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Chemosynthetic ecosystems are distributed worldwide in fragmented habitats harbouring seemingly highly specialized communities. Yet, shared taxa have been reported from highly distant chemosynthetic communities. These habitats are distributed in distinct biogeographical regions, one of these being the so-called Atlantic Equatorial Belt (AEB). Here, we combined genetic data (COI) from several taxa to assess the possible existence of cryptic or synonymous species and to detect the possible occurrence of contemporary gene flow among populations of chemosynthetic species located on both sides of the Atlantic. Several Evolutionary Significant Units (ESUs) of Alvinocarididae shrimp and Vesicomyidae bivalves were found to be shared across seeps of the AEB. Some were also common to hydrothermal vent communities of the Mid-Atlantic Ridge (MAR), encompassing taxa morphologically described as distinct species or even genera. The hypothesis of current or very recent large-scale gene flow among seeps and vents was supported by microsatellite analysis of the shrimp species Alvinocaris muricola/Alvinocaris markensis across the AEB and MAR. Two nonmutually exclusive hypotheses may explain these findings. The dispersion of larvae or adults following strong deep-sea currents, possibly combined with biochemical cues influencing the duration of larval development and timing of metamorphosis, may result in large-scale effective migration among distant spots scattered on the oceanic seafloor. Alternatively, these results may arise from the prevailing lack of knowledge on the ocean seabed, apart from emblematic ecosystems (chemosynthetic ecosystems, coral reefs or seamounts), where the widespread classification of endemism associated with many chemosynthetic taxa might hide wider distributions in overlooked parts of the deep sea., (© 2013 John Wiley & Sons Ltd.)

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.

The structure and familial-level composition of macrofaunal communities were examined within and at distance from the giant pockmark Regab in the Southern Gulf of Guinea, at 3200 m depth. The two main questions addressed in this study were (i) does habitat partitioning by large symbiont-bearing taxa influence macrofaunal assemblages and their environment? and (ii) to what extent does the chemosynthesis-based ecosystem influence the structure and the composition of nearby macrobenthic communities? Along two radials, at sites from 250 to 1000 m away from the active centre of the cold seep, the abundance and composition of the macrofauna were typical of a deep-sea community at that depth. Except for a few cores sampled in or near dead vesicomyid fields at the border of the pockmark area, the chemosynthetic ecosystem had no significant influence on macrofaunal communities beyond the edge of the giant pockmark. In seep sediments, the macrofauna had higher densities, reaching over 22,000 ind·m−2, but lower taxonomic richness than in background sediments. The polychaete assemblages associated with siboglinid tubeworms differed in their composition and vertical distribution from communities associated with vesicomyids or found at the border between vesicomyids and mytilids. The siboglinid habitat was dominated by surface-feeders (ampharetids) or small carnivores (dorvilleids, hesionids and syllids) concentrated in the top sediment layer, while the vesicomyid beds were dominated by deep-dwelling, subsurface feeders (capitellids and cossurids). This pattern may be related to bioturbation by the vesicomyids that allows an oxygenation of surface sediments while deepening down and enhancing the anaerobic oxidation of methane. Although alpha diversity of polychaete families is low in seep sediments, the variability of faunal assemblages and potentially high level of speciation in cold-seep environments point to high beta diversity. Comparable numbers of polychaete families coexist in a quarter square meter of seep or non-seep sediments, although the number of ecological niches that were sampled might be higher at seeps. Further integrative and comparative studies of the diversity and functioning of seep and non-seep communities would certainly provide clues to understanding both ecosystems and their interactions. [ABSTRACT FROM AUTHOR]

The high biodiversity found at cold seeps, despite elevated concentrations of methane and hydrogen sulfide, is attributed to multiple sources of habitat heterogeneity. In addition to geological and geochemical processes, biogenic habitats formed by large symbiont-bearing taxa, such as bivalves and siboglinid tubeworms, or by microbial mats drive the biodiversity of small-sized fauna. However, because these habitat-forming species also depend on geochemical gradients, the respective influence of abiotic and biotic factors in structuring associated macrofaunal communities is often unresolved. The giant pockmark Regab located at 3200 m depth on the Congo margin is characterized by different fluid-flow regimes, providing a mosaic of the most common biogenic habitats encountered at seeps: microbial mats, mussel beds, and vesicomyid clam beds; the latter being distributed along a gradient of environmental conditions from the center to the periphery of the pockmark. Here, we examined the structure of macrofaunal communities in biogenic habitats formed in soft sediment to (1) determine the influence of the habitats on the associated macrofaunal communities (inter-habitat comparison), (2) describe how macrofaunal communities vary among vesicomyid clam beds (intra-habitat comparison) and (3) assess the inter-annual variation in vesicomyid beds based on repeated sampling at a three-year interval. The highest densities were found in the microbial mat communities in intermediate fluid-flow areas, but they had low diversity — also observed in the sediment close to mussel beds. In contrast, vesicomyid beds harbored the highest diversity. The vesicomyid beds did not show a homogeneous macrofaunal community across sampled areas; instead, density and composition of macrofauna varied according to the location of the beds inside the pockmark. The clam bed sampled in the most active, central part of the pockmark resembled bacterial mat communities by the presence of highly sulfide-tolerant species living at the sediment surface, along with vesicomyid juveniles. This similarity suggests a gradual change in community composition from mats to clam beds. Inter-annual comparisons of the different clam beds highlighted that the most active central site had a more variable community than its peripheral counterparts. Finally, a rapid shift in community structure, particularly in polychaete families, in experimentally reduced oxygen concentrations in the central part of Regab, suggests that high beta-diversity communities can withstand intense variation in geochemical conditions. These community dynamics are likely related to the diversity and to the plasticity of the vesicomyids themselves, because they can cope with high spatial and temporal environmental variability at a very local scale. [ABSTRACT FROM AUTHOR]

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 macro- and 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, macro- and 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. • Environmental conditions of pockmarks affect benthic community composition. • Meiofaunal taxa are able to take advantage of pockmark activity, i.g Desmodora. • Desmodora nematode could be a benthic indicator of fluid emission. • Benthic community compositions are largely dissimilar between the two sites. [ABSTRACT FROM AUTHOR]