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.     

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.     

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.     

Effect of fish length and nutritional condition on the fecundity of distressed Atlantic cod Gadus morhua from the Baltic Sea

The disappearance of larger individuals and the decrease in individual body condition suffered by Atlantic cod Gadus morhua in the eastern Baltic during the past two decades can be expected to affect the stock reproductive output. To investigate this, female G. morhua were collected during the spawning and pre‐spawning period in 2015?2016. The current individual potential fecundity (F_P) of eastern Baltic G. morhua was estimated and analysed in relation to total length (L_T) and indices of nutritional status such as body condition (K) and hepato‐somatic index (I_H) using generalized linear models. In addition, the current prevalence of atresia and its potential relation to K were investigated. Moreover, a calibration curve to estimate F_P from oocyte diameter, based on the autodiametric oocyte counting method, was established for the first time for eastern Baltic G. morhua and can be used for future fecundity studies on this stock. The results showed that F_P was mainly positively related to fish length, but K and I_H also contributed significantly to the variation in F_P. The model predicted that fish with K = 1·2 have a F_P 51% higher than fish of the same L_T with K = 0·8. The prevalence of fecundity regulation by atresia was 5·8%, but it was found only in fish in the pre‐spawning maturity stage and with low K. Temporal changes in biological features such as the length composition and individual body condition of eastern Baltic G. morhua, should be accounted for when estimating stock reproductive potential.     

Genetic differentiation of southeast Baltic populations of sea trout inferred from single nucleotide polymorphisms

Sea trout (Salmo trutta m. trutta) is a migratory form of brown trout common in the Baltic Sea. Nine populations from the southeast Baltic (Poland; Lithuania; Denmark, Bornholm; Estonia and Russia) were genotyped using iPLEX Gold technology (Sequenom) with 62 informative SNPs. A diagnostic panel of 23 SNPs was applied to estimate genetic differentiation and assess the population structure of Baltic sea trout. The highest level of pairwise F_ST differences was observed between the Russian (East Gulf of Finland) and Polish (Baltic main basin) populations. The lowest differences were between the two Polish and the Polish and Lithuanian populations. A genetic similarity was noted between the Estonian Riguldi River and Danish Bornholm populations, and this finding was supported by a Bayesian and factorial correspondence analysis. Diversity within populations was highest for populations from Estonia and lowest for the Lithuanian population. Genetic structure analysis indicated that individuals from the nine populations were clustered into four groups.     

The reproductive biology of the two-banded sea bream (Diplodus vulgaris) from the southwest coast of Portugal

The study of the reproduction of Diplodus vulgaris (Geoff.) as part of a base‐line study of the fishery resources of the south‐west coast of Portugal, was based on the analysis of the spawning season, gonad maturation, size at maturity, fecundity and hermaphroditism. The spawning season is relatively long, from December to March, with peaks in January and February. No significant differences were found either in the sex ratio (M : F = 1.01) over the year or by size. The size at first maturity (L₅₀) for all sexes and undetermined individuals combined was 18.33 cm total length (TL), with no significant differences between males and females. The estimated L₅₀ is considerably greater than the minimum legal size in Portugal of 15.0 cm. Mean absolute fecundity (F_a) was 131 127 oocytes, ranging from 31 523 to 250 608. The relationship between absolute fecundity and total length (TL) (F_a = 25 398 TL – 484 426) and somatic weight (SW) (F_a = 878.8SW – 71 416) was of the linear type. The mean number of oocytes per gram of somatic weight was 526, ranging from 194 to 887. The reproductive strategy of this species is characterized by a rudimentary hermaphroditism with possible protandry, as evidenced by the existence of individuals in sexual transition.     

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.     

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.     

