Periostracum in Cambrian helcionelloid and rostroconch molluscs: comparison to modern taxa

Abstract

The periostracum, a non-calcified organic layer that envelops the mineralized shell layer (‘ostracum’) of molluscs, acts as an initial framework for the calcified ‘ostracum.’ Despite its significance in biomineralization in molluscs, fossil examples of the periostracum are limited in the geological record, especially from the Palaeozoic era, due to its susceptibility to post-mortem decay. In this study, we describe radial threads found on the outer surface of Cambrian molluscs, including helcionelloids and rostroconchs, which we interpret as fossilized periostracal structures. These radial threads are observed in 13 species from five valid genera, exhibiting a widespread palaeogeographical distribution encompassing Gondwana (Australia), Siberia, western Laurentia (Utah), and eastern Laurentia (Greenland), and spanning Cambrian Stage 4？Drumian deposits. These radial threads always cover the outermost surface of shells, dividing it into regular intervals, and develop through incremental growth. They originate from the apical area of helcionelloids or the umbo of rostroconchs. Detailed examination has revealed that the radial threads initiate at the apical region, but may be absent from the protoconch (embryonic shell). The radial threads are most likely preserved by mineral encrustation on the periostracum. Similar, but not homologous, radial threads are found in other fossil and modern molluscan taxa, such as the adhesive radial lines of anomalodesmatan bivalves or the periostracal ridges of viviparid gastropods.