Håkon Mosby Mud Volcano (HMMV, SW Barents Sea slope, 1280 m) is one of the numerous cold methane-venting seeps existing along the continental margins. Analyses of videoguided core samples revealed extreme differences in the diversity and density of the metazoan meiobenthic communities associated with the different sub-habitats (centre, microbial mats, Pogonophora field, outer rim) of this mud volcano. Diversity was lowest in the sulphidic, microbial mat sediments that supported the highest standing stock, with unusually high densities (11 000 ind. 10 cm -2 ) of 1 nematode species related to Geomonhystera disjuncta. Stable carbon isotope analyses revealed that this nematode species was thriving on chemosynthetically derived food sources in these sediments. Ovoviviparous reproduction has been identified as an important adaptation of parents securing the survival and development of their brood in this toxic environment. The proliferation of this single species in exclusive association with free-living, sulphide-oxidising bacteria (Beggiatoa) indicates that its dominance is strongly related to trophic specialisation, evidently uncommon among the meiofauna. This chemoautotrophic association was replaced by copepods in the bare, sulphidefree sediments of the volcano's centre, dominated by aerobic methane oxidation as the chemosynthetic process. Copepods and nauplii reached maximum densities and dominance in the volcano's centre (500 ind. 10 cm -2 ). Their strongly depleted carbon isotope signatures indicated a trophic link with methane-derived carbon. This proliferation of only selected meiobenthic species supported by chemosynthetically derived carbon suggests that, in addition to the sediment geochemistry, the associated reduced meiobenthic diversity may equally be related to the trophic resource specificity in HMMV sub-habitats.

The great variety of geological and hydrological conditions in the deep sea generates many different habitats. Some are only recently explored, although their true extent and geographical coverage are still not fully established. Both continental margins and mid-oceanic seafloors are much more complex ecologically, geologically, chemically and hydrodynamically than originally thought. As a result, fundamental patterns of species distribution first observed and explained in the context of relatively monotonous slopes and abyssal plains must now be re-evaluated in the light of this newly recognized habitat heterogeneity. Based on a global database of nematode genus composition, collected as part of the Census of Marine Life, we show that macrohabitat heterogeneity contributes significantly to total deep-sea nematode diversity on a global scale. Different deep-sea settings harbour specific nematode assemblages. Some of them, like coral rubble zones or nodule areas, are very diverse habitats. Factors such as increased substrate complexity in the case of nodules and corals seem to facilitate the co-existence of a large number of genera with different modes of life, ranging from sediment dwelling to epifaunal. Furthermore, strong biochemical gradients in the case of vents or seeps are responsible for the success of particular genera, which are not prominent in more typical soft sediments. Many

Cold seeps are among the most heterogeneous of all continental margin habitats. Abiotic Sources of heterogeneity in these systems include local variability in fluid flow, geochemistry, and substrate type, which give rise to different sets of microbial communities, microbial symbiont-bearing foundation species, and associated heterotrophic species. Biogenic habitats created by microbial mats and the symbiotic species including vesicomyid clams, bathymodiolin mussels, and siboglinid tubeworms add an additional layer of complexity to seep habitats. These forms of habitat heterogeneity result in a variety of macrofaunal and meiofaunal communities that respond to changes in structural complexity, habitat geochemistry, nutrient sources, and interspecific interactions in different ways and at different scales. These responses are predicted by a set of theoretical metacommunity models, the most appropriate of which for seep systems appears to be the 'species sorting' concept, an extension of niche theory. This concept is demonstrated through predictable patterns of community assembly, succession, and beta-level diversity. These processes are described using a newly developed analytical technique examining the change in the slope of the species accumulation curve with the number of habitats examined. The diversity response to heterogeneity has a consistent form, but quantitatively changes at different seep sites around the world as the types of habitats present and the size-classes of fauna analyzed change. The increase in beta diversity across seep habitat types demonstrates that cold seeps and associated biogenic habitats are significant sources of heterogeneity on continental margins globally

