Antibiotic production by a Roseobacter clade-affiliated species from the German Wadden Sea and its antagonistic effects on indigenous isolates

A strain affiliated with the Roseobacter clade and producing a new antibiotic named tropodithietic acid (L. Liang, Ph.D. thesis, University of G?ttingen, G?ttingen, Germany, 2003) was isolated from the German Wadden Sea. The compound showed strong inhibiting properties with respect to marine bacteria of various taxa and marine algae. Antibiotic production was found to occur during the complete growth phase. Strain mutants without antagonistic properties appeared several times spontaneously.     

Antibiotic Production by Actinomycete Species on Nitrogen-Deficient Media

Summary: Actinomycetes isolated from nitrogen deficient Azotobacter plating medium were checked for antibiotic production on nitrogen deficient media and on these media with yeast extract or ammonium sulphate as nitrogen sources. In nitrogen deficient liquid media cobalt, molybdenum and citrate were required for growth and antibiotic formation. Evidence is presented that the antibiotics resulting from growth under these conditions are different from those produced when combined nitrogen is included in the medium, and it is suggested that this is an area for exploration for new antibiotics.     

Distribution patterns of bacterioplankton and chlorophyll-a in the German Wadden Sea

 In a first synoptic evaluation, the temporal and spatial distribution of bacterioplankton and chlorophyll-a were determined in the German Wadden Sea. Three surveys were undertaken in winter, spring, and summer of 1994 using up to eight ships simultaneously between the river Ems and Sylt island. Despite intensive hydrodynamic mixing of the Wadden Sea water, spatial gradients were obvious. The abundance of bacterioplankton ranged from 0.4 to 26×10⁵ ml^–1 and chlorophyll-a varied between <0.1 and 79 μg l^–1. In winter, relatively homogeneous distribution patterns of both parameters with small gradients were found. Highest chlorophyll-a values connected with a highly patchy structure were observed in spring, while in summer both total chlorophyll-a values and the complexity of the distribution pattern had decreased. In contrast, bacterial numbers increased steadily from January to July with the highest bacterial densities and greatest patchiness observed in summer. Moreover, in some regions of the Wadden Sea, a trophic succession of algae as carbon producers and bacteria as consumers was evident. Correlation analysis verified the relationship between bacteria and chlorophyll a, indicating bottom-up control of bacterial abundance in the northern part of the German Wadden Sea. Since the observed regression slope is remarkably low (0.12–0.46) compared to literature values (0.5–0.8), we suggest that the link between phytoplankton and bacteria found here is a special characteristic of the Wadden Sea as a transition zone between the coastal region and the outer North Sea. Received: 26 May 1998 / Accepted:12 November 1998     

Tidal impact on planktonic primary and bacterial production in the German Wadden Sea

 Tidal variation of biological parameters was studied at three anchor stations in selected inlet channels of the northern German Wadden Sea in May and July 1994. Concentrations of bacteria, chlorophyll a and suspended matter as well as primary and bacterial production were assessed over a period of 25 h in the surface and in the bottom water. Diurnal variation in primary production was found both under in situ light conditions and under constant illumination. Tidal turbulence caused the introduction of detritus, bacteria and pigments from the sediment into the water column. The impact of sediment resuspension was most evident in the bottom water, leading to tidally oscillating bacterial production rates which were high during high stream velocity and low during the slack times. Estimations of the areal daily phytoplankton production and corresponding bacterial carbon demands were unbalanced. Primary production accounted for only 25–45% of the total bacterial carbon requirement. This discrepancy is due to the shallow euphotic depth in the Wadden Sea, allowing net primary production only in the upper 2–3 m of the water column, while the relatively high levels of bacterial activity do not show a vertical decline. Assuming that the specific biological activities in the water columns over the tidal flats are similar to those found in the inlet channels, it was found that production processes dominate in shallow areas whereas decomposition processes dominate in the deep channels. Moreover, the predominance of heterotrophic processes in the inlet channels means that additional organic carbon sources must contribute to the heterotrophic metabolism in the deep parts of the Wadden Sea, and that the horizontal flux of material is important in this turbid mesotidal ecosystem. Received: 18 April 1998 / Accepted: 12 November 1998     

