An overview of chemosynthetic symbioses in bivalves from the North Atlantic and Mediterranean Sea

Deep-sea bivalves found at hydrothermal vents, cold seeps and organic falls are sustained by chemosynthetic bacteria that ensure part or all of their carbon nutrition. These symbioses are of prime importance for the functioning of the ecosystems. Similar symbioses occur in other bivalve species living in shallow and coastal reduced habitats worldwide. In recent years, several deep-sea species have been investigated from continental margins around Europe, West Africa, eastern Americas, the Gulf of Mexico, and from hydrothermal vents on the Mid-Atlantic Ridge. In parallel, numerous, more easily accessible shallow marine species have been studied. Herein we provide a summary of the current knowledge available on chemosymbiotic bivalves in the area ranging west-to-east from the Gulf of Mexico to the Sea of Marmara, and north-to-south from the Arctic to the Gulf of Guinea. Characteristics of symbioses in 53 species from the area are summarized for each of the five bivalve families documented to harbor chemosynthetic symbionts (Mytilidae, Vesicomyidae, Solemyidae, Thyasiridae and Lucinidae). Comparisons are made between the families, with special emphasis on ecology, life cycle, and connectivity. Chemosynthetic symbioses are a major adaptation to ecosystems and habitats exposed to reducing conditions. However, relatively little is known regarding their diversity and functioning, apart from a few “model species” on which effort has focused over the last 30 yr. In the context of increasing concern about biodiversity and ecosystems, and increasing anthropogenic pressure on oceans, we advocate a better assessment of the diversity of bivalve symbioses in order to evaluate the capacities of these remarkable ecological and evolutionary units to withstand environmental change

DeepIso - A global open database of stable isotope ratios and elemental contents for deep-sea ecosystems.

Stable isotopes have been instrumental to many key-findings about deep-sea ecosystem functioning, particularly in chemosynthesis-based habitats (hydrothermal vents, cold seeps). However, constraining sampling logistics commonly limit the scope, extent, and therefore insights drawn from isotope-based deep-sea studies. Overall, much is left to discover about factors globally influencing food web structure in deep-sea ecosystems. In this context, it is crucial that all generated data are easily discoverable, available and reusable. DeepIso is a collaborative effort to produce a global compilation of stable isotope ratios and elemental contents in organisms from deep-sea ecosystems. In doing so, it aims to provide the deep-sea community with an open data analysis tool that can be used in the context of future ecological research, and to help deep-sea researchers to use stable isotope markers at their full efficiency. The database, accessible under CC-BY licence at https://doi.org/10.17882/76595, currently contains 18677 fully documented measurements. Archived parameters include δ13C (n = 4587), δ15N (n = 4388), δ34S (n = 951), %C (n = 2740), %N (n = 2741), %S (n = 752) and C/N ratio (n = 2518). Those measurements pertain to 4378 distinct samples belonging to 493 taxa, plus sediments, suspended particulate organic matter, plankton and detritus. Samples were taken between 1989 and 2018 in multiple environments (hydrothermal vents, cold seeps, cold water coral reefs, and other benthic or pelagic environments) and at depths ranging up to 5209 meters. To maximise the scope of the project, we are looking to integrate more data, either underlying published articles, from grey literature, or even unpublished. We’ll be happy to assist in data formatting and publication. If you are willing to contribute, or simply if you have feedback about the database, please get in touch via [email protected]

Building up DeepIso - A global open database of stable isotope ratios and elemental contents for deep-sea ecosystems.

Stable isotopes have been instrumental to many key-findings about deep-sea ecosystem functioning, particularly in chemosynthesis-based habitats (hydrothermal vents, cold seeps). However, constraining sampling logistics commonly limit the scope, extent, and therefore insights drawn from isotope-based deep-sea studies. Overall, much is left to discover about factors globally influencing food web structure in deep-sea ecosystems. In this context, it is crucial that all generated data are easily discoverable, available, and reusable. DeepIso is a collaborative effort to produce a global compilation of stable isotope ratios and elemental contents in organisms from deep-sea ecosystems. In doing so, it aims to provide the deep-sea community with an open data analysis tool that can be used in the context of future ecological research, and to help deep-sea researchers to use stable isotope markers at their full efficiency. The database, accessible under CC-BY licence at https://doi.org/10.17882/76595, currently contains 18677 fully documented measurements. Archived parameters include δ13C (n = 4587), δ15N (n = 4388), δ34S (n = 951), %C (n = 2740), %N (n = 2741), %S (n = 752) and C/N ratio (n = 2518). Those measurements pertain to 4378 distinct samples belonging to 493 taxa, plus sediments, suspended particulate organic matter, plankton and detritus. Samples were taken between 1989 and 2018 in multiple environments (hydrothermal vents, cold seeps, cold water coral reefs, and other benthic or pelagic environments) and at depths ranging up to 5209 meters. To maximise the scope of the project, we are looking to integrate more data, either underlying published articles, from grey literature, or even unpublished. We’ll be happy to assist in data formatting and publication. If you are willing to contribute, or simply if you have feedback about the database, please get in touch via [email protected]

