Bernhard Zipfel

Homo naledi, a new species of the genus Homo from the Dinaledi Chamber, South Africa

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. DOI: http://dx.doi.org/10.7554/eLife.09560.001

The Foot and Ankle of Australopithecus sediba

Australopithecus sediba had a human-like ankle and arch but an ape-like heel and tibia, implying that while bipedal, this species was also adept at climbing trees. 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 Lower Limb and Mechanics of Walking in Australopithecus sediba

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.

Recent origin of low trabecular bone density in modern humans

Significance The human skeleton is unique in having low trabecular density representing a lightly built human body form. However, it remains unknown when during human evolution this unique characteristic first appeared. To our knowledge, this study is the first to examine trabecular bone density throughout the skeleton of fossil hominins spanning several million years. The results show that trabecular density remained high throughout human evolution until it decreased significantly in recent modern humans, suggesting a possible link between changes in our skeleton and increased sedentism. Humans are unique, compared with our closest living relatives (chimpanzees) and early fossil hominins, in having an enlarged body size and lower limb joint surfaces in combination with a relatively gracile skeleton (i.e., lower bone mass for our body size). Some analyses have observed that in at least a few anatomical regions modern humans today appear to have relatively low trabecular density, but little is known about how that density varies throughout the human skeleton and across species or how and when the present trabecular patterns emerged over the course of human evolution. Here, we test the hypotheses that (i) recent modern humans have low trabecular density throughout the upper and lower limbs compared with other primate taxa and (ii) the reduction in trabecular density first occurred in early Homo erectus, consistent with the shift toward a modern human locomotor anatomy, or more recently in concert with diaphyseal gracilization in Holocene humans. We used peripheral quantitative CT and microtomography to measure trabecular bone of limb epiphyses (long bone articular ends) in modern humans and chimpanzees and in fossil hominins attributed to Australopithecus africanus, Paranthropus robustus/early Homo from Swartkrans, Homo neanderthalensis, and early Homo sapiens. Results show that only recent modern humans have low trabecular density throughout the limb joints. Extinct hominins, including pre-Holocene Homo sapiens, retain the high levels seen in nonhuman primates. Thus, the low trabecular density of the recent modern human skeleton evolved late in our evolutionary history, potentially resulting from increased sedentism and reliance on technological and cultural innovations.

The Foot of Homo naledi

Modern humans are characterized by a highly specialized foot that reflects our obligate bipedalism. Our understanding of hominin foot evolution is, although, hindered by a paucity of well-associated remains. Here we describe the foot of Homo naledi from Dinaledi Chamber, South Africa, using 107 pedal elements, including one nearly-complete adult foot. The H. naledi foot is predominantly modern human-like in morphology and inferred function, with an adducted hallux, an elongated tarsus, and derived ankle and calcaneocuboid joints. In combination, these features indicate a foot well adapted for striding bipedalism. However, the H. naledi foot differs from modern humans in having more curved proximal pedal phalanges, and features suggestive of a reduced medial longitudinal arch. Within the context of primitive features found elsewhere in the skeleton, these findings suggest a unique locomotor repertoire for H. naledi, thus providing further evidence of locomotor diversity within both the hominin clade and the genus Homo.

Possible brucellosis in an early hominin skeleton from sterkfontein, South Africa.

We report on the paleopathological analysis of the partial skeleton of the late Pliocene hominin species Australopithecus africanus Stw 431 from Sterkfontein, South Africa. A previous study noted the presence of lesions on vertebral bodies diagnosed as spondylosis deformans due to trauma. Instead, we suggest that these lesions are pathological changes due to the initial phases of an infectious disease, brucellosis. The macroscopic, microscopic and radiological appearance of the lytic lesions of the lumbar vertebrae is consistent with brucellosis. The hypothesis of brucellosis (most often associated with the consumption of animal proteins) in a 2.4 to 2.8 million year old hominid has a host of important implications for human evolution. The consumption of meat has been regarded an important factor in supporting, directing or altering human evolution. Perhaps the earliest (up to 2.5 million years ago) paleontological evidence for meat eating consists of cut marks on animal remains and stone tools that could have made these marks. Now with the hypothesis of brucellosis in A. africanus, we may have evidence of occasional meat eating directly linked to a fossil hominin.

