Maritime midge radiations in the Pacific Ocean (Diptera: Chironomidae)

Maritime chironomid midges (Diptera) are diverse, yet these 'pearls of the ocean' are little known. Emphasizing Pacific Ocean taxa, we used six genetic markers (18S, 28S, CAD1, CAD4, FolCOI and COI) and fossil calibrations to produce Bayesian time-calibrated phylogenies to date eight independent marine transitions in three subfamilies. Deep nodes involve subfamily Telmatogetoninae (originating mid-Cretaceous, 101-128, 114 Ma), with sister genera Telmatogeton Schiner and Thalassomya Schiner splitting later in the Cretaceous (56-82, 69 Ma). Two transitions in Orthocladiinae involve Clunio Haliday and Pseudosmittia Edwards, dating from the upper Cretaceous, both with Eocene crown groups. In subfamily Chironominae, transitions to marine occur in two tribes. Four transitions occur within the otherwise nonmarine crown groups Kiefferulus Goetghebuer, Dicrotendipes Kieffer, Polypedilum Kieffer and Ainuyusurika Sasa & Shirasaka. Two separate robust clades in tribe Tanytarsini involve: (1) a minor radiation within Paratanytarsus dated to the mid-Eocene around 43 Ma; and (2) an unexpected but fully supported diversification in Pontomyia Edwards plus Yaetanytarsus Sasa dated to around 47 Ma, with separation of Pontomyia from Yaetanytarsus around 40 Ma. Crown Pontomyia, represented by three species, was estimated to have diverged around 19 Ma, whereas the crown radiation of Yaetanytarsus, with 12 sampled species, dates to the mid-Eocene. In a comprehensive global review we concisely document new synonymies and new combinations revealed by the study. The evolutionary timing estimate provides insights into the frequency of marine transitions and diversifications in the Chironomidae in association with dynamic oceanic changes during the Oligocene and Miocene.