Mining various genomic resources to resolve old alpha-taxonomy questions: A test of the species hypothesis of the Proteocephalus longicollis species complex (Cestoda: Platyhelminthes) from salmonid fishes.

High-throughput sequencing strategies became commonly employed to study non-model parasites, but the corresponding genomes and transcriptomes were seldom mined following the original publication. Similar to the data generated with genome skimming techniques based on shallow-depth shotgun genomes, various genomic and transcriptomic resources can be screened for useful molecular phylogenetic markers traditionally characterised with Sanger sequencing. Here, we provide an example of a strategy using reduced-representation genomic as well as transcriptomic data to obtain broad insights into the molecular diversity of the cestode Proteocephalus longicollis, a common parasite of salmonids distributed throughout the Holarctic region. We extract popular mitochondrial and nuclear ribosomal markers from various genomic resources for hundreds of parasite specimens from multiple European whitefish populations and compare those with Proteocephalus representatives from other species of salmonids and various geographical regions. In contrast with the previous morphology-based assessments, molecular phylogeny reveals a high degree of genetic divergence between Proteocephalus isolates from different salmonids, contrastingly low genetic differentiation within the parasite's populations hosted by the European whitefish (Coregonus lavaretus species complex), and a sister species relationship of Proteocephalus from European whitefish and Proteocephalus percae, a parasite of European perch (Perca fluviatilis). Proteocephalus spp. from North American lake whitefish, brown trout and Arctic charr each formed clearly distinct lineages. These results advance our understanding of the interrelationships of the Proteocephalus-aggregate, a well-recognized clade of Holarctic freshwater fish proteocephalids, and support resurrection of some of the nominal species of Proteocephalus, including Proteocephalus exiguus La Rue, 1911 from North American coregonids and Proteocephalus fallax La Rue, 1911 from European C. lavaretus, reserving Proteocephalus longicollis (Zeder, 1800) exclusively for parasites of Salmo trutta.

Salmon-lice as a potential threat to anadromous Arctic charr populations.

Salmon-lice have the potential to change the behaviour and growth of their salmonid host species. Here, the baseline infection levels of salmon-lice of post-smolts (n = 815) and veteran migrants (n = 875) of sea-run Arctic charr (Salvelinus alpinus Linnaeus, 1758) were monitored over two successive years in a sub-Arctic Norwegian fjord without farming of salmonids. All Arctic charr were collected after the sea-migration period from a trap placed in the river, ascending to their overwintering freshwater habitat (Lake Laksvatn). The sea-lice infection showed a stable infection across the 2 years while increasing through the migration period and with the size of the wild sea-run Arctic charr. The prevalence of sea-lice infection was intermediate to high, and the intensities of sea-lice infections observed were generally modest, although some individuals had high infections. The relatively high infection of salmon-lice highlights the potential detrimental effects these parasites can have at both the individual and population level of such endangered sub-Arctic life-history strategies. A comparative study should be performed in fjords with aquaculture activity as focal points for salmon-lice, to investigate the impact farming have on sea-run Arctic charr populations.

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.

Cercariae of a Bird Schistosome Follow a Similar Emergence Pattern under Different Subarctic Conditions: First Experimental Study.

The emergence of cercariae from infected mollusks is considered one of the most important adaptive strategies for maintaining the trematode life cycle. Short transmission opportunities of cercariae are often compensated by periodic daily rhythms in the cercarial release. However, there are virtually no data on the cercarial emergence of bird schistosomes from freshwater ecosystems in northern latitudes. We investigated the daily cercarial emergence rhythms of the bird schistosome Trichobilharzia sp. "peregra" from the snail host Radix balthica in a subarctic lake under both natural and laboratory seasonal conditions. We demonstrated a circadian rhythm with the highest emergence during the morning hours, being seasonally independent of the photo- and thermo-period regimes of subarctic summer and autumn, as well as relatively high production of cercariae at low temperatures typical of northern environments. These patterns were consistent under both field and laboratory conditions. While light intensity triggered and prolonged cercarial emergence, the temperature had little effect on cercarial rhythms but regulated seasonal output rates. This suggests an adaptive strategy of bird schistosomes to compensate for the narrow transmission window. Our results fill a gap in our knowledge of the transmission dynamics and success of bird schistosomes under high latitude conditions that may serve as a basis for elucidating future potential risks and implementing control measures related to the spread of cercarial dermatitis due to global warming.

Somatic Dimorphism in Cercariae of a Bird Schistosome.

