Biomass and abundance of macrofauna in intertidal sediments of Königshafen in the northern Wadden Sea

Intertidal sediments of Königshafen (Island of Sylt, North Sea) were sieved for mesofauna (>0.25 mm) and macrofauna (>1 mm) in spring and autumn 1990. Although sediments are coarser than in other parts of the Wadden Sea, the macrobenthic fauna was very similar but with a tendency towards higher species density, abundance and biomass. Taking into account the areal size of sandy flats, seagrass beds, mud flats and mussel beds, the average biomass is calculated to be 65 g ash-free dry weight m^?2 The lugwormArenicola marina dominates the biomass (28%), followed by the bivalvesMytilus edulis (21%),Mya areanaria (16%),Cerastoderma edule (10%) and the mudsnailHydrobia ulvae (9%). While spring and autumn biomass are almost alike, abundance is highly variable and entirely dominated byH. ulvae. Mesofauna is mainly composed of oligochaetes, small and juvenile polychaetes. Abundance is similar to that of macrofauna, while biomass is only about 1 g m^?2. Macrophyte biomass amounted to 9% of that macrofauna. In the course of the centurym mussel beds expanded while muddy areas declined. The concomitant effects on biomass presumably compensated each other.     

Nutrient fluxes in intertidal communities of a South European lagoon (Ria Formosa) – similarities and differences with a northern Wadden Sea bay (Sylt-Rømø Bay)

During an annual cycle, flux rates of oxygen, nitrate, nitrite, ammonium, phosphate and silicate were measured in light and dark bell jars at three sites in Ria Formosa (Algarve, Portugal) enclosing either a natural macrophytic community (macroalgae on sand or mud, a seagrass bed of Zostera noltii) or bare sediments. The results are compared with a preceeding study in which the same bell jar technique has been applied in the Sylt-Rømø Bay of the northern Wadden Sea. Nitrate flux was mainly directed from the water column to the benthic communities in Ria Formosa, as well as in the Sylt-Rømø Bay. However, nitrate uptake was higher in the northern, more eutrophic study area. In Ria Formosa, nutrient concentrations were lower than in the Sylt-Rømø Bay possibly due to strong water exchange with Atlantic waters. High temperatures and strong insolation had a greater impact on nitrate fluxes in Ria Formosa than in the Sylt-Rømø Bay. Bioturbating macrofauna increased ammonium efflux in the Sylt- Rømø Bay while this effect was not as pronounced in the Ria Formosa study sites. Benthic phosphate uptake dominated in the Ria Formosa and was correlated to initial phosphate concentrations in incoming waters. At both study sites, oxygen and nutrient fluxes were correlated with temperature. Additionally, flux rates were strongly influenced by biotic components and levels of eutrophication. A literature survey showed that mainly in temperate regions, material fluxes increase with temperature, whereas in warmer areas, ammonium and phosphate fluxes between sediment and water were generally lower.     

Microbial–meiofaunal interrelationships in coastal sediments of the Red Sea

Population density and biomass of bacteria and meiofauna were investigated seasonally in the sediments of the north-western bank of Red Sea. Samples of sediments were collected seasonally from three different stations to determine microphytobenthic biomass (chlorophyll a), protein, lipid, carbohydrate, and total organic matter concentrations. These investigations revealed that microbial components tended to increase their dominancy, whereas sensitive meiofauna were extremely reduced during the entire study period. Thus a very low density of the total meiofauna (with an annual average of 109 ± 26 ind./10 cm²) was recorded whilst the benthic microbial population densities exhibited higher values (ranging from 0.31 ± 0.02 × 10⁸ to 43.67 ± 18.62 × 10⁸/g dry sediment). These changes in the relative importance analysis of benthic microbial components versus meiofaunal ones seem to be based on the impact of organic matter accumulation on the function and structure of these benthic communities. Proteins, lipids and carbohydrates showed very low concentration values, and the organic matter mostly consisted of carbohydrates, reflecting lower nutritional values for benthic fauna in general and meiofauna in particular. The distribution of microbial and meiofaunal communities seems to be dependent on the quality of the organic matter rather than on its quantity. Total organic matter concentrations varied between 5.8 and 7.6 mg/g, with organic carbon accounting for only 32% of the total organic matter. Chlorophyll a attained very low values, fluctuating between 0.11 and 0.56 μg/g, indicating the oligotrophy of the studied area. The very low concentration of chlorophyll a in the Red Sea sediment suggests that the sedimentary organic matter, heterotrophic bacteria and/or protozoa constitute an alternative resource that is consumed by meiofauna when algae are less abundant. Protozoa, therefore, represent the “missing link in bacteria–meiofauna interaction in the Red Sea marine sediment ecosystem.     

