Absence of co-phylogeny indicates repeated diatom capture in dinophytes hosting a tertiary endosymbiont

Tertiary endosymbiosis is proven through dinophytes, some of which (i.e. Kryptoperidiniaceae) have engulfed diatom algae containing a secondary plastid. Chloroplasts are usually inherited together permanently with the host cell, leading to co-phylogeny. We compiled a diatom sequence data matrix of two nuclear and two chloroplast loci. Almost all endosymbionts of Kryptoperidiniaceae found their closest relatives in free-living diatoms and not in other harboured algae, rejecting co-phylogeny and indicating that resident diatoms were taken up by dinophytes multiple times independently. Almost intact ultrastructure and insignificant genome reduction are supportive for young, if not recent events of diatom capture. With their selective specificity on the one hand and extraordinary degree of endosymbiotic flexibility on the other hand, dinophytes hosting diatoms share more traits with lichens or facultatively phototrophic ciliates than with green algae and land plants. Time estimates indicate the dinophyte lineages as consistently older than the hosted diatom lineages, thus also favouring a repeated uptake of endosymbionts. The complex ecological role of dinophytes employing a variety of organismic interactions may explain their high potential and plasticity in acquiring a great diversity of plastids.