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Until1985, seven vent fields were described from the Mid-Atlantic Ridge (MAR). An eighth field, Mount Saldanha (36° N), discovered in 1998, showed unusual geological and biologica! settings. Vent sites on the MAR exhibit varied environmental conditions, resulting from depth variation of the axis and associated physical parameters, and different source rocks. These could be considered as first order (i.e. most dominant) factors affecting the composition of vent communities on the MAR, in contrast to the East Pacific Rise (EPR) where geographical isolation appears to be a major determinant of fauna! differences. In this paper, the geological setting and vent fluid composition of the fields are considered together with their community composition to tentatively ascertain the order of a hierarchy between dispersa! and environmental control. The deepest fields (>3000 m) are rather stable systems. The shallower fields, especially Rainbow and Menez Gwen, present some evidence of instability in time and space. The variability in fluid composition is related to phase separation processes (boiling/distillation of subsurface vent fluids) and to the nature of the basement rocks. Depending on depth, phase separation produces gas-enriched and metal-depleted fluids (Menez Gwen, Lucky Strike) or metal-enriched brines (Rainbow, TAG). In addition, high methane content characterises the fluids formed in ultramafic rocks (Rainbow, Logatchev) compared to basaltic rocks. The discrepancy in mineral particulate fluxes at Lucky Strike and Menez Gwen, on one hand, and TAG and Rainbow, on the other, is correlated to the predominance of the vapour or brîne phase. The semi-quantitative description of the fauna! composition of the different vent fields displays a continuum from Rimicaris-dominated to Bathymodiolus-dominated assemblages. Rather than geographic or bathymetric zonation, this gradati an appears tobe related to the metal content of the fluids. In addition, the penetrati an of non vent species into the vent environment increases with decreasing hydrostatic pressure and/or metal content in the fluids. Similarity analysis between vent communities shows that similarity is strongest between Menez Gwen and Lucky Strike (the shallowest fields), less significant between these sites and Rainbow, and weakest for Snake Pit. The inverse relationship between filter feeding organisms and metal concentration in vent fluids could result from a hindrance of mussel bed development by particulate or toxic metal fluxes, and has to be further investigated. Conversely, high metal and particulate content would less affect the more mobile Rimicaris populations. Considering specific similarities of endemic fauna between the faur best known hydrothermal vents, the distance between vent fields appears tobe a first order parameter. Nevertheless, within the proximity of the Azores Triple Junction area, and in the absence of geographical discontinuity, the similarity between fields stays rather low suggesting fauna! islands that have distinct composition and h...

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Among hydrothermal vent species, Rimicaris exoculata is one of the most emblematic, hosting abundant and diverse ectosymbioses that provide most of its nutrition. Rimicaris exoculata co-occurs in dense aggregates with the much less abundant Rimicaris chacei in many Mid-Atlantic Ridge vent fields. This second shrimp also houses ectosymbiotic microorganisms but has a mixotrophic diet. Recent observations have suggested potential misidentifications between these species at their juvenile stages, which could have led to misinterpretations of their early-life ecology. Here, we confirm erroneous identification of the earliest stages and propose a new set of morphological characters unambiguously identifying juveniles of each species. On the basis of this reassessment, combined use of C, N and S stable isotope ratios reveals distinct ontogenic trophic niche shifts in both species, from photosynthesis-based nutrition before settlement, towards a chemosynthetic diet afterwards. Furthermore, isotopic compositions in the earliest juvenile stages suggest differences in larval histories. Each species thus exhibits specific early-life strategies that would, without our re-examination, have been interpreted as ontogenetic variations. Overall, our results provide a good illustration of the identification issues persisting in deep-sea ecosystems and the importance of integrative taxonomy in providing an accurate view of fundamental aspects of the biology and ecology of species inhabiting these environments.

