Homo naledi is a previously-unknown species of extinct hominin discovered within the Dinaledi Chamber of the Rising Star cave system, Cradle of Humankind, South Africa. This species is characterized by body mass and stature similar to small-bodied human populations but a small endocranial volume similar to australopiths. Cranial morphology of H. naledi is unique, but most similar to early Homo species including Homo erectus, Homo habilis or Homo rudolfensis. While primitive, the dentition is generally small and simple in occlusal morphology. H. naledi has humanlike manipulatory adaptations of the hand and wrist. It also exhibits a humanlike foot and lower limb. These humanlike aspects are contrasted in the postcrania with a more primitive or australopith-like trunk, shoulder, pelvis and proximal femur. Representing at least 15 individuals with most skeletal elements repeated multiple times, this is the largest assemblage of a single species of hominins yet discovered in Africa.

The difficult birth process of humans, often described as the ''obstetric dilemma,'' is commonly assumed to reflect antagonistic selective pressures favoring neonatal encephalization and maternal bipedal locomotion. However, cephalo-pelvic disproportion is not exclusive to humans, and is present in some primate species of smaller body size. The fossil record indicates mosaic evolution of the obstetric dilemma, involving a number of different evolutionary processes, and it appears to have shifted in magnitude between Australopithecus, Pleistocene Homo, and recent human populations. Most attention to date has focused on its generic nature, rather than on its variability between populations. We reevaluate the nature of the human obstetric dilemma using updated hominin and primate literature, and then consider the contribution of phenotypic plasticity to variability in its magnitude. Both maternal pelvic dimensions and fetal growth patterns are sensitive to ecological factors such as diet and the thermal environment. Neonatal head girth has low plasticity, whereas neonatal mass and maternal stature have higher plasticity. Secular trends in body size may therefore exacerbate or decrease the obstetric dilemma. The emergence of agriculture may have exacerbated the dilemma, by decreasing maternal stature and increasing neonatal growth and adiposity due to dietary shifts. Paleodemographic comparisons between foragers and agriculturalists suggest that foragers have considerably lower rates of perinatal mortality. In contemporary populations, maternal stature remains strongly associated with perinatal mortality in many populations. Long-term improvements in nutrition across future generations may relieve the dilemma, but in the meantime, variability in its magnitude is likely to persist. Yrbk Phys The notion that maternal pelvic dimensions are subject to powerful competing demands from reproduction and locomotion is widely accepted in the biomedical and anthropological literature (Krogman, 1951;Washburn, 1960;Schultz, 1969;Rosenberg, 1992;Rosenberg and Trevathan, 1996;Wittman and Wall, 2007;Walsh, 2008;Franciscus, 2009;Trevathan, 2011). The maternal pelvis is frequently considered to be subject to two counteracting evolutionary forces: decreased height and increased mediolateral breadth in order to optimize the biomechanics of locomotion, and increased anteroposterior dimensions in order to enable birth of the unusually encephalized human infant. The compromise imposed by these antagonistic demands manifests as a difficult passage of the fetal head through the birth canal (Wittman and Wall, 2007), resulting in the birth process being a more complex and lengthy procedure in humans than in closely related species of ape (Tevathan, 1988;Rosenberg, 1992;Rosenberg and Trevathan, 2002). The antagonistic interaction of bipedalism and encephalization has been assumed to have followed the emergence of the large Homo brain within the last 2 million years (Martin, 1983).In addition to shaping the unusual mechanism of birth, h...

Whether early hominins were adept tree climbers is unclear. Although some researchers have argued that bipedality maladapts the hominin skeleton for climbing, others have argued that early hominin fossils display an amalgamation of features consistent with both locomotor strategies. Although chimpanzees have featured prominently in these arguments, there are no published data on the kinematics of climbing in wild chimpanzees. Without these biomechanical data describing how chimpanzees actually climb trees, identifying correlates of climbing in modern ape skeletons is difficult, thereby limiting accurate interpretations of the hominin fossil record. Here, the first kinematic data on vertical climbing in wild chimpanzees are presented. These data are used to identify skeletal correlates of climbing in the ankle joint of the African apes to more accurately interpret hominin distal tibiae and tali. This study finds that chimpanzees engage in an extraordinary range of foot dorsiflexion and inversion during vertical climbing bouts. Two skeletal correlates of modern ape-like vertical climbing are identified in the ankle joint and related to positions of dorsiflexion and foot inversion. A study of the 14 distal tibiae and 15 tali identified and published as hominins from 4.12 to 1.53 million years ago finds that the ankles of early hominins were poorly adapted for modern ape-like vertical climbing bouts. This study concludes that if hominins included tree climbing as part of their locomotor repertoire, then they were performing this activity in a manner decidedly unlike modern chimpanzees.chimpanzee ͉ tibia ͉ talus

