Early euprimates already had a diverse locomotor repertoire: Evidence from ankle bone morphology

The morphological adaptations of euprimates have been linked to their origin and early evolution in an arboreal environment. However, the ancestral and early locomotor repertoire of this group remains contentious. Although some tarsal bones like the astragalus and the calcaneus have been thoroughly studied, the navicular remains poorly studied despite its potential implications for foot mobility. Here, we evaluate early euprimate locomotion by assessing the shape of the navicular-an important component of the midtarsal region of the foot-using three-dimensional geometric morphometrics in relation to quantified locomotor repertoire in a wide data set of extant primates. We also reconstruct the locomotor repertoire of representatives of the major early primate lineages with a novel phylogenetically informed discriminant analysis and characterize the changes that occurred in the navicular during the archaic primate-euprimate transition. To do so, we included in our study an extensive sample of naviculars (36 specimens) belonging to different species of adapiforms, omomyiforms, and plesiadapiforms. Our results indicate that navicular shape embeds a strong functional signal, allowing us to infer the type of locomotion of extinct primates. We demonstrate that early euprimates displayed a diverse locomotor behavior, although they did not reach the level of specialization of some living forms. Finally, we show that the navicular bone experienced substantial reorganization throughout the archaic primate-euprimate transition, supporting the major functional role of the tarsus during early primate evolution. This study demonstrates that navicular shape can be used as a reliable proxy for primate locomotor behavior. In addition, it sheds light on the diverse locomotor behavior of early primates as well as on the archaic primate-euprimate transition, which involved profound morphological changes within the tarsus, including the navicular bone.