Temperature acclimation and heat tolerance of photosynthesis in Norwegian Saccharina latissima (Laminariales,Phaeophyceae)

Kelps, seaweeds and seagrasses provide important ecosystem services in coastal areas, and loss of these macrophytes is a global concern. Recent surveys have documented severe declines in populations of the dominant kelp species, Saccharina latissima, along the south coast of Norway. S. latissima is a cold‐temperate species, and increasing seawater temperature has been suggested as one of the major causes of the decline. Several studies have shown that S. latissima can acclimate to a wide range of temperatures. However, local adaptations may render the extrapolation of existing results inappropriate. We investigated the potential for thermal acclimation and heat tolerance in S. latissima collected from three locations along the south coast of Norway. Plants were kept in laboratory cultures at three different growth temperatures (10, 15, and 20°C) for 4–6 weeks, after which their photosynthetic performance, fluorescence parameters, and pigment concentrations were measured. S. latissima obtained almost identical photosynthetic characteristics when grown at 10 and 15°C, indicating thermal acclimation at these temperatures. In contrast, plants grown at 20°C suffered substantial tissue deterioration, and showed reduced net photosynthetic capacity caused by a combination of elevated respiration and reduced gross photosynthesis due to lowered pigment concentrations, altered pigment composition, and reduced functionality of Photo‐system II. Our results support the hypothesis that extraordinarily high temperatures, as observed in 1997, 2002, and 2006, may have initiated the declines in S. latissima populations along the south coast of Norway. However, observations of high mortality in years with low summer temperatures suggest that reduced population resilience or other factors may have contributed to the losses.     

Development of Saccharina latissima (Phaeophyceae) kelp hatcheries with year-round production of zoospores and juvenile sporophytes on culture ropes for kelp aquaculture

Saccharina latissima is attractive for industrial cultivation for different usages, such as biofuels, feed supplements, and derivation of chemicals. A continuous supply of kelp sporelings throughout the year may ensure a year-round production of kelp juveniles on ropes. In this study, induction of sporangial areas (sorus portions) on the blade of S. latissima was performed throughout the year at three locations: Trondheim (Norway), Grenaa (Denmark), and Sylt island (Germany). The results indicate that a year-round sorus induction in S. latissima is possible and that this induction is controlled by applying short-day treatment of adult sporophytes throughout the year and by the removal of the basal blade meristem. The artificially induced and released zoospores formed viable sporelings at all seasons, but cultivation in the sea in Norway was successful only during autumn, winter, and spring, while the growth conditions were poor during the summer. The results are important for industrial scale-up and continuous production of kelp biomass.     

In a squeeze: Epibiosis may affect the distribution of kelp forests

The processes limiting the population recovery of the kelp Saccharina latissima after recent large‐scale loss from the south coast of Norway are poorly understood. Previous investigations do, however, suggest that the impacts of biotic interactions (epibiosis and competition) and increased water turbidity are important. We investigated the depth‐related patterns of growth, epibiosis, and mortality in two sample populations of kelp, from the south and the southwest coast of Norway. The investigations were performed over a period of seven months, in a crossed translocational study, where kelps were mounted on rigs at six depths (1, 3, 6, 9, 15, and 24 m). In a second experiment, the amounts of light blocked by different epibiont layers growing on the kelp frond were investigated. While growth decreased with depth in spring and summer, the kelp grew faster at 15 m than at shallower depths in fall. Survival was low both in shallow water and below 15 m depth. Epibionts covered the kelp growing at depths from 1 to 9 m, and the laboratory study showed that the coverage may have deprived the individuals of as much as 90% of the available light. Although the depth‐related results we present apply—in the strictest sense—only to kelp translocated on rigs, we argue that the relative patterns are relevant for natural populations. Growth and survival of S. latissima is likely to be reduced by heavy loads of epibionts, while depths where epibionts are sparse may be close to the lower limit of the kelps depth distribution along the south coast of Norway. This suggests that a vertical squeeze, or narrowing of the distribution range of kelp forests may be occurring in Norway.     