A well-preserved and articulated partial foot and ankle of Australopithecus sediba, including an associated complete adult distal tibia, talus, and calcaneus, have been discovered at the Malapa site, South Africa, and reported in direct association with the female paratype Malapa Hominin 2. These fossils reveal a mosaic of primitive and derived features that are distinct from those seen in other hominins. The ankle (talocrural) joint is mostly humanlike in form and inferred function, and there is some evidence for a humanlike arch and Achilles tendon. However, Au. sediba is apelike in possessing a more gracile calcaneal body and a more robust medial malleolus than expected. These observations suggest, if present models of foot function are correct, that Au. sediba may have practiced a unique form of bipedalism and some degree of arboreality. Given the combination of features in the Au. sediba foot, as well as comparisons between Au. sediba and older hominins, homoplasy is implied in the acquisition of bipedal adaptations in the hominin foot.

The midtarsal break was first described in this journal nearly 75 years ago to explain the ability of non-human primates to lift their heel independently of the rest of the foot. Since the initial description of the midtarsal break, the calcaneocuboid joint has been assumed to be the anatomical source of this motion. Recently, however, it has been suggested that the midtarsal break may occur at the cuboid-metatarsal joint, rather than at the calcaneocuboid joint. Data compiled from X-rays, dissections, manual manipulation of living primate feet, video of captive catarrhines, and osteological specimens concur that the midtarsal break is a complex motion caused by dorsiflexion at both joints with the cuboid-metatarsal joint contributing roughly 2/3 of total midfoot dorsiflexion, and the calcaneocuboid joint only about 1/3 of total midfoot dorsiflexion. The convexity of the proximal articular surface of the fourth and fifth metatarsals and corresponding concave cuboid facets provide skeletal correlates for the presence of midfoot dorsiflexion at the cuboid-metatarsal joint. Study of hominin metatarsals from Australopithecus afarensis, A. africanus, Homo erectus, and the metatarsals and a cuboid from the OH 8 foot show little capacity for dorsiflexion at the cuboid-metatarsal joint. These results suggest that hominins may have already evolved a stable midfoot region well adapted for the push-off phase of bipedalism by at least 3.2 million years ago. These data illuminate the evolution of the longitudinal arch and show further evidence of constraints on the arboreal capacity in early hominins.

The discovery of a relatively complete Australopithecus sediba adult female skeleton permits a detailed locomotor analysis in which joint systems can be integrated to form a comprehensive picture of gait kinematics in this late australopith. Here we describe the lower limb anatomy of Au. sediba and hypothesize that this species walked with a fully extended leg and with an inverted foot during the swing phase of bipedal walking. Initial contact of the lateral foot with the ground resulted in a large pronatory torque around the joints of the foot that caused extreme medial weight transfer (hyperpronation) into the toe-off phase of the gait cycle (late pronation). These bipedal mechanics are different from those often reconstructed for other australopiths and suggest that there may have been several forms of bipedalism during the Plio-Pleistocene.

The Rising Star cave system has produced abundant fossil hominin remains within the Dinaledi Chamber, representing a minimum of 15 individuals attributed to Homo naledi. Further exploration led to the discovery of hominin material, now comprising 131 hominin specimens, within a second chamber, the Lesedi Chamber. The Lesedi Chamber is far separated from the Dinaledi Chamber within the Rising Star cave system, and represents a second depositional context for hominin remains. In each of three collection areas within the Lesedi Chamber, diagnostic skeletal material allows a clear attribution to H. naledi. Both adult and immature material is present. The hominin remains represent at least three individuals based upon duplication of elements, but more individuals are likely present based upon the spatial context. The most significant specimen is the near-complete cranium of a large individual, designated LES1, with an endocranial volume of approximately 610 ml and associated postcranial remains. The Lesedi Chamber skeletal sample extends our knowledge of the morphology and variation of H. naledi, and evidence of H. naledi from both recovery localities shows a consistent pattern of differentiation from other hominin species.DOI: http://dx.doi.org/10.7554/eLife.24232.001

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