Cosmogenic nuclide dating of Australopithecus at Sterkfontein, South Africa.

Sterkfontein is the most prolific single source of Australopithecus fossils, the vast majority of which were recovered from Member 4, a cave breccia now exposed by erosion and weathering at the landscape surface. A few other Australopithecus fossils, including the StW 573 skeleton, come from subterranean deposits [T. C. Partridge et al., Science 300, 607-612 (2003); R. J. Clarke, K. Kuman, J. Hum. Evol. 134, 102634 (2019)]. Here, we report a cosmogenic nuclide isochron burial date of 3.41 ± 0.11 million years (My) within the lower middle part of Member 4, and simple burial dates of 3.49 ± 0.19 My in the upper middle part of Member 4 and 3.61 ± 0.09 My in Jacovec Cavern. Together with a previously published isochron burial date of 3.67 ± 0.16 My for StW 573 [D. E. Granger et al., Nature 522, 85-88 (2015)], these results place nearly the entire Australopithecus assemblage at Sterkfontein in the mid-Pliocene, contemporaneous with Australopithecus afarensis in East Africa. Our ages for the fossil-bearing breccia in Member 4 are considerably older than the previous ages of ca. 2.1 to 2.6 My interpreted from flowstones associated with the same deposit. We show that these previously dated flowstones are stratigraphically intrusive within Member 4 and that they therefore underestimate the true age of the fossils.

Bipedalism or bipedalisms: The os coxae of StW 573.

There has been a long debate about the possibility of multiple contemporaneous species of Australopithecus in both eastern and southern Africa, potentially exhibiting different forms of bipedal locomotion. Here, we describe the previously unreported morphology of the os coxae in the 3.67 Ma Australopithecus prometheus StW 573 from Sterkfontein Member 2, comparing it with variation in ossa coxae in living humans and apes as well as other Plio-Pleistocene hominins. Statistical comparisons indicate that StW 573 and 431 resemble humans in their anteroposteriorly great iliac crest breadth compared with many other early australopiths, whereas Homo ergaster KNM WT 15000 surprisingly also has a relatively anterioposteriorly short iliac crest. StW 573 and StW 431 appear to resemble humans in having a long ischium compared with Sts 14 and KNM WT 15000. A Quadratic Discriminant Function Analysis of morphology compared with other Plio-Pleistocene hominins and a dataset of modern humans and hominoids shows that, while Lovejoy's heuristic model of the Ardipithecus ramidus os coxae falls with Pongo or in an indeterminate group, StW 573 and StW 431 from Sterkfontein Member 4 are consistently classified together with modern humans. Although clearly exhibiting the classic "basin shaped" bipedal pelvis, Sts 14 (also from Sterkfontein), AL 288-1 Australopithecus afarensis, MH2 Australopithecus sediba and KNM-WT 15000 occupy a position more peripheral to modern humans, and in some analyses are assigned to an indeterminate outlying group. Our findings strongly support the existence of two species of Australopithecus at Sterkfontein and the variation we observe in os coxae morphology in early hominins is also likely to reflect multiple forms of bipedality.

Dart and the Taung juvenile: making sense of a century-old record of hominin evolution in Africa.

The announcement in 1925 by Raymond Dart of the discovery of the Taung juvenile's skull in a quarry in sub-Saharan Africa is deservedly a classic publication in the history of palaeoanthropology. Dart's paper-which designated Taung as the type specimen of the early hominin species Australopithecus africanus-provided the first fossil evidence supporting Charles Darwin's 1871 prediction that Africa was where the human lineage originated. The Taung juvenile's combination of ape and human characteristics eventually led to a paradigm shift in our understanding of human evolution. This contribution focuses on the milieu in which Dart's paper appeared (i.e. what was understood in 1925 about human evolution), the fossil evidence as set out by Dart, his interpretation of how a species represented by a fossilized juvenile's skull fitted within prevailing narratives about human evolution and the significance of the fossil being found in an environment inferred to be very different from that occupied by living apes. We also briefly review subsequent fossil finds that have corroborated the argument Dart made for having discovered evidence of a hitherto unknown close relative of humans, and summarize our current understanding of the earliest stages of human evolution and its environmental context.

A review of the distal femur in Australopithecus.