The animal communities associated with the deep-sea reducing environment have been studied for almost 30 years, but until now only a single species of ophiuroid, Ophioctenella acies, has been found at both hydrothermal vents and methane cold seeps. Since the faunal overlap between vent and seep communities is small and many endemic species have been found among other taxa (e.g. Mollusca, Crustacea), additional species of ophiuroids were expected at previously unstudied sites. Chemical compositions at reducing sites di¡er greatly from the nearby bathyal environment. Generally, species adapted to chemosynthetic environments are not found in non-chemosynthetic habitats, but occasional visitors of other bathyal species to vent and seep sites have been recorded among many taxa except ophiuroids. This paper presents an analysis of the ophiuroid fauna found at hydrothermal vents and non-reducing nearby sites on the Mid-Atlantic Ridge and on methane cold seeps in the Gulf of Mexico, at Blake Ridge o¡ South Carolina and south of Barbados. In addition to O. acies, four species were found at vents, Ophiactis tyleri sp. nov., Ophiocten centobi, Ophiomitra spinea and Ophiotreta valenciennesi rufescens. While Ophioctenella acies appears to be restricted to chemosynthetic areas, the other four species were also found in other bathyal habitats. They also occur in low numbers (mostly single individuals), whereas species adapted to hydrothermal areas typically occur in large numbers. Ophioscolex tripapillatus sp. nov. and Ophiophyllum atlanticum sp. nov. are described from nearby non-chemosynthetic sites. In a cold seep south of Barbados, three species of ophiuroids were found, including Ophioctenella acies, Amphiura sp., Ophiacantha longispina sp. nov. and Ophioplinthaca chelys. From the cold seeps at Blake Ridge and the Gulf of Mexico, Ophienigma spinilimbatum gen. et sp. nov. is described, likely restricted to the reducing environment. Ophiotreta valenciennesi rufescens occurred abundantly among Lophelia corals in the Gulf of Mexico seeps, which is the ¢rst record of this species from the West Atlantic. Habitat descriptions complement the taxonomic considerations, and the distribution of the animals in reducing environments is discussed.

Fatty acid biomarker analysis coupled with gas chromatography-isotope ratio mass spectrometry was used to confirm the presence of methanotrophic and thiotrophic bacterial endosymbionts in the tissues of a hydrothermal vent mussel (Bathymodiolussp.), collected from the Menez Gwen vent field on the mid-Atlantic ridge. Monounsaturated (n-8) fatty acids, which are diagnostic of methanotrophic bacteria, were detected in all three types of tissues examined (gill, posterior adductor, and mantle), although levels were highest in gill tissues where the bacteria were found. Stable-carbon-isotope compositions (δ-13C per mille relative to that of Peedee belemnite) of fatty acids for all three tissues ranged from −24.9 to −34.9‰, which encompasses the range predicted for both thiotroph- and methanotroph-based nutrition. The data suggest that these thio- and methanotrophic bacterial endosymbionts are equally important in the nutrition of the vent mussel at this particular vent site.

Mjrocaris fortunata were sampled from the Lucky Strike hydrothermal vent area (Eiffel Tower site) on the mid-Atlantic ridge during the French DlVA 2 cruise (June 1994). Small adults (17 to 22 mm total length), although morphologically identical, could be divided into 2 categones on the basis of pigmentat~on, llpid composition and I3C/l2c stable isotope ratios of fatty acids H~ghly pigmented small adults (8 6 to 9.2 pg carotenoid shrimp-') contamed higher levels of total lipid than similar-sized individuals containing lower levels of pigment (0.9 to 2 9 pg carotenoid shrimp-'). Lipid class analysis indicated that wax esters comprised 62.5% of total lipid in the former group. These p~gmented shrimp also contained high proportions of polyunsaturated fatty acids (PUFA), particularly the phototrophic microplanktonic markers 20:5(n-3) and 22:6(n-3) (14.0 and 33.5% respectively). By contrast small adults (22 mm) and adult shrimp (25 to 26 mm) with low levels of carotenoid pigmentation contained lower amounts of total lipid, little or no wax ester and low levels of 20:5(n-3) and 22:6(n-3), but did contain 16:2(n-4) and 18:2(n-4) and the non-methylene interrupted dienes 20:285,13 and 22:287,15. GC-IRMS analysis of all fatty acids and fatty alcohols in the pigmented small adults indicated 6I3C values of -18.2 to -27.7?&,, which is consistent with a photosynthetic carbon source for these compounds. The I3C/l2C Isotope composition of fatty acids from low-pigmented small adults and adults was more variable (-12 5 to -33.11:L) and suggests a bimodal distribution which may be attributable to differing nutritional sources or the physiological/reproductive status of these shnmp Samples of eggs, which are carried by the female on the pleopods, represented approximately 57 % of total somatic lipid which indicates a substantial reproductive investment by this species. The egg lipids comprised high proportions of triacylglycerols (64.4 to 78.0% of total I~pid) whilst the fatty acid composition was dominated by the monounsaturated fatty acids 16:l(n-71, 18:l(n-7) and 18:l(n-9). which accounted for 65.7 to 73.5"0 of total fatty acids. By contrast, PUFA were relatively minor components of egg lipids, particularly 20:5(n-3) and 22:6(n-3), which accounted for only 1.1 and 2.9% of total egg fatty acids respectively. This indicates that the reproductive investment by this species is supported mainly by material derived from bacterial chemosynthesis. The potential for M. fortunata hedge betting by producing larvae which either metamorphose at the vent site or adopt a bathypelagic lifestyle and delay metamorphosis to facilitate more widespread dispersal is discussed.