A new early permian fruit, Dengfengfructus maxima gen. et sp. nov., supports the pre-cretaceous origin of angiosperms

BACKGROUND: Angiosperms are the most important plant group for humans in the current earth’s ecosystem. Although angiosperms are clearly defined by enclosed seeds/ovules, the origin and early history of angiosperms remain elusive and controversial. An over-60-year-old model in botany hypothesizes that angiosperms cannot be older than the Cretaceous. However, this hypothesis is now facing new challenges from fossil evidence and molecular estimates. Fossil materials from the pre-Cretaceous strata would provide new evidence in resolving this academic debate. In recent years, a renewed wave of interest in Permian fossil plants in Cathaysian flora in Henan, China has been rekindled by the recent discovery of traces of angiosperms in the Permian. METHODS: During a recent field excursion in May 2025, we collected a new fossil organ from an outcrop of the Lower Shihezi (formerly Shihhotse) Formation (lower Permian) of Dengfeng, Henan, China. Observations with incident-light microscopic and SEM revealed the morphology and anatomy of this fossil organ, which lay the foundation for our treatment of the fossil organ. RESULTS: The fossil organ is a highly flattened compression preserved with cellular details, and its morphology and anatomy allow us to interpret it as a large angiosperm fruit named Dengfengfructus maxima gen. et sp. nov. The seed enclosed by the pericarp has a peripheral three-layered testa, which distinguishes the seed itself from a nucellus or other seed content. The good preservation allows the cellular details in the testa and seed content to be revealed. This organization distinguishes Dengfengfructus from all known gymnosperm seeds and makes it comparable to an angiosperm fruit. Our observations support Dengfengfructus is a large fruit with a thick pericarp. CONCLUSIONS: This new fossil organ apparently updates and enhances the current understanding of angiosperms and their diversity in the Permian. The history of angiosperms can thus be pushed back to the early Permian (Palaeozoic). Our discovery, together with the estimation of molecular clocks, challenges the current hypothesis that the angiosperms didn’t appear until the Cretaceous.