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Long-term changes in the age and size structure of animal populations are well documented, yet their impacts on population productivity are poorly understood. Fishery exploitation can be a major driver of changes in population age-size structure because fisheries significantly increase mortality and often selectively remove larger and older fish. Climate change is another potential driver of shifts in the demographic structure of fish populations. Northeast Arctic (NEA) cod is the largest population of Atlantic cod (Gadus morhua) and one of the world's most important commercial fish stocks. This population has experienced considerable changes in population age-size structure over the past century, largely in response to fishing. In this study, we investigate whether changes in spawner age structure have affected population productivity in NEA cod, measured as recruits per spawning stock biomass, over the past 75 years. We find evidence that shifts in age structure toward younger spawners negatively affect population productivity, implying higher recruitment success when the spawning stock is composed of older individuals. The positive effect of an older spawning stock is likely linked to maternal effects and higher reproductive output of larger females. Our results indicate a threefold difference in productivity between the youngest and oldest spawning stock that has been observed since the 1950s. Further, our results suggest a positive effect of environmental temperature and a negative effect of intraspecific cannibalism by older juveniles on population productivity, which partly masked the effect of spawner age structure unless accounted for in the model. Collectively, these findings emphasize the importance of population age structure for the productivity of fish populations and suggest that harvest-induced demographic changes can have negative feedbacks for fisheries that lead to a younger spawning stock. Incorporating demographic data into harvest strategies could thus facilitate sustainable fishery management., (© 2022 The Ecological Society of America.)

Sustainable human exploitation of living marine resources stems from a delicate balance between yield stability and population persistence to achieve socioeconomic and conservation goals. But our imperfect knowledge of how oceanic oscillations regulate temporal variation in an exploited species can obscure the risk of missing management targets. We illustrate how applying a management policy to suppress fluctuations in fishery yield in variable environments (prey density and regional climate) can present unintended outcomes in harvested predators and the sustainability of harvesting. Using Atlantic cod (Gadus morhua, an apex predatory fish) in the Barents Sea as a case study we simulate age-structured population and harvest dynamics through time-varying, density-dependent and density-independent processes with a stochastic, process-based model informed by 27-year monitoring data. In this model, capelin (Mallotus villosus, a pelagic forage fish), a primary prey of cod, fluctuations modulate the strength of density-dependent regulation primarily through cannibalistic pressure on juvenile cod survival; sea temperature fluctuations modulate thermal regulation of cod feeding, growth, maturation, and reproduction. We first explore how capelin and temperature fluctuations filtered through cod intrinsic dynamics modify catch stability and then evaluate how management to suppress short-term variability in catch targets alters overharvest risk. Analyses revealed that suppressing year-to-year catch variability impedes management responses to adjust fishing pressure, which becomes progressively out of sync with variations in cod abundance. This asynchrony becomes amplified in fluctuating environments, magnifying the amplitudes of both fishing pressure and cod abundance and then intensifying the density-dependent regulation of juvenile survival through cannibalism. Although these transient dynamics theoretically give higher average catches, emergent, quasicyclic behaviors of the population would increase long-term yield variability and elevate overharvest risk. Management strategies that overlook the interplay of extrinsic (fishing and environment) and intrinsic (life history and demography) fluctuations thus can inadvertently destabilize fish stocks, thereby jeopardizing the sustainability of harvesting. These policy implications underscore the value of ecosystem approaches to designing management measures to sustainably harvest ecologically connected resources while achieving socioeconomic security., (© 2021 The Authors. Ecological Applications published by Wiley Periodicals LLC on behalf of The Ecological Society of America.)

The Arctic Ocean is an early warning system for indicators and effects of climate change. We use a novel combination of experimental and time-series data on effects of ocean warming and acidification on the commercially important Northeast Arctic cod (Gadus morhua) to incorporate these physiological processes into the recruitment model of the fish population. By running an ecological-economic optimization model, we investigate how the interaction of ocean warming, acidification and fishing pressure affects the sustainability of the fishery in terms of ecological, economic, social and consumer-related indicators, ranging from present day conditions up to future climate change scenarios. We find that near-term climate change will benefit the fishery, but under likely future warming and acidification this large fishery is at risk of collapse by the end of the century, even with the best adaptation effort in terms of reduced fishing pressure., Competing Interests: The authors have declared that no competing interests exist.

