Mapping the missing branch on the neogastropod tree of life: molecular phylogeny of marginelliform gastropods
ABSTRACT Marginelliform gastropods are a heterogeneous and diverse group of molluscs encompassing over 1,600 living species, among which are the smallest known neogastropods. The relationships of marginelliform gastropods within the order Neogastropoda are controversial, and the monophyly of the two marginelliform families the Marginellidae J. Fleming, 1828 and the Cystiscidae Stimpson, 1865, remains unconfirmed. DNA sequence data have never been used to assess the relationships of the marginelliform gastropods, making this group the only major branch missing in our current understanding of the neogastropod tree of life. Here we report results of the first multilocus phylogenetic analysis of marginelliform gastropods, which is based on a dataset comprising 63 species (20 genera) of Marginellidae and Cystiscidae, and a wide range of neogastropod lineages. The Marginellidae and Cystiscidae form a moderately supported clade that is sister to the family Volutidae. Marginellona gigas appears to be sister to all other marginelliforms. The subfamily Marginellinae was recovered as a well-supported clade, and good resolution of this part of the tree makes it possible to propose amendments to the family-level classification of the group. The relationship between Granulina and other marginelliforms could not be resolved and requires further study. Due to poor resolution of basal relationships within the Marginellidae–Cystiscidae clade, the monophyly of the Cystiscidae was neither confirmed nor convincingly rejected. The shell morphology of most marginellid and cystiscid genera is taxonomically not very informative but, nevertheless, of the traditionally recognized genera only Gibberula and Dentimargo were shown to be polyphyletic. Although a comprehensive systematic revision of the group requires more extensive taxonomic sampling (e.g. with better representation of the type species of nominal genus-group names), our results support the superfamily Volutoidea, comprising four families (Volutidae, Cystiscidae, Marginellidae and Marginellonidae), with the placement of the Granulinidae uncertain for the time being.