Overgrazing of Kelp Beds Along the Coast of Norway

The aim of this study was to better understand the down-grazing of kelp beds by sea urchins (Strongylocentrotus droebachiensis) along the coast of Norway. Barren grounds were first observed in sheltered areas along the coast of the counties of Trø ndelag, Nordland and Troms in 1974. In the 1980s, the barren grounds spread to areas more heavily exposed to waves. In the 1990s, the kelp beds were re-established in some localities in southern Trø ndelag, initially in wave-exposed areas. In the northernmost parts of Norway, i.e. the counties of Troms and Finnmark, the barren ground areas may still be increasing. Crabs (Cancer pagurus) and common eiders are the most common predators on urchins. Predation on sea urchins in kelp beds is probably not among the factors that limit the sea urchin populations. Along the coast of Nordland and further north, sea urchins are infected by nematodes, resulting in a low, but significant increase in their mortality. No re-growth of kelp beds has been found in the most infected areas. In the late 1960s and the early 1970s, a high occurrence of echinoderm larvae was observed in deeper waters. This was a period with cold water, which may have caused high recruitment of sea urchins. The bet-hedging life strategy of sea urchins may account for the sudden increase in the size of the populations. In the present paper I propose the hypothesis that higher individual growth rates and higher mortality rates in the south than in the north may explain the decrease in the populations, which may in turn account for the re-growth of kelp in the southern areas.     

Strong population structure but no equilibrium yet: Genetic connectivity and phylogeography in the kelp Saccharina latissima (Laminariales,Phaeophyta)

Kelp aquaculture is globally developing steadily as human food source, along with other applications. One of the newer crop species is Saccharina latissima, a northern hemisphere kelp inhabiting temperate to arctic rocky shores. To protect and document its natural genetic variation at the onset of this novel aquaculture, as well as increase knowledge on its taxonomy and phylogeography, we collected new genetic data, both nuclear and mitochondrial, and combined it with previous knowledge to estimate genetic connectivity and infer colonization history. Isolation‐with‐migration coalescent analyses demonstrate that gene flow among the sampled locations is virtually nonexistent. An updated scenario for the origin and colonization history of S. latissima is developed as follows: We propose that the species (or species complex) originated in the northwest Pacific, crossed to the northeast Pacific in the Miocene, and then crossed the Bering Strait after its opening ~5.5 Ma into the Arctic and northeast Atlantic. It subsequently crossed the Atlantic from east to west. During the Pleistocene, it was compressed in the south with evidence for northern refugia in Europe. Postglacial recolonization led to secondary contact in the Canadian Arctic. Saccharina cichorioides is shown to probably belong to the S. latissima species complex and to derive from ancestral populations in the Asian North Pacific. Our novel approach of comparing inferred gene flow based on coalescent analysis versus Wright's island model suggests that equilibrium levels of differentiation have not yet been reached in Europe and, hence, that genetic differentiation is expected to increase further if populations are left undisturbed.     

qPCR-based relative quantification of the brown algal endophyte <Emphasis Type="Italic">Laminarionema elsbetiae</Emphasis> in <Emphasis Type="Italic">Saccharina latissima</Emphasis>: variation and dynamics of host—endophyte interactions

Morphological changes—such as dark spots, twisted stipes and deformed blades—have been observed in wild and cultivated Saccharina latissima. The putative cause for the disease symptoms is the filamentous endophytic brown alga Laminarionema elsbetiae, which is known to invade stipes and fronds of its hosts. Little is known about this interaction and its occurrence in the field, although former studies indicated high endophyte prevalence in kelp populations. Previous epidemiological studies on kelp endophytes were mainly based on the examination of microscopic sections, followed by time-consuming isolation and cultivation steps in order to identify the endophyte and a reliable method to quantify endophyte infections was missing. As a novel approach, we established and validated a qPCR assay for relative quantification of the endophyte L. elsbetiae within its host S. latissima, which allows to examine both, the prevalence of endophytic algae and the severity of infections. The assay was shown to be highly specific and suitable to reliably detect small amounts of endophyte DNA in the host. Using this method, we detected very high endophyte prevalence in the investigated kelp populations, up to 100% in young S. latissima sporophytes in Brittany during spring. Furthermore, our results suggest that Saccharina sporophytes are infected early in their life and that seasonality and environmental factors have a significant impact on infection rates. In the future, this approach could also be applied to study other host-endophyte pairs using specific primers.     

