Putting a crinoid on a stalk: new evidence on the Devonian diplobathrid camerate Monstrocrinus

The diplobathrid camerate crinoid genus Monstrocrinus is morphologically reinterpreted on the basis of new finds from the upper Emsian (Lower Devonian) to lower Eifelian (Middle Devonian) of Germany (Rhenish Slate Mountains, North Rhine-Westphalia, and Rhineland-Palatinate). The most complete, new specimen has a long segment of column projecting outward from the basal concavity, which confirms that the Monstrocrinus crown was attached to a column as an adult. Thus, a free-living life mode of a passively rolling or drifting crinoid is refuted and Monstrocrinus is reinterpreted as an attached, stalked echinoderm. This finding is supported by re-examination of the historical type material. A column attachment with a central pentalobate axial canal was recognized for the first time in the calyx of the holotype of the type species Monstrocrinus securifer. The holotype of Monstrocrinus granosus is far more complete than previously thought. It is a partially disarticulated crown embedded together with an associated longer part of the column. The extraordinary spines on the calyx plates of Monstrocrinus can be divided into eight morphological types and into two superordinate categories: "Category A" is a physical part of the calyx plate, whereas "Category B" is an attachment onto the calyx plate. Taxonomically, M. aliformis from the upper Emsian of Spain is treated herein as a subjective junior synonym of M. securifer.