Deep-Sea Hydrocarbon Seep Communities: Evidence for Energy and Nutritional Carbon Sources

Mussels, clams, and tube worms collected in the vicinity of hydrocarbon seeps on the Louisiana slope contain mostly ``dead'' carbon, indicating that dietary carbon is largely derived from seeping oil and gas. Enzyme assays, elemental sulfur analysis, and carbon dioxide fixation studies demonstrate that vestimentiferan tube worms and three clam species contain intracellular, autotrophic sulfur bacterial symbionts. Carbon isotopic ratios of 246 individual animal tissues were used to differentiate heterotrophic ($\delta ^{13}$C = -14 to -20 per mil), sulfur-based ($\delta ^{13}$C = -30 to -42 per mil), and methane-based ($\delta ^{13}$C = < -40 per mil) energy sources. Mussels with symbiotic methanotrophic bacteria reflect the carbon isotopic composition of the methane source. Isotopically light nitrogen and sulfur confirm the chemoautotrophic nature of the seep animals. Sulfur-based chemosynthetic animals contain isotopically light sulfur, whereas methane-based symbiotic mussels more closely reflect the heavier oceanic sulfate pool. The nitrogen requirement of some seep animals may be supported by nitrogen-fixing bacteria. Some grazing neogastropods have isotopic values characteristic of chemosynthetic animals, suggesting the transfer of carbon into the background deep-sea fauna.