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Investigations of genetic stock structure sometimes reveal a mismatch between management units and biological units. In Scandinavian waters, Norway lobster (Nephrops norvegicus) is divided into two management units (the Skagerrak–Kattegat and the Norwegian Deep). We have tested the population genetic structure of Nephrops within this region using microsatellite DNA markers, and compared the structure with the present management units. Our study suggests no population genetic structure of Nephrops within the Skagerrak, Kattegat, and Norwegian Deep region, whereas a shallow genetic structure was detected on a larger geographical scale when comparing outgroup samples from Scotland and Iceland. We found indications of sex-biased dispersal as the overall genetic differences were larger for females. Ocean current patterns suggest that Nephrops stocks in the region may be connected by larval drift. The two areas differ in fishing pressure, monitoring, assessment, and regulations, which is an argument for maintaining the present two-areas management regime despite the evidence for one biological population.

Environmental DNA (eDNA) has gained popularity as a tool for ecosystem biomonitoring and biodiversity assessment. Although much progress has been made regarding laboratory and fieldwork protocols, the issue of sampling efficiency requires further investigation, particularly in three-dimensional marine systems. This study focuses on fish community composition in marine ecosystems and aims to analyze the efficiency of sampling design given the sampling effort for distinguishing between different communities. We sampled three fjords in Northern Norway, taking samples along fjord transects and at three different depths, and amplified a fragment of the mitochondrial 12S rRNA gene of bony fishes using the MiFish primers. We evaluated the effect of (i) the number of sampling stations, (ii) samples' spatial distribution, and (iii) the data treatment approach (presence/absence versus semiquantitative) for maximizing the efficiency of eDNA metabarcoding sampling when inferring differences of fish community compositions between fjords. We found that the manner of data treatment strongly affected the minimum number of sampling stations required to detect differences among communities; because the semiquantitative approach retained some information about abundance of the underlying reads, it was the most efficient. Furthermore, we found little-to-no difference of fish communities in samples from intermediate depths when comparing vertical fish communities. Lastly, we found that the differences between fish communities at the surface were the highest across the horizontal distance and overall, samples ~30 km apart showed the highest variation in the horizontal distribution. Boosting sampling efficiency (reducing sampling effort without compromising ecological inferences) can significantly contribute to enhanced biodiversity management and efficient biomonitoring plans.

Marine fisheries are often allocated to stocks that reflect pragmatic considerations and may not represent the species’ spatial population structure, increasing the risk of mismanagement and unsustainable harvesting. Here we compile mark–recapture data collected across the North Atlantic to gain insight into the spatial population structure of Greenland halibut (Reinhardtius hippoglossoides), an issue that has been unresolved for decades. The dataset contains 168130 fish tagged from 1952 to 2021, with 5466 (3.3%) recaptured individuals. Our results indicate that fish tagged at

The cold-adapted polar cod Boreogadus saida, a key species in Arctic ecosystems, is vulnerable to global warming and ice retreat. In this study, 1257 individuals sampled in 17 locations within the latitudinal range of 75–81°N from Svalbard to East Siberian Sea were genotyped with a dedicated suite of 116 single-nucleotide polymorphic loci (SNP). The overall pattern of isolation by distance (IBD) found was driven by the two easternmost samples (East Siberian Sea and Laptev Sea), whereas no differentiation was registered in the area between the Kara Sea and Svalbard. Eleven SNP under strong linkage disequilibrium, nine of which could be annotated to chromosome 2 in Atlantic cod, defined two genetic groups of distinct size, with the major cluster containing seven-fold larger number of individuals than the minor. No underlying geographic basis was evident, as both clusters were detected throughout all sampling sites in relatively similar proportions (i.e. individuals in the minor cluster ranging between 4 and 19% on the location basis). Similarly, females and males were also evenly distributed between clusters and age groups. A differentiation was, however, found regarding size at age: individuals belonging to the major cluster were significantly longer in the second year. This study contributes to increasing the population genetic knowledge of this species and suggests that an appropriate management should be ensured to safeguard its diversity.

Blue ling is a deep-water species that has been severely fished upon into coastal and offshore fisheries since the early 1970s, thus causing the collapse of the populations in the last two decades. Genetic information is scarce in this species, and molecular markers are therefore needed to provide advice both for management and for rebuilding the stocks. A suite of 103 SNP markers was identified from ddRAD sequencing data. From those, 81 were organized in three multiplex reactions, and tested on 150 individuals from three different sampling locations. Good-quality amplification products were successfully obtained from 70 of the markers. All SNP loci were biallelic, with averaged He per locus ranging between 0.101 and 0.500.

Coastal Atlantic cod (Gadus morhua) in the Northeast Atlantic has seen a continuous decline since the industrialization of the coastal fishery, and management needs to address the spatial and temporal complexities of coexisting cod stocks. Toward that end, genetic analyses and oceanographic modelling of coastal and oceanic cod larval drift patterns were combined to elucidate the mechanisms responsible for an observed genetic cline over a >1500 km stretch along the coast of Norway. The results indicate that the north–south cline in coastal cod represents an extended contact zone between genetically divergent North Sea and Northeast Arctic cod and is maintained by two-way gene flow: by northward drift of pelagic eggs and larvae and by southward spawning migrations of Northeast Arctic cod. Computer simulations verify that the genetic cline can be established rapidly if gene flow into coastal populations is substantial. The shape of the cline, on the other hand, was found to be largely insensitive to the total amount of gene flow and therefore carries little information on extent of gene flow into and among coastal populations.

