Cold seeps occur in geologically active and passive continental margins, where pore waters enriched in methane are forced upward through the sediments by pressure gradients. The advective supply of methane leads to dense microbial communities with high metabolic rates. Anaerobic methane oxidation presumably coupled to sulphate reduction facilitates formation of carbonates and, in many places, generates extremely high concentrations of hydrogen sulphide in pore waters. Increased food supply, availability of hard substratum and high concentrations of methane and sulphide supplied to free-living and symbiotic bacteria provide the basis for the complex ecosystems found at these sites. This review examines the structures of animal communities in seep sediments and how they are shaped by hydrologic, geochemical and microbial processes. The full size range of biota is addressed but emphasis is on the mid-size sediment-dwelling infauna (foraminiferans, metazoan meiofauna and macrofauna), which have received less attention than megafauna or microbes. Megafaunal biomass at seeps, which far exceeds that of surrounding non-seep sediments, is dominated by bivalves (mytilids, vesicomyids, lucinids and thyasirids) and vestimentiferan tube worms, with pogonophorans, cladorhizid sponges, gastropods and shrimp sometimes abundant. In contrast, seep sediments at shelf and upper slope depths have infaunal densities that often differ very little from those in ambient sediments. At greater depths, seep infauna exhibit enhanced densities, modified composition and reduced diversity relative to background sediments. Dorvilleid, hesionid and ampharetid polychaetes, nematodes, and calcareous foraminiferans are dominant. There is extensive spatial heterogeneity of microbes and higher organisms at seeps. Specialized infaunal communities are associated with different seep habitats (microbial mats, clam beds, mussel beds and tube worms aggregations) and with different vertical zones in the sediment. Whereas fluid flow and associated porewater properties, in particular sulphide concentration, appear to regulate the distribution, physiological adaptations and sometimes behaviour of many seep biota, sometimes the reverse is true. Animal-microbe interactions at seeps are complex and involve symbioses, heterotrophic nutrition, geochemical feedbacks and habitat structure. Nutrition of seep fauna varies, with thiotrophic and methanotrophic symbiotic bacteria fueling most of the megafaunal forms but macrofauna and most meiofauna are mainly heterotrophic. Macrofaunal food sources are largely photosynthesis-based at shallower seeps but reflect carbon fixation by chemosynthesis and considerable incorporation of methane-derived C at deeper seeps. Export of seep carbon appears to be highly localized based on limited studies in the Gulf of Mexico. Seep ecosystems remain one of the ocean's true frontiers. Seep sediments represent some of the most extreme marine conditions and offer unbounded opportunities for discovery in the realms of animal-microbe-geochemical interactions, physiology, trophic ecology, biogeography, system-atics and evolution.

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Ecology of cold seep sediments: interactions of fauna with flow, chemistry and microbes

Although initially viewed as oases within a barren deep ocean, hydrothermal vent and methane seep communities are now recognized to interact with surrounding ecosystems on the sea floor and in the water column, and to affect global geochemical cycles. The importance of understanding these interactions is growing as the potential rises for disturbance from oil and gas extraction, seabed mining and bottom trawling. Here we synthesize current knowledge of the nature, extent and time and space scales of vent and seep interactions with background systems. We document an expanded footprint beyond the site of local venting or seepage with respect to elemental cycling and energy flux, habitat use, trophic interactions, and connectivity. Heat and energy are released, global biogeochemical and elemental cycles are modified, and particulates are transported widely in plumes. Hard and biotic substrates produced at vents and seeps are used by “benthic background” fauna for attachment substrata, shelter, and access to food via grazing or through position in the current, while particulates and fluid fluxes modify planktonic microbial communities. Chemosynthetic production provides nutrition to a host of benthic and planktonic heterotrophic background species through multiple horizontal and vertical transfer pathways assisted by flow, gamete release, animal movements, and succession, but these pathways remain poorly known. Shared species, genera and families indicate that ecological and evolutionary connectivity exists among vents, seeps, organic falls and background communities in the deep sea; the genetic linkages with inactive vents and seeps and background assemblages however, are practically unstudied. The waning of venting or seepage activity generates major transitions in space and time that create links to surrounding ecosystems, often with identifiable ecotones or successional stages. The nature of all these interactions is dependent on water depth, as well as regional oceanography and biodiversity. Many ecosystem services are associated with the interactions and transitions between chemosynthetic and background ecosystems, for example carbon cycling and sequestration, fisheries production, and a host of non-market and cultural services. The quantification of the sphere of influence of vents and seeps could be beneficial to better management of deep-sea environments in the face of growing industrialization.