Collapses and regime changes are pervasive in complex systems (such as marine ecosystems) governed by multiple stressors. The demise of Atlantic cod ( Gadus morhua) stocks constitutes a text book example of the consequences of overexploiting marine living resources, yet the drivers of these nearly synchronous collapses are still debated. Moreover, it is still unclear why rebuilding of collapsed fish stocks such as cod is often slow or absent. Here, we apply the stochastic cusp model, based on catastrophe theory, and show that collapse and recovery of cod stocks are potentially driven by the specific interaction between exploitation pressure and environmental drivers. Our statistical modelling study demonstrates that for most of the cod stocks, ocean warming could induce a nonlinear discontinuous relationship between fishing pressure and stock size, which would explain hysteresis in their response to reduced exploitation pressure. Our study suggests further that a continuing increase in ocean temperatures will probably limit productivity and hence future fishing opportunities for most cod stocks of the Atlantic Ocean. Moreover, our study contributes to the ongoing discussion on the importance of climate and fishing effects on commercially exploited fish stocks, highlighting the importance of considering discontinuous dynamics in holistic ecosystem-based management approaches, particularly under climate change.

Climate warming and harvesting affect the dynamics of species across the globe through a multitude of mechanisms, including distribution changes. In fish, migrations to and distribution on spawning grounds are likely influenced by both climate warming and harvesting. The Northeast Arctic (NEA) cod (Gadus morhua) performs seasonal migrations from its feeding grounds in the Barents Sea to spawning grounds along the Norwegian coast. The distribution of cod between the spawning grounds has historically changed at decadal scales, mainly due to variable use of the northern and southern margins of the spawning area. Based on historical landing records, two major hypotheses have been put forward to explain these changes: climate and harvesting. Climate could affect the distribution through, for example, spatial habitat shifts. Harvesting could affect the distribution through impacting the demographic structure. If demographic structure is important, theory predicts increasing spawner size with migration distance. Here, we evaluate these hypotheses with modern data from a period (2000-2016) of increasing temperature and recovering stock structure. We first analyze economic data from the Norwegian fisheries to investigate geographical differences in size of spawning fish among spawning grounds, as well as interannual differences in mean latitude of spawning in relation to changes in temperature and demographic parameters. Second, we analyze genetically determined fish sampled at the spawning grounds to unambiguously separate between migratory NEA cod and potentially smaller sized coastal cod of local origin. Our results indicate smaller spawners farther away from the feeding grounds, hence not supporting the hypothesis that harvesting is a main driver for the contemporary spawning ground distribution. We find a positive correlation between annual mean spawning latitude and temperature. In conclusion, based on contemporary data, there is more support for climate compared to harvesting in shaping spawning ground distribution in this major fish stock in the North Atlantic Ocean., (© 2018 John Wiley & Sons Ltd.)

Predicting fish stock variations on interannual to decadal time scales is one of the major issues in fisheries science and management. Although the field of marine ecological predictions is still in its infancy, it is understood that a major source of multi-year predictability resides in the ocean. Here we show the first highly skilful long-term predictions of the commercially valuable Barents Sea cod stock. The 7-year predictions are based on the propagation of ocean temperature anomalies from the subpolar North Atlantic toward the Barents Sea, and the strong co-variability between these temperature anomalies and the cod stock. Retrospective predictions for the period 1957-2017 capture well multi-year to decadal variations in cod stock biomass, with cross-validated explained variance of over 60%. For lead times longer than one year the statistical long-term predictions show more skill than operational short-term predictions used in fisheries management and lagged persistence forecasts. Our results thus demonstrate the potential for ecosystem-based fisheries management, which could enable strategic planning on longer time scales. Future predictions show a gradual decline in the cod stock towards 2024., Competing Interests: The authors have declared that no competing interests exist.

Sustainability of commercial fisheries is best achieved when fishing gears are selective and have low impacts on bottom habitat. Pots (baited traps) are a fishing technology that typically has lower impacts than many other industrial gears. In this study we compared the efficiency of five models of pots (baited traps) designed to catch Atlantic cod (Gadus morhua) for use in Newfoundland and Labrador (NL)'s expanding cod fishery. We compared catch per unit effort (CPUE) and total lengths of cod across each pot type, as well as bycatch rates of each model. All pot types were successful at catching cod, but two models (the modified Newfoundland pot, and a four-entrance pot of our design) had highest CPUE. Specifically, we found that modifying Newfoundland pots increased their CPUE by 145% without a corresponding increase in bycatch. None of the pot types produced substantial amounts of bycatch. This study demonstrated that potting gear is an effective way to catch cod in NL, and that there is flexibility in which pot fishers can use, depending on the layout of their fishing vessel., Competing Interests: The authors have declared that no competing interests exist.

