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The synthesis of organic molecules in hydrothermal fluids raises the question of what role hydrothermal vents may have played in the emergence of life on Earth. These deep‐sea ecosystems were most likely protected from extreme conditions at the surface (meteor showers and ultraviolet radiation). Since the middle of the 19th century, scientists have questioned the position of deep‐sea animals in the evolution of marine fauna, in the hope to discover the missing link, i.e. the species or group that may have survived mass extinctions in this protected and stable environment. This chapter discusses the geology and geochemistry of hydrothermal systems, microbial chemosynthesis, symbioses and trophic chains, and fauna at different spatial scales. Fauna assemblages colonize narrow mixing gradients between hydrothermal fluids and seawater, on hard substrataor in thick sediment layers. Most temporal studies are not based on continuous temporal monitoring but on regular visits by manned or unmanned submersibles.

Aim Deep-sea hydrothermal vents have now been reported along all active mid-ocean ridges and back-arc basins, but the boundaries of biogeographic entities remain questionable owing to methodological issues. Here we examine biogeographic patterns of the vent fauna along the East Pacific Rise (EPR) and determine the relative roles of regional and local factors on the distribution of biodiversity associated with mussel beds along a poorly explored zone, the southern EPR (SEPR). Location East Pacific Rise. Methods A species list of macrobenthic invertebrates along the EPR was compiled from the literature and supplemented with data recovered during the French research cruise BIOSPEEDO carried out in 2004 along the SEPR. Biogeographic patterns were assessed by combining the identification of morphological species with a molecular barcoding approach. A multivariate regression tree (MRT) analysis was performed to identify any geographic breaks, and an empirical distribution of species richness was compared with predictions provided by a mid-domain effect model. Macrofaunal community structure associated with mussel beds along the SEPR was analysed in relation to environmental factors using cluster and canonical redundancy analyses. Results Sequencing of the cytochrome c oxidase subunit I gene revealed the occurrence of several cryptic species complexes along the EPR, with the equator separating the southern and northern clades. Furthermore, during the BIOSPEEDO cruise at least 10 still unnamed species were collected between 7 degrees 25' S and 21 degrees 33' S. The shift in community structure identified by MRT analysis was located south of 17 degrees 34' S or south of 13 degrees 59' S, depending on the data used, suggesting that the southern part of the SEPR (17 degrees 25'-21 degrees 33' S) constitutes a biogeographic transition zone in the vent fauna along the EPR. At a regional scale, latitude combined with the type of venting was significantly correlated with the commu

The Eiffel Tower edifice is situated in the Lucky Strike hydrothermal vent field at a mean depth of 1690 m on the Mid-Atlantic Ridge (MAR). At this 11-m-high hydrothermal structure, different faunal assemblages, varying in visibly dominant species (mussels and shrimp), in mussel size and in density of mussel coverage, were sampled biologically and chemically. Temperature and sulphide (Sigma S) were measured on the different types of mussel-based assemblages and on a shrimp-dominated assemblage. Temperature was used as a proxy for calculating total concentrations of CH4. Based on the physico-chemical measurements, two microhabitats were identified, corresponding to (i) a more variable habitat featuring the greatest fluctuations in environmental variables and (ii) a second, more stable, habitat. The highest temperature variability and the highest maximum recorded temperatures were found in the assemblages visibly inhabited by alvinocaridid shrimp and dense mussel beds of large Bathymodiolus azoricus, whereas the less variable habitats were inhabited by smaller-sized mussels with increasing bare surface in between. Larger mussels appeared to consume more Sigma S compared with smaller-sized (< 1 cm) individuals and thus had a greater influence on the local chemistry. In addition, the mussel size was shown to be significantly positively correlated to temperature and negatively to the richness of the associated macrofauna. The presence of microbial mats was not linked to specific environmental conditions, but had a negative effect on the presence and abundance of macro-fauna, notably gastropods. Whereas some taxa or species are found in only one of the two microhabitats, others, such as polychaetes and Mirocaris shrimp, cross the different microhabitats. Temperature was proposed to be a more limiting factor in species distribution than Sigma S.

