Distribution and temporal variation of mega‐fauna at the Regab pockmark (Northern Congo Fan), based on a comparison of videomosaics and geographic information systems analyses

The Regab pockmark is a large cold seep area located 10 km north of the Congo deep sea channel at about 3160 m water depth. The associated ecosystem hosts abundant fauna, dominated by chemosynthetic species such as the mussel Bathymodiolus aff. boomerang, vestimentiferan tubeworm Escarpia southwardae, and vesicomyid clams Laubiericoncha chuni and Christineconcha regab. The pockmark was visited during the West African Cold Seeps (WACS) cruise with RV Pourquoi Pas? in February 2011, and a 14,000‐m² high‐resolution videomosaic was constructed to map the most populated area and to describe the distribution of the dominant megafauna (mussels, tubeworms and clams). The results are compared with previous published works, which also included a videomosaic in the same area of the pockmark, based on images of the BIOZAIRE cruise in 2001. The 10‐year variation of the faunal distribution is described and reveals that the visible abundance and distribution of the dominant megafaunal populations at Regab have not changed significantly, suggesting that the overall methane and sulfide fluxes that reach the faunal communities have been stable. Nevertheless, small and localized distribution changes in the clam community indicate that it is exposed to more transient fluxes than the other communities. Observations suggest that the main megafaunal aggregations at Regab are distributed around focused zones of high flux of methane‐enriched fluids likely related to distinct smaller pockmark structures that compose the larger Regab pockmark. Although most results are consistent with the existing successional models for seep communities, some observations in the distribution of the Regab mussel population do not entirely fit into these models. This is likely due to the high heterogeneity of this site formed by the coalescence of several pockmarks. We hypothesize that the mussel distribution at Regab could also be controlled by the occurrence of zones of both intense methane fluxes and reduced efficiency of the anaerobic oxidation of methane possibly limiting tubeworm colonization.     

Habitat,growth and physiological ecology of a basaltic community of Ridgeia piscesae from the Juan de Fuca Ridge

The vestimentiferan tubeworm Ridgeia piscesae is an ecosystem-structuring organism in the hydrothermal vent environments of the Northeast Pacific. During this study, a single representative aggregation of the long-skinny morphotype of R. piscesae from the main endeavor segment was monitored for 3 yr before being collected in its entirety with a hydraulically actuated collection device manipulated in situ by a research vehicle. Vestimentiferan growth rates in this aggregation were determined by staining the exterior of the tubes and measuring newly deposited tube sections. The average growth rate of R. piscesae in this aggregation was very low in both years of the growth study (3.2 mm yr⁻¹). Although the incidence of plume damage from partial predation was very high (>95%), mortality was very low (<4% yr⁻¹). The distribution and the very tight clustering of recently recruited individuals indicated gregarious settlement behavior that is hypothesized to be partly due to biotic cues from settled larvae. Coupled measurements of vent fluid sulfide concentration and temperature were used to calculate the exposure of the vestimentiferans to sulfide from short- and long-term temperature monitoring. Plume-level temperature records indicate that most of the time individuals in this aggregation were exposed to extremely low levels of vent fluid, and therefore sulfide (<0.1 μM), while their posterior sections were consistently exposed to sulfide concentrations in the 100 μM range. A rootball-like structure formed the common base of the aggregation. In contrast to the anterior sections of the tubeworm tubes, the portions of the tubes within the “rootball” were freely permeable to sulfide. The results of this study show that R. piscesae, unlike vestimentiferans from the East Pacific Rise, can survive and grow in areas of low diffuse vent flow with very low plume-level exposure to sulfide. We propose that this morphotype of R. piscesae has the ability to acquire sulfide from sources near their posterior ends, similar to some species of cold seep vestimentiferans from the Gulf of Mexico. The ability of this single species of vestimentiferan to survive low exposure to vent flow with low mortality coupled with sulfide uptake across posterior tube sections may help explain the occurrence of a single vent vestimentiferan species in a wide variety of habitat conditions at hydrothermal vent sites in the Northeast Pacific.     

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.     

