Living crinoids are exclusively passive suspension feeders and benthic as adults. However, in the past they adapted to a broad range of ecological niches. For instance, the stratigraphically important middle Paleozoic scyphocrinoids are hypothesized to have been planktonic, employing their inferred gas-filled globular, chambered structure at the distal end of the stem, the so-called lobolith, as a buoyancy device with the crinoid calyx suspended below it. Here, we evaluate this hypothesis using evidence from skeletal micromorphology and theoretical biomechanical modeling. Lobolith walls are typically composed of ossicles, which are exclusively composed of constructional labyrinthic stereom. In plates from the distal side of the lobolith, this stereom extends into microperforate stereom layer, forming wavy ridges and spines. No microscale adaptations for preventing gas leaks and/or ingress of water (such as internal and external imperforate stereom layers) are known. Furthermore, theoretical calculations suggest that the scyphocrinoid tow-net mode of feeding would have resulted in small relative velocities between the towed filter and the ambient water, thus making it an ineffective passive filter feeder. We suggest that the lobolith of these crinoids acted as a modified holdfast rather than as a floating buoy. Its globular shape and distally positioned microspines served as adaptations for living in unconsolidated sediments, analogous to iceberg- and snowshoe-like strategies used by some mollusks and brachiopods. Like modern isocrinids, scyphocrinoids could have maintained an upright feeding posture by extending the distal portion of the stalk along the bottom. In this recumbent posture, the distal part of the stalk with the lobolith might have functioned as a drag anchor. As a consequence of the ~3-m-long stem, even with this posture, the benthic scyphocrinoids could have risen to the highest epifaunal tier in the Paleozoic.

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