Jon Egil Skjæraasen

Synergies between climate and management for Atlantic cod fisheries at high latitudes

Significance Currently many exploited fish populations, including many of the Atlantic cod stocks, are at historically low levels with widespread concern about whether contemporary management is capable of facilitating population recovery. In contrast, the spawning stock biomass of Barents Sea cod is now at an historic high. Here we demonstrate that successful management actions interacting synergistically with prevailing climate caused this increase. Warming of water masses in the Barents Sea over the last decade positively reinforced management actions. A unique and possibly generic mechanism of climate affecting marine animals at high latitudes, especially when at the polar extreme of their distribution, is identified: adjustment of the suitable feeding area. This adjustment is linked closely to community dynamics and increased stock productivity. The widespread depletion of commercially exploited marine living resources is often seen as a general failure of management and results in criticism of contemporary management procedures. When populations show dramatic and positive changes in population size, this invariably leads to questions about whether favorable climatic conditions or good management (or both) were responsible. The Barents Sea cod (Gadus morhua) stock has recently increased markedly and the spawning stock biomass is now at an unprecedented high. We identify the crucial social and environmental factors that made this unique growth possible. The relationship between vital rates of Barents Sea cod stock productivity (recruitment, growth, and mortality) and environment is investigated, followed by simulations of population size under different management scenarios. We show that the recent sustained reduction in fishing mortality, facilitated by the implementation of a “harvest control rule,” was essential to the increase in population size. Simulations show that a drastic reduction in fishing mortality has resulted in a doubling of the total population biomass compared with that expected under the former management regime. However, management alone was not solely responsible. We document that prevailing climate, operating through several mechanistic links, positively reinforced management actions. Heightened temperature resulted in an increase in the extent of the suitable feeding area for Barents Sea cod, likely offering a release from density-dependent effects (for example, food competition and cannibalism) through prolonged overlap with prey and improved adult stock productivity. Management and climate may thus interact to give a positive outlook for exploited high-latitude marine resources.

Frequent skipped spawning in the world’s largest cod population

Life-history theory suggests that animals may skip reproductive events after initial maturation to maximize lifetime fitness. In iteroparous teleosts, verifying past spawning history is particularly difficult; the degree of skipped spawning at the population level therefore remains unknown. We unequivocally show frequent skipped spawning in Northeast Arctic cod (NEAC) in a massive field and laboratory effort from 2006 to 2008. This was verified by postovulatory follicles in temporarily arrested ovaries close to the putative spawning period. At the population level, “skippers” were estimated to be approximately equally abundant as spawning females in 2008, constituting ∼24% of the females 60–100 cm. These females never truly started vitellogenesis and principally remained on the feeding grounds when spawners migrated southward, avoiding any migration costs. The proximate cause of skipping seems to be insufficient energy to initiate oocyte development, indicating that skipped spawning may partly be a density-dependent response important in population regulation. Our data also indicate more skipping among smaller females and potential tradeoffs between current and future reproductive effort. We propose that skipped spawning is an integral life-history component for NEAC, likely varying annually, and it could therefore be an underlying factor causing some of the currently unexplained large NEAC recruitment variation. The same may hold for other teleosts.

Domestication causes rapid changes in heart and brain morphology in Atlantic cod (Gadus morhua)

Brain and heart development is very plastic in teleost fishes, and receptive to changes in social and environmental conditions. Domestication in salmonids has been reported to result in pronounced changes in both heart and brain morphology. In particular, a high prevalence of heart deformities has been reported in farmed salmonids, which has been linked to increased stress responsiveness that can impair survival of both farmed and escaped fish. Here we report for the first time that significant changes in heart and brain morphology occur following domestication of Atlantic cod (Gadus morhua), an emerging aquaculture species. Juvenile farmed cod developed significantly larger hearts and smaller brains, by weight, compared to their wild conspecifics. These differences occurred within the first captive generation, suggesting that they were driven largely by the strong contrast in environmental and social conditions experienced within their respective rearing environments. Changes in brain and heart morphology, as a consequence of domestication could affect the well-being and survival of Atlantic cod raised under intensive aquaculture conditions.

Reproductive interactions between fugitive farmed and wild Atlantic cod (Gadus morhua) in the field

Atlantic cod (Gadus morhua) are being increasingly farmed in net pens adjacent to coastal populations that are currently at historic lows. One concern is that farmed escapees enter local spawning shoals and mate with wild cod. We tested for the potential of escaped farmed cod to interact and hybridize with wild fish by examining the spatial dynamics of, and associations between, fish tagged with ultrasonic transmitters. Based on these data, we also investigated the basic mating system of cod in the field. The spawning ground was best described as a lekking arena. Wild males aggregated near the seafloor and associations between individuals were frequent. Wild females had a pelagic and dispersed distribution and rarely associated with each other. Associations between individual wild males and females were also infrequent. Farmed males rarely associated with wild fish and had core usage areas above the wild males, suggesting that they were not admitted into the spawning arena. Farmed females were over the sp...

Triploidy alters brain morphology in pre-smolt Atlantic salmon Salmo salar: possible implications for behaviour.

