Stock structure of Icelandic cod Gadus morhua L. based on otolith chemistry

Otolith chemistry was used to study the stock structure of Icelandic cod Gadus morhua . Otoliths were collected from spawning cod at 12 and 17 different spawning locations around Iceland during the spawning season in 2002 and 2003, respectively. The otolith elemental composition of cod spawning north and south of Iceland differed substantially from each other, as did spawning cod below and above 125 m depth at the main spawning ground in the south of Iceland. Otolith chemistry differed more between locations than among years within a location, indicating temporal stability between the 2 years. The Icelandic cod stock is managed as a single stock. These results suggest, however, that optimal fisheries management may require different management units than are currently present.     

Feeding behaviour of cod (Gadus morhua) in relation to spawning

The food consumption and egg production of 26 adult (13 female and 13 male) Atlantic cod ( Gadus morhua ) were monitored during prespawning, spawning and postspawning periods. Females spawned from late January to mid-April. Feeding activity occurred from December to early January and ceased for females, on average, 36 days (15–54 days) before the onset of spawning. The duration of spawning by females was, on average, 42 days (10–61 days) and feeding was suppressed by both sexes during the first three-quarters of each female's spawning period. Mature females went, on average, 70 days or 19% of the year without eating. An abrupt increase in feeding activity, particularly by females, occurred during the last quarter of spawning or shortly after the release of the last egg batch (on average, feeding started again after 91% of a female's eggs had been released or 82% of egg batches). Females consumed greater quantities of food than males during both winter and postspawning feeding periods. During spawning, females lost, on average, 29% of their body weight and males 14%. Fecundity ranged from 0.75 to 3.97 million eggs per female. The volume of eggs produced by four individual females (range = 1285–5995 ml in four to 11 batches) ranged from 99 to 195% (mean 150%) of a female's postspawning body volume. Six immature cod fed throughout the experimental period and gained, on average, 8% of initial body weight. Laboratory results were supported by stomach fullness index values of Georges Bank cod exhibiting different maturity states.     

Ticket to spawn: Combining economic and genetic data to evaluate the effect of climate and demographic structure on spawning distribution in Atlantic cod

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.     

Recolonization history and large-scale dispersal in the open sea: the case study of the North Atlantic cod, Gadus morhua L.

Most studies of the genetic structure of Atlantic cod have focused on small geographical scales. In the present study, the genetic structure of cod sampled on spawning grounds in the North Atlantic was examined using eight microsatellite loci and the Pan I locus. A total of 954 cod was collected from nine different regions: the Baltic Sea, the North Sea, the Celtic Sea, the Irish Sea and Icelandic waters during spring 2002 and spring 2003, from Norwegian waters and the Faroe Islands (North and West spawning grounds) in spring 2003, and from Canadian waters in 1998. Temporal stability among spawning grounds was observed in Icelandic waters and the Celtic Sea, and no significant difference was observed between the samples from the Baltic Sea and between the samples from Faroese waters. F -statistics showed significant differences between most populations and a pattern of isolation-by-distance was described with microsatellite loci. The Pan I locus revealed the presence of two genetically distinguishable basins, the North-west Atlantic composed of the Icelandic and Canadian samples and the North-east Atlantic composed of all other samples. Permutation of allele sizes at each microsatellite locus among allelic states supported a mutational component to the genetic differentiation, indicating a historical origin of the observed variation. Estimation of the time of divergence was approximately 3000 generations, which places the origin of current genetic pattern of cod in the North Atlantic in the late Weichselian (Wisconsinian period), at last glacial maximum.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 315–329.     

Survival, activity and feeding ability of Baltic cod (Gadus morhua) yolk-sac larvae at different salinities

Due to unfavourable conditions (declines in salinity and water oxygen content) in the spawning areas, there has been a considerable decrease in the Baltic cod stock since the beginning of the 1980s, and consequently a decrease in catches. In order to examine the feasibility of introducing yolk-sac larvae in areas of low salinity to improve the stock, laboratory experiments were performed on the effects of salinity on the survival, level of activity and feeding ability of larvae. Yolk-sac larvae from spawning cod caught off northern Gotland, Sweden, were exposed to four different salinities: 10 and 15%○ (salinities of the main spawning areas); and 5 and 7%○ (salinities in the Bothnian Sea and the Baltic proper respectively).
The survival of yolk-sac larvae was high at all salinities, even though there was an indication of higher mortality at low salinities in less viable larval groups. No differences were found in swimming speed or feeding ability at the four salinities, but a significant difference in vertical distribution was recorded. There were significant differences in survival, vertical distribution and feeding ability among larval groups, which indicates that larval quality or viability is of greater importance for larval survival than salinity, in the range of 5–15%○.     

