THE EARLIEST BATHYMODIOLIN MUSSELS: AN EVALUATION OF EOCENE AND OLIGOCENE TAXA FROM DEEP-SEA METHANE SEEP DEPOSITS IN WESTERN WASHINGTON STATE, USA

Bathymodiolin mussels are a group of bivalves associated with deep-sea hydrothermal vents and other reducing deep-sea habitats, and they have a particularly rich early Cenozoic fossil record in western Washington State, U.S.A. Here we recognize six species from middle Eocene to latest Oligocene deep-water methane seep deposits in western Washington. Two of them are new: Vulcanidas? goederti from the middle Eocene Humptulips Formation and Bathymodiolus (sensu lato) satsopensis from the late Oligocene part of the Lincoln Creek Formation. Very similar to the latter but more elongate are specimens from the early Oligocene Jansen Creek Member of the Makah Formation and are identified as B. (s.l.) aff. satsopensis. Bathymodiolus (s.l.) inouei Amano and Jenkins, 2011 is reported from the Lincoln Creek Formation. Idas? olympicus Kiel and Goedert, 2007 was previously known from late Eocene to Oligocene whale and wood falls in western Washington and is here reported from Oligocene seep deposits of the Makah and Pysht Formations. Vulcanidas? goederti occurs at a seep deposit from a paleodepth possibly as great as 2000 m, suggesting that its living relative, Vulcanidas insolatus Cosel and Marshall, 2010, which lives at depths of only 150-500 m, is derived from a deep-water ancestor. The bathymodiolins in western Washington indicate that the group originated at least in the middle Eocene and underwent a first diversification in the late Eocene to Oligocene. Early ontogenetic shells of all fossil species investigated so far, including the middle Eocene Vulcanidas? goederti, reflect planktotrophic larval development indicating that this developmental mode is an ancestral trait of bathymodiolins.