New investigation of the cyst–motile relationship for Votadinium spinosum reveals a Protoperidinium claudicans species complex (Dinophyceae, Peridiniales)

Protoperidinium claudicans is a planktonic, heterotrophic, bioluminescent dinoflagellate species commonly found in neritic waters. It has long been considered to display phenotypic variation in its second anterior intercalary plate, which could vary between quadra, penta and hexa. The equivalent spinose, cordate cyst goes under the name of Votadinium spinosum. Here we perform incubation experiments with cysts from France, Canada, China and Japan, which demonstrate that P. claudicans forms a species complex, with at least two ribotypes with a penta configuration (P. claudicans) and one with a quadra configuration (P. carriae sp. nov.). A fossil-based cyst, V. multispinosum sp. nov., is described as the equivalent of P. carriae. Molecular phylogenetics using large subunit ribosomal DNA supports these observations. The cyst–theca relationships for Votadinium psilodora and another, undescribed, cordate, spineless Votadinium species are also reported from China. Macromolecular analyses of the cyst wall of V. multispinosum reveal it is comprised of a protein-rich carbohydrate compound. We show that this compound is not uncommon in dinoflagellate and ciliate cysts and that it is unlikely to preserve very well upon sedimentation and burial.