Commercial fishing generally removes large and old individuals from fish stocks, reducing mean age and age diversity among spawners. It is feared that these demographic changes lead to lower and more variable recruitment to the stocks. A key proposed pathway is that juvenation and reduced size distribution causes reduced ranges in spawning period, spawning location, and egg buoyancy; this is proposed to lead to reduced spatial distribution of fish eggs and larvae, more homogeneous ambient environmental conditions within each year-class, and reduced buffering against negative environmental influences. However, few, if any, studies have confirmed a causal link from spawning stock demographic structure through egg and larval distribution to year class strength at recruitment. We here show that high mean age and size in the spawning stock of Barents Sea cod (Gadus morhua) is positively associated with high abundance and wide spatiotemporal distribution of cod eggs. We find, however, no support for the hypothesis that a wide egg distribution leads to higher recruitment or a weaker recruitment-temperature correlation. These results are based on statistical analyses of a spatially resolved data set on cod eggs covering a period (1959-1993) with large changes in biomass and demographic structure of spawners. The analyses also account for significant effects of spawning stock biomass and a liver condition index on egg abundance and distribution. Our results suggest that the buffering effect of a geographically wide distribution of eggs and larvae on fish recruitment may be insignificant compared with other impacts., Competing Interests: The authors declare no conflict of interest.

Pershing et al (Science, 13 November 2015, p. 809) concluded that recent warming in the Gulf of Maine contributed to the collapse of Gulf of Maine cod. We argue that this conclusion is based on a flawed analysis of the population dynamics of this cod stock. We believe that understanding the potential role of climate change in the collapse of this stock requires more defensible analyses., (Copyright © 2016, American Association for the Advancement of Science.)

Pershing et al (Reports, 13 November, p. 809) concluded that failure to account for temperature in the assessment and management of Gulf of Maine Atlantic cod caused overfishing. We argue that the "extra mortality" calculation driving this conclusion is an artifact. Environmental factors affect all stocks, but attribution of additional mortality to temperature alone by Pershing et al is unsupported by the data., (Copyright © 2016, American Association for the Advancement of Science.)

Palmer et al and Swain et al suggest that our "extra mortality" time series is spurious. In response, we show that including temperature-dependent mortality improves abundance estimates and that warming waters reduce growth rates in Gulf of Maine cod. Far from being spurious, temperature effects on this stock are clear, and continuing to ignore them puts the stock in jeopardy., (Copyright © 2016, American Association for the Advancement of Science.)

Marine recreational fishing is a popular outdoor activity. However, knowledge about the magnitude of recreational catches relative to commercial catches in coastal fisheries is generally sparse. Coastal Atlantic cod (Gadus morhua) is a target species for recreational fishers in the North Atlantic. In Norway, recreational fishers are allowed to use a variety of traps and nets as well as long-line and rod and line when fishing for cod. From 2005 to 2013, 9729 cod (mean size: 40 cm, range: 15-93 cm) were tagged and released in coastal Skagerrak, southeast Norway. Both high-reward (NOK 500) and low-reward tags (NOK 50) were used in this study. Because some harvested fish (even those posting high-reward tags) may go unreported by fishers, reporting rates were estimated from mark-recovery models that incorporate detection parameters in their structure, in addition to survival and mortality estimates. During 2005 to 2013, a total of 1707 tagged cod were recovered and reported by fishers. We estimate the overall annual survival to be 33% (SE 1.5). Recreational rod and line fishing were responsible for 33.7% (SE 2.4) of total mortality, followed by commercial fisheries (15.1% SE 0.8) and recreational fixed gear (6.8% SE 0.4). Natural mortality was 44.4% (SE 2.5) of total mortality. Our findings suggest that recreational fishing-rod and line fishing in particular-is responsible for a substantial part of fishing mortality exerted on coastal cod in southern Norway.