Many marine organisms have a permanent presence both inshore and offshore and spawn in multiple areas, yet their status as separate populations or stocks remain unclear. This is the situation for the northern shrimp (Pandalus borealis) around the Arctic Ocean, which in northern Norway represents an important income for a small-scale coastal fishery and a large-vessel offshore fleet. In Norwegian waters, we uncovered two distinct genetic clusters, viz. a Norwegian coastal and a Barents Sea cluster. Shrimps with a mixed heritage from the Norwegian coastal and the Barents Sea clusters, and genetically different from both, inhabit the fjords at the northernmost coast (Finnmark). Genetic structure between fjords did not display any general trend, and only the Varangerfjord in eastern Finnmark displayed significant genetic structure within the fjord. Shrimps in the Finnmark fjords differed in some degree from shrimps both in the adjacent Barents Sea and along the rest of the coast and should probably be considered a separate management unit.

Challenging long‐held perceptions of fish management units can help to protect vulnerable stocks. When a fishery consisting of multiple genetic stocks is managed as a single unit, overexploitation and depletion of minor genetic units can occur. Atlantic cod (Gadus morhua) is an economically and ecologically important marine species across the North Atlantic. The application of new genomic resources, including SNP arrays, allows us to detect and explore novel structure within specific cod management units. In Norwegian waters, coastal cod (i.e. those not undertaking extensive migrations) are divided into two arbitrary management units defined by ICES: one between 62° and 70°N (Norwegian coastal cod; NCC) and one between 58° and 62°N (Norwegian coastal south; NCS). Together, these capture a fishery area of >25,000 km2 containing many spawning grounds. To assess whether these geographic units correctly represent genetic stocks, we analysed spawning cod of NCC and NCS for more than 8,000 SNPs along with samples of Russian White Sea cod, north‐east Arctic cod (NEAC: the largest Atlantic stock), and outgroup samples representing the Irish and Faroe Sea's. Our analyses revealed large differences in spatial patterns of genetic differentiation across the genome and revealed a complex biological structure within NCC and NCS. Haplotype maps from four chromosome sets show regional specific SNP indicating a complex genetic structure. The current management plan dividing the coastal cod into only two management units does not accurately reflect the genetic units and needs to be revised. Coastal cod in Norway, while highly heterogenous, is also genetically distinct from neighbouring stocks in the north (NEAC), west (Faroe Island) and the south. The White Sea cod are highly divergent from other cod, possibly yielding support to the earlier notion of subspecies rank. publishedVersion

The bearded goby Sufflogobius bibarbatus is an abundant endemic small fish species on the continental shelf of the northern Benguela. The goby habitat is characterised by generally low bottom oxygen concentrations that vary spatially and seasonally. In the present study of population structure, 13 samples of S. bibarbatus from inner and outer shelf areas between 19°S and 32°S were screened using ten microsatellite loci. The genetic data were analysed in relation to isolation by distance and depth. Furthermore, for the first time, this study examined genetic data in relation to bottom oxygen concentration at the sampling locations. The data show low but significant genetic heterogeneity (G‐test; FST = 0.007, p < .05). There was weak but significant genetic differentiation along a latitudinal gradient across all sampling sites from 19.50°S to 32.37°S (Mantel test; r = .464, p = .001), but this disappeared when the southernmost sample was removed. On the other hand, a positive correlation of bottom oxygen concentration with pairwise FST (r = .336; p = .017) was observed among the sampling sites from the Northern Benguela shelf area. Overall, the data are complex but suggest that isolation by distance and bottom oxygen concentration may play a role in the genetic structuring of S. bibarbatus. The findings are discussed in relation to the species’ life history features and oceanographic characteristics of the Benguela upwelling ecosystem. Paid Open Access

Background Atlantic cod (Gadus morhua L.) has formed the basis of many economically significant fisheries in the North Atlantic, and is one of the best studied marine fishes, but a legacy of overexploitation has depleted populations and collapsed fisheries in several regions. Previous studies have identified considerable population genetic structure for Atlantic cod. However, within Norway, which is the country with the largest remaining catch in the Atlantic, the population genetic structure of coastal cod (NCC) along the entire coastline has not yet been investigated. We sampled > 4000 cod from 55 spawning sites. All fish were genotyped with 6 microsatellite markers and Pan I (Dataset 1). A sub-set of the samples (1295 fish from 17 locations) were also genotyped with an additional 9 microsatellites (Dataset 2). Otoliths were read in order to exclude North East Arctic Cod (NEAC) from the analyses, as and where appropriate. Results We found no difference in genetic diversity, measured as number of alleles, allelic richness, heterozygosity nor effective population sizes, in the north-south gradient. In both data sets, weak but significant population genetic structure was revealed (Dataset 1: global FST = 0.008, P

In management of fisheries, knowledge about the species is crucial due to differences in life history traits. The first step is therefore species identification. For many species this task is straightforward, however for some species e.g. the two species of Atlantic redfish Sebastes mentella and Sebastes norvegicus, it can be difficult for an untrained eye. With the goal to separate the two species, we analysed otolith shape variation of S. mentella and S. norvegicus caught on the continental slope of East Greenland using the R package “Shape R”. Results were evaluated against genetic analysis of the same fish, and compared to results of both a visual identification of the two species and a separation based on a linear discriminant analysis on standardised values of fish length, fish weight and otolith weight. It was concluded that the objective otolith shape analysis using the Shape R package analysis achieve a reasonable classification success, however with a clear bias towards S. mentella. Classification using the otolith weight achieved a slightly higher success than the shape analysis making it a promising method. Furthermore, the method is at the same time both objective, less time consuming than the otolith shape analysis and less expensive than genetic analysis. However, the visual classification method of the whole fish had the highest success rate of the three tested methods, which despite the need for trained technicians makes it the most successful method.

