Assessment of production of eggs of eastern Baltic cod (<Emphasis Type="Italic">Gadus morhua callarias</Emphasis> L.) on the basis of long-term ichthyoplankton data

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 abundance and patterns of the egg and larval spatial distribution of Eastern Baltic cod Gadus morhua callarias (Gadidae) in 1931–1996

According to ichthyoplankton surveys of AtlantNIRO and published sources, the data on abundance of eggs and larvae of eastern Baltic cod Gadus morhua callarias at different spawning grounds of the Baltic Sea in 1931–1938, 1949–1956, and 1991–1996 are considered. We show that a similar type of spatial distribution of eggs and larvae of cod with localization of the greatest amount of its early ontogenetic stages in the Bornholm Basin in the southwest of the sea was observed in the 1930s and 1990s. The extension of the area of distribution was accompanied by an increase in the numbers of eggs and larvae of cod in all areas of the sea, and its decline was followed by a decrease in numbers, in the first turn, at northeastern spawning grounds in the Gotland Basin. The period of high numbers of eggs in 1949–1956 coincided with the century maximum of salinity of the near-bottom water layer.     

Oxygen and salinity characteristics of predator–prey distributional overlaps shown by predatory Baltic cod during spawning

In the distributional overlap volume of Baltic cod Gadus morhua and its prey, studied in the Bornholm Basin in the southern Baltic Sea, only a fraction of the sprat Sprattus sprattus population vertically overlapped with the Baltic cod population. Sprat occurred in the intermediate water, in the halocline and in the bottom water, while herring Clupea harengus and Baltic cod occurred exclusively in the halocline and in the bottom water. Only parts of the sprat population were hence accessible for Baltic cod, and only a fraction of the sprat had access to the Baltic cod eggs below the halocline. Baltic cod–clupeid overlap volumes appeared to be determined by salinity stratification and oxygenation of the bottom water. Hydrography time series were used to estimate average habitat volumes and overlap from July to September in 1958–1999. In the 1999 survey spawning Baltic cod had greater ratios of empty stomachs and lower average rations than non-spawning Baltic cod. The average ration for Baltic cod caught within 11· 4 m from the bottom (demersal) did not differ from the average ration of Baltic cod caught in shallower waters (pelagic), because spawning and non-spawning Baltic cod in both strata were caught at equal rates. The diet of the Baltic cod caught demersally contained more benthic invertebrates, especially Saduria entomon, but Baltic cod caught pelagically also had fresh benthic food in their stomachs, indicating vertical migration of individual fish.     

Evidence of a hybrid-zone in Atlantic cod (Gadus morhua) in the Baltic and the Danish Belt Sea revealed by individual admixture analysis

The study of hybrid zones is central to our understanding of the genetic basis of reproductive isolation and speciation, yet very little is known about the extent and significance of hybrid zones in marine fishes. We examined the population structure of cod in the transition area between the North Sea and the Baltic Sea employing nine microsatellite loci. Genetic differentiation between the North Sea sample and the rest increased along a transect to the Baltic proper, with a large increase in level of differentiation occurring in the Western Baltic area. Our objective was to determine whether this pattern was caused purely by varying degrees of mechanical mixing of North Sea and Baltic Sea cod or by interbreeding and formation of a hybrid swarm. Simulation studies revealed that traditional Hardy-Weinberg analysis did not have sufficient power for detection of a Wahlund effect. However, using a model-based clustering method for individual admixture analysis, we were able to demonstrate the existence of intermediate genotypes in all samples from the transition area. Accordingly, our data were explained best by a model of a hybrid swarm flanked by pure nonadmixed populations in the North Sea and the Baltic Sea proper. Significant correlation of gene identities across loci (gametic phase disequilibrium) was found only in a sample from the Western Baltic, suggesting this area as the centre of the apparent hybrid zone. A hybrid zone for cod in the ecotone between the high-saline North Sea and the low-saline Baltic Sea is discussed in relation to its possible origin and maintenance, and in relation to a classical study of haemoglobin variation in cod from the Baltic Sea/Danish Belt Sea, suggesting mixing of two divergent populations without interbreeding.     

