Estimating estuarine residence times in the Westerschelde (The Netherlands) using a box model with fixed dispersion coefficients

The residence time of the water masses in the Westerschelde estuary was determined using a simple compartment-model that simulates the advective-diffusive transport of a conservative dissolved substance (chlorinity). The residence time of a water parcel in the upstream part of the estuary (i.e. the time needed for this water parcel to leave the estuary) varied from about 50 days in winter to about 70 days in summer. The most seaward compartment had residence times of about 10-15 days.Dispersive coefficients that are fixed in time were able to reproduce the observed salinity distributions very well in the Westerschelde. They were obtained by calibration on observed chlorinities. It is argued that the apparent relationship of dispersive coefficients with freshwater flow, which is observed in certain studies, could (partly) reflect the deviation from steady state conditions which are required assumptions to calculate these dispersive coefficients directly from salinity profiles.     

Carbon flows in the Westerschelde estuary (The Netherlands) evaluated by means of an ecosystem model (MOSES)

The autotrophic production and heterotrophic consumption of organic matter in the Westerschelde, a highly turbid and eutrophic estuary in the Southwest Netherlands is examined by means of a dynamic simulation model. The model describes the ecologically relevant processes in thirteen spatial compartments and adequately fits most observed data.Three autotrophic processes are included in the model. Net pelagic photosynthetic production is relatively low (average 41 gC m^–2 yr^–1) and three spatial compartments near the turbidity maximum zone are respiratory sinks of phytoplankton biomass. According to the model, net phytobenthic primary production is more important than pelagic primary production in the upstream half of the Westerschelde. On the scale of the entire estuary, benthic primary production amounts to about 60% of pelagic primary production. Water-column nitrification, which is very important in the nitrogen cycle, is most pronounced near the turbidity zone where it accounts for the major autotrophic fixation of carbon (up to 27 g C m^–2 yr^–1). Viewed on the scale of the total estuary, however, the process is not very important.Less than 20% of total organic carbon input to the estuary is primary produced, the remainder is imported from waste discharges and from the river.The degree of heterotrophy of the Westerschelde estuary proved to be one of the highest yet reported. On average 380 g carbon per square metre is net lost per year (range 200–1200 gC m^–2 yr^–1). The yearly community respiration (bacterial mineralization, respiration of higher trophic levels and sedimentation) is 4 to 35 times (estuarine mean of 6) higher than the net production. This degree of heterotrophy is highest near the turbidity maximum and generally decreases from the freshwater to the seaward boundary. About 75% of all carbon losses can be ascribed to pelagic heterotrophic processes; the sediment is only locally important.Mineralisation rates are highest in the turbidity region, but as only a fraction of total carbon resides here, less than 20% of all organic carbon is lost in this part of the estuary. This result is in contradiction with a previous budget of the estuary, based on data of the early seventies, where more than 80% of all carbon was estimated to be lost in the turbidity zone. Part of this discrepancy is probably caused by changes that have occurred in the estuary since that time.Due to the high heterotrophic activity, nearly all imported and in situ produced carbon is lost in the estuary itself and the Westerschelde is an insignificant source of organic matter to the coastal zone.The model estuary acts as a trap for reactive organic matter, both from the land, from the sea or in situ produced. Internal cycling, mainly in the water column, results in the removal of most of the carbon while the more refractory part is exported to the sea.     

Some reflections on the structure of the mandibleplate ofEurytemora affinis (Copepoda,Calanoida)

The mandibleplates of several specimen of the copepodEurytemora affinis collected from the Westerschelde and the Gironde estuary were studied. Using the Edge index (ITOH, 1970) this species could be classified as an omnivore. Pictures obtained by light and scanning electron microscopy reveal that the mandible teeth ofE. affinis have a molar-like structure, which likely changes in functional morphology with developmental stage, sex and age of the adults. Based on these observations, the desirability to consider the three dimensional structure of copepod mandibles for feeding mode classification is discussed.Communication no. 540 of the Delta Institute for Hydrobiological Research, Vierstraat 28, 4401 EA Yerseke, The Netherlands.     

