Morphology, fine structure, life cycle and phylogenetic analysis ofPhyllosiphon arisari, a siphonous parasitic green alga

Phyllosiphon arisari Kühn (Phyllosiphonaceae, Chlorophyta) commonly occurs in Arisarum leaves in coastal Mediterranean areas of the Iberian Peninsula and Balearic islands. The genus Phyllosiphon was first considered to be a member of the Xanthophyceae but was later transferred to the chlorophytes. However, there are few data about its morphology, ultrastructure, ecology or phylogenetic affinities. In this paper we describe the morphology of Phyllosiphon, as studied in field material and in culture; the fine structure, analysed by transmission electron microscopy; and phylogenetic relationships, inferred from DNA sequences. The siphonous filaments were seen to divide and penetrate leaf tissues. The cytoplasm divided into spherical or subspherical sporocysts producing autospores inside. Cytoplasmic remains could be observed between autospores or on their cell walls. Phylogenetic analysis of 18S rDNA and 16S rDNA sequences showed that the closest relatives of Phyllosiphon are subaerial strains of Heterochlorella, Heveochlorella and Kalinella, demonstrating that Phyllosiphon should be transferred to Trebouxiophyceae. An evolution from unicells to a siphonous thallus, and from aerophytic to endophytic and parasitic habits, is proposed for Trebouxiophyceae.