On the presence of Ichniotherium in the Coconino Sandstone (Cisuralian) of the Grand Canyon and remarks on the occupation of deserts by non-amniote tetrapods

Abstract

The colonization of deserts by tetrapods occurred for the first time in the late Paleozoic. In spite of amniotes being the most abundant and diverse taxon in such environments, fossil tracks indicate that anamniotes also inhabited late Paleozoic deserts. In this paper, the presence of the tetrapod-footprint ichnotaxa Ichniotherium sphaerodactylum and cf. Ichniotherium is documented in the eolian Coconino Sandstone (early Permian), based on 16 trackways found at several localities in Arizona, USA. Because there is a strong association between Ichniotherium and different species of diadectomorphs, we also discuss some aspects of the ichnotaxonomy of this ichnogenus. Diadectomorpha is considered the sister taxon of Amniota and, as a consequence, the tracks described in this paper represent the oldest evidence of occupation of deserts by non-amniote tetrapods. The presence of Ichniotherium in this environmental context also sheds light on the paleobiology of Diadectomorpha and, as a result, the emergence of features typically related to Amniota. The ichnofauna of the Coconino Sandstone has been used as a model for the Chelichnus ichnofacies, which supposedly indicates a low-diversity desert fauna. On the other hand, the tracks described here demonstrate that diadectomorphs were also important faunal components of such deserts and suggests that the significance of this ichnofacies should be reconsidered.