Bivalves on the move: the interplay of extrinsic and intrinsic factors on the morphology of the trace fossil Protovirgularia

Tide-influenced deposits of the Pennsylvanian Rock Lake Shale Member, Stanton Formation, Missouri and Kansas (U.S.A.), contain well-preserved evidence of the burrowing activities of protobranch bivalves, including locomotion (Protovirgularia) and locomotion-resting (Protovirgularia-Lockeia) trace fossils. Protovirgularia shows three distinct morphotypes, the morphological variability of which was controlled by external factors such as substrate conditions, toponomy, and undertrack deficiency. Extreme morphological modifications of Protovirgularia and/or Lockeia, represented by irregular bilobate structures, probably result from erosion and subsequent partial passive filling of the original biogenic structures. In addition, very well-preserved trace fossils showing intergradation of a fan-shaped cluster of ridges with Protovirgularia-Lockeia structures are interpreted as a compound biogenic structure that represents locomotion, resting, and feeding activities of burrowing protobranch bivalves.