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June 1997 - present
UiT The Arctic University of Norway (former University of Tromsoe)
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- Professor (Full)
Publications
Publications (155)
While it is likely that ecological context is important, the factors that facilitate and maintain variable levels of intraspecific diversity in Salmonidae fishes across environments remain unclear.
Using a meta‐analysis of sympatric ecotype assemblages from two salmonid genera— Salvelinus and Coregonus —we evaluated the importance of ecological fac...
Tapeworms of the genus Proteocephalus Weinland, 1858 (Cestoda: Proteocephalidae) are common and widespread intestinal parasites of whitefish (Coregonus spp., Salmonidae: Coregoninae). Previous taxonomic studies, based solely on morphology and inconsistently fixed specimens, concluded that all salmoniform fish, including whitefish, are parasitised b...
Reciprocal effects of adaptive radiations on the evolution of interspecific interactions, like parasitism, remain barely explored. We test whether the recent radiations of European whitefish (Coregonus spp.) across and within perialpine and subarctic lakes promote its parasite Proteocephalus fallax (Platyhelminthes: Cestoda) to undergo host reperto...
Aim
High repeatability among assemblages of closely related but ecologically distinct ecotypes implies predictability in evolution and assembly of communities. The conditions under which ecotype assemblages form predictably, and the reasons, have been little investigated. Here, we test whether repeatability declines as the number of ecotypes builds...
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 farm...
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...
Host density, host body size and ambient temperature have all been positively associated with increases in parasite infection. However, the relative importance of these factors in shaping long‐term parasite population dynamics in wild host populations is unknown due to the absence of long‐term studies. Here, we examine long‐term drivers of gill lic...
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 s...
Acanthobdellida gnaw into the sides of salmonid fishes in frigid Arctic lakes and rivers, latching on with fearsome facial hooks. Sister to leeches, they are an ancient lineage with two described species. Unfortunately, Acanthobdellida are rarely collected, leading to a paucity of literature despite their unique morphology. Populations range from E...
Disentangling the causes and consequences of ontogenetic niche shifts has been a pivotal challenge in ecology, aiming to enhance the understanding of biological processes that function at the individual, population, and community levels. Studies on ontogenetic dietary shifts have traditionally focused on short time scales, mostly including sampling...
Acanthobdellida gnaw into the sides of salmonid fishes in frigid Arctic lakes and rivers, latching on with fearsome
facial hooks. Sister to leeches, they are an ancient lineage with two described species. Unfortunately, Acanthobdellida
are rarely collected, leading to a paucity of literature despite their unique morphology. Populations range from E...
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 schist...
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 ba...
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/...
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 perio...
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 o...
Introduction: Polymorphic Arctic charr Salvelinus alpinus populations frequently display distinct differences in habitat use, diet and parasite communities. Changes to the relative densities and composition of fish communities has the potential to alter the habitat-niche of Arctic charr populations. This study investigated whether the restocking of...
Concomitant predation, which occurs when parasites are consumed and digested along with their hosts, has previously been suggested as a profound factor determining food web structure. Few studies have adressed the impact of concomitant predation in research on behaviourally parasite-modified prey or in biological control studies. However, empirical...
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 popula...
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...
Abstract 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 fo...
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,...
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 account...
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. Th...
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...
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 cha...
Trophically transmitted parasites are known to impair fish growth in experimental studies, but this is not well documented in natural populations. For Arctic charr [Salvelinus alpinus (L.)], individual growth is positively correlated with food consumption. However, increased food consumption will increase the exposure to trophically transmitted par...
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 trophical...
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 interconnected southern Norwegian lakes at different altitudes. A...
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...
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)...
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, ta...
Climate change and the intensification of land use practices are causing widespread eutrophication of subarctic lakes. The implications of this rapid change for lake ecosystem function remain poorly understood. To assess how freshwater communities respond to such profound changes in their habitat and resource availability, we conducted a space‐for‐...
We review three long‐term research programs performed over the last four decades on the ecology and management of oligotrophic lake systems with different fish communities at 69° N in Norway. Through whole‐lake perturbation experiments, intensive culling of stunted fish removed 35 tons (1984–1991) of Arctic charr Salvelinus alpinus in Takvatn (15 k...
