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.     

Reconstruction of food conditions for Northeast Atlantic bivalve species based on Dynamic Energy Budgets

Required assimilated energy to support observed growth was reconstructed for four common bivalve species (Mya arenaria, Cerastoderma edule, Mytilus edulis and Macoma balthica) from various Northeast Atlantic coastal areas, along the species distributional range. The approach applied was based on the Dynamic Energy Budget (DEB) theory whereby observed growth patterns in the field, in combination with prevailing temperatures, were used to reconstruct the average food intake experienced in the field scaled to the maximum possible. For all species, results suggest food limitation over the range of locations. In general, reconstructed food intake indicated better conditions for C. edule compared to the other species, while M. edulis presented the lowest food conditions in all the areas. Despite the indications for a latitudinal trend in primary production, no clear pattern or relationship between reconstructed food conditions and latitude was observed suggesting that any trend may be overruled by local conditions.     

Eiders Somateria mollissima scavenging behind a lugworm boat

The eider is one of the most important molluscivorous birds in the Wadden Sea, where it feeds mainly on blue mussels Mytilus edulis and edible cockles Cerastoderma edule. These prey species are within reach of the birds at all times. Other potential prey of suitable size that are abundantly present, such as several polychaete worms, or the clam Mya arenaria, are taken to a much lesser extent, possibly because they live buried in the sediment and digging them out would take too much effort. Mya may pose another problem because they grow to sizes that prevent eiders from swallowing them. Large Mya also live too deep down in the sediment, but young (small) specimens should be available to eiders. Yet, even these have only rarely been found as prey in eiders in the Wadden Sea. However, diet studies in relation to food abundance have been few, and may have missed prey that do not leave large shell fragments (i.e. in faeces studies). This paper describes observations on eiders taking both Mya and polychaete worms. The eiders fed on these prey in a fashion reminiscent of gulls that scavenge behind fishing vessels: some eiders have learnt to follow professional worm-digging boats that supply a bycatch of molluscs (mainly Mya arenaria) and polychaete worms (mainly Arenicola marina and Nephtys hombergii). Mya and worms were also the main targets of the eiders that fed in a dense flock close to the boat's stern. Faeces found on the flats at low tide comprised mainly cockle shell fragments, a prey rarely taken by the eiders behind the boat. Faeces studies may thus give a highly biased impression of local eider diet.     

Seasonal variability in somatic and reproductive investment of the bivalve Scrobicularia plana (da Costa, 1778) along a latitudinal gradient

Monthly investment in soma and gonads in the bivalve Scrobicularia plana is described for three populations along its distributional range: Minho estuary, Portugal; Westerschelde estuary, The Netherlands and Buvika estuary, Norway. Seasonal cycles in body mass (BMI), somatic mass (SMI) and gonadal mass (GMI) indices were observed for all populations. In Portugal, BMI and SMI peaked in mid-autumn, while in The Netherlands both indices were at their highest in mid-spring. Norway showed a different pattern with two distinct peaks: one in mid-autumn and a second peak in spring. GMI reached maximum values in July in Portugal and Netherlands and in June in Norway. Overall, mean BMI and SMI were lower in Portugal while mean GMI was lower in Norway. The spawning period lasted the whole summer in Portugal, but was shorter (only two months) in The Netherlands and Norway. The reproductive investment in The Netherlands was significantly higher than in Portugal and Norway, with the lowest values being observed in Norway. Differences in annual cycles between populations were attributed to environmental factors, namely temperature and food availability. Temperature seems important in shaping the reproductive pattern with more northern populations showing shorter reproductive periods starting later in the year, and a lower reproductive output. In addition, winter water temperatures can explain the lower mean body and somatic mass values observed in Portugal. Food availability influenced the physiological performance of the species with peaks in somatic mass coinciding with phytoplankton blooms. This relation between physiological performance and environmental factors influences S. plana distribution, densities and even survival, with natural consequences on its commercial importance.     

