Sympatric Arctic Charr Research Articles

Temporal stability of polymorphic Arctic charr parasite communities reflects sustained divergent trophic niches.

Polymorphic Arctic charr Salvelinus alpinus populations frequently display distinct differences in habitat use, diet, and parasite communities. Changes to the relative species densities and composition of the wider fish community have the potential to alter the habitat niche of sympatric Arctic charr populations. This study evaluated the temporal stability of the parasite community, diet, and stable isotopes (δ13C, δ15N) of three sympatric Arctic charr morphs (piscivore, benthivore, and planktivore) from Loch Rannoch, Scotland, in relation to changes to the fish community. All Arctic charr morphs displayed distinct differences in parasite communities, diet, and stable isotope signatures over time, despite the establishment of four new trophically transmitted parasite taxa, and increased fish and zooplankton consumption by the piscivorous and planktivore morphs, respectively. Native parasite prevalence also increased in all Arctic charr morphs. Overall, Loch Rannoch polymorphic Arctic charr morph populations have maintained their distinct trophic niches and parasite communities through time despite changes in the fish community. This result indicates that re-stocking a native fish species has the potential to induce shifts in the parasite community and diet of Arctic charr morphs.

Environmentally Modulated Repeat Evolution of Polymorphic Arctic Charr Life History Traits

Sympatric Arctic charr, Salvelinus alpinus (L. 1758), morphs have flexible but repeated life history strategies tested across five Norwegian lakes. In several Scandinavian polymorphic Arctic charr populations differentiated by their diet and habitat use, a large littoral omnivorous (LO) morph commonly cooccurs with a smaller profundal spawning (PB/PZ) morph. A third, large piscivorous (PP) morph is also known to occur within a portion of Arctic charr populations in the profundal habitat along with the PB/PZ individuals. Life history traits, such as age at maturity, growth, and diet are known to differ among coexisting morphs. Notably, the PP morph was the longest morph with the oldest age at maturity while the PB/PZ morph showed the shortest lengths overall and youngest age with LO morph being intermediate in both traits. Growth parameters differed across all the morphs. When examining growth within morph groups, the LO morph was found to have different growth across all lakes, while similar reproductive investments and different energy acquisition patterns were seen within the PB/PZ and PP morphs. These results suggest repeat evolution in several life history strategies of reproductively isolated Arctic charr sympatric morphs, notably for the first time in the PP morph, while also highlighting the importance of the local environment in modulating life history traits.

Multivariate analysis of morphology, behaviour, growth and developmental timing in hybrids brings new insights into the divergence of sympatric Arctic charr morphs

BackgroundStudying the development of fitness related traits in hybrids from populations diverging in sympatry is a fundamental approach to understand the processes of speciation. However, such traits are often affected by covariance structures that complicate the comprehension of these processes, especially because the interactive relationships between traits of different nature (e.g. morphology, behaviour, life-history) remain largely unknown in this context. In a common garden setup, we conducted an extensive examination of a large suit of traits putatively involved in the divergence of two morphs of Arctic charr (Salvelinus alpinus), and investigated the consequences of potential patterns of trait covariance on the phenotype of their hybrids. These traits were measured along ontogeny and involved growth, yolk sac resorption, developmental timing (hatching and the onset of exogeneous feeding), head morphology and feeding behaviour.ResultsGrowth trajectories provided the strongest signal of phenotypic divergence between the two charr. Strikingly, the first-generation hybrids did not show intermediate nor delayed growth but were similar to the smallest morph, suggesting parental biases in the inheritance of growth patterns. However, we did not observe extensive multivariate trait differences between the two morphs and their hybrids. Growth was linked to head morphology (suggesting that morphological variations in early juveniles relate to simple allometric effects) but this was the only strong signal of covariance observed between all the measured traits. Furthermore, we did not report evidence for differences in overall phenotypic variance between morphs, nor for enhanced phenotypic variability in their hybrids.ConclusionOur study shed light on the multivariate aspect of development in a context of adaptive divergence. The lack of evidence for the integration of most traits into a single covariance structure suggested that phenotypic constraints may not always favour nor impede divergence toward ecological niches differing in numerous physical and ecological variables, as observed in the respective habitats of the two charr. Likewise, the role of hybridization as a disruptive agent of trait covariance may not necessarily be significant in the evolution of populations undergoing resource polymorphism.

Extensive genetic differentiation between recently evolved sympatric Arctic charr morphs.

