Cod, Gadus morhua L., populations identified by mitochondrial DNA

Isozyme and haemoglobin analysis has shown that the cod, Gadus morhua L., in the NE Atlantic can be regarded as two populations: Arctic and coastal cod. One hundred and one individual cod from nine different locations were sampled and restriction fragment analysis uncovered 14 different mtDNA clones. Calculation of sequence divergence between localities displayed large divergence between samples from coastal and Arctic areas (l.77–5.62%), as opposed to the low intrapopulation divergence (∼0.10%) and the divergence between localities along the coast (∼0.17%) and in the Barents Sea (∼ 1.00%).     

Minisatellite DNA fingerprints of Atlantic cod (Gadus morhua)

The human minisatellite probes 33.6 and 33.15 cross–hybridized to Hae III and Hinf I digested cod DNA, revealing complex fragment patterns in both Arctic and coastal cod. The DNA fingerprints were highly polymorphic, individual specific and stable in the germline. The potential applications of multi locus probes in cod research are discussed.     

Under-ice distribution of polar cod <Emphasis Type="Italic">Boreogadus saida</Emphasis> in the central Arctic Ocean and their association with sea-ice habitat properties

In the Arctic Ocean, sea-ice habitats are undergoing rapid environmental change. Polar cod (Boreogadus saida) is the most abundant fish known to reside under the pack-ice. The under-ice distribution, association with sea-ice habitat properties and origins of polar cod in the central Arctic Ocean, however, are largely unknown. During the RV Polarstern expedition ARK XXVII/3 in the Eurasian Basin in 2012, we used for the first time in Arctic waters a Surface and Under Ice Trawl with an integrated bio-environmental sensor array. Polar cod was ubiquitous throughout the Eurasian Basin with a median abundance of 5000 ind. km^?2. The under-ice population consisted of young specimens with a total length between 52 and 140 mm, dominated by 1-year-old fish. Higher fish abundance was associated with thicker ice, higher ice coverage and lower surface salinity, or with higher densities of the ice-amphipod Apherusa glacialis. The fish were in good condition and well fed according to various indices. Back-tracking of the sea-ice indicated that sea-ice sampled in the Amundsen Basin originated from the Laptev Sea coast, while sea-ice sampled in the Nansen Basin originated from the Kara Sea. Assuming that fish were following the ice drift, this suggests that under-ice polar cod distribution in the Eurasian Basin is dependent on the coastal populations where the sea-ice originates. The omnipresence of polar cod in the Eurasian Basin, in a good body condition, suggests that the central Arctic under-ice habitats may constitute a favourable environment for this species survival, a potential vector of genetic exchange and a recruitment source for coastal populations around the Arctic Ocean.     

Genetic divergence at the synaptophysin (Syp I) locus among Norwegian coastal and north-east Arctic populations of Atlantic cod

Several nuclear RFLP loci have been discovered recently that exhibit extensive allele frequency variation among Norwegian coastal and north-east Arctic populations of Atlantic cod Gadus morhua. One of these polymorphisms was detected by hybridizing an anonymous cDNA clone (GM798) against genomic DNA digested with the restriction enzyme DraI. This cDNA clone has now been sequenced and identified as synaptophysin (Syp I), an integral synaptic vesicle membrane protein. Primers were constructed that amplify an intron of the Syp I gene that is polymorphic for the DraI site, thus making it possible to use a PCR-based assay to score the polymorphism. A total of 965 individuals sampled from the Barents Sea, coastal areas and fjords in northern Norway have been analysed for this polymorphism. The results confirm that highly significant differences exist between NE Arctic and coastal cod at the Syp I locus. A cluster analysis revealed a deep split between coastal and Arctic populations and hierarchical F-statistics indicated that about 40% of the total variation was attributable to differences between Arctic and coastal groups. The temporal stability of allele frequencies was assessed by comparing Syp I allele frequencies among samples of juveniles (0 group) captured at specific locations in fjords in consecutive years and among samples of adults and juveniles collected from the same fjord. Samples of juveniles collected in 1994 and 1995 in Malangen were genetically indistinguishable whereas juveniles sampled from Dønnesfjord and Ullsfjorden over the same 2-year period exhibited significant differences. Adults and 0-group individuals collected from the same fjord were found to be genetically indistinguishable in Malangen, but not in Balsfjorden. In addition to detecting large differences among Arctic and coastal groups, the Syp I locus suggests that genetic heterogeneity exists among resident populations of cod in different fjords and that gene flow among populations throughout northern Norway may be considerably lower than previously believed.     

