Palaeoenvironments of the Earliest (Middle Devonian) Tetrapod Trackways from the Holy Cross Mountains, Poland;Locomotion in a Terrestrial Setting?

Abstract: The palaeoenvironment from which early tetrapods emerged is crucial to comprehend in order to understand the mechanisms that drove and allowed the terrestrialization of vertebrates; one of the most important evolutionary ‘events’ in the history of animals. Nevertheless, much of the terrestrialization is shrouded in obscurity, inter alia, due to scarcity of early tetrapods in the fossil record. Each new discovery of anything linked to vertebrate terrestrialization is therefore of great importance. Here, I present new detailed analyses of the palaeoenvironmental conditions at the time of formation of the earliest (early Middle Devonian) tetrapod trackways found in the Zachełmie Quarry in the Holy Cross Mountains of southern Poland. The trackways are found in three horizons in a clay-rich dolomitic succession represented by the Lower Complex of the Wojciechowice Formation. This Lower Complex is composed of short shallowing upward sequences that often terminate with desiccation cracks and/or paleosols. Vertically fluctuating δ18O values in the complex suggest multiple episodes of closed hydrological systems. A model of ephemeral to perennial lakes in a terrestrial setting is therefore proposed. Such environment is in concordance with evidence of scarce bioturbation and a flourish of microbial communities that, in contrast to a normal marine setting, most likely represent an ecologically stressed ecosystem. Furthermore: non-marine rare earth element (REE) signals, desiccation events, fossils of green algae (charophytes), paleosol development, low energy cyclic deposits and general lack of marine taxa in the body and trace fossil records in this complex firmly establishes the interpretation of palustrine carbonates formed in a lacustrine-like setting. However, in the lower part of the complex, some highly fractioned marine fossils occur. These are suggested to have been transported during wash-up events which temporally places the setting in marine proximity. Seasons of monsoonal rainfall resulted in erosion and influx of detrital grains which is evidenced by fine planar lamination (of seasonal cyclicity) in most of the sediments and occasional occurrence of blackened clasts (and rain-drop imprints).Thus, none of the pre-existing palaeoenvironmental hypotheses of tetrapod emergence are in full concordance with the data from the Zachełmie Quarry. Instead, a new palaeoenvironment is proposed for the earliest tetrapods: schizohaline ephermal to perennial lakes with periodic desiccation. This infers that already in the early Middle Devonian, tetrapods had conquered the terrestrial realm and were perhaps already capable of terrestrial locomotion over quite substantial distances.