Studies on woloszynskioid dinoflagellates IX: ultrastructure, cyst formation and phylogeny of the ‘red-snow’ alga Borghiella pascheri (Suchlandt) Moestrup (= Glenodinium pascheri, Woloszynskia pascheri, Gyrodinium nivalis)

Abstract

ABSTRACTRed snow caused by dinoflagellates is a phenomenon rarely reported, described from the European Alps from 1914 onwards, and subsequently observed outside Europe on several occasions in Ontario, Canada. Considerable taxonomic confusion exists regarding the identity of the organism(s) causing red snow, but the most recent occurrence in 2016 in Ontario has now allowed detailed studies, including LM, SEM, TEM and molecular sequencing of the causative species. We conclude that the two species originally described as the cause of red snow, Glenodinium pascheri and Gyrodinium nivale, are synonymous and that the appropriate name for the organism is Borghiella pascheri (syn. Woloszynskia pascheri) as suggested by Moestrup & Calado in the recent volume of the Süsswasserflora. The central part of Borghiella pascheri cells is tomato red and filled with numerous organelles, whose ultrastructure indicates modified chloroplasts. Lack of cultures has prevented chemical characterization of the red pigment. Formation of temporary cysts was common in the samples. Transformation of the motile cells into temporary cysts was followed in detail, and the cysts were shown to be surrounded by the fused inner membranes of the amphiesmal vesicles, which became the cell membrane of the cysts, covered by the fused pellicle precursors. The cell membrane from the motile cell was discarded together with the outer amphiesmal vesicle membrane and the thin thecal plates, and the temporary cysts were therefore not surrounded by any pattern of vesicles. Sexual reproduction resulted in the formation of hypnozygotes. Although the species possessed several unusual features, DNA sequencing showed it to belong to Borghiella. The culture established in 1965 from the Botanical Garden in Göttingen, Germany and generally identified as Woloszynskia pascheri belongs to a separate species of Borghiella, to be described separately.The occurrence of red snow caused by dinoflagellates is discussed.