Long-term variation in dates of mass spawning of cod Gadus morhua callarias (Gadidae) in the southeastern part of the Baltic Sea

Long-term data on the numbers of eggs of cod Gadus morhua callarias in ichthyoplankton and the seasonal variation of their distribution in the Gdansk Deep of the Baltic Sea are considered. The spring peak of spawning was observed for the years 1948–1954 and 1960–1964; the spring-summer peak for 1968–1977; and the summer peak for 1955–1959 and 1995–2001. Maximum egg numbers of cod were recorded from the late 1940s to the early 1950s; minimums were recorded in the late 1980s and the late 1990s. The decrease in the numbers of cod eggs in the ichthyoplankton, in general, coincided with a decrease in the proportion of spawners maturing at the beginning of the season of mass reproduction and a decrease in the oxygen content in the near-bottom layer of the Gdansk Deep. It is suggested that a shift of the spawning peak of cod from spring to summer is related to a decrease in the volume of near-bottom waters providing favorable conditions for the reproduction of this population.     

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.     

Changes in the diet of the coastal cod <Emphasis Type="Italic">Gadus morhua marisalbi</Emphasis> in the Kandalaksha Gulf of the White Sea under conditions of increased abundance of three-spined stickleback <Emphasis Type="Italic">Gasterosteus aculeatus</Emphasis>

The data on feeding of coastal cod Gadus morhua marisalbi in the area of the Chupa Bay (Kandalaksha Gulf of the White Sea) are presented. In the food spectrum of cod, over 25 cm long fish and their eggs dominated (77.5% by frequency of occurrence and 91.7% by weight). Other groups of food organisms occupied a secondary place. Among fish, three-spined stickleback Gasterosteus aculeatus was the most prominent food item in the diet of cod (48.3% by weight). The published data on long-term changes of the diet of cod are considered, which are mostly related to considerable fluctuations of the abundance of three-spined stickleback in the White Sea. It is shown that, at present, three-spined stickleback again plays an important role in the diet of the White Sea cod     

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.     

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.     

The ability of Baltic cod eggs to maintain neutral buoyancy and the opportunity for survival in fluctuating conditions in the Baltic Sea

In the brackish water of the Baltic Sea successful spawning of Baltic cod Gadus morhua is restricted to the Bornholm, Gdansk and Gotland basins below the halocline, occurring at 50–80 m depth. Due to irregular mixing of the deep water, stagnant conditions occur regularly accompanied with unfavourable oxygen conditions. In avoiding stressful oxygen conditions maintenance of egg buoyancy is considered a major limiting factor for successful spawning of Baltic cod. Batches of eggs were incubated experimentally in a density gradient column. Egg specific gravity changed during development, decreasing from the time of gastrulation, then increasing prior to hatching. The changes in specific gravity varied among egg batches from different females and were related to egg quality, egg size and ambient salinity. Eggs achieve different specific gravity depending on incubation salinity. Initial egg specific gravity together with the ability of eggs to gain and maintain buoyancy up to hatching, determine larval specific gravity and the depth where hatching will occur, and thus opportunities for larval survival, avoiding stressful oxygen conditions and developing at favourable feeding conditions.     

