Benthos structure on tropical tidal flats of Australia

General features of benthos communities of tropical tidal flats are defined from northeast Australia, based on surveys from 1988 to 1991 in Hinchinbrook Channel and in the Haughton River estuary. A zonation of benthic communities is described. Total abundances for macrofauna averaged 31 individuals 200 cm⁻², mesofauna (defined as infauna smaller than 0.5 mm and retained on a 0.25 mm sieve) averaged 16 individuals 10 cm⁻² and meiofauna averaged 231 individuals 5 cm⁻². The two study sites were similar in their abundance structures, but species due to a higher diversity of polychaeta in the Channel (120 species, H′=3.80 vs. 29 species and H′=1.78 in the Haughton estuary). Species densities were comparable at both sites and rather low (6 species 177 cm⁻², 2–7 species 10 cm⁻² and 5 meiobenthic Plathelminth species 5 cm⁻²). Over 50% of the species encountered were represented by less than three individuals. Deposit feeders, dominated the assemblages both in terms of individuals as well as species numbers. Tropical tidal flats are compared with their temperate counterparts and approaches for future research in tropical benthos communities are recommended.     

Spatial and temporal patterns of platyhelminth assemblages in intertidal sediments of northeast Australia

The distribution of Platyhelminthes was surveyed along two transects in tidal flats of the northeast coast of Queensland, Australia, in May 1990 and September 1991. Species numbers and densities as well as individual abundances and diversity increased from the high towards the low intertidal in relation to sediment composition. Along this gradient, platyhelminth assemblages of low intertidal sandflats, muddy sandflats and mudflats were distinguished with multivariate analyses. The distribution pattern was affected by the sediment parameter, and also by interactions with macrofauna. Sampling and analysis of the platyhelminth assemblage in tidal flats of Hinchinbrook Channel over time (November 1988 to October 1991) revealed annual and interannual variation, but no seasonality. This revised version was published online in July 2006 with corrections to the Cover Date.     

Patterns in pelagic and benthic nanoflagellate densities in the coastal upwelling system along the Banc d'Arguin,Mauritania

We studied spatial variation in abundance of marine benthic and pelagic heterotrophic nanoflagellates in relation to abundances of autotrophic flagellates, bacteria and cyanobacteria in an upwelling area off the Banc d'Arguin, Mauritania.There was enormous spatial variation in densities. In the sediments these ranged from 8–219 × 10³ cm^–3 for heterotrophic flagellates. Maximum values are in the range of those for temperate shallow marine bottoms. Low densities (< 20 × 10³) were confined to the deep stations (> 1000 m). Over the shelf (10–100 m depth) densities were high but related to grain size rather than to concurrent upwelling phenomena or to the abundance of benthic macrofauna.Pelagic flagellate abundance appeared to be more indicative of contemporary hydrographic conditions, most obvious by an increase in the ratio heterotrophic/autotrophic nanoflagellates away from the area of most intense upwelling.     

Zonation of intertidal macrobenthos in the estuaries of Schelde and Ems

Based on data, collected in 1980–1990, the intertidal benthic macrofauna of the Schelde and Ems estuaries was compared. The spatial occurrence of the benthic macrofauna along the salinity gradient, including the freshwater tidal area was emphasized. Both estuaries appeared to have a very similar species composition, especially at genus level. The higher number of species observed in the Schelde estuary was probably due to a greater habitat diversity. In both estuaries species diversity decreased with distance upstream. The total density did not vary along the estuarine gradient, whereas biomass is highest in the polyhaline zone.In both estuaries distinct intertidal benthic communities were observed along the salinity gradient: a marine community in the polyhaline zone, a brackish community in the mesohaline zone, and a third community in the oligohaline and freshwater tidal zones of the estuary. These three communities were very similar between both estuaries. Their main characteristics were discussed together with the occurrence and distribution of the dominant species.For the Schelde estuary and to a lesser extent also for the Ems estuary, there was evidence that anthropogenic stress had a negative effect on the intertidal macrobenthic communities of the oligohaline/freshwater tidal zone. Only Oligochaeta were dominating, whereas the very euryhaline and/or true limnetic species were missing. In the mesohaline zone, the Schelde estuary was dominated by large numbers of short-living, opportunistic species, whereas in the Ems estuary relatively more stable macrobenthic communities were observed. A comparison with some other European estuaries showed in general similar trends as those observed for the Schelde and Ems estuaries.     

