“…Middle Devonian lycopsids evolved many key characteristics, e.g., the heterospory [ 4 , 5 ] and the tree habit [ 6 , 7 ]. Heterosporous lycopsids blossomed in the Late Devonian (e.g., [ 8 , 9 , 10 , 11 , 12 , 13 , 14 , 15 , 16 ]), which is regarded as a major step in the full colonization of land surfaces by plants [ 17 , 18 ]. In the past 20 years, over ten species of Late Devonian heterosporous lycopsids have been reported or restudied all around the world (e.g., [ 9 , 10 , 11 , 12 , 13 , 14 , 15 , 16 , 19 , 20 , 21 , 22 ]); especially, the South China Plate held eight species with monosporangiate strobili (e.g., [ 9 , 10 , 11 , 12 , 13 , 14 , 15 , 16 ]), suggesting a probable origin center and diversity hotspot of this group.…”