Mapping the spawning grounds of North Sea cod (Gadus morhua) by direct and indirect means

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.     

First evidence of spawning of eastern Baltic cod (Gadus morhua callarias) in the Belt Sea,the main spawning area of western Baltic cod (Gadus morhua L.)

Two cod stocks (western Baltic cod, WBC, and eastern Baltic cod, EBC) are managed in the Baltic Sea which is characterized by different main spawning areas and different main spawning periods. In this study we analyse the spatial and temporal occurrence of spawning individuals of both cod stocks in the main spawning grounds of the Baltic Sea based on eight microsatellite loci. Our results suggest that EBC (Gadus morhua callarias) has formed currently temporally stable, substantially homogeneous population not only in the Bornholm Sea (ICES SD: 25) but also in the Arkona Sea (ICES SD: 24). The presented analyses proved that EBC (G. m. callarias) can temporarily also spawn in the Belt Sea.     

Comparative analysis of genetic variability of white sea cod (Gadus morhua marisalbi) at allozyme and microsatellite markers

Variability of cod spawning and feeding schools from Kandalaksha Bay of the White Sea, was examined at six allozyme and eight microsatellite loci. The degree of genetic differentiation at allozyme loci constituted θ = 0.36% [95% bootstrap interval 0.0458; 0.6743]. The differentiation estimates obtained using microsatellite markers were higher, θ = 1.33% [0.057; 3.11]. It was demonstrated that the level of genetic diversity in the White Sea cod was lower than that established for the Atlantic cod from Barents Sea using the same set of allozyme and microsatellite markers. The genetic data obtained support the opinion that the White Sea cod is a reproductively independent group formed as a result of the Holocene dispersal of Atlantic cod.     

Evidence of Segregated Spawning in a Single Marine Fish Stock: Sympatric Divergence of Ecotypes in Icelandic Cod?

There is increasing recognition of intraspecific diversity and population structure within marine fish species, yet there is little direct evidence of the isolating mechanisms that maintain it or documentation of its ecological extent. We analyzed depth and temperature histories collected by electronic data storage tags retrieved from 104 Atlantic cod at liberty ≥1 year to evaluate a possible isolating mechanisms maintaining population structure within the Icelandic cod stock. This stock consists of two distinct behavioral types, resident coastal cod and migratory frontal cod, each occurring within two geographically distinct populations. Despite being captured together on the same spawning grounds, we show the behavioral types seem reproductively isolated by fine-scale differences in spawning habitat selection, primarily depth. Additionally, the different groups occupied distinct seasonal thermal and bathymetric niches that generally demonstrated low levels of overlap throughout the year. Our results indicate that isolating mechanisms, such as differential habitat selection during spawning, might contribute to maintaining diversity and fine-scale population structure in broadcast-spawning marine fishes.     

Diet of Murray cod (<Emphasis Type="Italic">Maccullochella peelii peelii</Emphasis>) (Mitchell) larvae in an Australian lowland river in low flow and high flow years

Researchers have hypothesised that influxes of pelagic zooplankton to river channels after floods and high flows are necessary for strong recruitment of some native fish species, including Murray cod (Maccullochella peelii peelii) (Mitchell), in the Murray–Darling river system, Australia. This study investigated the composition of the diet and gut fullness of drifting Murray cod larvae weekly during two spawning seasons with contrasting flows, to determine if pelagic zooplankton comprised a greater proportion of the gut contents and guts were fuller in a high flow (2000) than in a low flow (2001) year. Gut fullness and yolk levels of 267 larvae were ranked, and prey identified to family level. Approximately 40 and 70% of individuals had been feeding in 2000 and 2001, respectively. Gut fullness increased with declining yolk reserves. Larvae in both the years had an almost exclusively benthic diet, irrespective of the flow conditions at the time. Substantial inundation of dry ground in 2000, albeit restricted to in-channel benches, anastomosing channels and oxbow lakes, did not lead to an influx of pelagic, floodplain-derived zooplankton subsequently exploited by Murray cod larvae. These results have the implications for the management of regulated temperate lowland rivers: high flows cannot automatically be assumed to be beneficial for the fish larvae of all species and their food resources, and caution should be exercised with the timing of flow releases.     