Ashadze-1 (128 580N 448 510W, 4080 m) on the Mid-Atlantic Ridge (MAR) is the deepest known ctive hydrothermal vent ield. The first observations on this site were numerous clear and black smokers and surprisingly few nown symbiotic species ominant in other vent areas on the MAR. The species most abundant at Ashadze-1 are those usually ound at the periphery f hydrothermal communities: sea-anemones Maractis rimicarivora and chaetopterid polychaetes hyllochaetopterus sp. ov. This study comprised site mapping and faunal sampling and Ashadze-1 was completely mapped y using the Remote perated Vehicle ‘Victor 6000’ and a new high resolution tool available for deep-sea research. A photo-mapping survey was carried out with a long range optical black and white camera. Digitization of substrata and sea-anemones visible on the images was performed by GIS. Spatial distribution of Ma. rimicarivora was distinguished by high densities of 32 ind.m22 on the western side of the main smoker area. Submersible sampling operations allowed taxonomic identification within a 200 × 110 m area. Carbon, nitrogen and sulphur isotopic ratios were measured in four dominant species to identify their trophic position. The present paper gives the complete maps and describes the faunal community of the Ashadze-1 vent field. The results obtained led us to consider this site as an ecosystem in its declining stage. Finally we compare the similarities of this community to other hydrothermal communities on the northern MAR.

Like hydrothermal vents along oceanic ridges, cold seeps are patchy and isolated ecosystems along continental margins, extending from bathyal to abyssal depths. The Atlantic Equatorial Belt (AEB), from the Gulf of Mexico to the Gulf of Guinea, was one focus of the Census of Marine Life ChEss (Chemosynthetic Ecosystems) program to study biogeography of seep and vent fauna. We present a review and analysis of collections from five seep regions along the AEB: the Gulf of Mexico where extensive faunal sampling has been conducted from 400 to 3300m, the Barbados accretionary prism, the Blake ridge diapir, and in the Eastern Atlantic from the Congo and Gabon margins and the recently explored Nigeria margin. Of the 72 taxa identified at the species level, a total of 9 species or species complexes are identified as amphi-Atlantic. Similarity analyses based on both Bray Curtis and Hellinger distances among 9 faunal collections, and principal component analysis based on presence/absence of megafauna species at these sites, suggest that within the AEB seep megafauna community structure is influenced primarily by depth rather than by geographic distance. Depth segregation is observed between 1000 and 2000m, with the middle slope sites either grouped with those deeper than 2000m or with the shallower sites. The highest level of community similarity was found between the seeps of the Florida escarpment and Congo margin. In the western Atlantic, the highest degree of similarity is observed between the shallowest sites of the Barbados prism and of the Louisiana slope. The high number of amphi-atlantic cold-seep species that do not cluster according to biogeographic regions, and the importance of depth in structuring AEB cold-seep communities are the major conclusions of this study. The hydrothermal vent sites along the Mid Atlantic Ridge (MAR) did not appear as "stepping stones" for dispersal of the AEB seep fauna, however, the south MAR and off axis regions should be further explor