Production of Antibacterial Compounds and Biofilm Formation by Roseobacter Species Are Influenced by Culture Conditions

Bacterial communities associated with marine algae are often dominated by members of the Roseobacter clade, and in the present study, we describe Roseobacter phenotypes that may provide this group of bacteria with selective advantages when colonizing this niche. Nine of 14 members of the Roseobacter clade, of which half were isolated from cultures of the dinoflagellate Pfiesteria piscicida, produced antibacterial compounds. Many non-Roseobacter marine bacteria were inhibited by sterile filtered supernatants of Silicibacter sp. TM1040 and Phaeobacter (formerly Roseobacter) strain 27-4, which had the highest production of antibacterial compound. In contrast, Roseobacter strains were susceptible only when exposed to concentrated compound. The production of antibacterial compound was influenced by the growth conditions, as production was most pronounced when bacteria were grown in liquid medium under static conditions. Under these conditions, Silicibacter sp. TM1040 cells attached to one another, forming rosettes, as has previously been reported for Phaeobacter 27-4. A spontaneous Phaeobacter 27-4 mutant unable to form rosettes was also defective in biofilm formation and the production of antibacterial compound, indicating a possible link between these phenotypes. Rosette formation was observed in 8 of 14 Roseobacter clade strains examined and was very pronounced under static growth in 5 of these strains. Attachment to surfaces and biofilm formation at the air-liquid interface by these five strains was greatly facilitated by growth conditions that favored rosette formation, and rosette-forming strains were 13 to 30 times more efficient in attaching to glass compared to strains under conditions where rosette formation was not pronounced. We hypothesize that the ability to produce antibacterial compounds that principally inhibit non-Roseobacter species, combined with an enhancement in biofilm formation, may give members of the Roseobacter clade a selective advantage and help to explain the dominance of members of this clade in association with marine algal microbiota.     

A Polyphasic Approach To Study the Diversity and Vertical Distribution of Sulfur-Oxidizing Thiomicrospira Species in Coastal Sediments of the German Wadden Sea

Recently, four Thiomicrospira strains were isolated from a coastal mud flat of the German Wadden Sea (T. Brinkhoff and G. Muyzer, Appl. Environ. Microbiol. 63:3789–3796, 1997). Here we describe the use of a polyphasic approach to investigate the functional role of these closely related bacteria. Microsensor measurements showed that there was oxygen penetration into the sediment to a depth of about 2.0 mm. The pH decreased from 8.15 in the overlaying water to a minimum value of 7.3 at a depth of 1.2 mm. Further down in the sediment the pH increased to about 7.8 and remained constant. Most-probable-number (MPN) counts of chemolithoautotrophic sulfur-oxidizing bacteria revealed nearly constant numbers along the vertical profile; the cell concentration ranged from 0.93 × 10⁵ to 9.3 × 10⁵ cells per g of sediment. A specific PCR was used to detect the presence of Thiomicrospira cells in the MPN count preparations and to determine their 16S rRNA sequences. The concentration of Thiomicrospira cells did not decrease with depth. It was found that Thiomicrospira strains were not dominant sulfur-oxidizing bacteria in this habitat. Denaturing gradient gel electrophoresis (DGGE) of PCR-amplified 16S ribosomal DNA fragments followed by hybridization analysis with a genus-specific oligonucleotide probe revealed the diversity of Thiomicrospira strains in the MPN cultures. Sequence analysis of the highest MPN dilutions in which the genus Thiomicrospira was detected revealed that there were four clusters of several closely related sequences. Only one of the 10 Thiomicrospira sequences retrieved was related to sequences of known isolates from the same habitat. Slot blot hybridization of rRNA isolated from different sediment layers showed that, in contrast to the concentration of Thiomicrospira cells, the concentration of Thiomicrospira-specific rRNA decreased rapidly in the region below the oxic layer of the sediment. This study revealed the enormous sequence diversity of closely related microorganisms present in one habitat, which so far has been found only by sequencing molecular isolates. In addition, it showed that most of the Thiomicrospira populations in the sediment studied were quiescent.     

