Fertile Woodwardia from the middle Eocene of South China and its implications for palaeogeography and palaeoclimate

The genus Woodwardia, which together with the genera Anchistea and Lorinseria comprise the subfamily Woodwardioideae of Blechnaceae, has a disjunct distribution across Central and North America, Europe and the temperate to tropical areas of Asia. Fossil records of Woodwardia occur throughout the Paleogene and Neogene of North America, Europe and Asia. However, well-preserved fertile pinna fossils of this genus have not yet been reported in South China. In this paper, a new species, W. changchangensis Naugolnykh et Song, sp. nov. is described from the middle Eocene of the Changchang Basin, Hainan Island, South China. Macromorphological and micromorphological features of the fertile pinna show a straight pinna rachis, alternate, subtriangular pinnules, acute pinnule apices, almost entire or slightly undulate pinnule margins, long-ovoid sori, stalked sporangia and spores with wing-like folds on the surface, which are characterised in detail. Overall, the present fossil is most similar to the extant species Woodwardia japonica, which mostly grows in warm and moist environments. The discovery of this new species from the Changchang Basin of Hainan Island indicates that this genus has been distributed in the low-latitude tropical regions of South China from as early as the middle Eocene. Based on this find, and previous studies of other ferns from the same site, we infer that the climate of the Palaeo-Hainan landscape during deposition of the Changchang Formation was warm and humid, similar to conditions prevailing today across this region.