Cranial variation in the European badger Meles meles (Carnivora,Mustelidae) in Scandinavia

Morphometric variation in 30 craniometric characters of 465 skulls of the European badgers (Meles meles) from across Europe was analysed. Multivariate analyses revealed that the populations from Norway, Sweden, and Finland differ from other European populations in having smaller skulls. The analyses also revealed significant differences between the ‘south‐western Norwegian’ and ‘main Fennoscandian’ forms. On average, badgers from south‐west Norway were smaller than those of the remaining Fennoscandia. Morphological differences between the ‘south‐western Norwegian’ and ‘main Fennoscandian’ populations of M. meles suggest a possible in situ semisympatric divergence since the beginning of the Holocene warming, or a complex history of two groups involving at least two colonization routes. The small‐sized Scandinavian badgers may be close to the ancestral form that used to be widespread in Denmark and throughout Europe. The animals from south‐west Norway may instead be descendants of ancestors that were the first to penetrate the southern parts of the Scandinavian Peninsula. The ‘main Fennoscandian’ badgers are likely to have been the descendants of the second wave of recolonization of Scandinavia. Specifically, they might have colonized the Scandinavian Peninsula from the east after the last glaciation.     

Recruitment of the intertidal kelp Lessonia nigrescens Bory in northern Chile: successional constraints and opportunities

The strong 1982/83 El Niño event caused local extinction in populations of the low intertidal kelp Lessonia nigrescens Bory on northern Chilean coasts. The kelp has partially recolonized, but its recovery has apparently been low. This study documents the effect of biological factors that potentially decrease the velocity of kelp recolonization in northern Chile. A removal experiment showed that encrusting coralline algae, which dominate the lower intertidal, significantly reduce the recruitment of L. nigrescens in the presence of herbivores. Epithallium shedding by encrusting corallines is the most probable cause for this inhibition process. On the other hand, grazing on encrusting corallines by the chiton Enoplochiton niger (Barnes), a large-sized herbivore (10–20 cm length), could also affect kelp recruitment. The intestinal content of E. niger, where encrusting corallines are the main item (84.2%), revealed the presence of L. nigrescens. Among the natural substrata on which kelp recruits, measured at five different localities, the coarsely-branched alga Corallina officinalis (Decaisne) Kützing reached the highest frequency, despite its extremely low cover (< 1%) in the field. This suggests that settlement on turfs of C. officinalis allows the kelp to escape from herbivory, thus facilitating its recruitment. The roles of timing of kelp recruitment and seasonal grazing are also discussed.     

Genetic diversity of Saccharina latissima (Phaeophyceae) along a salinity gradient in the North Sea–Baltic Sea transition zone

The North Sea–Baltic Sea transition zone constitutes a boundary area for the kelp species Saccharina latissima due to a strong salinity gradient operating in the area. Furthermore, the existence of S. latissima there, along Danish waters, is fairly patchy as hard bottom is scarce. In this study, patterns of genetic diversity of S. latissima populations were evaluated along the salinity gradient area of Danish waters (here designated brackish) and were compared to reference sites (here designated marine) outside the gradient area, using microsatellite markers. The results showed that the S. latissima populations were structured into two clusters corresponding to brackish versus marine sites, and that gene flow was reduced both between clusters and between populations within clusters. In addition, results provided empirical evidence that marginal populations of S. latissima in the salinity gradient area exhibited a distinct genetic structure when compared to marine ones. Brackish populations were less diverse, more related, and showed increased differentiation over distance compared to marine populations. The isolation of the brackish S. latissima populations within the salinity gradient area of Danish waters in conjunction with their general low genetic diversity makes these populations vulnerable to ongoing environmental and climate change, predicted to result in declining salinity in the Baltic Sea area that may alter the future distribution and performance of S. latissima in the area.     

