Norwegian Coastal Cod Research Articles

Sex-specific vertical movements of spawning atlantic cod in coastal habitats inferred from acoustic telemetry.

Fish spawning location and behaviour can be challenging to detect, especially in deep water. Here we utilise two large acoustic telemetry datasets from western and mid Norway to study the vertical movement dimension of individual Atlantic cod Gadus morhua spawning behaviour in their natural habitats. The datasets comprise ~ 6million depth detections from 644 sexed, mature cod, collected at seven different Norwegian coastal cod spawning grounds during 2017-2021. During the spawning period, females were typically found at shallower depths compared to males, while this pattern was much less pronounced or absent outside this period. Furthermore, we identify a hitherto undescribed sex-dependent vertical movement behaviour: periodic descents, where females made deep dives, putatively associated with egg release and spawning, approximately every third day. Females not showing this behaviour tended to occupy deeper positions in the water column than the "periodic descent" females and hence may approach and retreat from spawning males more horizontally. These observed sex-specific behaviours correspond well to the lek-like description of the cod mating system and with cod reproductive physiology, where females are known to mature and release batches of eggs at intervals of several days as part of a bet-hedging strategy.

Quota Consolidation in Norwegian Coastal Fisheries

AbstractBalancing the trade-off between economic efficiency and social objectives has been a challenge for natural resource managers under rights-based management. While the actual prioritization should be guided by social preferences, the mechanisms and consequences of the quota transfer system need to be well understood. We investigate the effects of the quota transfer scheme implemented in the Norwegian coastal cod fishery in 2004. This is a small-scale fishery that has traditionally been important for employment in the northern part of Norway. Using vessel-level quota registry data, we estimate the effect of quota trading on vessel exit using a difference-in-differences approach that exploits variation in implementation timing between regulatory groups. In addition, we describe the outcome of quota consolidation with descriptive statistics. Our results confirm that the quota transfer scheme accelerates the exit of vessels from the fishery by at least 5% points in the short run. The descriptive analysis reveals great heterogeneity in both the size and geographic location of exiting vessels, suggesting that the scheme has distributional impacts. While the policy change has the expected effects in the short run, our results suggest that the implications of consolidation last longer. This has implications for policymakers trying to balance economic efficiency and social objectives of rational fishery management.

Understanding the organisational structure of fisheries crime in well-regulated fisheries

Illegal, Unreported and Unregulated (IUU) fishing is recognised as a global environmental, economic, and social problem, taking place in all kinds of fisheries. Preventing it is however difficult and there is a continuous need to expand the knowledge base on how the issue can be addressed. In this article we study fisheries crime from an environmental criminology approach by conducting a crime script analyses to describe the organisational structure of unreported fishing in a well-regulated fishery. The approach gives detailed insights into the different steps in the crime commission process in the Norwegian coastal cod fisheries. The crime script technique is expanded to also include an analysis of the regulatory framework designed to prevent illegal fishing activities. The main MCS mechanisms to prevent illegalities are present but the diversity in the industry makes the implementation of universal prevention mechanisms difficult. The analysis highlights the fisher-buyer dialogue and interactions prior to misreporting as a core aspect to the organisation of the violations, and difficult to regulate. When linking regulations and guardianship to different steps of the crime we discuss the identified vulnerabilities in the resource control system and present some possible intervention points and prevention mechanisms to be considered by policymakers.

Movement diversity and partial sympatry of coastal and Northeast Arctic cod ecotypes at high latitudes.

Movement diversity within species represent an important but often neglected, component of biodiversity that affects ecological and genetic interactions, as well as the productivity of exploited systems. By combining individual tracking data from acoustic telemetry with novel genetic analyses, we describe the movement diversity of two Atlantic cod Gadus morhua ecotypes in two high-latitude fjord systems: the highly migratory Northeast Arctic cod (NEA cod) that supports the largest cod fishery in the world, and the more sedentary Norwegian coastal cod, which is currently in a depleted state. As predicted, coastal cod displayed a higher level of fjord residency than NEA cod. Of the cod tagged during the spawning season, NEA cod left the fjords permanently to a greater extent and earlier compared to coastal cod, which to a greater extent remained resident and left the fjords temporarily. Despite this overall pattern, horizontal movements atypical for the ecotypes were common with some NEA cod remaining within the fjords year-round and some coastal cod displaying a low fjord fidelity. Fjord residency and exit timing also differed with spawning status and body size, with spawning cod and large individuals tagged during the feeding season more prone to leave the fjords and earlier than non-spawning and smaller individuals. While our results confirm a lower fjord dependency for NEA cod, they highlight a movement diversity within each ecotype and sympatric residency between ecotypes, previously undetected by population-level monitoring. This new knowledge is relevant for the management, which should base their fisheries advice for these interacting ecotypes on their habitat use and seasonal movements.

