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.     

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.     

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.     

The dangers of ignoring stock complexity in fishery management: the case of the North Sea cod

The plight of the marine fisheries is attracting increasing attention as unsustainably high exploitation levels, exacerbated by more extreme climatic conditions, are driving stocks to the point of collapse. The North Atlantic cod (Gadus morhua), a species which until recently formed a major component of the demersal fisheries, has undergone significant declines across its range. The North Sea stock is typical of many, with a spawning stock biomass that has remained below the safe biological limit since 2000 and recruitment levels near the lowest on record. Cod within the North Sea are currently managed as a single stock, and yet mounting empirical evidence supports the existence of a metapopulation of regionally variable, genetically distinct, sub-stocks. Applying the same management strategies to multiple stocks that differ in their resilience to exploitation inevitably results in the overfishing and likely collapse of the weaker components. Indeed, recent studies have identified two North Sea spawning stocks that have undergone disproportionally large collapses with very substantial reductions in egg production. Similarly affected cod stocks in the northwest Atlantic have shown little evidence of recovery, despite fishery closures. The possible implications of ignoring sub-structuring within management units for biocomplexity, local adaptation and ecosystem stability are considered.     

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.     

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.     

Climate,plankton and cod

While a few North Atlantic cod stocks are stable, none have increased and many have declined in recent years. Although overfishing is the main cause of most observed declines, this study shows that in some regions, climate by its influence on plankton may exert a strong control on cod stocks, complicating the management of this species that often assumes a constant carrying capacity. First, we investigate the likely drivers of changes in the cod stock in the North Sea by evaluating the potential relationships between climate, plankton and cod. We do this by deriving a Plankton Index that reflects the quality and quantity of plankton food available for larval cod. We show that this Plankton Index explains 46.24% of the total variance in cod recruitment and 68.89% of the variance in total cod biomass. Because the effects of climate act predominantly through plankton during the larval stage of cod development, our results indicate a pronounced sensitivity of cod stocks to climate at the warmer, southern edge of their distribution, for example in the North Sea. Our analyses also reveal for the first time, that at a large basin scale, the abundance of Calanus finmarchicus is associated with a high probability of cod occurrence, whereas the genus Pseudocalanus appears less important. Ecosystem‐based fisheries management (EBFM) generally considers the effect of fishing on the ecosystem and not the effect of climate‐induced changes in the ecosystem state for the living resources. These results suggest that EBFM must consider the position of a stock within its ecological niche, the direct effects of climate and the influence of climate on the trophodynamics of the ecosystem.     

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.     

Species assemblages of pelagic fish embryos in the southern North Sea between 1984 and 2000

The occurrence and abundance of the pelagic eggs of southern North Sea spring-spawning fish were analysed between 1984 and 2000. Species number varied between six (1986) and 14 (1999) and was positively correlated with sea surface temperature. With one exception, dab eggs were always the most abundant and usually highly dominant. Ranking of species depended on temperature, but no significant differences in ranking between years was discernible. Although with the increase in temperature in the 1990s a change in species assemblage was evident [species belonging to the boreal-Mediterranean (Lusitanian) group became more apparent in the species assemblage] this did not lead to an increased species diversity (Shannon Index) or a change in other community parameters. It appears that the recent developments regarding spawning stock biomass of commercial North Sea fish is reflected in the declining egg abundance of the respective commercial and larger species (i.e. cod, flounder, plaice) and an increase in abundance of the eggs of small species (i.e. long rough dab, rockling) over the years. Received in revised form: 5 November 2001 Electronic Publication     

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.     

Multi-torsking: simultaneous measurements of cod behaviour show differences between North Sea and Irish Sea stocks*

Fish behaviour can vary considerably in response to environmental and biological conditions. We used electronic Data Storage Tags (DSTs) to investigate the behaviour of cod in different environments; 74 North Sea cod and 20 Irish Sea cod were tagged and released in late-March 1999. So far, 32 tags (>2000 days of data) have been returned from the North Sea and four (>700 days) from the Irish Sea. The maximum recorded depth of a North Sea cod was 84 m, while Irish Sea cod utilised a depth range of 154 m. Active depth changes were of greater magnitude and rate in Irish Sea cod than North Sea cod. In addition, Irish Sea cod were continually active and made proportionally more active depth changes than North Sea cod, which were very active during migration (March–June) but exhibited little vertical movement during the summer months. To our knowledge, this is the first time that the behaviour patterns of cod from different stocks have been measured simultaneously over extended periods at such a fine temporal scale. In addition to describing considerable inter-stock differences in behaviour, our results suggest that while cod in the Irish Sea remain active and highly mobile throughout the year, North Sea cod reduce their movements during the summer feeding period. Our findings will contribute to the development of more biologically realistic management models used for evaluating the efficacy of fisheries technical measures, such as closed areas.     

