Palaeoecology of Middle Triassic tetrapod ichnoassociations (middle Muschelkalk, NE Iberian Peninsula) and their implications for palaeobiogeography in the western Tethys region

Tetrapod ichnology is a powerful tool to reconstruct the faunal composition of Middle Triassic ecosystems. However, reconstructions based on a single palaeoenvironment provide an incomplete and impoverished picture of the actual palaeodiversity. In this paper, we analyse Middle Triassic tetrapod ichnoassociations from the detrital Muschelkalk facies of the Catalan Basin of northeast Spain, ranging from terrestrial to coastal settings. We identified two main tetrapod ichnoassociations, preserved in two different palaeoenvironments, comprising the following ichnogenera and morphotypes: Procolophonichnium, Chelonipus, Rhynchosauroides, Rotodactylus, Chirotherium, Isochirotherium, Sphingopus, and indeterminate chirotheriids. We also statistically analyse a database of all known Middle Triassic tetrapod footprint localities worldwide; this database includes, for each track locality, the precise age, the palaeoenvironment and the presence/absence of ichnotaxa. Our results on the composition of ichnofauna within the palaeoenvironments of the Catalan Basin are integrated into this database. This approach allows us to revisit the palaeoenvironmental bias linked to the marine transgression that affected the Western Tethys region. Tetrapod ichnoassociations reveal the following palaeoenvironmental patterns: (1) in coastal settings, ichnoassociations are Rhynchosauroides-dominated and diversity is relatively low; (2) in terrestrial settings and those with less marine influences, ichnoassociations are non-Rhynchosauroides-dominated, usually characterised by more abundant chirotheriid tracks and, generally, a higher track diversity. The correlation between tetrapod ichnoassociations and sedimentary facies reveals how palaeoenvironmental constraints influenced faunal assemblages, especially those of the Middle Triassic of the Western Tethys region. Ichnoassociations allow the ecological response of tetrapod faunas to the environmental changes to be inferred for this critical time interval. Marine transgressions strongly influenced tetrapod ecosystems: environmental conditions were key for the faunal recovery in the aftermath of the end-Permian extinction, with the settlement of the so-called modern faunas and the rise of the dinosaur lineage.