The ecology and distribution of benthic foraminifera at the Håkon Mosby mud volcano (SW Barents Sea slope)

To investigate a possible influence of submarine methane seepage on benthic foraminiferal communities, Rose Bengal stained (“live”) and empty tests of benthic foraminifera were studied from the sediment surface down to 15 cm sub-bottom depth of 12 sites at the Håkon Mosby mud volcano (HMMV). In addition, one reference site well away from the seep sites, but from similar water depths and the same general hydrographic setting was occupied for comparison. The HMMV is located at 1265 m water depth on the SW Barents Sea continental slope. Distinct living foraminiferal associations at the HMMV are linked to specific sedimentary, microbial, and macrofaunal habitats. In the center of the crater, and in crater areas completely covered by bacterial mats, Cassidulina reniforme is the only, albeit rare, living species. Below the top few millimeters, sediments are anoxic and devoid of living specimens. At the rim of the mud volcano, at sites densely populated by pogonophoran tube worms, three benthic foraminiferal associations are found; (i) a Fontbotia wuellerstorfi–Lobatula lobatula association living attached to the upper parts of pogonophoran tubes, which protrude into oxic water, (ii) a diverse Cassidulina neoteretis association populating dysoxic sediments of the surface centimeter, and (iii) a species-poor Bolivina pseudopunctata association colonizing the subsurface sediments down to four centimeters. Generally, we did not find endemic or seep indicative species or associations at the HMMV. However, the HMMV live faunas dominated by B. pseudopunctata are not found at the reference site nor are they described from comparable water depths and environments without gas seepages from the Norwegian-Greenland Seas.In the center and outer rim of the mud volcano, a C. neoteretis–Reophax guttifer dead association, similar to the one at the reference site, characterizes an assemblage of strongly corroded and partly displaced tests. At bacterial mat sites, a C. reniforme dead association corresponds to the living one. Thus both the living and the dead associations are indicative of a specific bacterial mat environment at the HMMV.     

Nematode species distribution patterns at the Håkon Mosby Mud Volcano (Norwegian Sea)

The Håkon Mosby Mud Volcano (HMMV) is a methane seep that is densely populated by the bacteria Beggiatoa spp. as well as by tubeworms of the Family Siboglinidae. Analyses of samples from seven stations located in five different habitats (bare sediments, bacterial mats, siboglinid fields, sediments with small patches of siboglinids and areas of ‘normal’ sea floor) revealed differences in the density and species‐level diversity of nematode communities. Total densities increased from stations inside the mud volcano and on its rim towards the ‘normal’ areas outside the volcano. Nematode species diversity was similar in samples from the siboglinid fields and the bare HMMV sediments. However, the sediment with patches of siboglinids collected outside the volcano was characterised by a diverse nematode community comprising 47 species, whereas the control sediments without siboglinids yielded only 27 species. The nematode assemblage at the bacterial mat site included only two identified species, Halomonhystera disjuncta and Sabatieria ornata, with H. disjuncta being strongly dominant. Terschellingia distlamphida, S. ornata and Aponema ninae dominated nematode communities at the siboglinid fields and in bare HMMV sediments. Species dominating at stations inside the caldera were found in comparable low densities in the sediment with siboglinid patches collected outside the volcano, but were missing in the background samples, where species of Endeolophos, Acantholaimus and Desmoscolex were dominant. Species diversity generally decreased with sediment depth. A subsurface‐dwelling nematode community was observed in the siboglinid fields and the bare sediments. Background sediments showed a sharp decline with 18–20 species at 0–2 cm versus three species at 3–5 cm sediment depth. The trophic composition of the nematode fauna at the HMMV showed a prevalence of deposit feeders at almost all stations, whereas background sediments exhibited a uniform distribution of non‐selective and selective deposit feeders and epistrate feeders. The high percentage of deposit feeders inside the mud volcano could be related to the grain‐size distribution and the significantly higher bacterial biomass compared to the control sediments.     

