Geosiphon pyriforme, a fungus forming endocytobiosis with Nostoc (cyanobacteria), is an ancestral member of the Glomales: evidence by SSU rRNA analysis

Abstract

Geosiphon pyriforme inhabiting the surface of humid soils represents the only known example of endocytobiosis between a fungus (Zygomycotina; macrosymbiont) and cyanobacteria (Nostoc; endosymbiont). In order to elucidate the taxonomical and evolutionary relationship of Geosiphon pyriforme to fungi forming arbuscular mycorrhiza (AM fungi), the small-subunit (SSU) ribosomal RNA genes of Geosiphon pyriforme and Glomus versiforme (Glomales; a typical AM fungus) were analyzed and aligned with SSU rRNA sequences of several Basidiomycetes, Ascomycetes, Chytridiomycetes, and Zygomycetes, together with all AM-fungal (Glomales) sequences published yet. The distinct group of the order Glomales, which includes Geosiphon, does not form a clade with any other group of Zygomycetes. Within the Glomales, two main lineages exist. One includes the families Gigasporaceae and Acaulosporaceae; the other one is represented by the genus Glomus, the members of which are very divergent. Glomus etunicatum and Geosiphon pyriforme both form independent lineages ancestral to the Glomales. The data provided by the present paper confirm clearly that Geosiphon represents a fungus belonging to the Glomales. The question remains still open as to whether or not Geosiphon is to be placed within or outside the genus Glomus, since this genus is probably polyphyletic and not well defined yet. Geosiphon shows the ability of a Glomus-like fungus to form a "primitive" symbiosis with a unicellular photoautotrophic organism, in this case a cyanobacterium, leading to the conclusion that a hypothetical association of a Glomus-like fungus with a green alga as a step during the evolution of the land plants appears probable.