Material exchange and food web of seagrass beds in the Sylt-Rømø Bight: how significant are community changes at the ecosystem level?

Material exchange, biodiversity and trophic transfer within the food web were investigated in two different types of intertidal seagrass beds: a sheltered, dense Zostera marina bed and a more exposed, sparse Z. noltii bed, in the Northern Wadden Sea. Both types of Zostera beds show a seasonal development of above-ground biomass, and therefore measurements were carried out during the vegetation period in summer. The exchange of particles and nutrients between seagrass beds and the overlying water was measured directly using an in situ flume. Particle sedimentation [carbon (C), nitrogen (N) and phosphorus (P) constituents] from the water column prevailed in dense seagrass beds. In the sheltered, dense seagrass bed, a net particle uptake was found even on windy days (7–8 Beaufort). Dissolved inorganic N and orthophosphate were mainly taken up by the dense seagrass bed. At times of strong winds, nutrients were released from the benthic community to tidal waters. In a budget calculation of total N and total P, the dense seagrass beds were characterised as a material sink. The seagrass beds with sparse Z. noltii were a source of particles even during calm weather. The uptake of dissolved inorganic N in the sparse seagrass bed was low but significant, while the uptake of inorganic phosphate and silicate by seagrasses and their epiphytes was exceeded by release processes from the sediment into the overlying water. Estimates at the ecosystem level showed that material fluxes of seagrass beds in the Sylt-Rømø Bight are dominated by the dense type of Zostera beds. Therefore, seagrass beds act as a sink for particles and for dissolved inorganic nutrients. During storms, seagrass beds are distinct sources for inorganic nutrients. The total intertidal area of the Sylt-Rømø Bight could be described as a sink for particles and a source for dissolved nutrients. This balance of the material budget was estimated by either including or excluding seagrass beds. Including the subtidal part, the function of the ecosystem as a source for particles increased, supposing that all seagrass beds were lost from the area. During the vegetation period, seagrass beds act as a storage compartment for material accumulated in the living biomass of the community. There was great biodiversity among the plant and animal groups found in intertidal seagrass beds of the Sylt-Rømø Bay, representing 50–86% of the total number of species investigated, depending on the particular group. Since most species are not exclusively seagrass residents, the loss of intertidal seagrass beds would be of minor importance for biodiversity at the ecosystem level. Food web structure in seagrass beds is different from other intertidal communities. Primary production and detritus input is high, but secondary production is similar to that of unvegetated areas, although the relative importance of the trophic guilds is different. The loss of seagrass beds leads to profound alterations in the food web of the total ecosystem. Historical as well as recent changes in material fluxes and energy flow due to man-made alterations to the ecosystem are discussed.     

Effects of the severe winter 1995/96 on the biological oceanography of the Sylt-Rømø tidal basin

Water temperature, salinity and precipitation, micronutrients (N, P, Si) and chlorophyll a concentrations in the Sylt-R?m? tidal basin (German Bight) deviated between the early 1990s, with mild winters, and the years 1996–1997 with a severe winter and a moderate winter. As a consequence of low temperature, offshore winds and low precipitation in the severe winter 1995/96, salinity peaked in February 1996 and nitrate concentrations were low. The latter further decreased in March as chlorophyll a peaked with an exceptional bloom of the diatom Odontella aurita, probably triggered by low water temperature. Winter temperatures and spring chlorophyll a in the Sylt-R?m? tidal basin correlate well with the climatic North Atlantic Oscillation index. Received in revised form: 7 May 2001 Electronic Publication     