Moderate predation on meiofauna by the macrobenthos of the Wadden Sea

The role of meiofauna in the trophic web of marine ecosystems is a controversial topic. During an experimental study on tidal flats in the eastern part of the North Sea, potential predators on meiofauna were kept in enclosures. A gobiid fish, shrimp, crabs, an amphipod, four polychaete worms and a nemertine were tested for their ability to decimate nematodes, turbellarians, ostracods and benthic copepods. Only few macrobenthic species preyed heavily on permanent meiofauna: juveniles of the shore crabCarcinus maenas, the hermit crabPagurus bernhardus with the colonial hydrozoanHydractinia echinata on its shells, and the rag wormNereis diversicolor. When benthic infauna was protected from these predators with exclosures, juvenile macrofauna responded with a marked increase in number, while the permanent meiofauna remained unaltered except for a limited increase in nematodes. It is concluded that in the Wadden Sea abundance of permanent meiofauna (Nematoda, Turbellaria, Ostracoda, Copepoda) is only locally or temporarily regulated by macrobenthic predators.I acknowledge a grant from the Deutsche Forschungsgemeinschaft (DFG).     

The Benefit of a Roseobacter Species on the Survival of Scallop Larvae

A marine strain (BS107), identified as a Roseobacter species, was antagonistic to Vibrio species on agar plates. Results suggested that the inhibitory effect was displayed only in the presence of another bacterium. Quantification of the antibacterial activity showed that 48-hour-coculture supernatants from BS107 and another bacterial strain (V. anguillarum 408) reached the highest titers of bacterial inhibition. The antibacterial substance was also liberated when supernatants from V. anguillarum 408 were added to pure cultures of the inhibition-productive bacterium. The presence of a proteinaceous molecule may induce BS107 to display the inhibitory effect. The antibacterial substance was sensitive to trypsin (8000 U/ml) and stable at 100°C. Cell extracts of the isolate BS107 (10⁶ cells/ml) significantly enhanced scallop larval survival, thus being beneficial to the rearing process. Received December 8, 1997; accepted July 15, 1998.     

Production of a New Thiopeptide Antibiotic,TP-1161, by a Marine Nocardiopsis Species