Several studies have documented fish populations changing in response to long-term warming. Over the past decade, sea surface temperatures in the Gulf of Maine increased faster than 99% of the global ocean. The warming, which was related to a northward shift in the Gulf Stream and to changes in the Atlantic Multidecadal Oscillation and Pacific Decadal Oscillation, led to reduced recruitment and increased mortality in the region's Atlantic cod (Gadus morhua) stock. Failure to recognize the impact of warming on cod contributed to overfishing. Recovery of this fishery depends on sound management, but the size of the stock depends on future temperature conditions. The experience in the Gulf of Maine highlights the need to incorporate environmental factors into resource management., (Copyright © 2015, American Association for the Advancement of Science.)

Gadus morhua (Atlantic cod) stocks in the Barents Sea are currently at levels not seen since the 1950s. Causes for the population increase last century, and understanding of whether such large numbers will be maintained in the future, are unclear. To explore this, we digitised and interrogated historical cod catch and diet datasets from the Barents Sea. Seventeen years of catch data and 12 years of prey data spanning 1930-1959 cover unexplored spatial and temporal ranges, and importantly capture the end of a previous warm period, when temperatures were similar to those currently being experienced. This study aimed to evaluate cod catch per unit effort and prey frequency in relation to spatial, temporal and environmental variables. There was substantial spatio-temporal heterogeneity in catches through the time series. The highest catches were generally in the 1930s and 1940s, although at some localities more cod were recorded late in the 1950s. Generalized Additive Models showed that environmental, spatial and temporal variables are all valuable descriptors of cod catches, with the highest occurring from 15-45°E longitude and 73-77°N latitude, at bottom temperatures between 2 and 4°C and at depths between 150 and 250 m. Cod diets were highly variable during the study period, with frequent changes in the relative frequencies of different prey species, particularly Mallotus villosus (capelin). Environmental variables were particularly good at describing the importance of capelin and Clupea harengus (herring) in the diet. These new analyses support existing knowledge about how the ecology of the region is controlled by climatic variability. When viewed in combination with more recent data, these historical relationships will be valuable in forecasting the future of Barents Sea fisheries, and in understanding how environments and ecosystems may respond.

1. Exploitation of living marine resources has resulted in major changes to populations of targeted species and functional groups of large-bodied species in the ocean. However, the effects of overfishing and collapse of large top predators on the broad-scale biodiversity of oceanic ecosystems remain largely unexplored. 2. Populations of the Atlantic cod (Gadus morhua) were overfished and several collapsed in the early 1990s across Atlantic Canada, providing a unique opportunity to study potential ecosystem-level effects of the reduction of a dominant predator on fish biodiversity, and to identify how such effects might interact with other environmental factors, such as changes in climate, over time. 3. We combined causal modelling with model selection and multimodel inference to analyse 41 years of fishery-independent survey data (1970-2010) and quantify ecosystem-level effects of overfishing and climate variation on the biodiversity of fishes across a broad area (172 000 km(2) ) of the Scotian Shelf. 4. We found that alpha and beta diversity increased with decreases in cod occurrence; fish communities were less homogeneous and more variable in systems where cod no longer dominated. These effects were most pronounced in the colder north-eastern parts of the Scotian Shelf. 5. Our results provide strong evidence that intensive harvesting (and collapse) of marine apex predators can have large impacts on biodiversity, with far-reaching consequences for ecological stability across an entire ecosystem., (© 2015 The Authors. Journal of Animal Ecology published by John Wiley & Sons Ltd on behalf of British Ecological Society.)

Complex natural systems with eroded resilience, such as populations, ecosystems and socio-ecological systems, respond to small perturbations with abrupt, discontinuous state shifts, or critical transitions. Theory of critical transitions suggests that such systems exhibit fold bifurcations featuring folded response curves, tipping points and alternate attractors. However, there is little empirical evidence of fold bifurcations occurring in actual complex natural systems impacted by multiple stressors. Moreover, resilience of complex systems to change currently lacks clear operational measures with generic application. Here, we provide empirical evidence for the occurrence of a fold bifurcation in an exploited fish population and introduce a generic measure of ecological resilience based on the observed fold bifurcation attributes. We analyse the multivariate development of Barents Sea cod (Gadus morhua), which is currently the world's largest cod stock, over six decades (1949-2009), and identify a population state shift in 1981. By plotting a multivariate population index against a multivariate stressor index, the shift mechanism was revealed suggesting that the observed population shift was a nonlinear response to the combined effects of overfishing and climate change. Annual resilience values were estimated based on the position of each year in relation to the fitted attractors and assumed tipping points of the fold bifurcation. By interpolating the annual resilience values, a folded stability landscape was fit, which was shaped as predicted by theory. The resilience assessment suggested that the population may be close to another tipping point. This study illustrates how a multivariate analysis, supported by theory of critical transitions and accompanied by a quantitative resilience assessment, can clarify shift mechanisms in data-rich complex natural systems., (© 2014 John Wiley & Sons Ltd.)

