Phylogeny of some of the major genera of dinoflagellates based on ultrastructure and partial LSU rDNA sequence data, including the erection of three new genera of unarmoured dinoflagellates

Evidence from partial large-subunit (LSU) rDNA sequencing has been combined with ultrastructure, including details of the flagellar apparatus, in a number of phototrophic dinoflagellates, with the aim of trying to solve some of the most pressing taxonomic problems and to contribute to an improved understanding of the phylogeny within the group. Special attention has been paid to the unarmoured (naked) genera, many of which were described during the 1800s or early 1900s and whose taxonomy is artificial and misleading. This is particularly unsatisfactory because many of the species cause extensive plankton blooms, fish kills and other harmful events. Our studies have indicated that the path of the so-called apical groove (acrobase) is of particular importance for the taxonomy of the unarmoured genera of dinoflagellates. Features presently used to characterize many of the genera, such as the relative size of the epicone and hypocone, are misleading. Our data have resulted in the splitting of the large genus Gymnodinium into four genera. The fish-killing species are confined to two genera, Karenia G. Hansen & Moestrup gen. nov. and Karlodinium J. Larsen gen. nov. The paralytic shellfish poisoning-producing species Gymnodinium catenatum is retained within Gymnodinium, together with a number of harmless species. The fourth genus, Akashiwo G. Hansen & Moestrup gen. nov., presently comprises only the large nontoxic species previously known as Gymnodinium sanguineum. The genus Gyrodinium is redefined. The genus Amphidinium is artificial, but more data are needed before redescription of the genus can be made with any confidence. Within the armoured dinoflagellates, LSU and previously published small-subunit rDNA data show the Gonyaulacales to be a natural group. Peridiniella catenata, sometimes included in the Gonyaulacales based on gross morphology, falls outside this order both genetically and ultrastructurally. Based on the DNA data, the genus Peridinium appears to be artificial. Ultrastructure as well as gene sequences confirm that the genus Heterocapsa is unusual, since it includes both apparently unarmoured species (but with very thin plates) and armoured species.