A pioneering riparian community dominated by Nystroemiaceae gymnosperms from the Guadalupian (middle Permian) of China

A new fossil plant community is documented from the Upper Shihhotse Formation on the North China Block with an extremely unusual composition and diversity. Zircon U–Pb dating yields an age of 268.3 ± 2.3 Ma during the mid-Roadian to early Wordian stages of the Guadalupian. The community occurs in two beds and is dominated by fossils of the gymnosperm family Nystroemiaceae, which account for up to 99% of specimens recovered and comprise stems, branches, roots, ovuliferous complexes of Nystroemia pectiniformis and N. sp., and leaves of Chiropteris reniformis. The two different species of Nystroemia are distinguished on size and apical features of ovules, with N. pectiniformis having two apical bicornuate horns, whereas N. sp. has apical auricular extensions. Leaves of C. reniformis are megaphyllous and possess a long petiole and net venation. In the Cathaysian flora, different species of Nystroemiaceae are distinguished by features of their ovules and habit, but all bear leaves of C. reniformis which suggests evolutionary modularity and different rates of evolution between the ovules and leaves. Both N. pectiniformis and N. sp. are shown to have been short, uptight trees. Sedimentary evidence shows the community grew in a meandering fluvial setting with repeated river channel changes. Compared with contemporaneously diverse Cathaysian floral communities in the Upper Shihhotse Formation, the vegetation has a very restricted diversity and is a rare occurrence of a gymnosperm-dominated community. The Nystroemiaceae demonstrates an r-selective ecological strategy as pioneering plants adapted to living in unstable habitats and newly opened gaps in the riparian zone.