A threefold perspective on the role of a pockmark in benthic faunal communities and biodiversity patterns

Pockmarks are circular-shaped depressions that increase seabed heterogeneity and are characterized by discontinuous fluid emissions. To understand how environmental conditions of pockmarks affect the structure of macroand meiofauna, we investigated two sites in a pockmark field in the northwestern Madagascar margin. In a comparative approach, we explored the community structure of the dominant taxa (Polychaeta, Nematoda and hyaline foraminifera) in each component (macro-, metazoan meiofauna and foraminifera, respectively). The investigated active pockmark showed approximately two times higher meiofauna abundance compared to in a site away from another pockmark field, but macrofauna showed the opposite trend, with almost half density at the pockmark site. However, at both sites, macroand meiofauna showed higher richness and abundance values in the top well-oxygenated layers of the sediment than in the underlaying ones. Polychaeta and Nematoda showed lower richness in the pockmark, opposed to hyaline foraminiferans, but lower evenness in the pockmark was found for the three groups. The detection of gas flares in the water column attests of the recent activity within the pockmark. High amount of sulfur-bearing minerals (mainly pyrite) evidences a production of dissolved free sulfides (not detected at the time of sampling) by sulfate reduction process driven by organic matter degradation and anaerobic oxidation of methane. Furthermore, recent increase in sedimentation rates in the past 70 years and organic matter inputs could have led to higher organic matter degradation rates resulting in reduced conditions and a high oxygen consumption. All this together seem to act as key factors in the determination of variation in richness, abundance and community composition of macrofauna and meiofauna. Additionally, some taxa seem to be more tolerant to these extreme conditions, such as species belonging to the Nematoda genus Desmodora and the phylum Kinorhyncha, which are highly abundant in the pockmark, and hence, may be considered as potential bioindicators of pockmark activity in this area. Further studies are required for a better assessment

Multi-disciplinary investigation of fluid seepage on an unstable margin: The case of the Central Nile deep sea fan

We report on a multidisciplinary study of cold seeps explored in the Central Nile deep-sea fan of the Egyptian margin. Our approach combines in situ seafloor observation, geophysics, sedimentological data, measurement of bottom-water methane anomalies, pore-water and sediment geochemistry, and 230Th/U dating of authigenic carbonates. Two areas were investigated, which correspond to different sedimentary provinces. The lower slope, at ∼ 2100 m water depth, indicates deformation of sediments by gravitational processes, exhibiting slope-parallel elongated ridges and seafloor depressions. In contrast, the middle slope, at ∼ 1650 m water depth, exhibits a series of debris-flow deposits not remobilized by post-depositional gravity processes. Significant differences exist between fluid-escape structures from the two studied areas. At the lower slope, methane anomalies were detected in bottom-waters above the depressions, whereas the adjacent ridges show a frequent coverage of fractured carbonate pavements associated with chemosynthetic vent communities. Carbonate U/Th age dates (∼ 8 kyr BP), pore-water sulphate and solid phase sediment data suggest that seepage activity at those carbonate ridges has decreased over the recent past. In contrast, large (∼ 1 km2) carbonate-paved areas were discovered in the middle slope, with U/Th isotope evidence for ongoing carbonate precipitation during the Late Holocene (since ∼ 5 kyr BP at least). Our results suggest that fluid venting is closely related to sediment deformation in the Central Nile margin. It is proposed that slope instability leads to focused fluid flow in the lower slope and exposure of ‘fossil’ carbonate ridges, whereas pervasive diffuse flow prevails at the unfailed middle slope