Influence of mesh size and core penetration on estimates of deep-sea nematode abundance,biomass, and diversity

It is essential to maximize the information that can be gathered in deep-sea studies by thoroughly assessing sample processing methods. Nematodes are commonly used for the study and monitoring of deep-sea floor habitats, but the potential effects of different methods on the quantification of community attributes remain to be quantified. Here, we consider key methodological elements by comparing the effect of sediment depth and mesh size (63, 45, and 32 μm) on: (1) estimates of nematode community attributes, and (2) the sampling effort required to detect changes in these attributes at a bathyal site on the Chatham Rise, south-west Pacific Ocean. The 63 μm mesh retained most (95%) of the nematode biomass but a lower proportion (53–71%) of the nematode abundance. Retention efficiency of common species on this mesh ranged from 12 (Hapalomus sp.) to >88% (Comesomatidae spp.). The 63 μm mesh yielded significantly lower diversity estimates than the finer meshes, and failed to detect differences in community structure observed using the 45 and 32 μm mesh sizes. Sediment depth had a substantial effect on all measured community attributes, highlighting the importance of sufficient core penetration into the sediment (≥5 cm) for adequately characterizing nematode distribution. Power analysis showed that using a 32 μm mesh and deepest core penetration led to relatively few (3–8) samples being required to detect significant changes in nematode diversity indices relative to coarser mesh sizes. Characterization of nematode diversity and community structure using appropriate and robust methods of sampling is suggested as a sensitive and efficient tool for the assessment of anthropogenic impacts on deep-sea ecosystems.     

Macrofaunal succession in sediments around kelp and wood falls in the deep NE Pacific and community overlap with other reducing habitats

Sunken parcels of macroalgae and wood provide important oases of organic enrichment at the deep-sea floor, yet sediment community structure and succession around these habitat islands are poorly evaluated. We experimentally implanted 100-kg kelp falls and 200 kg wood falls at 1670 m depth in the Santa Cruz Basin to investigate (1) macrofaunal succession and (2) species overlap with nearby whale-fall and cold-seep communities over time scales of 0.25–5.5 yr. The abundance of infaunal macrobenthos was highly elevated after 0.25 and 0.5 yr near kelp parcels with decreased macrofaunal diversity and evenness within 0.5 m of the falls. Apparently opportunistic species (e.g., two new species of cumaceans) and sulfide tolerant microbial grazers (dorvilleid polychaetes) abounded after 0.25–0.5 yr. At wood falls, opportunistic cumaceans become abundant after 0.5 yr, but sulfide tolerant species only became abundant after 1.8–5.5 yr, in accordance with the much slower buildup of porewater sulfides at wood parcels compared with kelp falls. Species diversity decreased significantly over time in sediments adjacent to the wood parcels, most likely due to stress resulting from intense organic loading of nearby sediments (up to 20–30% organic carbon). Dorvilleid and ampharetid polychaetes were among the top-ranked fauna at wood parcels after 3.0–5.5 yr. Sediments around kelp and wood parcels provided low-intensity reducing conditions that sustain a limited chemoautrotrophically-based fauna. As a result, macrobenthic species overlap among kelp, wood, and other chemosynthetic habitats in the deep NE Pacific are primarily restricted to apparently sulfide tolerant species such as dorvilleid polychaetes, opportunistic cumaceans, and juvenile stages of chemosymbiont containing vesicomyid bivalves. We conclude that organically enriched sediments around wood falls may provide important habitat islands for the persistence and evolution of species dependent on organic- and sulfide-rich conditions at the deep-sea floor and contribute to β and γ diversity in deep-sea ecosystems.     

Fungi in deep-sea sediments of the Central Indian Basin

Although a great amount of information is available on bacteria inhabiting deep-sea sediments, the occurrence of fungi in this environment has been poorly studied and documented. We report here the occurrence of fungi in deep-sea sediments from ∼5000 m depth in the Central Indian Basin (9–16°S and 73–76°E). A total of 181 cultures of fungi, most of which belong to terrestrial sporulating species, were isolated by a variety of isolation techniques. Species of Aspergillus and non-sporulating fungi were the most common. Several yeasts were also isolated. Maximum species diversity was observed in 0–2 cm sections of the sediment cores. Direct staining of the sediments with Calcofluor, a fluorescent optical brightener, revealed the presence of fungal hyphae in the sediments. Immunofluorescence using polyclonal antibodies raised against a deep-sea isolate of Aspergillus terreus (# A 4634) confirmed its presence in the form of hyphae in the sub-section from which it was isolated. A total of 25 representative species of fungi produced substantial biomass at 200 bar pressure at 30° as well as at 5 °C. Many fungi showed abnormal morphology at 200 bar/5 °C. A comparison of terrestrial isolates with several deep-sea isolates indicated that the former could grow at 200 bar pressure when growth was initiated with mycelial inocula. However, spores of a deep-sea isolate A. terreus (# A 4634), but not the terrestrial ones, showed germination at 200 bar pressure and 30 °C. Our results suggest that terrestrial species of fungi transported to the deep sea are initially stressed but may gradually adapt themselves for growth under these conditions.     