Total brain mass and the volumes of five specific brain regions in diploid and triploid Atlantic salmon Salmo salar pre-smolts were measured using digital images. There were no significant differences (P > 0·05) in total brain mass when corrected for fork length, or the volumes of the optic tecta or hypothalamus when corrected for brain mass, between diploids and triploids. There was a significant effect (P < 0·01) of ploidy on the volume of the olfactory bulb, with it being 9·0% larger in diploids compared with triploids. The cerebellum and telencephalon, however, were significantly larger, 17 and 8% respectively, in triploids compared with diploids. Sex had no significant effect (P > 0·05) on total brain mass or the volumes of any measured brain region. As the olfactory bulbs, cerebellum and telencephalon are implicated in a number of functions, including foraging ability, aggression and spatial cognition, these results may explain some of the behavioural differences previously reported between diploids and triploids.

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.

Secondary sexual characteristics in codfishes (Gadidae) in relation to sound production, habitat use and social behaviour

Abstract Little is known about the reproductive biology of the codfishes (Gadidae). Lacking direct observations, the study of secondary sexual characteristics can provide cues to their reproductive biology and behaviour. We reviewed here published accounts on sexual dimorphisms in 25 gadids in light of their general lifestyle, i.e. pelagic or demersal, and social behaviour. In addition, complementary data on fin lengths and drumming muscle size in haddock (Melanogrammus aeglefinus), saithe (Pollachius virens), blue whiting (Micromesistius poutassou) and cod (Gadus morhua) are presented. Capacity for sound production occurred in almost half of the studied species, but was most prevalent in demersal species, where it is probably used in resource contests and to attract mates. For semi-pelagic gadids, we postulate that sound production may be linked to the formation of male-biased spawning shoals and the attraction of females towards such shoals; we identify candidate species to further test this hypothesis. Although rarely studied, sexual fin dimorphisms occur in several gadids. Cod, saithe and blue whiting males have longer pelvic fins than females, whereas no such dimorphism was observed in haddock. In cod and haddock, males use pelvic fins during courtship of females and agonistic encounters with other males. Pelvic fins probably also have a similar function in other gadids. The hitherto available information on sexually dimorphic traits and/or courtship behaviour in seven gadid species suggests that complex mating systems and non-random mate choice occurs frequently in this important group of exploited fishes.

Vaccination and triploidy increase relative heart weight in farmed Atlantic salmon, Salmo salar L.

Heart morphology is particularly plastic in teleosts and differs between farmed and wild Atlantic salmon. However, little is known about how different culture practices and sex affect heart morphology. This study investigated how vaccination, triploidy and sex affected heart size and heart morphology (ventricle shape, angle of the bulbus arteriosus) in farmed Atlantic salmon for 18 months following vaccination (from c. 50-3000 g body weight). In addition, hearts were examined histologically after 7 months in sea water. All fish sampled were sexually immature. Vaccinated fish had significantly heavier hearts relative to body weight and a more triangular ventricle than unvaccinated fish, suggesting a greater cardiac workload. Irrespective of time, triploids had significantly heavier hearts relative to body weight, a more acute angle of the bulbus arteriosus and less fat deposition in the epicardium than diploids. The ventricle was also more triangular in triploids than diploids at seawater transfer. Sex had transient effects on the angle of the bulbus arteriosus, but no effect on relative heart weight or ventricle shape. From a morphological perspective, the results indicate that vaccination and triploidy increase cardiac workload in farmed Atlantic salmon.

Shifting reproductive success in a shoal of Atlantic Cod, Gadus morhua L.

Individual and temporal variability in the spawning behaviour of Atlantic Cod (Gadus morhua L.) was examined by studying wild and reared individuals from the same putative population in a common spawning arena. The number of eggs fertilized per male was best explained by the number of aggressive interactions initiated by each male. A strong skew in male reproductive success and a temporal shift in male fertilization success were also observed. The latter co-occurred with an apparent similar temporal shift in the reproductive behaviour of the individual males. We hypothesize that energetic costs associated with reproduction, potentially mediated by sperm depletion in dominant individuals, may be responsible for temporal shifts in mating behaviour and reproductive success of male Atlantic Cod. We suggest that this hypothesis merits further study.

Multidimensionality of behavioural phenotypes in Atlantic cod, Gadus morhua

Much of the inter-individual variation observed in animal behaviour is now attributed to the existence of behavioural phenotypes or animal personalities. Such phenotypes may be fundamental to fisheries and aquaculture, yet there have been few detailed studies of this phenomenon in exploited marine animals. We investigated the behavioural and neuroendocrine responses of Atlantic cod (Gadus morhua L.), to situations reflecting critical ecological challenges: predator attacks and territorial challenges. Both hatchery-reared and wild fish were tested and behavioural profiles were compared with baseline conditions. We then used an objective, multivariate approach, rather than assigning individuals along one-dimensional behavioural axes, to examine whether distinct behavioural phenotypes were present. Our results indicate that two distinct behavioural phenotypes were evident in fish from each background. In hatchery-reared fish, phenotypes displayed divergent locomotor activity, sheltering, brain monoamine concentrations and responses to competitive challenges. In wild fish, phenotypes were distinguished primarily by locomotor activity, sheltering and responsiveness to predator stimuli. Hatcheries presumably represent a more stressful social environment, and social behaviour and neuroendocrine responses were important in discerning behavioural phenotypes in hatchery fish, whereas antipredator responses were important in discerning phenotypes in wild fish that have previously encountered predators. In both fish types, behavioural and physiological traits that classified individuals into phenotypes were not the same as those that were correlated across situations. These results highlight the multidimensionality of animal personalities, and that the processes that regulate one suite of behavioural traits may be very different to the processes that regulate other behaviours.