Twenty-seven marine sediment- and sponge-derived actinomycetes with a preference for or dependence on seawater for growth were classified at the genus level using molecular taxonomy. Their potential to produce bioactive secondary metabolites was analyzed by PCR screening for genes involved in polyketide and nonribosomal peptide antibiotic synthesis. Using microwell cultures, conditions for the production of antibacterial and antifungal compounds were identified for 15 of the 27 isolates subjected to this screening. Nine of the 15 active extracts were also active against multiresistant Gram-positive bacterial and/or fungal indicator organisms, including vancomycin-resistant Enterococcus faecium and multidrug-resistant Candida albicans. Activity-guided fractionation of fermentation extracts of isolate TFS65-07, showing strong antibacterial activity and classified as a Nocardiopsis species, allowed the identification and purification of the active compound. Structure elucidation revealed this compound to be a new thiopeptide antibiotic with a rare aminoacetone moiety. The in vitro antibacterial activity of this thiopeptide, designated TP-1161, against a panel of bacterial strains was determined.Natural products remain the most prolific source of new antimicrobials, and the chemical diversity of natural compounds is still unmatched by combinatorial chemistry approaches (9, 31). While the latter has been successfully applied for lead optimization, it basically failed to deliver genuinely new pharmacophores, especially in the field of antimicrobials (31), mainly due to limitations in the structural variety of compounds represented in combinatorial libraries.Most of the antibiotics in clinical use today have been developed from compounds isolated from bacteria and fungi, with members of the actinobacteria being the dominant source (34). Traditionally, most of these antimicrobials have been isolated from soil-derived actinomycetes of the genus Streptomyces. However, isolation strategies in recent years have been directed to unexploited environments like marine sources (40). Bioprospecting efforts focusing on the isolation and screening of actinobacteria from ocean habitats (25, 27) have added new biodiversity to the order Actinomycetales and revealed a range of novel natural products of pharmacological value. The existence of marine actinobacterial species physiologically and phylogenetically distinct from their terrestrial relatives is now widely accepted, and new taxonomic groups of marine actinomycetes have been described for at least six different families within the order Actinomycetales (12). Apart from being phylogenetically distinct from their terrestrial relatives, marine isolates have been shown to possess specific physiological adaptations (e.g., to high salinity/osmolarity and pressure) to their maritime surroundings and many were found to produce novel and chemically diverse secondary metabolites (10, 13, 35).Most streptomycetes and other filamentous actinomycetes possess numerous gene clusters for the biosynthesis of secondary metabolites (2, 32), and genome sequence studies have shown that large portions of their genomes are devoted to secondary metabolite biosynthesis. Twenty gene clusters coding for known or predicted secondary metabolites were identified in the 8.7-Mb genome of Streptomyces coelicolor A3(2) (2), and 6.4% of the 8.7-Mb genome of Streptomyces avermitilis is dedicated to gene clusters for secondary metabolite biosynthesis (32). The marine actinomycete Salinispora allocates nearly 10% of its 5.2-Mb genome to 17 diverse biosynthetic loci, including polyketide synthases (PKSs), nonribosomal peptide synthetases (NRPSs), and several hybrid clusters (4, 43). Many medicinally important natural products, including antibacterials and antifungals, are synthesized by these multimodular assembly lines (14), and genome mining for secondary metabolite gene clusters has become a common tool to assess the genetic capability of bacteria to produce novel bioactive compounds. However, even for well-studied model antibiotic producers like S. coelicolor A3(2), discrepancies between the number of known metabolites on the one hand and the number of pathways identified from genomic data on the other hand are tremendous (2). These discrepancies can only be explained by the facts that most gene clusters for secondary metabolites are silenced under standard laboratory cultivation conditions and that an expression or upregulation of these pathways is only triggered in response to certain environmental signals. It has been shown that by cultivating bacteria under a range of conditions, it is possible to obtain products of many of these “orphan” biosynthetic pathways (4). Using the OSMAC (one strain-many compounds) approach, Bode et al. were able to isolate more than 100 compounds comprising 25 structural classes from only six microorganisms (4).In this study, marine sediment-derived actinomycete isolates were analyzed for the production of antimicrobial secondary metabolites by using microwell plate fermentations and a range of media and conditions. This approach led to the isolation of a new thiopeptide antibiotic, designated TP-1161, produced by a marine sediment-derived Nocardiopsis isolate. Here we report the isolation and structural and biological characterization of TP-1161.     

Grazing on green algae by the periwinkleLittorina littorea in the Wadden Sea

On sedimentary tidal flats in the Wadden Sea near the Island of Sylt, the periwinkleLittorina littorea occurred preferentially on clusters and beds of mussels and on shell beds (100 to 350 m⁻²), achieved moderate densities on green algal patches or mats (20 to 50 m⁻²), and remained rare on bare sediments (<5 m⁻²). Green algae covering>10% of sediment surface appeared in summer on approximately one third of the tidal zone, mainly in the upper and sheltered parts and almost never on mussel and shell beds. In feeding experiments,L. littorea ingested more of the dominant alge,Enteromorpha, than ofUlva, irrespective of whether or not algae were fresh or decaying. The tough thalli ofChaetomorpha were hardly consumed. Snails feeding onEnteromorpha produced fecal pellets from which new growth ofEnteromorpha started. In the absence of periwinkles,Enteromorpha developed on mussels and the attached fucoids. Experimentally increased snail densities on sediments prevented green algal development, but the snails were unable to graze down established algal mats. It is concluded that natural densities ofL. littorea hardly affect the ephemeral mass development of green algae on sediments. However, where the snails occur at high densities, i.e. on mussel beds, green algal development may be prevented.     