Organic sedimentation and macrofauna as forcing factors in marine benthic nanofagellate communities

We investigated how benthic nanoflagellate communities in marine sediments respond to sedimentation of organic material and to the presence of macrofaunal organisms in controlled boxcosms. An input of 24 g C m^–2 resulted in a sharp increase in densities, from 93 to 477 × 10³ flagellates cm^–3 within 11 days. At the onset, this increase was paralleled by enhanced bacterial production and bacterial numbers. When bacterial production collapsed, flagellate ingestion rates, varying from 17 to 67 bact flag^–1 h^–1, were sufficient to control bacterial abundance. The presence of macrofauna accelerated the burst in flagellate densities. With macrofauna the same maximum densities were reached, but later densities dropped to relatively low levels. Macrofaunal bioturbation resulted in higher flagellate densities deeper in the sediment (up to 1200% at 3 cm and up to 460% at 6 cm deep). Correspondence to: R.P.M. Bak.     

Intertidal zone of svalbard

Summary In summer 1985–1991, the intertidal zone of the Svalbard archipelago was sampled in 242 localities. Thirty seven laxa of macrofauna and 22 of macrophytes were considered as littoral zone inhabitants. Four major littoral assemblages are described: Fucus-Balanus, Gammarus, Onisimus and Oligochaeta communities. More than 80% of the investigated coast is occupied by the Oligochaeta assemblage with mean biomass values less than 1 kJ/m². The richest benthos was found at Fucus-Balanus sites (8% of the coast line) with biomass values exceeding 2000 kJ/m². The southern tip of Spitsbergen is part of a major zoogeographical border in the littoral fauna distribution. Subarctic species like barnacles, periwinkles and Gammarus oceanieus predominate on the western coast whereas, on the Arctic East coast barren beaches, G. setosus predomination was found.     

Plathelminths in tropical intertidal sediments of northeastern Australia

Assemblages of free-living interstitial plathelminths are described from the tropical coast of northeastern Australia. About one hundred species were recorded from sediments of mangroves, tidal flats, and seagrass beds. The majority (88%) of the plathelminths occurred in only one havitat. Richest in species were Kalyptorhynchia (39), followed by Acoela (17) and Typhloplanoida (16). Over 50% of the species found were represented by one to three individuals, but a few species were frequently encountered. The preferred habitat was sand with 86% of all species, while abundances were higher in muddy sediments. The mean species density was 7 in 10 cm^–2 of sediment, and abundances ranged from 8 to 52 individuals 10 cm^–2 with an average plathelminth abundance of 25 individuals 10 cm^–2. Predatory plathelminths were predominant with up to 77% of the individuals. A comparison is made to plathelminth assemblages in temperate tidal flats.     

Sedimentation and overfishing drive changes in early succession and coral recruitment