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To estimate the degree of spatial heterogeneity of benthic deep-sea communities, we carried out a multiple-scale (from m's to 200 km) investigation in the Congo-Angola margins (Equatorial West African margin, 3150-4800 m) in which we examined the metazoan meiofauna at a variety of habitats along the Congo Channel system and in the associated cold seep. We investigate the structure, density, vertical distribution patterns in the sediment and biomass of meiofaunal communities in the Gulf of Guinea and how they are controlled by hydrologic and biogeochemical processes. The meiofaunal communities in the Gulf of Guinea were shaped by heterogeneous conditions on the margin, and reflect the multiple-scale spatial variability that corresponds with different identified habitats. The two control sites, located at >100 km away from the canyon, were inhabited by very dense and the most diverse meiobenthic communities. Similar meiobenthic communities inhabited the transition zone between the canyon and the cold seep. Sites located along the Congo Channel were obviously affected by the local high-velocity bottom currents and unstable sedimentary conditions in this active submarine system. Extremely low meiobenthic densities and very low proportions in the most surficial sediment layers provided evidence for recently highly disturbed sediments at these sites. The ROV Victor 6000 provided images of the cold seep, showing a patchy distribution of several types of patchy distributed megafaunal communities dominated by three key symbiotic taxa (Mytilidae, Vesicomyidae and Siboglinidae). These cold seep sediments were colonised by a unique meiobenthic community, characterised by a high small-scale (m's) patchiness, low species richness and the prominent dominance of two large-sized nematode species: Sabatieria mortenseni, which is a cosmopolitan nematode known from littoral habitats, and an undescribed Desmodora species. The high individual body weight of Sabatieria mortenseni and its dominance at the cold seep site resulted in a significantly higher nematode biomass at the seep compared to the surrounding sites. In addition, the vertical A c c e p t e d m a n u s c r i p t 2 nematode profiles, with maximum proportions in subsurficial layers, points to a chemosynthesis-based meiobenthic community in this cold seep, in contrast to the phytodetritus-based communities at the control sites and at the transition zone.

Cold-seep environments and their associated symbiont-bearing megafaunal communities create islands of primary production for macro-and meiofauna in the otherwise monotonous and nutrientpoor deep-sea environment. To examine the spatial variation and distribution patterns of metazoan meiobenthos in different seepage-related habitats, samples were collected in two regions off Norway: several pockmarks associated with the Storegga Slide including the Nyegga pockmark area (730 m; 64°N), and the active, methane-venting Håkon Mosby Mud Volcano (HMMV) west of the Barents Sea (1280 m; 72°N). Based on sediment geochemistry and associated epifauna, three different habitat types were distinguished across the two regions: (1) reduced sediment with suboxic conditions, sometimes covered by bacterial mats, (2) sediment colonised by chemosynthetic, siboglinid tubeworms, and (3) sediment outside the influence of seepage and without a large chemosynthetic fauna. Meiofaunal communities varied strongly in terms of generic diversity and dominance among the different habitat types. Control sites and Siboglinidae polychaete fields both supported high nematode genus richness similar to normal deep-sea sediments, whereas the reduced sediments yielded a genus-poor nematode community dominated by one or two successful species. Meiofaunal densities in the different habitats were negatively correlated with macrobenthic densities. An extremely dense (>11,000 ind. 10 cm-2), mono-specific nematode population appeared to be restricted to the bacterial mats at HMMV. It consisted of a new cryptic species of the Halomonhystera disjuncta complex, which has been described from intertidal habitats in the North Sea. The reduced seep sediments at Nyegga did not yield H. disjuncta but were dominated by Terschellingia longicaudata, another cosmopolitan nematode species known to be abundant in organic-rich, oxygen-poor, shallow-water environments. These observations point to a past or recent connection between margins and shallow-water habitats.