Effects of herbivorous birds on intertidal seagrass beds in the northern Wadden Sea

During autumn migration (September to December), brent geese (Branta b. bernicla) and wigeon (Anas penelope) feed on the seagrass Zostera noltii in the nearshore, upper tidal zone leeward of the island of Sylt (eastern North Sea). To graze on leaves and shoots above the sediment and on rhizomes and roots below, these birds reworked the entire upper 1 cm layer of sediment eight times within this 3-month period. In addition, brent geese excavated pits 3–10 cm deep by trampling in order to feed on below-ground phytomass. About 12% of the seagrass beds became pitted to an average depth of 4.5 cm. Using net exclosures, it was estimated that birds removed 34 g dry weight m^–2 of above-ground and 28 g of below-ground phytomass. This corresponds to 45% of the phytomass in September. Of the overall loss of phytomass from September to December, 63% was caused by birds. Roughly half of the leaves fell off anyway until December and the other half were taken by the birds. Below the ground, phytomass remained almost constant where birds were excluded, while with birds phytomass of rhizomes and roots was halved. In spite of this strong effect, in the next vegetation period the blade density was lower at former exclosure sites compared to the ambient seagrass bed. The underlying process seems to be a self-inhibition of dense overwintering seagrass by mud accretion. Assuming our experimental results can be scaled up to the entire seagrass bed, we hypothesize that in the sheltered upper intertidal zone, seasonal erosion caused by herbivorous geese and ducks is necessary for the persistence of Z. noltii. Received: 7 January 1999 / Received in revised form: 23 August 1999 / Accepted: 25 August 1999     

Antibiotic Resistance Patterns and Detection of blaDHA-1 in Salmonella Species Isolates from Chicken Farms in South Korea

Fifteen nonrepetitive ampicillin-resistant Salmonella spp. were identified among 91 Salmonella sp. isolates during nationwide surveillance of Salmonella in waste from 131 chicken farms during 2006 and 2007. Additional phenotyping and genetic characterization of these 15 isolates by using indicator cephalosporins demonstrated that resistance to ampicillin and reduced susceptibility to cefoxitin in three isolates was caused by TEM-1 and DHA-1 β-lactamases. Plasmid profiling and Southern blot analysis of these three DHA-1-positive Salmonella serovar Indiana isolates and previously reported unrelated clinical isolates of DHA-1-positive Salmonella serovar Montevideo, Klebsiella pneumoniae, and Escherichia coli from humans and swine indicated the involvement of the large-size plasmid. Restriction enzyme digestion of the plasmids from the transconjugants showed variable restriction patterns except for the two Salmonella serovar Indiana isolates identified in this study. To the best of our knowledge, this is the first report of the presence of the DHA-1 gene among Salmonella spp. of animal origin.Nontyphoidal Salmonella (NTS) strains are a significant cause of gastrointestinal infections of food origin. These microbes are a heterogeneous group of medically important Gram-negative bacteria and can infect a wide range of animals, including humans (3, 6, 9-11, 25).Currently, no antimicrobial therapies are recommended for the treatment of NTS infection unless a patient is of extreme age, has an underlying disease, or is infected with an invasive Salmonella sp. However, the use of antibiotics in treatment of clinical enteric infection has been heavily compromised by emerging multidrug-resistant microbes (4, 17, 18, 23). In particular, resistance due to extended-spectrum β-lactamases (ESBLs) and AmpC β-lactamases is of special concern as these enzymes confer resistance to some of the front-line antibiotics used to treat enteric infection in humans and animals (4, 13, 14, 19).Four classes of β-lactamases are known to confer resistance to β-lactam antibiotics. Among these, plasmid-mediated class A and class C β-lactamases have been frequently reported, whereas class B and class D β-lactamases are relatively rare (4). TEM and SHV enzymes of class A β-lactamases are generally found in Gram-negative bacteria and are derived by one or more amino acid substitutions around the active site of the enzyme that is responsible for the ESBL phenotype (4). Recently, the CTX-M enzyme of class A β-lactamases has been increasingly reported from enteric microbes, like Salmonella and Escherichia coli (4, 5, 9, 15). These have greater activity against cefotaxime than do other oxyimino-β-lactam substrates, like ceftazidime, ceftriaxone, or cefepime (4, 5). Plasmid-mediated AmpC β-lactamases, like DHA and CMY, are not inhibited by clavulanic acid and have been isolated from a wide variety of clinical and community-acquired microbes (2, 4, 13, 14, 16). These β-lactamases are native to the chromosomes of many Gram-negative bacilli but are missing in some genera, like Salmonella (4). The majority of β-lactamases reported in Salmonella to date have been derived from human clinical isolates, and only limited information is available regarding Salmonella spp. derived from farm animals, although isolates from both humans and animals are of clinical and epidemiological importance (4, 15, 25).In light of this knowledge gap, our study focused on assessing the distribution of Salmonella serovars in poultry farms in South Korea. Subsequently, isolates were analyzed for resistance to antibiotics commonly used in farms. Phenotypic and genetic characteristics of ampicillin-resistant Salmonella isolates were tested to gain insight into what β-lactamases were prevalent among these strains. We also characterized DHA-1-associated plasmids in these Salmonella spp. and compared them with clinical isolates of Salmonella, Klebsiella pneumoniae, and Escherichia coli from humans and from swine.     