Spreading and physico-biological reproduction limitations of the invasive American comb jelly Mnemiopsis leidyi in the Baltic Sea

Hydrodynamic drift modeling was used to investigate the potential dispersion of Mnemiopsis leidyi from the Bornholm Basin in the Baltic Sea where it has been observed since 2007 further to the east and north. In the brackish surface layer dispersion is mainly driven by wind, while within the halocline dispersion is mainly controlled by the baroclinic flow field and bottom topography. Model runs showed that the natural spreading via deep water currents from the Bornholm Basin towards north and east is limited by topographic features and low advection velocities. Based on the information on ranges of salinity and temperature, which limit survival and reproduction of this ctenophore within the Baltic Sea, areas have been identified where the American comb jelly, M. leidyi could potentially survive and reproduce. While, we could show that M. leidyi might survive in vast areas of the northern Baltic Sea its reproduction is prevented by low salinity (<10 psu) and temperature (<12°C). Thus, due to the combined effect of low salinity and temperature, it is not probable that M. leidyi could establish permanent populations in the central or northern Baltic Sea. However, it seems that in the southern parts of the Baltic Sea environmental conditions are suitable for a successful reproduction of M. leidyi.     

Cod migration and transplantation experiments in the Baltic

In the first half of the paper a review is given of the occurrence and migratory habits of Baltic cod based on experience since the 1950s, results of extensive tagging experiments combined with information on changes in allele frequency for haemoglobin types, in meristic characters, and in otolith types. The latter pan of the paper is an account of about 15 transplantation experiments with tagged cod to study a potential homing ability of the fish. The very great importance of the threshold areas inside the Baltic and out to the Kattegat is stressed, and so are the hydrographic conditions in general for cod recruitment and migration. The waters west of Bornholm constitute an area of hydrographic instability with varying cod migrations and passive transport of fry with the currents. The migration east of Bornholm refers—except for local stocks and a varying contribution from the west-mainly to fish grown up in the central Baltic proper and northern areas. The latter fish migrate primarily southward for spawning, and as adults they usually stay in the southern waters east and north of Bornholm. The results of the transplantation experiments support a strong direct linkage of cod migration and hydrographic factors. Cod that were tagged and transplanted to a new area behaved and moved in the same way as the local stock. Indications of “homing” can be found in areas with suitable hydrographic gradients, such as changes in salinity, for example in the Öresund.     

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.     

Fecundity and the survival conditions of the offspring of the Baltic Cod <Emphasis Type="Italic">Gadus morhua callarias</Emphasis> (Gadidae)

The interannual and ontogenetic dynamics of the fecundity of the Baltic cod Gadus morhua callarias and the dependence of the number of the offspring on the reproduction conditions were studied. The absolute individual fecundity and the relative individual fecundity had no pronounced interannual dynamics for the period of 1994?2013. The maximal population fecundity was observed in early 1980, the minimal, in 1990?2000. Nearly every year, the highest contribution to the population fecundity belongs to the 4?5-year old females. The ranking of the survival rates and the year-class strength for the females at the age of 2 years showed that the emergence of the generations of high and average productivity (1980?1986) referred to the unfavorable and moderate survival conditions during the early ontogenesis. During the period of small stock (after 1987), the conditions were mostly favorable and moderate. It is suggested that the significant role in the offspring dynamics belongs to the factors linked to the population density. After 2005, favorable survival conditions promoted the increase of the Baltic cod abundance; this increase was also supported by the decrease of the commercial fishing and by the sporadic large-scale advective processes.     

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.     

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.     

Effect of salinity on filtration rates of mussels Mytilus edulis with special emphasis on dwarfed mussels from the low-saline Central Baltic Sea

The effect of salinity on the filtration rate of blue mussels, Mytilus edulis, from the brackish Great Belt (Denmark) and the low-saline Central Baltic Sea, respectively, was studied. First, we measured the effect of long-term (weeks) constant ambient salinities between 5 and 30 psu on the filtration rate of M. edulis collected in the Great Belt where the mean salinity is 17 psu. At salinities between 10 and 30 psu, the filtration rates did not vary much, but at 5 psu the filtration rates were significantly lower. Next, we studied dwarfed M. edulis (<25 mm shell length) from Central Baltic Sea (Askö, Sweden) where the mean salinity is 6.5 psu. The maximum filtration rate (F, ml min^?1 ind.^?1) as a function of shell length (L, mm) and dry weight of soft parts (W, mg) were found to be: F = 0.003L ^2.71 and F = 0.478W ^0.92, respectively, and these results indicate that the filtration rates of dwarfed Baltic Sea mussels are comparable to filtration rates of Great Belt mussels of similar size exposed to salinities >10 psu. When Baltic Sea mussels acclimatized to 20 psu in the laboratory were exposed to 6.5 psu this caused a drastic reduction in the filtration rate, but after about 2 days the previous high filtration rate was regained at 6.5 psu, and further, a similar pattern was observed when the 6.5 psu exposed mussels were finally re-exposed to 20 psu. The observed lack of Great Belt mussels to completely adjust to 5 psu, in contrast to the ease of Baltic Sea mussels to adjust back and forth between 6.5 and 20 psu, is remarkable and may perhaps be explained by different genotypes of Great Belt and Baltic Sea mussels.     