A brackish-water cold seep on the North Anatolian Fault (NAF) in the Marmara Sea was investigated with the Nautile submersible during the MarNaut cruise in 2007. This active zone has already been surveyed and revealed evidence of active seeping on the seafloor, such as bubble emissions, patches of reduced sediments, microbial mats and authigenic carbonate crusts. MarNaut was the first opportunity to sample benthic communities in the three most common microhabitats (bioturbated and reduced sediments, carbonate crust) and to examine their relationships with environmental conditions. To do so, faunal communities were sampled and chemical measurements were taken close to the organisms. According to diversity indices, the bioturbated microhabitat exhibited the highest taxonomic diversity and evenness despite a lower number of samples. Conversely, the reduced sediment microhabitat exhibited the lowest taxonomic diversity and evenness. The carbonate crust microhabitat was intermediate although it had the highest biomass. Multivariate analyses showed that (1) fauna were relatively similar within a single microhabitat; (2) faunal community structure varied greatly between the different microhabitats; (3) there was a link between faunal distribution and the type of substratum; and (4) chemical gradients (i.e. methane, oxygen and probably sulphides) may influence faunal distribution. The estimated fluid flow velocity (0.4-0.8 m/yr) confirmed the presence of fluid emission and provided evidence of seawater convection in the two soft-sediment microhabitats. Our results suggest that the reduced sediments may represent a harsher environment with high upward fluid flow, which restrains seawater from penetrating the sediments and inhibits sulphide production, whereas bioturbated sediments can be viewed as a bio-irrigated system with sulphide production occurring at greater depths. Therefore, the environmental conditions in reduced sediments appear to prevent the colonization of symb

Author Posting. © American Geophysical Union, 2004. This article is posted here by permission of American Geophysical Union for personal use, not for redistribution. The definitive version was published in Journal of Geophysical Research 109 (2004): B12102, doi:10.1029/2004JB003141., Mid-ocean ridge volcanic activity is the fundamental process for creation of ocean crust, yet the dynamics of magma emplacement along the slow spreading Mid-Atlantic Ridge (MAR) are largely unknown. We present acoustical, seismological, and biological evidence of a magmatic dike intrusion at the Lucky Strike segment, the first detected from the deeper sections (>1500 m) of the MAR. The dike caused the largest teleseismic earthquake swarm recorded at Lucky Strike in >20 years of seismic monitoring, and one of the largest ever recorded on the northern MAR. Hydrophone records indicate that the rate of earthquake activity decays in a nontectonic manner and that the onset of the swarm was accompanied by 30 min of broadband (>3 Hz) intrusion tremor, suggesting a volcanic origin. Two submersible investigations of high-temperature vents located at the summit of Lucky Strike Seamount 3 months and 1 year after the swarm showed a significant increase in microbial activity and diffuse venting. This magmatic episode may represent one form of volcanism along the MAR, where highly focused pockets of magma are intruded sporadically into the shallow ocean crust beneath long-lived, discrete volcanic structures recharging preexisting seafloor hydrothermal vents and ecosystems., This study was made possible through the support of the U.S. National Science Foundation (grants OCE-9811575, OCE- 0137164, and OCE-0201692) and the NOAA Vents Program.

The discovery of deep-sea hydrothermal vent fauna, kilometres deep in the oceans, is a great achievement of 20th-century marine biology. The deep-sea hydrothermal food web does not directly depend on the sun's energy. Vent communities rely primarily on trophic associations between chemoautotrophic bacteria and consumers A small. number of endemic taxa are adapted to the inhospitable vent environments that are distributed along ridge crests. Where these vent communities originated and how they dispersed are among the important questions ecologists must answer. Here, by statistical analysis of the most comprehensive database ever assembled about deep-sea hydrothermal fauna, we delineate six major hydrothermal provinces in the world ocean and identify seven possible dispersal pathways between adjacent provinces. Our model suggests that the East-Pacific Rise may have played a pivotal role as a centre of dispersal for the hydrothermal fauna. Our data-driven conclusion will have to be tested by phylogenetic studies and completed by surveys of less-explored fields.

Whilst the fauna inhabiting hydrothermal vent structures in the Atlantic Ocean is reasonably well known, less is understood about the spatial distributions of the fauna in relation to abiotic and biotic factors. In this study, a major active hydrothermal edifice (Eiffel Tower, at 1690 m depth) on the Lucky Strike vent field (Mid-Atlantic Ridge (MAR)) was investigated. Video transects were carried out by ROV Victor 6000 and complete image coverage was acquired. Four distinct assemblages, ranging from dense larger-sized Bathymodiolus mussel beds to smaller-sized mussel clumps and alvinocaridid shrimps, and two types of substrata were defined based on high definition photographs and video imagery. To evaluate spatial variation, faunal distribution was mapped in three dimensions. A high degree of patchiness characterizes this 11 m high sulfide structure. The differences observed in assemblage and substratum distribution were related to habitat characteristics (fluid exits, depth and structure orientation). Gradients in community structure were observed, which coincided with an increasing distance from the fluid exits. A biological zonation model for the Eiffel Tower edifice was created in which faunal composition and distribution can be visually explained by the presence/absence of fluid exits.

