Long-term effects of mechanical harvesting of lugworms Arenicola marina on the zoobenthic community of a tidal flat in the Wadden Sea

More than half of the annual catch of about 30 million lugworms Arenicola marina from the Dutch Wadden Sea originates from digging machines which make 40-cm deep guilles in a few restricted tidal-flat areas (Texel, Balgzand) in the westernmost part of the Wadden Sea. Four successive years (1978–1982) of frequent disturbance by a lugworm dredge of one of the 15 sampling stations involved in a long-term study of the dynamics of the macrozoobenthos on Balgzand allowed a study of long-term effects of mechanical lugworm digging.Within an area of about 1 km², a near-doubling of the annual lugworm mortality rate resulted in a gradual and substantial decline of the local lugworm stock from more than twice the overall Balgzand mean at the start of the 4-year digging period to a value close to this mean at the end of the period (when the dredge moved to a richer area). Simultaneously, total zoobenthic biomass declined even more by the almost complete extinction of the population of larger gaper clams Mya arenaria that initially comprised half of the total biomass. Of the other, mostly short-lived, species only Heteromastus filiformis showed a clear reduction during the dredging period. Recovery of the biomass of the benthos took several years, particularly by the slow re-establishment of a Mya population with a normal size and age structure.     

Year-to-year variability in the biomass of macrobenthic animals on tidal flats of the Wadden Sea: How predictable is this food source for birds?

The Wadden Sea is important as a stop-over and wintering area for several species of waders, foraging for shorter or longer times on its tidal flats. The size of the food stocks these birds encounter varies from place to place and from year to year. We studied characteristics of the variability in time of such prey stocks, using long-term data series of annual estimates of biomass of macrobenthic animals collected on tidal flats in various parts of the Dutch and German Wadden Sea. Year-to-year fluctuations were stronger in nearly all individual benhos species than in total macrozoobenthic biomass. The various species differed significantly in their year-to-year variability. Everywhere the tellinid bivalve Macoma balthica was relatively stable, whereas species such as the cockle Cerastoderma edule and the polychaetes Lanice conchilega, Nephtys hombergii and Anaitides mucosa fluctuated heavily and rapidly in all parts of the Wadden Sea where they were found. Within species, fluctuations in biomass of individual age or size classes were stronger than those in total biomass of the species.Several species showed minimal biomass values in the same years over vast areas. This synchronization of scarcity was caused particularly by similar responses to winter character, which was each year similar over the entire Wadden Sea. Such similar responses included low over-winter survival during severe winters and recruitment failure after exceptionally mild winters. Immediately after severe winters, such cold-sensitivee species as C. edule, L. conchilega and N. hombegii were scarce all over the Waden Sea, thus liimiting the possibilities for birds to switch to other parts of the Wadden Sea to find their preferred prey in sufficient quantitu. Simultaneous recruitment failure in several important bivalve species (C. edule, Mytilus edulis, and Mya arenaria limited the possibility for specialized bivalve consumers to switch to alternative prey types in certain years.     

Distribution and abundance of the zooplankton of the Ems estuary (North Sea)

From 1974 to 1977 a study was made of the abundance and the distribution of the zooplankton species of the Ems estuary (The Netherlands and the Federal Republic of Germany), an area of about 500 km² with extensive tidal flats.The most important component of the zooplankton consisted of holoplanktonic calanoid copepods with, during the summer, a significant contribution of meroplankton, mainly consisting of polychaete and cirripede larvae. Zooplankton abundance showed a marked seasonality, with a pronounced spring peak and a smaller late summer/autumn peak. In the low salinity area the spring bloom was dominated by Eurytemora affinis, which persisted in the salinities below S=5 upriver during the whole year. In the polyhaline area Acartia bifilosa was the main component of the spring bloom. The late summer maximum from August to October was dominated by Acartia tonsa in the meso- and polyhaline area and by A. discaudata and Centropages hamatus in the euhaline part of the estuary bordering on the Wadden Sea proper. The number of abundant (> 1000 ind·m⁻³) species increased from the inner, fresher part of the estuary towards the Wadden Sea but, except for April, average zooplankton density in salinities <18 was similar (2768 ind·m⁻³) to the density in salinities >18 (2817 ind·m⁻³).The distribution patterns show that there are only a few autochthonous species and that the majority of species is allochthonous and penetrates more or less deeply into the estuary from the Wadden Sea and North Sea, varying with species and season.     