The isolation of Atlantic cod, Gadus morhua (Gadiformes), populations in Northern Meromictic lakes—A recurrent arctic phenomenon

Although it has previously been considered to be a rare phenomenon, this paper provides evidence of eight occurrences of Atlantic cod in northern coastal saline lakes in Arctic Canada, Greenland, Norway, and Russia. Historic and current habitat features that are necessary for Atlantic cod to colonize and persist in Arctic meromictic lakes are discussed. Data analyses confirm that the persistence of Atlantic cod in Arctic lakes is a recurrent phenomenon, and these populations represent an important component of intraspecific biodiversity. Published in Russian in Voprosy Ikhtiologii, 2008, Vol. 48, No. 2, pp. 179–190. The text was submitted by the authors in English.     

Distributional patterns of fishes in an Alaskan Arctic Lagoon

Summary In summer, the fish community of Simpson Lagoon and adjacent coastal waters of the Beaufort Sea was dominated by two marine species (Arctic cod, fourhorn sculpin) and three anadromous species (Arctic and least cisco, Arctic char). The anadromous species remained in the relatively warm and brackish waters near shore and demonstrated an affinity for shoreline edges, particularly the mainland shoreline where species occurrence and catch per unit effort (CPUE) were highest. Spatial segregation was low, presumably reflecting the migratory nature of these species. Marine species were less restricted to nearshore waters in summer and were typically the only species present in winter because anadromous species return to rivers, lakes and deltas to spawn or overwinter. Winter CPUE was low and consisted primarily of Arctic cod and fourhorn sculpin.     

Temperature-dependent growth and behavior of juvenile Arctic cod (<Emphasis Type="Italic">Boreogadus saida</Emphasis>) and co-occurring North Pacific gadids

The thermal sensitivity of Arctic fish species is poorly understood, yet such data are a critical component of forecasting and understanding ecosystem impacts of climate change. In this study, we experimentally measured temperature-dependent growth and routine swim activity in the juvenile stage of two Arctic gadids (Arctic cod, Boreogadus saida and saffron cod, Eleginus gracilis) and two North Pacific gadids (walleye pollock, Gadus chalcogrammus and Pacific cod, Gadus macrocephalus) over a 6-week growth period across five temperatures (0, 5, 9, 16 and 20 °C). Arctic cod demonstrated a cold-water, stenothermic response in that there was relatively high growth at 0 °C (0.73 % day^?1), near-maximal growth at 5 °C (1.35 % day^?1) and negative impacts on activity, growth and survival at 16 °C. In contrast, saffron cod demonstrated a warmer-water, eurythermic response, and temperature had a positive effect on growth and condition beyond 16 °C. However, despite these distinct thermal responses, walleye pollock and Pacific cod grew 2–3 times faster than Arctic gadids across a relatively broad temperature range above 5 °C. These results, coupled with possible northward expansion by both Pacific cod and walleye pollock, suggest Arctic cod are highly vulnerable to continued climate change in the Arctic, especially in coastal areas of the Beaufort and Chukchi Seas where temperatures already exceed 14 °C in the summer growth period.     

Pantophysin (Pan I) locus divergence between inshore v. offshore and northern v. southern populations of Atlantic cod in the north‐east Atlantic

More than 6000 cod Gadus morhua , sampled in coastal and offshore waters stretching from the Barents Sea down to the North Sea, were analysed for frequencies of alleles at the scnDNA pantophysin locus ( Pan I)[formerly called synaptophysin ( Syp I)]. The significant allele frequency difference between the two major stocks of cod in Norway, north‐east Arctic cod (NEAC) and Norwegian coastal cod (NCC), was upheld in all years of the investigation (1993 to 2001), and applied both to larval cod and post‐juveniles of various ages. On a north‐south axis, the appearance of a latitudinal cline of post‐juvenile (≥1 year) allele frequencies was exposed. The intermediate allele frequencies in coastal areas of northern Norway, seem to a large extent to be caused by intermingling of the two stocks, although the existence of populations of coastal cod with alternative Pan I frequencies could not be ruled out. The role of selection is yet unresolved. Depth of the sampling location seemed to have an effect on the allele frequencies and their temporal stability, while there was no indication of seasonal variation in the frequencies. Breeding structure was the most likely cause for upholding the extreme divergence in Pan I frequencies between NEAC and NCC.     