Temporal and spatial variations of biodeposits—A preliminary investigation of the role of fecal pellets in the Sylt-Rømø tidal area

The fecal pellets ofHeteromastus filiformis andHydrobia ulvae are abundant on the sediment surface of the Sylt-Rømø tidal flats. AlthoughCerastoderma edule is present in most parts of the study area, only small amounts of its pellets could be found. Fecal pellets are non-randomly distributed in the study area. Large amounts ofH. ulvae pellets are found in the mud flats and smaller amounts in the mised flats and sand flats, whereH. filiformis pellets predominate as biogenic component. This distribution pattern varies with time under the influence of environmental factors. Marked temperature fluctuations provide seasonal signals reflecting distinct patterns. The results show a net balance of biodeposit components in the sediment. This could be a result of the sheltered location of the area and this fits in well with the mostly stable sedimentological picture.     

Microbial community structure across the tree of life in the extreme Río Tinto

Understanding biotic versus abiotic forces that shape community structure is a fundamental aim of microbial ecology. The acidic and heavy metal extreme Río Tinto (RT) in southwestern Spain provides a rare opportunity to conduct an ecosystem-wide biodiversity inventory at the level of all three domains of life, because diversity there is low and almost exclusively microbial. Despite improvements in high-throughput DNA sequencing, environmental biodiversity studies that use molecular metrics and consider entire ecosystems are rare. These studies can be prohibitively expensive if domains are considered separately, and differences in copy number of eukaryotic ribosomal RNA genes can bias estimates of relative abundances of phylotypes recovered. In this study we have overcome these barriers (1) by targeting all three domains in a single polymerase chain reaction amplification and (2) by using a replicated sampling design that allows for incidence-based methods to extract measures of richness and carry out downstream analyses that address community structuring effects. Our work showed that combined bacterial and archaeal richness is an order of magnitude higher than eukaryotic richness. We also found that eukaryotic richness was highest at the most extreme sites, whereas combined bacterial and archaeal richness was highest at less extreme sites. Quantitative community phylogenetics showed abiotic forces to be primarily responsible for shaping the RT community structure. Canonical correspondence analysis revealed co-occurrence of obligate symbionts and their putative hosts that may contribute to biotic forces shaping community structure and may further provide a possible mechanism for persistence of certain low-abundance bacteria encountered in the RT.     

Microbial community composition of anoxic marine sediments in the Bay of Cádiz (Spain)

The composition of the microbial community inhabiting the anoxic coastal sediments of the Bay of Cádiz (southern Spain) was investigated using a molecular approach consisting of PCR cloning and denaturing gradient gel electrophoresis (DGGE), based on 16S rRNA sequences. The total cell count was 1-5 × 10? cells/g sediment and, as determined by catalyzed reporter deposition-fluorescent in situ hybridization (CARD-FISH), the proportion of Bacteria to Archaea was about 70:30. The analysis of 16S-rRNA gene sequences revealed a wide spectrum of microorganisms, which could be grouped into 111 operational taxonomic units (OTUs). Many of the OTUs showed high phylogenetic similarity to microorganisms living in marine sediments of diverse geographic origin. The phylogenetic groups that were predominantly detected were Firmicutes, Deltaproteobacteria, and Gammaproteobacteria, accounting for 23, 15, and 14% of the clones, respectively. Diversity in the domain Archaea was significantly lower than in the domain Bacteria. The majority of the archaeal OTUs belonged to the Crenarchaeota phylum. Since most of the sequences could not be identified precisely at the genus/species level, the functional roles of the microorganisms in the ecosystem could not be inferred. However, seven OTUs affiliated with the Delta- and Epsilonproteobacteria were identified down to the genus level, with all of the identified genera known to occur in sulfate-rich marine environments.     

