Reconstructing articular cartilage in the Australopithecus afarensis hip joint and the need for modeling six degrees of freedom

The postcranial skeleton of Australopithecus afarensis (AL 288-1) exhibits clear adaptations for bipedality, although there is some debate as to the efficiency and frequency of such upright movement. Some researchers argue that AL 288-1 walked with an erect limb like modern humans do, whilst others advocate for a "bent-hip bent-knee" (BHBK) gait, although in recent years the general consensus favors erect bipedalism. To date, no quantitative method has addressed the articulation of the AL 288-1 hip joint, nor its range of motion (ROM) with consideration for joint spacing, used as a proxy for the thickness of the articular cartilage present within the joint spacing which can affect how a joint moves. Here, we employed ROM mapping methods to estimate the joint spacing of AL 288-1's hip joint in comparison to a modern human and chimpanzee. Nine simulations assessed different joint spacing and tested the range of joint congruency (i.e., ranging from a closely packed socket to loosely packed). We further evaluated the sphericity of the femoral head and whether three rotational degrees of freedom (DOFs) sufficiently captures the full ROM or if translational DOFs must be included. With both setups, we found that the AL 288-1 hip was unlikely to be highly congruent (as it is in modern humans) because this would severely restrict hip rotational movement and would severely limit the capability for both bipedality and even arboreal locomotion. Rather, the hip was more cartilaginous than it is in the modern humans, permitting the hip to rotate into positions necessitated by both terrestrial and arboreal movements. Rotational-only simulations found that AL 288-1 was unable to extend the hip like modern humans, forcing the specimen to employ a BHBK style of walking, thus contradicting 40+ years of previous research into the locomotory capabilities of AL 288-1. Therefore, we advocate that differences in the sphericity of the AL 288-1 femoral head with that of a modern human necessitates all six DOFs to be included in which AL 288-1 could osteologically extend the hip to facilitate a human-like gait