Deep-sea habitat heterogeneity influence on meiofaunal communities in the Gulf of Guinea

To estimate the degree of spatial heterogeneity of benthic deep-sea communities, we carried out a multiple-scale (from m's to 200 km) investigation in the Congo-Angola margins (Equatorial West African margin, 3150–4800 m) in which we examined the metazoan meiofauna at a variety of habitats along the Congo Channel system and in the associated cold seep. We investigate the structure, density, vertical distribution patterns in the sediment and biomass of meiofaunal communities in the Gulf of Guinea and how they are controlled by hydrologic and biogeochemical processes. The meiofaunal communities in the Gulf of Guinea were shaped by heterogeneous conditions on the margin, and reflect the multiple-scale spatial variability that corresponds with the different identified habitats. The two control sites, located at >100 km away from the canyon, were inhabited by very dense and the most diverse meiobenthic communities. Similar meiobenthic communities inhabited the transition zone between the canyon and the cold seep. Sites located along the Congo Channel were obviously affected by the local high-velocity bottom currents and unstable sedimentary conditions in this active submarine system. Extremely low meiobenthic densities and very low proportions in the most surficial sediment layers provided evidence for recently highly disturbed sediments at these sites. The remote operated vehicle (ROV) Victor 6000 provided images of the cold seep, showing a patchy distribution of several types of patchy distributed megafaunal communities dominated by three key symbiotic taxa (Mytilidae, Vesicomyidae and Siboglinidae). These cold seep sediments were colonised by a unique meiobenthic community, characterised by a high small-scale (m's) patchiness, low species richness and the prominent dominance of two large-sized nematode species: Sabatieria mortenseni, which is a cosmopolitan nematode known from littoral habitats, and an undescribed Desmodora species. The high individual body weight of S. mortenseni and its dominance at the cold seep site resulted in a significantly higher nematode biomass at the seep compared to the surrounding sites. In addition, the vertical nematode profiles, with maximum proportions in subsurficial layers, points to a chemosynthesis-based meiobenthic community in this cold seep, in contrast to the phytodetritus-based communities at the control sites and at the transition zone.     

Abundance of small individuals influences the effectiveness of processing techniques for deep-sea nematodes

Nematodes are the most abundant metazoans of deep-sea benthic communities, but knowledge of their distribution is limited relative to larger organisms. Whilst some aspects of nematode processing techniques, such as extraction, have been extensively studied, other key elements have attracted little attention. We compared the effect of (1) mesh size (63, 45, and 32 μm) on estimates of nematode abundance, biomass, and body size, and (2) microscope magnification (50× and 100×) on estimates of nematode abundance at bathyal sites (250–3100 m water depth) on the Challenger Plateau and Chatham Rise, south-west Pacific Ocean. Variation in the effectiveness of these techniques was assessed in relation to nematode body size and environmental parameters (water depth, sediment organic matter content, %silt/clay, and chloroplastic pigments). The 63-μm mesh retained a relatively low proportion of total nematode abundance (mean±SD=55±9%), but most of nematode biomass (90±4%). The proportion of nematode abundance retained on the 45-μm mesh in surface (0–1 cm) and subsurface (1–5 cm) sediment was significantly correlated (P<0.01) with %silt/clay (R²=0.39) and chloroplastic pigments (R²=0.29), respectively. Variation in median nematode body weight showed similar trends, but relationships between mean nematode body weight and environmental parameters were either relatively weak (subsurface sediment) or not significant (surface sediment). Using a low magnification led to significantly lower (on average by 43%) nematode abundance estimates relative to high magnification (P<0.001), and the magnitude of this difference was significantly correlated (P<0.05) with total nematode abundance (R²_p=0.53) and the number of small (≤250 μm length) individuals (R²_p=0.05). Our results suggest that organic matter input and sediment characteristics influence the abundance of small nematodes in bathyal communities. The abundance of small individuals can, in turn, influence abundance estimates obtained using different mesh sizes and microscope magnifications.     

