Palaeoenvironmental reconstruction of a lower to middle Permian terrestrial composite succession from the Catalan Pyrenees: Implications for the evolution of tetrapod ecosystems in equatorial Pangaea

Tetrapod diversity in Permian terrestrial ecosystems of southwestern Europe is poorly recorded by bone specimens, but it is better represented by an important tetrapod ichnological record that is relevant to our understanding of vertebrate communities in equatorial Pangaea. Herein, two tetrapod ichnoassociations from three new ichnosites, within Cisuralian and Guadalupian volcanosedimentary terrestrial successions (the Lower Red Unit and the Upper Red Unit deposits) of the Castellar de n'Hug sub-basin (Catalan Pyrenees, NE Iberian Peninsula) are presented. Tetrapod ichnology in combination with stratigraphic and facies analyses permit a characterisation of these ecosystems. The sedimentary deposits show an evolution from fluvial meandering systems to playa-lake floodplains, denoting increased aridification and seasonality under a monsoonal regime. The ichnofossil record shows how tetrapod assemblages shifted from a prevalence of non-amniotes and eureptiles to a dominance of therapsids, eureptiles and parareptiles. The reported ichnotaxa are distributed in two ichnoassociations. The first preserves Batrachichnus, Dromopus, Hyloidichnus, Characichnos and an indeterminate tetrapod morphotype, whereas the second is defined by the presence of Dromopus, Hyloidichnus, Brontopus antecursor and Pachypes. The stratigraphically older ichnoassociation, preserved in volcaniclastic and fluvial deposits, presents features of the early Erpetopus biochron, whereas the stratigraphically younger one, preserved in playa-lake deposits, is constrained to the Brontopus sub-biochron. Biostratigraphic and magnetostratigraphic data suggest a late Cisuralian to middle Guadalupian age for the whole succession. The ichnoassemblage highlights a palaeobiogeographical connection between the Pyrenean Basin and other peri-Tethyan basins, which allows us to expand our knowledge about the palaeoecology and palaeobiodiversity of the Cisuralian to Guadalupian terrestrial ecosystems.