Transition between terrestrial-submerged walking and swimming revealed by Early Permian amphibian trackways and a new proposal for the nomenclature of compound trace fossils

Exceptionally preserved Early Permian tetrapod trackways from the Orobic Basin (Central–Western Southern Alps) offer a unique opportunity to investigate in detail locomotion in fossil vertebrates that lived on continental European landmasses. Herein are reported the results of a study on several tetrapod trackways that display a large variety of behavioral, gait and substrate related extramorphologies. They clearly document the transition from terrestrial–underwater walking to swimming and are assigned to the compound ichnotaxon Batrachichnus C Lunichnium. The use of the “C” symbol is here introduced for the first time as nomenclatural indication of a Com- pound trace. Producers were probably small-sized temnospondyl or lepospondyl (microsaurs) amphibians. Com- parisons with living urodelan anatomy and mechanics provide evidence for conservatism of locomotor mechanics in evolutionary history among amphibians. The derived model for locomotor kinematics in Early Permian amphibians provides a reference for interpreting transitional land-to-water trackways. The shift from walking to swimming behavior in early tetrapods, as in extant urodelan amphibians, is described as a complex balance between different dynamics.