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.     

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.     

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.     

Consequences of the Sudden Removal of Nearly All Mussels and Cockles from the Dutch Wadden Sea

Abstract. Three successive years of recruitment failure, intensive fishery, and high rates of natural mortality eliminated all intertidal mussel beds and nearly all cockle beds from the Dutch Wadden Sea in the course of 1990. In late 1990 and early 1991, zoobenthic biomass and particularly bivalve stocks were unprecedentedly low. The following unusual events were noted:

• 1)

  Relatively high diatom and chlorophyll concentrations in the autumn of 1990 and an exceptionally early and dense spring bloom of diatoms in March 1991;
• 2)

  Exceptionally early onset of rapid growth in the bivalve Macoma balthica in March 1991;
• 3)

  Above-average weights of the soft parts of bivalves in the 1990/91 winter;
• 4)

  Elevated mortality rates in bivalves that could serve as alternative prey for birds specialised on big bivalves (oystercatchers and eiders);
• 5)

  Signs of food shortage in such birds: high mortality rates and emigration to other feeding areas;
• 6)

  A rapid recovery of the benthos started in the summer of 1991, accelerated by an exceptionally high recruitment in some of the affected species.

It is concluded that the Wadden Sea ecosystem responded in an elastic way to the removal of two of its key species. The events are not seen as merely coincidental but are regarded as a chain of causes and effects.     

Numbers of larvae and primary plantigrades of the mussel Mytilus edulis in the western Dutch Wadden Sea

Abundance of larvae and primary plantigrades in the blue mussel Mytilus edulis L. was studied quantitatively in the western part of the Dutch Wadden Sea. Observations were made in seven years between 1981 and 1996. In general, a first large peak of larvae appeared in May, followed by one or more much smaller peaks until September. For pelagic larvae an approximate life time of four to six weeks was calculated (on average about five weeks). Pelagic larvae appeared to have a relatively higher survival chance at low larval concentrations than at high concentrations. No relation could be demonstrated between time of larval peaks and spring tides. Year-to-year differences in abundance of plantigrades did not predict mussel recruitment success on nearby tidal flats.     

Seasonal variation of hepatic EROD activity in flounder (Platichthys flesus) in the Dutch Wadden Sea

The use of hepatic 7-ethoxyresorufin O-deethylase (EROD) activity in flounder (Platichthys flesus) as a potential biomarker of marine pollution by organic chemicals (as indicated by hepatic PCB-153 concentration) was investigated in the Dutch Wadden Sea in 1989 and 1990. Particular attention was paid to the seasonal variation. The results indicate that EROD activity was reasonably stable during the year, except for a period in January–February when a marked peak occurred. This peak coincided with the absence of mature flounder from the sampling area probably due to the spawning migration. Generally a good relationship existed between hepatic EROD activity and PCB-153 concentration, with one exception. During the period January–April 1990 a peak in PCB-153 concentration occurred immediately after the observed peak in EROD activity. It is concluded that the best period for monitoring EROD activity in flounder is the summer, corresponding to the non-migratory period of the species.     

Seasonality of microzooplankton grazing in the northern Wadden Sea

A sequence of nine dilution experiments was conducted according to Landry and Hassett [Landry, M.R., Hassett, R.P., 1982. Estimating the grazing impact of marine microzooplankton. Mar. Biol. 67, 283–288] in the northern Wadden Sea from March until October 2004 to investigate the seasonality of microzooplankton grazing. From March until April, no grazing was observed. Microzooplankton grazing started in May (0.66 d^− 1) and increased until August (1.22 d^− 1). In October microzooplankton grazing was low again (0.17 d^− 1). Phytoplankton growth rates varied between 0 and 1.1 d^− 1. Since the reliability of dilution experiments is still frequently discussed in literature, we tested if our data obtained by dilution experiments reflected short-term in situ phytoplankton dynamics of the study site. We scaled experimental growth rates to water column irradiance, calculated short-term chlorophyll-a dynamics and compared the results to in situ measured chlorophyll-a concentrations. Calculated chlorophyll-a concentrations correlated significantly with in situ measured chlorophyll-a concentrations but slightly overestimated the in situ measured chlorophyll-a. This overestimation was in the range of phytoplankton assimilation reported for the Wadden Sea benthos. We will show that microzooplankton grazing had a large impact during the Phaeocystis bloom and during summer suggesting that a large proportion of phytoplankton biomass remained the pelagic food web. Microzooplankton grazing did not impact the diatom spring bloom and its demise.     

