Description: The phylogenetic relationship of sulphur-oxidising endosymbiotic bacteria from bivalves of the families Vesicomyidae (Calyptogena sp. C1, Calyptogena sp. C3), Solemyidae (Acharax sp.) and Thyasiridae (Conchocele sp.) from cold-seep habitats were determined by 16S rDNA nucleotide sequence analyses. The endosymbiotic bacteria form distinct groups within the gamma-Proteobacteria and are well separated from each other and from free-living sulphur-oxidising bacteria of the genera Beggiatoa, Halothiobacillus and Thiomicrospira. The endosymbiotic bacteria of Acharax sp. from cold seeps off Oregon, Indonesia and Pakistan have sequences highly similar to each other but quite distinct from other thiotrophic endosymbionts. This includes endosymbionts from Solemya spp., to which they are distantly related. Symbiotic bacteria of Conchocele sp. from a cold seep in the Sea of Okhotsk are similar to those of Bathymodiolus thermophilus and related species, as shown by their overall sequence similarity and by signature sequences. The endosymbiotic bacteria of Calyptogena spp. from cold seeps off Oregon and Pakistan are closely related to those of other vesicomyids. Endosymbiont species found off Oregon corresponded to 2 different clusters of Calyptogena spp. symbionts in the same samples. The results corroborate the hypothesis of a monophyletic origin of the symbionts in vesicomyid clams, and support the existence of deeply branching groups in solemyid symbionts and of divergent lines and distribution for thyasirid symbionts. The results also indicate that certain symbiont species cluster according to the depth distribution of their hosts, and that in consequence host species together with their symbionts may have undergone depth-specific adaptation and evolution.

Description: Gas hydrates occur at the sediment surface on the southern summit of Hydrate Ridge, Cascadia convergent margin. The hydrates are found in mounds several meters in diameter and up to 2 m high, and are covered by sediment and mats of the filamentous sulfur-oxidizing bacteria Beggiatoa. The mounds are surrounded by vesicomyid clams (Calyptogena pacifica, C. kilmeri), which in turn are encircled by solemyid bivalves (Acharax sp.). The zonation pattern of 3 species (Calyptogena spp. and Acharax sp., which harbor chemoautotrophic bacteria in their gills, and the chemoautotrophic Beggiatoa), is also reflected in a change in the entire community structure. Beggiatoa, Calyptogena spp. and Acharax sp. are shown to be characteristic species for the different communities. The Beggiatoa community directly overlaying the gas hydrates consists of seep endemic species in high densities: gastropods (Provanna laevis, P. lomana, Pyropelta corymba, Hyalogyrina sp. nov.), bivalves (Nuculana sp. nov.) and polychaetes (Ampharetidae, Polynoidae, Dorvilleidae). Based on pooled samples, the rarefaction curves show a decrease in species diversity in the Beggiatoa and Calyptogena communities. The hydrogen sulfide gradients in the porewater of sediments below the different communities dominated by either Beggiatoa, Calyptogena spp. or Acharax sp. vary by 3 orders of magnitude. The diffusive sulfide flux based on the measured sulfide concentration gradients is highest in Beggiatoa sp. communities (23 ± 13 mol m-2 yr-1), slightly less in Calyptogena communities (6.6 ± 2.4 mol m-2 yr-1), and low in Acharax communities (0.05 ± 0.05 mol m-2 yr-1). The difference in the sulfide environment is a factor influencing the distribution patterns of the chemoautotrophy-dependant and heterotrophic species at the deep-sea sediments containing gas hydrate.

Description: At Hook Ridge hydrothermal vent, a new species of Sclerolinum (Monilifera, Siboglinidae) was found at a water depth of 1,045 m. On the basis of investigations of multicores and gravity cores, the species habitat is characterized. Sclerolinum does not occur in sediments that are most strongly influenced by hydrothermal fluids, probably because of high temperature (up to 49°C) and precipitation of siliceous crusts. About 800 individuals m-2 occur in sediments that are only weakly exposed to hydrothermal flow and have the following characteristics: 20°C (15 cm sediment depth) to 21.5°C (bottom water), 18-40 cm yr-1 advection rates, pH 5.5, 〈25 µmol L-1 methane, 〈170 µmol L-1 sulfide, and 〈0.0054 mol m-2 yr-1 sulfide flux. Comparison with geochemical data from other reducing sediments indicates that the two groups of Siboglinidae, Monilifera and Frenulata, occur in sediments with low sulfide concentration and flux. In contrast, sulfurbased chemosynthetic organisms that typically occur at hydrothermal vents and cold seeps (e.g., Vestimentifera, vesicomyid clams, and bacterial mats) occur in sediments with higher sulfide availability; threshold values are around 500 µmol L-1 sulfide and 0.1 mol m-2 yr-1 sulfide fluxes. We did not find typical hydrothermal vent species at Hook Ridge hydrothermal vent, which might be explained by the unfavorable physicochemical habitat: At sites inhabited by Sclerolinum, sulfide availability appears to be too low, whereas at sites with higher sulfide availability, the temperatures might be too high, siliceous crust precipitation could preclude their occurrence, or both.

Description: The family Solemyidae represents ancestral protobranch bivalves with the shallow-water genus Solemya and the deep-sea genus Acharax. All known members of this family host symbiotic sulfur-oxidizing bacteria in their gill filaments. Analysis of 18S rRNA gene sequences of Acharax specimens from methane-seeps off Makran (Pakistan), Java (Indonesia), the Aleutian Trench and off the Oregon, Costa Rica, and Peru margins revealed that Solemya spp. and Acharax spp. are well-separated genetically. This supports the current systematic distinction based on morphological criteria. We found 2 clearly distinct clusters within the genus Acharax, with specimens from the Makran, Oregon and Peru (MOP) margins in one (MOP–Acharax) cluster, and those from Java, the Aleutian Trench and Costa Rica (JAC) in the other (JAC–Acharax) cluster. The separation of MOP– and JAC–Acharax clusters from each other and from Solemya (S. reidi and S. velum) is well-supported by phylogenetic calculations employing maximum likelihood and maximum parsimony. Compared to genetic distances among other protobranch groups, distances between the MOP– and JAC–Acharax clusters would justify the affiliation of these clusters to separate species. This implies that species differentiation in Acharax based on shell morphology is likely to underestimate true species diversity within this taxon. Furthermore, our results support the hypothesis that genetic separation of Solemya and Acharax is congruent with the phylogeny of their bacterial endosymbionts.