The food and growth of 0-group flatfish on nursery grounds in the Clyde Sea Area

In early summer plaice (Pleuronectes platessa L.) in the Clyde Sea Area eat Tellina siphons and spionid palps. As they grow, they take progressively larger siphons, fragments of spionids and some crustaceans. In late summer they eat entire spionids and other sedentary polychaetes. There is no significant predation on fish. The common dab [Limanda limanda (L.)] took a wider range of prey species than the plaice, but also depended mainly on polychaetes, crustaceans and bivalves. Depth distribution may limit both intra- and interspecific competition between these fish and also influence the choice of prey. The results from the main nursery areas of Irvine and Ayr Bays were similar to those from the other bays throughout the Clyde Sea Area and support earlier work at Firemore Bay in north-west Scotland and in the Dutch Wadden Sea.The growth rate of the fish increases during the early summer and may be related to increasing food availability, feeding experience of the fish and longer day length. Growth rate between areas appeared to be correlated with benthic productivity. 0-group plaice in Irvine, Ayr and Firemore Bays grew to different mean lengths by the end of the summer growth season, being 82 mm, 73 mm and 61 mm, respectively, for the period 1972–1974. This agrees with the known differences in standing stocks of intertidal macrobenthos in these bays (17·0, 11·1 and 1·3 g m^?2 dry weight, respectively). The significance of this in relation to survival is discussed. Offshore migration of the larger fish in the autumn may result in apparent slowing of growth rates prematurely in inshore areas. The year-class strength depends more on the food availability in the nursery areas than on the number of plaice that settle on the bottom.     

Genetic structure of juvenile plaice Pleuronectes platessa on nursery grounds within the Irish Sea

As a preliminary investigation into the genetic structure of Irish Sea plaice we genotyped samples of juvenile plaice from six inshore areas within the Irish Sea across eight microsatellite loci and compared them with fish from two sites from the Dutch Wadden Sea (North Sea stocks). Genetic variability in all samples was generally low for that typically observed at microsatellite loci. The number of alleles per locus varied between two and nine (average 5.5) and the observed heterozygosity ranged from 0.080 up to a maximum of 0.909 (average 0.382). Few significant heterozygote deficits were observed, even when the data set was pooled. The majority (98%) of genetic variation present was within, rather than between, populations. None of the pairwise comparisons of population differentiation (F_ST) were significant (P>0.05) and a Bayesian analysis of population structure provided no evidence for a partitioning of the samples. Since juveniles arriving at nursery grounds in the Irish Sea are not distinct, it is likely that adult plaice form a single stock (perhaps with some weak differentiation). However, if plaice eggs and larvae do not disperse as predicted by a particle tracking model, then it is possible that the juveniles represent a mixture of several distinct stocks. Further work is therefore required to determine whether the phenotypic variation observed between female plaice from the eastern and western Irish Sea is the product of reproductive isolation or of the environment.     

Growth changes in plaice,cod, haddock and saithe in the North Sea: a comparison of (post-)medieval and present-day growth rates based on otolith measurements

Fishing effort has strongly increased in the North Sea since the mid-19th century, causing a substantial reduction in the population size of exploited fish stocks. As fisheries research has developed simultaneously with the industrialisation of the fisheries, our knowledge of population dynamics at low levels of exploitations is limited. Otoliths retrieved from archaeological excavations offer a unique opportunity to study growth rates in the past. This study compares historical and present-day growth rates for four commercially important demersal fish species. A total of 2532 modern otoliths (AD 1984–1999) and 1286 historical otoliths (AD 1200–1925) obtained from archaeological excavations in Belgium and Scotland were analysed. Comparison of the growth patterns between eras revealed a major increase in growth rate of haddock, whereas growth changes were not observed in saithe and only in the smaller size classes of plaice and cod. Comparison of our results with literature data indicates that the observed growth rate changes in plaice and cod occurred within the 20th century. Apparently the onset of industrialised fisheries has not greatly affected the growth of plaice, cod and saithe populations in the North Sea. This result contradicts the expectation of density-dependent limitation of growth during the era of pre-industrialised fishing, but is in agreement with the concentration hypothesis of Beverton (Neth. J. Sea Res. 34 (1995) 1) stating that species which concentrate spatially into nursery grounds during their early life-history may ‘saturate’ the carrying capacity of the juvenile habitat even though the adult part of the population is not limited by the adult habitat.     