Production of a Naphthoquinone Pigment by a Species of Streptoverticillium and Its Accumulation by a Streptomycete

A naphthoquinone pigment produced by a species of Streptoverticillium was accumulated by a Streptomyces sp. when the two organisms were grown in mixed cultures.     

Ecology, Inhibitory Activity, and Morphogenesis of a Marine Antagonistic Bacterium Belonging to the Roseobacter Clade

Roseobacter strain 27-4 has been isolated from a turbot larval rearing unit and is capable of reducing mortality in turbot egg yolk sac larvae. Here, we demonstrate that the supernatant of Roseobacter 27-4 is lethal to the larval pathogens Vibrio anguillarum and Vibrio splendidus in a buffer system and inhibited their growth in marine broth. Liquid chromatography (LC) with both UV spectral detection and high-resolution mass spectrometry (HR-MS) identified the known antibacterial compound thiotropocin or its closely related precursor tropodithietic acid in the bioactive fractions. Antibacterial activity correlated with the appearance of a brownish pigment and was only formed in marine broth under static growth conditions. A thick biofilm of multicellular star-shaped aggregated cells formed at the air-liquid interface under static growth conditions. Here, the bioactive compound was the base peak in the LC-UV chromatograms of the extracts where it constituted 15% of the total peak area. Aerated conditions results in 10-fold-higher cell yield, however, cultures were nonpigmented, did not produce antibacterial activity, and grew as single cells. Production of antibacterial compounds may be quorum regulated, and we identified the acylated homoserine lactone (3-hydroxy-decanoyl homoserine lactone) from cultures of Roseobacter 27-4 using LC-HR-MS. The signal molecule was primarily detected in stagnant cultures. Roseobacter 27-4 grew between 10 and 30°C but died rapidly at 37°C. Also, the antibacterial compounds was sensitive to heat and was inactivated at 37°C in less than 2 days and at 25°C in 8 days. Using Roseobacter 27-4 as a probiotic culture will require that is be established in stagnant or adhered conditions and, due to the temperature sensitivity of the active compound, constant production must be ensured.     

Against the trend: increasing numbers of breeding Northern Lapwings Vanellus vanellus and Black-tailed Godwits Limosa limosa limosa on a German Wadden Sea island

Capsule The increase in population sizes over the last 30 years cannot be explained by reproductive success.

Aims To establish whether the positive population trends are due to increasing and self-sustaining populations or to immigration.

