Search

1. Competition for shared resources is commonly assumed to restrict population-level niche width of coexisting species. However, the identity and abundance of coexisting species, the prevailing environmental conditions, and the individual body size may shape the effects of interspecific interactions on species’ niche width. 2. Here we study the effects of interspecific and intraspecific interactions, lake area and altitude, and fish body size on the trophic niche width and resource use of a generalist predator, the littoral-dwelling large, sparsely rakered morph of European whitefish (Coregonus lavaretus; hereafter LSR whitefish). We use stable isotope, diet and survey fishing data from 14 subarctic lakes along an environmental gradient in northern Norway. 3. The isotopic niche width of LSR whitefish showed a humped-shaped relationship with increasing relative abundance of sympatric competitors, suggesting widest population niche at intermediate intensity of interspecific interactions. The isotopic niche width of LSR whitefish tended to decrease with increasing altitude, suggesting reduced niche in colder, less productive lakes. 4. LSR whitefish typically shifted to a higher trophic position and increased reliance on littoral food resources with increasing body size, although between-lake differences in ontogenetic niche shifts were evident. In most lakes, LSR whitefish relied less on littoral food resources than coexisting fishes and the niche overlap between sympatric competitors was most evident among relatively large individuals (>250 mm). Individual niche variation was highest among >200 mm long LSR whitefish, which likely have escaped the predation window of sympatric predators. 5. We demonstrate that intermediate intensity of interspecific interactions may broaden species’ niche width, whereas strong competition for limited resources and high predation risk may suppress niche width in less productive environments. Acknowledging potential humped-shaped relationships between population niche width and interspecific interactions can help us understand species’ responses to environmental disturbance (e.g. climate change and species invasions) as well as the driving forces of niche specialization.

Competition is assumed to shape niche widths, affecting species survival and coexistence. Expectedly, high interspecific competition will reduce population niche widths, whereas high intraspecific competition will do the opposite. Here we test in situ how intra- and interspecific competition affects trophic resource use and the individual and population niche widths of two lacustrine fish species, Arctic charr and brown trout, covering a 40 year study period with highly contrasting competitive impacts prior to and following a large-scale fish culling experiment. Initially, an overcrowded Arctic charr population dominated the study system, with brown trout being nearly absent. The culling experiment reduced the littoral Arctic charr density by 80%, whereupon brown trout gradually increased its density in the system. Thus, over the study period, the Arctic charr population went from high to low intraspecific competition, followed by increasing interspecific competition with brown trout. As hypothesized, the relaxed intraspecific competition following the experimental culling reduced individual diet specialization and compressed population niche width of Arctic charr. During the initial increase of the brown trout population, there was a large dietary overlap between the two species. Over the subsequent intensified interspecific competition from the population build-up of brown trout, their trophic niche overlap chiefly declined due to a dietary shift of Arctic charr towards enhanced zooplankton consumption. Contrary to theoretical expectations, the individual and population niche widths of Arctic charr increased with intensified interspecific competition. In contrast, the diet and niche width of brown trout remained stable over time, confirming its competitive superiority. The large-scale culling experiment and associated long-term research revealed pronounced temporal dynamics in trophic niche and resource use of the inferior competitor, substantiating that intra- and interspecific competition have large and contrasting impacts on individual and population niches.

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.

The use of translocations to establish new or ‘refuge’ populations for species with high conservation value is controversial but widely used in conservation management. One of the risks of this approach is that an establishing population does not adequately capture the genetic diversity of the donor gene pool. This effect, rarely examined, is tested here. In this study the genetic consequences of two conservation translocations after five generations (16 years) of the European whitefish, Coregonus lavaretus, were quantified. Both translocations were made using almost the same genetic groups and thus represent a partly replicated natural study. Analysis of 12 informative microsatellites showed that expected heterozygosity, the mean number of alleles per locus and allelic richness did not differ between donor and translocated populations. There was also no loss of heterozygosity in the translocated populations, nor deviations from Hardy–Weinberg equilibrium expectations, nor signs of linkage disequilibrium. All populations were genetically differentiated but pairwise FST values were low, indicating that the magnitude of divergence was small. There was no evidence of inbreeding but there were significant differences in private allelic richness between donor and translocated populations. Of 50 alleles found in the donor population, 16% of the rarer alleles were lost in one translocated population and 8% in the other. Allele loss without a reduction in heterozygosity strongly points to stochastic drift effects having occurred following translocation. The evidence indicates that alleles that were not detected in the donor population have arisen de novo in the translocated populations. It is concluded that conservation translocations comprising even a modest number of propagules can successfully capture a high proportion of genetic variation of the host population, and that reduced genetic variation in the translocated population may be mitigated by the emergence of new variation over short time periods.