Patterns of growth in cold-seep vestimenferans including Seepiophila jonesi: a second species of long-lived tubeworm

Seepiophila jonesi is a vestimentiferan tubeworm (Siboglinidae: Polychaeta) inhabiting the cold seeps of the upper slope of the Gulf of Mexico. It commonly co‐occurs with Lamellibrachia luymesi, which is among the most long‐lived non‐clonal animals known. The growth pattern of S. jonesi is best described by a model including a size‐specific probability of growth and an average growth rate that does not vary with individual size. This model, based on growth data from in situ staining and collection approximately 1 year later, predicts that S. jonesi is very slow growing and may attain ages comparable with L. luymesi. The efficacy of this model in describing L. luymesi growth rate was assessed, but the previously employed model of declining growth rate with individual size provided the better fit to the empirical data. Comparisons of both S. jonesi and L. luymesi growth rates among sites and among aggregations within a site indicate that there is some degree of habitat variability contributing to differences in growth rates. However, position of the anterior end of the worm within an aggregation did not have a significant effect on growth rate in comparisons among groups of L. luymesi from different distances from the center of an aggregation. The evolution of longevity in these species of vestimentiferans was favored by the relative stability of the seep habitat and sulfide sources, in contrast to the hydrothermal vent environment inhabited by relatively short‐lived and fast‐growing vestimentiferan species.     

Phylogenetic relationship between symbionts of tubeworm <Emphasis Type="Italic">Lamellibrachia satsuma</Emphasis> and the sediment microbial community in Kagoshima Bay

Vestimentiferan tubeworms acquire their symbionts through horizontal transmission from the surrounding environment. In the present study, we constructed a 16S rRNA gene clone library to investigate the phylogenetic relationship between diverse microbes in the sediment and symbiotic bacteria in the trophosome of the tubeworm, Lamellibrachia satsuma, from Kagoshima Bay, Japan. Two symbiotic bacterial phylotypes belonging to the classes γ- and ε-Proteobacteria were found from this tubeworm trophosome. They were very closely related to the symbionts of several other marine invertebrates. The most predominant bacteria in the sediment were ε-Proteobacteria. A broad diversity of bacteria belonged to non-proteobacterial phyla such as Planctomycetes, Acidobacteria, and Chloroflexi was observed. The presence of sulfur oxidizers (i.e., ε-Proteobacteria and γ-Proteobacteria) and sulfur reducers (i.e., δ-Proteobacteria) may play a significant role in the sulfur cycle in these habitats and provide multiple sources of nutrition to the cold-seep communities. Closely related clones of ε-Proteobacteria symbiont in the species level and of γ-Proteobacteria symbiont in the genus level were found in the surrounding sediment. The similarity of symbiont clones of L. satsuma with other symbionts and free-living bacteria suggests the possibility of opportunistic symbiosis in ε-Proteobacteria and the co-evolution of γ-Proteobacteria having occurred after symbiosis with the tubeworms.     

The Co-occurrence of Methanotrophic and Chemoautotrophic Sulfur-Oxidizing Bacterial Symbionts in a Deep-sea Mussel

Abstract. In April 1990 a new species of mytilid containing bacterial endosymbionts was discovered at a deep-water seep site within Alaminos Canyon in the Gulf of Mexico. Activities of ribulose bisphosphate carboxylase/oxygenase and ATP sulfurylase, as well as the presence of substantial levels of elemental sulfur in the gill tissues indicate the presence of chemoautotrophic sulfur-oxidizing symbionts in the gills. Methanol dehydrogenase activity and the tissue stable carbon isotope ratios indicate the presence of methanotrophic bacteria in the gills of the same animals. Two distinct size classes and morphological types of gram negative bacteria are visible in transmission electron micrographs of the gill tissue, one of which contains the complex internal membranes typical of methanotrophs. Both general types of symbionts have been demonstrated singly in related species of deep-sea mytilids. In this species, however, both types are found in single individuals, often within the same cell vacuole.     