The availability of diverse ecological niches can promote adaptation of trophic specializations and related traits, as has been repeatedly observed in evolutionary radiations of freshwater fish. The role of genetics, environment, and history in ecologically driven divergence and adaptation, can be studied on adaptive radiations or populations showing ecological polymorphism. Salmonids, especially the Salvelinus genus, are renowned for both phenotypic diversity and polymorphism. Arctic charr (Salvelinus alpinus) invaded Icelandic streams during the glacial retreat (about 10,000 years ago) and exhibits many instances of sympatric polymorphism. Particularly, well studied are the four morphs in Lake Þingvallavatn in Iceland. The small benthic (SB), large benthic (LB), planktivorous (PL), and piscivorous (PI) charr differ in many regards, including size, form, and life history traits. To investigate relatedness and genomic differentiation between morphs, we identified variable sites from RNA‐sequencing data from three of those morphs and verified 22 variants in population samples. The data reveal genetic differences between the morphs, with the two benthic morphs being more similar and the PL‐charr more genetically different. The markers with high differentiation map to all linkage groups, suggesting ancient and pervasive genetic separation of these three morphs. Furthermore, GO analyses suggest differences in collagen metabolism, odontogenesis, and sensory systems between PL‐charr and the benthic morphs. Genotyping in population samples from all four morphs confirms the genetic separation and indicates that the PI‐charr are less genetically distinct than the other three morphs. The genetic separation of the other three morphs indicates certain degree of reproductive isolation. The extent of gene flow between the morphs and the nature of reproductive barriers between them remain to be elucidated.

Exploring trophic niches and parasite communities of sympatric Arctic charr and brown trout populations of southern Norway

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.

Reproductive strategies of Arctic charr Salvelinus alpinus (L.) forms in Kiryalta lakes, Transbaikalia, Russia

The divergence of reproductive strategies is an important development toward intraspecific diversification. We describe contrasting spawning patterns of parapatric and sympatric Arctic charr forms in two interconnected lakes Kiryalta-3 and Kiryalta-4 in Transbaikalia for comparison to charr reproduction in nine more lakes of this region. In each Kiryalta lake, two charr forms (small and large) differing in growth and diet are present, in Kiryalta-3 they are morphologically more distinct than in Kiryalta-4. In Kiryalta-3, the spawning of the small form was recorded in autumn and spring suggesting two subpopulations; spawning time of the large form is unknown. In Kiryalta-4, both forms spawn in autumn. Overall, reproductive patterns of Transbaikalian charr vary greatly in spawning time, duration, location, depth, and bottom substrate. Different populations and sympatric forms spawn in early or late autumn, spring, summer, or during more than one season, including the littoral or profundal zone, over rocks or silt, and from 1–2 weeks to at least 7 months. Peak spawning time difference between sympatric forms can reach up to 5 months, and is correlated with morphological and genetic differences. Diversification of reproductive strategies largely contributed to the emergence of exceptionally high diversity of Transbaikalian charr.

Differential gene expression during early development in recently evolved and sympatric Arctic charr morphs.

Phenotypic differences between closely related taxa or populations can arise through genetic variation or be environmentally induced, leading to altered transcription of genes during development. Comparative developmental studies of closely related species or variable populations within species can help to elucidate the molecular mechanisms related to evolutionary divergence and speciation. Studies of Arctic charr (Salvelinus alpinus) and related salmonids have revealed considerable phenotypic variation among populations and in Arctic charr many cases of extensive variation within lakes (resource polymorphism) have been recorded. One example is the four Arctic charr morphs in the ∼10,000 year old Lake Thingvallavatn, which differ in numerous morphological and life history traits. We set out to investigate the molecular and developmental roots of this polymorphism by studying gene expression in embryos of three of the morphs reared in a common garden set-up. We performed RNA-sequencing, de-novo transcriptome assembly and compared gene expression among morphs during an important timeframe in early development, i.e., preceding the formation of key trophic structures. Expectedly, developmental time was the predominant explanatory variable. As the data were affected by some form of RNA-degradation even though all samples passed quality control testing, an estimate of 3′-bias was the second most common explanatory variable. Importantly, morph, both as an independent variable and as interaction with developmental time, affected the expression of numerous transcripts. Transcripts with morph effect, separated the three morphs at the expression level, with the two benthic morphs being more similar. However, Gene Ontology analyses did not reveal clear functional enrichment of transcripts between groups. Verification via qPCR confirmed differential expression of several genes between the morphs, including regulatory genes such as AT-Rich Interaction Domain 4A (arid4a) and translin (tsn). The data are consistent with a scenario where genetic divergence has contributed to differential expression of multiple genes and systems during early development of these sympatric Arctic charr morphs.