Harvesting may be a potent driver of demographic change and contemporary evolution, which both may have great impacts on animal populations. Research has focused on changes in phenotypic traits that are easily quantifiable and for which time series exist, such as size, age, sex, or gonad size, whereas potential changes in behavioural traits have been under-studied. Here, we analyse potential drivers of long-term changes in a behavioural trait for the Northeast Arctic stock of Atlantic cod Gadus morhua, namely choice of spawning location. For 104 years (1866-1969), commercial catches were recorded annually and reported by county along the Norwegian coast. During this time period, spawning ground distribution has fluctuated with a trend towards more northerly spawning. Spawning location is analysed against a suite of explanatory factors including climate, fishing pressure, density dependence, and demography. We find that demography (age or age at maturation) had the highest explanatory power for variation in spawning location, while climate had a limited effect below statistical significance. As to potential mechanisms, some effects of climate may act through demography, and explanatory variables for demography may also have absorbed direct evolutionary change in migration distance for which proxies were unavailable. Despite these caveats, we argue that fishing mortality, either through demographic or evolutionary change, has served as an effective driver for changing spawning locations in cod, and that additional explanatory factors related to climate add no significant information., (© 2014 The Authors. Global Change Biology Published by John Wiley & Sons Ltd.)

To estimate the impact of accidental oil-spills on cod fisheries a model framework is developed in which a Dynamic Energy Budget (DEB) model is applied to assess mortality caused by petroleum substances in early life stages. In this paper we report on a literature search and DEB analyses, aiming for cod specific DEB-parameters. Furthermore, we explored the relevance of Fathead minnow DEB-parameters as surrogate by comparing LC50 values calculated from DEB-parameters with literature. Cod specific DEB-parameters could not be estimated based on available literature. LC50 values calculated from Fathead minnow DEB-parameters were higher than literature LC50 for early life stages of fish. Applying an extrapolation factor of 50 to the DEB-parameters resulted in LC50 values that were below literature irrespective of life stage. Therefore, we propose to use the last as an estimate for early life stages in cod and recommend relevant experiments with individual petroleum substances on cod., (Copyright © 2014 Elsevier Ltd. All rights reserved.)

The northern Gulf of St. Lawrence (NGSL) stock of Atlantic cod (Gadus morhua), historically the second largest cod population in the Western Atlantic, has known a severe collapse during the early 1990 s and is currently considered as endangered by the Committee on the Status of Endangered Wildlife in Canada. As for many fish populations over the world which are currently being heavily exploited or overfished, urgent management actions in the form of recovery plans are needed for restoring this stock to sustainable levels. Stochastic projections based on a statistical population model incorporating predation were conducted over a period of 30 years (2010-2040) to assess the expected outcomes of alternative fishing strategies on the stock recovery under different scenarios of harp seal (Pagophilus groenlandicus) abundance and environmental conditions. This sensitivity study shows that water temperature is key in the rebuilding of the NGSL cod stock. Model projections suggest that maintaining the current management practice under cooler water temperatures is likely to maintain the species in an endangered status. Under current or warmer conditions in the Gulf of St. Lawrence, partial recovery might only be achieved by significant reductions in both fishing and predation pressure. In the medium-term, a management strategy that reduces catch could be favoured over a complete moratorium so as to minimize socio-economic impacts on the industry.