Methods We studied the population development of breeding lapwings from 1971 until 2005 and of godwits from 1977 until 2005 on Wangerooge, a German Wadden Sea island. Both species increased significantly during the last three decades. For each species we used a logistic growth model to derive the reproductive output required to explain the past population development without assuming immigration. We compare the values derived by this model with empirical findings of reproductive output of the respective populations.

Results For neither lapwings nor godwits can empirical reproductive success explain the observed population development.

Conclusion Our results imply that the increase in breeding pairs of Northern Lapwing and Black-tailed Godwit on Wangerooge Island is not due to reproductive output. We propose that it is mainly caused by immigration onto the island.     

Investigation of a Great Number of Actinomycete Isolates on Their Antagonistic Effects Against Soil-Borne Fungal Plant Pathogens by an Improved Method

By means of a modified agar-ring method, the antibiotic efficacy of 300 isolates of actinomycetes, obtained from soil samples from Turkey, against six different test fungi was investigated. A wide range in their degree of sensitivity was evident. Whereas Sclerotinia sclerotiorum was completely inhibited by more than 90 % of the tested isolates, Rhizoctonia solani and Alternaria alternata were suppressed by only 17 and 14 %, respectively; Pythium debaryanum, Cochliobulus sativ, us and Macrophomina phaseolina held an intermediate position. Isolates which completely prevented the growth of Rhizoctonia solani and Alternaria alternata, were usually also highly effective against the other test fungi. Possible conclusions which may be drawn from these results are discussed.     

Large-Scale Spatial Dynamics of Intertidal Mussel (Mytilus edulis L.) Bed Coverage in the German and Dutch Wadden Sea

Intertidal blue mussel beds are important for the functioning and community composition of coastal ecosystems. Modeling spatial dynamics of intertidal mussel beds is complicated because suitable habitat is spatially heterogeneously distributed and recruitment and loss are hard to predict. To get insight into the main determinants of dispersion, growth and loss of intertidal mussel beds, we analyzed spatial distributions and growth patterns in the German and Dutch Wadden Sea. We considered yearly distributions of adult intertidal mussel beds from 36 connected tidal basins between 1999 and 2010 and for the period 1968–1976. We found that in both periods the highest coverage of tidal flats by mussel beds occurs in the sheltered basins in the southern Wadden Sea. We used a stochastic growth model to investigate the effects of density dependence, winter temperature and storminess on changes in mussel bed coverage between 1999 and 2010. In contrast to expectation, we found no evidence that cold winters consistently induced events of synchronous population growth, nor did we find strong evidence for increased removal of adult mussel beds after stormy winter seasons. However, we did find synchronic growth within groups of proximate tidal basins and that synchrony between distant groups is mainly low or negative. Because the boundaries between synchronic groups are located near river mouths and in areas lacking suitable mussel bed habitat, we suggest that the metapopulation is under the control of larval dispersal conditions. Our study demonstrates the importance of moving from simple habitat suitability models to models that incorporate metapopulation processes to understand spatial dynamics of mussel beds. The spatio-dynamic structure revealed in this paper will be instrumental for that purpose.     

Macrofauna distribution and bioturbation on tidal confluences of the dutch Wadden Sea

Macrofauna density and bioturbation intensity (measured with X-ray radiography), were studied in the Dutch Wadden Sea near Ameland and in Mok Bay, Texel, in September 1988. The bivalveMacoma balthica and the polychaeteHeteromastus filiformis were widely distributed in the areas studied. The cockleCerastoderma edule and the polychaeteScoloplos armiger did not occur in areas with high clay content, while the molluscsHydrobia ulvae andMya arenaria preferred fine sediments. The polychaeteNereis diversicolor preferred the higher parts of the intertidal. In the Ameland area disturbance of primary sediment structures was highest near the shore and near the middle of the tidal confluence where physical reworking was low. The sheltered Mok Bay sediments were completely reworked by deposit-feeders. Bioturbation intensity and deposit-feeder (bioturbator) density were, however, not highly correlated. A number of stations showed higher bioturbation towards the surface and this may be related to the high abundance of near surface dwelling macrofauna, due to eutrophication and organic enrichment of the sediments in recent years.