Identification of cryptic species can have profound implications in fishery management, conservation and biodiversity contexts. In the North Atlantic, the genus Sebastes is currently represented by four species, although additional cryptic species have been assumed. The connectivity of the gene-pools within the genus in Greenland waters, in particular, remains largely unexplored. Using a panel of 13 microsatellite markers for 720 fish, we explored the species complex of Sebastes norvegicus in Greenland waters. Genetic analyses provided evidence for three cryptic species in samples that were morphologically identified as S. norvegicus. They were termed S. norvegicus-A, S. norvegicus-B, and S. norvegicus-giants. A few phenotypic features exist to identify adult S. norvegicus giants, but no characteristics have been identified for the two other cryptic species. The proposed cryptic species should be recognized in the management regime to ensure sustainable exploitation and conservation of Sebastes species in Greenland waters.

Exploited marine resources can be managed more effectively when accurate information on geographic population structure is available. Genetic markers offer a powerful tool for fisheries management, because they reveal biologically sound management units. Single nucleotide polymorphisms (SNPs) markers derived from restriction-site associated DNA sequencing (RAD-seq) were developed and used to investigate the stock structure of Greenland halibut (Reinhardtius hippoglossoides). A total of 96 SNPs were analyzed from 384 individuals and eight locations across the Atlantic. Our results suggest a subdivision of Greenland halibut into two populations, an eastern Atlantic population and a western Atlantic population, with a proposed border across the Denmark Strait. In general, Greenland halibut display weak but significant population structure (overall FST= 0.003; p < 0.001), which can be explained by connectivity among populations owing to the migratory behavior or egg and larval drift.

Science-based management of marine fisheries and effective ecosystem monitoring both require the analysis of large amounts of often complex and difficult to collect information. Legislation also increasingly requires the attainment of good environmental status, which again demands collection of data to enable efficient monitoring and management of biodiversity. Such data is traditionally obtained as a result of research surveys through the capture and/or visual identification of organisms. Recent years have seen significant advances in the utilisation of environmental DNA (eDNA) in the marine environment in order to develop alternative cost-effective ways to gather relevant data. Such approaches attempt to identify and/or quantify the species present at a location through the detection of extra-organismal DNA in the environment. These new eDNA based approaches have the potential to revolutionise data collection in the marine environment using non-invasive sampling methods and providing snapshots of biodiversity beyond the capacity of traditional sampling. Here we present a non-technical summary of different approaches in the field of eDNA, and emphasise the broad application of this approach, with value for the governance and management of marine aquatic ecosystems. The review focuses on identifying those tools which are now readily applicable and those which show promise but are currently in development and require further validations. The aim is to provide an understanding of techniques and concepts that can be used by managers without genetic or genomic expertise when consulting with specialists to perform joint evaluations of the utility of the approaches. Spanish Ministry of Science, Innovation and Universities through the EDAMAME ("Environmental DNA based approaches for marine and aquatic monitoring and evaluation") project [CTM2017-89500-R]; Scientific Research Network "Eco-evolutionary dynamics in natural and anthropogenic communities" (Research Foundation - Flanders) [W 0.037.10N] info:eu-repo/semantics/publishedVersion

Fish stocks represent fundamental units in fisheries management, and their identification, especially in mixed-fisheries, represents one of the primary challenges to sustainable harvest. Here, we describe the first “real-time” genetic management program used to manage a mixed-stock fishery of a non-salmonid and commercially significant marine fish, the Atlantic cod (Gadus morhua L). Based upon the analysis of >18 000 fish sampled from the commercial catch in Lofoten (Norway), which represents the largest remaining cod fishery in the Atlantic, we estimated the fraction of North East Arctic cod (NEAC), and Norwegian Coastal cod (NCC), just 24 h post-landing. These estimates, based upon the analysis of the Pantophysin gene, were performed weekly in the winter and spring of each year in the period 2007–2017. The program has successfully permitted the Norwegian Directorate of Fisheries to actively manage the commercial exploitation of the highly abundant NEAC stock, while simultaneously limiting exploitation of the fragile NCC stock, both of which overlap at the spawning grounds. Data from this program have also revealed a distinct temporal increase in the fraction of NEAC on the spawning grounds in this region, which is consistent with the overall increased abundance of this stock as estimated by ICES. publishedVersion

The identification of isolated populations in widely distributed marine species is often impeded by low levels of genetic differentiation. However, modern genetic approaches now allow for the efficient detection of potentially large numbers of novel genetic variations, thereby improving the power to identify biologically meaningful management units. To investigate the genetic structuring of saithe (Pollachius virens L.), we applied seascape genetic approaches to 131 single nucleotide polymorphism loci genotyped in 584 fish collected from 11 locations across the North Atlantic. Wombling analysis and the Monmonier algorithm revealed four genetic clusters (Barents Sea, Central Northeast Atlantic, Rockall, and Canada) across the species distribution range. These results imply that genetic structuring in saithe may be influenced by abiotic factors such as geographical distance, and bathymetry as well as biotic factors such as sex-biased migration, and natal homing. The results suggest a potential mismatch between management and biological units across the Northeast Atlantic, which may have implications for sustainable exploitation of the species.