Time of start of spawning in Atlantic cod (Gadus morhua) females in relation to vitellogenic oocyte diameter, temperature, fish length and condition

Spawning time of female Atlantic cod ( Gadus morhua L.) can be predicted to within 3 days using vitellogenic oocyte diameters measured over the last month before spawning. Simulations show that a l°C drop in temperature during vitellogenesis delays spawning by about 8–10 days. Similar results were obtained with fish on low ration for 4–8 months before spawning. Moderately fed females held over two to four consecutive spawning seasons demonstrated a significant delay (⋍3–40 days) in the date of first spawning as the fish grew older and more fecund. Forecasting studies on spawning migrating Arcto-Norwegian cod gave no evidence of a significant effect of fish length on the time of spawning. Warmer water temperature during vitellogenesis for the larger fish appears to compensate for the negative effect of fecundity On spawning time.     

The trophic ecology of Atlantic cod: insights from tri-monthly, localized scales of sampling

The meso-scale trophic dynamics of cod Gadus morhua were examined based upon tri-monthly stomach sample collections from a nearshore, localized ( c. 800 km²) region off Cape Cod, Massachusetts, U.S.A. The major objective for this work was to relate any changes in cod diet and amount of food eaten to seasonal variations in prey availability, water temperature and spawning at a spatial scale between broad-scale and laboratory studies. Results suggested that the type and amount of food eaten by cod was generally consistent throughout a year and repeatable across years. Cod feeding was marked by two periods of increased feeding, corresponding to the arrival of small pelagic fishes in the area. This pelagic migration and subsequent increased feeding by cod occurred during important periods in the life history of cod ( e.g . spawning and overwintering). Similar annual patterns in food consumption and diet composition were remarkably consistent over the 2·5 years of the project, suggesting important feeding periods for cod that correspond to environmental and biological events. The diet of cod was composed primarily of several species of forage fishes [ e.g. herrings (predominantly Atlantic herring Clupea harengus ), sand lance Ammodytes sp. and Atlantic mackerel Scomber scombrus ], ophiuroids, Cancer sp. crabs and other small crustaceans. It was inferred that cod exhibited a maintenance diet on local forage fishes and benthic macroinvertebrates, augmenting their diet by seasonally gorge feeding upon migrating pelagic species.     

Historical DNA documents long‐distance natal homing in marine fish

The occurrence of natal homing in marine fish remains a fundamental question in fish ecology as its unequivocal demonstration requires tracking of individuals from fertilization to reproduction. Here, we provide evidence of long‐distance natal homing (>1000 km) over more than 60 years in Atlantic cod (Gadus morhua), through genetic analysis of archived samples from marked and recaptured individuals. Using a high differentiation single‐nucleotide polymorphism assay, we demonstrate that the vast majority of cod tagged in West Greenland and recaptured on Icelandic spawning grounds belonged to the Iceland offshore population, strongly supporting a hypothesis of homing. The high degree of natal fidelity observed provides the evolutionary settings for development of locally adapted populations in marine fish and emphasize the need to consider portfolio effects in marine fisheries management strategies.     