In 1938, the first distal femur of a fossil Australopithecus was discovered at Sterkfontein, South Africa. A decade later, another distal femur was discovered at the same locality. These two fossil femora were the subject of a foundational paper authored by Kingsbury Heiple and Owen Lovejoy in 1971. In this paper, the authors discussed functionally relevant anatomies of these two fossil femora and noted their strong affinity to the modern human condition. Here, we update this work by including eight more fossil Australopithecus distal femora, an expanded comparative dataset, as well as additional linear measurements. Just as Heiple and Lovejoy reported a half-century ago, we find strong overlap between modern humans and cercopithecoids, except for inferiorly flattened condyles and a high bicondylar angle, both of which characterize modern humans and Australopithecus and are directly related to striding bipedalism. All other measured aspects of the femora are by-products of these key morphological traits. Additional fossil material from the early Pliocene will help to inform the evolution of the hominin distal femur and its condition in the Pan-Homo common ancestor that preceded bipedal locomotion.

Canine sexual dimorphism in Ardipithecus ramidus was nearly human-like.

Body and canine size dimorphism in fossils inform sociobehavioral hypotheses on human evolution and have been of interest since Darwin's famous reflections on the subject. Here, we assemble a large dataset of fossil canines of the human clade, including all available Ardipithecus ramidus fossils recovered from the Middle Awash and Gona research areas in Ethiopia, and systematically examine canine dimorphism through evolutionary time. In particular, we apply a Bayesian probabilistic method that reduces bias when estimating weak and moderate levels of dimorphism. Our results show that Ar. ramidus canine dimorphism was significantly weaker than in the bonobo, the least dimorphic and behaviorally least aggressive among extant great apes. Average male-to-female size ratios of the canine in Ar. ramidus are estimated as 1.06 and 1.13 in the upper and lower canines, respectively, within modern human population ranges of variation. The slightly greater magnitude of canine size dimorphism in the lower than in the upper canines of Ar. ramidus appears to be shared with early Australopithecus, suggesting that male canine reduction was initially more advanced in the behaviorally important upper canine. The available fossil evidence suggests a drastic size reduction of the male canine prior to Ar. ramidus and the earliest known members of the human clade, with little change in canine dimorphism levels thereafter. This evolutionary pattern indicates a profound behavioral shift associated with comparatively weak levels of male aggression early in human evolution, a pattern that was subsequently shared by Australopithecus and Homo.

A comparative approach for characterizing the relationship among morphology, range-of-motion and locomotor behaviour in the primate shoulder.

Shoulder shape directly impacts forelimb function by contributing to glenohumeral (GH) range-of-motion (ROM). However, identifying traits that contribute most to ROM and visualizing how they do so remains challenging, ultimately limiting our ability to reconstruct function and behaviour in fossil species. To address these limitations, we developed an in silico proximity-driven model to simulate and visualize three-dimensional (3D) GH rotations in living primate species with diverse locomotor profiles, identify those shapes that are most predictive of ROM using geometric morphometrics, and apply subsequent insights to interpret function and behaviour in the fossil hominin Australopithecus sediba. We found that ROM metrics that incorporated 3D rotations best discriminated locomotor groups, and the magnitude of ROM (mobility) was decoupled from the anatomical location of ROM (e.g. high abduction versus low abduction). Morphological traits that enhanced mobility were decoupled from those that enabled overhead positions, and all non-human apes possessed the latter but not necessarily the former. Model simulation in A. sediba predicted high mobility and a ROM centred at lower abduction levels than in living apes but higher than in modern humans. Together these results identify novel form-to-function relationships in the shoulder and enhance visualization tools to reconstruct past function and behaviour.

An updated analysis of hominin phylogeny with an emphasis on re-evaluating the phylogenetic relationships of Australopithecus sediba.

The discovery and description of Australopithecus sediba has reignited the debate over the evolutionary history of the australopiths and the genus Homo. It has been suggested that A. sediba may be an ancestor of Homo because it possesses a mosaic of derived Homo-like and primitive australopith-like traits. However, an alternative hypothesis proposes that the majority of the purported Homo-like craniodental characters can be attributed to the juvenile status of the type specimen, MH1. We conducted an independent character assessment of the craniodental morphology of A. sediba, with particular emphasis on evaluating whether the ontogenetic status of MH1 may have affected its purported Homo-like characteristics. In doing so, we have also expanded fossil hypodigms to incorporate the new Australopithecus anamensis cranium from Woranso-Mille (MRD-VP-1/1), as well as recently described Paranthropus robustus cranial remains from Drimolen (DNH 7, DNH 155). Morphological character data were analyzed using both standard parsimony and Bayesian techniques. In addition, we conducted a series of Bayesian analyses constrained to evaluate the hypothesis that Australopithecus africanus and A. sediba are sister taxa. Based on the results of the parsimony and Bayesian analyses, we could not reject the hypothesis that A. sediba shares its closest phylogenetic affinities with the genus Homo. Therefore, based on currently available craniodental evidence, we conclude that A. sediba is plausibly the terminal end of a lineage that shared a common ancestor with the earliest representatives of Homo. We caution, however, that the discovery of new A. sediba fossils preserving adult cranial morphology or the inclusion of postcranial characters may ultimately necessitate a re-evaluation of this hypothesis.