Trophic relationships in Bathymodiolus azoricus mussel bed communities on the Tour Eiffel hydrothermal edifice (Lucky Strike) were assessed using delta C-13 and delta N-15 signatures from 14 hydrothermal species. The nutritional basis of B. azoricus was also investigated with delta S-34. Faunal samples and environmental data (temperature, pH, total dissolved sulfide, iron and copper concentrations) were collected from 12 different locations on the edifice. Chemical conditions varied between microhabitats, and were all correlated to temperature. Carbon and nitrogen isotopic results revealed the presence of two, apparently independent, trophic groups. The first was composed of symbiont-bearing fauna (B. azoricus and their associated polychaetes Branchipolynoe seepensis), while the second enclosed heterotrophic fauna (bacterivores, cletritivores, scavengers, predators). A majority of mussels displayed delta C-13 values ranging from -27 parts per thousand to -34 parts per thousand, supporting thiotrophy as the dominant nutritional pathway at Tour Eiffel, with methanotrophy and filter feeding emerging as secondary strategies. This result was corroborated by delta S-34 signatures. However, higher delta C-13 values in larger mussels suggested that, as they grow, B. azoricus mussels rely more heavily on their methanotrophic enclosymbionts. Significant spatial variability in isotopic signatures for single faunal species was observed at the scale of the edifice for three species (B. azoricus, B. seepensis, Amathys lutzi), and environmental conditions explained variation in isotopic signatures for one-third of the species. This confirms the hypothesis raised by several authors on the role of hydrothermal fluids on the trophic network at small spatial scales. We suggest that vent fluid characteristics, by influencing microbial production, are key factors in the variation of local carbon sources at vents.

In the framework of the deep-sea environmental programme BIOZAIRE (Ifremer-Total), colonization trays were deployed for 283-433 days at three sites along the equatorial West African margin: ZA at 1300-m depth, ZC at 4000-m depth far from the Congo canyon and ZD at 4000-m depth close to the Congo canyon. The experiments aimed at determining the influence of depth and local environmental settings on macrofaunal colonization patterns and organic carbon degradation rates. The trays were filled with glass beads and this artificial substrate was enriched with ground particulate organic matter in a gradient of 0%, 0.34%, 1.02% and 3.43% organic carbon. The highest rates of organic carbon degradation ranged, according to the duration of the experiments, from 1.59 to 2.36 gC m(-2) day(-1) but were independent of depth or location. Colonization rates, conversely, varied by one order of magnitude between bathyal and abyssal experiments. The influence of experimental treatments on the structure of the colonizing macrofauna also varied according to location and depth. At ZA, colonization patterns were highly predictable and driven by a shift in dominance of opportunistic taxa along the enrichment gradient. To a lesser extent, this was also true at ZD, near the Congo canyon, while at ZC the treatments had no significant effect on the composition of the colonizing fauna. At abyssal depth, high rates of organic matter degradation associated with low rates of colonization suggested that pulse of organic matter would mainly benefit the resident community. At bathyal depth, high colonization rates of a specialized fauna might conversely play an important role in the functioning of the ecosystem. The regional and local coexistence of an opportunistic fauna via a spatial storage effect associated with dispersal might significantly contribute to the maintenance of high diversity on continental margins.