International audienceHåkon Mosby mud volcano (HMMV) is one of the most active and most studied seep sites in European waters. Many authors have described its thermal activity, dynamic of mud flows, and geochemical and microbial processes. It is characterised by a concentric zonation of successive biogenic habitats related to an activity and geochemical gradient from its centre to its periphery. Around the central area covered by mud flows, white and grey microbial mats occur among areas of bare sediment, whereas siboglinid tubeworm fields of Sclerolinum contortum and/or Oligobrachia haakonmosbiensis colonise the peripheral areas. The meiofaunal community is known to be structured among habitats, but the macrofauna has rarely been investigated and has never been sampled in situ. As part of the European project HERMES, using the ROVs Victor 6000 and Quest 4000, we sampled quantitatively the different habitats of the volcano for macrofauna sensus lato, retained on a 250- or 500-μm sieve. We also sampled a newly discovered pockmark on Storegga slide (cne 5.6) and two pockmarks (G11, G12) in the Nyegga area. Macrofauna was identified and counted from phylum to family level. Our results on HMMV showed a gradient of increasing density and diversity from the volcano centre (1-3 taxa; 260 ind*m−2) to the peripheral siboglinid fields (8-14 taxa, 93,000 ind*m−2), with an intermediate situation for microbial mats. For macrofauna ≥500 μm, non-siboglinid polychaetes dominated the communities of the central mud volcano area, white mats and S. contortum fields (83, 89 and 37% of the total, respectively), whereas gastropods dominated grey mats and O. haakonmosbiensis fields (89 and 44% of the total, respectively). Polychaete families followed the same pattern of diversity according to habitats within HMMV. Of 23 polychaete families identified, only one occurred in the centre, and three in the microbial mats. Capitellidae and Dorvilleidae (typical of organically and sulphide-enriched areas) occurred at remarkably high densities in white microbial mats and in O. haakonmosbiensis fields. The S. contortum fields were the most diverse habitat with 12 polychaete families. The 250-μm fraction showed similar taxa dominating the habitats, but taking meiofauna into account, nematodes became the major taxon in white mats and in S. contortum fields, where they were particularly large in size, whereas copepods dominated in other habitats. Meiofauna and macrofauna did not show the same patterns of density according to habitats. Using principal components analysis the habitats at HMMV were clearly distinct, and clustered according to dominant species of siboglinids and type of microbial mats. Pockmarks at Nyegga showed a similar concentric pattern of habitats around fluid sources as on the volcano, which seemed similarly to influence macrofauna composition, but at a much smaller scale. Total taxa and polychaete diversity are high in the S. contortum fields in these pockmarks as well. Regional-scale comparisons including HMMV and Storegga sugge...

We report the results of a detailed investigation on the trophoecology of two dominant meiofaunal species at the HAyenkon Mosby Mud Volcano (HMMV), a deep-sea cold methane-venting seep. Analyses of fatty acids (FAs) and their stable carbon isotopes were used to determine the importance of chemosynthetic nutritional pathways for the dominant copepod species (morphologically very similar to Tisbe wilsoni) inhabiting the volcano's centre and the abundant nematode Halomonhystera disjuncta from the surrounding microbial mats. The strong dominance of bacterial biomarkers (16:1 omega 7c, 18:1 omega 7c and 16:1 omega 8c) coupled with their individual light carbon isotopes signatures (delta C-13 ranging from -52 to -81aEuro degrees) and the lack of symbiotic relationships with prokaryotes (as revealed by molecular analyses and fluorescent in situ hybridisation) indicated that chemosynthetically derived carbon constitutes the main diet of both species. However, the copepod showed a stronger reliance on the utilisation of methanotrophic bacteria and contained polyunsaturated FAs of bacterial origin (20:5 omega 3 and 22:6 omega 3 with isotope signatures delta C-13 < -80aEuro degrees). Instead, the FA profiles of H. disjuncta suggested that sulphide-oxidising bacteria constituted the main diet of this nematode. Therefore, HMMV can be regarded as a persistent deep-sea cold seep, allowing a chemosynthesis-based trophic specialisation by the dominant meiofaunal species inhabiting its sediments. The present investigation, through the determination of the fatty acid profiles, provides the first evidence for trophic specialisation of meiofauna associated with sub-habitats within a cold seep