The soil environment of the intertidal area in the Westerschelde

Hydrodynamic forces and sediment discharges determine the sedimentary environment and surface morphology of the intertidal area in the Westerschelde estuary in the S.W. Netherlands. Sandflats (clay content <8%) are found in the central part, mudflats (>8% clay) and especially salt marshes (>15% clay) occur in sheltered places.The semi-terrestrial soil of the intertidal area is a complex environment where hydrosphere, biosphere, lithosphere, and atmosphere interact. Surface morphology and the semi-diurnal tide may significantly modify this interaction pattern. Organic matter and constituents of the clay-size fraction are the most reactive compounds in the soil.The pathway and rate of organic matter decomposition determines the nature of many biogeochemical processes. Sulfur transformations are dominant processes in the sulfidic grey-black coloured horizons underneath the generally thin yellow-brown surface layer. The sulfur geochemistry also affects the sorption-behaviour of metals. Oxygen intrusion by diffusion, burrowing fauna and plant roots cause a local and sometimes temporary oxidation of reduced inorganic compounds. Thus, in the surface layers, and in and around tubes and roots at greater depth in the soil, a dynamic cycling occurs of redox active compounds, which affects the sorption mechanism ofe.g. metals-ions.     

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.     

Scale-dependent patterns of macrofaunal distribution in soft-sediment intertidal habitats along a large-scale estuarine gradient

We investigated the pattern of distribution of intertidal soft-bottom fauna in streams and lagoons of the Uruguayan coast at three spatial scales. The Río de la Plata and the Atlantic Ocean produce on this coast a large-scale gradient in salinity, defining a freshwater (west), an estuarine (central) and a marine (east) region. Within each region, there are several streams and coastal lagoons (sites) that define a second scale of variability. A third scale is given by intertidal gradients within each site. Species richness and total abundance was low in the freshwater west region and high in the central and east regions. The community in the west region was characterized by the clam Curbicula fluminea; in the other regions, it was dominated mainly by the polychaete Heteromastus similis. The polychaete Nephtys fluviatilis was more abundant in the east region, while another polychaete, Laeonereis acuta, characterized the central region. Sediment fractions did not vary significantly at this scale. At the scale of the sites, species richness and total macrofaunal abundance were higher in coastal lagoons than in streams. Coarse sands were more common in coastal lagoons, while medium and fine sand characterized the sediment in streams. Within each site, species richness and total abundance increased towards the lower intertidal level; the macrofauna of the upper levels were a subsample of the fauna occurring at the lower levels. There was also a significantly lower proportion of fine sand at the upper level. At regional scales, the observed patterns may be indirectly or directly related to the gradient in salinity, through differential physiological tolerance to osmotic stress. At the scale of the sites, variability may be explained mainly by geomorphological and sedimentological differences between lagoons and streams. Variation among levels may be related to gradients in desiccation, colonization and predation.     

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.     

Habitat segregation and temporal variation in some tropical intertidal populations

Observations were made to document habitat segregation and seasonal changes in density for the most common intertidal organisms at a study site on the Pacific coast of Costa Rica. The pulmonate limpets Siphonaria gigas (Sowerby) and Siphonaria maura (Sowerby) were more abundant in the mid-high- and high-intertidal zone, whereas the keyhole limpet Fissurella virescens (Sowerby) was more abundant in the mid zone. Coiled gastropods were found in the mid- and mid-high-intertidal zone but were absent or scarce in the high zone. Barnacles and coralline algae were found throughout the intertidal zone and foliose algae were mostly present in the splash zone. Abundance of all species changed over the 2-yr period of the study. S. gigas and S. maura and the barnacle Chthamalus fissus Darwin exhibited well-defined fluctuations in density related to recruitment. No recruitment was observed for F. virescens or Tetraclita panamensis Pilsbry and populations of both species declined. Changes also occurred for coiled gastropods and were probably related to movement of adults. Coralline and foliose algae were more abundant during the rainy season than during the dry months. Far from being constant, populations on these tropical shores were highly dynamic with temporal changes rivaling those in the temperate zone.     