Natural selection and the genetic differentiation of coastal and Arctic populations of the Atlantic cod in northern Norway: a test involving nucleotide sequence variation at the pantophysin (PanI) locus

To examine the role of contemporary selection in maintaining significant allele frequency differences at the pantophysin (PanI) locus among populations of the Atlantic cod, Gadus morhua, in northern Norway, we sequenced 127 PanIA alleles sampled from six coastal and two Barents Sea populations. The distributions of variable sites segregating within the PanIA allelic class were then compared among the populations. Significant differences were detected in the overall frequencies of PanIA alleles among populations within coastal and Arctic regions that was similar in magnitude to heterogeneity in the distributions of polymorphic sites segregating within the PanIA allelic class. The differentiation observed at silent sites in the PanIA allelic class contradicts the predicted effects of widescale gene flow and suggests that postsettlement selection acting on cohorts cannot be responsible for the genetic differences described between coastal and Arctic populations. Our results suggest that the marked differences observed between coastal and Arctic populations of G. morhua in northern Norway at the PanI locus reflect the action of recent diversifying selection and that populations throughout the region may be more independent than suggested by previous studies.     

Mixed stock analysis and the power of different classes of molecular markers in discriminating coastal and oceanic Atlantic cod (<Emphasis Type="Italic">Gadus morhua</Emphasis> L.) on the Lofoten spawning grounds,Northern Norway

Atlantic cod (Gadus morhua) encompasses many different populations or stocks, which in part are managed separately. In the northeast Atlantic cod is divided into two main management units; northeast Arctic cod and coastal cod. These two groups have traditionally been separated by otolith classification. In this study, the power of different classes of genetic markers in separating the two cod groups was investigated. The variation in thirteen genetic markers, including allozymes, haemoglobin, the scDNA locus Pantophysin (Pan I) and a number of microsatellites was investigated, and mixed stock analysis and individual assignment tests were performed on samples comprising a mixture of individuals of putative coastal and oceanic type cod. The genetic analyses showed a large genetic differentiation between outer stations and stations located closer to the mainland shore. Mixed stock analysis and individual assignment tests used for estimation of stock proportions gave results similar to those obtained by otolith classification. Guest editors: J. Davenport, G. Burnell, T. Cross, M. Emmerson, R. McAllen, R. Ramsay & E. Rogan Challenges to Marine Ecosystems     

Occurrence of a gelatinous predator (<Emphasis Type="Italic">Cyanea capillata</Emphasis>) may affect the distribution of <Emphasis Type="Italic">Boreogadus saida</Emphasis>, a key Arctic prey fish species

Although the Arctic cod (Boreogadus saida) has a pan-Arctic distribution, little is known about its occurrence in near-shore waters where this species is the principal prey for seabirds, marine mammals and other fish. Published research describes the scyphomedusa Cyanea capillata as an Arctic cod predator, and this paper presents observations from long-term investigations using active hydroacoustics that suggest the Arctic cod avoided C. capillata in two small bays of Cornwallis Island (Canadian High Arctic archipelago). Distribution patterns in echograms suggested that features such as boundary layer fronts restricted jellyfish movements and Arctic cod were often abundant on the side of fronts where C. capillata were absent. Thus, habitat partitioning allowed Arctic cod to share habitat with its predator, albeit exceptions to this sharing occurred when jellyfish abundance was high and Arctic cod were displaced. Thus, if a warmer Arctic triggers an increase in C. capillata abundance, it is possible that small-scale aspects of Arctic cod distribution could be affected. This in turn could have significant ripple effects within the Arctic food web, an additional and previously unrecognized consequence of climate change.     

Variation in the diet of Arctic Cod (<Emphasis Type="Italic">Boreogadus saida</Emphasis>) in the Pacific Arctic and Bering Sea

Arctic cod, Boreogadus saida (Lepechin, 1774), is a nodal species in Arctic marine foodwebs as an important prey of many birds, marine mammals, and other fishes, as well as an abundant predator of zooplankton and epibenthic fauna. We examined the summer diet of Arctic cod across a latitudinal gradient extending from the southern limit of their distribution in the eastern Bering Sea to the northern margins of the eastern Chukchi Sea (ECS) continental shelf. Specimens were collected from demersal and pelagic trawls conducted between 1999 and 2012, and across a range of predator sizes (3–26 cm). Arctic cod diets vary with body size and between regions within the study area, and appear to vary between years in the eastern Bering Sea, indicating opportunistic feeding habits. Constrained Analysis of Principal Coordinates was conducted on ECS demersal samples and revealed consumption of fish and decapod crustacea were positively correlated with Arctic cod length while consumption of euphausiids and copepods had the opposite relationship. The demersal Arctic cod diet in the northern latitudes of the ECS was dominated by copepod consumption (47% by weight, %W), but copepods were less important (12–26%W) in the central and southern latitudes of the ECS and in the northern and eastern Bering Sea—areas where diets were more varied in their composition. High levels of variation in the diet of Arctic cod highlights the need to monitor Arctic cod diets to identify consistently dominant prey types and potential future changes to trophic relationships related to climate change or increasing anthropogenic activity.     