Changes in distribution patterns of 0-group bivalves in the Wadden Sea: Byssus-drifting releases juveniles from the constraints of hydrography

The large-scale (km) distributional patterns of juvenile bivalves are established by larval settlement and subsequently changed due to actively initiated postlarval migrations (byssus-drifting), resuspension during sediment disturbance, and local differences in mortality. Repeated mapping of 0-group bivalve distribution during two summers was combined with simultaneous registrations of the numbers of drifting specimens. Species differed in their susceptibility to passive resuspension and their activity in byssus-drifting, but the two ways of entry into the water column were independent of each other. As a result, the relative magnitude of byssus-drifting and passive resuspension varied with the species. While hydrographical conditions always determined the changes in the distributional patterns of passively eroded species, this was not the case in active migrations of byssus-drifting bivalves. A comparison of the distributional patterns of juveniles over two consecutive years showed similar patterns in the tellinid clam Macoma balthica and the razor clam Ensis americanus. In these two species spatfall mainly occurred around mean low-tide level. Subsequently, juvenile M. balthica rapidly accumulated in the upper intertidal, whereas juvenile E. americanus accumulated subtidally. In mussels Mytilus edulis the distribution of juveniles only changed in a longshore direction, not in the tidal level occupied. The spatial pattern of cockles Cerastoderma edule changed from aggregation of the early spat in the mid and lower intertidal towards a more uniform distribution of recruits over these tidal flats. There were only a few patches of high abundance left at the end of summer. Finally, in juvenile clams Mya arenaria the spatial patterns of both spatfall and redistribution were unpredictable.On a km-scale, abundance of some species correlated with sediment granulometry. Presumably, this does not reflect a causal relationship but is a consequence of the correlation between grain size distribution and tidal level within the area studied. On a scale of hundreds of metres, there were no consistent correlations between juvenile abundance and sediment granulometry in any species. In summary, it seems that hydrography ruled the initial settlement of larvae to the sediment and strongly influenced the subsequent redistribution of juveniles caused by passive resuspension. The outcome of actively initiated migrations, on the other hand, was only weakly influenced by hydrography. Therefore it is suggested that hydrography plays a dominant role in the initial development of km-scale distributional patterns of just-settled bivalves in the Wadden Sea, while habitat selection is delayed to the byssus-drifting postlarvae phase.     

Erodibility of a mixed mudflat dominated by microphytobenthos and Cerastoderma edule, East Frisian Wadden Sea,Germany

Sediment erodibility and a range of physical and biological parameters were measured at an intertidal site in the German Wadden Sea area in June, September and November 2002 and February and April 2003 in order to examine the influence of macrozoobenthos and microphytobenthos on sediment erodibility and the temporal variation. The study site was a mixed mudflat situated in the mesotidal Baltrum–Langeoog tidal basin at the East Frisian barrier coast. The mud content at the site was about 35% and the filter-feeding cockle Cerastoderma edule was the dominating macrozoobenthic species (by biomass). The erodibility of the sediment showed strong temporal variation with high erosion thresholds in spring and late summer and significantly lower thresholds during the rest of the study period. The erosion thresholds were strongly dependent on the contents of chlorophyll a (chl a) and colloidal carbohydrates, both indicators of the content of microphytobenthos, in this environment primarily benthic diatoms. The content of microphytobenthos was high in September 2002 and April 2003, and regression analysis indicated that this was the only likely reason for the low erodibility found at these times. A biostabilisation index of about 4.5 was found for a situation with both abundant biofilms and cockles.     