Low‐coverage whole‐genome sequencing reveals molecular markers for spawning season and sex identification in Gulf of Maine Atlantic cod (Gadus morhua,Linnaeus 1758)

Atlantic cod (Gadus morhua, Linnaeus 1758) in the western Gulf of Maine are managed as a single stock despite several lines of evidence supporting two spawning groups (spring and winter) that overlap spatially, while exhibiting seasonal spawning isolation. Low‐coverage whole‐genome sequencing was used to evaluate the genomic population structure of Atlantic cod spawning groups in the western Gulf of Maine and Georges Bank using 222 individuals collected over multiple years. Results indicated low total genomic differentiation, while also showing strong differentiation between spring and winter‐spawning groups at specific regions of the genome. Guided regularized random forest and ranked F _ST methods were used to select panels of single nucleotide polymorphisms (SNPs) that could reliably distinguish spring and winter‐spawning Atlantic cod (88.5% assignment rate), as well as males and females (95.0% assignment rate) collected in the western Gulf of Maine. These SNP panels represent a valuable tool for fisheries research and management of Atlantic cod in the western Gulf of Maine that will aid investigations of stock production and support accuracy of future assessments.     

Seasonal maturity development of Baltic cod in different spawning areas: importance of the Arkona Sea for the summer spawning stock

We investigated the seasonal maturity development of cod in four areas of the Baltic Sea. Two different spawning peaks were identified and found to be consistent over the period 1992–2005. In the Kiel Bight and Mecklenburg Bight (ICES SD 22) a spawning peak was observed from March to April (spring spawning). In the areas of the Arkona Sea (ICES SD 24) and Bornholm Sea (ICES SD 25) the spawning peak occurred during summer. In the Bornholm Sea, the main spawning activities began in June and ended in September, with a spawning peak in June–August (summer spawning). In the Arkona Sea, which is a transition area between the Mecklenburg Bight and the central Baltic Sea, spawning began in March and lasted until July, with a spawning peak in June–July (summer spawning). Seasonal maturity development and proportions of spawning cod in June in the Arkona Sea were similar to that of the Bornholm Sea. In addition, the proportion of spawning cod in the Arkona Sea was positively correlated with the size of the spawning stock in the Bornholm Sea. Our results provide evidence of a spatial expansion of spawning activities of the summer spawning stock from the eastern Baltic Sea into the Arkona Sea. Therefore, the Arkona Sea should be considered as one of the spawning habitats of the summer spawning stock of Baltic cod.     

Mitochondrial DNA sequence variation and genetic stock structure of Atlantic cod (Gadus morhua) from bay and offshore locations on the Newfoundland continental shelf

Bay cod, Atlantic cod (Gadus morhua) that over-winter in the deep-water bays of northeastern Newfoundland, have historically been regarded as distinct in migration and spawning behaviour from offshore (Grand Bank) cod stocks. To investigate their genetic relationships, we determined the DNA sequence of a 307-base-pair portion of the mitochondrial cytochrome b gene for 236 adult cod taken from the waters off northeastern Newfoundland, including fish found over-wintering and spawning in Trinity Bay. Although 17 genotypes were found, a single common genotype occurs at a frequency of greater than 80% in all samples, and no alternative genotype occurs at a frequency of greater than 3%. Genotype proportions did not differ significantly among samples. Measures of genetic subdivision among sampling locations are nil. Cod over-wintering in Trinity Bay are not genetically distinct from offshore cod. In combination with tagging and physiological studies, these data suggest that there is sufficient movement of cod between bay and offshore locations to prevent the development or maintenance of independent inshore stocks. Adult cod that over-winter in Trinity Bay appear to represent an assemblage of temporarily nonmigratory fish that have become physiologically acclimated to cold-water inshore environments. The pattern of genetic variation in northern cod suggests a recent population structure characterized by extensive movement of contemporary individuals superimposed on an older structure characterized by a bottleneck in the population size of cod in the north-western Atlantic.     

Genetic Analysis of the Populations of Japanese Anchovy (Engraulidae: Engraulis japonicus) Using Microsatellite DNA