Population structure of two deep sea tubeworms,Lamellibrachia luymesi and Seepiophila jonesi,from the hydrocarbon seeps of the Gulf of Mexico

Vestimentiferan tubeworms are a group of large sessile marine polychaete annelids (family Siboglinidae) found in the regions of hydrothermal venting or seepage of the reduced chemical hydrogen sulfide. Hydrocarbon seeps on the Louisiana Slope of the Gulf of Mexico support large communities of the co-occurring vestimentiferan species Lamellibrachia luymesi and Seepiophila jonesi. These sessile species have the opportunity to disperse between the patchy sites of active seepage on the seafloor during a planktonic larval stage. However, it is unclear whether dispersal occurs at a local or global scale. Four (L. luymesi) and seven (S. jonesi) microsatellite loci were used to test for population substructure among ten hydrocarbon seep sites on the Louisiana Slope. Both species showed high levels of allelic diversity, averaging 18.5 (L. luymesi) and 22 (S. jonesi) alleles/locus, respectively, and high observed heterozygosity at all microsatellite loci (0.71–0.9 in L. luymesi, 0.27–0.84 in S. jonesi). The two species showed a significant deficiency in heterozygotes compared to that predicted under the Hardy–Weinberg equilibrium. L. luymesi showed a small but significant amount of population structure, with a positive correlation between genetic and geographic distance among the sample sites spanning 540 km. S. jonesi, in contrast, showed no evidence for isolation by distance, but did show a significant genetic difference between aggregations of different ages. These results suggest that these two species differ in how larvae are able to colonize new seep sites through space (L. luymesi) and though time (S. jonesi).     

Mineralization of vestimentiferan tubes at methane seeps on the Congo deep-sea fan

Vestimentiferan tube worms are prominent members of modern methane seep communities and are totally reliant as adults on symbiotic sulphide-oxidizing bacteria for their nutrition. The sulphide is produced in the sediment by a biochemical reaction called the anaerobic oxidation of methane (AOM). A well-studied species from the Gulf of Mexico shows that seep vestimentiferans ‘mine’ sulphide from the sediment using root-like, thin walled, permeable posterior tube extensions, which can also be used to pump sulphate and possibly hydrogen ions from the soft tissue back into the sediment to increase the local rate of AOM. The ‘root-balls’ of exhumed seep vestimentiferans are intimately associated with carbonate nodules, which are a result of AOM. We have studied vestimentiferan specimens and associated carbonates from seeps at the Kouilou pockmark field on the Congo deep-sea fan and find that some of the posterior ‘root’ tubes of living specimens are enclosed with carbonate indurated sediment and other, empty examples are partially or completely replaced by the carbonate mineral aragonite. This replacement occurs from the outside of the tube wall inwards and leaves fine-scale relict textures of the original organic tube wall. The process of mineralization is unknown, but is likely a result of post-mortem microbial decay of the tube wall proteins by microorganisms or the precipitation from locally high flux of AOM derived carbonate ions. The aragonite-replaced tubes from the Kouilou pockmarks show similar features to carbonate tubes in ancient seep deposits and make it more likely that many of these fossil tubes are those of vestimentiferans. These observations have implications for the supposed origination of this group, based on molecular divergence estimates.     

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.     

Formation of seep bubble plumes in the Coal Oil Point seep field

The fate of marine seep gases (transport to the atmosphere or dissolution, and either bacterial oxidation or diffusion to the atmosphere) is intimately connected with bubble and bubble-plume processes, which are strongly size-dependent. Based on measurements with a video bubble measurement system in the Coal Oil Point seep field in the Santa Barbara Channel, California, which recorded the bubble-emission size distribution (Φ) for a range of seep vents, three distinct plume types were identified, termed minor, major, and mixed. Minor plumes generally emitted bubbles with a lower emission flux, Q, and had narrow, peaked Φ that were well described by a Gaussian function. Major plumes showed broad Φ spanning very small to very large bubbles, and were well described by a power law function. Mixed plumes showed characteristics of both major and minor plume classes, i.e., they were described by a combination of Gaussian and power law functions, albeit poorly. To understand the underlying formation mechanism, laboratory bubble plumes were created from fixed capillary tubes, and by percolating air through sediment beds of four different grain sizes for a range of Q. Capillary tubes produced a Φ that was Gaussian for low Q. The peak radius of the Gaussian function describing Φ increased with capillary diameter. At high Q, they produced a broad distribution, which was primarily described by a power law. Sediment-bed bubble plumes were mixed plumes for low Q, and major plumes for high Q. For low-Q sediment-bed Φ, the peak radius decreased with increasing grain size. For high Q, sediment-bed Φ exhibited a decreased sensitivity to grain size, and Φ tended toward a power law, similar to that for major seep plumes.     