Impact of hypoxia on consumption of Baltic cod in a multispecies stock assessment context

The Baltic Sea is characterised by a heterogeneous oceanographic environment. The deep water layers forming the habitat of Baltic cod (Gadus morhua callarias L.) are subjected to frequently occurring pronounced anoxic conditions. Adverse oxygen conditions result in physiological stress for organisms living under these conditions. For cod e.g. a direct relationship between oxygen availability and food intake with a decreasing ingestion rate at hypoxia could be revealed. In the present study, the effects of oxygen deficiency on consumption rates were investigated and how these translate to stock size estimates in multi‐species models. Based on results from laboratory experiments, a model was fitted to evacuation rates at different oxygen levels and integrated into the existing consumption model for Baltic cod. Individual mean oxygen corrected consumption rates were 0.1–10.9% lower than the uncorrected ones. At the currently low predator stock size, however, the effect of oxygen‐reduced consumption on the total amount of eaten prey biomass and thus predation mortalities was only marginal. But should successful management lead to higher cod stock sizes in the future, then total predation mortalities will greatly increase and thus improved precision of these estimates would be valuable for the assessment of prey stocks.     

Life history characteristics of an invasive cyprinid fish (Carassius gibelio) in Chimaditis Lake (northern Greece)

The life history characteristics of an invasive cyprinid fish, the Prussian carp (Carassius gibelio) were examined in Lake Chimaditis (northern Greece). The population is dominated by females that are able to reproduce gynogenetically. Fish samples were collected using trammel nets from August 2004 to July 2005. Their total length (TL) ranged between 21.9 and 37.0 cm. The length–weight relationship was W = 0.0336TL^2.81 (r² = 0.92, n = 600) and the mean Fulton condition factor exhibited monthly variation from 0.172 (May) to 0.186 (August). According to the annual growth marks present, the lifespan of the Prussian carp in Lake Chimaditis is 6 years. The growth parameters were estimated as L_∞ = 34.46 cm, K = 0.297 year^?1, t₀ = ?1.994 year. Absolute fecundity (F_A) ranged between 26 000 and 176 600 oocytes (mean = 66990) and relative fecundity between 78 and 251 oocytes per gram of total weight (mean = 158). Absolute fecundity increased exponentially with length (F_A = 0.286 × TL^3.66, r² = 0.66) and weight (F_A = 17.93 × W^1.36, r² = 0.73) of the fish.     

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.     

Variability of microsatellite loci of Greenland cod <Emphasis Type="Italic">Gadus ogac</Emphasis> Richardson 1836: Comparison with other species of <Emphasis Type="Italic">Gadus</Emphasis> genus (Gadidae)

Comparative analysis of variability of seven microsatellite loci—Gmo3, Gmo-G12, Gmo-G18, Gmo19, Gmo34, Gmo35 and Pgmo32—was performed for the Greenland cod Gadus ogac, Pacific cod G. macrocephalus, Atlantic cod G. morhua, and White Sea cod G. morhua marisalbi. High genetic identity was observed between the Greenland cod and Pacific cod (I = 0.9520). Pair analysis of genetic differentiation was performed on the studied microsatellite loci according to θ (analogue of F _ST). The Greenland cod differed significantly from the Pacific, Atlantic, and the White Sea cod; however, the differentiation level varied. The lowest value was observed for the pair Greenland cod-Pacific cod (0.123), and the highest levels were registered for the pairs Greenland cod-Atlantic cod (0.605) and Greenland cod-White Sea cod (0.535).