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.     

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.     

Species and length composition of fish in the south-western Baltic Sea

Length distributions of fish were analysed based on data from the German annual (November/December) bottom trawl surveys in the Mecklenburg Bight and Arkona Basin [ICES Subdivisions (SD) 22 and 24, respectively] of the southwest Baltic Sea, covering the period from 1991 to 2000. The length spectra for all fish species, and those for demersal and pelagic species separately, all showed a consistent pattern of declining logarithmic numbers with increasing length over the fully selected size range. Such consistency allowed the shape of the spectra to be used as a community property that could be tracked in space and time. There was a rapid reduction in the number of specimens in the larger size classes, resulting in steep slopes of the spectra, which may be due to intensive fishing pressure in the Baltic. The species composition of fish communities in the southwest Baltic Sea was stable during the period considered. No consistent species replacement was found, although the clupeid group dropped sharply in the catch during the 1996 survey, which might have been due to sudden environmental changes in that year.     

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.     

Differentiation of western and eastern Baltic Sea cod stocks (Gadus morhua) by means of stable isotope ratios in muscles and otoliths

Stable isotope analysis of Baltic Sea cod from the Bornholm Sea (ICES‐SD 25) and western Baltic (Belt Sea, ICES‐SD 22) revealed significant differences in the δ¹⁵N and δ¹³C values of the dorsal muscle, as well as in the δ¹⁸O values of otoliths. The method pledges to become especially appropriate to the Baltic due to the high variability in oxygen isotope ratios associated with its estuarine nature.     

Sexual maturation of cod (Gadus morhua L.) in the southern Baltic (1990–2006)

Baltic cod, like other species, is susceptible to inter‐annual fluctuations in sexual maturation, depending on the length, age, sex, extent of the habitat area, and stock abundance of the cod population. Maturity is one of the biological indicators used to detect changes in a stock that can be caused by fishing. To address these issues specifically for the eastern Baltic cod stock, long‐term data (1990–2006) from Polish research vessels in the southern Baltic were examined. To date, the ICES has used the same maturity ogives over extended periods and assumed invariant sex ratios for the assessment of eastern Baltic cod. The combined maturity ogives calculated in the present study were markedly lower, particularly for age groups 2–4 (5), in all periods, than those used in the ICES assessment. Moreover, the proportion of females increased with length and age, suggesting that annual verification of the sex ratio is needed. The present study also revealed that the total length (L_50%) and the age (mean age‐at‐maturity; MAM_50%) at which 50% achieved first sexual maturity were higher for females than for males in the study period. The long‐term mean L_50% and MAM_50% for females were 43.9 cm and 4.3 years, respectively, and for males 34.8 cm and 3.4 years. There was also a spatial difference between calculated maturity ogives, with slightly lower L_50% (range: 1.4–8.6 cm) in the Gdańsk Basin than in the Bornholm Basin. The increasing trend in fishing mortality observed in 1993–2004 (ICES data) did not translate into a temporal trend in calculated maturity ogives. However, changes in L_50% and MAM_50% reflected recruitment variations (ICES data). The significance of these findings is discussed in the context of the environment and recruits abundance.     

Phytoplankton spring blooms in the southern Baltic Sea--spatio-temporal development and long-term trends

The seasonal and long-term development of the phytoplanktonspring bloom in different regions of the southern Baltic Seawas investigated on the basis of monitoring data. The developmentof a spring bloom starts when the upper mixed layer becomesshallower than the euphotic zone, as proved also by a mesocosmexperiment. This already happens in March in Mecklenburg Bightand the western part of the Arkona Sea, leading to a diatombloom, but only in April in the Bornholm Sea, increasingly givingrise to a dinoflagellate bloom. The new production of the springphytoplankton may be calculated from the decrease in nutrientsduring spring. In comparison with the Redfield ratio, phosphorusis taken up in excess (N:P = 9.2–10.2). The consumptionof silicate in spring has been reduced in the southern Balticproper since 1989, pointing to a decline in diatoms. The increasein chlorophyll a in the Bornholm and the southern Gotland Seasis related to eutrophication, whereas the decrease in diatomsin favour of the dinoflagellates is related to mild winters.The lack of deep-reaching circulation after mild winters maybe one reason for the suppression of the non-motile diatoms.     

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.     

Fecundity of Baltic cod: temporal and spatial variation

Potential and relative potential fecundity of 701 cod Gadus morhua caught in the Bornholm Basin (ICES Sub-division 25) during 1987–1996 and 106 from the Gdansk Deep (ICES Sub-division 26) in 1995 and 1996 varied significantly between years but neither between different spawning grounds nor between different sampling months within one spawning season. Growth and water temperature are factors identified to influence the mean relative fecundity explaining 66% of the variance between years.     

