Identifying signatures of the earliest benthic bulldozers in emergent subaerial conditions during the colonization of land by animals.

The colonization of land by animals was a milestone in the history of life. Approximately 100 million years before full terrestrialization, early animals sporadically traversed emergent subaerial substrates, leaving behind trace fossils recording their activities. However, identifying temporarily emergent environments and determining the affinities, motility and subaerial endurance of the trace-makers, and the timing and magnitude of their impacts on marginal-marine environments, are challenging. Here, we used semi-resolved computational fluid dynamics-discrete element method coupling to simulate trace formation on non-cohesive sediments in submerged and emergent subaerial conditions. This revealed instability-induced morphological signatures that allow us to identify the earliest terrestrial trace fossils. Quantitative metrics enable us to infer that the putative earliest terrestrial trace-makers were molluscs, and dimensional analysis suggests that their subaerial excursions could last at least 15 min. These organisms navigated emergent environments from the early Cambrian (stage 2), tens of millions of years earlier than arthropods. This quantitative paradigm provides new insights into the palaeobiology of the earliest subaerial bulldozers and highlights that mollusc-like animals were among the first ecosystem engineers to enter marginal-marine settings. They may thus have contributed to the establishment of marginal-marine biogeochemical cycles, laying the groundwork for subsequent terrestrialization by other animals.