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A new molecular phylogeny of the chemosymbiotic bivalve family Lucinidae is presented. Using sequences from the nuclear 18S and 28S rRNA genes and the mitochondrial gene cytochrome b, 105 specimens were analysed representing 87 separate species classified into 47 genera. Samples were collected from a wide range of habitats including mangroves, seagrass beds, shallow sands, offshore muds, and hydrocarbon seeps at depths ranging from the intertidal to over 2000 m. A chronogram, derived from the combined molecular tree, was calibrated using ten lucinid fossils. The trees show five well-supported clades and two single branches of Fimbria fimbriata (Linnaeus, 1758) and Monitilora ramsayi (Smith, 1885). A new classification of Lucinidae is proposed with seven subfamilial divisions: three new subfamilies ‐ Pegophyseminae, Leucosphaerinae, and Monitilorinae ‐ are introduced and Codakiinae, usually treated as a synonym of Lucininae, is revived to include the Lucinoma, Codakia, and Ctena subclades. Membership of the Lucininae and Myrteinae is considerably revised compared with Chavan’s commonly employed ‘Treatise’ classification. Previously considered as a separate family, Fimbriinae is now regarded as a subfamily within Lucinidae. The status of Milthinae is presently equivocal pending further analysis and Divaricellinae is recognized as polyphyletic, and is therefore abandoned, with species and genera now grouped in various places within the Lucininae. Deeper water Lucinidae mainly belong to Leucosphaerinae, Codakiinae (Lucinoma clade), and Myrteinae, with Lucinoma species being most frequently associated with hydrocarbon seeps. Species occurring in seagrass habitats derive largely from Pegophyseminae, Codakiinae, and Lucininae, and species from mangrove habitats derive from the Pegophyseminae and Lucininae.

Recent molecular analyses have demonstrated that the traditional Lucinoidea, comprising the extant families Lucinidae, Thyasiridae, Ungulinidae, Fimbriidae, and Cyrenoididae, is not monophyletic. Thyasiridae and Ungulinidae are unrelated to Lucinidae, a result corroborated by clear morphological differences between the groups. Chemosymbiosis in Thyasiridae and Lucinidae has been independently derived. Within the family Lucinidae, previous ideas of relationship and subfamilial divisions based on shell characters have little support from molecular results. Anatomical characters of the ctenidia, mantle gills, and posterior apertures have potential in phylogenetic analysis but rigorous analysis of shell characters is also needed. Although there is a good fossil record of Lucinidae throughout the Cenozoic and Mesozoic, in the Palaeozoic fossils are less frequent and most need reappraisal. The Silurian Ilionia prisca is probably the earliest fossil with convincing lucinid features, followed in the Devonian by Phenacocyclas and some Paracyclas species. © 2006 The Linnean Society of London, Zoological Journal of the Linnean Society, 2006, 148, 421–438.