Genetic population structure is often used to identify management units in exploited species, but the extent of genetic differentiation may be inflated by geographic variation in the level of hybridization between species. We identify the genetic population structure of Sebastes mentella and investigate possible introgression within the genus by analyzing 13 microsatellites in 2562 redfish specimens sampled throughout the North Atlantic. The data support an historical divergence between the ‘shallow’ and ‘deep’ groups, beyond the Irminger Sea where they were described previously. A third group, ‘slope’, has an extended distribution on the East Greenland Shelf, in addition to earlier findings on the Icelandic Slope. Furthermore, S. mentella from the Northeast Arctic and Northwest Atlantic waters are genetically different populations. In both areas, interspecific introgression may influence allele frequency differences among populations. Evidence of introgression was found for almost all the identified Sebastes gene pools, but to a much lower extent than suggested earlier. Greenland waters appear to be a sympatric zone for many of the genetically independent Sebastes groups. This study illustrates that the identified groups maintain their genetic integrity in this region despite introgression. This article is protected by copyright. All rights reserved.

With the decline of many of the world’s fisheries, increased regulation, including marine protected areas (MPA), forms an increasingly important role in promoting sustainable resource use. Here, we present a novel “real-time” genetic monitoring programme used to protect the depleted Norwegian coastal cod stock (NCC) in an MPA during the spawning season, while a fishery targeted at the sustainable Northeast Arctic cod stock (NEAC) operates immediately outside. In the period 2009–2016, >6800 cod from the fishery were genotyped with the PanI locus that is discriminatory between these two stocks. The estimated fraction of NEAC increased during the study period until 2014; however, it did not exceed 70% for any sustained period. Therefore, the MPA remained closed for commercial harvest. Genetic analysis of eggs revealed a distinctly lower fraction of NEAC than in the catch from the adult stock, both immediately outside and within the MPA itself. We suggest that this discrepancy is likely to reflect differences in spawning areas used by NCC and NEAC. Estimated fractions of NEAC/NCC using PanI, otolith classification, and 39 single nucleotide polymorphisms were similar, thus validating the use of PanI to estimate NEAC/NCC composition.

The Northeast Arctic cod (Gadus morhua L.: NEAC) remains the most abundant cod stock in the North Atlantic, while the catches of the partially co-occurring Norwegian coastal cod (NCC) stocks have dramatically decreased in recent years. To ensure effective management of the two stocks, it is necessary to know if the population genetic structure is associated with any pattern in the spatial dynamics or whether it is affected by any distinct environmental factors. By combining information from electronic data storage tags (DST) and molecular genetics methods with statistical tools, we have been able to associate spatial dynamics and distinct environmental factors to the two cod stocks. In general, adult NEAC migrate between deep, warm overwintering grounds and shallow summer feeding grounds where water temperatures maybe low. In contrast, NCC do not undertake large-scale seasonal migrations, show little seasonal variation in depth distribution, and experience the opposite seasonal change in temperature compared with NEAC. However, within the NCC group, some individuals did conduct longer horizontal movements than others. Even though the distances calculated in this study represent the shortest distance between release and recapture positions, they are far higher than previously reported by NCC. Distinctive depth profiles indicate that this migrant NCC have moved out of the area, passing the deep trenches outside Lofoten while more stationary NCC occupies shallower depths throughout the year. The temperature profiles also indicate that migrant and stationary NCC has occupied different areas during the year. We demonstrate that the combination of information from DSTs and molecular genetics offers a deeper understanding of individual cod behaviour, provides an insight in the spatial dynamics of the species, and ultimately, improves the scientific basis for management of a complex mixed fishery of Atlantic cod.

Genetic population structure is often used to identify management units in exploited species, but the extent of genetic differentiation may be inflated by geographic variation in the level of hybridization between species. We identify the genetic population structure of Sebastes mentella and investigate possible introgression within the genus by analyzing 13 microsatellites in 2,562 redfish specimens sampled throughout the North Atlantic. The data support an historical divergence between the “shallow” and “deep” groups, beyond the Irminger Sea where they were described previously. A third group, “slope,” has an extended distribution on the East Greenland Shelf, in addition to earlier findings on the Icelandic slope. Furthermore, S. mentella from the Northeast Arctic and Northwest Atlantic waters are genetically different populations. In both areas, interspecific introgression may influence allele frequency differences among populations. Evidence of introgression was found for almost all the identified Sebastes gene pools, but to a much lower extent than suggested earlier. Greenland waters appear to be a sympatric zone for many of the genetically independent Sebastes groups. This study illustrates that the identified groups maintain their genetic integrity in this region despite introgression.