Earliest complete hominin fifth metatarsal : implications for the evolution of the lateral column of the foot

StW 114/115, from Sterkfontein, South Africa, is the earliest complete hominin fifth metatarsal. Comparisons of StW 114/115 to modern humans, extant apes, and partial hominin metatarsals AL 333-13, AL 333-78, SKX 33380, OH 8, and KNM-ER 803f reveal a similar morphology in all six fossils consistent with habitual bipedality. Although StW 114/115 possesses some primitive characters, the proximal articular morphology and internal torsion of the head are very human-like, suggesting a stable lateral column and the likely presence of lateral longitudinal and transverse tarsal arches. We conclude that, at least in the lateral component of the foot of the StW 114/115 individual, the biomechanical pattern is very similar to that of modern humans. This, however, may not have been the case in the medial column of the foot, as a mosaic pattern of hominin foot evolution and function has been suggested. The results of this study may support the hypothesis of an increased calcaneo-cuboid stability having been an early evolutionary event in the history of terrestrial bipedalism.

A complete second metatarsal (StW 89) from Sterkfontein Member 4, South Africa

The functional anatomy of the hominin foot has played a crucial role in studies of locomotor evolution in human ancestors and extinct relatives. However, foot fossils are rare, often isolated, and fragmentary. Here, we describe a complete hominin second metatarsal (StW 89) from the 2.0-2.6 million year old deposits of Member 4, Sterkfontein Cave, South Africa. Like many other fossil foot bones, it displays a mosaic of derived human-like features and primitive ape-like features. StW 89 possesses a domed metatarsal head with a prominent sulcus, indicating dorsiflexion at the metatarsophalangeal joint during bipedal walking. However, while the range of motion at the metatarsophalangeal joint is human-like in dorsiflexion, it is ape-like in plantarflexion. Furthermore, StW 89 possesses internal torsion of the head, an anatomy decidedly unlike that found in humans today. Unlike other hominin second metatarsals, StW 89 has a dorsoplantarly gracile base, perhaps suggesting more midfoot laxity. In these latter two anatomies, the StW 89 second metatarsal is quite similar to the recently described second metatarsal of the partial foot from Burtele, Ethiopia. We interpret this combination of anatomies as evidence for a low medial longitudinal arch in a foot engaged in both bipedal locomotion, but also some degree of pedal, and perhaps even hallucal, grasping. Additional fossil evidence will be required to determine if differences between this bone and other second metatarsals from Sterkfontein reflect normal variation in an evolving lineage, or taxonomic diversity.

Brief communication: radiographic study of metatarsal one basal epiphyseal fusion: a note of caution on age determination.

This study examines radiographs of first metatarsals of 131 individuals from age 17-88 years to determine whether internal basal epiphyseal lines may be visible past the age of metatarsal fusion, which usually occurs between 14 and 16 years of age (Scheuer and Black: The juvenile skeleton. San Diego: Elsevier Academic Press,2004). In 29% (38 out of 131) of the radiographed first metatarsals (MT1s) the basal epiphyseal scar is visible, including in one individual who was 80 years old. Statistically, there was no relationship between the loss of the epiphyseal scar and age. Thus, the presence of the epiphyseal scar does not necessarily indicate subadult age. These data suggest that OH 8s radiographically visible basal epiphyseal line has no bearing on whether it is a subadult or not.

The Olduvai Hominid 8 foot: Adult or subadult?

Olduvai Hominid 8 (OH 8), an articulating set of fossil hominin tarsal and metatarsal bones, is critical to interpretations of the evolution of hominin pedal morphology and bipedal locomotion. It has been suggested that OH 8 may represent the foot of a subadult and may be associated with the OH 7 mandible, the type specimen of Homo habilis. This assertion is based on the presence of what may be unfused distal metatarsal epiphyses. Accurately assessing the skeletal maturity of the OH 8 foot is important for interpretations of the functional morphology and locomotor behavior of Plio-Pleistocene hominins. In this study, we compare metatarsal fusion patterns and internal bone morphology of the lateral metatarsals among subadult hominines (85 modern humans, 48 Pan, and 25 Gorilla) to assess the likelihood that OH 8 belonged to either an adult or subadult hominin. Our results suggest that if OH 8 is indeed from a subadult, then it displays a metatarsal developmental pattern that is unobserved in our comparative sample. In OH 8, the fully fused base of the first metatarsal and the presence of trabecular bone at the distal ends of the second and third metatarsal shafts make it highly improbable that it belonged to a subadult, let alone a subadult that matches the developmental age of the OH 7 mandible. In total, the results of this study suggest that the OH 8 foot most likely belonged to an adult hominin.