Cold-seep assemblages on a giant pockmark off West Africa: spatial patterns and environmental control

A giant pockmark colonised by dense cold‐seep assemblages near 3160 m depth along the Congo‐Angola margin has been surveyed by the ROV Victor 6000. The quantitative distribution of chemosynthetic communities was mapped along the dive tracks from a video study using GIS and image mosaicking. Several types of faunal assemblages, either dominated by bivalves of the families Mytilidae (Bathymodiolus sp.) or Vesicomyidae (Calyptogena sp., ‘Vesicomya’ aff. chuni), or by Siboglinidae polychaetes (Escarpia southwardae) were mapped over the 800‐m diameter pockmark area and sampled for fauna, water and sediment. The isotopic analyses (δ¹³C) of tissues from symbiont‐bearing species were within the range typical of nutrition via symbiosis using methane for mussels and sulphide for vesicomyids and siboglinids. The living chemosynthetic communities were distributed on a SW‐NE axis, corresponding to the expression at the sediment surface of a main buried channel providing fluids to the pockmark. The site was characterised by a more active central part in a depression with abundant carbonate concretions where high‐density clusters of siboglinids and mytilids dominate. Large fields of dead and live vesicomyids with a lower mean density were observed in the external areas. The mean coverage of each of the three symbiotic taxa in these two contrasted areas was estimated from mosaic analysis and was up to 30% in the central area dominated by E. southwardae bushes (23%). Symbiont‐bearing species distribution was consistent with methane concentrations in seawater that were generally higher in mytilid beds than in the vicinity of siboglinids and vesicomyids. A Principal Component Analysis performed on environmental factors at the ten sampling sites revealed that 37% of the observed variance in the distribution of symbiont‐bearing species may be explained by variation in both methane and oxygen concentrations, while a Canonical Redundancy Analysis selected methane concentration as the only variable which explains symbiont‐bearing species densities. This spatial distribution of chemosynthetic species at the pockmark scale may reflect temporal patterns of succession of both substrate and fauna, and may be related to different individual pockmarks visible on the microbathymetry mapped using ROV data.     

Ecophysiological differences between vesicomyid species and metabolic capabilities of their symbionts influence distribution patterns of the deep‐sea clams

This study provides an analysis of vesicomyid bivalve–symbiont community distribution across cold seep and hydrothermal vent areas in the Guaymas Basin (Gulf of California, Mexico). Using a combination of morphological and molecular approaches including fluorescent in situ hybridization (FISH), and electronic microscopy observations, vesicomyid clam species and their associated symbionts were characterized and results were analyzed in light of geochemical conditions and other on‐site observations. A greater diversity of vesicomyids was found at cold seep areas, where three different species were present (Phreagena soyoae [syn. kilmeri], Archivesica gigas, and Calyptogena pacifica). In contrast, A. gigas was the only species sampled across the hydrothermal vent area. The same haplotype of A. gigas was found in both hydrothermal vent and cold seep areas, highlighting possible contemporary exchanges among neighboring vents and seeps. In either ecosystem, molecular characterization of the symbionts confirmed the specificity between symbionts and hosts and supported the hypothesis of a predominantly vertical transmission. In addition, patterns of clams could reflect potential niche preferences for each species. The occurrence of numerous traces of vesicomyid movements on sediments in the sites colonized by A. gigas seemed to indicate that this species might have a better ability to move. Furthermore, variation in gill sulfur content could reveal a higher plasticity and sulfur storage capacity in A. gigas. Thus, the distribution of vesicomyid species across the chemosynthetic areas of the Guaymas Basin could be explained by differences in biological traits of the vesicomyid species that would allow A. gigas to more easily exploit transient and punctual sources of available sulfide than P. soyoae.     

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.     