Bathymodiolus azoricus is a mussel from vent fields in the south-west of the Azores Triple junction (Mid-Atlantic Ridge-MAR). Experimental evidence indicates that B. azoricus is a mixotrophic organism, which obtains energy from a dual endosymbiosis and filter-feeding. Yet the relative contribution of symbiosis and filter-feeding to B. azoricus nutrition is still unclear. To address this question, we developed and individual-based model which describes sulphide and methane uptake by endosymbionts, the energy gained through microbial oxidations, the transfer of energy from endosymbionts to B. azoricus, filter-feeding of particulate organic matter (POC) by B. azoricus and the energetic wastes of the mytilid with respiration. The model accounts for size-dependent relationships obtained from empirical data. External concentrations of H2S and CH4 correspond to estimated values for the Menez Gwen vent field, maximal and minimal values measured at MAR. From in situ observed densities of B. azoricus, productivity predictions at the individual level were upscale to the mytilid population at Menez Gwen and compared to estimated values. Predicted biomass of B. azoricus and its endosymbionts show a very high fitting level with estimated values. Results suggest that the relative contribution of filter-feeding and endosymbiosis varies with B. azoricus size, with small mytilids being strongly dependent on filter-feeding, whilst larger mussels obtain a significant portion of its energy from endosymbiosis. This is related with the variation of gill weight with total weight. Results also suggest that, an individual of a certain size can potentially regulate the relative contribution of filter-feeding and endosymbiosis according to external conditions. However, large B. azoricus exhibit a higher level of nutritional flexibility than small mytilids. The relative contribution of endosymbioisis and filter-feeding to the total energy budget of B. azoricus, as well as the mytilid particulat

The polar lipid fatty acid (PLFA) profiles of invertebrates living in chemosynthetic communities can indicate the degree to which these animals depend on specific types of bacteria. To identify the nutritional sources of various species from deep-sea hydrothermal vents of the Mid-Atlantic Ridge, a Principal Component Analysis was performed using individual PLFA profiles as descriptors. Two associations representing different feeding groups were identified: (i) mussels, commensal polychaetes and gastropods, (ii) shrimps and crabs. The first association relies more on sulphide-oxidizing bacteria, while the second one has more anaerobic sulphate-reducing bacteria biomarkers. Other small invertebrates reveal different diets. The polychaete Amathys lutzi shows the most diversified bacterial diet, with fatty acid biomarkers from both S-oxidizing and S-reducing bacteria.

The composition, biomass and diversity of the fauna in a Bathymodiolus thermophilus mussel assemblage colonizing diffuse flow areas on the SEPR is described and compared with biological characteristics of other hydrothermal assemblages. The spatio-temporal dynamics of temperatures over fine scales within the mussel habitat are characterized using newly-developed statistical approaches. Temperature data obtained from two adjacent habitats (a cirriped assemblage and a bare substratum) are compared. The results of the temperature data show that the mean temperature and the temperature variations were significantly higher in the mussel assemblage (mean temperature: 4.12 +/- 1.85 degrees C) in comparison with 2.40 +/- 0.14 degrees C in the cirriped assemblage and 2.20 +/- 0.23 degrees C on the bare substratum. Discrete temperature measurements showed that temperature data varied both at a broad and fine scales in the mussel assemblage. Finally, analysis of the temperature periodic variability in the mussel assemblage during a 4-day period showed significant periodic modulations near T = 24h. Although not significant, a second trend is also observed around T = 12h. This work represents a step forward to the understanding of species distribution patterns at vents.

Malacoceros samurai, new species, (Spionidae) is described from deep-sea hydrothermal vents of the southern East Pacific Rise, at 17 degrees 25'S. It is a large species, living on chimney walls along with Avinella spp. Although this species is described in the genus Malacoceros, it possesses unidentate hooded hooks and anterior scalpel-like aristate neurosetae, both characters which clearly differentiate it from other species of the same genus. These unusual characters require the modification of the generic diagnosis of Malacoceros.

