mRNA editing and spliced-leader RNA trans-splicing groups Oxyrrhis, noctiluca, Heterocapsa, and Amphidinium as basal lineages of dinoflagellates

Identification of novel dinoflagellate taxa through molecular analysis is hindered by lack of well-defined basal lineages. To address this issue, we attempted to reassess the phylogenetic status of Oxyrrhis marina Dujard. as well as other potentially basal taxa. The analysis was based on two newly established premises: (1) editing density of mitochondrial cob and cox1 mRNA increases from basal to later diverging lineages; (2) nuclear-encoded mRNA in dinoflagellates is trans-spliced to receive a 22 bp spliced leader (SL) at the 5'-end. We analyzed these two genetic traits in O. marina, Noctiluca scintillans (Macartney) Kof. et Swezy, Heterocapsa triquetra (Ehrenb.) F. Stein, H. rotundata (Lohmann) Ge. Hansen, Amphidinium carterae Hulburt, and A. operculatum Clap. et J. Lachm. Surprisingly, no editing was detected in cob and cox1 mRNAs in these lineages, except for a small number of editing events in Amphidinium. However, nuclear-encoded mRNAs in these species contained the SL sequence at the 5'-end, indicative of SL RNA trans-splicing. These findings, together with the recent cob-cox1-18S rRNA three-gene phylogeny, suggest the following: (1) O. marina is a basal dinoflagellate; (2) Heterocapsa, Amphidinium, and Noctiluca likely are also early diverging lineages of dinoflagellates, and the position of Heterocapsa is inconsistent with literature and needs further investigation; and (3) the presence of the 22 bp SL and mitochondrial (mt) mRNA editing can be considered a landmark of dinoflagellate splits.