Exploring interactions among intertidal macrozoobenthos of the Dutch Wadden Sea using population growth models

Using data taken from three long-term monitoring programs, we modelled the population dynamics of 13 common macrozoobenthic species of the Balgzand, an approximately 50 km² intertidal area in the Dutch part of the Wadden Sea. In order to identify likely interactions among species, while accounting as much as possible for other factors influencing population dynamics, models included both ‘environmental’ (water temperature, and phytoplankton concentration) and ‘biological’ (biomass of potentially interacting species) variables. This approach appeared to be effective at identifying certain types of interactions acting over relatively short time-frames, such as predation and competition for food. Among the species we considered, the strongest effects of this kind appeared to be from what we presume to be predation by Nephtys hombergii, and from competition for food among bivalves. Several of the strongest results of our analysis corroborated expectations derived from previous small-scale surveys or experimental studies, but there were also a number of highly significant results indicating possible interactions which were not immediately explicable, including the apparent positive effect of Macoma balthica on Arenicola marina, and the apparent negative effect of Nephtys hombergii on Mya arenaria. Our results also suggest that year to year changes in Cerastoderma edule and Arenicola marina populations did not have substantial and widespread influences on the population dynamics of the majority of other common infauna, which is not to deny the likely critical importance of recruitment effects and biogenic transformation of the intertidal environment by these species over longer time frames. Our modelling approach could be applied to other long-term data sets, and will become more useful as longer time series become available.     

The role of ecological divergence in speciation between intertidal and subtidal Scoloplos armiger (Polychaeta,Orbiniidae)

The concept of ecological speciation implies that habitat differences may split a species by strong selection and rapid adaptation even under sympatric conditions. Studies on the polychaete Scoloplos armiger in the Wadden Sea (North Sea) indicate sibling species existing in sympatry: the intertidal ‘Type I’ with holobenthic development out of egg cocoons and the subtidal ‘Type S’ producing pelagic larvae. In the current study, Types I and S are compared in habitat-related traits of reproductive timing and physiological response to hypoxia and sulphide. Spawnings of Type I and Type S recorded over six years overlap in spring and both appear to be triggered by a rise in seawater temperature above 5 °C. Type S exhibits an additional autumn spawning (at seawater temperatures around 10 °C) which was previously unknown and is absent in Type I. The overall abundance of pelagic larvae in the Wadden Sea is higher in spring than in autumn. Tolerance of both sulphide and hypoxia was lower in Type S than in Type I. This correlates with a 5 to 10-fold lower sulphide concentration in the subtidal compared to the intertidal habitat. Physiological tolerance and divergence in developmental mode appear as traits which may have led to reproductive isolation between Type I and Type S. Their role in allopatric and sympatric speciation scenarios in S. armiger is discussed. Since the pelagic dispersal mode has been neglected so far, a reassessment of population dynamics models for S. armiger is suggested.     

Dense aggregations of Pygospio elegans (Claparède): effect on macrofaunal community structure and sediments

Epibenthic biogenic structures such as polychaete tubes are conspicuous features of many marine soft-bottom habitats. This paper compares the benthic macrofauna in patches with high and low densities of the tube-dweller Pygospio elegans on intertidal sandflats in eastern Scotland (UK). The main aim of this study was to determine potential differences in the macrofaunal community structure, the size distribution of individual species and sediment properties.Multivariate data analyses revealed that the macrofaunal community composition (excluding P. elegans) within patches was always significantly different from outside patches, mainly due to variability in the abundances of Cerastoderma edule and Corophium volutator. In addition to P. elegans, 5 taxa were sufficiently abundant for univariate analyses, 4 of these (Capitella capitata, C. edule, Macoma balthica and C. volutator) being significantly more abundant within P. elegans patches than in surrounding, non-patch sediments. The size distribution of P. elegans was significantly different between patches (bimodal distribution) and non-patches (skewed distribution). Similarly, there was a greater proportion of larger C. capitata individuals within patches compared to non-patch sediments. Sediment organic content and silt/clay fraction were always significantly higher in patch sediments while redox profiles showed no differences except at the end of the study period when the top 2 cm within patches were more positive and more negative at 4 cm. These results imply that even relatively small (1–1.5 m²) P. elegans patches can have large effects on the spatial variability of macrofaunal community structure on intertidal sandflats.Towards the end of the study there were marked visual changes in the P. elegans patches, such as wave-ripple marks on the surface, which signified their demise. This coincided with dramatic changes in the invertebrate community structure within patches. Along with the decline in P. elegans numbers, dramatic increases in the densities of the 2 bivalve species C. edule and M. balthica occurred, while remaining comparatively stable outside patches. This suggested that the conditions within P. elegans patches were particularly favourable to successful bivalve spat settlement. We postulate that P. elegans patches have limited longevity and propose that enhanced bivalve competition within them leads to rapid decreases in P. elegans numbers.     