Natural resource management requires approaches to understand and handle sources of uncertainty in future responses of complex systems to human activities. Here we present one such approach, the "biological ensemble modeling approach," using the Eastern Baltic cod (Gadus morhua callarias) as an example. The core of the approach is to expose an ensemble of models with different ecological assumptions to climate forcing, using multiple realizations of each climate scenario. We simulated the long-term response of cod to future fishing and climate change in seven ecological models ranging from single-species to food web models. These models were analyzed using the "biological ensemble modeling approach" by which we (1) identified a key ecological mechanism explaining the differences in simulated cod responses between models, (2) disentangled the uncertainty caused by differences in ecological model assumptions from the statistical uncertainty of future climate, and (3) identified results common for the whole model ensemble. Species interactions greatly influenced the simulated response of cod to fishing and climate, as well as the degree to which the statistical uncertainty of climate trajectories carried through to uncertainty of cod responses. Models ignoring the feedback from prey on cod showed large interannual fluctuations in cod dynamics and were more sensitive to the underlying uncertainty of climate forcing than models accounting for such stabilizing predator-prey feedbacks. Yet in all models, intense fishing prevented recovery, and climate change further decreased the cod population. Our study demonstrates how the biological ensemble modeling approach makes it possible to evaluate the relative importance of different sources of uncertainty in future species responses, as well as to seek scientific conclusions and sustainable management solutions robust to uncertainty of food web processes in the face of climate change.

In March 2012, fishermen operating in a fjord in Northern Norway reported catching Atlantic cod, a native fish forming an economically important marine fishery in this region, with unusual prey in their stomachs. It was speculated that these could be Atlantic salmon, which is not typical prey for cod at this time of the year in the coastal zone. These observations were therefore reported to the Norwegian Directorate of Fisheries as a suspected interaction between a local fish farm and this commercial fishery. Statistical analyses of genetic data from 17 microsatellite markers genotyped on 36 partially-degraded prey, samples of salmon from a local fish farm, and samples from the nearest wild population permitted the following conclusions: 1. The prey were Atlantic salmon, 2. These salmon did not originate from the local wild population, and 3. The local farm was the most probable source of these prey. Additional tests demonstrated that 21 of the 36 prey were infected with piscine reovirus. While the potential link between piscine reovirus and the disease heart and skeletal muscle inflammation is still under scientific debate, this disease had caused mortality of large numbers of salmon in the farm in the month prior to the fishermen's observations. These analyses provide new insights into interactions between domesticated and wild fish.

Common haematological [haematocrit (Hct)], primary (serum cortisol) and secondary (serum glucose and plasma lactate) analytes were utilized to compare blood biochemical status of Gadus morhua captured rapidly by jig with that of G. morhua captured by commercial demersal longline. In general, the physiological status of G. morhua, despite blind hook times, was significantly more disrupted (pronounced haemo-concentration and significantly elevated concentrations of cortisol, glucose and lactate) following longline capture relative to capture by jig, while no differences were detected among longline-caught fish as a function of dehooking method (or concomitant extent of overt physical trauma). Blood profiles from the more stressed G. morhua, a possible function of more extended longline hook times, were similar to the most stressed values reported for this species. The results also demonstrate that, although acute blood biochemical status is an effective gauge of relative stress, it does not reflect physical injury status, which has been shown to exert a strong influence on delayed mortality in previous studies in this species. Thus, acute blood chemical status alone may not be the most complete predictor of mortality. Future studies should evaluate physiological repercussions from capture-handling against physical trauma during more extended post-release periods for this species., (© 2012 The Authors. Journal of Fish Biology © 2012 The Fisheries Society of the British Isles.)

Proceeding from long-term data on the numbers of eggs of cod in ichthyoplankton, the total annual production of cod eggs at four main spawning grounds of the Baltic Sea was calculated. It was shown that the long-term fluctuations of cod egg production were positively related to the dynamics of the volume of waters coming to the Baltic Sea in years of the North Sea advections. It is suggested that this dependence was determined by a set of adaptations providing the extension of cod reproduction upon the improvement of the environment.

The behavioural responses of wild (predator-experienced) and hatchery-reared (predator-naive) cod Gadus morhua to standardized mechano-acoustic (MA) stimuli were compared in the laboratory. Wild fish responded mainly with freezing and fast-start escapes away from the stimulus, whereas hatchery-reared fish often ignored or approached the stimulus. Wild fish also had stronger responses, turning faster during escapes and reducing activity immediately after the stimulus. Both fish types were less active on a 'risky' bare substratum after the stimulus. The antipredator responses of wild fish were consistent to repeated stimuli, whereas hatchery-reared fish that had generally only encountered harmless stimuli showed more variable responses with lower repeatability. This suggests that experience plays a role in shaping the behavioural response of fishes to MA stimuli., (© 2011 The Authors. Journal of Fish Biology © 2011 The Fisheries Society of the British Isles.)

