Ancient noeggerathialean reveals the seed plant sister group diversified alongside the primary seed plant radiation

There were two heterosporous lignophyte lineages of which only one, the seed plants, survived the Permian-Triassic mass extinction. Based on exceptionally complete fossil trees from a 300-My-old volcanic ash, the enigmatic Noeggerathiales are now recognized as belonging to the other lineage. They diversified alongside the primary seed plant radiation and constitute seed plants' closest relatives. Noeggerathiales are reconstructed as members of a plexus of free-sporing woody plants called progymnosperms, extending their age range by 60 My. Following the origin of seed plants, progymnosperms were previously thought to have become gradually less abundant before dying out in Carboniferous. We show they diversified and evolved complex morphologies including cone-like structures from modified leaves before going extinct at the Permian-Triassic extinction.