Observations of a mass occurrence of Macoma balthica larvae in midsummer

In 1995 the seasonal development of concentrations of both phytoplankton and larvae of the bivalve Macoma balthica was studied in the coastal zone behind the back-barrier island of Spiekeroog (German Wadden Sea). In July/August larvae reached maximum concentrations of about 1000 to 4200 ind. m⁻³ (depending on the sampling site and tidal period), probably in relation to a phytoplankton bloom in July. This observation of an unusually late maximum of Macoma larvae is discussed in detail in connection with the recent literature available about spawning of M. balthica in the Wadden Sea.     

Reproductive investment of the American razor clam Ensis americanus in the Dutch Wadden Sea

The American razor clam Ensis americanus is a marine bivalve indigenous from the American coast that was introduced in Europe in the late 1970s and has managed to build up strong populations along the European coast. In the present paper, we studied energy investment into soma and gonads by following the seasonality in growth and reproduction of this species in a subtidal area of the western Dutch Wadden Sea. Body and somatic mass indices showed a seasonal pattern with maximal values in August and minimal in April. Gonadal mass index showed the opposite trend with highest values in April. Spawning started in April/May and a second, but weaker, spawning event apparently occurred in August/September. Gonadosomatic ratio values were very low. Just before the spawning period, only 2.5% of the body mass of E. americanus consisted of gonads.     

Population dynamics of the venerid bivalve Callista chione (L.) in a coastal area of the eastern Mediterranean

The venerid bivalve Callista chione is the most prominent suspension-feeding bivalve inhabiting the soft bottom of coastal areas in the northern Euboikos Gulf (Aegean Sea), where it is exploited by small-scale fishery. The parameters of the von Bertalanffy growth function were estimated and the somatic production-to-biomass ratio was found to be 0.45 y⁻¹. The maximum age of adult individuals was 17 years. The species is gonochoristic, its oocyte diameter ranged from 54.5 to 120 μm with a conspicuous jelly-coat. Microscopical examination of the gonads showed that gonadal production did not start until the second year of life. The observations on the gonadal cycle indicate that this bivalve is capable of year-round reproduction.     

Reproductive investment in the intertidal bivalve Macoma balthica

Bivalve eggs generally contain large amounts of lipids which, in comparison with proteins and carbohydrates, have high energy contents and are thus costly in energetic terms. As lipid contents vary between species, comparisons of reproductive investments should not only include numbers and sizes of eggs, but also their energy content. We estimated the investment in egg material of mature females of the Baltic tellin Macoma balthica (L.) in terms of both mass and energy content. All mass below a minimum body mass (below which no eggs are produced) was defined as structural mass. This threshold amounts to a body mass index (BMI) of 5.6 (ash-free dry mass per cubic shell length in mg cm⁻³). More than half (55%) of the mass above the structural mass was invested in egg material and 45% in extra somatic tissue and tissue for production and storage of gametes. This means that the amount of eggs spawned ranged from 0 (at BMI = 5.6 mg cm⁻³) to 33% of the total ash-free dry mass (at a high BMI value of 14 mg cm⁻³). Eggs contained a relatively large amount of lipids, about 30% of their ash-free dry mass, whereas non-egg material contained only about 7% lipids. Eggs of two other bivalves in the Wadden Sea, the cockle Cerastoderma edule and the mussel Mytilus edulis, were smaller and contained only about 11% and 20% lipids, respectively. Energy content of M. balthica eggs amounted to ∼0.006 J, in the other two species to ∼0.002 J. The function of the more expensive eggs in M. balthica may be related to its early spawning in spring, causing slower larval development until first feeding.     