Sedimentation and overfishing are important local stressors on coral reefs that can independently result in declines in coral recruitment and shifts to algal-dominated states. However, the role of herbivory in driving recovery across environmental gradients is often unclear. Here we investigate early successional benthic communities and coral recruitment across a sediment gradient in Palau, Micronesia over a 12-month period. Total sedimentation rates measured by ‘TurfPods’ varied from 0.03 ± 0.1 SE mg cm⁻² d⁻¹ at offshore sites to 1.32 ± 0.2 mg cm⁻² d⁻¹ at inshore sites. To assess benthic succession, three-dimensional settlement tiles were deployed at sites with experimental cages used to exclude tile access to larger herbivorous fish. Benthic assemblages exhibited rapid transitions across the sediment gradient within three months of deployment. At low levels of sedimentation (less than 0.6 mg cm⁻² d⁻¹), herbivory resulted in communities dominated by coral recruitment inducers (short turf algae and crustose coralline algae), whereas exclusion of herbivores resulted in the overgrowth of coral inhibitors (encrusting and upright foliose macroalgae). An ‘inducer threshold’ was found under increasing levels of sedimentation (greater than 0.6 mg cm⁻² d⁻¹), with coral inducers having limited to no presence in communities, and herbivore access to tiles resulted in sediment-laden turf algal assemblages, while exclusion of herbivores resulted in invertebrates (sponges, ascidians) and terrestrial sediment accumulation. A ‘coral recruitment threshold’ was found at 0.8 mg cm⁻² d⁻¹, below which net coral recruitment was reduced by 50% in the absence of herbivores, while recruitment was minimal above the threshold. Our results highlight nonlinear trajectories of benthic succession across sediment gradients and identify strong interactions between sediment and herbivory that have cascading effects on coral recruitment. Local management strategies that aim to reduce sedimentation and turbidity and manage herbivore fisheries can have measurable effects on benthic community succession and coral recruitment, enhancing reef resilience and driving coral recovery.     

Algae, macrofaunal assemblages and temperature: a quantitative approach to intertidal ecosystems of Iceland

Algae and the associated macrofauna in two Icelandic intertidal ecosystems under cold and warm influence, respectively, were studied with respect to algae-macrofauna relationships and a possible effect of temperature on community structure. Two sites in Iceland were selected, Sandgerdi ligthhouse (64°8′N 22°40′W) on the southwestern coast, and Grimsey Island (66°33′N 18°04′W), in the north, on the Arctic Circle, where sea temperature is considerably lower (5° approximately). The biomass of algae and the number of species of algae and macrofauna were higher in Sandgerdi than in Grimsey, and the patterns of diversity, evenness, biomass and abundance also differed between the sites. In the intertidal zone of Sandgerdi, a total of 28 species of algae and 45 species of macrofauna were identified whereas only 16 algal species and 27 macrofaunal species were found in Grimsey. Canonical correspondence analysis (CCA) using algal biomass as the environmental variable were conducted, and revealed significant relationships between algae composition and the associated macrofauna; some macrofauna taxa showed specific trophic or refuge relationships with algal species. According to the CCA, Corallina officinalis showed the highest correlation with macrofaunal assemblages in both study sites. However, correlations between macrofauna and other algae differed between Grimsey and Sandgerdi. The present study, together with additional observations in Greenland waters, shows a general decrease of species richness and diversity towards the north which may primarily be due to the temperature regime.     

Bacterial productivity and microbial biomass in tropical mangrove sediments

Bacterial productivity (³H-thymidine incorporation into DNA) and intertidal microbenthic communities were examined within five mangrove estuaries along the tropical northeastern coast of Australia. Bacteria in mangrove surface sediments (0–2 cm depth) were enumerated by epifluorescence microscopy and were more abundant (mean and range: 1.1(0.02–3.6)×10¹¹ cells·g DW^–1) and productive (mean: 1.6 gC·m^–2· d^–1) compared to bacterial populations in most other benthic environments. Specific growth rates (¯x=1.1) ranged from 0.2–5.5 d^–1, with highest rates of growth in austral spring and summer. Highest bacterial numbers occurred in winter (June–August) in estuaries along the Cape York peninsula north of Hinchinbrook Island and were significantly different among intertidal zones and estuaries. Protozoa (10⁵–10⁶·m^–2, pheopigments (0.0–24.1g·gDW^–1) and bacterial productivity (0.2–5.1 gC·m^–2·d^–1) exhibited significant seasonality with maximum densities and production in austral spring and summer. Algal biomass (chlorophylla) was low (mean: 1.6g·gDW^–1) compared to other intertidal sediments because of low light intensity under the dense forest canopy, especially in the mid-intertidal zone. Partial correlation analysis and a study of possible tidal effects suggest that microbial biomass and bacterial growth in tropical intertidal sediments are regulated primarily by physicochemical factors and by tidal flushing and exposure. High microbial biomass and very high rates of bacterial productivity coupled with low densities of meiofaunal and macroinfaunal consumers observed in earlier studies suggest that microbes may be a sink for carbon in intertidal sediments of tropical mangrove estuaries.     