Gametogenic development and spawning of the razor clam,Zenatia acinaces in northeastern New Zealand

The potential exists for New Zealand to exploit already established markets for razor clams through development of fisheries or aquaculture industries for the New Zealand razor clam, Zenatia acinaces. However, fishery or aquaculture development for Z. acinaces requires an understanding of the reproductive cycle including the timing of gametogenic development and spawning. The reproductive cycle of Z. acinaces was studied over an 11‐month period from May 2000 to March 2001 at Kennedy Bay, Coromandel Peninsula, using qualitative standard histological analysis and quantitative measures of oocytes. Histological analysis indicated that Z. acinaces is dioecious and gametogenic development was synchronous between the sexes. Gametogenesis began in June with gametes maturing quickly and by August/September (late winter/ early spring) most razor clams were ripe. Spawning began as early as September (spring) although spawning mainly occurred during October. By December (summer), nearly all clams were completely spent. From January 2001 most clams could not be sexed as all residual gametes were resorbed. Razor clams remained in this stage during March 2001. Spawning began when the water temperature was around its lowest, c. 15°C. Monthly mean number of eggs/follicle was sensitive to changes in reproductive development, closely following patterns observed in the qualitative stagings. Patterns of monthly mean oocyte diameters did not adequately describe the spawning events observed in qualitative analyses. Sex ratios were equal over the size range (69–99 mm shell length) of clams that could be sexed. The data presented in this study provide valuable information on the timing of spawning events for Z. acinaces, necessary for developing sustainable management strategies and selecting broodstock for aquaculture.     

The impact of water jet dredging for razor clams,Ensis spp., in a shallow sandy subtidal environment

The effects of water jet dredging for Ensis spp. on the seabed and benthos were examined through experimental fishing. Immediate physical effects were apparent, with the dredge leaving visible trenches in the seabed. While these trenches had started to fill after five days, and were no longer visible after 11 weeks, the sediment in fished tracks remained fluidised beyond this period. The majority of the studied infaunal community is adapted morphologically and behaviourally to a dynamic environment, and other than initial removal through dispersal, is not greatly affected by the dredge at the site studied. Species that are likely to be affected (e.g. the heart urchin Echinocarium cordatum, Arctica islandica and other large bivalves) were very rare in infaunal samples, but present in dredge catches, where damage was noted, and ranged on average from 10 to 28% of individuals. Epifauna were scarce in the study area, and unaffected by the fishing, except that epifaunal scavenging species were attracted to the fished tracks. On the evidence of the present and previous studies, it would appear that there was little difference between the biological impact of hydraulic and suction dredging, although the latter may have a greater physical effect (larger trenches).     

Benthic photosynthesis in submerged Wadden Sea intertidal flats

In this study we compare benthic photosynthesis during inundation in coarse sand, fine sand, and mixed sediment (sand/mud) intertidal flats in the German Wadden Sea. In situ determinations of oxygen-, DIC- and nutrient fluxes in stirred benthic chamber incubations were combined with measurements of sedimentary chlorophyll, incident light intensity at the sediment surface and scalar irradiance within the sediment. During submergence, microphytobenthos was light limited at all study sites as indicated by rapid response of gross photosynthesis to increasing incident light at the sea floor. However, depth integrated scalar irradiance was 2 to 3 times higher in the sands than in the mud. Consequently, gross photosynthesis in the net autotrophic fine sand and coarse sand flats during inundation was on average 4 and 11 times higher than in the net heterotrophic mud flat, despite higher total chlorophyll concentration in mud. Benthic photosynthesis may be enhanced in intertidal sands during inundation due to: (1) higher light availability to the microphytobenthos in the sands compared to muds, (2) more efficient transport of photosynthesis-limiting solutes to the microalgae with pore water flows in the permeable sands, and (3) more active metabolic state and different life strategies of microphytobenthos inhabiting sands.     