Size and visibility of Baltic cod eggs with reference to size-selective and stage dependent predation mortality

Size of Baltic cod eggs from incubation experiments and from field samples was determined by microscopic analysis. Results from plankton samples were compared with corresponding size distributions of cod eggs found in herring stomachs. The influence of fixation on size of different developmental stages was studied. Live eggs from incubation experiments were also sized repeatedly throughout the developmental period with an optical plankton counter (OPC) based on light attenuance measurements as this was assumed to be more closely related to the visibility of the eggs for potential predators than egg diameter as obtained by microscopic analysis.
Preservation in formaldehyde solution caused a small reduction in egg diameter (2.2%) whereby no differences between the developmental stages were detected. Egg size decreased slightly during incubation (6.9%) while the OPC measurements revealed a substantial increase in light attenuance during egg development (42.2%). In the field, a general decrease in egg size with increasing depth was observed while no change between the developmental stages was detectable. The mean size of eggs ingested by herring was slightly lower than in the water column which was most pronounced for the late stages containing a well-developed embryo. The frequency of eggs in an advanced stage of development was considerably higher in the stomachs than in corresponding plankton samples. Therefore, it is suggested that the selection of further developed egg stages by predatory fish in the central Baltic Sea, i.e. herring and sprat, is due to an increase of visibility during egg development in relation to growth and pigmentation of the embryo. Thus it is likely that egg mortality due to predation is stage-dependent rather than strictly size-selective.     

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.     

Differences in Salinity Tolerance and Gene Expression Between Two Populations of Atlantic Cod (Gadus morhua) in Response to Salinity Stress

Populations of marine fish, even from contrasting habitats, generally show low genetic differentiation at neutral genetic markers. Nevertheless, there is increasing evidence for differences in gene expression among populations that may be ascribed to adaptive divergence. Studying variation in salinity tolerance and gene expression among Atlantic cod (Gadus morhua) from two populations distributed across a steep salinity gradient, we observed high mortality (45% North Sea cod and 80% Baltic Sea cod) in a reciprocal common garden setup. Quantitative RT-PCR assays for expression of hsp70 and Na/K-ATPase α genes demonstrated significant differences in gene regulation within and between populations and treatment groups despite low sample sizes. Most interesting are the significant differences observed in expression of the Na/K-ATPase α gene in gill tissue between North Sea and Baltic cod. The findings strongly suggest that Atlantic cod are adapted to local saline conditions, despite relatively low levels of neutral genetic divergence between populations.     

On differentiation of cod (Gadus morhua L.) groups in Baltic Sea

Russian Journal of Genetics - Using the AGP*, PGI-1*, PGI-2*, LDH*, IDH*, and PGM* allozyme markers, the differentiation of cod groups during the spawning period in Baltic Sea was evaluated. It was...     

Long-term prediction on Baltic fish stocks based on periodicity of solar activity

The methods presently in wide use for the assessment of marine ecosystems and fish stocks cannot provide the high-quality long-term prognoses urgently needed for improved management of marine ecosystems and their fishery resources. A novel method of forecasting the long-term qualitative composition of ecosystem and fish stocks in the Baltic Sea based on the periodicity and succession of the processes: extraterrestrial factors????climate changes????regime forming factors (chiefly salinity and temperature)????ecosystems????fish stocks was used in 1989 to predict the approximate year class abundances of cod (Gadus morhua callarias), sprat (Sprattus sprattus balticus), sea herring and gulf herring (Clupea harengus membras) in the Baltic Sea for the years 1989?C2008. A comparison with actual development up to 2008 show that this prognosis was confirmed to a considerable extent. The method based on periodical fluctuations of freshwater input and the thermal regime is described. It predicts a new regime shift in the 2020s to a higher salinity and more acceptable conditions for the organisms of marine origin.     

The potential fecundity of cod in the Baltic Sea from 1993 to 1999

Investigations on the reproduction biology of cod in the Baltic Sea (Gadus morhua L. and G. morhua callarias L.) were carried out beginning in 1993, with yearly sampling of gonads to estimate the potential absolute and relative individual fecundity in the Belt Sea (subdivision, SD: 22) and in the Arkona Sea (SD: 24). Additional samples were taken in the Bornholm Sea (SD: 25). The potential absolute fecundity of the total area of investigation is described by the regression models: F_a = 6.89 × TL^3.09 and F_a = ?40 586 + 977.7 × W. As a further result the mean relative fecundity was 978 oocytes per gram body weight. Furthermore, it was shown that there was a difference in development of the potential individual fecundity between the Belt Sea cod and the original eastern Baltic cod stock. In contrast to the stable fecundity of cod in the Belt Sea, the fecundity of the large females in the Bornholm Sea significantly increased from 1993 to 1999. This change is related to a decrease in the dry weight of oocytes in the ovaries. In the same period this cod stock dramatically decreased.     