Role of Avian Seed Dispersers in Tree Recruitment in Woodland Pastures

The rewilding of abandoned agricultural lands opens up opportunities for the recovery of forest ecosystem extent. Frugivorous animals not only take part in the regeneration of unaltered forests, but leave a visible footprint in restoring areas in the form of the number and spatial distribution of new trees recruited from dispersed seeds. Nevertheless, their contribution is conditioned by how environmental factors affect both the patterns of seed dispersal and the fate of post-dispersal regeneration stages throughout the whole ecosystem. Here, we evaluated the role of avian seed dispersers in tree regeneration in woodland pastures resulting from anthropic deforestation. Using an integrative approach, considering the different tree regeneration stages, we dissected the ways in which forest loss conditioned the contribution of frugivores. Habitat structure influenced bird activity, mainly restricting seed dispersal to forested areas. Tree recruitment was severely reduced during early regeneration stages, but maintained the initial forest-biased spatial distribution. However, the presence of scrub in deforested areas, which protect against grazing at late regeneration stages, drastically increased the relevance of tree recruitment outside the forest. Frugivorous birds made a significant contribution to tree regeneration in the woodland pastures under study. The interplay between seed dispersal by birds and the protective role of scrub was fundamental in facilitating the recolonization of deforested areas. If we wish to encourage this natural reforestation, we will need to preserve populations of frugivorous birds while favoring landscape configurations that encourage seed dispersal outside the forest and species that promote tree establishment (like nurse scrubs).     

Nuclear genetic markers indicate Danish origin of the Norwegian beech (Fagus sylvatica L.) populations established in 500�C1,000?AD

The northernmost range of beech (Fagus sylvatica L.) is in southern Norway and consists of two distinct and isolated distributions, a single population at Seim in West Norway and several adjacent populations in Vestfold, East Norway. The modest beech pollen deposits beyond these main distributions suggest that the Norwegian beech distribution has never been an extension of the south Scandinavian range. We used genetic markers and historical sources to trace the ancestor populations for the beech at Seim and Vestfold, hypothesising Denmark as the most likely source. Nuclear inter-simple sequence repeat markers, amplified by polymerase chain reaction (PCR), were applied to estimate genetic distances between beech populations in Norway, England and Denmark. The variation in chloroplast DNA polymorphism was estimated using PCR-restriction fragment length polymorphism. The nuclear genetic data indicate Denmark as a source for the beech in Norway, although the data are less certain in the case of Seim than in that of Vestfold. The populations from South England were genetically different from most Scandinavian populations. The genetic variation within Norwegian populations was only slightly lower than that of the English and Danish populations, questioning birds as vectors for dispersal. Thus, the pollen data and our results are in accordance with the intentional introduction and documented human migrations across Skagerrak before and during the Viking Age.     

Modelling seasonal growth and composition of the kelp Saccharina latissima

A dynamical model for simulating growth of the brown macroalga Saccharina latissima is described. In addition to wet and dry weights, the model simulates carbon and nitrogen reserves, with variable C/N ratio. In effect, the model can be used to emulate seasonal changes in growth and composition of the alga. Simulation results based on published, environmental field data are presented and compared with corresponding data on growth and composition. The model resolves seasonal growth, carbon and nitrogen content well, and may contribute to the understanding of how seasonal growth in S. latissima depends simultaneously on a combination of several environmental factors: light, nutrients, temperature and water motion. The model is applied to aquaculture problems such as estimating the nutrient scavenging potential of S. latissima and estimating the potential of this kelp species as a raw material for bioenergy production.     

Assimilation of inorganic nutrients from salmon (Salmo salar) farming by the macroalgae (Saccharina latissima) in an exposed coastal environment: implications for integrated multi-trophic aquaculture

This paper investigated the assimilation of dissolved inorganic nitrogen (DIN) in Saccharina latissima in proximity to salmon cages in coastal waters. The bioassays were performed on plants from three stations located in the vicinity of a salmon farm (Salmo salar) in exposed waters at Tristein (63° 52′ N, 9° 37′ E) in Central Norway. The growth, the C and N content, and the nitrogen isotope ratios (δ¹⁵N) of S. latissima were monitored over 1 year. The DIN concentrations in seawater were higher at the salmon farm stations than at the reference station during the winter, and the N/P ratio at the salmon farm stations was higher from September to January and in June. S. latissima at the salmon farm stations grew faster than at the reference station. The length of S. latissima increased by 50 % when integrated with the salmon farm compared to the reference station. The N content of S. latissima was positively correlated to the DIN concentration in seawater (p?<?0.05), but the increased N supply from salmon did not result in N accumulation in S. latissima at the salmon farm station because of the dilution by a higher growth rate. The δ¹⁵N in S. latissima was higher at the salmon farm station from April to June and changed in the direction of the δ¹⁵N signature in urine. This indicated that N in S. latissima at the salmon farm station partly originated from the salmon. One hectare of S. latissima may absorb 0.8～1.2 t N during one growth season. Large-scale cultivation of S. latissima should be considered to mitigate the environmental effects of DIN wastes from salmon farms.     

