Inorganic sulfate-based signatures of chemosymbiosis in modern infaunal lucinids

Bivalves that host sulfur-oxidizing bacterial gill-hosted endosymbionts can inhabit low diversity, sulfidic environmental niches. However, understanding the history of this life strategy is limited by the lack of a robust method that can be applied to fossils. Measurements of carbonate-associated sulfate S isotope ratios (CAS -634S) in carbonate fossils could fill this void by fingerprinting symbiont-driven oxidation of environmental sulfide. We begin to evaluate this prediction using modern lucinid bivalves, a useful test case because: (1) all modern genera host symbionts and live in sulfidic sediments, and (2) morphological evidence suggests that this has been true since the earliest ancestral lucinids. We measured S speciation, abundance, and CAS -634S values in the shells of a suite of modern infaunal lucinids, in addition to epifaunal bivalves with and without S-oxidizing symbionts as controls. For infaunal lucinids, CAS concentrations were at most one-third of those of non-symbiotic epifaunal bivalves, and CAS -634S values were lower (9.2%0-18.5%0) than in modern seawater (21%0) or epifaunal bivalves (20.8%0-21%0). These observations indicate that lucinids with symbionts incorporate sulfide-derived sulfate into their shells as a direct consequence of their chemosymbiosis. We argue that both the concentration and the magnitude of 34S depletion in infaunal lucinid CAS reflect environmental sulfide concentrations and could viably reveal chemosymbiosis in fossils.