Analysis of a PAH-degrading bacterial population in subsurface sediments on the Mid-Atlantic Ridge

Little is known about the types and concentrations of polycyclic aromatic hydrocarbons (PAHs) existing in the deep-sea subsurface environment, which is believed to be cold, oligothrophic and of high static pressure. PAHs in the upper layers of the water column are unavoidably subjected to degradation while they are deposited to the sea floor and become embedded in the deep-sea sediment. In this report, a high concentration of PAHs was discovered in the sediment 2.7 m beneath the bottom surface at a water depth of 3962 m on the Mid-Atlantic Ridge (MAR). The total concentration of PAHs was 445 ng (g dry wt sediment)^-1. Among the seven detected PAHs, the concentrations of phenanthrene (222 ng g^-1) and fluorene (79 ng g^-1) were relatively high. In addition, PAH-degrading bacteria were found within the sediments. As in a previously detected site on the MAR, in the PAH-enriched region of this site, a bacterium of the genus Cycloclasticus was found to be the predominant isolate detected by PCR-DGGE analysis. In addition, bacteria of the Halomonas, Marinobacter, Alcanivorax, Thalassospira and Maricaulis genera, were also included in the PAH-degrading community. In summary, a high concentration of PAHs was detected in the subsurface of the deep-sea sediment, and once again, the Cycloclasticus bacterium was confirmed to be a ubiquitous marine PAH degrader even in the subsurface marine environment. Considering the abundance of PAHs therein, biodegradation is thus thought to be inactive, probably because of the low temperature, limited oxygen and/or limited nutrients.     

Nucleic acid concentrations (DNA,RNA) in the continental and deep-sea sediments of the eastern Mediterranean: relationships with seasonally varying organic inputs and bacterial dynamics

In order to study temporal variations of the genetic material in the continental shelf and deep-sea sediments of the extremely oligotrophic Cretan Sea, samples were collected on seasonal basis from August 1994 to September 1995, with a multiple corer, at seven stations (from 40 to 1540 m depth). Surface sediments (0–1 cm) were sub-sampled and analyzed for nucleic acid content (DNA, RNA) and bacterial density. DNA concentrations in the sediments were high (on annual average, 25.0 μg g^-1) and declined with increasing water depth, ranging from 3.5 to 55.2 μg g^-1. DNA concentrations displayed wide temporal changes also at bathyal depths confirming the recent view of the large variability of the deep-sea environments. Also RNA concentrations decreased with increasing water depth (range: 0.4–29.9 μg g^-1). The ratio of RNA to DNA did not show a clear spatial pattern but was characterized by significant changes between sampling periods. DNA concentrations were significantly correlated with protein and phytopigment concentrations in the sediment, indicating a possible relationship with the inputs of primary organic matter from the photic layer. Bacterial densities were generally high (range: 0.9–4.6×10⁸ cells g^-1) compared to other deep-sea environments and decreased with increasing water depth. Estimates of the bacterial contribution to the sedimentary genetic material indicated that bacterial-DNA accounted, on annual average, for a small fraction of the total DNA pool (4.3%) but that bacterial-RNA represented a significant fraction of the total sedimentary RNA (26%). Bacterial contribution to nucleic acids increased, even though irregularly, with increasing depth. In deep-sea sediments, changes in RNA concentrations appear to be largely dependent upon bacterial dynamics. Estimates of the overall living contribution to the DNA pools (i.e. microbial plus meiofaunal DNA) indicated that the large majority (about 90%) of the DNA in continental and deep-sea sediments of the eastern Mediterranean was detrital. The non-living DNA pools reach extremely high concentrations up to 0.41 g DNA m^-2 cm^-1. Thus, especially in deep benthic habitats, characterized by low inputs of labile organic compounds, detrital DNA could represent a suitable and high quality food source or a significant reservoir of nucleic acid precursors for benthic metabolism.     

Diversity and distribution of methane-oxidizing microbial communities associated with different faunal assemblages in a giant pockmark of the Gabon continental margin

A giant 800-m-diameter pockmark named REGAB was discovered on the Gabon continental margin actively emitting methane at a water depth of 3200 m. The microbial diversity in sediments from four different assemblages of chemosynthetic organisms, Mytilidae, Vesicomyidae, Siboglinidae and a bacterial mat, was investigated using comparative 16S rRNA gene sequence analysis. Aggregates of anaerobic methanotrophic archaea (ANME-2) and bacteria of the Desulfosarcina/Desulfococcus cluster were found in all four chemosynthetic habitats. Fluorescence in situ hybridization targeting the ANME-2/Desulfosarcina/Desulfococcus aggregates showed their presence few centimeters (3–5 cm) below the surface of sediment. 16S rRNA gene sequences from all known marine ANME groups were detected in the pockmark sediments, as well as from both known bacterial partners. The archaeal diversity was limited to the ANME cluster for all investigated samples. The bacterial diversity included members of the Proteobacteria, Bacilliales, Cytophaga/Flavobacteria, Verrucomicrobia, JS1 and Actinobacteria clusters. Bacterial 16S rRNA gene sequences related to those of known sulphide-oxidizing symbionts were recovered from tissues of several invertebrates including vesicomyid clams and siboglinid tubeworms of REGAB.     

