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, ∼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.

Permian vegetational Pompeii from Inner Mongolia and its implications for landscape paleoecology and paleobiogeography of Cathaysia.

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.

Nystroemiaceae, a new family of Permian gymnosperms from China with an unusual combination of features.

Nystroemiaceae is proposed as a new family of gymnosperms from the Permian of Cathaysia that adds to the diversity of gymnosperms known from this critical time in seed plant evolution. This family is characterized by bifurcating and highly branched pinnate ovuliferous organs bearing bicornute ovules (seeds) and entire leaves with anastomozing veins that are born on complex and modern-looking branching systems with clear axillary branching. The reconstruction is based on numerous large specimens from two localities in North China, in which the different plant parts are attached to each other. The ovulate structures show some apparently plesiomorphic (primitive) character states more typical of early seed plants, whereas the leaves and branches show the clearly apomorphic (derived) character states of broad-leaved gymnosperms.

Paratingia wudensis sp. nov., a whole noeggerathialean plant preserved in an earliest Permian air fall tuff in Inner Mongolia, China.

Noeggerathiales are a little known group of Carboniferous and Permian plants of uncertain systematic position that have been variously considered to be ferns, sphenopsids, progymnosperms, or a separate group. These heterosporous plants carry adaxial sporangia on leaf-like or disk-shaped sporophylls that form cones. Leaves are pinnate with a rather stiff appearance, and pinnules can be attached in either two or four rows. In the present report, we present the top of a noeggerathialean plant with leaves and strobili attached, Paratingia wudensis Wang, Pfefferkorn et Bek sp. nov., from an earliest Permian volcanic ash fall tuff in Inner Mongolia. The excellent preservation allows the reconstruction of the whole plant, the complex three-dimensional leaves with anisophyllous pinnules, the heterosporous strobili, and the spores in situ. The homology of leaves and strobili can be elucidated and contributes to an understanding of the debated taxonomic position of Noeggerathiales. The "anisophyllous" leaves carry pinnules arranged in four rows. The strobili are bisporangiate and have disk-shaped sporophylls, each with one ring of 10-14 adaxial sporangia around the strobilus axis. Megaspores have an equatorial bulge. This new species expands the known diversity of Noeggerathiales. It grew in a peat-forming forest, thus changing earlier interpretations of the growth of noeggerathialean plants with anisophyllous pinnules.

A Late Devonian isoetalean lycopsid, Otzinachsonia Beerboweri, gen. et sp. nov., from north-central Pennsylvania, USA.

Compressions and impressions of an isoetalean lycopsid, comprising lower portions of stems, lobed bases, attached rootlets, and rounded rootlet scars, discovered in Late Devonian (Famennian) rocks of Clinton County, north-central Pennsylvania, Appalachian Basin, USA, are here described as Otzinachsonia beerboweri, gen. et sp. nov. These specimens demonstrate unequivocally the existence of the isoetalean lobe-and-furrow rhizomorphic growth pattern as early as the Late Devonian. They were found in an Archaeopteris- and Rhacophyton-dominated flora at Red Hill, an outcrop of the Duncannon Member of the Catskill Formation. The fossils were found in a dark-gray to greenish-gray lenticular siltstone layer that has an average thickness of 1.0 m. This deposit is interpreted as a floodplain pond. The low-energy nature of the deposit and the fine preservation of the intact rootlets of the specimens imply little or no transport. The plants were probably growing along the edge of the floodplain pond with their lower portions submerged for at least part of the year.