Phylogeny and Diversification Patterns among Vesicomyid Bivalves

Vesicomyid bivalves are among the most abundant and diverse symbiotic taxa in chemosynthetic-based ecosystems: more than 100 different vesicomyid species have been described so far. In the present study, we investigated the phylogenetic positioning of recently described vesicomyid species from the Gulf of Guinea and their western Atlantic and Pacific counterparts using mitochondrial DNA sequence data. The maximum-likelihood (ML) tree provided limited support for the recent taxonomic revision of vesicomyids based on morphological criteria; nevertheless, most of the newly sequenced specimens did not cluster with their morphological conspecifics. Moreover, the observed lack of geographic clustering suggests the occurrence of independent radiations followed by worldwide dispersal. Ancestral character state reconstruction showed a significant correlation between the characters “depth” and “habitat” and the reconstructed ML phylogeny suggesting possible recurrent events of ‘stepwise speciation’ from shallow to deep waters in different ocean basins. This is consistent with genus or species bathymetric segregation observed from recent taxonomic studies. Altogether, our results highlight the need for ongoing re-evaluation of the morphological characters used to identify vesicomyid bivalves

The smaller vesicomyid bivalves in the genus Isorropodon (Bivalvia, Vesicomyidae, Pliocardiinae) also harbour chemoautotrophic symbionts

Species of Isorropodon are vesicomyid bivalves for which little information is available regarding host phylogeny and bacterial symbioses. In this study we investigated the symbioses in three Isorropodon species from three cold seep areas: Isorropodon bigoti (Gulf of Guinea), Isorropodon megadesmus (Gulf of Cadiz) and Isorropodon perplexum (Eastern Mediterranean). Analysis of bacterial 16S ribosomal RNA gene sequences demonstrated that each vesicomyid species harbours a single symbiont phylotype, that symbionts from the three species cluster together, and that they are closely related to other known vesicomyid symbionts. These results are confirmed by other marker genes (encoding 23S rRNA and APS reductase) and by fluorescence in situ hybridization. Due to their extended depth range and transoceanic distribution Isorropodon species are interesting examples to further study evolutionary processes in bivalve hosts and their associated symbionts

Deep-Sea Biodiversity in the Mediterranean Sea: The Known, the Unknown, and the Unknowable

Deep-sea ecosystems represent the largest biome of the global biosphere, but knowledge of their biodiversity is still scant. The Mediterranean basin has been proposed as a hot spot of terrestrial and coastal marine biodiversity but has been supposed to be impoverished of deep-sea species richness. We summarized all available information on benthic biodiversity (Prokaryotes, Foraminifera, Meiofauna, Macrofauna, and Megafauna) in different deep-sea ecosystems of the Mediterranean Sea (200 to more than 4,000 m depth), including open slopes, deep basins, canyons, cold seeps, seamounts, deep-water corals and deep-hypersaline anoxic basins and analyzed overall longitudinal and bathymetric patterns. We show that in contrast to what was expected from the sharp decrease in organic carbon fluxes and reduced faunal abundance, the deep-sea biodiversity of both the eastern and the western basins of the Mediterranean Sea is similarly high. All of the biodiversity components, except Bacteria and Archaea, displayed a decreasing pattern with increasing water depth, but to a different extent for each component. Unlike patterns observed for faunal abundance, highest negative values of the slopes of the biodiversity patterns were observed for Meiofauna, followed by Macrofauna and Megafauna. Comparison of the biodiversity associated with open slopes, deep basins, canyons, and deep-water corals showed that the deep basins were the least diverse. Rarefaction curves allowed us to estimate the expected number of species for each benthic component in different bathymetric ranges. A large fraction of exclusive species was associated with each specific habitat or ecosystem. Thus, each deep-sea ecosystem contributes significantly to overall biodiversity. From theoretical extrapolations we estimate that the overall deep-sea Mediterranean biodiversity (excluding prokaryotes) reaches approximately 2805 species of which about 66% is still undiscovered. Among the biotic components investigated (Prokaryotes excluded), most of the unknown species are within the phylum Nematoda, followed by Foraminifera, but an important fraction of macrofaunal and megafaunal species also remains unknown. Data reported here provide new insights into the patterns of biodiversity in the deep-sea Mediterranean and new clues for future investigations aimed at identifying the factors controlling and threatening deep-sea biodiversity