Episodic Mass Invasions of Juvenile Gadoids into the Wadden Sea and their Consequences for the Population Dynamics of Brown Shrimp (Crangon crangon)

Abstract. The brown shrimp Crangon crungon is considered to be a key species in the coastal waters of the North Sea and in particular in the Wadden Sea, since it occurs in masses and acts as both a highly efficient predator and important prey. The Wadden Sea is the nursery ground for brown shrimp. In 1990, a mass invasion of O-group whiting ( Merlangius merlangus ) reduced Crangon numbers to almost zero. Similar events, which had previously been reported in the 19th century, were observed in 1959 (whiting), 1970 (cod, Gadus morhua ), and 1983 (cod and whiting).
The 1990 invasion and the decline of the shrimp stock are documented on the basis of surveys that are compared with the years preceding and following. Consequences for the food web in the Wadden Sea and the reasons for the complete recovery of the shrimp stock in each case within one year after its collapse are discussed with regard to the Crangon crungon behaviour and reproductive potential. Sampling error and the impact of fisheries are also considered.     

Endoparasites in common eiders Somateria mollissima from birds killed by an oil spill in the northern Wadden Sea

Mass mortalities of common eiders Somateria mollissima have been ascribed to high parasite loads. However, the actual role of parasites in mortalities is disputed as in the case of a mass mortality of eiders in the Wadden Sea in the winter of 1999/2000. A critical evaluation of the role of parasites in eider mass mortalities is hampered by (1) a lack of data on actual parasite loads of the birds involved, (2) missing regional data for comparison, and (3) a lack of unbiased samples: investigations are often based on dead beached individuals, which are presumably the more heavily infected birds of a population and thus more likely to die and be washed ashore. Although published data on parasite loads in birds of the winter 1999/2000 mortality are available, no data on background parasitism in eiders from the Wadden Sea exist, making an evaluation of the potential role of parasites in this mortality event difficult. By investigating endoparasites of 102 eiders affected by an oil spill in the northern Wadden Sea in winter 1998/1999, we provide a data set of background parasitism in wintering eiders from the Wadden Sea. We found 13 different parasite taxa with high prevalence values (% infected birds) in the acanthocephalan Profilicollis botulus, the nematode Amidostomum acutum, cestodes and trematodes. In some taxa we observed pronounced differences in prevalence values between juvenile eiders and adults, as well as between adult sexes. The parasite composition shows that bivalves, crabs (Carcinus maenas) and other crustaceans are important sources of infections by being intermediate hosts. This is partly mirrored in the food content of eider stomachs where bivalves and crabs were predominantly found. Intensities of the acanthocephalan P. botulus, suspected of causing eider mortalities, were especially high in juveniles (1112 ± 416 ind per infected host), but lower in adult males (40 ± 7) and adult females (81 ± 18). However, no extraordinary mortality event was observed in the winter of 1998/1999, indicating no or a very weak effect of the parasites on host condition. A comparison with the parasite loads of eiders from the mass mortality in the winter of 1999/2000 shows that parasite numbers were by no means exceptional for birds from the area. Hence, parasites alone are unlikely to have caused this mortality. Regional background parasite loads are important to differentiate between primary and secondary roles of parasites in anomalous mortality events.     