Divergent functional traits in three sympatric Arctic charr Salvelinus alpinus morphs are not coupled with the age of the lineage divergence

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.

Visual pigments of Arctic charr (Salvelinus alpinus (L.)) and whitefish (Coregonus lavaretus (L.)) morphs in subarctic lakes

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.

Differential expression of the aryl hydrocarbon receptor pathway associates with craniofacial polymorphism in sympatric Arctic charr.

BackgroundThe developmental basis of craniofacial morphology hinges on interactions of numerous signalling systems. Extensive craniofacial variation in the polymorphic Arctic charr, a member of the salmonid family, from Lake Thingvallavatn (Iceland), offers opportunities to find and study such signalling pathways and their key regulators, thereby shedding light on the developmental pathways, and the genetics of trophic divergence.ResultsTo identify genes involved in the craniofacial differences between benthic and limnetic Arctic charr, we used transcriptome data from different morphs, spanning early development, together with data on craniofacial expression patterns and skeletogenesis in model vertebrate species. Out of 20 genes identified, 7 showed lower gene expression in benthic than in limnetic charr morphs. We had previously identified a conserved gene network involved in extracellular matrix (ECM) organization and skeletogenesis, showing higher expression in developing craniofacial elements of benthic than in limnetic Arctic charr morphs. The present study adds a second set of genes constituting an expanded gene network with strong, benthic–limnetic differential expression. To identify putative upstream regulators, we performed knowledge-based motif enrichment analyses on the regulatory sequences of the identified genes which yielded potential binding sites for a set of known transcription factors (TFs). Of the 8 TFs that we examined using qPCR, two (Ahr2b and Ap2) were found to be differentially expressed between benthic and limnetic charr. Expression analysis of several known AhR targets indicated higher activity of the AhR pathway during craniofacial development in benthic charr morphotypes.ConclusionThese results suggest a key role of the aryl hydrocarbon receptor (AhR) pathway in the observed craniofacial differences between distinct charr morphotypes.Electronic supplementary materialThe online version of this article (doi:10.1186/s13227-015-0022-6) contains supplementary material, which is available to authorized users.

Bones in motion: Ontogeny of craniofacial development in sympatric arctic charr morphs.

The impressive diversity in the feeding apparatus often seen among related fish species clearly reflects differences in feeding modes and habitat utilization. Such variation can also be found within species. One example of such intraspecific diversity is the Arctic charr in Lake Thingvallavatn, where four distinct morphs coexist: two limnetic, with evenly protruding jaws, and two benthic, with subterminal lower jaws. We used these recently evolved morphs to study the role of ontogenetic variation in shaping craniofacial diversity. The segmental development of the pharyngeal arches and the order of events in craniofacial development is the same as has been described for teleosts, emphasizing the conserved nature of this process. However, our morphometric analyses reveal differences between morphs. Hatching is accompanied by increase in size and allometric shape changes in Arctic charr. Ontogenetic trajectories of craniofacial shape also differ significantly between morphs. The results point to hatching as a significant developmental event in Arctic charr and possibly other fishes. Also, the developmental origins of limnetic and benthic specializations in the craniofacial elements of Arctic charr may stem from events around hatching. This calls for investigations of the mechanisms and consequences of hatching, in the context of development and evolution. Developmental Dynamics 244:1168-1178, 2015. © 2015 Wiley Periodicals, Inc.

Divergent growth patterns between juveniles of two sympatric Arctic charr morphs with contrasting depth gradient niche preferences

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.

Water temperatures influence the marine area use of Salvelinus alpinus and Salmo trutta.

The migratory behaviour and spatial area use of sympatric Arctic charr Salvelinus alpinus and brown trout Salmo trutta were investigated during their marine feeding migration. The likelihood of finding individuals of both species in the inner or outer fjord areas was dependent on water temperature in the inner area (especially for S. alpinus), the temperature difference between the inner and outer areas (especially for S. trutta) and fish fork length (both species). The strongest predictor was the water temperature in the inner area, and particularly S. alpinus left this area and moved to the outer areas with increasing temperatures in the inner area. At 8° C in the inner area, the likelihood of finding S. alpinus in the outer areas was >50%. This predictor had a smaller effect on S. trutta, and the likelihood of finding S. trutta in the outer areas only started to increase at around 14° C. The relationships between temperature and area use did not correspond to the species' optimal growth temperatures, but to their previously documented temperature preferences. Individuals of both species used mainly the littoral fjord areas, and to a lesser extent the pelagic areas. In conclusion, temperature differences between the inner and outer marine areas probably resulted in the segregated area use between the species, because water temperatures or factors influenced by temperature affected their migratory behaviour and habitat use differently. The results indicate that increased marine temperatures with global warming may lead to increased spatial overlap between S. trutta and S. alpinus, which again may lead to increased interspecific competition during their marine phase, and with S. alpinus probably being the more negatively affected.