A long-term comparison of the benthic algal flora of Clare Island, County Mayo, western Ireland

The marine benthic algal flora of Clare Island, off County Mayo, western shore of Ireland, was investigated; collections of intertidal and subtidal marine algae were made at 16 sites along the eastern and southern shores in the years 1990, 1993 and 2000–2002. The data and observations obtained were compared with the results of a similar survey conducted by Arthur Disbrowe Cotton in 1910–1911. Considering the results of the original survey and the new survey together, the marine algal flora of the island currently totals 293 species; 224 species were recorded by Cotton in the original survey, whereas 223 species were identified in the present study. Most species are common to the original and the new list and the main differences are easily explainable; the new survey used SCUBA diving, which allowed the collection of several subtidal species not collected in 1910, and Cotton reported several microscopic green and brown algae, usually difficult to recognise in the field, which were not rediscovered. The most remarkable differences consist in the current presence of some large intertidal brown algae (Bifurcaria bifurcata, Cystoseira foeniculacea and Cystoseira nodicaulis) that were not reported in the survey of 1910. Two algae, Codium fragile subsp. tomentosoides and Asparagopsis armata, were introduced in Europe after the original survey. At present, the benthic algal assemblages of Clare Island still have basically the same structure and distribution as in 1910 and, if compared with other coastal areas of Europe, the intertidal marine environment of Clare Island appears remarkably well conserved.     

Spatial structure of a subtidal macrobenthic community in the Bay of Veys (western Bay of Seine, English Channel)

The spatial distribution of the muddy fine sand community from the Bay of Veys (western English Channel) were investigated during spring and autumn 1997. A grid of 55 and 54 sites was sampled in March and October, respectively, using two replicates per site of a Hamon grab (0.25 m²) for macrofauna collection and an additional one for sediment analysis. A total of 172 species were sampled with a dominance of polychaetes, followed by crustaceans and bivalves. The species richness and abundance show low temporal changes despite higher values in October than in March. In March, the mean abundance was 165 ind. 0.5 m⁻²; in October, the mean abundance was 212 ind. 0.5 m⁻². Four assemblages from the Abra alba-Pectinaria koreni community were identified corresponding to a bathymetric and sedimentary gradient from muddy fine sands with high levels of fine particles in shallow water to fine sands in deeper water. The discussion focuses on factors prevailing on the spatial structure of sandy communities in the English Channel.     

Faunal utilization of constructed intertidal oyster (Crassostrea rivularis) reef in the Yangtze River estuary,China

An intertidal oyster reef (～260 ha) was created by planting hatchery-reared seed oysters (Crassostrea rivularis) on an artificial concrete modular reef in the Deepwater Navigation Channel Regulation Project of the Yangtze River estuary. We examined the development of reef communities (oyster, barnacle and motile epibenthic macrofauna), characterized nekton use and assessed the habitat value of the constructed reef. The C. rivularis oyster population showed a rapid exponential increase with time and reached maximum density (3410 ± 241 ind./m²) and biomass (3175 ± 532 g/m²) after one year of restoration. The barnacle Balanus albicostatus was the most abundant sessile macrofauna and had a significantly greater density in the high intertidal zone than in the low intertidal zone (P < 0.05). The reef also supported diverse motile epibenthic macrofauna (11 mollusks, 11 crustaceans, 4 annelids and 2 fishes), and the reef-associated communities were numerically dominated by Neanthes japonica, Perinereis aibuhitensis, Nerita yoldi and Littorinopsis intermedia. A total of 50 nekton species (31 fishes, 9 shrimps and 10 crabs) utilized the constructed intertidal oyster reef, and grass shrimp Palaemon spp. dominated the nekton communities in term of abundance. Since the constructed intertidal oyster reef supports a variety of reef communities and abundant nektons, it should be recognized as an important and protective fish habitat in the Yangtze River estuary.     