Diseases and Injuries Associated with Mortality of Hatchery Reared Baltic Cod (Gadus morhua L.) Larvae

A cod hatching plant was established in 1992 on the island of Bornholm in the Baltic Sea in order to elucidate the possibilites for restocking of cod fry in this brackishwater system. The disease prevalence in 3 batches of hatchery-reared yolksac larvae from the Baltic cod (Gadus morhua L.) was monitored during the posthatch period. High prevalences of bacteriosis/mycosis, lordosis/scoliosis, injuries and protozoan endoparasitism were recorded. Vibrio sp. and Vibrio anguillarum serovar 04, 06, 08 in addition to nontypable strains and saprolegniaceous fungi were isolated from the larvae. The dinoflagellate-like endoparasites were located in the yolksac of the cod larvae.     

Effects of varying natural mortality and selectivity on the assessment of eastern Baltic cod (Gadus morhua Linnaeus, 1758) stock

In 2014, the International Council for the Exploration of the Sea (ICES) was unexpectedly unable to provide an analytical assessment of eastern Baltic cod stock; factors such as data issues, assessment methodology, and the ecological situation of cod were indicated as the reasons for this failure. Some evidence suggests that the natural mortality (M) of cod could increase substantially in forthcoming years and that the selectivity could change. In this paper, age‐structured and stock‐production assessment models were applied to simulate the dynamics of cod stock; in the models, both constant and increasing natural mortalities were permitted. In the age‐structured model, the effects of selectivity related to the cod size on the cod assessment were also analysed. In addition, stock with characteristics similar to Baltic cod stock and increasing natural mortality was generated and assessed with the age‐structured model using both constant and increasing M. It was shown that models with increasing natural mortality of cod in recent years perform much better than models with constant natural mortality in terms of the distribution of residuals and retrospective patterns. The models with size‐dependent selectivity did not perform better than other standard assessments. The assessment of generated stock (where natural mortality was increasing) with constant natural mortality in the assessment model showed a poor distribution of residuals and strong retrospective patterns, similar to the ICES assessment with constant M. The conducted simulations strongly suggest that the main reason for the poor recent cod assessment is the increase in natural mortality, which is not considered in the assessment methodology.     

Could Seals Prevent Cod Recovery in the Baltic Sea?

Fish populations are increasingly affected by multiple human and natural impacts including exploitation, eutrophication, habitat alteration and climate change. As a result many collapsed populations may have to recover in ecosystems whose structure and functioning differ from those in which they were formerly productive and supported sustainable fisheries. Here we investigate how a cod (Gadus morhua) population in the Baltic Sea whose biomass was reduced due to a combination of high exploitation and deteriorating environmental conditions might recover and develop in the 21st century in an ecosystem that likely will change due to both the already started recovery of a cod predator, the grey seal Halichoerus grypus, and projected climate impacts. Simulation modelling, assuming increased seal predation, fishing levels consistent with management plan targets and stable salinity, shows that the cod population could reach high levels well above the long-term average. Scenarios with similar seal and fishing levels but with 15% lower salinity suggest that the Baltic will still be able to support a cod population which can sustain a fishery, but biomass and yields will be lower. At present knowledge of cod and seal interactions, seal predation was found to have much lower impact on cod recovery, compared to the effects of exploitation and salinity. These results suggest that dual management objectives (recovery of both seal and cod populations) are realistic but success in achieving these goals will also depend on how climate change affects cod recruitment.