Spatial and temporal variation in the sediment properties of an intertidal mangrove forest: implications for sampling

Studies of the structure or functioning of intertidal soft sediments often involves collection of biogeochemical data over tidal, diel and seasonal time-scales. Little effort has, however, been made to quantify accurately the time-scales at which these properties vary. Many previous studies collected samples from different sites at different times, potentially resulting in the confounding of spatial and temporal variation. This experiment was designed to determine if time of day or time within the exposure period had any significant effect on measurements of 7 different properties of sediments. Samples of sediment were collected using contact cores at the beginning, middle and end of tidal emersion at each of 9 a.m., 12 p.m. and 3 p.m., with two replicate days of each condition, from mangrove forests fringing Glades Bay, Sydney, Australia. These samples were analysed for water-content, pigments, carbohydrates, grain-size and loss on ignition (LOI), to determine the potential effects of time of day and time within the tidal cycle on these properties of the sediments.Whilst both time of day and time within the tidal cycle were found to occasionally have a significant effect upon the measured properties, most of the variation occurred among sites and between replicate days of each set of conditions. The minimal influence of time of day and time within the tidal cycle show that sampling effort should be preferentially placed into replicating days of sampling and sites because these are the scales with the greatest variation. Differences in the patterns found also depended on whether the data were expressed as content or concentration, the consequences of which are briefly described.     

The effects of tidally-driven temporal variation on measuring intertidal cohesive sediment erosion threshold

Accurate measurement of intertidal sediment erodibility is essential for the development of meaningful and accurate models of sediment dynamics. Despite considerable advances in technology and methodology, the measurement of cohesive intertidal sediment erosion remains problematic. Sediment erodibility varies according to both physical and biological properties and processes. These cannot be considered in isolation, as they can interact to create both positive and negative feedbacks, resulting in seemingly idiosyncratic responses in the system. If working models of estuarine sediment dynamics are to be made, it is essential that the influence of these processes on both the measurements and the system itself be considered. Recent developments in measurement technology enable rapid measurement of sediment stability allowing temporal and spatial variability to be measured on a time scale of minutes. This paper reports temporal variability in cohesive intertidal sediment erosion threshold related to immersion and emersion, and the concomitant responses in selected sediment properties (carbohydrates, water content and chlorophyll). Erosion threshold tended to increase over emersion and decrease over immersion, although the patterns of change varied depending upon local conditions, and in one case there was no temporal trend. Temporal changes resulted in a range of measured erosion threshold, dependant upon the erosion device used. Modifications to existing methodology, in order to account for this variation, are proposed and implications for modelling erosion processes are considered.     

A functional and morphological approach to evaluate the vertical migration of estuarine intertidal nematodes during a tidal cycle

We tested herein the hypothesis that exposure time significantly contributes to the vertical distribution profile of nematodes during a tidal cycle as a function of distinct feeding and locomotion behaviors, conditioned by body morphology. We showed that the vertical distribution profile of the slender with filiform tail, numerically dominant Terschellingia longicaudata is in fact significantly correlated with sediment changes induced by tidal variation. Conversely, none of the other nematode species showed unequivocal evidence of vertical migration. Horizontal spatial heterogeneity also influenced the vertical distribution of nematode associations, probably as a response to varying temperature and desiccation levels at the sediment surface. The resulting vertical profiles for individual or species groups are a trade-off among locomotory and feeding strategies and concordant morphological adaptations.     

Seasonal composition and spatial distribution of hyperbenthic communities along estuarine gradients in the Westerschelde

The hyperbenthic fauna of the Westerschelde estuary was sampled in spring, summer and winter of 1990 at 14 stations along the salinity gradient. Mysids dominated the hyperbenthos in each season. Other important species, either permanently (e.g. amphipods and isopods) or temporarily (e.g. fish larvae and decapod larvae) hyperbenthic, belong to a variety of faunistic groups. Spatial structure was stable through time: the estuary could be divided in the same geographically defined zones in each season. Each zone had a characteristic fauna. Gradients in salinity, dissolved oxygen and turbidity correlated strongly with the observed variation in community structure. The spatial patterns dominated over the temporal patterns, especially in the brackish part of the estuary. In the marine part, seasonal differences in the communities were more pronounced due to the occurrence of a series of temporary hyperbenthic species in spring and summer. In each season, the upstream (brackish) communities were characterized by few species occurring in very high numbers, whereas the downstream (marine) communities were composed of many species but at lower densities.     