Distinctly variable mudscapes: Distribution gradients of intertidal macrofauna across the Dutch Wadden Sea

The Wadden Sea is a shallow coastal region, with a large area of sedimentary tidal flats that extends from The Netherlands to Denmark and has been declared a site of international importance in the Dutch and German parts (Ramsar status and UNESCO World Heritage Site). Benthic macrofauna are central to the ecosystem functioning of this area, as they recycle nutrients, decompose organic matter and are an important food source for many secondary consumers, like fish and waterbirds. Due to the environmental gradients characteristic of estuarine systems, it is expected that changes in assemblage composition will be observed across the physical and environmental gradients of the Wadden Sea. First, we explored the spatial variation in assemblage composition of benthic macrofauna across the intertidal part of the Dutch Wadden Sea using 3 years of biomass data. Then, we identified the relative importance of six environmental variables for explaining and predicting changes in assemblage composition across the intertidal areas of the Wadden Sea using generalised dissimilarity modelling (GDM). In accordance with the environmental gradients across this system, the biomass contributed by a few common species differed from west to east and were distinct in the Dollard. In the west, bivalves Mya arenaria, Cerastoderma edule and Ensis directus contributed a relatively large and equal share of the total biomass, whereas C. edule contributed the sole largest share of the total biomass towards the east. The polychaete Alitta succinea became a large share of the total biomass in the upper Ems and in the Dollard estuary, but contributed little elsewhere. Similar to the observed differences in species composition, the spatial patterns in assemblage composition, as predicted by the GDM models, identified the Dollard as distinct and that the prevalence of assemblage types in the west differed to the east. Median grain size, followed by microphytobenthic biomass, and exposure time were the most important variables describing differences in assemblage composition. That the Wadden Sea forms a heterogeneous landscape where assemblage composition varies across multiple gradients has repercussions for management and monitoring.     

Changes in potential North Sea spawning grounds of plaice (Pleuronectes platessa L.) based on early life stage connectivity to nursery habitats

We explored the hypothesis that spawning ground locations of North Sea plaice reflect the locations of nursery grounds using drift scenarios based on a baroclinic, shallow-water circulation model (HAMSOM). The transport of pelagic eggs and larvae was simulated each year from 1975 to 2006 using in situ forcing, temperature-dependent development and stage-specific behaviour of eggs and larvae. This long-term simulation period also allowed us to explore climate effects. A release position was considered a potential and suitable spawning site if larvae from that area reached coastal nurseries after the onset of metamorphosis. In general, larvae were transported in an anti-clockwise direction and settled in nurseries that were relatively close to the release positions. Spawning locations that were offshore were poorly connected to nursery grounds while those closer to the shore had higher connectivity. Simulated suitable spawning locations broadly agreed with the main centres of egg production (English Channel, Southern Bight, German Bight), except for the known spawning grounds south of Dogger Bank. Over the 31-year simulation period, positive and negative trends in transport success were found for the western and eastern parts of the North Sea, respectively. Changes in the west (Flamborough Head) were mainly due to changes in water circulation patterns whereas those in the east (northern German Bight) were induced by changes in both currents and water temperature. The implications of these findings, and the significant correlation between changes in drift and recruitment, suggest that climate-driven changes in the suitability of nursery grounds will directly affect the distribution and productivity of plaice in the North Sea.     

