Origins of the terrestrial flora: A symbiosis with fungi?

Land phototrophs need to exploit both atmosphere (providing gas and light) and substrate (furnishing water and minerals). Yet, their algal ancestors were poorly pre-adapted to such a life at the interface. We review the paleontological evidence that fungal symbioses which can exploit substrate resources, helped adaptation to land constraints. Diverse structures dating back to the Devonian present convincing evidence for lichens, (symbioses between fungi and microscopic algae) but fossils remain scarce, so that early lichen abundance and ecological relevance remain questionable. Several enigmatic but abundant fossils from the Siluro-Devonian, such as Spongiophyton or the giant Prototaxites (Nematophytes), likely represent fungus-algal symbioses, which shaped early terrestrial ecosystems. Yet, these taxa are fully extinct, and do not have clear affinities with extant groups. Finally, terrestrialization of Embryophyta (land plants), which currently dominate land ecosystems, is linked to a symbiosis with Glomeromycetes. Today, these fungi form arbuscular mycorrhizae, which help most Embryophyta to exploit soil, and molecular data combined with paleontological evidence support the idea that this type of association is ancestral. The role of symbiotic Mucoromycetes during terrestrialization is not fully understood and mycorrhizal association diversified later in the evolution of Embryophyta. Fungal-algal symbioses thus recurrently contributed to terrestrialization of phototrophs.