Characterizing coastal cod vocalization using a towed hydrophone array

Abstract To better understand spawning vocalizations of Norwegian coastal cod (Gadus morhua), a prototype eight-element coherent hydrophone array was deployed in stationary vertical and towed horizontal modes to monitor cod sounds during an experiment in spring 2019. Depth distribution of cod aggregations was monitored concurrently with an ultrasonic echosounder. Cod vocalizations recorded on the hydrophone array are analysed to provide time–frequency characteristics, and source level distribution after correcting for one-way transmission losses from cod locations to the hydrophone array. The recorded cod vocalization frequencies range from ∼20 to 600 Hz with a peak power frequency of ∼60 Hz, average duration of 300 ms, and mean source level of 163.5 ± 7.9 dB re 1 μPa at 1 m. Spatial dependence of received cod vocalization rates is estimated using hydrophone array measurements as the array is towed horizontally from deeper surrounding waters to shallow water inlet areas of the experimental site. The bathymetric-dependent probability of detection regions for cod vocalizations are quantified and are found to be significantly reduced in shallow-water areas of the inlet. We show that the towable hydrophone array deployed from a moving vessel is invaluable because it can survey cod vocalization activity at multiple locations, providing continuous spatial coverage that is complementary to fixed sensor systems that provide continuous temporal coverage at a given location.

Detecting significant retrospective patterns in state space fish stock assessment

Retrospective patterns are commonly investigated to validate fish stock assessment models. A widely applied measure for retrospective bias is Mohn’s ρ and corresponding retrospective plots. However, retrospective patterns can be interpreted differently by experts. To make decisions regarding significant retrospective patterns less subjective, we proposed a post-sample Mohn’s ρ significance test. As case studies, we applied the state space assessment model SAM with data on Northeast Arctic cod and Norwegian coastal cod north of 67°N. We showed that the acceptance regions of Mohn’s ρ depends on both the data available and the assessment model complexity. We also assessed the test power under a range of assumption violations and conclude that Mohn’s ρ is useful for detecting violations associated with bias, but not for violations associated with variances and correlations.

Exploring fishery history in game form: ‘Never again April 18!’

ABSTRACT This article studies the ways in which Norwegian fishery history can be explored through games. Using the 1989 closure of the Norwegian coastal cod commons as a case study, issues related to historical thinking and game studies are discussed. The main focus is on understanding history through serious games, but theoretical considerations for presenting the case in any game format are discussed. The case involves a historical resource crisis, and the article traces how a serious game can frame counterfactual imagination for questioning the institutional politics of resource management as well as for producing historical empathy with stakeholders in resource crises.

Genomic analysis reveals neutral and adaptive patterns that challenge the current management regime for East Atlantic cod Gadus morhua L.

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.

Ontogenetic movements of cod in Arctic fjords and the Barents Sea as revealed by otolith microchemistry

The distribution of Atlantic cod (Gadus morhua) in northern Norwegian waters is expanding eastward and northward in the Barents Sea and along western Svalbard. In the Arctic fjords of Svalbard, cod has become abundant, but little is known about the biology, origin, or residence patterns of these populations. To address this issue, we used laser ablation inductively coupled plasma mass spectrometry to quantify the trace elemental composition of cod otoliths at age-0, age-3 and the year of spawning at five distinct locations in northern Norway and western Svalbard. Chemical composition data was used to identify natal sources of cod, their broad-scale migration patterns, and to determine if cod are currently resident in Arctic fjords. Our results suggest that cod collected at Kongsfjord, Isfjord, outside Svalbard, Lofoten, and Porsangerfjord were recruited mainly from the Barents Sea, conforming to the Northeast Arctic cod ecotype. The degree of chemical overlap between Porsangerfjord and Isfjord cod, however, varied with fish age, suggesting individual movements consistent with the Norwegian coastal cod ecotype. Finally, the chemical composition of mature fish at Isfjord, and to a lesser extent Kongsfjord, suggests that cod from the Barents Sea might have recently established residency in these two Arctic fjords.

