by Sonja Kistenich, Einar Timdal, Mika Bendiksby, Stefan Ekman
The Ramalinaceae is the fourth largest family of lichenized ascomycetes with 42 genera and 913 species exhibiting considerable morphological variation. Historically, generic boundaries in the Ramalinaceae were primarily based on morphological characters. However, molecular systematic investigations of subgroups revealed that current taxonomy is at odds with evolutionary relationships. Tropical members of the family remain particularly understudied, including the large genus Phyllopsora. We have generated and collected multilocus sequence data (mtSSU, nrITS, nrLSU, RPB1, RPB2) for 149 species associated with the Ramalinaceae and present the first comprehensive molecular phylogeny of the family. We used ancestral state reconstructions on our molecular family phylogeny to trace the evolution of character states. Our results indicate that the Ramalinaceae have arisen from an ancestor with long, multiseptate ascospores living in humid temperate forests, and that the phyllopsoroid growth form has evolved multiple times within the family. Based on our results using integrative taxonomy, we discuss sister-relations and taxon-delimitation within five well-supported clades: The Bacidia , Biatora-, Ramalina-, Rolfidium-, and Toninia-groups. We reduce six genera into synonymy and make 50 new nomenclatural combinations. The genera Bacidia, Phyllopsora, Physcidia and Toninia are polyphyletic and herein split into segregates. We describe the two genera Bellicidia and Parallopsora and resurrect the genera Bibbya, Kiliasia, Thalloidima and Sporacestra. According to our new circumscription, the family Ramalinaceae now comprises 38 genera.