Microphytobenthic dynamics in a Wadden Sea intertidal flat – Part II: Seasonal and spatial variability of non-diatom community components in relation to abiotic parameters

The microphytobenthos colonizing the intertidal flats forms an important component of the Wadden Sea. Ten sampling points along a 1-km transect were studied in a fringe area of the Solthörn tidal flat, southern North Sea, in order to determine seasonal differences in the microphytobenthos. An accompanying paper deals with the major component of the flora, the diatoms; here we, focus on the minor taxonomic groups. From May 2008 to May 2009 surface sediments were collected during low tide. Variation of environmental factors as well as microphytobenthic density (abundance and chlorophyll a) were monitored. The area investigated was a mixed-sediment mudflat, with a gradient from coarse to fine. Highest biomass was obtained in summer 2008 with 215.9?±?12.6?mg chlorophyll a m^–2. In late autumn the chlorophyll a concentration decreased continuously at all investigated stations. Lowest values were detected in December 2008. Species abundances varied considerably both along the transect and seasonally, depending on species-specific requirements as well as hydrodynamic conditions (tidal currents). Higher densities of benthic pro- and eukaryotic microalgae were observed in sites characterized by fine sediments. Apart from the diatoms, the most abundant microphytobenthic group was the cyanophytes. Coccoid cyanophytes, mainly Merismopedia sp., were most abundant during summer, with cell numbers up to 5.72?×?10⁶ cells cm^?2, while diatoms dominated in winter, spring and autumn. Filamentous cyanophytes, particularly Microcoleus chthonoplastes, were most abundant during autumn, while coccoid chlorophytes (spring: Chlorococcum submarinum, Crucigenia tetrapedia, Tetraselmis suecica), euglenophytes (summer: Euglena obtusa), dinophytes (autumn: Amphidinium operculatum, A. herdmanii) and cryptophytes (autumn: Hillea marina, Hemiselmis virescens) contributed to the microphytobenthos during warmer seasons. The statistical analysis confirmed that the composition of the microphytobenthos was related to sediment features and to characteristics of particular seasons.     

The phosphorus budget of the Marsdiep tidal basin (Dutch Wadden Sea) in the period 1950–1985: importance of the exchange with the North Sea

Nutrient and carbon budgets of the western part of the Dutch Wadden Sea are based on the concept formulated by Postma (1954): import of organic compounds from the North Sea, mineralization in the estuary ans subsequently an export of dissolved inorganic nutrients to the North Sea. In this paper the phosphorus budget of the westernmost part of the Wadden Sea (Marsdiep basin) during the period 1950–1985 is considered to evaluate this concept and to investigate whether eutrophication processes have changed these P-budgets during the last decades. Most attention is paid to the importance of the exchange of P-components between the tidal basin and the open sea. The budgets are based on a general mass balance equation in which the main entries are the loadings via the freshwater sources, the net sediment-water exchange, the exchange with the North Sea, the input from the adjacent Vlie basin, and a transformation term representing the interchange between dissolved and particulate phosphorus. Averaged over the entire period 55% of the phosphate input (SRP) to the basin is delivered by the fresh water sources, and 45% by the Vlie basin. For non-SRP components these numbers are 50% and 40% respectively, while only 10% comes from the North Sea. The import from the North Sea, as calculated by our model thus seems less important than previously documented. There is a clear effect of the eutrophication during the last decades. In the first 20 years of the period investigated there was a moderate increase in the TP input to the basin, partly caused by an increasing input of non-SRP (TP minus soluble reactive phosphorus) from the North Sea. In 1970/71 almost 30% of the non-SRP loading originated from the North Sea. From approximately 1975 onwards this import of non-SRP turns into an export, while the TP-loading from the other sources inclined, until a maximum was reached in 1981. Recently a decrease in the TP-inputs has been observed. It is concluded that the recent output of phosphorus from the Marsdiep basin may have a considerable impact on the adjacent area of the North Sea.Publication no. 2 of the project Applied Scientific Research Neth Inst. for Sea Res. (BEWON).     