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.

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.

The European whitefish Coregonus lavaretus complex represents one of the most diverse radiations within salmonids, with extreme morphological and genetic differentiation across its range. Such variation has led to the assignment of many populations to separate species. In Great Britain, the seven native populations of C. lavaretus (two in Scotland, four in England, one in Wales) were previously classified into three species, and recent taxonomic revision resurrected the previous nomenclature. Here we used a dataset of 15 microsatellites to: (1) investigate the genetic diversity of British populations, (2) assess the level of population structure and the relationships between British populations. Genetic diversity was highest in Welsh (HO = 0.50, AR = 5.29), intermediate in English (HO = 0.41–0.50, AR = 2.83–3.88), and lowest in Scottish populations (HO = 0.28–0.35, AR = 2.56–3.04). Population structure analyses indicated high genetic differentiation (global FST = 0.388) between all populations but for the two Scottish populations (FST = 0.063) and two English populations (FST = 0.038). Principal component analysis and molecular ANOVA revealed separation between Scottish, English, and Welsh populations, with the Scottish populations being the most diverged. We argue that the data presented here are not sufficient to support a separation of the British European whitefish populations into three separate species, but support the delineation of different ESUs for these populations.

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

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.

Catchment-scale variation between lake habitats has the potential to simultaneously influence the trophic niche and parasite community of fish hosts. In this study, we investigated the trophic niche and parasite community of sympatric Arctic charr and brown trout populations from two inter-connected southern Norwegian lakes at different altitudes. Arctic charr and brown trout occupied profundal and littoral habitats in each lake, respectively, whereas brown trout replaced Arctic charr in pelagic habitats of the lower altitude lake. Distinct between-lake differences in diet and parasite community composition were noted for brown trout; however, both fish species showed highly overlapping trophically transmitted parasite communities regardless of the habitats each species used. Our results suggest that environmental differences over relatively limited geographical distances have the potential to influence fish habitat use and parasite community structure.

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/s10750-019-3969-9. Parallelism in trophic niches of polymorphic populations of Arctic charr was investigated in two similar subarctic lakes, Tårnvatn and Skøvatn, in northern Norway. Analysis of eleven microsatellite loci confirmed, respectively, the existence of three and two genetically differentiated morphs. Three methods were used to describe their trophic niches: habitat choice and stomach contents for the recent feeding behaviour, and trophically transmitted parasites and stable isotopes (δ13C and δ15N) as proxies for the longer term trophic niche differences. The results showed a distinct segregation in trophic resource utilization of the different morphs. Tårnvatn has three morphs: a littoral omnivorous (LO), a small-sized profundal benthivorous (PB), and a large-sized profundal piscivorous (PP). In contrast, a novel Arctic charr morph was discovered in Skøvatn: a small-sized profundal zooplanktivorous-morph (PZ), which when compared to the sympatric LO-morph, had distinct stable isotope values and a contrasting parasite community. A parallelism in habitat choice and external morphology was found among the small-sized, deep-water morphs and between the upper-water, omnivorous LO-morphs in both lakes. There was a no parallel pattern in diet choice between the PB- and the PZ-morphs. These findings show how evolution can produce diverse outcomes, even among systems with apparently similar environmental and ecological conditions.

A major goal of evolutionary science is to understand how biological diversity is generated and altered. Despite considerable advances, we still have limited insight into how phenotypic variation arises and is sorted by natural selection. Here we argue that an integrated view, which merges ecology, evolution and developmental biology (eco evo devo) on an equal footing, is needed to understand the multifaceted role of the environment in simultaneously determining the development of the phenotype and the nature of the selective environment, and how organisms in turn affect the environment through eco evo and eco devo feedbacks. To illustrate the usefulness of an integrated eco evo devo perspective, we connect it with the theory of resource polymorphism (i.e. the phenotypic and genetic diversification that occurs in response to variation in available resources). In so doing, we highlight fishes from recently glaciated freshwater systems as exceptionally well‐suited model systems for testing predictions of an eco evo devo framework in studies of diversification. Studies on these fishes show that intraspecific diversity can evolve rapidly, and that this process is jointly facilitated by (i) the availability of diverse environments promoting divergent natural selection; (ii) dynamic developmental processes sensitive to environmental and genetic signals; and (iii) eco evo and eco devo feedbacks influencing the selective and developmental environments of the phenotype. We highlight empirical examples and present a conceptual model for the generation of resource polymorphism – emphasizing eco evo devo, and identify current gaps in knowledge., Biological Reviews, 94 (5), ISSN:1469-185X, ISSN:1464-7931, ISSN:0006-3231