Oceans are experiencing increasing acidification in parallel to a distinct warming trend in consequence of ongoing climate change. Rising seawater temperatures are mediating a northward shift in distribution of Atlantic cod (Gadus morhua), into the habitat of polar cod (Boreogadus saida), that is associated with retreating cold water masses. This study investigates the competitive strength of the co-occurring gadoids under ocean acidification and warming (OAW) scenarios. Therefore, we incubated specimens of both species in individual tanks for 4 months, under different control and projected temperatures (polar cod: 0, 3, 6, 8 °C, Atlantic cod: 3, 8, 12, 16 °C) and PCO2 conditions (390 and 1170 µatm) and monitored growth, feed consumption and standard metabolic rate. Our results revealed distinct temperature effects on both species. While hypercapnia by itself had no effect, combined drivers caused nonsignificant trends. The feed conversion efficiency of normocapnic polar cod was highest at 0 °C, while optimum growth performance was attained at 6 °C; the long-term upper thermal tolerance limit was reached at 8 °C. OAW caused only slight impairments in growth performance. Under normocapnic conditions, Atlantic cod consumed progressively increasing amounts of feed than individuals under hypercapnia despite maintaining similar growth rates during warming. The low feed conversion efficiency at 3 °C may relate to the lower thermal limit of Atlantic cod. In conclusion, Atlantic cod displayed increased performance in the warming Arctic such that the competitive strength of polar cod is expected to decrease under future OAW conditions.

We developed and characterized 20 microsatellite primer loci for the northern shrimp Pandalusborealis. All 20 loci were polymorphic with number of alleles ranging from 3 to 36 and with observed heterozygosity between 0.04 and 0.93. In addition, we tested the utility of these markers in three related shrimp species, P. montagui, Atlantopandaluspropinqvus and Dichelopandalusbonnieri. These new markers will prove useful in the identification of stock structure and hence, assessment of the commercially important species P. borealis.

Summary This study presents a molecular genetic characterization of Atlantic cod reared in commercial marine farms. Samples consisted of approximately 47 fish collected from nine cages located on four farms throughout Norway. In addition, 28 farmed escapees were recaptured in the sea (443 fish in total). Nine microsatellite loci and the Pan I gene were analysed, revealing a total of 181 alleles. Each sample contained 43‐63% of total allelic variation. Comparing variation with published data for wild cod indicates that lower genetic variation exists within single cages than in wild populations. Significant linkage disequilibrium was observed amongst pairs of loci in all samples, suggesting a low number of contributing parental fish. Global FST was 0.049, and the highest pairwise FST value (pooled loci) was 0.085. For single loci, the Pan I gene was the most diagnostic, displaying a global FST of 0.203. Simulations amongst the samples collected on farms revealed an overall correct self-assignment percentage of 75%, demonstrating a high probability of identifying individuals to their farm of origin. Identification of the 28 escapees revealed a single cage as the most likely source of origin for half of the escapees, whilst the remaining fish were assigned to a mixture of samples, suggesting more than one source of escapees.

The deepwater redfish Sebastes mentella has a wide distribution in the North Atlantic Ocean. In the Irminger Sea, there is evidence for two phenotypes (deep-sea and oceanic) of deepwater redfish. The two phenotypes have overlapping geographic distributions but differ in depth preferences. There are two hypotheses on deepwater redfish stock structure in the Irminger Sea. One suggests that mature individuals of a single stock segregate according to size and age and therefore that the phenotypes represent different life stages of the same stock. The other hypothesis suggests that there are two different stocks and that these stocks segregate mainly according to depth. Additionally, it is not clear whether the fish of the deep-sea phenotype in the Irminger Sea and those on the Icelandic continental shelf represent one stock. Analysis of genetic variability at eight allozyme markers in 1,763 deepwater redfish from 26 samples collected at different depths in the Irminger Sea and the Icelandic continent...

Hypoxia can influence fish growth, survival and on larger scales, population structure. These effects may be influenced by water temperature, and may vary intra-specifically with genotype. In Atlantic cod (Gadus morhua L.), the two haemoglobin homozygotes (Hb-I⁎11 and Hb-I⁎22) vary in oxygen affinity at different temperatures, which is thought to correspond to variation in hypoxia tolerance. We therefore tested if hypoxic avoidance behaviour in cod 1) depends on ambient temperature and 2) is modified by haemoglobin genotype. In a laminar flow choice box, we subjected juvenile cod to an initial phase of non-escapable hypoxia, and a subsequent recovery phase, where one habitat was kept at 20% O2 saturation while the other was raised in steps to full saturation. The experiment was performed at 5 and 15 °C with Hb-I⁎11 and Hb-I⁎22 cod. Cod responded to inescapable hypoxia by reducing their overall swimming speed and then, at the initial levels of the recovery phase, avoiding the most hypoxic habitat, irrespective of temperature or genotype. Fish recovered quickly as O2 levels increased, as evidenced by increased swimming speed and time spent in the most hypoxic habitat. The avoidance response depended strongly on temperature: the relative reduction in speed and avoidance of the most hypoxic habitat was more pronounced at 15 than at 5 °C. During the recovery phase, stressed fish initially maintained a higher swimming speed in the most hypoxic habitat. However, as O2 increased, swimming speed in both habitats converged. This point of convergence occurred at a lower O2 saturation at 5 °C. Fish ventilation rate in inescapable hypoxia was also higher at 15 °C. Haemoglobin genotype did not influence either ventilation rates or the nature of the hypoxic avoidance response at either temperature, but Hb-I⁎11 cod swam faster than Hb-I⁎22 cod in normoxia at 15 °C. Our results indicate that increased temperature limits the ability of cod of both haemoglobin genotypes to exploit hypoxic habitats. This may have negative future consequences for coastal cod stocks in light of increasing global temperatures and eutrophication in coastal waters. © 2008 Elsevier B.V. All rights reserved.

