Complex behavioural patterns and ethological analysis of the trace fossil Zoophycos: evidence from the Lower Devonian of South China

The trace fossil Zoophycos is abundant in the shallow-marine deposits (tempestites) of the Lower Devonian (Emsian) Yangmaba Formation in Ganxi of Sichuan, South China. It often occurs as part of complex trace fossils that comprise different integrated elements: scratch traces, simple to complex spreiten structures with marginal tubes (Zoophycos) and vertical tunnels. The complex Zoophycos burrows consist of spreiten with a marginal tube, preserved as convex hyporeliefs on the sole of an erosion surface. The exquisite, complex spreiten are interpreted to have been formed by deposit-feeding behaviour, where the animal constructed the trace upwards without leaving faeces in the spreiten. The width of the marginal tube in different whorls is almost constant. The scratches are observed on the wall of the marginal tubes. The Zoophycos intergrades with Spongeliomorpha and Chondrites and was later cut by vertical shafts. All these features together indicate that the Zoophycos-maker might have been a vermiform polychaete instead of a predator such as a decapod crustacean (Spongeliomorpha producer). Based on stratigraphical and ichnological features, the complex trace fossils resulted from the complex activity of different opportunistic organisms (r-strategist) that quickly occupied and thrived within the quiet, nutrient-rich environment after storm events.