Hyper-oceanic liverwort species of conservation concern: evidence for dispersal limitation and identification of suitable uncolonised regions

In order to successfully manage and conserve species and plant communities, it is important to have a good understanding of their ecology and distributions. The three liverwort species Anastrophyllum donnianum, Scapania ornithopodioides and Scapania nimbosa, are restricted to the mixed northern hepatic mat community found in the most oceanic parts of north-western Europe. These species are of conservation concern because they are globally rare with strict environmental requirements and a limited dispersal potential, which makes them vulnerable to disturbance and climate change. In this study we used species distribution modelling to (1) predict their potential distribution in Norway (2) to assess whether they are limited by dispersal or suitable climate, (3) identify which climatic factors are most important in determining their distribution and (4) suggest regions for further field based surveys. Maximum entropy (MaxEnt) models were developed for each species using target-group background data, and five environmental coverage layers. Our results indicate that all three species are limited by dispersal rather than the availability of suitable areas in Norway. In particular, A. donnianum seems to be limited from reaching uncolonised highly suitable areas in northern Norway due to a barrier unsuitable region with insufficient summer rain. S. ornithopodioides is absent from northern Norway despite the presence of highly suitable regions scattered along the coast. The models locate highly suitable areas where conservation measures should be focused when they overlap with known populations. Areas of interest for targeting searches for potentially undiscovered populations are indicated.     

Population genomic evidence for plant glacial survival in Scandinavia

Quaternary glaciations have played a major role in shaping the genetic diversity and distribution of plant species. Strong palaeoecological and genetic evidence supports a postglacial recolonization of most plant species to northern Europe from southern, eastern and even western glacial refugia. Although highly controversial, the existence of small in situ glacial refugia in northern Europe has recently gained molecular support. We used genomic analyses to examine the phylogeography of a species that is critical in this debate. Carex scirpoidea Michx subsp. scirpoidea is a dioecious, amphi‐Atlantic arctic–alpine sedge that is widely distributed in North America, but absent from most of Eurasia, apart from three extremely disjunct populations in Norway, all well within the limits of the Weichselian ice sheet. Range‐wide population sampling and variation at 5,307 single nucleotide polymorphisms show that the three Norwegian populations comprise unique evolutionary lineages divergent from Greenland with high between‐population divergence. The Norwegian populations have low within‐population genetic diversity consistent with having experienced genetic bottlenecks in glacial refugia, and host private alleles that probably accumulated in long‐term isolated populations. Demographic analyses support a single, pre‐Weichselian colonization into Norway from East Greenland, and subsequent divergence of the three populations in separate refugia. Other refugial areas are identified in North‐east Greenland, Minnesota/Michigan, Colorado and Alaska. Admixed populations in British Columbia and West Greenland indicate postglacial contact. Taken together, evidence from this study strongly indicates in situ glacial survival in Scandinavia.     

Open-sea cultivation by transplanting young fronds of the kelp Saccharina latissima

Saccharina latissima is an economically and ecologically important native kelp. As its limited supply from wild stock cannot meet increasing current and future demands, methods for its cultivation in the ocean need to be developed. This kelp is now beginning to be farmed off the Atlantic coast of Spain using a regular method similar to the “forced cultivation” technique used with Asian kelps (kombu). Its cultivation is also a growing enterprise in other European countries. In this study, the open-sea farming of S. latissima using the transplanting method is tested on a commercial scale. This cultivation method has not been studied with kelp species outside Asian waters. The tested method includes the following steps: indoor production of seedlings, pre-culture in greenhouse tanks, and open-sea cultivation by transplanting young fronds. Results demonstrate that open-sea cultivation using transplanted young fronds is a technically and biologically viable method. The mean yield obtained (7.8 kg fresh wt per meter rope equivalent to 45.6 t fresh wt per hectare farm) is satisfactory, considering the low densities of transplanted fronds (25–30 fronds per meter rope). Moreover, these values are comparable to those reported in previous cultivations with this species, as well as in the farming of similar kelps. The transplanting method used in conjunction with the regular cultivation method has valuable practical applications for the commercial farming of S. latissima.     