Sources and fate of manganese in a tidal basin of the German Wadden Sea

Dissolved and particulate Mn concentrations were investigated on a seasonal scale in surface waters of the NW German Wadden Sea (Spiekeroog Island) in 2002 and 2003. As the Wadden Sea forms the transition zone between the terrestrial and marine realms, Mn was analysed in coastal freshwater tributaries and in the adjoining German Bight as well. Additionally, sediments and porewaters of the tidal flat sediments were analysed for Mn partitioning and microbial activity.Dissolved Mn concentrations show strong tidal and seasonal variation with elevated concentrations during summer at low tide. Summer values in the Wadden Sea (av. 0.7 μM) are distinctly higher than in the central areas of the German Bight (av. 0.02 μM), suggesting a possible impact of the Wadden Sea environment on the Mn budget of the North Sea. Seasonality is also observed for particulate Mn in the Wadden Sea (winter av. 800 mg kg^− 1; summer av. 1360 mg kg^− 1). Although particles are relatively Mn-poor during winter, the high SPM load during this season causes elevated excess concentrations of particulate Mn, which in part exceed those of the dissolved phase. Therefore, winter values cannot be ignored in balance calculations for the Wadden Sea system.Porewater Mn concentrations differ depending on sediment type and season. Maximum concentrations are found in surface sediments at a mixed flat site (190 μM) during summer, while winter values are distinctly lower. This indicates that enhanced microbial activity owing to higher temperature during summer leads to increased reduction of Mn-oxides in surface sediments and enhances the corresponding diffusive and advective Mn flux across the sediment-water interface. Draining of Mn-rich porewaters from sediments is also documented by analyses of tidal creek waters, which are highly enriched in Mn during summer.Furthermore, an important Mn source is freshwater discharged into the Wadden Sea via a flood-gate. The concentration of dissolved Mn in freshwater was highly variable during the sampling campaigns in 2002 and 2003, averaging 4 μM. In contrast, particulate Mn displayed a seasonal behaviour with increasing contents during summer. On the basis of salinity variations in the Wadden Sea, the total amount of Mn contributed to the Wadden Sea via freshwater was estimated. This balance shows the importance of the freshwater environment for the Mn inventory of the Wadden Sea. During winter the total Mn inventory of the Wadden Sea water column may be explained almost completely by freshwater discharge, whereas in summer the porewater system forms the dominating source.     

Seasonal patterns in the fish and epibenthic crustaceans community of an intertidal zone with particular reference to the population dynamics of plaice and brown shrimp

The intertidal zone of a sandy beach located on the French coast of the Eastern Channel, was sampled during spring and summer 2000 to analyse the community structure of fish and epibenthic crustaceans. The presence of many juvenile fish (mainly O-group) and crustaceans indicated the important role played by the intertidal zone as a nursery ground. The brown shrimp, Crangon crangon and O-group plaice, Pleuronectes platessa are the two most abundant species of the intertidal ecosystem.Plaice settlement period extended from mid-March to early June. Over the survey period, densities increased to maximum numbers of about 27 ind. 10 m⁻² in mid-April. In the following week, density rapidly decreased due to mortality and migration into deeper waters. The mortality was attributed mainly to predation by brown shrimp (C. crangon) and to a lesser extent by the shore crab (Carcinus maenas). The mean size of 0-group plaice increased from 19 mm in mid-April to 58 mm in July. Growth of juvenile 0-group plaice is described by an exponential equation: total length (TL, mm) = 12.602 e^{0.022 (post-settlement age)}. Post-settlement growth rates, estimated by otolith microstructure analysis, were 0.38 mm d⁻¹ for plaice ≤30 mm and 0.61 mm d⁻¹ for plaice >30 mm. Settlement of juvenile brown shrimp started in mid-April, peaked in early June (93 ind. 10 m⁻²) and continued with fluctuating intensity throughout the summer. Growth rate of juvenile C. crangon, estimated after the settlement peak, was 0.163 mm d⁻¹. Growth conditions of juvenile plaice and C. crangon were analysed by comparing estimated growth in the field with predicted maximum growth according to temperature-growth rate models from experimental studies of growth with unlimited food supply. For plaice, the estimated growth rate was lower (plaice ≤30 mm) but similar (plaice >30 mm) to the predicted maximum growth suggesting a food limitation only for newly settled individual. The observed increase in mean length of juvenile C. crangon was lower than the maximum possible growth. The function of the intertidal zone in the early juvenile stages of marine species is discussed.     

Diel feeding of the transparent goby Aphia minuta (Pisces,Gobiidae) in the Northwestern Adriatic Sea in spring time

The present study provides the first estimate of the daily ration for a goby species in the Mediterranean Sea, using a new approach to determine the mass of fish stomach contents through the sum of individual prey dry mass derived indirectly from prey size. Diel feeding activity and daily ration of the pelagic goby Aphia minuta were studied under natural conditions in the coastal waters off Comacchio (Northwestern Adriatic Sea) during the spring season. A total of 318 individuals of A. minuta, collected by 14 trawls carried out on 4 consecutive days, was examined for gut contents. The diel pattern of the vacuity index and the stomach fullness index indicated that A. minuta has nocturnal feeding behaviour. The daily ration, computed over a period of 16 h, was equivalent to 2.23% wet body mass.     

