New insight into the soft anatomy and shell microstructures of early Cambrian orthothecids (Hyolitha)

Hyoliths (hyolithids and orthothecids) were one of the most successful early biomineralizing lophotrochozoans and were a key component of the Cambrian evolutionary fauna. However, the morphology, skeletogenesis and anatomy of earliest members of this enigmatic clade, as well as its relationship with other lophotrochozoan phyla remain contentious. Here, we present a new orthothecid, Longxiantheca mira gen. et sp. nov. preserved as part of secondarily phosphatized small shelly fossil assemblage from the lower Cambrian Xinji Formation of North China. Longxiantheca mira retains some ancestral traits of the clade with an undifferentiated disc-shaped operculum, a simple conical conch with apical septa and a two-layered microstructure of aragonitic fibrous bundles. The operculum interior exhibits impressions of soft tissues, including muscle attachment scars, mantle epithelial cells and a central kidney-shaped platform interpreted as a support structure in association with its presumptive feeding apparatus. The muscular system in orthothecids appears to be similar to that in hyolithids, suggesting a consistent anatomical configuration among the total group of hyoliths. The new finding of shell secreting cells demonstrates a mantle regulating the mode of growth for the operculum. Investigations of shell microstructures support the placement of hyoliths as total group molluscs with an unsettled position within the phylum Mollusca.