Phylogenetic comparative morphological analysis of fern spores in subfamily Pteridoideae (Pteridaceae, Pteridophyta)

Scanning electron microscopy (SEM) was used to characterize spore morphology of 24 taxa of the subfamily Pteridoideae C. Chr. ex Crabbe, Jermy & Mickel of the family Pteridaceae E.D.M. Kirchn. The family is considered to be one of the most taxonomically confusing families due to its high level of polymorphism. The standardized data on spore morphology of the subfamily Pteridoideae were projected onto the final phylogenetic tree in the Mesquite program. This approach made it possible to carry out a comprehensive interdisciplinary analysis of the evolution of spore morphology characters of the subfamily Pteridoideae, as well as to assess the relationships in the family Pteridaceae. The equatorial ridge (cingulum, "flange") has been proven as one of the key spore morphology features, which confirms the close relationship of "Onychium clade" with Pteris. The species-specific characters of the subfamily are fold and tubercle along laesura, equatorial ridge on proximal and distal side, tubercle, and folds on proximal and distal side. The knowledge will help to solve the problems of taxonomy in the family Preridaceae and to supplement the information on the natural classification of the subfamily Pteridoideae.