Mussel fishery affects diet and reduces body condition of Eiders Somateria mollissima in the Wadden Sea

Although the Danish Wadden Sea is of international importance for several bird species, large-scale blue mussel Mytilus edulis fishing took place from 1984–1987, ceasing thereafter due to low mussel stocks. Mussel fishing removes much of the blue mussel biomass, especially larger individuals. Hence we predict that intensive mussel fishing will affect their predators, such as the Eider Somateria mollissima, which is predominantly a blue mussel feeder by, 1) reducing the amount of blue mussels in their diet relative to alternative prey items, 2) exploitation of smaller blue mussel shell classes, 3) loss of body condition, 4) changing feeding distribution to aggregate to the remaining mussel stocks, and 5) decreasing numbers. Before winter 1986/87 blue mussel biomass was estimated at 40,600 tons, decreasing to 15,400 tons in 1987/88 due to mussel fishery. We collected Eiders in both periods to monitor their diet and body mass and used aerial surveys to determine changes in numbers and distribution. Between the two periods, blue mussels declined in the Eiders diet, numbers of Eiders with empty stomachs increased and the mean length of blue mussel taken by Eiders decreased. Eider body condition declined from 1986/87 to 1987/88, mostly the result of the reduction in numbers of individuals with blue mussel remains in their gizzards and in better body condition compared to those taking alternative food items or having empty gizzards. Eiders shifted their distribution from the southern part of the Danish Wadden Sea to the northern part, where the remaining blue mussel stocks were situated. Eider numbers were lowest in 1987/88, the year of lowest blue mussel stocks. We conclude that intensive mussel fishery affected the Eider's diet, reduced their body condition and affected distribution and abundance. The results also showed that availability of blue mussels may have a key role in building up and maintaining body condition in Eiders during winter.     

Effects of current exposure on habitat preference of mobile 0-group epibenthos for intertidal seagrass beds (Zostera noltii) in the northern Wadden Sea

In the northern Wadden Sea, the extent of intertidal seagrass beds, their plant biomass and shoot density highly depends on local current regimes. This study deals with the role of intertidal Zostera noltii beds as nursery for mobile epibenthic macrofauna and the impact of seagrass bed characteristics on their abundance and distribution patterns. According to their exposure to the main tidal gullies, sampling sites were separated into exposed, semi-exposed and sheltered. Dominant species of crustaceans and demersal fish were studied in respect of their abundances within seagrass beds and adjacent unvegetated areas. Quantitative sampling was performed at day and night high tide using a portable drop trap. In general, species composition varied little between seagrass beds and bare sand. However, the presence of vegetation had a quantitative effect increasing individual numbers of common epifaunal species. Abundances of 0-group shore crabs (Carcinus maenas), common gobies (Pomatoschistus microps) and brown shrimps (Crangon crangon) were highest within sheltered seagrass beds. With decreasing plant density habitat preference of epibenthos changed on species level. By regulating the habitat complexity the currents regime is profoundly influencing the nursery function of intertidal seagrass beds in the Wadden Sea.     

Dynamic interpretation of grain size relations in silt

In the salt marsh deposits of the Danish Wadden Sea, a separation has successfully been made of two log-normally distributed grain size populations, which make up the major part of these fine-grained sediments. The coarse population consists almost exclusively of silt and is interpreted as material either deposited as resistant flocs or as single particles. The fine population consists of approximately 50% silt and 50% clay and is interpreted as particles settled as easily destroyable flocs. These interpretations are confirned by the fact that the relative abundance of the two populations is demonstrated to be related to the degree of exposure of salt marsh areas.     