Hydrothermal Vents and Methane Seeps: Rethinking the Sphere of Influence

Mining of deep-sea seafloor massive sulfides: A review of the deposits, their benthic communities, impacts from mining, regulatory frameworks and management strategies

The 3- to 5-m-thick Doushantuo cap carbonate in south China overlies the glaciogenic Nantuo Formation (ca. 635 Ma) and consists of laterally persistent, thinly laminated and normally graded dolomite and limestone indicative of relatively deep-water deposition, most likely below storm wave base. The basal portion of this carbonate contains a distinctive suite of closely associated tepee-like structures, stromatactis-like cavities, layer-parallel sheet cracks, and cemented breccias. The cores of tepees are composed of stacked cavities lined by cements and brecciated host dolomicrite. Onlap by laminated sediment indicates synsedimentary disruption of bedding that resulted in a positive seafloor expression. Cavities and sheet cracks contain internal sediments, and they are lined by originally aragonitic isopachous botryoidal cements with acicular radiating needles, now replaced by dolomite and silica. Pyrite and barite are common, and calcite is locally retained as a primary mineral. These features share morphological and petrographic attributes with modern and ancient methane seeps in which methane gas and fluids provide both a force for physical disruption from buoyancy and a source of alkalinity for significant cementation. The presence of δ13C values as low as −41‰ in well preserved limestone crusts and cements within and immediately above the tepee-like structures provides unequivocal evidence for methane influence, and the widespread distribution of identical sedimentary structures and paragenetic cement sequences across the entire basin at the same basal cap carbonate level is consistent with gas hydrate destabilization and the development of methane seeps as a result of postglacial warming of the ocean. Considering the broad distribution of similar features at the same stratigraphic level in other cap carbonates globally, we suggest that the late Neoproterozoic postglacial methane release may have influenced the oceanic oxygen level as well as contributed to postglacial warming via the greenhouse effects of methane.

Stratigraphy, Sedimentary Structures, and Textures of the Late Neoproterozoic Doushantuo Cap Carbonate in South China

Relationships among fluid flow, sulfide concentration, sulfur bacteria and macrofaunal assemblages were examined at methane seeps on the northern California margin, near the mouth of the Eel River (512 to 525 m). Over a 6 mo period, sediments covered with microbial mats exhibited significant but variable outflow of altered fluids, with no flow reversals. This fluid flow was associated with high porewater sulfide concentrations (up to 20 mM) and almost no oxygen penetration of sedi- ments ( 300 µm), but biomass and diversity (no. species per core, E(S100), H ') were lower and composition varied in the sulfidic microbial mat sediments relative to clam-bed and non-seep sediments. The community in microbial mat-covered sediments consisted largely (82%) of 6 species in the polychaete family Dorvilleidae, whereas the clam-bed and non-seep microhabitats supported a mixture of annelids, peracarid crustaceans, nemerteans, and mollusks. Vertical microprofiling of sulfide in animal cores indicated that most taxa avoid H2S concentrations >1 mM. However, sulfide-oxidizing filamentous bacteria, dorvilleid polychaetes and bivalves (mainly V. pacifica) exhibited highest densities at sul- fide concentrations of 1 to 5 mM sulfide. Horizontal and vertical patterns of sulfide availability have a strong influence on the fine-scale distribution, structure and composition of macrofaunal assem- blages inhabiting methane seeps and must be accounted for when characterizing the microbiology and ecology of seep habitats.

