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.

在贝尔吉施格拉德巴赫（德国莱茵地块）沙区的发掘中发现了上弗拉斯阶“原角藻”变异带（上泥盆纪 IK，两个 Kellwasser 层之间的间隔）的丰富的菊花类动物群。先前未知的组合包括6个tornoceratid属20种，其中包括7个新种（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 aequilobumsp. nov.) 和开放命名法中描述的分类群。有 5 个相关的 gephuroceratid 属与 9 个物种。在统计指数的支持下，Sand 出人意料的高属和物种多样性与其他同时期的菊花动物群相比是前所未有的。它突出了目前在全球范围内对顶级弗拉斯尼亚菊石动物群的零碎知识。凤蝶，欧洲和北非缺氧和富含有机物的 Kellwasser 相的主要龙角类动物，在 Sand 完全消失了。当地的组合结构进行了统计分析，并根据古生态学进行了解释。一种新型“Housean Pits”的出现，可能是由寄生虫引起的，在5属9种中发现，主要是在龙角动物中。沙地动物群表明，在Lower Kellwasser 事件之后，龙角动物的物种起源率仍然很高。