Reproductive biology of Ilisha elongata (Teleostei: Pristigasteridae) in Ariake Sound,Japan: Implications for estuarine fish conservation in Asia

Elongate ilisha (Ilisha enlongata) is a commercially important species that contributes to clupeoid fisheries in Asian countries. In the present study, the reproductive biology of I. elongata in Ariake Sound, Japan is determined for the first time. Six maturity stages were described using ovarian and testicular histology throughout the annual cycle. The spawning season in Ariake Sound lasts from May to July, with peak spawning activity in May and June. Age at first maturity was estimated to be 2 years, with a few exceptions of 1 year in well-developed males. Ovaries that contained both tertiary yolk oocytes and postovulatory follicles occurred from late May to late July, indicating that I. elongata is a multiple spawner. The size–frequency distribution of oocytes provided evidence for its multiple spawning and accuracy of the fecundity estimates. The batch fecundity of this species was estimated at between 22,200 and 270,900 eggs per individual, increasing with age between two and six years. The present findings on the reproductive strategy of I. elongata in Ariake Sound are generally consistent with those in temperate or subtropical populations, but quite different from those of tropical population where first maturation occurs around 200 days and life spans are shorter, with a maximum age less than 3 years. The conservation implications of this reproductive strategy in a harsh, variable environment in Asian countries are also discussed.     

Effect of macrofaunal bioturbation on bacterial distribution in marine sandy sediments, with special reference to sulphur-oxidising bacteria

We have studied the impact of the bioturbating macrofauna, in particular the lugworm Arenicola marina and the bivalve Cerastoderma edule, on abundances and distribution patterns of total bacteria and of bacteria of selected functional groups in sandy intertidal sediments. The selected groups comprised the colourless sulphur-oxidising bacteria and the anoxygenic phototrophic bacteria, which are expected to occupy small zones at the oxygen–sulphide interface in stable (non-bioturbated) sediments. The presence of a wooden wreck buried in the sediment at 10 cm depth within a large area of intertidal sand flat colonised by lugworms provided a unique opportunity to confront field observations with laboratory simulations. The site with the wooden wreck, which was used as control site, was devoid of both A. marina and C. edule, while the composition of the rest of the zoobenthic community was rather similar to that of the surrounding area. In the field, the density of total bacteria was approximately one order of magnitude higher in the control site than in the natural (bioturbated) site. This can be explained by the higher contents of silt and clay particles (higher surface-area/volume ratio) and higher total organic-carbon contents found at the control site. It appears that the presence of macrofauna affects sedimentation processes, which indirectly influence bacterial dynamics. Samples from the control site have been incubated in the laboratory with A. marina and C. edule added (bioturbated core), while an unamended core served as a control. The laboratory experiments contrasted with the field observations, because it was found that total bacteria were actually higher in the deeper layers of the bioturbated core. Moreover, the populations were more homogeneous (less stratified) and colourless sulphur bacteria were on average less numerous in the bioturbated core. In general, laboratory incubations resulted in a decrease of total bacteria with a concomitant increase of colourless and phototrophic sulphur-oxidising bacteria and thus in modifications of the bacterial community structure. Hence, our results demonstrate that care must be taken in extrapolating results from laboratory experiments (e.g. mesocosm research) to field situations.     

