Phylogenetic relationships and convergent evolution of ocean-shore ground beetles (Coleoptera: Carabidae: Trechinae: Bembidion and relatives)

Through phylogenetic analysis of seven genes, we show that there have been at least six independent entries into intertidal habitats in the history of bembidiine carabids, in the ancestors of: (i) Orzolina Machado, (ii) Bembidion (Desarmatocillenus Netolitzky), (iii) Bembidion laticeps (LeConte) + palosverdes Kavanaugh & Erwin, (iv) Bembidion laterale (Samouelle), (v) Bembidion umi Sasakawa and Bembidion quadriimpressum (Motschulsky) (which may represent two separate entries), and (vi) B. nigropiceum (Marsham). The following lineages are widely separated within the subtribe Bembidiina: Orzolina is sister to the genus Ocys Stephens; subgenus Desarmatocillenus appears to be sister to all Bembidion Latreille excluding subgenus Phyla Motschulsky; B. laticeps + B. palosverdes is a clade in the Bembidion series; B. laterale is a member of the Princidium Motschulsky complex; B. umi and B. quadriimpressum are related to the Nearctic Clade of the Ocydromus Clairville complex; B. nigropiceum is sister to B. praeustum Dejean among sampled species. There are three separate lineages of ocean-shore bembidiines that are known to prey on amphipods, and adults of these lineages [Bembidion (Desarmatocillenus), B. laterale, and B. mandibulare Solier] have unusually wide heads with long mandibles. Also common among independent lineages restricted to ocean shores are prominent front angles of the prothorax, larger numbers of setae on the elytral disc, a notable sinuation in the margin of each elytron near its apex, and short, wide mesotarsi. The reasons for the repeated evolution of these features are not evident. Our results also suggest that inland species of the Nearctic Clade may have arisen from an ocean-shore ancestor. The close genetic similarities between the gravel river shore dwelling B. praeustum and the intertidal specialist B. nigropiceum suggest that the striking morphological adaptations of B. nigropiceum to the intertidal zone arose rapidly. We make one nomenclatural change: we resurrect the subgenus Lymneops Casey to accommodate B. palosverdes and B. laticeps.