Dispersal,survival and delayed growth of benthic foraminiferal propagules

New data support our previously published propagule dispersal hypothesis and show that propagules of some benthic foraminiferal species can survive for two years before growth commences. Following exposure to simulated shallow-water conditions, shallow-water species of benthic foraminifera appeared and grew in large numbers (commonly >100 ind/12 ml sediment) in the <32 µm-size sediment fraction collected from 320 m water depth in the Skagerrak basin (North Sea). None of the shallow-water species that grew abundantly (Planorbulina mediterranensis, Morulaeplecta bulbosa, Bolivina pseudoplicata, Cuneata arctica, Eggerelloides scaber, Gavelinopsis praegeri) seem to grow or reproduce at or in the vicinity of the sampling site. Consequently, they must have been transported there as <32 µm-sized individuals. Their sudden appearance when exposed to shallow-water conditions suggests that they had been transported to the sampling site as propagules and that they could survive in the sediments until conditions became suitable for growth and, for some, reproduction. The lack of agglutination on the proloculi of the agglutinated taxa that appeared in the growth-chambers may enhance their passive transport via currents and, thereby, dispersal. Of all the indigenous foraminiferal species that occur at the sampling site, only Textularia earlandi and Bolivinellina pseudopunctata continued to grow and reproduce when transferred from bathyal (320 m) to simulated shallow-water (0 m) conditions. The former is considered a highly opportunistic species. According to the literature, most of the morphospecies which grew in the experiments are cosmopolitan. Our results indicate substantial inter-specific differences in dispersal potential and support previous suggestions that among free-living species, some serial forms have the potential for long-distance dispersal. Still, oceanographic, physical and ecological boundaries and barriers constrain the distribution of most species. In addition to benthic foraminifera, Gromia spp. (rhizarian protists related to the foraminifera) grew in >60% of the experimental growth-chambers.     

Impact of lugworms (Arenicola marina) on mobilization and transport of fine particles and organic matter in marine sediments

We investigated the impact of sediment reworking fauna and hydrodynamics on mobilization and transport of organic matter and fine particles in marine sediments. Experiments were conducted in an annular flume using lugworms (Arenicola marina) as model organisms. The impact of lugworms on sediment characteristics and particle transport was followed through time in sediments experimentally enriched with fine particles (< 63 μm) and organic matter. Parallel experiments were run at low and high water current velocity (11 and 25 cm s^− 1) to evaluate the importance of sediment erosion at the sediment–water interface. There was no impact of fauna on sediment composition and particle transport at current velocity below the sediment erosion threshold. At current velocity above the erosion threshold, sediment reworking by lugworms resulted in dramatic particle transport (12 kg dry matter m^− 2) to an adjacent particle trap within 56 days. The transported matter was enriched 6–8 times in fine particles and organic matter when compared to the initial sediment. This study suggests that sediment reworking fauna is an important controlling factor for the particle composition of marine sediments. A. marina mediated sediment reworking greatly increases the sediment volume exposed to hydrodynamic forcing at the sediment–water interface, and through sediment resuspension control the content of fine particles and organic matter in the entire reworked sediment layer (> 20 cm depth).     

Biodiversity and community composition of sediment macrofauna associated with deep-sea Lophelia pertusa habitats in the Gulf of Mexico