Community structure of infaunal macrobenthos around vestimentiferan thickets at the San Clemente cold seep,NE Pacific

The San Clemente cold seep lies within 100–200 km of other reducing habitats in the NE Pacific, offering an opportunity to compare diversity and species overlap among reducing habitats (i.e. whale‐, kelp‐, and wood‐falls) at similar depths within a single region. Video observations from the research submersible Alvin at the San Clemente seep (1800 m depth) indicated clumps (‘thickets’) of vestimentiferans distributed as meter‐scale patches interspersed with vesicomyid clam beds and black sediments. Sediment‐core samples were collected at distances of 0 to 80–200 m along randomly oriented transects radiating outward from vestimentiferan thickets to evaluate changes in macrofaunal community structure from thickets into the background community. Macrofaunal abundance was elevated at distances of 0–1 m compared to 80–200 m (i.e. the ‘background’ community). The tube‐building frenulate worms Siboglinum spp., along with peracarid crustaceans, dominated sediments within 1 m of vestimentiferan thickets. Species diversity was depressed within 1 m of thickets but with high rates of species accumulation, suggesting that seep sites greatly increase sediment heterogeneity and facilitate colonization by non‐background macrofaunal species. Stable isotope data indicate chemosynthetic nutrition for some dominant macrofaunal species within 1 m of tubeworm thickets. The macrofaunal community near vestimentiferan thickets in San Clemente seep contains intermediate levels of species richness and diversity compared to other deep‐sea seep areas in the northeast Pacific. There was low species overlap between the San Clemente seep macrofauna and communities in reducing habitats near wood‐, whale‐, and kelp‐falls at similar depths within the region, suggesting that seeps harbor a distinct infaunal community.     

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.     

Effects of the oophagous bivalve Acesta oophaga on the morphology and fecundity of its deep‐sea tubeworm host,Lamellibrachia luymesi

The deep‐sea bivalve Acesta oophaga lives attached to the anterior end of the vestimentiferan tubeworm, Lamellibrachia luymesi, at cold methane seeps. The bivalve is found almost exclusively on female tubeworms, where it consumes the lipid‐rich eggs of L. luymesi that are spawned year round (Biological Bulletin, 209, 2005, 87). It is apparent that A. oophaga benefits directly from this close association, but the consequences for the tubeworm host may be more complicated than just a simple predator–prey interaction. Since A. oophaga completely surrounds the tube opening and plume of the worm, it is likely that its presence would limit oxygen uptake by L. luymesi, thereby inhibiting worm growth and reproduction. We hypothesized that occupied tubeworms would compensate for this by growing larger plumes for oxygen uptake. To explore the effects of bivalve presence/absence on female tubeworms, several morphological features, including body size, plume length, tube diameter, and tube segment length, as well as instantaneous fecundity, were compared. Results suggest that the mere presence of A. oophaga has a significant impact on the morphology of its host worm, as all measures of worm size, except for tube segment length, were significantly greater with clams present. Additionally, instantaneous fecundity was 3.5 times higher in occupied worms, implying that tubeworms are not oxygen‐deprived or energy limited as a result of bivalve presence. Our findings suggest that the association between these two deep‐sea organisms may be a more complex form of symbiosis than the simple predator–prey relationship, as previously thought.     

ROV study of a giant pockmark on the Gabon continental margin

A giant, 800-m wide pockmark, called Regab, was discovered along the Equatorial African margin at 3160-m water depth and was explored by remote operated vehicle (ROV) as part of the Zaiango (1998–2000) and Biozaire (2001–2003) projects carried out conjointly by TOTAL and a number of French research institutes. A microbathymetric map obtained using the ROV sensors shows that the pockmark actually consists of a cluster of smaller pockmarks aligned N70 along a 15-m deep depression. Methane was recorded all over the pockmark, the highest values along the axis of the depression where massive carbonate crusts and dense seep communities were also found. Several faunal species belong to the Vesicomyidae and Mytilidae bivalve families, as well as to Siboglinidae (Vestimentifera) tubeworms. Preliminary analyses confirm their association with symbiotic bacteria, thus documenting their dependence on fluid seeps. The pockmark appears to be related to an infilled channel, visible on the seismic data 300 m below the seafloor, which may act as a reservoir for biogenic fluids supplied to the trap from the surrounding sediments.     