Interspecific morph variations in trophic morphology related to skull-bones and head traits is associated to ecological segregation of Arctic charr morphs (genus Salvelinus) in two sub-arctic lakes (Fjellfrosvatn and Skogsfjordvatn, Norway). The replicated morph pair, the profundal spawning benthivorous PB-morph and the littoral spawning omnivorous LO-morph of Arctic charr, diverge along the shallow-deep-water resource axis. In Skogsfjordvatn there is also a profundal spawning piscivorous PP-morph. The PB-morphs from both lakes have similar skull-bone traits and head morphology such as elongated jaw-bones, small opercular bones and relatively longer heads. The PP-morph also has an elongated head, relatively small opercular bones as well as larger jaw-bones. In contrast, the LO-morphs in both lakes have shorter jaw-bones, larger opercular bones in addition to relatively small heads. However, some small non-parallel differences exist among the morphs from the two lakes. Overall, all profundal morphs (PB and PP) have relatively similar skull-bone structures, suggesting adaptations to the deep-water environment but also to their separated dietary niches. There is strong evidence for parallel evolution with some local adaptations in skull-bones and head morphology of the PB-morph and the LO-morph from separate lakes.

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.

Parasite communities can show large differences in species composition between sympatric host species. Here, we assessed how divergent resource use of polymorphic populations of Arctic charr from two environmentally similar sub-arctic lakes was related to parasite community composition. Large similarities in parasite infections were found for both the reproductively isolated littoral-spawning omnivore LO-morph and the profundal-spawning benthivore PB-morph, reflecting cross-lake similarities in resource use. Furthermore, whereas the PB-morph had the lowest richness and abundance of parasite species in both lakes, a third morph, the profundal-spawning piscivorous PP-morph (one lake only), had the highest parasite richness and accumulated parasites by preying upon fish. These results highlight that host ecology and abiotic factors are important for structuring parasite communities. The dissimilarities in the local parasite species pool between the lakes suggest that stochastic events as well as transmission abilities are involved in structuring the parasite communities. The parallel divergent parasite infections between morphs may form an arena for parasite-mediated selection promoting the incipient speciation process in concert with other factors. The deep-water PB-morphs may avoid parasites by specialising in parasite-poor but low-productive habitats, while the piscivorous PP-morph aggregates infections of detrimental parasites through specialising on energetically rich but generally more heavily parasitised prey.

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.

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.

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.

Submitted manuscript version. Published version available at https://doi.org/10.1016/j.tree.2018.01.011. Species interactions can influence ecosystem functioning by enhancing or suppressing the activities of species that drive ecosystem processes, or by causing changes in biodiversity. However, one important class of species interactions – parasitism – has been little considered in biodiversity and ecosystem functioning (BD-EF) research. Parasites might increase or decrease ecosystem processes by reducing host abundance. Parasites could also increase trait diversity by suppressing dominant species or by increasing within-host trait diversity. These different mechanisms by which parasites might affect ecosystem function pose challenges in predicting their net effects. Nonetheless, given the ubiquity of parasites, we propose that parasite–host interactions should be incorporated into the BD-EF framework.

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.

This article is also available at onlinelibrary.wiley.com Understanding how a monophyletic lineage of a species diverges into several adaptive forms has received increased attention in recent years, but the underly- ing mechanisms in this process are still under debate. Postglacial fishes are excellent model organisms for exploring this process, especially the initial stages of ecological speciation, as postglacial lakes represent replicated discrete envi- ronments with variation in available niches. Here, we combine data of niche utilization, trophic morphology, and 17 microsatellite loci to investigate the diversification process of three sympatric European whitefish morphs from three northern Fennoscandian lakes. The morphological divergence in the gill raker number among the whitefish morphs was related to the utilization of dif- ferent trophic niches and was associated with reproductive isolation within and across lakes. The intralacustrine comparison of whitefish morphs showed that these systems represent two levels of adaptive divergence: (1) a consistent littoral – pelagic resource axis; and (2) a more variable littoral – profundal resource axis. The results also indicate that the profundal whitefish morph has diverged repeatedly from the ancestral littoral whitefish morph in sympatry in two different watercourses. In contrast, all the analyses performed revealed clus- tering of the pelagic whitefish morphs across lakes suggesting parallel postglacial immigration with the littoral whitefish morph into each lake. Finally, the analy- ses strongly suggested that the trophic adaptive trait, number of gill rakers, was under diversifying selection in the different whitefish morphs. Together, the results support a complex evolutionary scenario where ecological speciation acts, but where both allopatric (colonization history) and sympatric (within watercourse divergence) processes are involved.