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Spatial heterogeneity of macrofauna at northern California methane seeps: influence of sulfide concentration and fluid flow

Observations from the first submersible reconnaissance of the Blake Ridge Diapir provide the geological and ecological contexts for chemosynthetic communities established in close association with methane seeps. The seeps mark the loci of focused venting of methane from the gas hydrate reservoir, and, in one location (Hole 996D of the Ocean Drilling Program), methane emitted at the seafloor was observed forming gas hydrate on the underside of a carbonate overhang. Megafaunal elements of a chemosynthetically based community mapped onto dive tracks provide a preliminary overview of faunal distributions and habitat heterogeneity. Dense mussel beds were prominent and covered 20 � 20 m areas. The nearly non-overlapping distributions of mussels and clams indicate that there may be local (meter-scale) variations in fluid flux and chemistry within the seep site. Preliminary evidence suggests that the mussels are host to two symbiont types (sulfide-oxidizing thiotrophs and methanotrophs), while the clams derive their nutrition only from thiotrophic bacteria. Invertebrate biomass is dominated by mussels (Bathymodiolus heckerae) that reach lengths of up to 364 mm and, to a lesser extent, by small (22 mm length) vesicomyid clams (Vesicomya cf. venusta). Taking into account biomass distributions among taxa, symbiont characteristics of the bivalves, and stable-isotope analyses, the relative importance of methanotrophic vs thiotrophic bacteria in the overall nutrition of the invertebrate

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Blake Ridge methane seeps: characterization of a soft-sediment, chemosynthetically based ecosystem

Estimations of biodiversity and species richness in deep-sea marine ecosystems are impeded by time-consuming methods of species identification. In conservation biology, in environmental monitoring, and in paleontology, a higher-taxon approach (e.g., identification to genera or families) can be used as a surrogate for species richness. We applied a higher-taxon approach to well-documented chemosynthetic communities associated with seep and vent mussel beds to test its applicability in these systems. Significant positive correlations between cumulative number of species and cumulative number of higher taxa were found at the generic, family, and order levels. The number of these higher taxa can be used to predict species richness in vent and seep mussel beds.

Higher-taxon richness as a surrogate for species richness in chemosynthetic communities

Species lists for vent fields on the Mid-Atlantic Ridge (MAR) from 14°N to 38°N suggest that there is a northern (>27°N), shallow ( 3000 m) endemic vent fauna, but little is known about how community structure varies along the ridge axis. In this study, quantitative samples of macrofaunal invertebrates associated with mussels (Bathymodiolus puteoserpentis) were collected at Logatchev (14°45′N), the southern-most explored vent field on the MAR. Community structure (including species composition, species richness, diversity, and relative species abundances) in mussel beds at Logatchev was compared with that of Snake Pit (23°22′N) and Lucky Strike (37°17′N) mussel beds. The most striking feature of the Logatchev mussel-bed macrofaunal invertebrate community was the tremendous abundance (up to 2390 individuals per liter of mussel-volume sampled) and biomass of the ophiuroid, Ophioctenella acies. Logatchev and Snake Pit mussel beds share >50% of their associated macrofaunal species; these two sites share only 20–25% of their macrofaunal species with Lucky Strike. Species–effort curves and univariate measures of diversity (H′, J′) do not support the claim that diversity of vent organisms on the MAR is highest at Logatchev, at least when one assesses this within a habitat type. Multivariate analysis readily differentiates the species-abundance characteristics of Logatchev, Snake Pit, and Lucky Strike mussel-bed macrofaunas. The relationship between sea-floor spreading rate and diversity was explored through comparison of species richness in mussel-bed habitats on slow-spreading (MAR), fast-spreading [northern East Pacific Rise (EPR)], and ultra-fast-spreading (southern EPR) mid-ocean ridges. Species richness was greater in samples from the faster-spreading ridge axes, where vents are more closely spaced but shorter lived, than on slow-spreading centers, where vents are further apart but longer lived.

Community structure in mussel beds at Logatchev hydrothermal vents and a comparison of macrofaunal species richness on slow‐ and fast‐spreading mid‐ocean ridges

M.B. Doerries

Papers

Blake Ridge methane seeps: characterization of a soft-sediment, chemosynthetically based ecosystem

Higher-taxon richness as a surrogate for species richness in chemosynthetic communities

Community structure in mussel beds at Logatchev hydrothermal vents and a comparison of macrofaunal species richness on slow‐ and fast‐spreading mid‐ocean ridges