Scleractinian corals create three-dimensional reefs that provide sheltered refuges, facilitate sediment accumulation, and enhance colonization of encrusting fauna. While heterogeneous coral habitats can harbor high levels of biodiversity, their effect on the community composition within nearby sediments remains unclear, particularly in the deep sea. Sediment macrofauna from deep-sea coral habitats (Lophelia pertusa) and non-coral, background sediments were examined at three sites in the northern Gulf of Mexico (VK826, VK906, MC751, 350–500 m depth) to determine whether macrofaunal abundance, diversity, and community composition near corals differed from background soft-sediments. Macrofaunal densities ranged from 26 to 125 individuals 32 cm⁻² and were significantly greater near coral versus background sediments only at VK826. Of the 86 benthic invertebrate taxa identified, 16 were exclusive to near-coral habitats, while 14 were found only in background sediments. Diversity (Fisher’s α) and evenness were significantly higher within near-coral sediments only at MC751 while taxon richness was similar among all habitats. Community composition was significantly different both between near-coral and background sediments and among the three primary sites. Polychaetes numerically dominated all samples, accounting for up to 70% of the total individuals near coral, whereas peracarid crustaceans were proportionally more abundant in background sediments (18%) than in those near coral (10%). The reef effect differed among sites, with community patterns potentially influenced by the size of reef habitat. Taxon turnover occurred with distance from the reef, suggesting that reef extent may represent an important factor in structuring sediment communities near L. pertusa. Polychaete communities in both habitats differed from other Gulf of Mexico (GOM) soft sediments based on data from previous studies, and we hypothesize that local environmental conditions found near L. pertusa may influence the macrofaunal community structure beyond the edges of the reef. This study represents the first assessment of L. pertusa-associated sediment communities in the GOM and provides baseline data that can help define the role of transition zones, from deep reefs to soft sediments, in shaping macrofaunal community structure and maintaining biodiversity; this information can help guide future conservation and management activities.     

Effects of the invasive polychaete Marenzelleria spp. on benthic processes and meiobenthos of a species-poor brackish system

The effects of the invasive polychaete Marenzelleria spp. on sediment processes and meiobenthos with an emphasis on free-living nematodes of the Vistula Lagoon (southern Baltic Sea) were investigated in a laboratory microcosm experiment. Marenzelleria occupies an open niche and its deep burying behaviour and feeding strategy represent a new function in the study area. Halos of oxidized sediment along Marenzelleria burrow walls indicated oxygen penetration into the burrows but the polychaete had no significant effect on porewater nutrient concentrations. The results showed, however, the density dependent effects of Marenzelleria on ammonium transport. An enhanced ammonium efflux was recorded at high polychaete densities (2000 ind. m^− 2) but not at low polychaete densities (300 ind. m^− 2).There was no observable impact of the polychaete on total meiobenthic numbers. There was, therefore, no indication that Marenzelleria caused meiofauna mortality. On the contrary, the polychaete significantly affected vertical distribution of meiofauna facilitating the colonization of deeper sediment depths and thus extending the habitat to be used by meiobenthos. In addition, Marenzelleria had a positive impact on the survival of turbellarians.Nevertheless, there was no effect of Marenzelleria on nematode assemblage structure and diversity, indicating that neither the physical presence nor the biological activity of the worm affected the nematode community. This suggests either 1. the limited impact of Marenzelleria on nematodes, resulting from the creation of simple, narrow and un-branched burrows, 2. poor response of nematode community resulting from their low abundance and diversity in the study area, or 3. the overriding role of the harsh chemical environment typical of sediments of the Vistula Lagoon, masking the effect of the bioturbator.     

Sulfate reduction and anaerobic methane oxidation in Black Sea sediments

Beyond the shelf break at ca. 150 m water depth, sulfate reduction is the only important process of organic matter oxidation in Black Sea sediments from the surface down to the sulfate–methane transition at 2–4 m depth. Sulfate reduction rates were measured experimentally with ³⁵SO₄²⁻, and the rates were compared with results of two diffusion-reaction models. The results showed that, even in these non-bioirrigated sediments without sulfide reoxidation, modeling strongly underestimated the high reduction rates near the sediment surface. A hybrid modeling approach, in which experimentally measured rates in the upper sediment layers force a model that includes also the deeper layers, probably provides the most realistic estimate of sulfate reduction rates. Areal rates of sulfate reduction were 0.65–1.43 mmol SO₄²⁻ m⁻² d⁻¹, highest in sediments just below the chemocline. Anaerobic methane oxidation accounted for 7–11% of the total sulfate reduction in slope and deep-sea sediments. Although this methane-driven sulfate reduction shaped the entire sulfate gradient, it was only equivalent to the sulfate reduction in the uppermost 1.5 cm of surface sediment. Methane oxidation was complete, yet the process was very sluggish with turnover times of methane within the sulfate–methane transition zone of 20 yr or more.     