An evaluation of intertidal feeding habitats from a shorebird perspective: Towards relevant comparisons between temperate and tropical mudflats

Macrozoobenthic communities of intertidal soft sediments are reviewed worldwide from the perspective of a mollusc-eating shorebird species. Based on 19 sites, total biomass figures varied between 5 and 80 g AFDM per m² (average 24 g AFDM per m²); no latitudinal trends are apparent. The contribution made by bivalves and gastropods varies between 1% and 99%, north-temperate intertidal flats having relatively more molluscs than tropical flats. Intertidal flats in the tropics contain a greater variety of taxa, with brachiopods in Indonesia and echinoderms in northwest Australia contributing significantly to biomass only there. Limits to the occurrence of avian predators of intertidal benthos are set by the harvestable fraction of the biomass on offer and the costs of living at a particular site. No systematic differences in the harvestable fraction of the total mollusc-biomass for a worldwide occurring shorebird species specializing on molluscs (knots Calidris canutus) were apparent between temperate and tropical intertidal areas, in spite of large differences in maintenance metabolism incurred by these birds. The harvestable fractions of bivalves in the two West African areas (Banc d'Arguin, Mauritania and Guinea-Bissau) tended to be high (23–84% of total biomass in six species), they were relatively low (2–52% in five species) in the temperate Wadden Sea and the tropical northwest Australian site. Harvestable biomass determines the intake rate of shorebirds, as illustrated by functional-response curves of knots feeding on two bivalves species. We argue that the collection of information on size-depth relationships along with faunal and biomass surveys at a range of sites is bound to greatly increase our understanding of both the biology of tidal-flat invertebrates and the resource base underpinning the spectacular seasonal migrations of shorebirds.     

How the food supply harvestable by waders in the Wadden Sea depends on the variation in energy density,body weight,biomass, burying depth and behaviour of tidal-flat invertebrates

For several reasons, waders in the Wadden Sea face a large seasonal and annual variation in their food supply. Observations on a tidal flat in the Dutch Wadden Sea have shown that:

• 1.- (1) The average energy density of ten invertebrate prey species varies between 21 and 23 kJ·g⁻¹ AFDW. In Scrobicularia plana and Mya arenaria, but not in Macoma balthica, the energy density is 10% lower in winter than in summer.
• 2.- (2) Depending on the species, body weights of prey of similar size are 30 to 60% lower in winter than in summer.
• 3.- (3) The year-to-year fluctuation in standing-crop biomass is larger in some species than in others, the difference depending mainly on the frequency of successful recruitment. The overall biomass of the macrobenthos in winter is half of that in summer, but the timing of the peak biomass differs per species.
• 4.- (4) The burying depth varies per species: Cerastoderma edule live just beneath the surface, while M. balthica, S. plana, M. arenaria, Arenicola marina and Nereis diversicolor bury more deeply and the majority of these prey live out of reach of the bird's bill. In all six species, burying depth increases with size. There is no seasonal variation in depth of C. edule and M. arenaria, but the four other species live at most shallow depth in early summer and most deeply in midwinter. Burying depths in winter vary from year to year, but are unrelated to temperature. Neither has temperature any effect on depth within months. For knot Calidris canutus feeding on M. balthica, the fluctuation in the accessible fraction was the main source of variation in the biomass of prey that is actually harvestable, i.e. the biomass of prey of suitable size that is accessible.Accordingly, the paper reviews the available data on the temporal variations in accessibility, detectability, ingestibility, digestibility and profitability of prey for waders. Only a small part of the prey is harvestable since many accessible prey are ignored because of their low profitability, while many profitable prey are inaccessible. The profitability of prey depends on their size and weight but also on their depth in the mud, since handling time increases with burying depth. A simple biomechanical rule explains why the handling time of small prey increases with bill length and why large, long-billed waders ignore a disproportionately larger part of the small prey. The fraction detectable for visually feeding waders is usually very low, especially when the temperature of the substrate is below 3–6°C. Waders vary their prey choice over the year in response to the changes in the availability and profitability of their different prey species. The food supply harvestable by waders is much lower in winter than in summer. For waders wintering in the Wadden Sea, the food supply may be characterized as unpredictable and usually meagre. Waders wintering in NW Europe are concentrated in coastal sites where the average surface temperature is above 3°C. This probably cannot be explained by a greater burying depth, and only partly by a lower body condition, of prey in colder areas. Yet the harvestable fraction is lower in colder sites, especially for sight-feeding waders, as invertebrates are less active at low temperatures. However, the lower energetic cost of living and reduced chances of the prey being covered by ice may also contribute to the waders' preference for warmer sites.