The Role of Substrate Type on Benthic Diatom Assemblages in the Daly and Roper Rivers of the Australian Wet/Dry Tropics

The selection of one or more river substrata for the collection of benthic diatoms is fundamental to any monitoring or research programme because it can potentially bias the diatom data set. In the wet/dry tropics of Australia, where the use of benthic diatoms for river health assessment is in its infancy, the comparability of diatom assemblages on river substrata has been assessed. Benthic diatoms were sampled from seven river sites, with a range of ionic chemistries (conductivities 27–6500 μS cm⁻¹) but low nutrient concentrations. At each site, triplicate samples were collected from 3 to 6 substrata. The diatom assemblages sampled were: epilithon (assemblages on rock), epiphytes on macroalgae and macrophytes, epidendron (assemblages on wood), epipsammon (assemblages on sand), epipelon (assemblages on mud) and bacterial slime. The variability between substrate assemblages, at each site, were assessed according to the following: (1) a multivariate analysis of diatom assemblages, (2) ANOVA tests of species richness, (3) ANOVA tests of the relative abundance of common species (defined by an abundance of at least 10% in any one sample), and (4) a comparison of the number of species unique to a substrate. A total of 198 taxa were identified, with some taxa common to temperate Australia. Common species were found on all substrata, with sometimes statistically significantly different relative abundances. Taxa unique to a substrate had low relative abundances (0.1–2%), were most often found on only one replicate, and are unlikely to be substrate specific because many are known to occur on other substrata. The assemblages on hard substrata, epilithon and epidendron, were found to be most similar. Diatom assemblages on macroalgal and macrophyte substrata, compared to other substrata, were highly variable. This is attributed to the loss of diatoms from grazing and sloughing, followed by recolonisation of newly exposed substrate. Other assemblages, notably epipsammon, were similar to epilithon and epidendron but sometimes differed in their relative abundance of common species. The principal finding of the study was the similarity of the epilithon and epidendron, which are considered to be indistinguishable. Rock and wood hard substrata can be substituted for one another during field surveys, thereby increasing the number of potential sample sites available for monitoring activities that standardise to a hard substrate.     

Distribution of live benthic Foraminifera at three oceanographically dissimilar sites in the northeast Atlantic: preliminary results

Live (i.e. rose Bengal stained) benthic foraminiferal assemblages in the 0–1 cm layer of multiple core samples were examined at three contrasting sites in the northeast Atlantic as part of the Natural Environment Research Council Deep Ocean Benthic Boundary Layer (BENBO) Thematic Programme. Sites A (3600 m water depth) and C (1900 m) were located in the Rockall Trough while Site B (1100 m) was in the Hatton–Rockall Basin. Wet-sorting sediment residues (>125 m fraction) revealed more abundant (98–190 individuals/10 cm²) and diverse (71–99 species /27 cm²) assemblages than found previously in studies employing dry-sorting techniques. At all three sites, the assemblages were numerically dominated by delicate soft-bodied and agglutinated species, most of which are undescribed. Calcareous taxa formed a relatively small proportion of the assemblages (23% of individuals at Site B, 13% at Site C and 7% at Site A). Conversely, the agglutinated hormosinaceans (mainly Reophax species) became more prominent with increasing depth and accounted for 8% of the assemblage at Site B, 33% at Site C and 51% at Site A. Foraminifera represented 67–81% of the fauna in the samples and greatly outnumbered the metazoans. The assemblage at Site A has similarities with abyssal assemblages, while those at Sites B and C contain characteristic bathyal species. At Site B, Uvigerina peregrina and Melonis barleeanum are abundant, suggesting that there are high organic carbon inputs at this site. Reduced species diversity was observed at Site A following the spring phytodetritus bloom due to the presence of large numbers of juvenile Hoeglundina elegans, although it is not clear whether this species is responding to phytodetritus inputs.     