Biomass yield and morphological features of the seaweed Saccharina latissima cultivated at two different sites in a coastal bay in the Atlantic coast of Spain

The kelp Saccharina latissima is an economically important edible seaweed with great potential for use in other applications. The goal of this study was to identify locations in a coastal bay with better conditions for commercial cultivation of this species. We compared the biomass yield and morphological features of S. latissima cultivated at two locations in a bay of Galicia (northwest Spain): in an external moderately exposed site and in an internal sheltered site. Biomass yield was significantly higher at the moderately exposed site than at the sheltered site (16 fresh kg vs. 12 fresh kg per linear meter of rope). Fronds were significantly larger at the moderately exposed site compared to that at the sheltered site. However, the “substantiality value” (around 40 mg cm^?2), which is an index used to assess blade quality for human consumption, did not differ significantly between the two locations. The effect of light exposure and water velocity seemed to be the most important environmental factors that explain the differences in the results obtained at both culture sites. In summary, both the moderately exposed site and the sheltered site were suitable for cultivation of S. latissima, as indicated by the high yields and similar blade quality obtained at both locations.     

Glacial survival or late glacial colonization? Phylogeography of the root vole (Microtus oeconomus) in north‐west Norway

Aim It has been proposed that the root vole subspecies, Microtus oeconomus finmarchicus, survived the last glacial period on islands on the north‐west coast of Norway. The Norwegian island of Andøya may have constituted the only site with permanent ice‐free conditions. Geological surveys and fossil finds from Andøya demonstrate that survival throughout the last glacial maximum was probably possible for some plants and animals. In this study we aim to infer the recent evolutionary history of Norwegian root vole populations and to evaluate the glacial survival hypothesis. Methods DNA sequence variation in the mitochondrial cytochrome b gene was studied in 46 root voles from 19 localities. Location Northern Fennoscandia and north‐west Russia with a focus on islands on the north‐west coast of Norway. Results The phylogeographical analyses revealed two North European phylogroups labelled ‘Andøya’ and ‘Fennoscandia’. The Andøya phylogroup contained root voles from the Norwegian islands of Andøya, Ringvassøya and Reinøya and two localities in north‐west Russia. The Fennoscandian phylogroup encompassed root voles from the three Norwegian islands of Kvaløya, Håkøya and Arnøya and the remaining specimens from Norway, northern Sweden and Finland. Nucleotide diversity within the Andøya and Fennoscandian phylogroups was similar, ranging from 0.5% to 0.7%. Main conclusions Both our genetic data and previously published morphological data are consistent with in situ glacial survival of root voles on Andøya during the last glacial maximum. However, the level of genetic diversity observed in the extant island populations, the past periods of severe climatic conditions on Andøya and the ecology of the root vole are somewhat difficult to reconcile with this model. A biogeographical scenario involving late glacial recolonization along the northern coasts of Russia and Norway therefore represents a viable alternative. Our results demonstrate that complex recolonization and extinction histories can generate intricate phylogeographical patterns and relatively high levels of genetic variation in northern populations.     

Structure and specific features of formation of aggregations of deepwater redfish <Emphasis Type="Italic">Sebastes mentella</Emphasis> (Scorpaenidae) in the Norwegian Sea pelagial

Results of studies of the structure and specific features of formation of aggregations of deepwater redfish Sebastes mentella in the Norwegian Sea pelagial are provided. Data on the biological characteristics of S. mentella from different regions of the Norwegian Sea pelagial do not allow us to consider its aggregations as separate populations. On the basis of comparative analysis of the structure of aggregations in the Norwegian Sea and of population parameters and specific features of the life cycle of S. mentella in different parts of the Atlantic Ocean, its complex intraspecies organization at sites of overlapping of two largest populations—North Atlantic and Norwegian-Barents Sea-was established. The formation of pelagic aggregations in the water area of the Norwegian Sea occurs due to migration of maturing and mature individuals from the adjoining water areas. In the southern areas of the Norwegian Sea, dominant recruitment comes from the pelagial of the northeastern part of the Irminger Sea, which is promoted by the direction of streams of the North-Atlantic Current; it is not excluded either that fish can migrate to the southern part of the sea from the slopes of the Faeroes, Faeroes-Icelandic threshold, and the western coast of Norway. Migration to the northern areas of the sea is mainly performed by rapidly maturing fish from the rearing region from the western slopes of the Barents Sea and Spitsbergen Archipelago. The presence of seasonal migrations hinders the formation in the pelagial of perch isolated groups and promotes their migration.