     

Chattonella and Fibrocapsa (Raphidophyceae): First observation of,potentially harmful,red tide organisms in Dutch coastal waters

Species of the potentially toxic and red-tide-forming marine-phytoplankton genera Chattonella and Fibrocapsa (Raphidophyceae) were observed for the first time in 1991 in samples taken in Dutch coastal waters; they were again recorded and enumerated in the following years. Chattonella spp. cell numbers varied with the season, with a maximum in May or June in the Dutch Wadden Sea. Cell numbers of Chattonella and F. japonica Toriumi et Takano were up to 6.0·10³ cells·dm⁻³ in the Dutch Wadden Sea, except at one station in June 1993 when over 10⁴ cells·dm⁻³ Chattonella were counted. In May 1993, a minor bloom (over 2.0·10⁵ cells·dm⁻³) was observed at a station in the southern central North Sea, 100 km northwest of the island of Terschelling. The potentially neurotoxic species Chattonella marina (Subrahmanyan) Hara et Chihara was identified and discriminated from morphologically related species within the class of Raphidophyceae by immunofluorescence. F. japonica could only be clearly identified in live samples; in fixed samples cell morphology was severely affected. The identification of this species was supported by the presence of mucocysts, structures that can be observed readily by optical and electron microscopy.     

Adaptation to anaerobic metabolism in two mussel species,Mytilus edulis and Mytilus galloprovincialis,from the tidal zone at Arcachon Bay,France

Aspects of anaerobic metabolism were investigated in two sympatric mussel species, viz. Mytilus edulis and Mytilus galloprovincialis, living in the tidal zone in Arcachon Bay, France. Specific activities of pyruvate kinase (PK) and phosphoenolpyruvate kinase (PEP-CK) were remarkably similar in the two sympatric species and generally corresponded more closely to those observed in M. galloprovincialis in the Mediterranean than with M. edulis in the Dutch Wadden Sea. However, the values for the radio PK: PEP-CK for the two species in Arcachon Bay agreed with those of intertidal M. edulis from the Dutch Wadden Sea.Succinate accumulation during the first 24 h of anaerobicsis was about the same as in M. galloprovincialis in the Mediterranean, but decreased during the second 24 h, particularly in M. edulis, obviously due to propionate formation. Decrease in ATP concentrations in the tissues during anaerobiosis corresponded to that of intertidal M. edulis from the Dutch Wadden Sea. With the exception of specific activities of PK and PEP-CK, all properties investigated in both species were as expected in intertidal mussels.     

Trophic resource partitioning between two coexisting flatfish species off the North coast of Anglesey,North Wales

Interspecific and intraspecific trophic resource partitioning between coexisting plaice, Pleuronectes platessa L. and dab, Limanda limanda (L.) populations were investigated at three subtidal locations in the Irish Sea: Two inshore locations off the coast of Anglesey, North Wales and one off-shore location. In shallow inshore waters, the two species had similar diets dominated by the polychaete Pectinaria koreni. Offshore dab appeared to concentrate on Ensis ensis whilst the plaice took a wider variety of prey including bivalves, polychaetes, crustaceans and sand eels. Trawls made over a 24-hour period indicated some crepuscular and nocturnal feeding by individuals of both species. The observed patterns of resource partitioning were due to the abundance of prey species which vary spatially and seasonally and are discussed in relation to current work on resource partitioning between fishes.     