Mid-ocean ridge volcanic activity is the fundamental process for creation of ocean crust, yet the dynamics of magma emplacement along the slow spreading Mid-Atlantic Ridge (MAR) are largely unknown. We present acoustical, seismological, and biological evidence of a magmatic dike intrusion at the Lucky Strike segment, the first detected from the deeper sections (>1500 m) of the MAR. The dike caused the largest teleseismic earthquake swarm recorded at Lucky Strike in >20 years of seismic monitoring, and one of the largest ever recorded on the northern MAR. Hydrophone records indicate that the rate of earthquake activity decays in a nontectonic manner and that the onset of the swarm was accompanied by 30 min of broadband (>3 Hz) intrusion tremor, suggesting a volcanic origin. Two submersible investigations of high-temperature vents located at the summit of Lucky Strike Seamount 3 months and 1 year after the swarm showed a significant increase in microbial activity and diffuse venting. This magmatic episode may represent one form of volcanism along the MAR, where highly focused pockets of magma are intruded sporadically into the shallow ocean crust beneath long-lived, discrete volcanic structures recharging preexisting seafloor hydrothermal vents and ecosystems.

1. Les préoccupations communautaires prioritaires, et les enjeux. " La biodiversité peut être vue sous deux angles complémentaires. Le premier, familier au public, est celui de la recherche spécifique des écosystèmes; le public est sensibilisé à la disparition d'espèces comme aux invasions biologiques. Mais la biodiversité, c'est aussi la multiplicité des interactions dynamiques entre des gènes et protéines dans les organismes, des espèces dans un milieu, des bactéries aux grands mammifères et du plus petit écosystème jusqu'à la biosphère dans son ensemble. Si le développement durable est une façon d'aborder conjointement la croissance, l'environnement et la question sociale, alors le concept de biodiversité introduit une façon nouvelle de considérer les interactions entre les espèces y compris l'espèce humaine, à toutes les échelles." Depuis la Conférence des Nations Unies sur l'Environnement et le Développement (Rio, 1992), la diversité du vivant est devenue une préoccupation planétaire : 188 Etats Parties y ont signé la Convention sur la Diversité Biologique qui, depuis, coordonne des travaux scientifiques et techniques réguliers. L'Union Européenne est clairement et fortement engagée dans ce processus et a rappelé à plusieurs reprises cet engagement : • Déclaration du Conseil de Gothenburg (juin 2001) •Conclusions du sommet sur le développement durable de Johannesburg (août 2002) • Conférence de Kuala Lumpur (mai 2004) (" Sustaining Livelihoods and Biodiversity : attaining the 2010 Target in the European Bodiversity Strategy ”). • Conclusion du Conseil de l'Europe (Bruxelles, 28 juin 2004) : " mettre un terme au processus de diminution de la biodiversité à l'horizon 2010 ". 2. Les domaines et thèmes de recherche. Dans ce domaine, la communauté scientifique française a conduit une réflexion prospective , permettant d'aboutir à un important rapport " Stratégie nationale de recherche pour la biodiversité au service du développement durable " (Paris septembre 2003).

In this study we assessed the phylogenetic relationships of the hydrothermal vent polychaete group Alvinellidae, based on parsimony analyses of combined morphological and molecular data. Morphological data were obtained from newly examined specimens and literature information of 16 terminal taxa belonging to Alvinellidae, Ampharetidae, Pectinariidae, Terebellidae, Trichobranchidae, and the outgroups Oweniidae and Sabellidae. Molecular data were based on 28S rRNA from 13 of the 16 morphological terminals (10 previously published sequences plus three new ones). The combined analysis indicated the clades ((Alvinellidae, Trichobranchidae) Pectinariidae) and (Ampharetidae, Terebellidae). Alvinellidae, Ampharetidae and Terebellidae, as currently delineated, are monophyletic. The positions of Trichobranchidae and Pectinariidae contradicted traditional views, and they also had low Bremer support and merit further studies. Well-supported clades included Alvinellidae and Terebellinae. Previous statements that Alvinellidae are either nested within Ampharetidae or the sister to this taxon were not supported. The traditional but here contradicted view that Terebellidae and Trichobranchidae are closely related may be based on plesiomorphic similarities between these two taxa.