Synergistic Effects of Climate and Fishing in a Marine Ecosystem

Current climate change and overfishing are affecting the productivity and structure of marine ecosystems. This situation is unprecedented for the marine biosphere and it is essential to understand the mechanisms and pathways by which ecosystems respond. We report that climate change and overfishing are likely to be responsible for a rapid restructuring of a highly productive marine ecosystem with effects throughout the pelagos and the benthos. In the mid-1980s, climate change, consequent modifications in the North Sea plankton, and fishing, all reduced North Sea cod recruitment. In this region, production of many benthic species respond positively and immediately to temperature. Analysis of a long-term, spatially extensive biological (plankton and cod) and physical (sea surface temperature) dataset suggests that synchronous changes in cod numbers and sea temperature have established an extensive trophic cascade favoring lower trophic level groups over economic fisheries. A proliferation of jellyfish that we detect may signal the climax of these changes. This modified North Sea ecology may provide a clear indication of the synergistic consequences of coincident climate change and overfishing. The extent of the ecosystem restructuring that has occurred in the North Sea suggests we are unlikely to reverse current climate and human-induced effects through ecosystem resource management in the short term. Rather, we should understand and adapt to new ecological regimes. This implies that fisheries management policies will have to be fully integrated with the ecological consequences of climate change to prevent a similar collapse in an exploited marine ecosystem elsewhere. Author Contributions  RRK conceived the project and GB analysed the data. RRK, GB and JAL co-wrote the paper.     

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.     

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 spawning activity of cod, Gadus morhua L.

Aspects of the reproduction of reared cod, Gadus morhua L., with special emphasis on the females, were studied under laboratory conditions. The fecundity and condition factor were 2–5 and 1–5 times, respectively, that of wild cod. A total of 18 spawning females were kept in separate tanks/ chambers, each with one or two males. Seven of the 18 females were classified as stressed, based upon behaviour, irregular spawning intervals and low fertilization rates of the eggs. The reared cod were found to spawn 17–19 batches. The number of eggs liberated in each batch normally followed a smooth, dome-shaped curve with time. The fertilization rate was normally 100%. Egg size decreased from first to last batch and the egg dry weight decreased by about 20–30%. The reared cod showed the same egg diameter to dry weight relation as wild cod. Egg diameter of first batch and maternal fish length were significantly positively correlated. The mean spawning interval for the female and the mean water temperature during its spawning were negatively correlated. The reared cod spawned in both the night and the day for about 50–60 days.     

TaqMan DNA technology confirms likely overestimation of cod (Gadus morhua L.) egg abundance in the Irish Sea: implications for the assessment of the cod stock and mapping of spawning areas using egg-based methods

Recent substantial declines in northeastern Atlantic cod stocks necessitate improved biological knowledge and the development of techniques to complement standard stock assessment methods (which largely depend on accurate commercial catch data). In 2003, an ichthyoplankton survey was undertaken in the Irish Sea and subsamples of 'cod-like' eggs were analysed using a TaqMan multiplex, PCR (polymerase chain reaction) assay (with specific probes for cod, haddock and whiting). The TaqMan method was readily applied to the large number of samples (n = 2770) generated during the survey and when combined with a manual DNA extraction protocol had a low failure rate of 6%. Of the early stage 'cod-like' eggs (1.2-1.75 mm diameter) positively identified: 34% were cod, 8% haddock and 58% whiting. As previous stock estimates based on egg surveys for Irish Sea cod assumed that the majority of 'cod-like' eggs were from cod, the TaqMan results confirm that there was probably substantial contamination by eggs of whiting and haddock that would have inflated estimates of the stock biomass.     

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.     

The influence of maternal length and age on the size and weight of the eggs and the relative fecundity of the haddock, Melanogrammus aeglefinus, in British waters

The diameter and dry weight of the eggs of haddock, Melanogrammus aeglefinus , are positively correlated with fish length. The correlations are largely due to the fact that many of the smaller fish that were sampled were 2 years old, and the eggs of these young haddock, which can be regarded as precocious spawners, are significantly smaller and lighter than those of older fish. The relative fecundity of 2–year–old haddock (274 eggs g⁻¹) is also lower than that of the other age classes (493 eggs g⁻¹); this has important implications for the estimation of egg production from female spawning stock biomass. It is pointed out that in some years precocious spawners represent a large component of the North Sea haddock spawning stock. When the annual egg production of this stock is calculated, it may be appropriate to apply a weighting factor to the number of eggs contributed by the 2–year–olds, on the assumption that the small eggs of these fish produce larvae that are less viable than those of older haddock.     

Temperature-dependent development rates of plaice (Pleuronectes platessa L.) eggs from the Irish Sea

Experimentally derived temperature-dependent development ratesfor planktonic fish eggs are required for developing IndividualBased Models (IBMs) of early life history stages and for estimatingdaily egg production in ichthyoplankton surveys. Developmentrates of Irish Sea plaice eggs were experimentally determinedfor temperatures between 4.5 and 12° C. Differences in developmentrates were apparent when these results were compared with literaturedata for eggs from North Sea parents. On average, plaice eggsfrom the Irish Sea developed more rapidly and hatched up to2 days earlier compared with North Sea eggs incubated at thesame temperature. In terms of parental effects, larger femalestended to produce larger eggs but egg size decreased over successivebatches from the same female within a year. Temperature-dependentegg development rates were influenced by egg size in a non-linearmanner for eggs incubated at 6° C, but egg size had no effectat higher temperatures. Since most plaice eggs sampled fromthe Irish Sea during ichthyoplankton surveys in 1995 and 2000were found in waters cooler than 8° C, such effects maymodify egg development rates in the wild.