Scale and intensity of intertidal habitat use by knots Calidris canutus in the Western Wadden Sea in relation to food,friends and foes

In August–October 1988–1992 we studied the distribution and abundance of knots Calidris canutus around Griend in the western Wadden Sea, and the extent to which these can be explained by benthic prey availability and presence of avian predators. Numbers in the nonbreeding season showed monthly averages of 10 000 to 25 000 birds. Over 100 000 knots were recorded on three occassions. Knots feed in large flocks, individual birds usually experiencing 4 000 to 15 000 flock-mates. The Siberian-breeding/west-African wintering canutus subspecies passed through in late July and early August. Otherwise the Greenlandic/Canadian breeding islandica subspecies was present. Over the period 1964–1992 there were no clear trends in the number of knots, but canutus-knots were particularly abundant in July–August 1991, whereas in 1992 both subspecies were absent.Macoma balthica was the preferred prey of both subspecies. Hydrobia ulvae, Mytilus edulis and Cerastoderma edule were eaten when Macoma was absent close to the surface of the sediment. As Macoma buried deeper from July onwards, canutus faced better average feeding conditions than islandica later in the year. The spatial distributon of knots feeding on the intertidal flats around Griend was best explained by the harvestable biomass of the prevalent prey species in a particular year and season, i.e. Macoma (main prey when their harvestable biomass densities were greater than ca 0.8 g AFDM per m²) and Cerastoderma, and by the avoidance of situations where they run the risk of attack by bird-eating birds. Flocks of knots covered most of the intertidal flats in the Western Dutch Wadden Sea in a couple of tidal cycles. This is about 800 km², much larger than the equivalent area used by knots on their wintering grounds in Mauritania (10–15 km²), a difference that is correlated with prey spectrum, prey availability and predictability.     

Increased mortality in alternative bivalve prey during a period when the tidal flats of the Dutch Wadden Sea were devoid of mussels

In the course of 1990, stocks of mussels (Mytilus edulis) declined to unprecedentedly low levels in the Dutch Wadden Sea. Hardly a wild mussel bed was left on the tidal flats as a consequence of three years (1988, 1989, and 1990) with failing recruitment and intensive fishing for seed mussels. During these three years, recruitment of cockles (Cerastoderma edule) also failed, whereas fishing was continued. Bird species taking these bivalves as staple food, the oystercatcher (Haematopus ostralegus) and the eider (Somateria mollissima), experienced food shortage. Significant numbers of eiders left the Dutch Wadden Sea area or died, whereas oystercatchers remained abundant trhoughout the winter in most of the Dutch Wadden Sea.Alternative prey species of oystercatchers experienced unsually high mortality rates in the appropriate size classes. This was so in all other common species of bivalves, viz. first-year and older cockles, adult Macoma balthica, and juvenile Mya arenaria. This led to minimal stocks of food for oystercatchers in the late winter of 1991. In March 1991, cockles were depleted and the combined stocks Mya and Macoma would soon have run out of food supply to the overwintering oystercatcher population. Apparently, oystercatchers are able to reduce the stocks of their various bivalve prey species to very low levels.     

Community structure and biomass distribution of seagrasses and macrofauna in the flores sea,Indonesia

In several locations in the Flores Sea region the community structure and the biomass distribution of seagrasses were studied along transects perpendicular to the shoreline. The share of each species within a sample plot was estimated, divided in above- and below-ground biomass. Statistics regarding substrate coverage, shoot density and leaf-area index were sampled. A standard relation was calculated between seagrass dry weight, ash-free dry weight and organic carbon content.The biotic data were related to environmental factors: DOC and nutrients in the water, salinity, tidal amplitude, sediment composition. A relation was estimated between bottom coverage of seagrasses and standing stock. Further calculations of biomass-production ratios allow a quick and rough estimate of seagrass productivity. Maximum above-ground biomass values (500–700 g AFDW·m⁻²) together with qualitative data indicate resource (= space) partitioning among the component seagrasses within a community, and suggest a carrying capacity of the reefflat habitat for seagrass density and biomass.A tentative model was constructed, starting from a constant, non-distributed multispecies vegetation in the lower intertidal and subtidal zone on sand and coral rubble, and moving into several suboptimal situations. The upper shore carries an impoverished, constrained vegetation (irregular tides, desiccation, harvesting). Sediment reworking by animals and physical displacement of sand disturbs the vegetation and favours pioneer species. Muddy habitats bordering mangroves carry monospecific stands showing extremely high biomass (e.g. below-ground Enhalus acoroides 3500 g AFDW·m⁻²). Thalassia hemprichii and Enhalus acoroides are the most constant species in all habitats mentioned.Macrofauna biomass within the seagrass beds fluctuated widely (maximum values 50–70 g AFDW·m⁻² in mixed seagrass vegetations) and only a weak relation between benthic macrofauna biomass and seagrass community structure and biomass was found.     