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...
The eye represents an immune privileged organ where parasites can escape host reactions. This study provides the first systematic evidence of the pathology associated with Diplostomum sp. infection in the eye retina of fish (i.e. Arctic charr). Histological sections showed that the trematodes caused mechanical disengagement between the retinal pigm...
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 mig...
From 2011-2013, anadromous brown trout Salmo trutta L. (213-730 mm) were collected during or shortly after their marine feeding migration at seven different localities in Central Norway. The mean volume of stomach content (%) of marine fish prey eaten by S. trutta captured in marine waters varied from 34 % to 89 %. There was a high prevalence (67-1...
The littoral benthivorous charrs Salvelinus malma with subterminal mouth distributed in Lake Kronotskoe basin are separated into three distinct morphs. These morphs significantly differ in head proportions, jaws' length, skull structure, and degree of gill rakers' reduction. The morphs spawn in distant lake tributaries at the spawning grounds of va...
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...
The nosed charr Salvelinus malma 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 t...
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...
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 (Fjellfrøsvatn and Skogsfjordvatn, Norway). The replicated morph pair, the profundal spawning benthivorous PB-morph and the littoral spawning omnivorous...
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...
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 sui...
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...
The trophic niches of Arctic charr and brown trout differ when the species occur in sympatry. Their trophically transmitted parasites are expected to reflect these differences. Here, we investigate how the infections of Diphyllobothrium dendriticum and D. ditremum differ between charr and trout. These tapeworms use copepods as their first intermedi...
We contrast the genetic consequences of allopatric and sympatric divergence from the littoral spawning Arctic charr morph from Lake Fjellfrøsvatn. The littoral spawning Arctic charr has sympatrically diverged into a natural profundal adapted morph and via a recent (1930) translocation of about 40 adult Arctic charr established a new allopatric popu...
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, Fjellfrøsvatn. Fjellfrøsvatn has one littoral omnivorous and one small profundal benthivor...
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 t...
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 Galggojav...
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 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)...
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 li...
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 f...
The new data on the parasites fauna of the Lake Kronotskoe sympatric charr forms (genus Salvelinus (Nilsson) Richardson, 1836) is presented. Parasites fauna of Bigmouth and Smallmouth charr forms are described for the first time. The information about Longhead charr, Nosed charr and White charr parasites is added. 29 species of parasites from 9 cla...
Some fish parasites constitute severe management problems as they may cause mortality of their fish host or are important zoonoses of humans. Parasite assessments are therefore critical to keep track of infections. If conventional sampling techniques can be simplified, parasite assessments might be easier to obtain, less time-consuming and more ext...
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...
Subarctic populations of brown trout (Salmo trutta) are often heavily infected with cestodes of the genus Diphyllobothrium, assumedly because of their piscivorous behaviour. This study explores possible associations between availability of fish prey and Diphyllobothrium spp. infections in lacustrine trout populations. Trout in (i) allopatry (group...
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....
G) имеют высокие показатели инвазии Cystidicola farionis, Cyathocephalus truncatus и Crepidostomum metoecus. Рыбы, питающиеся хирономидами и другим бен-тосом (группа А), практически не заражены этими видами паразитов, для них характерны Phyllodisto-mum umblae и Proteocephalus longicollis. Рыбы группы G имеют высокую условную биомассу паразитов по с...
Parasite−host systems show markedly variable patterns in isotopic fractionation: parasites can be either depleted or enriched in 15N and 13C as compared to their hosts. However, it remains unknown whether isotopic fractionation patterns are similar in comparable parasite−host systems from markedly different ecosystems. Results of this study show th...
Tracking individual variation in the dynamics of parasite infections in wild populations is often complicated by lack of knowledge of the epidemiological history of hosts. Whereas the dynamics and development of Gyrodactylus salaris Malmberg, 1957, on Atlantic salmon, Salmo salar L., are known from laboratory studies, knowledge about infection deve...
The ecological and life-history knowledge of anadromous stocks of Arctic charr (Salvelinus alpinus) from Greenland is almost lacking. This study focus on comparative results from three watercourses that supposedly contain both resident and anadromous population of charr. The three size distributions observed in the Arctic charr populations in the K...