Does small-scale vertical distribution of juvenile schooling fish affect prey availability to surface-feeding seabirds in the Wadden Sea?

Food availability is a key variable influencing breeding performance and demography of marine top predators. Due to methodological problems, proportionality between fish abundance and availability is often assumed without being explicitly tested. More specifically, better breeding performance of surface-feeding seabirds at times of large prey stocks suggests that prey availability is also a function of prey abundance. Using vertically resolved stow net sampling we tested whether local abundance and length composition of pelagic fish are reliable predictors of the availability of these fish to surface-feeding Common Terns (Sterna hirundo) breeding in the German Wadden Sea. Prey fish were found to concentrate below the maximum diving depth of the terns. Individuals caught close to the surface were in most cases smaller than conspecifics caught at greater depth. Correlations between fish abundance within and out of reach of the terns appeared to be both species- and site-specific rather than driven by overall fish abundance. Vertical distribution patterns of the terns' main prey fish could be explained as anti-predator behavior, reducing prey availability to the terns. In 2007, when breeding performance was much better than in 2006, herring and whiting were much more abundant, suggesting that overall prey abundance may also increase prey availability in habitats other than those represented by the stow net sampling.     

Intra- and interspecies comparison of energy flow in North Atlantic flatfish species by means of dynamic energy budgets

The quantitative aspects of growth and reproduction in four flatfish species (plaice, flounder, dab, sole) in terms of energy flow are described on the basis of a dynamic energy budget (DEB theory). This theory consists of general assumptions about energy uptake, storage and utilisation and describes an individual by two state variables: structure and reserve, whereby body size exerts its influence through the ratio between surface area and volume. Comparison between model estimates and field data shows that the DEB model successfully describes the energetics of growth and reproduction in a number of flatfish species. Differences between species could be captured in the same model using different parameter values. Intraspecific differences in growth between males and females are mainly caused by differences in maximum surface area-specific ingestion rates. Differences between species are reflected in the surface area-specific maximum ingestion rate, the energy partitioning over growth and reproduction, and in egg volume. According to these parameters at 283 K (10°C), the species could be ranked as follows: surface area-specific maximum ingestion rate (W m⁻²) plaice: 56.6; flounder: 54.5; sole: 45.1 and dab: 36.1 W m⁻². Fraction of energy allocated to reproduction (–): flounder: 0.35; plaice: 0.15; dab: 0.15 and sole 0.10. As a consequence of these differences in surface area-specific maximum ingestion rate and in the fraction of utilised energy allocated to reproduction, the gonad masses (g) of females of 0.5 kg wet mass differ considerably: flounder: 149 g; plaice: 86 g; sole: 70 g; and dab: 69 g. However, due to differences in egg size between species, the potential annual egg production shows a completely different pattern: dab: 2200 10³; flounder: 1560 10³; sole: 343 10³ and plaice: 130 10³ eggs.     

Potential transport of plaice eggs and larvae between two apparently self-contained populations in the Irish Sea

A coupled physics particle-tracking model, driven by realistic meteorological forcing was used to examine the dispersal and transport of plaice eggs and larvae in the year 2000 from two spawning grounds in the Irish Sea. The model included passive transport of eggs and early stage larvae, diel vertical movements for larvae between 7 and 10.5 mm in body-length and tidally synchronised, vertical movements for larger larvae (>9 mm body-length). The year 2000 was chosen because of the availability of ichthyoplankton data with which to initialise the model. The majority of larvae originating from spawning in the eastern Irish Sea settled into nursery grounds along the Scottish, English and Welsh coasts, in agreement with previous findings. In contrast, a significant portion of larvae originating from spawning in the western Irish Sea was transported eastwards to these same nursery grounds. Transport across the Irish Sea resulted from the onset of tidally synchronised vertical behaviour encoded in the model for older larvae. Settlement of larvae into local nursery grounds along the Irish coast was limited. Because of the prevailing winds and currents in the region, plaice eggs and larvae are unlikely to be transported from east to west; in most years spawning in the western Irish Sea probably acts as an additional source of juveniles for nursery grounds along the Scottish, English and Welsh coasts.     

