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Book Chapter
Levin, LA.  2005.  Ecology of cold seep sediments: Interactions of fauna with flow, chemistry and microbes. Oceanography and Marine Biology - an Annual Review, Vol. 43. 43( Gibson RN, Atkinson RJA, Gordon JDM, Eds.).:1-46., Boca Raton: Crc Press-Taylor & Francis Group Abstract

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, systematics and evolution.

Journal Article
Levin, LA, Ziebis W, Mendoza GF, Growney VA, Tryon MD, Brown KM, Mahn C, Gieskes JM, Rathburn AE.  2003.  Spatial heterogeneity of macrofauna at northern California methane seeps: influence of sulfide concentration and fluid flow. Marine Ecology-Progress Series. 265:123-139.   10.3354/meps265123   AbstractWebsite

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 sediments (<0.1 mm). Vesicomya pacifica (clam) bed and non-seep sediments exhibited little net fluid outflow and similar oxygen penetration (3 and 4 mm, respectively); however, sulfide concentrations were higher in subsurface clam-bed sediments (up to 2 mM) than in non-seep sediments (<200 muM). Macrofaunal densities did not differ among the 3 habitats (13 800 to 16 800 ind. m(-2); >300 mum), but biomass and diversity (no. species per core, E(S-100), 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 sulfide 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 assemblages inhabiting methane seeps and must be accounted for when characterizing the microbiology and ecology of seep habitats.