Closing the gap: a new phylogeny and classification of the chemosymbiotic bivalve family Lucinidae with molecular evidence for 73% of living genera

New molecular phylogenies of the chemosymbiotic bivalve family Lucinidae, using 18S rRNA, 28S rRNA and cytochrome b genes, include species from genera not previously analysed. Notable additions from Myrteinae are sequences from Rostrilucina, Solelucina and Taylorina species, species of Ustalucina, Gonimyrtea from Leucosphaerinae and additional species of Ctena, Codakia, Lucinoma and Divalucina from Codakiinae. New sequences of Lucininae include the type species of Parvilucina (P. tenuisculpta), Liralucina, Falsolucinoma, Easmithia, Jallenia, Radiolucina and Cardiolucina as well as samples of Loripes orbiculatus from multiple localities. Five major clades, defined as subfamilies, are recognized: Pegophyseminae, Myrteinae, Leucosphaerinae, Codakiinae and Lucininae. Two branches, Fimbriinae and Monitilorinae, are represented by single species. Pegophyseminae are an extremely long-branched group with a sister-group relationship to Leucosphaerinae, while Codakiinae are a sister clade to the Lucininae. In various gene trees, the position of Myrteinae is unstable in relation to Pegophyseminae + Leucosphaerinae, Monitilorinae and Fimbriinae. The Myrteinae are not well resolved, with an ambiguous correlation of molecular and morphological characters. Codakiinae now include Divalucina cumingi, shown to be related to Lucinoma rather than Divaricella and Divalinga of the Lucininae. Leucosphaerinae are a well-supported clade but morphologically disparate, with the positions of Gonimyrtea and Callucina unresolved. Several molecularly distinct subclades are recognized within the Lucininae, especially the Lucinisca, Loripes and Parvilucina groups. Parvilucina species are paraphyletic with P. tenuisculpta, the type species, distinct from the western Atlantic species. Codakia, Ctena and Pegophysema have pan-tropical distributions with former connections disrupted by vicariant events of the closure of the eastern Tethyan and Central American Seaways. Species of Radiolucina, Pleurolucina and Lucinisca are present on either side of the Isthmus of Panama. A new classification of the 96 living lucinid genera is presented, providing a framework for future studies of systematics, ecology, biogeography and bacterial symbioses.