Atlantic cod is composed of multiple migratory and stationary populations widely distributed in the North Atlantic Ocean. The Northeast Arctic cod (NEAC) population in the Barents Sea undertakes annual spawning migrations to the northern Norwegian coast. Although spawning occurs sympatrically with the stationary Norwegian coastal cod (NCC), phenotypic and genetic differences between NEAC and NCC are maintained. In this study, we resolve the enigma by revealing the mechanisms underlying these differences. Extended linkage disequilibrium (LD) and population divergence were demonstrated in a 17.4-Mb region on linkage group 1 (LG1) based on genotypes of 494 SNPs from 192 parents of farmed families of NEAC, NCC or NEACxNCC crosses. Linkage analyses revealed two adjacent inversions within this region that repress meiotic recombination in NEACxNCC crosses. We identified a NEAC-specific haplotype consisting of 186 SNPs that was fixed in NEAC sampled from the Barents Sea, but segregating under Hardy–Weinberg equilibrium in eight NCC stocks. Comparative genomic analyses determine the NEAC configuration of the inversions to be the derived state and date it to ~1.6–2.0 Mya. The haplotype block harbours 763 genes, including candidates regulating swim bladder pressure, haem synthesis and skeletal muscle organization conferring adaptation to long-distance migrations and vertical movements down to large depths. Our results suggest that the migratory ecotype experiences strong directional selection for the two adjacent inversions on LG1. Despite interbreeding between NEAC and NCC, the inversions are maintaining genetic differentiation, and we hypothesize the co-occurrence of multiple adaptive alleles forming a ‘supergene’ in the NEAC population.

The large-scale population genetic structure of northern shrimp, Pandalus borealis, was investigated over the species’ range in the North Atlantic, identifying multiple genetically distinct groups. Genetic divergence among sample localities varied among 10 microsatellite loci (range: FST = −0.0002 to 0.0475) with a highly significant average (FST = 0.0149; P

The nucleotide sequence of the internal transcribed region (ITS) of ribosomal RNA genes from Atlantic cod (Gadus morhua L.) was determined. The complete ITS region spanned approximately 1113 base pairs. The ITS1 region comprised 532 base pairs, the 5.8S region 159, and the ITS2 region contained 422 base pairs. Sequence data were obtained from a total of 12 samples, one pool from six cod and 11 individuals. The sequencing was carried out in two separate experimental periods employing slightly different methodology. The samples were from two different cod stocks, Norwegian costal cod and North East Arctic cod. The sequence analysis showed that in the 12 samples, the ITS region, including the 5.8S RNA, was identical. The ITS region is thus totally conserved in these two cod stocks. The extreme conservation of the ITS regions in the cod rDNA could reflect the small genome size of cod and/or indicate a specific critical role in the processing of the ribosomal units in cold-adapted species.

The levels of heat-shock proteins of the 70 kDa family (Hsp70s) were measured in different soft tissues of Atlantic cod Gadus morhua from different locations and after exposure to various thermal conditions: acute temperature increments (1° C day -1 ), mid-term (73 days at 4-15° C) and long-term thermal acclimation (278 days at 8-15° C), and seasonal and latitudinal temperature variations (field samples). Tissue specific distribution patterns of Hsp70s were observed: liver > gills > red blood cells > brain > white muscle. Thus, different tissues may have required different levels of protection by Hsp70s, and possibly this was related to the rate of protein synthesis. There were no differences in tissue Hsp70s between Arctic cod populations (Arctic, i.e. Barents and White Seas, Norwegian coast, and North or Baltic Seas). No changes in Hsp70s levels were observed in response to temperature variation of any intensity (acute fluctuation or seasonal and latitudinal) within the range of physiological temperatures (4-15° C) in wild and laboratory Atlantic cod. This confirms previous observations that changes in Hsp70 caused by such temperature variation are often small in fishes. Probably, the constitutive level of Hsp70s in Atlantic cod was high enough to overcome potentially harmful effects of temperature variations within the physiological range. A suppressing effect of high temperature (15° C) has already been observed at a systematic level (as reduced rate of somatic growth), whereas it is not reflected in modified Hsp70s. Therefore, Hsp70s apparently played a secondary role in defining thermal tolerance limits in Atlantic cod. These conclusions are in line with a recent concept of thermal tolerance which indicated that the first line of thermal limitation in the cold and warm is a loss in aerobic scope.

Genetic variation among and within species of the genus Sebastes [ S. viviparus , S. fasciatus , S. mentella (including deep-sea and oceanic types) and S. marinus (both ordinary and giant types)] from the North Atlantic was investigated by polymerase chain reaction (PCR)-based random amplification of polymorphic DNA (RAPD). Initial analyses were based on 80 different 10 base pair primers applied to a small number of individuals of each of the four species. Four of the most variable primers were selected for analysis of the total samples from Norway, Iceland and Canada. Only one primer, OPA20, could be used to diagnostically distinguish among species. Statistical analysis clustered all samples in relation to known species structure. No genetic distance was found between ordinary and giant-type S. marinus and minor differences were observed between S. viviparus from Iceland and Norway. Sebastes mentella from the different locations clustered together, but some structuring was shown within this species. Twel...