Ecology of Arctic cod <Emphasis Type="Italic">Boreogadus saida</Emphasis> (Gadiformes,Gadidae) and its fishery potential in Kara Sea

According to the bottom trawl-survey data, 97% of the ichthyomass in the southwestern region of the Kara Sea are composed of the Arctic cod Boreogadus saida; its stock is significantly higher than the previously registered resources. The Arctic cod is most unevenly distributed across the water area and capable to form the high-density aggregations, which can be caught by the targeted trawls. A wide range of the age composition (0+?6+), the size-age composition, and the growth rates of the Arctic cod in the trawl catches in the Kara Sea, which are different from those in the fish in the adjacent Barents Sea, can indicate their assignment of the Arctic cod in these seas to different populations.     

Gastric evacuation rates and daily rations of Arctic cod (Boreogadus saida) at low temperatures

Gastric evacuation rates were determined for different sizes of Arctic cod (Boreogadus saida) at sub-zero temperatures (−1.4 and −0.5°C). These temperatures represent ambient conditions for Arctic cod in the Canadian high Arctic. Evacuation half-times, the time required for half of the content of the stomach to be evacuated, were longer (36–70 h; mean=51 h) than those reported in studies carried out on other fish species. Gastric evacuation rates at low temperatures were equal to, or below, those predicted by extrapolation from experiments conducted at higher temperatures. There were no significant differences in evacuation rates among fish size-groups or diets, but evacuation rates were slower for fish that had been starved prior to experiments. Estimated daily rations for Arctic cod in Resolute Bay, N.W.T., were 0.51% body weight for small fish (4.5 g) and 1.13% body weight for large fish (51 g). Slow stomach evacuation rates at low temperatures may limit daily food intake when food is seasonally abundant. This may contribute to slow growth rates and limited maximum size of Arctic cod in Canadian high Arctic waters. Received: 14 July 1997 / Accepted: 15 November 1997     

Distinct patterns of Arctic cod (<Emphasis Type="Italic">Boreogadus saida</Emphasis>) presence and absence in a shallow high Arctic embayment,revealed across open-water and ice-covered periods through acoustic telemetry

With climate change resulting in unpredictable sea ice conditions between years, it is crucial to gain a more comprehensive understanding of the subsequent effects on Arctic marine ecosystems. Arctic cod (Boreogadus saida) play a key role in the Arctic marine food web, serving as a food source that is estimated to contribute up to 75 % of energy transfer to higher trophic levels. To investigate Arctic cod residency and distribution in Resolute Bay (74°44′N, 095°04′W), 85 individuals from four locations in the bay were captured, measured, weighed, implanted with acoustic tags and subsequently tracked on an acoustic array of 49 receivers. Two main periods of residence in the bay were identified, the first in open water and the second under ice cover, and both concluded with a collective mass departure of fish. A generalised linear mixed model was used to investigate the influence of variables on Arctic cod presence/absence in the bay, indicating that ingress and egress were influenced by environmental changes, particularly those associated with the transition from open-water to the ice-covered period. Timing and distribution, during the study period, appeared to be influenced by a combination of physiological acclimation, and a balance between resource availability and refuge from predators. Receiver site Residence Index (RI) analysis revealed strong site fidelity of fish towards the northern areas of the bay, and this behaviour was consistent between tagging groups and individuals, indicating that the majority of tagged cod were representative of a single school. This study represents the first employment of acoustic telemetry to monitor the movements of individual Arctic cod over 9 months, incorporating both open-water and ice-covered periods.     

