The intricacies of dinoflagellate pellicle cysts: The example of Alexandrium minutum cysts from a bloom-recurrent area (Bay of Baiona, NW Spain)

The terms “temporary”, “pellicle”, and “ecdysal” cyst have been employed arbitrarily in the literature of the dinoflagellate life cycle to describe a non-motile and single-layered-wall stage with no mandatory dormancy period, of asexual or sexual origin. These three terms have been used more or less synonymously, but more specific definitions, taking into account morphological and physiological aspects and their roles in dinoflagellate population dynamics, are still needed. To clarify the current terminology, we examine and discuss the usages and foundations of those terms. The background for this discussion is provided by a comparison of the morphology and germination times of three different types of Alexandrium minutum cysts collected during a seasonal bloom in the Bay of Baiona (NW Spain). The double-walled cysts were similar to the resting cysts reported for this species, but other, thin-walled and thecate cysts were also observed. These latter cyst types needed between 1 and 17 days to germinate and were therefore considered as short-term cysts, in contrast to the 1.5-month dormancy period of resting (hypnozygotic) cysts. Our results showed that the temporal distribution of these short-term cysts during the bloom period followed a pattern very similar to that of vegetative cells. However, resting cysts were only detected at the end of the bloom. In the context of our present knowledge regarding the dormancy and quiescence of dinoflagellate cysts, “temporary” is a very misleading and uncertain term and must be rejected. The term “ecdysal” has been used in reference to thin-walled cysts when ecdysis has been proven; however, ecdysis is not unique to this type of cysts as thick-walled zygotic cysts can be formed thorough ecdysis of a thecate planozygote. In conclusion, based on our current understanding of cysts, the term “pellicle” more appropriately describes single-layered-wall stages.