A Three-dimensional Geometric Morphometric Study of Miocene Ape Lumbar Vertebrae, with Implications for Hominoid Locomotor Evolution

Miocene apes represent snapshots in time of key transitions in hominoid evolution. While all extant apes are adapted to orthograde posture and suspensory behavior, many Miocene apes demonstrate evidence for pronogrady and habitual arboreal quadrupedalism or present 'mosaic' morphologies suggestive of locomotion and posture unlike any extant catarrhine. Here, we use three-dimensional geometric morphometrics to study penultimate lumbar vertebrae of extant anthropoids and those of three well-preserved Miocene apes: Ekembo nyanzae (KNM-MW 13142), Morotopithecus bishopi (UMP 67-28), and Pierolapithecus catalaunicus (IPS 21350-64), which have been interpreted as a pronograde arboreal quadruped, an orthograde suspensory or vertical climbing ape, and an orthograde vertical climber that was not adapted to suspensory behavior, respectively. Our results show that E. nyanzae shares three-dimensional shape space with terrestrial papionins, whereas M. bishopi and P. catalaunicus fall within overlapping morphospace shared by Ateles and hylobatids. Morotopithecus bishopi and P. catalaunicus share with hylobatids and brachiating atelids (Ateles and Brachyteles) well-established features such as dorsal lumbar transverse (costal) processes and a newly identified feature in this study, the presence of a convex pillar along the pars interarticularis that forms the lateral borders of the laminae. The latter feature is also shared with E. nyanzae. Together with their large body size estimates, we interpret these results to indicate that E. nyanzae was primarily a pronograde quadruped that may have been semiterrestrial rather than strictly arboreal, while M. bishopi and P. catalaunicus were adapted to both orthogrady and forelimb-dominated climbing and suspension.