Plant communities of the geologic past can be reconstructed with high fidelity only if they were preserved in place in an instant in time. Here we report such a flora from an early Permian (ca. 298 Ma) ash-fall tuff in Inner Mongolia, a time interval and area where such information is filling a large gap of knowledge. About 1,000 m 2 of forest growing on peat could be reconstructed based on the actual location of individual plants. Tree ferns formed a lower canopy and either Cordaites, a coniferophyte, or Sigillaria, a lycopsid, were present as taller trees. Noeggerathiales, an enigmatic and extinct spore-bearing plant group of small trees, is represented by three species that have been found as nearly complete specimens and are presented in reconstructions in their plant community. Landscape heterogenity is apparent, including one site where Noeggerathiales are dominant. This peat-forming flora is also taxonomically distinct from those growing on clastic soils in the same area and during the same time interval. This Permian flora demonstrates both similarities and differences to floras of the same age in Europe and North America and confirms the distinct character of the Cathaysian floral realm. Therefore, this flora will serve as a baseline for the study of other fossil floras in East Asia and the early Permian globally that will be needed for a better understanding of paleoclimate evolution through time.coal-swamp plant community | plant paleoecology | volcanic ash-fall tuff | Wuda

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.

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A new species Lepidopteris baodensis sp. nov. belonging to the family Peltaspermaceae and represented by two ultimate pinnae in the collection under study, was recently discovered at the Baijiagou of Baode, Shanxi, China, from the Upper Permian Sunjiagou Formation. The lower surface of the ultimate rachis, the midrib and secondary veins is covered with triangular, trapezoid, rounded, or ligulate subepidermal swellings, which show different natures from intercalary pinnules. Lepidopteris is one of typical elements of the Late Permian Euramerican flora. Since Schimper erected the genus Lepidopteris in 1869, the entire epidermal structure of subepidermal swellings had been unclear. The new species L. baodensis clearly showing the distinguished epidermal structure of subepidermal swellings, not only enlarges and supplements our knowledge in biology and taxonomy of Lepidopteris as well as the Upper Permian stratigraphy of China, but also provides an opportunity to understand the relationship between Euramerican floras and Cathaysian floras in paleoclimatic, paleoenvironmental and paleogeographic context.