Upper Frasnian Tornoceratidae (Ammonoidea) from the Sand Formation (Bergisch Gladbach-Paffrath Syncline, Rhenish Massif)

Excavations in the Sand district of Bergisch Gladbach (Rhenish Massif, Germany) yielded a rich ammonoid fauna of the upper Frasnian “Archoceras” varicosum Zone (Upper Devonian I-K, interval between the two Kellwasser levels). The previously unknown assemblages include six tornoceratid genera with 20 species, including seven new species (Aulatornoceras steinhauseni sp. nov., Aul. frenklerae sp. nov., Aul. ventrosulcatum sp. nov., Crassotornoceras nudum sp. nov., Cr. hetzeneggeri sp. nov., Retrotornoceras juxi sp. nov., Tornoceras aequilobum sp. nov.) and taxa described in open nomenclature. There are five associated gephuroceratid genera with nine species. The unexpectedly high genus- and species-level diversity at Sand, supported by statistical indices, is unprecedented compared to other contemporaneous ammonoid faunas. It highlights the currently fragmentary knowledge of top-Frasnian ammonoid faunas on a global scale. Phoenixites frechi, the dominant tornoceratid of hypoxic and organic-rich Kellwasser facies of Europe and North Africa, is completely missing at Sand. The local assemblage structure is analyzed statistically and interpreted in terms of palaeoecology. The occurrence of a new type of “Housean Pits”, probably caused by parasitism, is found in nine species of five genera, mostly in tornoceratids. The Sand fauna indicates that the species origination rate in tornoceratids remained high after the Lower Kellwasser Event.