not extend to cisco and vendace, thereby infirming transcriptional convergence between limnetic whitefish species and their limnetic congeners for these genes. As recently proposed (Lynch 2007a. The evolution of genetic networks by non-adaptive processes. Nat Rev Genet. 8:803-813), these results may suggest that convergent phenotypic evolution can result from nonadaptive shaping of genome architecture in independently evolved coregonine lineages.

Gut-tract morphology differed between benthivorous and zooplanktivorous Arctic charr Salvelinus alpinus caught in the littoral and pelagic zones of a sub-Arctic lake. The differences were related to trophic niche and infection with the cestodes Cyathocephalus truncatus and Eubothrium salvelini. Measurements that were unrelated to cestode infection, including intestinal length, numbers of pyloric caeca and posterior pyloric caeca morphology, differed between S. alpinus that had consumed different prey types, suggesting an adaptive response to divergent trophic niches. The anterior pyloric caeca of S. alpinus with high numbers of cestodes were significantly wider and shorter than those in fish with lower levels of infection. The differences were strongly associated with infection by C. truncatus, which was site-selective and attached to the anterior caeca with a large scolex. Differences in caecal measurements were probably the result of mechanical damage caused by scolex attachment and host responses to infection with C. truncatus. The differences in anterior caeca were also an indirect indicator of trophic specialization. Benthivorous S. alpinus were more exposed to, and more heavily infected with C. truncatus than were pelagic zooplanktivorous fish, because this cestode had Gammaruslacustris as its intermediate host. Nevertheless, care should be exercized when using gut measurements for making inferences about the trophic ecology of fish because a history of infection with cestodes may have profound effects on caecal morphology.

The seasonal occurrence of the monogenean ectoparasite Gyrodactylus salaris Malmberg infecting Arctic charr, Salvelinus alpinus (L.) in the two rivers Skibotnelva and Signaldalselva in northern Norway was studied in the period from autumn 2003 to autumn 2005. Skibotnelva has been infected with the parasite since 1979, and treated with rotenone twice. Most likely resident Arctic charr avoided the rotenone treatment in small tributary streams, and thus was the source of the repeated re-infection of this river. G. salaris was first recorded in Signaldalselva in the year 2000 and it is still untreated. Unlike Atlantic salmon, which is highly susceptible to G. salaris, Arctic charr can display a wide range of host-responses to G. salaris infections. Arctic charr were sampled by electro fishing with a total sample of 681 Arctic charr. The results from this study demonstrate an evident seasonal dynamic in G. salaris infection in charr in both rivers. Parasite intensities fluctuated with the rise and fall in temperature through the year, with an autumn high and spring low. There was a significantly lower prevalence and mean intensity of G. salaris in Skibotnelva than in Signaldalselva. There were also a lower prevalence and intensity of G. salaris in the older than in the youngest charr. The different history of infection and treatment in the two rivers might be the underlying cause of these observed dissimilarities. The current study indicates that Arctic charr is a good natural host for G. salaris.

– In 2000, the Atlantic salmon (Salmo salar L.) stock in the river Signaldalselva, North Norway was found to be infected with the monogenean Gyrodactylus salaris (Malmberg). This river system has a multispecies fish community of nine species in total. Low densities of Atlantic salmon parr were found during electrofishing in October (2003) with a prevalence of 94% and a mean abundance of 848 G. salaris. This watercourse also holds a riverine stock of anadromous Arctic charr (Salvelinus alpinus), which is not common in Norway. Among the parr of Arctic charr, 64% were infested with an abundance of 23 parasites per fish. The most heavily infected 0+ and 1+ juvenile Arctic charr had 469 and 534 parasites, respectively, indicating that the parasites are able to reproduce on Arctic charr in the wild. There was little variation in the rates of infestation of Arctic charr along the studied stretch of the river, even in areas with very low densities of Atlantic salmon parr. The parr of anadromous Arctic charr seem to be suitable as long-term hosts for G. salaris. Moreover, a few (22%) infested adult sea-running Arctic charr were captured after ascending the river in the autumn, which shows that large fish may also act as carriers of G. salaris. However, no parasites were recorded on potential anadromous Arctic charr before descending during early spring (April 2003 and April 2004). Thus, it is still not clear whether anadromous Arctic charr are able to disperse the parasite between watercourses. So far, it is uncertain whether G. salaris may increase the mortality rates of Arctic charr and thereby be a threat to these unusual occurring riverine anadromous Arctic charr stocks.

The habitat and diet choice and the infection (prevalence and abundance) of trophically transmitted parasites were compared in Arctic charr and brown trout living sympatrically in two lakes in northern Norway. Arctic charr were found in all main lake habitats, whereas the brown trout were almost exclusively found in the littoral zone. In both lakes the parasite fauna reflected the niche segregation between trout and charr. Surface insects were most common in the diet of trout, but transmit few parasites, and accordingly the brown trout had a relatively low diversity and abundance of parasites. Parasites transmitted by benthic prey such as Gammarus and insect larva, were common in both salmonid host species. Copepod transmitted parasites were much more common in Arctic charr, as brown trout did not include zooplankton in their diets. Parasite species that may use small fish as transport hosts, were far more abundant in piscivorous fish, especially brown trout. The seasonal dynamics in parasite infection were also consistent with the developments in the diet throughout the year. The study demonstrates that the structure of parasite communities of charr and the trout is highly dependent on shifts in habitat and diet of their hosts both on an annual base and through the ontogeny, in addition to the observed niche segregation between the two salmonid species.

Published version available in https://doi.org/10.1007/s10750-015-2589-2 The trophic niches of Arctic charr and brown trout differ when the species occur in sympatry . Their trophically transmitted parasit es are expected to reflect these differences. Here , we investigate how the infections of Diphyllobothrium dendriticum and Diphyllobothrium ditremum differ between charr and trout. These tapeworms use copepods as their first intermediate hos ts and fish can become infected as second intermediate hosts by consuming either infected copepods or infected fish . We examined 767 charr and 368 trout for Diphyllobothrium plerocercoids in a subarctic lake. The prevalence of D. ditremum was higher in charr (61.5 %) than in trout , (39.5 %), but the prevalence of D. dendriticum was higher in trout (31.2 %) than in charr (19.3 %). Diphyllobothrium spp. intensities were elevated in trout compared to charr, particularly for D. dendriticum . Large fish with massive parasite bur dens were responsible for the high Diphyllobothrium spp. loads in trout. We hypothesize that fish prey may be the most important source for the Diphyllobothrium spp. infection s in trout, whereas charr predominantly acquire Diphyllobothrium spp. by feeding on copepods . Our findings support previous suggestions, that the ability to establish in a second piscine host is greater for D. dendriticum than for D. ditremum .

– 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 Fjellfrosvatn, 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.