Hindlimb functional morphology and locomotor biomechanics of the small Late Triassic pseudosuchian reptile Gracilisuchus stipanicicorum (Archosauria: Gracilisuchidae)

Abstract

Gracilisuchus stipanicorum was a pseudosuchian archosaur from the Late Triassic period in Argentina. Because it was small-bodied with relatively long, slender limbs, traits that are potentially ancestral for archosaurs, such as its locomotor functions, are important to estimate. It has been illustrated as a quadruped with plantigrade autopodia, and probably with an ‘erect’ or ‘semi-erect’ stance, because it is a suchian archosaur, but there has been no deep analysis of these reconstructions. Here, we detail our reconstruction of a three-dimensional digital skeleton of Gracilisuchus from scans of the bones of four main specimens, including the holotype. In this procedure, we found hitherto unrecognised elements of the manus (metacarpals) and incorporated them in our model of the whole organism. We added estimated hindlimb musculature and body segment mass properties to form a musculoskeletal model. This model allowed us to address three key questions: Was it quadrupedal or bipedal; plantigrade or digitigrade; and more sprawling or more erect? Furthermore, we examine how its hindlimb muscle moment arms compare to those of three other small-bodied Triassic archosauriforms and an extant juvenile Nile crocodile in order to assess the diversity and potential evolutionary polarity of these traits. Our analyses of the model support the inferences that Gracilisuchus was quadrupedal (but facultative bipedalism cannot be ruled out) and plantigrade, and not strongly sprawling, but probably not strongly erect hindlimbs; although terming this posture ‘semi-erect’ would be an oversimplification. Gracilisuchus, as modelled here, seems to roughly be a reasonable approximation of the ancestral state of the archosaurian locomotor system. Our synthesis of numerous lines of evidence, from qualitative functional morphology to whole-body centre of mass and muscle moment arms, forms a new reconstruction of Gracilisuchus that future analyses can build on, both biomechanically and comparatively, in order to better understand archosauriform locomotor evolution.