The effect of two disinfectants on eggs and larvae of Baltic cod, Gadus morhua, was investigated. The eggs were disinfected for 10 min using various concentrations of either glutaraldehyde (100, 200, 400, 600 and 800 mg L(-1)) or iodophor (10, 50, 100 and 150 mg L(-1)), 1-4-days post-fertilization. Bactericidal effect of disinfection, survival to hatching, hatching success and larval abnormalities were assessed. Larval survival was recorded at 5-, 10- and 15-days post-hatch (dph). Although Baltic cod eggs have an unusually thin chorion, they could tolerate surface disinfection. A reduction in bacterial growth was observed with increased concentrations of disinfectant (3.0 x 10(7)-1.6 x 10(1) CFU mL(-1)). Abnormalities in newly hatched larvae were not related to disinfection. Survival of the yolk sac larvae was significantly better for eggs treated with 400 mg L(-1) glutaraldehyde for 10 min at 10 and 15 dph. Effective disinfection was also recorded using 100 mg L(-1) Actomar K30. Egg batch effect rather than initial bacterial concentration, disinfectant type or incubation method determined the survival of the eggs to hatching and survival of larvae. Because of the carcinogenic effect of glutaraldehyde, iodophor is recommended for routine disinfection of cod eggs.

Prior to the 1920s, the northeast Arctic (NA) cod were caught at spawning grounds ranging from the southernmost to the northernmost parts of the Norwegian coast, but have for the last 50 yr mainly been caught around the Lofoten archipelago and northwards. The NA cod have their feeding and nursery grounds in the Barents Sea, and migrate south towards the Norwegian coast in the winter to spawn. This study uses commercial fisheries' data from landing ports along the entire Norwegian coast during the period 1866-1969 as evidence of long-term truncation and northerly shift of spawning grounds. Nearly all spawning grounds south of Lofoten have been abandoned, while an increasing proportion of the spawning stock only uses the northernmost areas of the Norwegian coast, Troms and Finnmark. The truncation can hardly be attributed to long-term climatic variations, but may result from an intensive size-selective trawl fishery in the Barents Sea causing a sudden increase in fishing mortality, probably altering the size structure and migratory capacity of the stock.

During the course of the last century, populations of Atlantic cod Gadus morhua L. have undergone dramatic declines in abundance across their biogeographic range, leading to debate about the relative roles of climatic warming and overfishing in driving these changes. In this chapter, we describe the geographic distributions of this important predator of North Atlantic ecosystems and document extensive evidence for limitations of spatial movement and local adaptation from population genetic markers and electronic tagging. Taken together, this evidence demonstrates that knowledge of spatial population ecology is critical for evaluating the effects of climate change and commercial harvesting. To explore the possible effects of climate change on cod, we first describe thermal influences on individual physiology, growth, activity and maturation. We then evaluate evidence that temperature has influenced population-level processes including direct effects on recruitment through enhanced growth and activity, and indirect effects through changes to larval food resources. Although thermal regimes clearly define the biogeographic range of the species, and strongly influence many aspects of cod biology, the evidence that population declines across the North Atlantic are strongly linked to fishing activity is now overwhelming. Although there is considerable concern about low spawning stock biomasses, high levels of fishing activity continues in many areas. Even with reduced fishing effort, the potential for recovery from low abundance may be compromised by unfavourable climate and Allee effects. Current stock assessment and management approaches are reviewed, alongside newly advocated methods for monitoring stock status and recovery. However, it remains uncertain whether the rebuilding of cod to historic population sizes and demographic structures will be possible in a warmer North Atlantic.

Individuals migrate to exploit heterogeneities between spatially separated environments to modulate growth, survival, or reproduction. We devised a bioenergetics model to investigate the evolution of migration distance and its dependence on individual states. Atlantic cod Gadus morhua ranges from sedentary populations to stocks that migrate several thousand kilometers annually. We focused on the Northeast Arctic cod stock, which migrates south to spawn. A linear relationship between migration distance and the expected survival of offspring was assumed, here understood as the prospects for future survival and development that a fertilized egg faces at a particular spawning location. Reasons for why it may increase southward include warmer water that increases development rates, and thereby survival, along the pelagic drift trajectory. In the model, ingested energy can either be allocated to growth or stored for migration and reproduction. When migrating, individuals forgo foraging opportunities and expend energy. Optimal energy allocation and migration strategies were found using state-dependent optimization, with body length, age, condition, and current food availability as individual states. For both a historical and contemporary fishing regime we modeled two behaviors: (1) homing cod returning to the same spawning site each year and (2) roaming cod with no such constraints. The model predicted distinct regions of locally high spawning stock biomass. Large individuals in good condition migrated farthest, and these also tended to mature later in life. The roaming cod spread farther south as they grew larger and older. Homing cod did not have this freedom, and spawning was generally concentrated along a narrower stretch of the coastline. Under contemporary fishing, individuals matured earlier at a smaller size, had shorter migrations, spawned over a contracted geographical range, and tended to be in poorer condition. The effects were most pronounced for the homing behavior.