Methane gas Seeps Along the Oxic/Anoxic Gradient in the Black Sea: Manifestations, Biogenic Sediment Compounds and Preliminary Results on Benthic Ecology

Abstract. Sediment parameters (grain size, organic content, chloroplastic pigments, Adenylates, Potential hydrolytic activity and electron transport system activity) and benthic biota were studied in methane seep area south-west of the Crimean Peninsula in the Black Sea over a depth range from 60 to 260m. A control transect with similar depths was performed in an adjacent nonseep area. Methane seepage in this region occurs from 35 to ˜ 85m depth, passing the oxic anoxic interface zone at 130 to 180m. The methane seep areas were characterized by abundant carbonate precipitates which occurred in various shapes from small, flat structures to tall chimneys with increasing anoxia. The carbonates were associated with distinct bacterial mats. Most of the measured biochemical parameters in the sediment were quite similar in the seep and nonseep areas. The content of organic matter was higher and grain size was more uniform in the nonseep areas. However, the seep areas were charaterized by highter proportions of the larger fauna (size classes 0.5–1 mm and ≥ 1mm) as well as increased total numbers of benthic fauna in the suboxic and upper anoxic zone. The animals did not show any seep-specific adaptations. In addition to methane seepage, highly variable hydrochemical and sedimentary conditions on the lower shelf and upper slope may also play an important role in structuring the composition and distribution of the benthic fauna.     

Importance of seep primary production to Lophelia pertusa and associated fauna in the Gulf of Mexico

To investigate the importance of seep primary production to the nutrition of Lophelia pertusa and associated communities and examine local trophic interactions, we analyzed stable carbon, nitrogen, and sulfur compositions in seven quantitative L. pertusa community collections. A significant seep signature was only detected in one of the 35 species tested (Provanna sculpta, a common seep gastropod) despite the presence of seep fauna at the three sample sites. A potential predator of L. pertusa was identified (Coralliophila sp.), and a variety of other trophic interactions among the fauna occupying the coral framework were suggested by the data, including the galatheid crab Munidopsis sp. 2 feeding upon hydroids and the polychaete Eunice sp. feeding upon the sabellid polychaete Euratella sp. Stable carbon abundances were also determined for different sections of L. pertusa skeleton representing different stages in the growth and life of the aggregation. There was no temporal trend detected in the skeleton isotope values, suggesting that L. pertusa settles in these areas only after seepage has largely subsided. Isotope values of individual taxa that were collected from both L. pertusa and vestimentiferan habitats showed decreasing reliance upon seep primary production with average age of the vestimentiferan aggregation, and finally, no seep signature was detected in the coral collections. Together our data suggest that it is the presence of authigenic carbonate substrata, a product of past seep microbial activity, as well as hydrodynamic processes that drive L. pertusa occurrence at seep sites in the Gulf of Mexico, not nutritional dependence upon primary production by seep microbes.     

Macrobenthic infaunal communities associated with deep‐sea hydrocarbon seeps in the northern Gulf of Mexico

There are thousands of seeps in the deep ocean worldwide; however, many questions remain about their contributions to global biodiversity and the surrounding deep‐sea environment. In addition to being globally distributed, seeps provide several benefits to humans such as unique habitats, organisms with novel genes, and carbon regulation. The purpose of this study is to determine whether there are unique seep macrobenthic assemblages, by comparing seep and nonseep environments, different seep habitats, and seeps at different depths and locations. Infaunal community composition, diversity, and abundance were examined between seep and nonseep background environments and among three seep habitats (i.e., microbial mats, tubeworms, and soft‐bottom seeps). Abundances were higher at seep sites compared to background areas. Abundance and diversity also differed among microbial mat, tubeworm, and soft‐bottom seep habitats. Although seeps contained different macrobenthic assemblages than nonseep areas, infaunal communities were also generally unique for each seep. Variability was 75% greater within communities near seeps compared to communities in background areas. Thus, high variability in community structure characterized seep communities rather than specific taxa. The lack of similarity among seep sites supports the idea that there are no specific infauna that can be used as indicators of seepage throughout the northern Gulf of Mexico, at least at higher taxonomic levels.