The structure of the plankton community of the Öregrundsgrepen (southwest Bothnian Sea)

Taxonomic composition and variations in density and biomass of the plankton community in the Öregrundsgrepen, a shallow coastal area, were investigated from June 1972 to November 1973. The phytoplankton biomass was large in spring but small during the rest of the year. The spring bloom was dominated by diatoms and dinoflagellates, especially byThalassiosira spp. which were also important during other seasons. Small forms, such asCryptomonas spp.,Rhodomonas spp. and monads, dominated during summer. Blue-green algae were never of any major importance. During the summer, the trophogenic layer exceeded 10 m in thickness. The metazoan fauna was of lower diversity than the plankton flora. The dominating species, the copepodsAcartia bifilosa andEurytemora affinis, constituted on the average 83% of the standing crop. The low salinities, 5–6 S, were regarded as the principal pertinent limiting factor. The metazoan fauna reached large biomass values from July to October. The protozoan fauna (in the case of ciliates), obtained biomass maxima during the spring bloom. It is suggested that the Öregrundsgrepen represents an area of elevated productivity within a region of low overall production, presumably due to local upwelling. From June 1972 to May 1973, the average biomasses were: phytoplankton 0.464 g C m^–2, ciliates 0.040 g C m^–2, copepod nauplii 0.010 g C m^–2, micro-rotifers 0.004 g C m^–2, and mesozooplankton (larger than 0.2 mm) 0.312 g C m^–2. It is estimated that about than 60% of the phytoplankton production is consumed by the microzooplankton (<0.2 mm).     

Juvenile-adult distribution of the bivalve Mya arenaria on intertidal flats in the Wadden Sea: why are there so few year classes?

Patchy distribution is frequently observed in benthic marine invertebrates. In order to indentify factors causing spatial patterns in the bivalve Mya arenaria, abundances of juveniles and adults, as well as death assemblages, were recorded on a 20-km scale in the intertidal zone of the Sylt-Rømø Bight. Both adults and juveniles exhibited pronounced patchiness. Shell length of juveniles rarely exceeded 2 mm in 1995, which was most likely a consequence of epibenthic predators truncating the size spectrum. Only a few yearclasses dominated the adult population. While the northern part of the Bight was colonized mainly by a 1993-cohort, most M. arenaria in the southern part were from the mid-1980s. It is hypothesized that epibenthic predation is a major cause of the lack of dense M. arenaria beds from other years. However, examination of the length–frequency distribution of death assemblages revealed that other unidentified causes of mortality exist. High abundances of adults were found in the mid and lower intertidal zone but not in the high intertidal zone. There was no indication that dispersal of M. arenaria spat in a landward or seaward direction contributed significantly to the observed distribution pattern, since spat occurred abundantly at all tidal levels except in the high intertidal zone. There was no evidence of negative adult-juvenile interaction. M. arenaria was not attracted by seagrass or projecting shell beds – the latter indicating erosion of the sediment – as abundances of adults and juveniles were generally low in these habitats. The effects of sediment type and of the bioturbating lugworm Arenicola marina were inconsistent. While adults were more abundant on muddy sand than on sand, recruitment was independent of sediment type. At all high density sites of adults (>50 ind m^–2), lugworm densities were below 5 ind m^–2, which may indicate a negative interaction. However, lugworm densities >30 ind m^–2 did not prevent relatively high M. arenaria recruitment of >500 ind m^–2.     

Effect of the Razdol’naya River on structure of the free-living nematode community of the Amursky Bay,Sea of Japan

The effect of the Razdol’naya River on the littoral and sublittoral nematode community was studied in the innermost part of the Amursky Bay. Thanks to the seasonal abundance of juveniles of macrobenthic animals, pseudomacrobenthic species prevailed in the meiobenthos littoral community. Juvenile amphipods dominated (39.5%), the percentage of nematodes was 18.4% (45 species). The nematodes dominated in the meiobenthos community (65.7%, 71 species) at the sublittoral zone. The similarity of the species structure of littoral and sublittoral nematodes reached 36.5%. Low population density (2.6±1.7 thousand ind/m²) and poor species diversity of nematodes (7 species), the dominating form being Oncholaimium paraolium, were recorded at the littoral zone, which was subjected to direct riverine influence (1‰salinity). The correlation analysis revealed no relationship between salinity and population density of nematodes at the littoral zone. At the sublittoral zone (at stations of salinity less than 2.85‰) the parameters of population density (73.9±32.3 thousand ind/m²) and species diversity (12 species) of nematodes were the lowest, Parodontophora timmica dominated. A poor expressed dependence between the density of nematode populations and salinity was revealed. The correlation between the average size of ground particles and population density of nematodes was recorded. The greatest similarity of the species structure of littoral and sublittoral nematodes (26.67%) was recorded on gravel-sand bottoms.     