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.

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.

Accepted manuscript version. Published version available in Temporal changes and between-host variation in the intestinal parasite community of Arctic charr in a subarctic lake (2016) 783(1), s.79-91. . The occurrence of trophically transmitted intestinal parasites in Arctic charr was analyzed from data material collected over two decades from Lake Takvatn, northern Norway. The main objectives were to investigate (i) between-year variation in parasite infracommunity composition, (ii) between-host variation in infracommunity composition, and (iii) possible interspecific interaction between parasites. The trematode Crepidostomum spp. and the cestode Eubothrium salvelini were most prevalent and had the highest mean intensities whereas the cestodes Proteocephalus sp. and Cyathocephalus truncatus were uncommon taxa. No clear indication of interspecific parasite interactions was evident. By far the most frequent infracommunity composition was E. salvelini co-existing with only Crepidostomum spp. Despite some indications of individual dietary specialization in Arctic charr, this strong species cooccurrence suggests that the majority of fish had been preying on the very different intermediate hosts of both parasites. Overall, even though some sporadic between-year variation was present, no long-term or cyclical variation in infracommunity composition and mean intensity was revealed, suggesting that the host– parasite system is in a steady state. Minor between-year variation was likely the result of changes in habitat and dietary utilization by Arctic charr, possibly linked to documented changes in the fish community structure of the lake.

Published version available in http://dx.doi.org/10.1127/fal/2016/0839 . Benthic invertebrate communities play a fundamental role in lake ecosystems, and the understanding of how those benthic communities are structured, particularly in terms of the identity and spatiotemporal distribution of their functional traits, is key to our understanding of how lake ecosystems work. In Takvatn, a subarctic lake in northern Norway, we identified the taxonomic and functional identity of the species characterizing benthic communities across three seasons and six different depths. Invertebrates were sampled using vacuum pump and Ekman grab. Despite the relative low species richness, we found large variation across depths and seasons in species functional composition using multivariate analyses. In all seasons and habitats, gathering traits were the most common feeding traits within the benthic community. The upper littoral habitat shifted from a community characterized by particle gathers and algal scrapers in spring and summer, to a community largely characterized by leaf shredders in autumn. There was also high relative occurrence of predator traits towards the deeper areas of the littoral zone, as well as in the sub-littoral and profundal habitats. The deeper habitats showed high variation in their taxonomic composition, but a stable composition of functional traits throughout all seasons. In conclusion, our study demonstrates large spatio-temporal differences in functional trait diversity and composition of the benthic invertebrates in Takvatn.

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.

Growth, feed intake and maturation were monitored in a fast-growing benthivore morph of European whitefish [Coregonus lavaretus (L.)] held under constant light (LD 24:0) and temperature conditions (10°C) for 415 days. Their growth was best described by polynomial regression model and no indication of seasonal patterns (e.g. winter depression) in growth rates was found. There was no significant difference in food conversion ratio (FCR) between periods during the experiment and the FCR was favourable (0.84). Furthermore, there were no differences in growth between males and females. None of the fish became sexually mature by the time the experiment was terminated, although they had reached a mean size of about 444 g. The European whitefish seems to be a promising aquaculture species as the growth rate was relatively high with a remarkable homogenous growth performance between individual fish within each experimental unit, a relatively favourable FCR, a low degree of agonistic behaviour and negligible growth depression due to internal biological rhythms or gonad maturation. These results were obtained at relatively low temperatures that should be beneficial for sustainable cost-effective farming conditions in cold climate condition in northern Scandinavia.

– 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.

– We explored the incidence of individual feeding specialisation among a naive predator (non-native rainbow trout postsmolts) and two native experienced predators (sea-run Arctic charr and sea-trout) in a subarctic Norwegian fjord. Interindividual foraging niche stability was obtained by combining information on stomach contents (recent dietary niche) with trophically transmitted parasite infestation (time-integrated historical dietary niche) of individual predators. Individual fish showed a high degree of resource specialisation as prey items such as gammarids and small fish (both potential intermediate host of parasites) rarely co-occurred in stomachs. In both naive and veteran predators, positive associations between the intensity of a specific parasite species and the occurrence of their respective intermediate host (gammarids or fish) in the stomachs of individual predators demonstrated temporally interindividual feeding specialisations. Several behavioural phenotypes clearly co-existed in both naive and veteran predator populations, including gammaridivore (benthic feeders), piscivore (pelagic feeders) or insectivore (pleuston feeders) individuals. The likely mechanism of this observed interindividual resource specialisation in the non-native naive predators involves a behavioural component of which rapid learning seems to be a key factor.