Shell microstructures of mussels (Bivalvia: Mytilidae: Bathymodiolinae) from deep-sea chemosynthetic sites: Do they have a phylogenetic significance?

Abstract

The increasing number of bathymodiolin mussel species being described from deep-sea chemosynthetic environments worldwide has raised many questions about their evolutionary history, and their systematics is still being debated. Mussels are also abundant in fossil chemosynthetic assemblages, but their identification is problematic due to conservative shell morphology within the group and preservation issues. Potential resolution of bathymodiolin taxonomy requires new character sets, including morphological features that are likely to be preserved in fossil specimens. To investigate the phylogenetic significance of shell microstructural features, we studied the shell microstructure and mineralogy of 10 mussel species from hydrothermal vents and hydrocarbon seeps, and 15 taxa from sunken wood and bone habitats, and compared these observations with current molecular phylogenies of the sub-family Bathymodiolinae. In addition, we analyzed the shell microstructure in Adipicola chickubetsuensis from fossil whale carcasses, and in Bathymodiolus cf. willapaensis and "Modiola exbrocchii" from fossil cold seeps, and discussed the usefulness of these characters for identification of fossil chemosymbiotic mussels. Microstructural shell features are quite uniform among vent, seep, wood and bone mussel taxa, and therefore established bathymodiolin lineages cannot be discriminated, nor can the relations between fossil and modern species be determined with these characters. Nevertheless, the uniformity of shell microstructures observed among chemosymbiotic mussels and the similarity with its closest relative, Modiolus modiolus, does not challenge the monophyly of the group. Slight differences are found between the large vent and seep mussels and the small mytilids commonly found in habitats enriched in organic matter. Together with previous data, these results indicate that a repeated pattern of paedomorphism characterizes the evolutionary history of deep-sea mussels, and the occurrence of neotenous features should be considered in the taxonomic revision of this group. (C) 2012 Elsevier Ltd. All rights reserved.