Tufa deposition and periphyton overgrowth as factors affecting the ciliate community on travertine barriers in different current velocity conditions

A characteristic of aquatic systems in karstic region is the formation of tufa – the product of calcium carbonate precipitation. Artificial substrata (glass slides) were used to investigate the influence of tufa deposits at two different current velocities (5 cm s^–1 and 50 cm s^–1) on the ciliate assemblages in periphyton. After two-month exposure periods, periphyton biomass and tufa deposit were c. three times greater at 50 cm s^–1 than at 5 cm s^–1. Ciliate population density was also higher on artificial substrata exposed in a lotic than in a lentic microhabitat (the overall mean number of ciliates at 5 cm s^–1 was 122 ind. cm^–2, and at 50 cm s^–1 497 ind. cm^–2). At each of the two observed current velocities, a Principal Components (PCA) ordination of the colonized ciliate associations showed a spatial separation of the associations that developed under two different conditions of tufa deposition. During the period of greater tufa deposit, associated with greater periphyton overgrowth rate, the ciliate communities had higher species diversity (a higher number of species and a lower number of individuals). Species diversity of ciliates had a positive nonlinear relation to tufa deposition rate. These results suggest that artificial surfaces covered by a rough tufa layer associated with greater periphyton biomass offer diverse conditions for colonization by ciliates.     

Composition,distribution and biomass of meiobenthos in the Oosterschelde estuary (SW Netherlands)

Meiofauna composition, abundance, biomass, distribution and diversity were investigated for 31 stations in summer. The sampling covered the whole Oosterschelde and comparisons between the subtidal — intertidal and between the western-central — eastern compartment were made.Meiofauna had a community density ranging between 200 and 17 500 ind 10 cm^–2, corresponding to a dry weight of 0.2 and 8.4 gm^–2. Abundance ranged between 130 and 17 200 ind 10 cm^–2 for nematodes and between 10 and 1600 ind 10 cm^–2 for copepods. Dry weight biomass of these taxa was between 0.5–7.0 gm^–2 and 0.008–0.3 gm^–2 for nematodes and copepods respectively.The meiofauna was strongly dominated by the nematodes (36–99%), who's abundance, biomass and diversity were significantly higher intertidally than subtidally and significantly higher in the eastern part than in the western part. High numbers were positively correlated with the percentage silt and negatively with the median grain size of the sand fraction. The abundance and diversity of the copepods were highest in the subtidal, but their biomass showed an inverse trend being highest on the tidal flats.The taxa diversity of the meiofauna community and species diversity of both the nematodes and the copepods were higher in subtidal stations than on tidal flats. In the subtidal, the meiofauna and copepod diversity decreased from west to east, whereas nematode diversity increased.The vertical profile clearly reflected the sediment characteristics and could be explained by local hydrodynamic conditions.Seasonal variation was pronounced for the different taxa with peak abundance in spring, summer or autumn and minimum abundance in winter.Changes in tidal amplitude and current velocity enhanced by the storm-surge barrier will alter the meiofauna community structure. As a result meiofauna will become more important in terms of density and biomass, mainly due to increasing numbers of nematodes, increasing bioturbation, nutrient mineralisation and sustaining bacterial growth. A general decrease in meiofauna diversity is predicted. The number of copepods is expected to decrease and interstitial species will be replaced by epibenthic species, the latter being more important in terms of biomass and as food for the epibenthic macrofauna and fishes.     