Estimated intensity needed for sampling flatfish assemblages in reference areas of tidal mud flats systems may be disproportionately costly and deleterious

Beam trawl surveys were carried out in a tidal stream system of the German Wadden Sea on two to three successive days of comparable weather conditions in three summers. The variability in time and space detected for plaice (Pleuronectes platessa), flounder (Platichthys flesus), dab (Limanda limanda), sole (Solea solea) and lemon sole (Microstomus kitt) was considerable even for species with a very high abundance. Other flatfish species such as turbot (Scophthalmus maximus) and brill (Scophthalmus rhombus) occurred only occasionally in the catches. Sampling effort calculated to detect a 50 or 20% difference between the surveys caused by immigration, emigration or mortality appears to be disproportionate. It is argued that such a major sampling effort of reference areas in tidal mud flat systems is likely to influence the results, to harm the environment, and to be very expensive.     

Differential reproductive strategies of two bivalves in the Dutch Wadden Sea

Cerastoderma edule and Mya arenaria are two common bivalve species in European waters. Longevity and maximum size are much greater in the latter species. Because comparison of species life-history strategies states that a long life span (i.e. high annual survival) generally goes with lower fecundity, we hypothesise that reproductive output would be lower in M. arenaria than in C. edule. In the present paper, we studied the reproductive strategies of these two species in an intertidal and a subtidal area of the western Dutch Wadden Sea, by following seasonal changes in absolute and relative weights of somatic and gonadal tissues in these bivalves. Starting of spawning was similar in the two species, around May, except for intertidal M. arenaria, which initiated spawning in August. Individual energy investment in reproduction was similar for the two species but, unlike M. arenaria, C. edule spawned completely, releasing all energy of gonadal mass in the form of gametes. Mya arenaria used the gonad not only for reproduction but also for storage. In the intertidal area, we found a trade-off between longevity and reproduction, i.e. maximum reproductive output (expressed as a proportion of body mass) was higher in C. edule than in M. arenaria. However, since body size is larger and life span longer in M. arenaria than in C. edule, mean lifetime reproductive output per individual must be higher in the first than in the latter. Based on the differences in reproductive strategies of these two species, we hypothesise that the negative effects of warming climate on bivalve population dynamics in the Wadden Sea will be stronger in C. edule than in M. arenaria.     

Spatial variability in growth and reproduction of the Pacific oyster Crassostrea gigas (Thunberg, 1793) along the west European coast

The Pacific oyster Crassostrea gigas was introduced in Europe for commercial purposes in the mid 1960s. It was initially thought that low winter temperatures would restrain this species' reproduction and settlement; however, its present distribution in areas where no introduction has taken place suggests that natural invasion and expansion has occurred. Along the European coast, wild populations of Pacific oysters are already found from northern Germany to southern Portugal. Whether C. gigas will continue to further expand through northern waters will depend on its physiological performance. In this study, the performance of wild oyster populations has been studied in terms of growth and reproduction at three stations: La Rochelle (France; 46°N), Yerseke (Oosterschelde estuary, The Netherlands, 51°N), and Texel (Wadden Sea estuary, The Netherlands, 53°N). The French population had the lowest somatic-shell mass ratio and an increase in maximum shell length, somatic and gonadal mass was observed from France to the Netherlands. In addition, mean oocyte diameter decreased significantly from south to north. The combination of increasing gonadal mass and decreasing oocyte volume suggests an increasing reproductive output in terms of egg numbers from France to The Netherlands. Differences in temperature between locations will at least be partly responsible for the observed patterns; however, other environmental factors (such as food availability, predation pressure, sediment type and/or seston concentration) cannot be excluded. Since smaller eggs (oocytes) are thought to have a longer development time, the environmental conditions along the Dutch coast may result in increased larval dispersal and possibly in further population expansion.