In the present paper are described the growth properties of three different haemoglobin genotypes of juvenile Atlantic cod (Gadus morhua) reared at 7, 10, 13 and 16 °C. In addition one group was reared under “temperature steps” i.e. moved successively from 16 to 13 and 10 °C. The genotype Hb-I(2/2)displayed the overall highest growth rate in the temperature range 13–16 °C, whereas the Hb- I(1/1)genotype showed the highest overall growth at the lowest temperature (7 °C). Accordingly, we found a significant interaction between genotype and temperature. The differences in growth were largest when cod were reared under the temperature step regime where the Hb-I(2/2)genotype displayed 17 and 24% higher growth than Hb-I(1/1)and Hb-I(1/2),respectively. Optimal temperature for growth (Topt.G) varied between the genotypes with the genotype Hb-I(1/2)displaying the highest (mean ± SE) Topt.G (14.5 ± 1:0.8 °C) and Hb-I(1/1)the lowest (12.5 ± 0.2 °C). The biological significance of this link between biochemical genetic variation and physiological properties might be the influences on growth pattern, ultimate size and age at first maturity.

As part of a more extensive study on the inter- and intraspecific genetic variation of redfish, genus Sebastes, a total sample of 593 individuals of the species Sebastes viviparus were collected from several locations in Norwegian waters and off the south-west coast of Iceland. Haemoglobin and five polymorphic enzymes were analysed. A new haemoglobin polymorphism was observed, and unique alleles were found in Icelandic and Norwegian samples. The results of the enzyme analyses were largely consistent with previous results, but interpopulation variation was observed in the frequencies of IDHP* and Hb* alleles. S. viviparus displayed higher similarity with S. fasciatus from the West Atlantic than with the other Sebastes species.

Fourteen polymorphic microsatellite markers with tetranucleotide repeats were developed for the bearded goby (Sufflogobius bibarbatus) from partial genomic DNA libraries using a repeat enrichment protocol, and characterized using two putative populations from the northern Benguela. The average number of alleles per locus ranged from 4 to 34, and the observed heterozygosities across loci were between 0.237 and 0.983. We also tested the utility of these markers in two other marine gobies; sand goby Pomatoschistus minutus and two-spotted goby Gobiusculus flavescens. These polymorphic markers can be employed to investigate population structure and related questions of the bearded goby.

Effects of global warming on animal distribution and performance become visible in many marine ecosystems. The present study was designed to develop a concept for a cause and effect understanding with respect to temperature changes and to explain ecological findings based on physiological processes. The concept is based on a wide comparison of invertebrate and fish species with a special focus on recent data obtained in two model species of fish. These fish species are both characterized by northern and southern distribution limits in the North Atlantic: eelpout (Zoarces viviparus), as a typical non-migrating inhabitant of the coastal zone and the cod (Gadus morhua), as a typical inhabitant of the continental shelf with a high importance for fisheries. Mathematical modelling demonstrates a clear significant correlation between climate induced temperature fluctuations and the recruitment of cod stocks. Growth performance in cod is optimal at temperatures close to 10°C, regardless of the population investigated in a latitudinal cline. However, temperature specific growth rates decrease at higher latitudes. Also, fecundity is less in White Sea than in North and Baltic Sea cod or eelpout populations. These findings suggest that a cold-induced shift in energy budget occurs which is unfavorable for growth performance and fecundity. Thermal tolerance limits shift depending on latitude and are characterized by oxygen limitation at both low or high temperatures. Oxygen supply to tissues is optimized at low temperature by a shift in hemoglobin isoforms and oxygen binding properties to lower affinities and higher unloading potential. Protective stimulation of heat shock protein synthesis was not observed. According to a recent model of thermal tolerance the downward shift of tolerance limits during cold adaptation is associated with rising mitochondrial densities and, thus, aerobic capacity and performance in the cold, especially in eurythermal species. At the same time the costs of mitochondrial maintenance reflected by mitochondrial proton leakage should rise leaving a lower energy fraction for growth and reproduction. The preliminary conclusion can be drawn that warming will cause a northern shift of distribution limits for both species with a rise in growth performance and fecundity larger than expected from the Q10 effect in the north and lower growth or even extinction of the species in the south. Such a shift may heavily affect fishing activities in the North Sea.

Summary PCR-RFLP analysis of mitochondrial DNA (NADH-3/4 gene and the control region) of the Benguelan bearded goby Sufflogobius bibarbatus revealed weak, but significant genetic differentiation (overall FST = 0.137) that could not be predicted by hydrodynamics alone, thus warranting further investigations to understand the forces behind population divergence within an ecosystem having complex dynamics.

A joint venture between Iceland and Norway to study the beaked redfish, Sebastes mentella , in the Irminger Sea was undertaken in 1995. The genetic relationship of deep-sea and oceanic S. mentella in the Irminger Sea and their relationship to redfish groups in adjacent waters were studied by gel electrophoresis and isoelectric focusing of haemoglobin and allozymes. The oceanic type of redfish in the Irminger Sea occurred as a homogenous group, but differed from the deep-sea type. This result was mainly based on frequency distributions of the enzyme loci MEP-1* and IDHP-2* and minor haemoglobin components. Heterogeneity was found within the deep-sea type, which also showed similarities with beaked redfish from the Icelandic shelf area. One sample of “untyped” S. mentella found below 600 m resembled the oceanic type in morphology and gene frequencies. Both types of S. mentella differed from S. mentella from Canada and Norway, but they could not be distinguished from immature S. mentella from Greenland as far as could be judged from previous studies.