Otolith growth and zone formation during first maturity and spawning of Atlantic cod (Gadus morhua)

Specific impacts of somatic growth, sexual maturation, and spawning events on otolith zone formation in Atlantic cod (Gadus morhua) were assessed in a 33-month tank experiment, using Barents Sea cod and Norwegian coastal cod. High and low feeding ration combinations were used to mimic environmental stressors in the field. For both stocks, apparent macrostructural “spawning zones” in otoliths are registered in statutory stock monitoring programs to estimate age at maturity, thus adding key information to stock biomass assessments. We found that substantial energy investments in reproduction caused reductions in otolith growth and altered proportional width between translucent and opaque zones. These effects, however, were only statistically significant among individuals with high reproductive investments, while otoliths from individuals with low investments did not differ from the otoliths for immatures. Reproduction may thus not necessarily induce spawning zones, and alternatively, spawning zones may not necessarily reflect reproduction. Altogether, this suggests that the individual energy level, as a premise for metabolic activity, plays a key role in the formation of such zones and thus is related to environmental conditions.

Closely-related Photobacterium strains comprise the majority of bacteria in the gut of migrating Atlantic cod (Gadus morhua)

BackgroundThe population of Atlantic cod (Gadus morhua), also known as Northeast Arctic cod, migrating Atlantic cod, or simply “skrei,” lives mainly in the Barents Sea and Svalbard waters and migrates in annual cycles to the Norwegian coast in order to spawn eggs during late winter. It is the world’s largest population of Atlantic cod, and the population is distinct from the Norwegian coastal cod (or “fjord” cod). Despite the biological, economic, and cultural importance of migrating Atlantic cod, current knowledge on the associated microbiota is very limited. Using shotgun metagenomics and metaproteomics approaches, we present here the gut microbiota, metagenome-assembled genomes (MAGs) of the most abundant bacterial species, DNA-based functional profile, and the metaproteome of Atlantic cod specimens caught at a spawning area in an open ocean outside of Tromsø, Norway.ResultsOur analyses identified 268 bacterial families in DNA isolated from feces of 6 individual migrating Atlantic cod. The most abundant family was Vibrionaceae (52%; 83% if unclassified reads are excluded), with Photobacterium (genus) representing the vast majority. The recovery of metagenome-assembled genomes provided further details and suggests that several closely related Photobacterium strains from the Photobacterium phosphoreum clade are the most abundant. A genomic-based functional profiling showed that the most abundant functional subsystems are “Carbohydrates”; “Amino Acids and Derivatives”; “Protein Metabolism”; “Cofactors, Vitamins, Prosthetic, Groups, and Pigments”; and “DNA Metabolism,” which is in agreement with other studies of gut microbiomes of marine organisms. Finally, the MS-based metaproteomic dataset revealed that the functional category “Protein Metabolism” is highly overrepresented (3×) when compared to the genome-based functional profile, which shows that ribosomal proteins are rich in the bacterial cytosol.ConclusionWe present here the first study of bacterial diversity of the gut of migrating Atlantic cod using shotgun sequencing and metagenome-assembled genomes (MAGs). The most abundant bacteria belong to the Photobacterium genus (Vibrionaceae family). We also constructed functional profiles of the gut microbiome. These may be used in future studies as a platform for mining of commercially interesting cold-active enzymes.

Analysis of coastal cod (Gadus morhua L.) sampled on spawning sites reveals a genetic gradient throughout Norway\u2019s coastline

BackgroundAtlantic 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.ResultsWe 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 < 0.0001. Dataset 2: global FST = 0.004, P < 0.0001). While no clear genetic groups were identified, genetic differentiation increased among geographically-distinct samples. Although the locus Gmo132 was identified as a candidate for positive selection, possibly through linkage with a genomic region under selection, overall trends remained when this locus was excluded from the analyses. The most common allele in loci Gmo132 and Gmo34 showed a marked frequency change in the north-south gradient, increasing towards the frequency observed in NEAC in the north.ConclusionWe conclude that Norwegian coastal cod displays significant population genetic structure throughout its entire range, that follows a trend of isolation by distance. Furthermore, we suggest that a gradient of genetic introgression between NEAC and NCC contributes to the observed population genetic structure. The current management regime for coastal cod in Norway, dividing it into two stocks at 62°N, represents a simplification of the level of genetic connectivity among coastal cod in Norway, and needs revision.

Genetic management of mixed-stock fisheries “real-time”: The case of the largest remaining cod fishery operating in the Atlantic in 2007–2017

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.

“Real-time” genetic monitoring of a commercial fishery on the doorstep of an MPA reveals unique insights into the interaction between coastal and migratory forms of the Atlantic cod

AbstractWith 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, &amp;gt;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.