Spatial and temporal variability in bio-optical properties of the Wadden Sea

The Wadden Sea, a shallow coastal area bordering the North Sea, is optically a complex area due to its shallowness, high turbidity and fast changes in concentrations of optically active substances. This study gathers information from the area on concentrations of suspended particulate matter (SPM), Chlorophyll-a (Chl-a), and Coloured Dissolved Organic Matter (CDOM), on total absorption and beam attenuation, and on reflectances from the whole area. It examines the processes responsible for variations in these. Sampling took place at 156 stations in 2006 and 2007. At 37 locations also the specific inherent optical properties (SIOPs) were determined. Results showed large concentration ranges of 2–450 (g m^-3) for SPM, 2–67 (mg m^-3) for Chl-a, and 0–2.5 m⁻¹ for CDOM(440) absorption. Tides had a large influence on the SPM concentration, while Chl-a had a mainly seasonal pattern. Resuspension lead to a correlation between SPM and Chl-a. The absorption of CDOM had a spatial variability with extremely high values in the Dollard, although the slope of CDOM absorption spectra was comparable with that of the North Sea. The Chl-a specific pigment absorption proved to be influenced by phytoplankton species and specific absorption of non-algal particles at 440 nm was correlated with the mud content of the soil at the sample locations. SPM specific absorption was not found to correlate with any measured factor. As the concentrations of optically active substances changed, we also found spatial and temporal variability in the absorption, beam attenuation and reflectances. Reflectance spectra categorized in groups with decreasing station water depths and with extreme CDOM and SPM concentrations showed distinguishable shapes.     

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.     

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.     

缢蛏耐高盐性能及其与生长性状的相关性分析

缢蛏(Sinonovacula constricta)是我国重要的水产经济贝类,开展耐高盐新品种选育对平衡不同地区缢蛏良种养殖意义重大。本研究对不同家系缢蛏幼贝进行高盐胁迫,运用统计学方法和相关性分析探究了缢蛏耐高盐性能及其与生长性状的相关性。结果显示,缢蛏家系间壳长、壳高、壳宽和体质量表型值差异较大,体质量变异系数达15.53%。不同家系对高盐耐受性差异较大,家系间死亡率范围在0～100%。胁迫实验共持续9 d,各家系缢蛏首粒死亡时间在第1～9 d之间,而且死亡率较高的家系内缢蛏死亡时间离散程度较大,说明不同个体间高盐耐受性差异也较大。缢蛏开始出现大量死亡的时间主要在胁迫后第8 d,第9 d部分家系死亡率高于80%,说明缢蛏对高盐耐受存在一个阈值,超过该阈值会开始或大量死亡。皮尔逊和斯皮尔曼相关性分析显示,缢蛏存活时间和生长性状为正相关,其中存活时间与缢蛏壳长、体质量相关性均达到显著水平(P<0.05)。缢蛏存活率与体质量、生长速度均显著正相关(P<0.05)。整体而言,壳长较长和体质量较重的缢蛏个体耐高盐性能较强,在高盐环境中具有生长优势。本研究揭示了缢蛏的高盐耐受性及...     

Infection by gymnophallid metacercariae enhances predation mortality of SW Atlantic stout razor clam Tagelus plebeius

Parasite life cycles are frequently completed in different hosts, thus the parasites have its life cycle overlapped to natural trophic webs. The family Gymnophallidae (Class: Trematoda; Subclass: Digenea) includes digenetic parasites whose larval stages occur on bivalves and may affect bivalve predation by the final host of these parasites. In this work we evaluated: (a) if individuals of the razor clam Tagelus plebeius with higher parasite intensity suffer higher predation by the oystercatcher Haematopus palliatus and, (b) if there is any effect of parasite intensity on burrowing and escape behaviours of these razor clams which may enhance exposure to predators. Field experiments (oystercatcher exclusion vs. open access) showed that clams with higher parasite intensity support higher predation by oystercatchers, which suggests a higher consumption of more parasitized clams and thus, a more successful reproduction of parasites linked to the intensity of infection. However, clam burrowing and escape behaviours did not show differences related to different parasite intensity, suggesting that the commonly believed mechanisms are not responsible in this case.