Anatomically preserved seeds of Nuphar (Nymphaeaceae) from the Early Eocene of Wutu, Shandong Province, China

Well-preserved seeds from the early Eocene of Wutu, Shandong, China are assigned to the genus Nuphar (Nymphaeaceae) based on morphology and anatomy. The seeds of Nuphar wutuensis sp. nov. are ellipsoidal to ovoid, 4-5 mm long with a clearly visible raphe ridge, and a truncate apex capped by a circular operculum ca. 1 mm in diameter bearing a central micropylar protrusion. These features, along with the testa composed of a uniseriate outer layer of equiaxial pentagonal to hexagonal surface cells and a middle layer 4-6 cells thick composed of thick-walled, periclinally elongate sclereids, correspond to the morphology and anatomy of extant Nuphar and distinguish this fossil species from all other extant and extinct genera of Nymphaeales. These seeds provide the oldest record for the genus in Asia and are supplemented by a similar well-preserved specimen from the Paleocene of North Dakota, USA. These data, together with the prior recognition of Brasenia (Cabombaceae) in the middle Eocene, indicate that the families Nymphaeaceae and Cabombaceae had differentiated by the early Tertiary.