Rocky intertidal fish communities of California: temporal and spatial variation

Synopsis We examined data from our own and published collections of intertidal and shallow littoral fishes of the North American Pacific Coast with respect to temporal and spatial trends in species composition and dominance. We compared (1) recent and past intertidal collections made five and seven years apart, respectively, for two California localities, (2) intertidal collections from twelve localities in California, Oregon, and British Columbia, and (3) intertidal versus subtidal collections at one California locality. Temporal comparisons indicated substantially lower abundance of the cottid Oligocottus snyderi at two California localities during 1984, at one locality due to depressed abundance of young. Interannual differences in abundances suggested that other tidepool fish assemblages undergo significant changes as well. Geographical comparisons indicated general similarity in species composition, with cottids predominating in tidepools although several other families also were well represented (e.g., Stichaeidae, Scorpaenidae). Eel-shaped stichaeids and pholids occurred at high densities in exposed boulder fields. Both the tidepool and boulder field assemblages showed north-south changes in species abundances. Comparison of collections from the intertidal and subtidal zones at one California locality demonstrated that fishes of these habitats form two essentially distinct assemblages, with most species restricted to or concentrated in one or the other habitat.     

Spatial and temporal variability of early post-settlement survivorship and growth in the barnacle Balanus glandula along an estuarine gradient

The performance of a species can be significantly altered by subtle changes in the physical environmental. The intertidal barnacle Balanus glandula is predominantly an open coast species in the Northeast Pacific. However, B. glandula commonly inhabits estuaries where environmental conditions such as salinity and temperature drastically differ from the open coast. We used survivorship and growth rates as a measure of performance in recently metamorphosed laboratory reared juvenile B. glandula outplanted along an environmental gradient at the mouth, mid-estuarine, and riverine end of the South Slough Estuary, Oregon, USA. Juvenile performance was highly variable over spatial and temporal scales and dependent upon existing environmental conditions. Surprisingly, along this estuarine gradient, juveniles performed better at a mid-estuarine location than at the mouth of the estuary. Typically, the riverine end of the estuary was the least suitable habitat along the estuarine gradient due to high juvenile mortality and a low growth rate. Although seasonally variable, survivorship and growth decreased with height along a vertical intertidal gradient as well. In a reciprocal transplant experiment, populations from both ends of the estuarine gradient displayed similar survivorship and growth rates. Our results demonstrate that the interactions of environmental conditions that vary temporally and spatially along a gradient strongly affect the success of an individual surviving and prospering during the early juvenile period.     

Heritable variation in resistance to gastro-intestinal nematodes in an unmanaged mammal population.

The impact of parasites on natural populations has received considerable attention from evolutionary biologists in recent years. Central to a number of theoretical developments during this period is the assumption of additive genetic variation in resistance to parasites. However, very few studies have estimated the heritability of parasite resistance under field conditions, and those that have are mainly restricted to birds and their ectoparasites. In this paper, to our knowledge, we show for the first time in a free-ranging mammal population, Soay sheep (Ovis aries) living on the islands of St Kilda, that there is significant heritable variation in resistance to gastrointestinal nematodes. This result is consistent with earlier studies on this population which have indicated locus-specific associations with parasite resistance. We discuss our results in the context of current studies examining heritable resistance to parasites in domestic sheep and the possible mechanisms of selective maintenance of genetic variation for resistance to gastrointestinal nematodes in the St Kilda Soay sheep population.     

Laboratory experiments on the infaunal activity of intertidal nematodes

The impact of oxygen on the vertical distribution of an intertidal nematode community was investigated in a manipulation experiment with sediments collected from the Oosterschelde (The Netherlands). The vertical distribution of nematodes was examined in response to sediment inversion in perspex cores with or without the presence of an air-flushed silicone tube introducing oxygen to the bottom sediment layer. Following an incubation of 1 week, peak densities of nematodes were recorded in the deep layers of the sediment in cores with subsurface oxygenation. In contrast, this concentration of nematodes was absent in the cores that lacked bottom oxygenation and the majority of the total assemblage was still located in anoxic sediment layers. This suggests that oxidised sediments with traces of oxygen represent favourable conditions for nematodes and indicates that these nematodes are tolerant to short-term anoxia and burial. Observations on the species level suggest that oxygenation, primarily through its direct impact on geochemical properties of the sediment, may be an important factor governing the subsurface activity of nematodes.     