The importance of rivers as nursery grounds for 0- and 1-group flounder (Platichthys flesus L.) in comparison to the Wadden sea

From May 1988 to June 1989 selected areas of the German Wadden Sea, the Ems, Weser, Elbe and Eider estuaries as well as in Elbe tributaries were sampled for 0- and 1-group flounder (Platichthys flesus L.). Estuarine and Wadden Sea sampling was conducted on board commercial shrimp vessels equipped with beam trawls. The Elbe river system was sampled on board a research boat equipped with a similar 3-m beam trawl. Density indices were calculated for the various areas. To estimate the importance of rivers as nursery areas for flounder, river surface area was compared to the area of tidal flats in the Wadden Sea.In all surveys, there was no relationship between fish densities and tow directions with respect to tide. In the Elbe river system flounder densities were not correlated to substrate types but increased significantly with decreasing salinity. Abundances were always lowestt in polyhaline habitats and increased up to 10-fold in mesohaline estuarine areas. In the limnetic tidal sections of the Elbe river and its tributaries densities increased again by factors of 3 to 10. The succession of 1-group modal lengths from limnetic to polyhaline habitats demonstrated that smaller fish preferred less saline waters. 0-group specimens lagging behind in growth did not leave the limnetic river section in winter. The onset of the spawning migration in November was determined by monitoring the length-frequency distributions throughout the year.The possible sources of bias are discussed and gear efficiency is estimated from literature sources. The contribution of rivers to the 0- and 1-group flounder population on the tidal flats of the Wadden Sea is estimated at about 35.1%.     

Large-scale distribution patterns of the mussel Mytilus edulis in the Wadden Sea of Schleswig-Holstein: Do storms structure the ecosystem?

The distribution of mussel beds in the Wadden Sea of Schleswig-Holstein was mapped by aerial surveys from 1989 to 1991. The number of mussel beds decreased from 94 in 1989 to 49 in 1990, as a result of severe storms in early 1990. Thereafter only small changes were observed. The mussel beds that remained in 1990 were found only in the shelter of islands; all beds in exposed areas had disappeared between the surveys of 1989 and 1990, leaving large areas without mussel beds. Storms are thus identified as a major factor limiting the distribution of mussel beds to the sheltered parts of the Wadden Sea. Beds in the exposed parts of the Wadden Sea are highly dynamic, whereas beds in sheltered areas may persist over long times. A comparison with distribution patterns of older surveys (from 1937, 1968 and 1978) revealed great similarities with the results of recent investigations, indicating a constant distribution pattern over a long period.The results are discussed in relation to eutrophication and the structure of the benthic communities of the Wadden Sea. It is concluded that any eutrophication-induced increase of the mussel population would be restricted to the sheltered parts of the Wadden Sea. Storms will largely determine whether the communities of a given area have to compete with mussels, which are most important filter feeders of the ecosystem. As competition for food is a major factor structuring the benthic communities of the Wadden Sea, it is assumed that storms indirectly affect all other communities, giving deeper-burying, storm-tolerant species a competitive advantage in exposed areas where epibenthic mussels are excluded. The impact of mussel fisheries will be different for persisting and dynamic beds: fishing on persisting beds in sheltered areas may remove the crucial reserve which mussel-feeding birds such as eiders or oystercatchers need in times of low mussel populations.     

