Continued evolutionary surprises among dinoflagellates -- Morden and Sherwood 99 (18): 11558 -- Proceedings of the National Academy of Sciences
"It is well established that chloroplasts in green and red algae are derived from a primary endosymbiotic event between a cyanobacterium and a eukaryotic organism 1 billion years ago (Fig. 1; refs. 1 and 2). Although these two groups account for many of the world's photosynthetic species, most other major taxonomic groups of photosynthetic organisms (stramenopilesincluding diatoms, phaeophytes, chrysophytesand haptophytes) have plastids derived from a photosynthetic eukaryote implying a secondary endosymbiosis (1, 2). Still other groups, such as the dinoflagellates, have more complicated associations believed to be derived from tertiary endosymbioses involving the engulfment of a secondary endosymbiont. Each endosymbiotic event has characteristic structural changes associated with it, the most notable of which is the addition of two membranes surrounding the plastid (the inner representing the cell membrane of the engulfed organism and the outer representing the phagocytosis vacuole membrane) (2). Dinoflagellates, although believed to be tertiary endosymbionts, have only 3 membranes surrounding their plastids (1, 2), suggesting that the acquisition of too many membranes may be functionally unstable and can cause some to be lost. "
Clifford W. Morden, and Alison R. Sherwood Continued evolutionary surprises among dinoflagellates PNAS September 3, 2002 vol. 99 no. 18 11558-11560