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 ident...
p>Amundsen, P.-A., Smalås, A., Knudsen, R., Kristoffersen, R., Siwertsson, A. & Klemetsen, A. 2015. Takvatnprosjektet. Erfaringer fra forskning og kultivering av en overbefolka røyebestand. Rapport, UiT Norges arktiske universitet. 53 s.
Overbefolka fiskebestander er et stort problem i mange av våre innsjøer. Det typiske kjennetegnet er en stor te...
Prey preference of top predators and energy flow across habitat boundaries are of fundamental importance for structure and function of aquatic and terrestrial ecosystems, as they may have strong effects on production, species diversity, and food-web stability. In lakes, littoral and pelagic food-web compartments are typically coupled and controlled...
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 (...
Co-introduction and colonization of parasites with the introduction of new host species into aquatic habitats may depend on the host specificity and dispersal capabilities of the parasites. We compared the metazoan parasite community of an introduced three-spined stickleback (Gasterosteus aculeatus) population with that of the nearby source populat...
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 environmen...
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,...
Invasion of exotic species has caused the loss of biodiversity and imparts evolutionary and ecological changes in the introduced systems. In northern Fennoscandia, European whitefish (Coregonus lavaretus (L.)) is a highly polymorphic species displaying adaptive radiations into partially reproductively isolated and thus genetically differentiated sy...
The numbers of divers, ducks, gulls, terns and waders in the 15 km2 oligotrophic lake Takvatn, North Norway, were estimated six times during 1983-2012. Systematic mapping surveys were done by boat within the first week after the ice-break in June. Twenty-one species were observed over the years and 12 were regarded as breeding on the lake. Red-brea...
Understanding how a monophyletic lineage of a species diverges into several adaptive forms has received increased attention in recent years, but the underlying 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 pos...
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 tr...
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 Arc...
Coregonids constitute one of the most diverse fi sh families in the northern freshwater systems and several species are highly endangered mainly due to anthropogenic pressure. Cost effective and powerful polymerase chain reaction (PCR) microsatellite multiplex assays were established for genetic studies of the population structure, hybridization an...
Invasions of non-native species represent a global problem of great scientific interest. Here we study in detail the response in population and life history characteristics of closely related native species, with divergent habitat preferences, that are impacted by an invading species over a sufficient time period to allow a new stable state to beco...
Parallel phenotypic evolution occurs when independent populations evolve similar traits in response to similar selective regimes. However, populations inhabiting similar environments also frequently show some phenotypic differences that result from non-parallel evolution. In this study, we quantified the relative importance of parallel evolution to...
Natural populations often vary in their degree of ecological, morphological and genetic divergence. This variation can be arranged along an ecological speciation continuum of increasingly discrete variation, with high inter-individual variation at one end and well defined species in the other. In postglacial fishes, evolutionary divergence has comm...
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) (pred...
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. T...
... Original Research. You have full text access to this OnlineOpen article Parallel and non - parallel morphological divergence among foraging specialists in European whitefish ( Coregonus lavaretus ). Anna Siwertsson 1,* ,; Rune ...
The effect of three different temperatures on growth in a first progeny generation, hatchery reared, subarctic population of European whitefish (Coregonus lavaretus L.) were investigated. The whitefish (start weight 444 g, ±SD 125 g) were reared for 60 days at three constant temperatures; 15, 18 and 21°C and under ambient light regimes for 70°N lat...
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 generali...
Introduced species can alter the topology of food webs. For instance, an introduction can aid the arrival of free-living consumers using the new species as a resource, while new parasites may also arrive with the introduced species. Food-web responses to species additions can thus be far more complex than anticipated. In a subarctic pelagic food we...
We explore the long-term developments in population biology and life history during the invasion and establishment of the fish species vendace Coreg-onus albula in a subarctic watercourse by comparing life-history traits and molecular genetic estimates between the source and the colonist population. The two populations exhibited highly contrasting...
The temporal stability of the distribution of gill raker number in European whitefish (Coregonus lavaretus) populations was monitored through a long-term study of seven subarctic lakes. There is long tradition to identify different morphs of whitefish and other coregonids by the number of gill rakers, which is a heritable morphological trait. This...