Structure and diversity of shallow soft-bottom benthic macrofauna in the Gulf of Lions (NW Mediterranean)

Samples of soft-sediment macrobenthos from 92 sites between 10 and 50 m depth were used to assess (1) the main soft-bottom macrofauna communities in the Gulf of Lions, (2) the different components of the diversity of benthic macrofauna in this area, and (3) the relevance of the use of major taxonomic groups as surrogates for the analysis of the structure and diversity of total macrofauna. Three main communities were identified by cluster analysis and associated procedures. These communities corresponded well to the assemblages recently identified on the basis of polychaete composition. The α-diversity indices were in accordance with those reported for similar communities in the Mediterranean. Conversely, the β-diversity value was higher than the few other data available in the literature for marine soft-bottom macrofauna. The total number of species in the studied area estimated by the “total species accumulation curve” (TS) method was 2,319, which was only 10% higher than the number obtained by extrapolation of the species–area curve. The similarity matrix based on polychaetes correlated best with the one based on total macrofauna. Polychaetes and crustaceans were also the best surrogates of total macrofauna when assessing α-diversity (except in the case of Δ*). Conversely, molluscs were the best surrogates of total macrofauna β-diversity. Our results show that the choice of an optimal surrogate for total benthic macrofauna depends on the characteristic of the benthic macrofauna to be studied. Moreover, this choice is also dependent on the environment to be studied.     

Spatial variability in structural and functional aspects of macrofauna communities and their environmental parameters in the Jade Bay (Wadden Sea Lower Saxony, southern North Sea)

Spatial distribution and functional structure of intertidal benthic macrofauna in relation to environmental variables in the Jade Bay (southern North Sea) were studied and compared with other intertidal areas of the Wadden Sea. A total of 128 stations covering the whole Jade Bay were sampled in summer 2009. A total of 114 taxa were found. Highest species numbers occurred in the subtidal areas, whereas highest mean abundances were found in the upper intertidal areas. Based on species abundance data, six significantly distinct macrofauna communities in the Jade Bay were identified and evaluated with multivariate statistics, univariate correlations and canonical correspondence analysis. Differences in these community patterns were caused by the response of the dominant species (Hydrobia ulvae, Tubificoides benedii, Pygospio elegans, Caulleriella killariensis, Scoloplos armiger, Urothoe poseidonis, Microprotopus maculatus) to prevailing environmental conditions along the gradient from the lower and exposed sandy intertidal areas via intermediate mixed sediments to the upper mudflat areas. Distribution patterns in relation to tidal zonation were best explained by variability in submergence time, Chlorophyll a (chl a) content and sediment composition (mud content), which are proxies for hydrodynamic conditions and food availability. Species inventory and species richness were comparable with other intertidal areas of the Wadden Sea, but the Jade Bay differs from these areas regarding dominant species. Differences in sediment composition and morphological characteristics (macrotidal versus mesotidal Wadden Sea areas) are discussed for comparison of regional differences.     

Counting Viruses and Bacteria in Photosynthetic Microbial Mats

Viral abundances in benthic environments are the highest found in aquatic systems. Photosynthetic microbial mats represent benthic environments with high microbial activity and possibly high viral densities, yet viral abundances have not been examined in such systems. Existing extraction procedures typically used in benthic viral ecology were applied to the complex matrix of microbial mats but were found to inefficiently extract viruses. Here, we present a method for extraction and quantification of viruses from photosynthetic microbial mats using epifluorescence microscopy (EFM) and flow cytometry (FCM). A combination of EDTA addition, probe sonication, and enzyme treatment applied to a glutaraldehyde-fixed sample resulted in a substantially higher viral (5- to 33-fold) extraction efficiency and reduced background noise compared to previously published methods. Using this method, it was found that in general, intertidal photosynthetic microbial mats harbor very high viral abundances (2.8 × 10¹⁰ ± 0.3 × 10¹⁰ g⁻¹) compared with benthic habitats (10⁷ to 10⁹ g⁻¹). This procedure also showed 4.5- and 4-fold-increased efficacies of extraction of viruses and bacteria, respectively, from intertidal sediments, allowing a single method to be used for the microbial mat and underlying sediment.