Contrasting life history strategies of sympatric Arctic charr morphs, Salvelinus alpinus

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.

Habitat use and diet of sympatric Arctic charr (Salvelinus alpinus) and whitefish (Coregonus lavaretus) in five lakes in southern Norway: not only interspecific population dominance?

Stable coexistence of Arctic charr and whitefish does occur in a number of native lake fish communities in Scandinavia. Even so, whitefish introductions into Arctic charr lakes have resulted in serious decline and possibly local extinction of Arctic charr. In this article, we analyze the habitat use and diet of the two species in five Norwegian lakes differing in basin shape and environmental conditions. In two of the lakes, both species are native, and appear to live in a relatively stable coexistence. Here, whitefish mainly occupy the littoral and upper pelagic zone, while Arctic charr live in the deeper habitats. Diets are generally quite different in terms of the zooplankton species eaten. In the three other lakes, either whitefish or both species have been introduced. In the shallowest lake, habitat segregation is similar to that seen in the pristine lakes, although Arctic charr appears to be on the brink of extinction. In the remaining two lakes, however, Arctic charr dominates, and occurs in higher numbers than whitefish in all the habitats. Our observations indicate that coexistence of the two species in oligotrophic and relatively pristine lakes requires an extensive profundal zone to serve as a refugium for Arctic charr. If the littoral zone is rendered inaccessible or unprofitable for whitefish due to dominance of a third competitor or predator, or as a result of lake regulation, then Arctic charr may be the dominant species.

Late ontogeny growth acceleration and size form transformations in Transbaikalian Arctic charr, Salvelinus alpinus complex: evidence from fin ray cross section growth layers

In some polymorphic populations of Arctic charr in Transbaikalia, an individual can transform from a smaller to larger size form during their lifetime as a result of accelerated growth that follows a period of slow growth and reproduction as a small size form. Alternating periods of slow and fast growth are reflected in growth layer patterns visible in fin ray cross sections. Stained microtome fin ray cross sections were used to reveal the incidence of transformations from one form to another. Data were collected from 14 northern Transbaikalian lakes containing two or three sympatric Arctic charr forms (‘dwarf’, ‘small’, and ‘large’) exhibiting varying levels of morphological separation. Individuals recruited from the dwarf or small form were found in varying proportions among the small and/or large form in 12 lakes. Small or large form charr that grew without noticeable acceleration to the adult size typical of the form or experienced accelerated growth as juveniles prior to maturation were also observed. There were no transformations between sympatric forms that differed in the length and number of gill rakers and in some other meristic characters. Results indicate that in the region under study, transformations of sympatric Arctic charr size forms are a widespread, but not a ubiquitous, phenomenon. Such transformations reflect the plasticity of the developmental channels of the forms. In the course of intra-lacustrine form divergence and genetic differentiation, the frequency of the observed transformations decreases to zero.

Genetic divergence between sympatric Arctic charr Salvelinus alpinus morphs in Gander Lake, Newfoundland: roles of migration, mutation and unequal effective population sizes

A suite of 10 microsatellite loci was used to examine genetic divergence between two sympatric morphs of Arctic charr Salvelinus alpinus (‘dark’ and ‘pale’) inhabiting Gander Lake, Newfoundland. Results can be summarized as follows: (1) the morphs are strongly reproductively isolated – gene flow–migration estimates were consistently low in long and short‐term evolutionary timescales of analysis; (2) intermorph divergence based on allele size (RST) was significantly larger than those based on allele state (θ) implying a cumulative effect of stepwise‐like mutations; (3) historical (coalescent) and current (linkage disequilibrium) point estimates of effective population size (Ne) were consistently higher for dark than for pale S. alpinus. The first and second findings lend support to the hypothesis that divergence between forms may have preceded the last glacial period (ending c. 12 000 years bp). The third finding argues for significant differences in habitat quantity and quality between morphs, which were emphasized in a previous study. Overall, these analyses underscore the importance of genetic assessment and monitoring in the conservation of fish diversity, with emphasis on ‘rare’ or under‐represented forms among temperate species pairs.