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.     

Separating two herring stocks from growth data: long‐term changes in survey indices for Western Baltic Spring Spawning Herring (Clupea harengus) after application of a stock separation function

In the Baltic Sea, herring stocks are surveyed and managed according to a spatial allocation based on ICES (International Council for the Exploration of the Sea) subdivisions. In the western Baltic, the distribution areas of two stocks overlap: the Western Baltic Spring Spawning Herring (WBSSH) and the Central Baltic Herring (CBH). Survey results of length‐at‐age data indicate in Subdivision (SD) 24, which is a part of the WBSSH management area, that a considerable fraction of CBH is present and correspondingly erroneously allocated to WBSSH stock indices. Accordingly, a stock Separation Function (SF) based on growth parameters was established to identify the fraction of CBH in the WBSSH area. In the present study, the SF was applied to 8 years of data from the GERman Acoustic Survey (GERAS), which is conducted annually in autumn in ICES subdivisions 21‐24. Results showed a distinct fraction of CBH in SD 24, and exclusion of the CBH greatly improved the quality of the GERAS index used in the assessment of the WBSSH stock.     

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.     

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.     

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.     

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.     

Reproduction of the summer flounder, Paralichthys dentatus (L.)

The reproduction of summer flounder, Paralichthys dentatus (L.), occurring in the Middle Atlantic Bight was studied from 1974–1979. Males dominated the length interval between 21–35 cm t.l. and were essentially absent in samples >55 cm t.l. Females were more abundant in all length intervals >45 cm t.l. Length at maturity ranged from 24–27 cm t.l. for males and from 30–33 cm t.l. for females which coincided with length at age two. Ovarian egg diamerter frequencies indicated summer flounders are serial spawners and the trend in mean and maximum maturity indexes (% ovary weight of total fish weight) indicated spawning began in September and continued through February in 1975 and probably into March in 1976. Annual condition cycle of males peaked in September and was lowest in April while females' cycle reached a maximum in October which coincided with peak spawning time. Fecundity was related to length, weight and ovary weight for 1974–1977 and the length-fecundity equation F = 0.0007975 L ^3.402 was found to be the best predictive relationship. The overall reproductive strategy of summer flounder was discussed and apparently tends to maximize reproductive potential through an extended spawning season, early maturation, high fecundity, serial spawning and extensive spawning migrations.     

Evidence for metapopulation structuring in cod from the west of Scotland and North Sea

The scale of population structuring in cod Gadus morhua from the west of Scotland (ICES stock area VIa) and northern North Sea was investigated using a combination of non-genetic methods. Site fidelity of spawning aggregations was examined using historic tag-recapture experiments and individual geolocation estimates from data storage tagged fish. The extent of movements from spawning areas indicated by these two tagging approaches was broadly similar. Between 67 and 97% of adult cod remained within 100 km of spawning areas throughout the year, suggesting resident spawning groups. A small proportion of cod, however, did appear to stray between spawning areas. A comparison of the elemental signature of the otoliths of 0 year-group and the 0 year-group component from the same year-class at age 2 years indicated that most adults originate from local nursery areas. Moreover, there did not appear to be detectable exchange between spawning areas >200 km apart, such as the Clyde and Minch or Shetland and the Inner Hebrides. As such, population processes may operate at a smaller spatial scale than the stock level, with spawning aggregations functioning as local populations within a metapopulation. The relevance of this scale of dynamics is discussed in relation to recently imposed closed areas in the region.     

Ecological and genetic impact of Atlantic cod larval drift in the Skagerrak

We evaluate the hypothesis that Atlantic cod larvae are passively transported by sea currents from off-shore spawning areas to settle in coastal waters, a hypothesis which has recently gained support from genetic analysis of cod in the North Sea-Skagerrak area. Such larval transport has been suggested to be an important mechanism behind the commonly observed low spatial genetic differentiation in many marine organisms. Here, we apply an ARMAX(2,2) model for juvenile abundance and use long-term monitoring data from the Skagerrak coast, constituting 54 continuous annual series from 1945 to 1997. Analysing the model, we find that the product of the size of the North Sea breeding stock and the strength of the net inflow of North Sea waters had a significant, positive effect on the abundance of coastal juvenile cod. The peak effect occurs during the month of March, just after spawning, when eggs and larvae remain pelagic and sensitive to currents. In contrast, we find no evidence of any direct effect of the North Sea spawning stock alone. Our analyses indicate that 15-20,000 0-group larvae from the North Sea reach each fjord per year, on average. This corresponds to about 1-10% of the total 0-group population in each fjord on average. These findings clearly demonstrate a direct link between larval drift and gene flow in the marine environment.     

Assessment of production of eggs of Eastern Baltic cod (Gadus morhua callarias 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.