Wind effects on prey availability: How northward migrating waders use brackish and hypersaline lagoons in the sivash,Ukraine

Large numbers of waders migrating northward in spring use the Sivash, a large system of shallow, brackish and hypersaline lagoons in the Black Sea and Azov Sea region (Ukraine). The bottoms of these lagoons are often uncovered by the wind. Hence, for waders the time and space available for feeding depend on wind conditions. In hypersaline lagoons the benthic and pelagic fauna was very poor, consisting mainly of chironomid larvae (0.19 g AFDM·m⁻²) and brine shrimps Artemia salina, respectively. Brine shrimp abundance was correlated with salinity, wind force, wind direction and water depth. Dunlin Calidris alpina and curlew sandpiper Calidris ferruginea were the only species feeding on brine shrimp. As brine shrimp densities are higher in deeper water, smaller waders such as broad-billed sandpipers Limicola falcinellus are too short-legged to reach exploitable densities of brine shrimp. In brackish lagoons the benthic and pelagic fauna was rich, consisting of polychaetes, bivalves, gastropods, chironomid larvae, isopods and amphipods (8.9 to 30.5 g AFDM·m⁻²), but there were no brine shrimps. Prey biomass increased with the distance from the coast, being highest on the site that was most frequently inundated. Dunlin, broad-billed sandpiper and grey plover Pluvialis squatarola were the most abundant birds in the brackish lagoon. Due to the effects of wind-tides only a small area was usually available as a feeding site. Gammarus insensibilis was the alternative prey resource in the water layer, and their density varied with wind direction in the same way as brine shrimp. Curlew sandpipers and dunlins in the hypersaline lagoons and broad-billed sandpipers in the brackish lagoons often changed feeding sites, probably following the variation in prey availability. Only because of the large size and variety of lagoons are waders in the Sivash always able to find good feeding sites.     

Phytoplankton primary production and biomass in the western Wadden Sea (The Netherlands); A comparison with an ecosystem model

Dissolved nutrient concentrations (Si, P, N), algal biomass (separated into diatoms and micro-flagellates), and total primary production were determined in 1986 during 7 cruises in both the inner and outer parts of the estuary of the western Wadden Sea. The biomass data, obtained after converting cell biovolume and cell counts into carbon, were compared with those simulated by a two-dimensional multitrophic ecosystem model.The general pattern of the succession of algal species was the same for all compartments: a bloom of diatoms in early spring followed by one of microflagellates, predominantly the colonial alga Phaeocystis pouchetii. Throughout the year the biomass of the two algal groups differed between inner and outer compartments of the estuary. In spring the carbon biomass of the microflagellates in the compartments bordering on the North Sea exceeded that of the inner compartments by a factor of 2. The reverse was found for the diatoms in summer.A reasonable fit between the field data and simulated data was only obtained in spring; large differences were found for the summer period. During that period an extensive diatom population characterized by a low primary production was observed, which did not correspond with the model simulation.Possible factors causing the differences between observed and simulated data, such as nutrient regeneration, were analysed.     

Long-term studies on Noctiluca scintillans in the German Bight population dynamics and red tide phenomena 1968–1988

The seasonal development of Noctiluca scintillans has been studied by continuous monitoring at Helgoland-Roads over a period of 20 years. These long-term studies have revealed a temporal distribution pattern of seasonal periodicity with maximal abundances in June/July, and with a low but pronounced peak in December/January. Additionally, over a 21-year term a periodicity of high abundances in three years intervals is observed. In Helgoland waters the mean near surface abundances are higher than the near sea bed values. Apart from physico-chemical influences, buoyancy in Noctiluca is also controlled by its nutritional status.Since 1981, investigations have been extended to the German Bight. Potential centres of Noctiluca growth are found within certain areas near the east-frisian Wadden Sea, persisting from April/May until late July when the centres of high cell concentrations are separated from the coastline. Simultaneously, there is a strong tendency for vertical separation, generating the well-known red-tide phenomena with large mass aggregations at the sea surface, often misleadingly called Noctiluca-blooms, although the cell masses are irreversibly damaged and die off very rapidly.The spatial distribution of Noctiluca as determined every year in July, serves as a realistic base for biomass calculations. From 1981 to 1988 the total per season produced biomass (ashfree dry-weight) of Noctiluca ranges between 30000 (1981) and 69000 tons (1982).     