Monitoring in the western part of the Dutch Wadden: Sea—sea level and morphology

This paper deals with the methods and results of monitoring sea level (tidal gauges) and with the analysis of depth sounding data. Possible future monitoring by menas of remote sensing techniques will be presented. Some trends, based on water level and morphological monitoring, have been established in the western part of the Dutch Wadden sea: (a) the relative mean sea level is rising persistently by 15 cm/century, observed since the middle of the 19th century. The tidal range is increasing as well; (b) the cross-sectional areas of the most western Dutch Wadden tidal inlets have been increasing over the last two centuries; (c) the intertidal zones of the Texelstroom and Vliestroom tidal basins have shown an increase of surface area over the last decades. The Borndiep tidal basin shows a decrease of 4% of the area shallower than 5 m — Dutch Ordinance Level (DOL) over the last two centuries. Several factors prevent serious predictions about future morphological developments of the tidal Wadden flats: (a) man induced interfering factors, e.g. the construction of the harbour revetments of Den Helder, the construction of the Enclosure Dike and construction of sand drift dikes stabilising the Wadden islands; (b) the limited scope of the present analysis that deals only with the western part of the Dutch Wadden Sea. Presented at the VI International Wadden Sea Symposium (Biologische Anstalt Helgoland, Wattenmeerstation Sylt, D-2282 List, FRG, 1–4 November 1988)     

Biological monitoring of fish and crustaceans in the Wadden Sea — potential and problems

Monitoring of fish and crustaceans in the Wadden Sea (WS) must cope with rapid changes in distribution patterns, access to certain areas and gear efficiency. Application and limitations of a variety of fishing devices (fyke nets, gill nets, enclosures, stow nets, purse seines, beam trawls, push nets, beach seines, bottom trawls, pelagic trawls) are discussed with regard to different objectives of monitoring. Furthermore, the validity of data from three current monitoring programmes is also discussed. Presented at the VI International Wadden Sea Symposium (Biologische Anstalt Helgoland, Wattenmeerstation Sylt, D-2282 List, FRG, 1–4 November 1988)     

Microbial Community Structure Associated with Biogeochemical Processes in the Sulfate–Methane Transition Zone (SMTZ) of Gas-hydrate-bearing Sediment of the Ulleung Basin,East Sea

We investigated the biogeochemical constituents, microbial communities and functional genes (mcr and dsr) associated with anaerobic methane oxidation and sulfate reduction, and metabolic activities by sulfate reduction in the sulfate–methane transition zone (SMTZ) of gas-hydrate-bearing sediment of the Ulleung Basin in the East Sea. Maxima in the sulfate reduction rate (12.6 nmol cm^?3 d^?1), CO concentration (83 μM), and gene abundances of dsrA (9.1 × 10⁶ copies cm^?3) and mcrA (11.6 × 10⁶ copies cm^?3) occurred in the SMTZ. The peaks of CO consistently found in the SMTZ suggested that CO is an intermediate metabolic product related to methane oxidation. Candidate division JS1, the predominant bacterial group that comprised 59.0–63.7% of the 16S rRNA gene sequences, was recognized as an important organic carbon oxidizer. Both Marine Benthic Group D (MBGD) and Marine Benthic Group B (MBGB), which constituted 40.8–52.9 and 10.3–43.9% of the 16S rRNA gene sequences, respectively, were the dominant archaeal groups. Analysis of functional gene diversity revealed that anaerobic methanotroph-1-related phylotypes appeared to be the major CH₄ oxidizer, whereas Firmicutes-like group was a predominant sulfate reducer in the 0.8 mbsf in SMTZ with low SO₄^2? concentration. Overall results indicated that JS1 and two archaeal groups (MBGB and MBGD) seem to play a significant role in carbon and elements cycles in the gas-hydrate-bearing subsurface sediment of the Ulleung Basin.