Phenotypic polymorphism is a commonly observed phenomenon in nature, but extremely rare in free-living stages of parasites. We describe a unique case of somatic polymorphism in conspecific cercariae of the bird schistosome Trichobilharzia sp. "peregra", in which two morphs, conspicuously different in their size, were released from a single Radix balthica snail. A detailed morphometric analysis that included multiple morphological parameters taken from 105 live and formalin-fixed cercariae isolated from several naturally infected snails provided reliable evidence for a division of all cercariae into two size groups that contained either large or small individuals. Large morph (total body length of 1368 and 1339 µm for live and formalin-fixed samples, respectively) differed significantly nearly in all morphological characteristics compared to small cercariae (total body length of 976 and 898 µm for live and formalin samples, respectively), regardless of the fixation method. Furthermore, we observed that small individuals represent the normal/commonly occurring phenotype in snail populations. The probable causes and consequences of generating an alternative, much larger phenotype in the parasite infrapopulation are discussed in the context of transmission ecology as possible benefits and disadvantages facilitating or preventing the successful completion of the life cycle.

Taxa-specific activity loss and mortality patterns in freshwater trematode cercariae under subarctic conditions.

Cercarial activity and survival are crucial traits for the transmission of trematodes. Temperature is particularly important, as faster depletion of limited cercarial energy reserves occurs at high temperatures. Seasonal climate conditions in high latitude regions may be challenging to complete trematode life cycle during the 6-month ice-free period, but temperature effects on the activity and survival of freshwater cercariae have not been previously identified. After experimentally simulating natural subarctic conditions during warmer and colder months (13 and 6°C), a statistical approach identifying changes in the tendency of cercarial activity loss and mortality data was used to detect differences in three trematode genera, represented by four taxa (Diplostomum spp., Apatemon spp., small- and large-sized Plagiorchis spp.). A strong temperature-dependent response was identified in both activity loss and mortality in all taxa, with Diplostomum spp. cercariae showing the most gradual changes compared to other taxa. Furthermore, whilst activity loss and mortality dynamics could not be divided into 'fish- vs invertebrate-infecting cercariae' groups, the detected taxa-specific responses in relation to life-history traits indicate the swimming behaviour of cercariae and energy allocation among larvae individuals as the main drivers. Cercariae exploit the short transmission window that allows a stable continuance of trematodes' life cycles in high-latitude freshwater ecosystems.

Species interactions, environmental gradients and body size shape population niche width.

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. 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. 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. 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. 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.

Cercarial behaviour alters the consumer functional response of three-spined sticklebacks.

Free-living parasite life stages may contribute substantially to ecosystem biomass and thus represent a significant source of energy flow when consumed by non-host organisms. However, ambient temperature and the predator's own infection status may modulate consumption rates towards parasite prey. We investigated the combined effects of temperature and predator infection status on the consumer functional response of three-spined sticklebacks towards the free-living cercariae stages of two common freshwater trematode parasites (Plagiorchis spp., Trichobilharzia franki). Our results revealed genera-specific functional responses and consumption rates towards each parasite prey: Type II for Plagiorchis spp. and Type III for T. franki, with an overall higher consumption rate on T. franki. Elevated temperature (13°C) increased the consumption rate on Plagiorchis spp. prey for sticklebacks with mild cestode infections (<5% fish body weight) only. High consumption of cercarial prey by sticklebacks may impact parasite population dynamics by severely reducing or even functionally eliminating free-living parasite life stages from the environment. This supports the potential role of fish as biocontrol agents for cercariae with similar dispersion strategies, in instances where functional response relationships have been established. Our study demonstrates how parasite consumption by non-host organisms may be shaped by traits inherent to parasite transmission and dispersal, and emphasises the need to consider free-living parasite life stages as integral energy resources in aquatic food webs.

High parasite diversity in the amphipod Gammarus lacustris in a subarctic lake.

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.

Temperature does not influence functional response of amphipods consuming different trematode prey.

Direct consumption on free-living cercariae stages of trematodes by non-host organisms interferes with trematode transmission and leads to reduced infections in the next suitable hosts. Consumer functional responses provide a useful tool to examine relationships between consumption rates and ecologically relevant prey densities, whilst also accounting for abiotic factors that likely influence consumption rates. We investigated how temperature influences the consumer functional response of the amphipod Gammarus lacustris towards the cercariae of three freshwater trematodes (Diplostomum, Apatemon and Trichobilharzia). Amphipods displayed different functional responses towards the parasites, with Type II responses for Diplostomum and Type I responses for Apatemon prey. Temperature did not alter the consumption rate of the amphipod predator. Trichobilharzia was likely consumed at similar proportions as Diplostomum; however, this could not be fully evaluated due to low replication. Whilst Type II responses of invertebrate predators are common to various invertebrate prey types, this is the first time a non-filter feeding predator has been shown to exhibit Type I response towards cercarial prey. The prey-specific consumption patterns of amphipods were related to cercarial distribution in the water column rather than to the size of cercariae or temperature influence. The substantial energy flow into food webs by non-host consumer organisms highlights the importance of understanding the mechanisms that modulate functional responses and direct predation in the context of parasitic organisms.

