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

Noeggerathiales are enigmatic plants that existed during Carboniferous and Permian times, approximately 323–252 million years ago. Although their morphology, diversity and distribution are well known, their systematic affinity remained enigmatic because their anatomy was unknown. Here we report from a 298 million-year-old volcanic ash deposit, an in-situ, complete, anatomically preserved noeggerathialean. The plant resolves the group’s affinity and places it in a key evolutionary position within the seed plant sister group. Paratingia wuhaia sp. nov. is a small tree producing gymnospermous wood with a crown of pinnate, compound megaphyllous leaves and fertile shoots each with omega-shaped vascular bundles. The heterosporous (containing both micro- and megaspores), bisporangiate fertile shoots appear cylindrical and cone-like, but their bilateral vasculature demonstrates that they are complex, three-dimensional sporophylls, representing leaf homologues that are unique to Noeggerathiales. The combination of heterospory and gymnospermous wood confirms that Paratingia, and thus the Noeggerathiales, are progymnosperms. Progymnosperms constitute the seed plant stem group, and Paratingia extends their range 60 million years, to the end of the Permian. Cladistic analysis resolves the position of the Noeggerathiales as the most derived members of a heterosporous progymnosperm clade that are the seed plant sister group, altering our understanding of the relationships within the seed plant stem lineage and the transition from pteridophytic spore-based reproduction to the seed. Permian Noeggerathiales show that the heterosporous progymnosperm sister group to seed plants diversified alongside the primary radiation of seed plants for approximately 110 million years, independently evolving sophisticated cone-like fertile organs from modified leaves.