On the cultivation of free-living marine and estuarine nematodes

Although a large body of literature exists on the systematics and ecology of free-living marine and brackish-water nematodes, key questions on the nature and magnitude of interactions between nematodes and other organisms in the benthos remain unanswered. Relatively few authors have investigated live nematodes in food web studies or in experiments dealing with the nematodes’ response to a varying environment. It is mainly for the latter purpose that attempts have been made to maintain, rear and cultivate selected species. This paper describes the methodology used for the maintenance, rearing, and eventual permanent agnotobiotic cultivation of a variety of estuarine nematodes. Spot plates, where small samples of sediment or macrophyte material are inoculated on a sloppy agar layer, have been used for the purpose of maintenance and initial cultivation. Those species that reproduce on spot plates are then selected for monospecific cultivation on agar layers with different nutrient enrichments and with micro-organisms cotransferred from the spot plates as food. Mixtures of bacto and nutrient agar prepared in artificial seawater were specifically suitable for the xenic cultivation of nine bacterivorous and, when supplied with Erdschreiber nutrients, two algivorous/bacterivorous nematode species. Up to three generations of five other nematode species have been reared under laboratory conditions, and several more were kept alive and active for variable periods of time on agar. Generation times observed on spot plates forAdoncholaimus fuscus andOncholaimus oxyuris were substantially shorter than previously published estimates and suggest a correspondingly higher predatory and scavenging potency for these and related enoplids. A procedure for the long-term storage of nematodes at −80°C with glycerol as a cryoprotectant was successfully used forDiplolaimella dievengatensis, Panagrolaimus sp. 1, andPellioditis marina, but not forDiplolaimelloides meyli. The authors have also summarized the existing literature on the cultivation of marine and brackish-water nematodes. Continuous cultivation appears to have been successful mainly for Aufwuchs and epiphytic nematodes; only few sediment-dwellers have been established in permanent culture. Of only just over 30 species that have ever been cultivated, more than half belong to one family (Monhysteridae) and three are Rhabditida, an order poorly represented in the marine environment. Four species have been grown in monoxenic and one in axenic culture, the latter though with limited success. It is concluded that our understanding of the basic nutritional requirements of marine nematodes is as yet insufficient, and that the culture techniques which have so far mainly deployed agar or liquid substrates, while being suitable for the cultivation of Aufwuchs and epiphytic nematodes, do not accurately enough mimic gradients specific of the natural habitat of many sediment-dwellers.     

Nitrogen dynamics in the Westerschelde estuary (SW Netherlands) estimated by means of the ecosystem model MOSES

A tentative nitrogen budget for the Westerschelde (SW Netherlands) is constructed by means of a simulation model with thirteen spatial compartments. Biochemical and chemical processes in the water column are dynamically modeled; fluxes of dissolved constituents across the water-bottom interface are expressed by means of diagenetic equations.The model is calibrated on a large amount of observed variables in the estuary (1980–1986) with relatively fine temporal and spatial detail. Additional constraints are imposed by the stoichiometric coupling of carbon, nitrogen and oxygen flows and the required conservation of mass. The model is able to reproduce rather well the observed distributions of nitrate, ammonium, oxygen and Kjeldahl nitrogen both in time and space. Also, model output of biochemical oxygen demand and total organic carbon falls within observed ranges.By far the most pervasive process in the nitrogen cycle of the estuary is nitrification which mainly takes place in the water column of the upper estuarine part. On average about three times as much nitrate is leaving the estuary at the sea side compared to what enters from the river and from waste discharges. Ammonium on the other hand is consumed much faster (nitrification) than it is regenerated and only about one third of the total import leaves the estuary at the sea side. The budget for detrital nitrogen reveals import from the river, from wastes and from the sea. Phytoplankton uptake of inorganic nitrogen is negligible in the model.About 21% of total nitrogen, 33% of inorganic nitrogen, is removed from the estuary (mainly to the atmosphere through denitrification) and the load of nitrogen net exported to the sea amounts to about 51 000 tonnes per year. Total denitrification in our model is lower than what was estimated in the literature from the late seventies, where a nitrogen removal up to 40–50% of the total inorganic load was reported. Part of the differences could be methodological, but inspection of the nutrient profiles that led to these conclusions show them to be different to the ones used in our study. The oxygen deficient zone has moved upstream since the late seventies, entrailing the zone of denitrification into the riverine part of the Schelde. The nitrification process now starts immediately upon entering the estuary.     

