Inter-ray variation in metatarsal strength properties in humans and African apes: Implications for inferring bipedal biomechanics in the Olduvai Hominid 8 foot.

When measured as a ratio of mean midshaft diameter to bone length, the OH 8 fossil hominin foot exhibits a metatarsal (Mt) robusticity pattern of 1 > 5 > 3 > 4 > 2, which differs from the widely perceived "common" modern human pattern (1 > 5 > 4 > 3 > 2); African apes generally exhibit a third pattern (1 > 2 > 3 > 4 > 5). Largely because of the relative ranking of Mt2 and Mt5, OH 8 metatarsals structurally resemble the pattern exhibited by bipedal humans more than the pattern of quadrupedal and climbing African apes. Considering only these three phenotypes, however, discounts the potentially important functional implications of variation in modern human (and African ape) metatarsal robusticity patterns, suggesting that they are not useful for interpreting the specific biomechanics of a bipedal gait in fossils (i.e., whether it was modern human-like or not). Using computed tomography scans to quantify metatarsal midshaft cross-sectional geometry in a large sample of Homo (n=130), Gorilla (n=44) and Pan (n=80), we documented greater variation in metatarsal robusticity patterns than previously recognized in all three groups. While apes consistently show a 1 > 2 > 3 > 4 > 5 pattern in our larger sample, there does not appear to be a similarly precise single "common" human pattern. Rather, human metatarsals converge towards a 1 > 4/5 > 2/3 pattern, where metatarsals 4 and 5, and metatarsals 2 and 3, often "flip" positions relative to each other depending on the variable examined. After reassessing what a "common" human pattern could be based on a larger sample, the previously described OH 8 pattern of 1 > 5 > 3 > 4 > 2 is only observed in some humans (<6%) and almost never in apes (<0.5%). Although this suggests an overall greater similarity to (some) humans than to any ape in loading of the foot, the relatively rare frequency of these humans in our sample underscores potential differences in loading experienced by the medial and lateral columns of the OH 8 foot compared to modern humans.

Metatarsal fusion pattern and developmental morphology of the Olduvai Hominid 8 foot: Evidence of adolescence.

The morphology of the Olduvai Hominid (OH) 8 foot and the sequence of metatarsal epiphyseal fusion in modern humans and chimpanzees support the hypothesis that OH 8 belonged to an individual of approximately the same relative age as the OH 7 subadult, the holotype of Homo habilis. Modern humans and chimpanzees exhibit a variety of metatarsal epiphyseal fusion patterns, including one identical to that observed in OH 8 in which metatarsal 1 fuses before metatarsals 2-5. More than the metatarsal fusion sequence, however, the principal evidence of the youthful age of OH 8 lies in the morphology of metatarsals 1, 2, and 3. Because both OH 8 and OH 7 come from the same stratum at the FLK NN type site, the most parsimonious explanation of the OH 8 and OH 7 data is that this material belonged to the same individual, as originally proposed by Louis Leakey. The proposition that OH 8 belonged to an adult is unsupported by morphology, including radiographic evidence, and the fusion sequences in human and chimpanzee skeletal material reported here and in the literature.

Terrestrial adaptations in the hands of Equatorius africanus revisited.

Interpretations of the postcranial anatomy of East African early and middle Miocene large-bodied hominoids (e.g., Proconsul, Afropithecus, Turkanapithecus, Nacholapithecus) have suggested that these diverse primates utilized positional behaviors dominated by arboreal quadrupedalism. Preliminary descriptions of the Equatorius africanus partial skeleton (KNM-TH 28860) and other forelimb specimens, however, have argued that this animal relied more on terrestrial locomotion compared to its contemporaries, possibly similar to extant large papionin monkeys. In this paper, we reevaluate this interpretation by examining intrinsic hand proportions based on the lengths of the third proximal phalanx and fifth metacarpal in Equatorius in reference to a large sample of extant catarrhine primate taxa. We focused on the lengths of these hand bones because the ratio between phalanx and metacarpal lengths has been previously documented to discriminate terrestrial from arboreal mammalian taxa, including primates. The Equatorius hand displays semi-terrestrial hand proportions with a relatively shorter proximal phalanx compared to most arboreal monkeys. Its proximal phalanx, however, is relatively longer than those of habitually terrestrial monkeys (e.g., Theropithecus, Papio). Accordingly, although Equatorius retains some arboreal quadrupedal characteristics, these results corroborate the previous inference that it engaged in more terrestrial locomotion than earlier Miocene apes such as Proconsul. We suggest that the postcranial skeleton of Equatorius evinces the earliest signs of semi-terrestriality in the hominoid fossil record. It is likely that the terrestrial specialization utilized by living hominoids, e.g., knuckle-walking, evolved separately.

Brief communication: evidence bearing on the status of Homo habilis at Olduvai Gorge.

Students of the early hominin career have debated the status of Homo habilis since its discovery in 1960. Today discussion centers on which specimens should be included in the species and what constitutes the holotype. Recent reviews of early Homo suggest that the Olduvai Hominid 8 foot may sample Paranthropus while the OH 7 skull bones, mandible, and hand sample H. habilis. Moreover, some suggest that while H. habilis in Middle Bed I at Olduvai is craniodentally Homo-like, the postcranial skeleton of H. habilis is more like that of Australopithecus. Evidence presented here indicates not only that OH 7 and OH 8 represent H. habilis but also that they come from a single individual. The association of OH 35 with OH 7 and OH 8 is less certain. Morphological, pathological, and taphonomic evidence favors the inclusion of OH 35 in the holotype. However, stratigraphic evidence suggests that OH 35 and OH 8 are not coterminous. With or without OH 35, the holotype of H. habilis ranks as one of the most complete early hominin skeletons and the most complete and functionally informative specimen of early Homo.

New hominin first metatarsal (SK 1813) from Swartkrans.

A recently recognized hominin hallucal metatarsal, SK 1813, from Swartkrans bears a suite of primitive and derived traits. Comparisons with extant apes, modern humans, SKX 5017, and Stw 562 reveals similar morphology in all three fossils and that these early hominins, while bipedal, possessed a unique toe-off mechanism. The implications of this are that both primitive and derived traits must be used to establish the total biomechanical pattern.

Oreopithecus bambolii: an unlikely case of hominid-like grip capability in a Miocene ape.

Oreopithecus bambolii, an ape from the late Miocene of Italy, is said to possess a hand capable of a precision grip like that of humans. Relative hand length, proportions of the thumb, and morphological features of the thumb and wrist were adduced to support the idea that Oreopithecus had a hand that closely matched the pattern in Australopithecus. A reappraisal of earlier arguments and comparisons of Oreopithecus with humans, apes, and Old World monkeys, reveals that Oreopithecus had an essentially ape-like hand that emphasized ape-like power grasping over human-like precision grasping.