An assessment of mitochondrial variation in Arctic gadoids

Climatic changes during the quaternary history in Arctic regions have shaped the genetic variation and genealogies of Arctic species. Several studies have been conducted in recent years on genetic diversity of Arctic organisms, but marine fishes are largely underrepresented in these studies. Here, we present a study on mitochondrial variation in three Arctic gadoids: Arctic cod (Arctogadus glacialis), Greenland cod (Gadus ogac), and Polar cod (Boreogadus saida). In addition, geographic variation in Polar cod is presented. The sequence variation at the mtDNA presents similar patterns as observed for other related marine fishes. Variation in these three species reflects rather different historic processes, due to colonization and climatic changes than differences in life histories. In Polar cod, a deeper genealogy is observed and variation is dependent on both latitude and longitude. The deep genealogy indicates either admixture of separate lineages or a population, which has been stable in size during alternating cold and warm periods of the pleistocene. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

New data on the longevity of coastal cod Gadus morhua Linnaeus, 1758 in the White Sea

The cod, Gadus morhua, is a common and abundant demersal fish in the White Sea coastal zone. The published data on the maximum age of White Sea cod still requires additional documentation, because some authors have noted difficulties in age determination by otoliths of large specimens. To obtain accurate data on the longevity of this species in the White Sea, an image analysis of thin‐sections of otoliths was made for age estimation. Research surveys in the Chupa Inlet and adjacent waters of the Kandalaksha Bay were conducted in June to August from 2007 to 2013. The five largest specimens of a total of 3564 captured fish were selected for age determination. Age of these largest individuals ranged between 7 and 12 years, total length and weight varied from 60.2 to 77.0 cm and from 2.4 to 6.1 kg, respectively. According to the data, maximal White Sea cod longevity is greater than in previously published data, and demonstrates similar longevity to conspecifics from the Baltic and North seas, whereas its life cycle is much shorter than cod from the Northeast Arctic, Iceland, Greenland, Newfoundland and Labrador stocks.     

Summer diet of beluga whales inferred by fatty acid analysis of the eastern Beaufort Sea food web

Beluga whales (Delphinapterus leucas) are the most abundant odontocetes in Arctic waters and are thus thought to influence food web structure and function. The diet of the Beaufort Sea beluga population is not well known, partly due to the inherent difficulty of observing feeding behaviour in Arctic marine cetaceans. To determine which prey items are critical to the Beaufort Sea beluga diet we first examine and describe the Mackenzie Delta and Beaufort Sea food web using fatty acid analyses. Fatty acid profiles effectively partitioned prey items into groups associated with their habitat and feeding ecology. Next, the relative contribution of various prey items to beluga diet was investigated using fatty acids. Finally, beluga diet variability was examined as a function of body size, a known correlate of habitat use. Beluga appeared to feed predominantly on Arctic cod (Boreogadus saida) collected from near shore and offshore regions. Size related dietary differences suggested larger sized beluga preferred offshore Arctic cod given the shared high levels of long chain monounsaturates, whereas smaller sized beluga appeared to feed on prey in their near shore habitats that included near shore Arctic cod. The presence of Arctic cod groups in shallow near shore and deep offshore habitats may facilitate the behavioural segregation of beluga habitat use as it relates to their size and resource requirements. Given Arctic cod are a sea ice associated fish combined with the accelerated sea ice loss in this region, beluga whales may need to adapt to new dietary regimes.     

Comparing microsatellite variation in north-east Atlantic cod (Gadus morhua L.) to genetic structuring as revealed by the pantophysin (Pan I) locus

Variation at seven microsatellite loci was compared with variation observed at Pan I, a single copy nuclear DNA gene coding for pantophysin, in 14 samples of Atlantic cod ( Gadus morhua ) stretching from Spitsbergen to the North Sea. Population structuring indicated by the two types of markers was concordant and in agreement with the traditional grouping of cod in the study area into three main populations: north-east Arctic cod (NEAC), Norwegian coastal cod (NCC) and North Sea cod (NSC). Microsatellites, however, revealed genetic heterogeneity not only within NCC, as did Pan I, but also within NEAC and NSC, which appeared to be homogenous when analysed for Pan I. Moreover, microsatellites displayed lower levels of differentiation than Pan I between NEAC and two other groups. Differences in the magnitude of differentiation for the two types of markers may be attributable to higher levels of polymorphism and alleged selective neutrality of microsatellites. Isolation by distance was clearly apparent for microsatellites but was less evident for Pan I, indicating that environmentally induced selection appears to shape the patterns of genetic differentiation for this marker. Even though the population structure of north-east Atlantic cod, as revealed by microsatellites and Pan I, appears to be maintained largely by restricted gene flow, selection acting on a recent historical time scale probably contributes to the observed geographic pattern of Pan I frequencies.