New postcranial fossils of Australopithecus afarensis from Hadar, Ethiopia (1990-2007).

Renewed fieldwork at Hadar, Ethiopia, from 1990 to 2007, by a team based at the Institute of Human Origins, Arizona State University, resulted in the recovery of 49 new postcranial fossils attributed to Australopithecus afarensis. These fossils include elements from both the upper and lower limbs as well as the axial skeleton, and increase the sample size of previously known elements for A. afarensis. The expanded Hadar sample provides evidence of multiple new individuals that are intermediate in size between the smallest and largest individuals previously documented, and so support the hypothesis that a single dimorphic species is represented. Consideration of the functional anatomy of the new fossils supports the hypothesis that no functional or behavioral differences need to be invoked to explain the morphological variation between large and small A. afarensis individuals. Several specimens provide important new data about this species, including new vertebrae supporting the hypothesis that A. afarensis may have had a more human-like thoracic form than previously appreciated, with an invaginated thoracic vertebral column. A distal pollical phalanx confirms the presence of a human-like flexor pollicis longus muscle in A. afarensis. The new fossils include the first complete fourth metatarsal known for A. afarensis. This specimen exhibits the dorsoplantarly expanded base, axial torsion and domed head typical of humans, revealing the presence of human-like permanent longitudinal and transverse arches and extension of the metatarsophalangeal joints as in human-like heel-off during gait. The new Hadar postcranial fossils provide a more complete picture of postcranial functional anatomy, and individual and temporal variation within this sample. They provide the basis for further in-depth analyses of the behavioral and evolutionary significance of A. afarensis anatomy, and greater insight into the biology and evolution of these early hominins.

Complete fourth metatarsal and arches in the foot of Australopithecus afarensis.

The transition to full-time terrestrial bipedality is a hallmark of human evolution. A key correlate of human bipedalism is the development of longitudinal and transverse arches of the foot that provide a rigid propulsive lever and critical shock absorption during striding bipedal gait. Evidence for arches in the earliest well-known Australopithecus species, A. afarensis, has long been debated. A complete fourth metatarsal of A. afarensis was recently discovered at Hadar, Ethiopia. It exhibits torsion of the head relative to the base, a direct correlate of a transverse arch in humans. The orientation of the proximal and distal ends of the bone reflects a longitudinal arch. Further, the deep, flat base and tarsal facets imply that its midfoot had no ape-like midtarsal break. These features show that the A. afarensis foot was functionally like that of modern humans and support the hypothesis that this species was a committed terrestrial biped.

Was Australopithecus anamensis ancestral to A. afarensis? A case of anagenesis in the hominin fossil record.

We tested the hypothesis that early Pliocene Australopithecus anamensis was ancestral to A. afarensis by conducting a phylogenetic analysis of four temporally successive fossil samples assigned to these species (from earliest to latest: Kanapoi, Allia Bay, Laetoli, Hadar) using polarized character-state data from 20 morphological characters of the dentition and jaws. If the hypothesis that A. anamensis is ancestral to A. afarensis is true, then character-state changes between the temporally ordered site-samples should be congruent with hypothesized polarity transformations based on outgroup (African great ape) conditions. The most parsimonious reconstruction of character-state evolution suggests that each of the hominin OTUs shares apomorphies only with geologically younger OTUs, as predicted by the hypothesis of ancestry (tree length=31; Consistency Index=0.903). This concordance of stratigraphic and character-state data supports the idea that the A. anamensis and A. afarensis samples represent parts of an anagenetically evolving lineage, or evolutionary species. Each site-sample appears to capture a different point along this evolutionary trajectory. We discuss the implications of this conclusion for the taxonomy and adaptive evolution of these early-middle Pliocene hominins.