Structure and energy demand of the benthic soft-bottom communities in the Java Sea and around the islands of Madura and Bali,Indonesia

In July 1984 benthic research was carried out in the Java Sea and around the islands of Madura and Bali.The aim was to describe the structure of the benthic communities in terms of numerical density (numbers of dominant macrofaunal groups and meiofauna per square metre) and biomass (g ashfree dry weight (AFDW) per square metre), and in relation to geographical position and main ambient factors, i.e. water depth, bottom temperature and sediment characteristics.In addition attempts were made to estimate the activity of these benthic ecosystems, and to show their relation to the pelagic.Box-core samples, collected along west-east axis in the Java Sea and in the Strait of Madura, showed relatively poor benthic communities. The average densities of macrofaunal organisms (> 1 mm) did not exceed 250 specimens·m⁻²; the total biomass was below 1 g AFDW·m⁻²; small polychaetes and crustaceans were the dominant groups.A number of larger organisms have developed very remarkable adaptations to the muddy environment. Meiofaunal organisms (nematodes being the dominant group) numbered 0.06·10⁶ to 0.46·10⁶ specimens·m⁻². In general there is a trend from west to east towards somewhat richer communities. The actual carbon demand of the benthic ecosystem in the area investigated is tentatively estimated at an average of 38 g C·m⁻²·y⁻¹, which amounts to about 40% of the primary production.Both the very soft fluid-mud bottoms in the central Java Sea and in Strait Madura, and the relatively low amount of energy available for growth, probably prevent the establishment of well developed benthic communities.     

Seasonal variation in growth and biomass of an intertidal Zostera noltii stand in the Dutch wadden sea

To assess relationships between the life cycle of the seagrass Zostera noltii and light conditions in its habitat, the seasonal dynamics of a seagrass-dominated community on a tidal flat off Terschelling were studied. The main components of this community were seagrass, periphyton and the periphyton grazing mudsnail Hydrobia ulvae. Total biomass of the seagrass stand showed a unimodal curve with a maximum of more than 110 gADW·m⁻² in August and a minimum of less than 10 gADW·m⁻² in January. Chlorophyll density of periphyton on the seagrass leaves followed a more or less similar pattern, ranging from 0.4 μg chlorophyll·cm⁻² in May to more than 3 μg chlorophyll·cm⁻² at the end of August. Periphyton biomass was, however, already maximal in May with almost 1.6 mgADW·cm⁻² and subsequently decreased to less than 0.6 mgADW·cm⁻² in August. The total weight of H. ulvae was more or less stable, varying between more than 150 and less than 400 gDW·m⁻², although significant changes were observed within size classes.Light is assumed to be the primary limiting factor for seagrass distribution in the Wadden Sea. The light conditions of seagrass in the study area were influenced by periphyton and mudsnails. Leaf growth rates and biomass development appeared to be related with light conditions in the seagrass habitat. Shading caused by periphyton during the growing season was estimated at 10 to 90% of incident light, resulting in a reduction of about 2 to 80% of the yearly period during which the light compensation point (LCP) of the seagrass is exceeded. The mudsnails were found to be theoretically able to ingest daily 25 to 100% of the standing stock of periphyton and microphytobenthos. It is concluded that shading by periphyton and grazing by mudsnails play an important role in the seasonal biomass development and survival of Z. noltii in the seagrass-dominated community on a tidal flat off Terschelling.     

Temporal and lateral fluctuations in production and biomass of bacterioplankton in the western Dutch Wadden Sea

To contribute to the validation of a recently developed ecosystem model of the western Wadden Sea (EON, 1988), data on bacterial biomass and production were acquired. Seven field stations, spread over the two main basins of the estuarine system, were sampled monthly in 1986. Between these basins significant differences were found in counts, biovolume, biomass and production of bacteria (measured by the ³H-thymidine method) with consistently higher mean values of bacterial variables in the Vlie basin. Bacterial production rates of 2 to 175 mg C· m⁻³· d⁻¹ were obtained for the Vlie basin, with an annual production of 10 to 11 g C· m⁻³, while the production in the Marsdiep basin did not exceed 45 mg C·m⁻³ ·d⁻¹, with an annual production of g C·m⁻³. Bacterial biomass varied over the year from 2 to 140 mg C·m⁻³ in the study area, with a mean biomass of 39 mg C·m⁻³ in the Vlie basin and 23 mg C·m⁻³ in the Marsdiep basin. Blooms of bacteria occurred in May and July–August. Spatial and temporal fluctuations in bacterial variables are discussed, taking into account different environmental factors and the availability of food for bacteria in relation to transport and exchange of water masses between the two basins and the North Sea. Results are compared with the results as simulated by the ecosystem model.