Factors predicting male fertilization success in an external fertilizer

In postcopulatory sexual selection both sperm competition and cryptic female choice are considered to be important selective agents, but their relative importance for male fertilization success has received little attention. We tested whether sperm quality, male spawning coloration, male heterozygosity, and genetic overlap with the female explained a male’s fertilization success in controlled in vitro fertilization competition trials between equal numbers of sperm from pairs of male Arctic charr (Salvelinus alpinus), an external fertilizer. Offspring were genotyped to determine each males’ share of paternity. The velocity of a male’s sperm relative to the velocity of the competing male’s sperm was the best predictor of male fertilization success. Yet, sperm velocity was not related to spawning coloration or male heterozygosity. In fact, the most brightly colored male in a pair had the lowest fertilization probability. This could result from cryptic female choice for pale males, but might rather be a result of paler males producing more competitive sperm than more colored males. Furthermore, the more microsatellite alleles a male shared with the female relative to the competing male, the higher fertilization success he had. We argue that this latter may be an effect of assortative cryptic female choice, which might prevent hybridization with sympatric Arctic charr morphs or one form of kin selection.

Summer stream habitat partitioning by sympatric Arctic charr, Atlantic salmon and brown trout in two sub‐arctic rivers

Summer habitat use by sympatric Arctic charr Salvelinus alpinus, young Atlantic salmon Salmo salar and brown trout Salmo trutta was studied by two methods, direct underwater observation and electrofishing, across a range of habitats in two sub‐arctic rivers. More Arctic charr and fewer Atlantic salmon parr were observed by electrofishing in comparison to direct underwater observation, perhaps suggesting a more cryptic behaviour by Arctic charr. The three species segregated in habitat use. Arctic charr, as found by direct underwater observation, most frequently used slow (mean ±s.d. water velocity 7·2 ± 16·6 cm s−1) or often stillwater and deep habitats (mean ±s.d. depth 170·1 ± 72·1 cm). The most frequently used mesohabitat type was a pool. Young Atlantic salmon favoured the faster flowing areas (mean ±s.d. water velocity 44·0 ± 16·8 cm s−1 and depth 57·1 ± 19·0 cm), while brown trout occupied intermediate habitats (mean ±s.d. water velocity 33·1 ± 18·6 cm s−1 and depth 50·2 ± 18·0 cm). Niche overlap was considerable. The Arctic charr observed were on average larger (total length) than Atlantic salmon and brown trout (mean ±s.d. 21·9 ± 8·0, 10·2 ± 3·1 and 13·4 ± 4·5 cm). Similar habitat segregation between Atlantic salmon and brown trout was found by electrofishing, but more fishes were observed in shallower habitats. Electrofishing suggested that Arctic charr occupied habitats similar to brown trout. These results, however, are biased because electrofishing was inefficient in the slow‐deep habitat favoured by Arctic charr. Habitat use changed between day and night in a similar way for all three species. At night, fishes held positions closer to the bottom than in the day and were more often observed in shallower stream areas mostly with lower water velocities and finer substrata. The observed habitat segregation is probably the result of interference competition, but the influence of innate selective differences needs more study.

Seasonal and ontogenetic variations in resource use by two sympatric Arctic charr morphs

The study compares the resource utilization of two sympatric Arctic charr morphs over an annual period in a subarctic lake. The two morphs are reproductively isolated in time and place of spawning, and are referred to as the littoral and profundal morphs (L-morph and P-morph) according to their spawning habitats. Fish were sampled monthly (ice-free season) or bimonthly (winter) using gillnets in the main lake habitats. The spatial range of the P-morph was restricted to the profundal zone throughout the whole annual period. The L-morph in contrast utilized all main habitats, exhibiting distinct seasonal and ontogenetic variations in habitat distribution. In the spring, the whole L-morph population was located along the bottom profile of the lake, in profundal and littoral habitats. During summer and autumn, habitat segregation occurred between different life-stages, juveniles mainly utilizing the profundal, pre-adults the pelagic and adult fishes the littoral zone. During winter the whole population was assembled in the littoral habitat. The L-morph also had large seasonal and ontogenetic variations in their feeding ecology, with littoral zoobenthos, zooplankton and surface insects being important prey. The P-morph had a narrower diet niche mainly consisting of chironomid larvae and other profundal zoobenthos. Hence, the two Arctic charr morphs exhibited a consistent resource differentiation during all annual seasons and throughout their life cycles, except for a dietary overlap between P-morph and juvenile L-morph charr in the profundal during summer. The findings are discussed in relation to resource polymorphism and incipient speciation.