We analyzed the population structure of the Japanese anchovy (Engraulis japonicus), a small pelagic fish, using 6 microsatellite DNA loci. The anchovy is known to have 2 separate spawning populations, one near northeastern Taiwan in the Pacific Ocean and the other near southwestern Taiwan in the Taiwan Strait. The planktonic larvae then drifted north to the feeding grounds in the East China Sea to advance in their life history. Three populations of the anchovy were analyzed, including 2 temporal population from the northeastern spawning ground (I-Lan 1999 and I-Lan 2000) and one population from the southwestern spawning ground (Peng-Hu 2000). The genetic variability of the 6 loci was high for all the populations. The average numbers of alleles per population ranged from 25.5 to 32.3, and the average observed heterozygosity ranged from 0.559 to 0.650. A significant population differentiation was found between geographic populations but not between the temporal populations. However, the level of geographic differentiation was weak, average FST 0.0088. The significant geographic population structure indicated that the populations of 2 spawning grounds belonged to separate stocks. Moreover, 16 of the 18 population-locus cases showed significant departure from Hardy-Weinberg equilibrium, implying that each spawning population in turn consisted of mixed native stocks. Finally, we posed 3 population models to be evaluated against the genetic data disclosed with the microsatellite markers.     

Spatio-temporal overlap of the alien invasive ctenophore <Emphasis Type="Italic">Mnemiopsis leidyi</Emphasis> and ichthyoplankton in the Bornholm Basin (Baltic Sea)

In 2007 the alien invasive ctenophore Mnemiopsis leidyi A. Agassiz 1865 was recorded for the first time in the Bornholm Basin, an area which serves as important spawning ground for Baltic fish stocks. Since M. leidyi is capable of preying upon early life stages of fish and further might act as food competitor for fish larvae, it is of major concern to investigate the potential threat that this non-indigenous species poses to the pelagic ecosystem of the Baltic Sea. The present study investigates the temporal and spatial overlap of M. leidyi with eggs and larvae of Baltic cod (Gadus morhua L.) and sprat (Sprattus sprattus L.) in order to assess the potential impact of this new invader on two of the most important Baltic fish stocks. Results show variable inter-seasonal distribution and overlap dynamics and thus different seasonal threat-scenarios for the early life stages of cod and sprat. The spatial overlap between M. leidyi and ichthyoplankton was low for most of the period observed, and we conclude that M. leidyi presently does not have a strong impact. However, we detected situations with high overlaps, e.g. for sprat larvae and cod eggs in spring. As the population dynamics of M. leidyi in the central Baltic are not yet fully understood, a future population explosion of the alien ctenophore with possible effects on fish recruitment cannot be ruled out. Furthermore, a possible shift in peak spawning of cod to the early season, when ctenophore abundances were relatively high, might increase the impact of M. leidyi on cod.     

Humpback whale (Megaptera novaeangliae) and killer whale (Orcinus orca) feeding aggregations for foraging on herring (Clupea harengus) in Northern Norway

Aggregations of predators on food patches have been documented for both terrestrial and marine animals. Here, we documented for the first time, and investigated, non-predatory aggregations occurring between humpback whales (Megaptera novaeangliae) and killer whales (Orcinus orca) while feeding on wintering Norwegian spring spawning herring (Clupea harengus) in Andfjord, northern Norway. Observational data were collected during 109 opportunistic surveys through three seasons 2013–2016. Killer whales were observed feeding on 59 occasions, with one to three humpback whales involved in 47 of these feeding events (79.7%), and there was an increased probability of finding feeding humpback whales when feeding killer whales also were observed. With killer whales identified as the initiating species in 94.4% of the feeding aggregations for which the first species was known, and with humpback whales joining and feeding on the fish ball afterwards, we suggest that humpback whales may benefit more from these aggregations than the opposite.     

Effects of varying temperature and food availability on growth and reproduction in first‐time spawning female Atlantic cod

The somatic growth, sexual maturation and fecundity of individually marked first‐time spawning female Atlantic cod Gadus morhua were examined under different varying temperature and feeding regimes over the months preceding spawning. A negative correlation between somatic and oocyte growth was found, reflecting the changing energy allocation pattern. Nevertheless, the somatic growth of mature individuals was at least as high as those of immature fish over the period of vitellogenesis. Potential fecundity was positively correlated with body size, but neither temperature or feeding regime significantly affected this relationship. Consequently, fish with unlimited feeding opportunity invested more energy into somatic growth during vitellogenesis compared to those held under a restricted ration. This indicated that once Atlantic cod had made the decision to invest in first reproduction, they allocated a certain amount of energy relative to their size into egg production and any surplus was invested into somatic growth. Low temperature led to an arrest in the onset of vitellogenesis and significantly affected the number of females that matured.