Impacts of ontogenetic dietary shifts on the food-transmitted intestinal parasite communities of two lake salmonids.

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

Seasonal dietary shifts enhance parasite transmission to lake salmonids during ice cover.

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

Marine trophic niche use and life history diversity among Arctic charr Salvelinus alpinus in southwestern Greenland.

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 δ34 S, the Arctic charr were categorized as either marine- or freshwater-dependent feeders. The use of time-integrated trophic tracers (stable isotopes of δ13 C, δ15 N, δ34 S) 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.

A way forward with eco evo devo: an extended theory of resource polymorphism with postglacial fishes as model systems.

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.

Parasitism and the Biodiversity-Functioning Relationship.

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.

Allometric trajectories of body and head morphology in three sympatric Arctic charr (Salvelinus alpinus (L.)) morphs.

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.

The trade-off between fecundity and egg size in a polymorphic population of Arctic charr (Salvelinus alpinus (L.)) in Skogsfjordvatn, subarctic Norway.

Reproductive traits differ between intralacustrine Arctic charr morphs. Here, we examine three sympatric lacustrine Arctic charr morphs with respect to fecundity, egg size and spawning time/site to assess reproductive investments and trade-offs, and possible fitness consequences. The littoral omnivore morph (LO-morph) utilizes the upper water for feeding and reproduction and spawn early in October. The large profundal piscivore morph (PP-morph) and the small profundal benthivore morph (PB-morph) utilize the profundal habitat for feeding and reproduction and spawn in December and November, respectively. Females from all morphs were sampled for fecundity and egg-size analysis. There were large differences between the morphs. The PB-morph had the lowest fecundity (mean = 45, SD = 13) and smallest egg size (mean = 3.2 mm, SD = 0.32 mm). In contrast, the PP-morph had the highest fecundity (mean = 859.5, SD = 462) and the largest egg size (mean = 4.5 mm, SD = 0.46 mm), whereas the LO-morph had intermediate fecundity (mean = 580, SD = 225) and egg size (mean = 4.3, SD = 0.24 mm). Fecundity increased with increasing body size within each morph. This was not the case for egg size, which was independent of body sizes within morph. Different adaptations to feeding and habitat utilization have apparently led to a difference in the trade-off between fecundity and egg size among the three different morphs.

Molecular analyses reveal high species diversity of trematodes in a sub-Arctic lake.

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.

Resource Partitioning in Food, Space and Time between Arctic Charr (Salvelinus alpinus), Brown Trout (Salmo trutta) and European Whitefish (Coregonus lavaretus) at the Southern Edge of Their Continuous Coexistence.

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.

Morphological divergence between three Arctic charr morphs - the significance of the deep-water environment.

Morphological divergence was evident among three sympatric morphs of Arctic charr (Salvelinus alpinus (L.)) that are ecologically diverged along the shallow-, deep-water resource axis in a subarctic postglacial lake (Norway). The two deep-water (profundal) spawning morphs, a benthivore (PB-morph) and a piscivore (PP-morph), have evolved under identical abiotic conditions with constant low light and temperature levels in their deep-water habitat, and were morphologically most similar. However, they differed in important head traits (e.g., eye and mouth size) related to their different diet specializations. The small-sized PB-morph had a paedomorphic appearance with a blunt head shape, large eyes, and a deep body shape adapted to their profundal lifestyle feeding on submerged benthos from soft, deep-water sediments. The PP-morph had a robust head, large mouth with numerous teeth, and an elongated body shape strongly related to their piscivorous behavior. The littoral spawning omnivore morph (LO-morph) predominantly utilizes the shallow benthic-pelagic habitat and food resources. Compared to the deep-water morphs, the LO-morph had smaller head relative to body size. The LO-morph exhibited traits typical for both shallow-water benthic feeding (e.g., large body depths and small eyes) and planktivorous feeding in the pelagic habitat (e.g., streamlined body shape and small mouth). The development of morphological differences within the same deep-water habitat for the PB- and PP-morphs highlights the potential of biotic factors and ecological interactions to promote further divergence in the evolution of polymorphism in a tentative incipient speciation process. The diversity of deep-water charr in this study represents a novelty in the Arctic charr polymorphism as a truly deep-water piscivore morph has to our knowledge not been described elsewhere.