Nutrients,light and primary production by phytoplankton and microphytobenthos in the eutrophic,turbid Westerschelde estuary (The Netherlands)

Abiotic factors and primary production by phytoplankton and microphytobenthos was studied in the turbid Westeschelde estuary. Because of the high turbidity and high nutrient concentrations primary production by phytoplankton is light-limited. In the inner and central parts of the estuary maximum rates of primary production were therefore measured during the summer, whereas in the more marine part spring and autumn bloom were observed. Organic loading is high, causing near anaerobic conditions upstream in the river Schelde. Because of this there were no important phytoplankton grazers in this part of the estuary and hence the grazing pressure on phytoplankton was minimal. As this reduced losses, biomass is maximal in the river Schelde, despite the very low growth rates.On a number of occasions, primary production by benthic micro-algae on intertidal flats was studied. Comparison of their rates of primary production to phytoplankton production in the same period led to the conclusion that the contribution to total primary production by benthic algae was small. The main reason for this is that the photosynthetic activity declines rapidly after the flats emerged from the water. It is argued that CO₂-limitation could only be partially responsible for the noticed decrease in activity.     

Short-term fluctuations in benthic diatom numbers on an intertidal sandflat in the Westerschelde estuary (Zeeland,The Netherlands)

During the period March–May 1991, sediment samples were taken every two or three days at one intertidal station in the brackish part of the Westerschelde estuary. Quantitative cell counts were made in order to investigate the short-term temporal changes in diatom numbers and assemblage structure.Throughout the whole sampling period, the diatom assemblage was dominated by epipsammic diatoms. Three species, Achnanthes delicatula, Opephora cf. perminuta and Catenula adhaerens on average accounted for almost 67% of all valves counted. The epipsammic diatom fraction showed no significant changes in absolute numbers; its species composition appeared relatively stable. In contrast, epipelic diatom densities significantly increased towards the end of the study period. Species composition within this fraction was less stable. Multivariate analysis (Principal Components Analysis), in combination with multiple regression, indicated that total sky irradiance (on the second and third day preceding sampling) and percentage organic matter were related to the short-term fluctuations of the epipelic diatom fraction.     

Applicability of the abundance/biomass comparison method to detect pollution effects on intertidal macrobenthic communities

Data are presented to evaluate further the abundance/biomass comparison method for the detection of pollution disturbance in marine benthic communities. The results suggest that this technique cannot be used in estuarine intertidal communities. The grossly polluted configurations appeared under unpolluted conditions, while undisturbed patterns were found under polluted conditions. The results support similar conclusions by BEUKEMA (1988) for the intertidal flats of the Dutch Wadden Sea.Communication nr. 497 of the Delta Institute for Hydrobiological Research, Yerseke.     

Growth patterns of Mytilopsis leucophaeata,an invasive biofouling bivalve in Europe

Abstract

For the first time, growth of Mytilopsis leucophaeata, an important European fouling species, was investigated. By means of growth cages, individual shell growth of three cohorts, with, respectively, initial shell lengths of ≤5 mm, 10 mm and 15 mm, was monitored in the harbour of Antwerp, Belgium, during 2003 – 2004. M. leucophaeata followed an oscillatory growth pattern with a single summer growing period per year (May to August). Growth decreased during wintertime, but never ceased completely. M. leucophaeata has an average growth rate of <3 – 6 mm year^?1. Temperature was found to be the main environmental factor affecting growth. The von Bertalanffy growth function was used to model growth of individuals ≤5 mm, resulting in L_∞ = 16.7 mm and K = 0.56. Based on a combination of growth of all three cohorts, the hypothetical growth of an average individual mussel could be modelled over a 5-year period, resulting in a maximum length >19 mm with a growth rate of 0.41. Its longevity (more than 5 years) and the positive effect of higher water temperatures on growth, combined with its high resistance to chlorination, provides M. leucophaeata with a high potential for severe and long-lasting biofouling     

The salt marshes of the Westerschelde and their role in the estuarine ecosystem

The salt marshes along the Westerschelde estuary have been influenced by various human activities of which reclamation has been a major cause for the loss of salt marsh area. The salinity gradient in the aquatic system is also mirrored in the vegetation of the salt marshes.The role of the salt marshes for the estuary as a whole is manifold but a major importance is their function as a sink for anthropogenic substances.The possible role as a carbon and mineral source for the estuary is discussed in this paper. It is estimated that the total area of salt marsh adds about 8% to the organic matter input in the estuary while the nutrient input may be as high as 25%.Communication nr. 403 of the Delta Institute, Yerseke.