Spatial and temporal variation in community composition and photosynthetic characteristics of phytoplankton in the upper Westerschelde estuary (Belgium,SW Netherlands)

Community composition, biomass and primary production of phytoplankton were studied in the east- ernmost section of the Westerschelde estuary in 1984. Photosynthetic characteristics were compared with distribution of some dominant phytoplankton species along a salinity gradient from 5 to 22 Spring phytoplankton, with Cyclotella meneghiniana (freshwater) and Skeletonema costatum (marine) as the dominant species grew faster than summer phytoplankton. In summer, biomass achieved its maximum, due to the riverine Scenedesmus species and the marine diatoms Thalassiosira levanderi and Ditylum brightwellii, as dominants. Growth conditions were more favourable to phytoplankton communities above 15%, than below this salinity. The data were compared with previous studies (1972) of species composition in the area.     

Benithic-pelagic coupling of nutrient metabolism along an estuarine eutrophication gradient

The shallow, brackish (11–18% salinity) Roskilde Fjord represents a eutrophication gradient with annual averages of chlorophyll, ranging from 3 to 25 mg chl a m^–3. Nutrient loadings in 1985 were 11.3–62.4 g N m^–2 yr^–1 and 0.4–7.3 g P m^–2 yr^–1. A simple one-layer advection-diffusion model was used to calculate mass balances for 7 boxes in the fjord. Net loss rates varied from –32.2 to 17.9 g P m^–2 yr^–1 and from –3.3 to 66.8 g N m^–2, corresponding to 74% of the external P-loading and 88% of the external N-loading to the entire estuary.Gross sedimentation rates measured by sediment traps were between 7 and 52 g p m^–2 yr^–1 and 50 and 426 g N M^–2 yr^–1, respectively. Exchangeable sediment phosphorus varied in annual average between 2.0 and 4.8 g P m^–2 and exchangeable sediment nitrogen varied from 1.9 to 33.1 g N m–1. Amplitudes in the exchangeable pools followed sedimentation peaks with delays corresponding to settling rates of 0.3 m d^–1. Short term nutrient exchange experiments performed in the laboratory with simultaneous measurements of sediment oxygen uptake showed a release pattern following the oxygen uptake, the changes in the exchangeable pools and the sedimentation peaks.The close benthic-pelagic coupling also exists for the denitrification with maxima during spring of 5 to 20 mmol N m^–2 d–1. Denitrification during the nitrogen-limited summer period suggests dependence on nitrification. Comparisons with denitrification from other shallow estuaries indicate a maximum for denitrification in estuaries of about 250 µmol N m^–2 h^–2 achieved at loading rates of about 25–125 g N m^–2 yr^–1.     

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.     

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.     

Temporal variation and succession in an algal-dominated high intertidal assemblage

We determined whether temporal variation and succession were similar among sites with similar species composition by sampling unmanipulated and cleared plots in a high intertidal assemblage dominated by Endocladia muricata and Mastocarpus papillatus. Sampling was done for 6 years at six sites spanning over 4° of latitude in California. Ten 1×2-m permanent plots were chosen in the central portion of the assemblage at each site. Four of these served as unmanipulated controls, three were cleared (scraped and burned) in the spring of 1985, and three were cleared in the fall of 1985. The cover of sessile and density of motile species were determined by subsampling within the plots from 1985 until 1991. Recovery of the clearings was determined by their similarity to the controls. The algae E. muricata, M. papillatus, and Fucus gardneri, and the barnacle Balanus glandula, were the most abundant sessile organisms in the control plots, although the latter never exceeded 12% cover at any site. The grazing gastropods Littorina scutulata/plena, various limpets, and Tegula funebralis were the most common mobile organisms. The species composition of the common species remained constant in the control plots over the study period and there were few large changes in relative abundance. Significant seasonal variation was detected in 11 species but variation was commonly site-specific.Ephemeral algae were abundant during early succession at only two of the six sites, and barnacle cover was low (<15% cover) at four sites and moderate (15-50% cover) at the remaining two throughout succession. Recovery rate varied considerably among sites and between times of clearing (1-10%/month). Correlations between ephemeral algae and grazer abundance, and between these variables and recovery rate were not significant. The effects of grazers on recovery rate were only evident at one site where they appeared to reduce an initially high cover of ephemeral algae and delay the establishment of perennials. Some of the largest differences in recovery rate were between clearing times, associated with differences in the phenology of the dominant perennial algae. In spite of these differences, most plots recovered by the end of the study period.These results indicate that the assemblages in the control plots at each site were relatively stable and, while successional pathways and processes varied, the assemblage at most sites still recovered. Current models, based largely on biological interactions, that attempt to explain within assemblage structure and succession were not broadly applicable.