Knuckle-walking in Sahelanthropus? Locomotor inferences from the ulnae of fossil hominins and other hominoids.

Because the ulna supports and transmits forces during movement, its morphology can signal aspects of functional adaptation. To test whether, like extant apes, some hominins habitually recruit the forelimb in locomotion, we separate the ulna shaft and ulna proximal complex for independent shape analyses via elliptical Fourier methods to identify functional signals. We examine the relative influence of locomotion, taxonomy, and body mass on ulna contours in Homo sapiens (n = 22), five species of extant apes (n = 33), two Miocene apes (Hispanopithecus and Danuvius), and 17 fossil hominin specimens including Sahelanthropus, Ardipithecus, Australopithecus, Paranthropus, and early Homo. Ulna proximal complex contours correlate with body mass but not locomotor patterns, while ulna shafts significantly correlate with locomotion. African apes' ulna shafts are more robust and curved than Asian apes and are unlike other terrestrial mammals (including other primates), curving ventrally rather than dorsally. Because this distinctive curvature is absent in orangutans and hylobatids, it is likely a function of powerful flexors engaged in wrist and hand stabilization during knuckle-walking, and not an adaptation to climbing or suspensory behavior. The OH 36 (purported Paranthropus boisei) and TM 266 (assigned to Sahelanthropus tchadensis) fossils differ from other hominins by falling within the knuckle-walking morphospace, and thus appear to show forelimb morphology consistent with terrestrial locomotion. Discriminant function analysis classifies both OH 36 and TM 266 with Pan and Gorilla with high posterior probability. Along with its associated femur, the TM 266 ulna shaft contours and its deep, keeled trochlear notch comprise a suite of traits signaling African ape-like quadrupedalism. While implications for the phylogenetic position and hominin status of S. tchadensis remain equivocal, this study supports the growing body of evidence indicating that S. tchadensis was not an obligate biped, but instead represents a late Miocene hominid with knuckle-walking adaptations.

Challenges and perspectives on functional interpretations of australopith postcrania and the reconstruction of hominin locomotion.

In 1994, Hunt published the 'postural feeding hypothesis'-a seminal paper on the origins of hominin bipedalism-founded on the detailed study of chimpanzee positional behavior and the functional inferences derived from the upper and lower limb morphology of the Australopithecus afarensis A.L. 288-1 partial skeleton. Hunt proposed a model for understanding the potential selective pressures on hominins, made robust, testable predictions based on Au. afarensis functional morphology, and presented a hypothesis that aimed to explain the dual functional signals of the Au. afarensis and, more generally, early hominin postcranium. Here we synthesize what we have learned about Au. afarensis functional morphology and the dual functional signals of two new australopith discoveries with relatively complete skeletons (Australopithecus sediba and StW 573 'Australopithecus prometheus'). We follow this with a discussion of three research approaches that have been developed for the purpose of drawing behavioral inferences in early hominins: (1) developments in the study of extant apes as models for understanding hominin origins; (2) novel and continued developments to quantify bipedal gait and locomotor economy in extant primates to infer the locomotor costs from the anatomy of fossil taxa; and (3) novel developments in the study of internal bone structure to extract functional signals from fossil remains. In conclusion of this review, we discuss some of the inherent challenges of the approaches and methodologies adopted to reconstruct the locomotor modes and behavioral repertoires in extinct primate taxa, and notably the assessment of habitual terrestrial bipedalism in early hominins.

Pliocene hominins from East Turkana were associated with mesic environments in a semiarid basin.