In 1996 a new fishery along the Reykjanes Ridge below 500 m was initiated. A Sebastes species similar in morphology to S. marinus and with average length above 60 cm (called giants) was one of the main targets. In three seasons; spring, summer and fall, in the period of 1995–1997, 560 tissue and blood samples were collected and in addition some whole redfish for morphometric studies were taken. The genetic relationship of this type of redfish to related Sebastes species was studied by starch and polyacrylamid gel electrophoresis of haemoglobin and allozymes. The results are compared to earlier studies on redfish from Greenland and Icelandic waters. Although morphologically similar to S. marinus, these giant redfish displayed haemoglobin patterns diagnostically different from the ones usually seen in S. marinus and S. mentella. Low level of IDHP-1* variation in liver tissue was found, in contrast to the ordinary S. marinus. On the other hand monomorphic MEP indicated relationship to S. marinus.

During an attempt to develop a molecular method for identification of species and morphs of Atlantic Sebastes, extremely low levels of interspecific sequence variation in a mitochondrial large subunit (16S) ribosomal RNA were found. Nucleotide sequencing of a 480 base pair portion of the gene from representatives of six phenotypes of four species of Sebastes was executed. No variation was found among the individuals of ‘Regular’ S. marinus, ‘giant’S. marinus, ‘oceanic’S. mentella, and ‘deepsea’ S. mentella analyzed. Sequences from S. fasciatis and S. viviparus differed with only one base each from the marinuslmentella haplotype. The low level of genetic variation at this gene makes it unsuitable as a target for methods to identify North Atlantic Sebastes. Level of variation may indicate recent evolutionary divergence between the species.

The genomic architecture underlying ecological divergence and ecological speciation with gene flow is still largely unknown for most organisms. One central question is whether divergence is genome-wide or localized in 'genomic mosaics' during early stages when gene flow is still pronounced. Empirical work has so far been limited, and the relative impacts of gene flow and natural selection on genomic patterns have not been fully explored. Here, we use ecotypes of Atlantic cod to investigate genomic patterns of diversity and population differentiation in a natural system characterized by high gene flow and large effective population sizes, properties which theoretically could restrict divergence in local genomic regions. We identify a genomic region of strong population differentiation, extending over approximately 20 cM, between pairs of migratory and stationary ecotypes examined at two different localities. Furthermore, the region is characterized by markedly reduced levels of genetic diversity in migratory ecotype samples. The results highlight the genomic region, or 'genomic island', as potentially associated with ecological divergence and suggest the involvement of a selective sweep. Finally, we also confirm earlier findings of localized genomic differentiation in three other linkage groups associated with divergence among eastern Atlantic populations. Thus, although the underlying mechanisms are still unknown, the results suggest that 'genomic mosaics' of differentiation may even be found under high levels of gene flow and that marine fishes may provide insightful model systems for studying and identifying initial targets of selection during ecological divergence.

Haemoglobins of 0-group cod from 30 to 150 mm length were analysed by agar gel electrophoresis and isoelectric focusing to identify the lower size limit for determination of the haemoglobin polymorphisms in cod; both the main system which has been known for more than 30 years and the recently described subgroups. The juvenile fish studied had higher concentration of the HbII-component (split into three bands at these fish sizes) compared to older fish. The main haemoglobin types could be distinguished clearly and reliably at a fish size of about 35 mm and above. The subgroups of the HbI-components, however, could not be identified reliably until the cod were about 80 mm.

To study the population structure of tusk Brosme brosme in the North Atlantic, samples from six fishing grounds were collected: St Kilda; Storegga; Faroe Islands; Rockall bank; Davis Strait and East Greenland. In addition two samples from the Norwegian coast; Kragero and Oygarden, were included. Nine enzymes and haemoglobin were analysed by starch gel and agar gel electrophoresis. In addition isoelectric focusing (IEF) was carried out. At least 22 loci were represented. Polymorphisms were shown in eight loci and average heterozygosity was found to be about 0.05. Highest heterozygosity was found in haemoglobin analysed by Smithies buffer pH 8.6. Malate dehydrogenase (MDH-2) analysed in IEF (pH 3.5-9.5), and haemoglobin were found as valuable tools for studies of stock structure in tusk. In the Greenland samples only one of the MDH-2 heterozygotes was found, while in the East Atlantic both the observed heterozygotes were more frequent. The frequencies of the the Hb*b allele were about 0.40 and 0.70 in the samples from east and west Atlantic respectively.

Samples of blue mouth, Helicolenus dactylopterus (De La Roche, 1809). were collected in the areas between Shetland and the Faroe Islands, and analysed by starch gel electrophoresis and isoelectric focusing of haemoglobins and tissue enzymes. Moderate intraspecific variation was found, but most enzyme patterns were diagnostic for this species compared to the Sebastes species. The haemoglobin patterns were similar to the pattern of Sebastes marinus and S. viviparus, and thus diagnostic in relation to S. mentella. The estimated genetic distances between blue mouth and the Sebastes species were high (0.92–1.05) compared to corresponding values within the latter genus (0.03–0.13).