Reproduction in the intertidal limpet Cellana ornata in southern New Zealand

The reproductive cycle of the abundant intertidal limpet Cellana ornata (Dillwyn 1817) was examined over 19 months, covering two breeding periods, on a rocky intertidal platform in southern New Zealand. A gonad index was calculated, and histological sections were used to count the numbers of mature and immature eggs present. Fecundity was estimated using counts of eggs before and after spawning. To determine any variation between sites, the gonad index of limpets at a second site dominated by boulders was examined over 6 months during the second breeding period. C. ornata had a single spawning period annually during summer, with greatest gonad sizes in January‐February. There were significant differences in peak gonad indices between two breeding seasons on the platform. Within the second breeding period, both the timing and magnitude of peak gonad development differed between sites, with lower reproductive output at the boulder site. Fecundity increased with increasing size, but individuals are probably capable of reproducing after their first year on the shore. The life history of this species is discussed in comparison to other limpets in New Zealand.     

Variability in the fractionation of stable isotopes during degradation of two intertidal red algae

Macroalgae contribute to intertidal food webs primarily as detritus, with unclear implications for food web studies using stable isotope analysis. We examined differences in the thallus parts of two South African rhodophytes (Gelidium pristoides and Hypnea spicifera) and changes in overall δ¹³C, δ¹⁵N signatures and C:N ratios during degradation in both the field and laboratory. We hypothesized that both degrading macroalgal tissue and macroalgal-derived suspended particulate material (SPM) would show negligible changes in δ¹³C, but enriched δ¹⁵N signatures and lower C:N ratios relative to healthy plants. Only C:N laboratory ratios conformed to predictions, with both species of macroalgae showing decomposition related changes in δ¹³C and significant depletions in δ¹⁵N in both the field and laboratory. In the laboratory, algal tissue and SPM from each species behaved similarly (though some effects were non-significant) but with differing strengths. Gelidium pristoides δ¹³C increased and C:N ratios decreased over time in tissue and SPM; δ¹⁵N became depleted only in SPM. Hypnea spicifera, δ¹³C, δ¹⁵N and C:N ratios all decreased during degradation in both SPM and algae.     

Effects of suction-dredging for cockles on non-target fauna in the Wadden Sea

Suction dredging for cockles removes large cockles from tidal flats and may also cause mortality of non-target fauna and make the habitat less suitable for some species. This study examines whether suction dredging for cockles on tidal flats of the Dutch Wadden Sea had affected densities of non-target fauna, directly after fishing and one year later. Densities of non-target fauna in two randomly chosen undredged locations were compared to densities at the surrounding heavily commercially dredged area. A significant negative effect of cockle dredging on densities of 0-group Macoma balthica was observed and this effect persisted one year after dredging. The dredged area appeared to be less suitable for settlement of mussels Mytilus edulis. No significant effects of dredging on the mudsnail Hydrobia ulvae and on 0 and 1-group C. edule were found. For the mobile young Macoma balthica it seems unlikely that the effect found after one year was still due to the mortality caused by dredging and this suggests that the habitat was less suitable as a consequence of dredging. Thus, even in the highly dynamic ecosystem of the Wadden Sea, effects of bottom disturbance by cockle dredging may persist after one year.     

Histopathological alterations in Mya arenaria following a # 2 fuel oil spill in the arthur kill, Elizabeth, New Jersey

Following a 2270000-liter #2 fuel oil spill on 1 January 1990 studies were carried out to evaluate histopathological alterations in a population of Mya arenaria inhabiting an impacted tidal mud flat in the Arthur Kill, Elizabeth, New Jersey. Approximately 30 clams were collected 1 week after the spill and on a monthly basis thereafter. The lesion incidence found in M. arenaria following the spill was compared to a base-line incidence established at the same location during 1988 and 1989. There was an increased incidence of moderate and severe lesions of the digestive gland, gills, and mantle 1 week after the spill, returning to pre-spill levels by 1 month. During the first 3 months post-spill an increase in lesion incidence of the intestines was observed which returned to previous levels by 6 months post-spill. The heart and kidney showed a latent increase in lesion incidence 2 and 3 months following the spill, which by 4 months returned to previous levels. Examination of the gonadal development indicated that ova and sperm development were not adversely affected and spawning occurred both in late spring and late summer. The siphon, gill, stomach, foot muscle, and body epithelium were not found to be adversely affected.     

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.