First reliable Miocene fossil winged fruits record of Engelhardia in Asia through anatomical investigation

Fossil genera with similar features to the winged fruits of the living Engelhardia Lesch. ex Blume (e.g., Palaeocarya G. Saporta) have been widely reported in Cenozoic fossil floras of the Northern Hemisphere. However, fossil winged fruits of Engelhardia with detailed anatomical structures have only been found in the upper Eocene of North America. This study reports the first Engelhardia fossil winged fruits with detailed anatomical structures in East Asia from the Miocene Erzitang Formation of Guangxi, South China. The anatomical and morphological features of the new fossils, including the unique structure of secondary septa, clearly distinguish them from other fossil genera and show unambiguously their attribution to the genus Engelhardia. This discovery suggests that Engelhardia had reached its modern distribution during the Miocene and the climate of the Guiping Basin in Guangxi during the Miocene was similar to that of present-day tropical and subtropical regions in Asia.