Gregarious behaviour among non-avian theropods inferred from trackways: A case study from the Cretaceous (Cenomanian) Candeleros Formation of Patagonia, Argentina

Gregarious behaviour among dinosaurs has been inferred from several lines of evidence (monospecific bonebeds, skeletal morphology, phylogenetic inferences, comparison with modern ecosystems and parallel trackways with particular characteristics), but is relatively poorly documented for non-avian theropods. Here, we report five parallel theropod trackways of large track size (average length of 28 cm) in the Cretaceous (Cenomanian) Candeleros Formation from northwestern Patagonia, Argentina. The tracks are provisionally assigned to aff. Asianopodus pulvinicalx and considering the autopod morphology of the theropod taxa documented in the Candeleros Formation, abelisaurid theropods are suggested as trackmakers. The trackways possess similar stride, speed estimation, direction and preservational features, and track show uniform depth and do not overlap. Physical barriers (i.e., large fluvial channels or perennial lake) that could influence the direction of the theropod trackmaker gaits were not recognized in the section. Taking into account these considerations, a gregarious behaviour for the abelisaurid theropod trackmakers is proposed. The tracks are preserved as shallow undertracks in a medium-grained sandstone bed deposited in a floodplain setting. The tracking surface is interpreted as the overlaying layer of muddy-siltstone. The occurrence of swelling clays and microbial mats in the track-bearing level could have contributed to the substrate stabilization and their role in the preservation is discussed.