Seafloor geological studies above active gas chimneys off Egypt (Central Nile Deep Sea Fan)

Four mud volcanoes of several kilometres diameter named Amon, Osiris, Isis, and North Alex and located above gas chimneys on the Central Nile Deep Sea Fan, were investigated for the first time with the submersible Nautile. One of the objectives was to characterize the seafloor morphology and the seepage activity across the mud volcanoes. The seepage activity was dominated by emissions of methane and heavier hydrocarbons associated with a major thermal contribution. The most active parts of the mud volcanoes were highly gas-saturated (methane concentrations in the water and in the sediments, respectively, of several hundreds of nmol/L and several mmol/L of wet sediment) and associated with significantly high thermal gradients (at 10 m below the seafloor, the recorded temperatures reached more than 40 °C). Patches of highly reduced blackish sediments, mats of sulphide-oxidizing bacteria, and precipitates of authigenic carbonate were detected, indicative of anaerobic methane consumption. The chemosynthetic fauna was, however, not very abundant, inhibited most likely by the high and vigorous fluxes, and was associated mainly with carbonate-crust-covered seafloor encountered on the southwestern flank of Amon. Mud expulsions are not very common at present and were found limited to the most active emission centres of two mud volcanoes, where slow extrusion of mud occurs. Each of the mud volcanoes is fed principally by a main narrow channel located below the most elevated areas, most commonly in the centres of the structures. The distribution, shape, and seafloor morphology of the mud volcanoes and associated seeps over the Central Nile Deep Sea Fan are clearly tectonically controlled.     

Modelling phytoplankton deposition to Chesapeake Bay sediments during winter–spring: interannual variability in relation to river flow

The often-rapid deposition of phytoplankton to sediments at the end of the spring phytoplankton bloom is an important component of benthic–pelagic coupling in temperate and high latitude estuaries and other aquatic systems. However, quantifying the flux is difficult, particularly in spatially heterogeneous environments. Surficial sediment chlorophyll-a, which can be measured quickly at many locations, has been used effectively by previous studies as an indicator of phytoplankton deposition to estuarine sediments. In this study, surficial sediment chlorophyll-a was quantified in late spring at 20–50 locations throughout Chesapeake Bay for 8 years (1993–2000). A model was developed to estimate chlorophyll-a deposition to sediments using these measurements, while accounting for chlorophyll-a degradation during the time between deposition and sampling. Carbon flux was derived from these estimates via C:chl-a = 75.Bay-wide, the accumulation of chlorophyll-a on sediments by late spring averaged 171 mg m⁻², from which the chlorophyll-a and carbon sinking fluxes, respectively, were estimated to be 353 mg m⁻² and 26.5 gC m⁻². These deposition estimates were ∼50% of estimates based on a sediment trap study in the mid-Bay. During 1993–2000, the highest average chlorophyll-a flux was in the mid-Bay (248 mg m⁻²), while the lowest was in the lower Bay (191 mg m⁻²). Winter–spring average river flow was positively correlated with phytoplankton biomass in the lower Bay water column, while phytoplankton biomass in that same region of the Bay was correlated with increased chlorophyll-a deposition to sediments. Responses in other regions of the Bay were less clear and suggested that the concept that nutrient enrichment in high flow years leads to greater phytoplankton deposition to sediments may be an oversimplification. A comparison of the carbon flux associated with the deposition of the spring bloom with annual benthic carbon budgets indicated that the spring bloom did not contribute a disproportionately large fraction of annual carbon inputs to Chesapeake Bay sediments. Regional patterns in chlorophyll-a deposition did not correspond with the strong regional patterns that have been found for plankton net community metabolism during spring.     

Depth-related structure and ecological significance of cold-seep communities—a case study from the Sea of Okhotsk

We discovered and investigated several cold-seep sites in four depth zones of the Sea of Okhotsk off Northeast Sakhalin: outer shelf (160–250 m), upper slope (250–450 m), intermediate slope (450–800 m), and Derugin Basin (1450–1600 m). Active seepage of free methane or methane-rich fluids was detected in each zone. However, seabed photography and sampling revealed that the number of chemoautotrophic species decreases dramatically with decreasing water depth. At greatest depths in the Derugin Basin, the seeps were inhabited by bacterial mats and bivalves of the families Vesicomyidae (Calyptogena aff. pacifica, C. rectimargo, Archivesica sp.), Solemyidae (Acharax sp.) and Thyasiridae (Conchocele bisecta). In addition, pogonophoran tubeworms of the family Sclerolinidae were found in barite edifices. At the shallowest sites, on the shelf at 160 m, the seeps lack chemoautotrophic macrofauna; their locations were indicated only by the patchy occurrence of bacterial mats.Typical seep-endemic metazoans with chemosynthetic symbionts were confined to seep sites at depths below 370 m. A comparative analysis of the structure of seep and background communities suggests that differences in predation pressure may be an important determinant of this pattern. The abundance of predators such as carnivorous brachyurans and asteroids, which can invade seeps from adjacent habitats and efficiently prey on sessile seep bivalves, decreased very pronouncedly with depth. We conclude from the obvious correlation with the conspicuous pattern in the distribution of seep assemblages that, on the shelf and at the upper slope, predator pressure may be high enough to effectively impede any successful settlement of viable populations of seep-endemic metazoans. However, there was also evidence that other depth-related factors, such as bottom-water current, sedimentary regimes, oxygen concentrations and the supply of suitable settling substrates, may additionally regulate the distribution of seep fauna in the area.As a consequence of the pronounced pattern in the distribution of seep communities, their ecological significance as food sources of surrounding background fauna increased with water depth. Isotopic analyses suggest that in the Derugin Basin seep colonists feed on chemoautotrophic seep organisms, either directly or by preying on metazoans with chemosynthetic symbionts. In contrast, seep organisms apparently do not contribute to the nutrition of the adjacent background fauna on the shelf and at the slope. In this area, elevated epifaunal abundances at seep sites were caused primarily by the availability of suitable settling substrates rather than by an enrichment of food supply.     

