Noeggerathiales are enigmatic plants that existed during Carboniferous and Permian times, ∼323 to 252 Mya. 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-My-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 Ω-shaped vascular bundles. The heterosporous (containing both microspores and megaspores), bisporangiate fertile shoots appear cylindrical and cone-like, but their bilateral vasculature demonstrates that they are complex, three-dimensional sporophylls, representing leaf homologs 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 My, 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 ∼110 My, independently evolving sophisticated cone-like fertile organs from modified leaves.

Birmingham Institute of Forest Research University of Birmingham Edgbaston Birmingham B15 2TT United Kingdom

Center for Excellence in Life and Paleoenvironment Chinese Academy of Sciences Nanjing 210008 China

Centre of Palaeobiodiversity West Bohemian Museum in Plzeň 301 36 Plzeň Czech Republic

College of Paleontology Shenyang Normal University Key Laboratory for Evolution of Past Life in Northeast Asia Ministry of Natural Resources Shenyang 110034 China

Department of Earth and Environmental Science University of Pennsylvania Philadelphia PA 19104 6316

Department of Geology and Atmospheric Science Indiana University Bloomington IN 47405

Department of Palaeobiology and Palaeoecology Institute of Geology v v i Academy of Sciences of the Czech Republic 165 00 Praha 6 Czech Republic

Department of Paleontology Geozentrum University of Vienna 1090 Vienna Austria

Indiana Geological and Water Survey Bloomington IN 47404;

School of Geography Earth and Environmental Sciences University of Birmingham Edgbaston Birmingham B15 2TT United Kingdom

State Key Laboratory of Palaeobiology and Stratigraphy Nanjing Institute of Geology and Palaeontology Chinese Academy of Sciences Nanjing 210008 China;

State Key Laboratory of Systematic and Evolutionary Botany Institute of Botany Chinese Academy of Sciences Xiangshan Beijing 100093 China

University of Chinese Academy of Sciences Shijingshan District Beijing 100049 China

Proc Natl Acad Sci U S A. 2021 Apr 27;118(17):e2104256118. doi: 10.1073/pnas.2104256118. PubMed

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