A morphometric analysis of Tobleria bicuspis, a Voltziales seed cone from the early Permian Jambi palaeoflora, Sumatra (Indonesia)

Tobleria bicuspis, a coniferophyte seed cone, is described from the Jambi Palaeoflora, Sumatra of Asselian (early Permian) age. A morphometric analysis based on cones, paired fertile units, and fertile and sterile scales, demonstrates their close relationship. Small paired fertile units occur mainly in cones. Medium-sized paired fertile units occur mainly on scales. And large paired fertile units are mainly dispersed. The cones are considered female and the paired fertile units are considered to represent the seeds. The cones are composed of helicoidal, bilaterally symmetrical and deeply incised scales with paired seeds. A comparison can be made with the Voltziales female taxon Schizolepis from the Triassic and Jurassic. Tobleria is regarded as having a voltzian Voltziales affinity and dates from approximately 16 to 26 million years before any other such cones.