Short-term environmental variability in cold-water coral habitat at Viosca Knoll,Gulf of Mexico

The Lophelia pertusa community at Viosca Knoll (VK826) is the most extensive found to date in the Gulf of Mexico. As part of a multi-disciplinary study, the physical setting of this area was described using benthic landers, CTD transects and remotely operated vehicle observations. The site was broadly characterised into three main habitats: (1) dense coral cover that resembles biogenic reef complexes, (2) areas of sediment, and (3) authigenic carbonate blocks with sparse coral and chemosynthetic communities. The coral communities were dominated by L. pertusa but also contained numerous solitary coral species. Over areas that contained L. pertusa, the environmental conditions recorded were similar to those associated with communities in the north-eastern Atlantic, with temperature (8.5–10.6 °C) and salinity (～35) falling within the known species niche for L. pertusa. However, dissolved oxygen concentrations (2.7–2.8 ml l^?1) and density (σ_Θ, 27.1–27.2 kg m^?3) were lower and mass fluxes from sediment trap data appeared much higher (4002–4192 mg m^?2 d^?1). Yet, this species still appears to thrive in this region, suggesting that L. pertusa may not be as limited by lower dissolved oxygen concentrations as previously thought. The VK826 site experienced sustained eastward water flow of 10–30 cm s^?1 over the 5-day measurement period but was also subjected to significant short-term variability in current velocity and direction. In addition, two processes were observed that caused variability in salinity and temperature; the first was consistent with internal waves that caused temperature variations of 0.8 °C over 5–11 h periods. The second was high-frequency variability (20–30 min periods) in temperature recorded only at the ALBEX site. A further pattern observed over the coral habitat was the presence of a 24 h diel vertical migration of zooplankton that may form part of a food chain that eventually reaches the corals. The majority of detailed studies concerning local environmental conditions in L. pertusa habitats have been conducted within the north-eastern Atlantic, limiting most knowledge of the niche of this species to a single part of an ocean basin. Data presented here show that the corals at VK826 are subjected to similar conditions in temperature, salinity, and flow velocity as their counterparts in the north-east Atlantic, although values for dissolved oxygen and density (sigma-theta: σ_Θ) are different. Our data also highlight novel observations of short-term environmental variability in cold-water coral habitat.     

Comparative study on microphytobenthic pigments of muddy and sandy intertidal sediments of the Tagus estuary

The abundance and distribution of microphytobenthic pigments determined by HPLC (chlorophylls and carotenoids) were compared between muddy and sandy sediments of the Tagus estuary (Portugal). In the two types of sediment, with similar periods of illuminated emersion, chlorophyll a concentrations on a per area basis (mg m⁻²) were comparable (down to 2 mm). Pigment analysis also revealed similar microphytobenthic communities in terms of algal classes. Diatoms were the dominant microalgae, but cyanophytes, euglenophytes and phanerogam debris were also present. For both muddy and sandy sediments, microphytobenthic biomass showed a high level of variability both within and between two consecutive years. Microphytobenthos was highly stratified in the mud, with most of the chlorophyll a occurring in the top 500 μm. In the sand, relatively constant concentrations were found throughout the sediment profile down to 3 mm. This is probably related to deeper light penetration in sandy sediment and/or increased physical mixing caused by invertebrate activity or overlying currents, leading to the burial of an important fraction of the microphytobenthic cells. Differences observed in the intensity of sediment coloration of muddy and sandy sediments might have resulted from the different vertical distribution of benthic biomass.