During the middle Pliocene (∼3.8-3.2 Ma), both Australopithecus afarensis and Kenyanthropus platyops are known from the Turkana Basin, but between 3.60 and 3.44 Ma, most hominin fossils are found on the west side of Lake Turkana. Here, we describe a new hominin locality (ET03-166/168, Area 129) from the east side of the lake, in the Lokochot Member of the Koobi Fora Formation (3.60-3.44 Ma). To reconstruct the paleoecology of the locality and its surroundings, we combine information from sedimentology, the relative abundance of associated mammalian fauna, phytoliths, and stable isotopes from plant wax biomarkers, pedogenic carbonates, and fossil tooth enamel. The combined evidence provides a detailed view of the local paleoenvironment occupied by these Pliocene hominins, where a biodiverse community of primates, including hominins, and other mammals inhabited humid, grassy woodlands in a fluvial floodplain setting. Between <3.596 and 3.44 Ma, increases in woody vegetation were, at times, associated with increases in arid-adapted grasses. This suggests that Pliocene vegetation included woody species that were resilient to periods of prolonged aridity, resembling vegetation structure in the Turkana Basin today, where arid-adapted woody plants are a significant component of the ecosystem. Pedogenic carbonates indicate more woody vegetation than other vegetation proxies, possibly due to differences in temporospatial scale and ecological biases in preservation that should be accounted for in future studies. These new hominin fossils and associated multiproxy paleoenvironmental indicators from a single locale through time suggest that early hominin species occupied a wide range of habitats, possibly including wetlands within semiarid landscapes. Local-scale paleoecological evidence from East Turkana supports regional evidence that middle Pliocene eastern Africa may have experienced large-scale, climate-driven periods of aridity. This information extends our understanding of hominin environments beyond the limits of simple wooded, grassy, or mosaic environmental descriptions.

The estimation and evolution of hominin body mass.

Body mass is a critical variable in many hominin evolutionary studies, with implications for reconstructing relative brain size, diet, locomotion, subsistence strategy, and social organization. We review methods that have been proposed for estimating body mass from true and trace fossils, consider their applicability in different contexts, and the appropriateness of different modern reference samples. Recently developed techniques based on a wider range of modern populations hold promise for providing more accurate estimates in earlier hominins, although uncertainties remain, particularly in non-Homo taxa. When these methods are applied to almost 300 Late Miocene through Late Pleistocene specimens, the resulting body mass estimates fall within a 25-60 kg range for early non-Homo taxa, increase in early Homo to about 50-90 kg, then remain constant until the Terminal Pleistocene, when they decline.

Comparative description and taxonomic affinity of 3.7-million-year-old hominin mandibles from Woranso-Mille (Ethiopia).

Fossil discoveries of early Australopithecus species from Woranso-Mille have played a significant role in improving our understanding of mid-Pliocene hominin evolution and diversity. Here, we describe two mandibles with dentitions, recovered from sediments immediately above a tuff radiometrically dated to 3.76 ± 0.02 Ma, and assess their taxonomic affinity. The two mandibles (MSD-VP-5/16 and MSD-VP-5/50) show morphological similarities with both Australopithecus anamensis and Australopithecus afarensis. Some of the unique features that distinguish Au. anamensis from Au. afarensis are present in the mandibles, which also share a few derived features with Au. afarensis. Their retention of more Kanapoi Au. anamensis-like traits, compared to the fewer derived features they share with Au. afarensis, and the presence of Au. anamensis at Woranso-Mille in 3.8-million-year-old deposits, lends support to their assignment to Au. anamensis. However, it is equally arguable that the few derived dentognathic features they share with Au. afarensis could be taxonomically more significant, making it difficult to conclusively assign these specimens to either species. Regardless of which species they are assigned to, the mosaic nature of the dentognathic morphology and geological age of the two mandibles lends further support to the hypothesized ancestor-descendant relationship between Au. anamensis and Au. afarensis. However, there is now limited fossil evidence indicating that these two species may have overlapped in time. Hence, the last appearance of Au. anamensis and first appearance of Au. afarensis are currently unknown. Recovery of Australopithecus fossils from 4.1 to 3.8 Ma is critical to further address the timing of these events.

Mosaic habitats at Woranso-Mille (Ethiopia) during the Pliocene and implications for Australopithecus paleoecology and taxonomic diversity.

