Hyolithid-like hyoliths without helens from the early Cambrian of South China, and their implications for the evolution of hyoliths

Abstract

Background: A small hyolith, with a triangular operculum and a conical-pyramidal conch with a sharp apex, originally documented as Ambrolinevitus ventricosus, is revised based on new material from the Chengjiang biota. The operculum of ‘Ambrolinevitus’ ventricosus displays strong morphological similarities with the operculum of Paramicrocornus from the Shuijingtuo Formation (Cambrian Series 2), indicating that the species should be reassigned to Paramicrocornus. Results: Based on the unusual morphology of Paramicrocornus, we herein propose a new family Paramicrocornidae fam. nov. A cladistic analysis of Cambrian and Ordovician hyoliths clearly delineates hyolithids as a monophyletic group which evolved from the paraphyletic orthothecids in the early Cambrian and with Paramicrocornidae as its closest relative. Conclusions: The phylogenetic analysis, together with the distribution of hyoliths from the Cambrian to the Ordovician, reveals the presumptive evolution model of both the skeleton and soft-part anatomy of hyoliths. The Family Paramicrocornidae plays an intermediate role in hyolith evolution, representing the transitional stage in the evolution from orthothecids to hyolithids.