Despite recent evidence for sub-stock structuring, North Sea cod are assessed as a single unit. As a consequence, knowledge of sub-stock trends is poor. In particular, there are no recent evaluations of which spawning grounds are active. Here we report results from the first ichthyoplankton survey to cover the whole North Sea. Also, this survey, conducted in 2004, was the first to make extensive use of DNA-based molecular methods to unambiguously identify early developmental stage cod eggs. We compare the findings from the plankton survey with estimated egg production inferred from the distribution of mature cod in contemporaneous trawl surveys. Results from both approaches were in general agreement and showed hot spots of egg production around the southern and eastern edges of the Dogger Bank, in the German Bight, the Moray Firth and to the east of the Shetlands. These areas broadly coincide with known spawning locations from the period 1940 to 1970. We were, however, unable to directly detect significant numbers of cod eggs at the historic spawning ground off Flamborough (northeast coast of England). The results demonstrate that most of the major spawning grounds of cod in the North Sea are still active but that some localized populations may have been reduced to the point where it is now difficult to detect the presence of eggs in the plankton.

World-wide many cetaceans drown incidentally in fishing nets. To reduce the unwanted bycatch in gillnets, pingers (acoustic alarms) have been developed that are attached to the nets. In the European Union, pingers will be made compulsory in some areas in 2005 and in others in 2007. However, pingers may effect non-target marine fauna such as fish. Therefore in this study, the effects of seven commercially-available pingers on the behaviour of five North Sea fish species in a large tank were quantified. The species tested were: sea bass (Dicentrarchus labrax), pout (Trisopterus luscus), thicklip mullet (Chelon labrosus), herring (Clupea harengus), and cod (Gadus morhua). The fish were housed as single-species schools of 9-13 individuals in a tank. The behaviour of fish in quiet periods was compared with their behaviour during periods with active pingers. The results varied both between pingers and between fish species. Sea bass decreased their speed in response to one pinger and swam closer to the surface in response to another. Thicklip mullet swam closer to the bottom in response to two pingers and increased their swimming speed in response to one pinger. Herring swam faster in response to one pinger, and pout and cod (close relatives) showed no behavioural responses to any of the pingers. Of the seven pingers tested, four elicited responses in at least one fish species, and three elicited no responses. Whether similar responses would be elicited in these fish species in the wild, and if so, whether such responses would influence the catch rate of fisheries, cannot be derived from the results of this study. However, the results indicate the need for field studies with pingers and fish. Based on the small number of fish species tested, the present study suggests that the higher the frequency of a pinger, the less likely it is to affect the behaviour of marine fish.

The majority of survival analyses focus on temporal scales. Consequently, there is a limited understanding of how species survival varies over space and, ultimately, how spatial variability in the environment affects the temporal dynamics of species abundance. Using data from the Barents Sea, we study the spatiotemporal variability of the juvenile Atlantic cod (Gadus morhua) survival. We develop an index of spatial survival based on changes of juvenile cod distribution through their first winter of life (from age-0 to age-1) and study its variability in relation to biotic and abiotic factors. Over the 25 years analyzed (1980-2004), we found that, once the effect of passive drift due to dominant currents is accounted for, the area where age-0 cod survival was lowest coincided with the area of highest abundance of older cod. Within this critical region, the survival of age-0 cod was negatively affected by its own abundance, by that of older cod, and by bottom depth. Furthermore, during cold years, age-0 cod survival increased in the eastern and coldest portion of the examined area, which was typically avoided by older conspecifics. Based on these results we propose that within the examined area top-down mechanisms and predation-driven density dependence can strongly affect the spatial pattern of age-0 cod survival. Climate-related variables can also influence the spatial survival of age-0 cod by affecting their distribution and that of their predators. Results from these and similar studies, focusing on the spatial variability of survival rates, can be used to characterize species habitat quality of marine renewable resources.