Structurally preserved Zamites bayeri Kvaček from the Coniacian Kashima Formation (Yezo Group) of Hokkaido, Japan

A permineralized frond from the Coniacian Kashima Formation (Yezo Group) of Hokkaido, Japan, is identified as Zamites bayeri based on leaf shape, transverse orientation of stomata, papillate epidermal surface, and hollow subsurface bodies formed only on the abaxial epidermis. The specimen has an articulation at the base of the leaflet from which the leaflet was detached by means of an abscission zone. This finding suggests that Z. bayeri has compound leaves that are functionally equivalent to a twig bearing simple leaves. This functional conversion is often found in extant compound-leaved ferns or cycads growing in infertile conditions; thus Z. bayeri might have lived in a similar habitat. Leaflet articulation is also found in other Zamites species including the type species of the genus, whereas it is unknown in other bennettitalean foliar morphogenera. The rachis of the specimen has a columnar vascular bundle, which is common in bennettitalean foliage.