Diet of Norwegian coastal cod (Gadus morhua) studied by using citizen science

Abstract The Norwegian coastal cod ( Gadus morhua ) is a keystone species in the food web of northern Norwegian fjords. Their relatively stationary populations might specifically depend on local food resources, but the diet of cod has rarely been studied in fjord systems. Using a citizen science approach, where recreational anglers and tourists participated in the sampling, we studied small-scale differences in the diet composition of cod in a fjord system in northern Norway. We compared the cod diet from the MPA Saltstraumen, characterised by strong tidal currents and a highly diverse and abundant fauna, with the inner fjord area of Skjerstadfjord. The diet composition of cod significantly differed between both areas within the fjord. Although fish was the dominant prey in both areas, cod consumed > 40% invertebrates in terms of weight, even in the cod size class of 70–99 cm. The invertebrate prey also caused the observed spatial differences. In Saltstraumen, brittle stars (Ophiuroidea), crabs (Brachyura) and sea cucumbers (Holothuroidea) were important food sources for cod, while sea urchins (Echinoidea), clams (Bivalvia), shrimps (Caridea) and krill (Euphausiacea) dominated the diet in the inner Skjerstadfjord. The high densities of sessile fauna in the dynamic environment of Saltstraumen, was only partly reflected in the diet of cod, with only Holothuroidea found in 17% of the stomachs. High rates of empty stomachs (24%), cannibalism as well as a higher proportion of low-energy prey in the diet of large cod, may indicate a shortage of high quality food in Skjerstadfjord. The samples for this study were collected through a citizen science campaign. This approach might provide opportunities to be used for coastal ecological monitoring with potential applications in local ecosystem management strategies through public involvement.

Environmental, population and life-stage plasticity in the visual system of Atlantic cod.

The visual system is for many fishes essential in guiding behaviors, such as foraging, predator avoidance and mate choice. The marine environment is characterized by large spatio-temporal fluctuations in light intensity and spectral composition. However, visual capabilities are restricted by both space limitations set by eye size and by the genomic content of light-absorbing opsin genes. The rich array of visual opsins in teleosts may be used differentially to tune vision towards specific needs during ontogeny and to changing light. Yet, to what extent visual plasticity is a pre-programmed developmental event, or is triggered by photic environment, is unclear. Our previous studies on Atlantic cod revealed an evolutionary genomic loss of UV-sensitive sws1 and red-sensitive lws opsin families, while blue-sensitive sws2 and green-sensitive rh2 opsins had duplicated. The current study has taken an opsin expression approach to characterize visual plasticity in cod towards different spectral light during the larval stage, to maturation and extreme seasonal changes in the Barents Sea. Our data suggest that opsin plasticity in cod larvae is controlled by developmental programme rather than immediate light environment. The lack of expressional changes during maturation suggests a less important role for visual modulation related to mate choice. Although no seasonal effects on visual opsins were detected in migratory Northeast Arctic cod, the expressed opsin subset differed from the more stationary Norwegian coastal cod described in previous studies. Interestingly, these data provide the first indications of a population difference in actively used visual opsins associated with cod ecotypes.

Spatial variation in fecundity of Norwegian coastal cod, Gadus morhua (Linnaeus), along the coast of Norway

Fecundity of Norwegian coastal cod (NCC) was estimated at several locations along the coast of Norway. There was no significant difference in carcass weight, liver weight or potential fecundity (Fp) between Lofoten and Verrabotn (Northern region) or between Bømlo and Langesund (Southern region). Fish caught in the Northern region had a higher liver weight and potential fecundity than fish caught in the Southern region.

Two adjacent inversions maintain genomic differentiation between migratory and stationary ecotypes of Atlantic cod.

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 ~1.6-2.0Mya. 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.

Three chromosomal rearrangements promote genomic divergence between migratory and stationary ecotypes of Atlantic cod.

Identification of genome-wide patterns of divergence provides insight on how genomes are influenced by selection and can reveal the potential for local adaptation in spatially structured populations. In Atlantic cod – historically a major marine resource – Northeast-Arctic- and Norwegian coastal cod are recognized by fundamental differences in migratory and non-migratory behavior, respectively. However, the genomic architecture underlying such behavioral ecotypes is unclear. Here, we have analyzed more than 8.000 polymorphic SNPs distributed throughout all 23 linkage groups and show that loci putatively under selection are localized within three distinct genomic regions, each of several megabases long, covering approximately 4% of the Atlantic cod genome. These regions likely represent genomic inversions. The frequency of these distinct regions differ markedly between the ecotypes, spawning in the vicinity of each other, which contrasts with the low level of divergence in the rest of the genome. The observed patterns strongly suggest that these chromosomal rearrangements are instrumental in local adaptation and separation of Atlantic cod populations, leaving footprints of large genomic regions under selection. Our findings demonstrate the power of using genomic information in further understanding the population dynamics and defining management units in one of the world’s most economically important marine resources.

Growth of the sword razor (Ensis magnus) early stages: modelling juvenile seasonal growth

Growth of the sword razor (Ensis magnus) early stages: modelling juvenile seasonal growth