Dinoflagellate Evolution: Speculation and Evidence*†

SYNOPSIS. Nuclear features of dinoflagellates that were used originally to support the Mesocaryota concept are reviewed. The fibrillar diameter of dinoflagellate chromatin, low level of chromosomal basic proteins, membrane attachment of chromosomes and swirl pattern observed in sectioned chromosomes are features that support a prokaryotic affinity. The presence of repeated and highly complex DNA, a S‐phase of DNA synthesis in the cell cycle, presence of basic proteins, and the reinterpretation of extranuclear microtubules as a spindle support the contention that dinoflagellates are eukaryotes. This combination of prokaryotic and eukaryotic features suggests that dinoflagellates are a geologically old group and that perhaps they diverged from the higher eukaryotic lineage before evolution of eukaryotic chromatin but after the evolution of repeated DNA. The 2 patterns of carotenoid composition exemplified by the presence of peridinin or fucoxanthin suggest separate origins of dinoflagellate plastids, perhaps by prokaryotic and eukaryotic capture. It is suggested that the species possessing fucoxanthin obtained their plastids by capture of photosynthetic eukaryotes. A new class and order, Syndiniophyceae and Syndiniales, are proposed for the dinoflagellates with low chromosome numbers, V‐shaped chromosomes, chromosomes containing a sufficient quantity of basic proteins detectable histochemically, possession of centrioles associated with mitosis, intracellular parasitism as a mode of nutrition, and lack of a cellular covering containing plates. Ultrastructural and paleontologic evidence indicates that the thecate is more primitive than the nonthecate condition. The Prorocentrales are considered to be primitive and their thecal construction is reinterpreted as having epithecal and hypothecal regions surrounding a flagellar pore region containing 7 plates. Acritarchs resemble cysts of modern dinoflagellates in size, structure, and chemical composition except for the absence of a polygonal excystment aperture and lack of any indication of transverse and longitudinal flagellar grooves on the acritarchs. The suggestion that some acritarchs may have dinoflagellate affinities is supported by the occurrence of modern dinoflagellates (Prorocentrales) which lack a theca of numerous polygonal plates and lack transverse and longitudinal flagellar arrangement. The Prorocentrales, as opposed to the more typical Dinophyceae, perhaps represent the type of organism that produced some acritarchs.