Many important Pliocene hominin specimens have been recovered from Woranso-Mille, a paleontological research area in the Afar region of Ethiopia, including the complete cranium of Australopithecus anamensis, a partial skeleton of Australopithecus afarensis, mandibular and maxillary elements representing a new species, Australopithecus deyiremeda, and a partial foot of an as-yet-unnamed species. Woranso-Mille is the only site, so far, to have reported the co-existence of more than one early hominin species between 3.8 and 3.3 Ma and the temporal overlap between A. anamensis and A. afarensis. Thus, the site has important implications for our understanding of the paleoecology and taxonomic diversity of early hominins and their ecological niche. This paper explores the paleohabitats of Woranso-Mille through its faunal community ecological structure and taxonomic composition using correspondence analysis and Forbes modified similarity index. The results suggest that Pliocene Woranso-Mille was a mosaic of different habitat types, with riparian woodland and floodplain grassland along rivers draining into a lake, along with less mesic habitats such as woodland, grassland, and shrubland. The apparent high level of vegetation heterogeneity may have promoted dietary specializations and niche differentiation among the different Australopithecus species at Woranso-Mille and allowed for their co-existence at the site.

Estimating origination times from the early hominin fossil record.

The age of the earliest recovered fossil evidence of a hominin taxon is all too often equated with that taxon's origination. However, the earliest known fossil record nearly always postdates, sometimes by a substantial period of time, the true origination of a taxon. Here we evaluate the first appearance records of the earliest potential hominins (Sahelanthropus, Ardipithecus, Orrorin), as well as of the genera Australopithecus, Homo, and Paranthropus, to illustrate the considerable uncertainty regarding the actual timing of origin of these taxa. By placing confidence intervals on the first appearance records of early hominin taxa, we can better evaluate patterns of hominin diversity, turnover, and potential correlations with climatic and environmental changes.

The hominid ilium is shaped by a synapomorphic growth mechanism that is unique within primates.

The human ilium is significantly shorter and broader than those of all other primates. In addition, it exhibits an anterior inferior iliac spine (AIIS) that emerges via a secondary center of ossification, which is unique to hominids (i.e., all taxa related to the human clade following their phyletic separation from the African apes). Here, we track the ontogeny of human and other primate ossa coxae. The human pattern is unique, from anlage to adulthood, and fusion of its AIIS is the capstone event in a repositioning of the anterior gluteals that maximizes control of pelvic drop during upright walking. It is therefore a hominid synapomorphy that can be used to assess the presence and age of bipedal locomotion in extinct taxa.

Body mass estimation from footprint size in hominins.

Although many studies relating stature to foot length have been carried out, the relationship between foot size and body mass remains poorly understood. Here we investigate this relationship in 193 adult and 50 juvenile habitually unshod/minimally shod individuals from five different populations-Machiguenga, Daasanach, Pumé, Hadzabe, and Samoans-varying greatly in body size and shape. Body mass is highly correlated with foot size, and can be predicted from foot area (maximum length × breadth) in the combined sample with an average error of about 10%. However, comparisons among populations indicate that body shape, as represented by the body mass index (BMI), has a significant effect on foot size proportions, with higher BMI samples exhibiting relatively smaller feet. Thus, we also derive equations for estimating body mass from both foot size and BMI, with BMI in footprint samples taken as an average value for a taxon or population, estimated independently from skeletal remains. Techniques are also developed for estimating body mass in juveniles, who have relatively larger feet than adults, and for converting between foot and footprint size. Sample applications are given for five Pliocene through Holocene hominin footprint samples from Laetoli (Australopithecus afarensis), Ileret (probable Homo erectus), Happisburgh (possible Homo antecessor), Le Rozel (archaic Homo sapiens), and Barcin Höyük (H. sapiens). Body mass estimates for Homo footprint samples appear reasonable when compared to skeletal estimates for related samples. However, estimates for the Laetoli footprint sample using the new formulae appear to be too high when compared to skeletal estimates for A. afarensis. Based on the proportions of A.L. 288-1, this is apparently a result of relatively large feet in this taxon. A different method using a ratio between body mass and foot area in A.L. 288-1 provides estimates more concordant with skeletal estimates and should be used for A. afarensis.

The pectoral girdle of StW 573 ('Little Foot') and its implications for shoulder evolution in the Hominina.

The ca. 3.67 Ma adult skeleton known as 'Little Foot' (StW 573), recovered from Sterkfontein Member 2 breccia in the Silberberg Grotto, is remarkable for its morphology and completeness. Preservation of clavicles and scapulae, including essentially complete right-side elements, offers opportunities to assess morphological and functional aspects of a nearly complete Australopithecus pectoral girdle. Here we describe the StW 573 pectoral girdle and offer quantitative comparisons to those of extant hominoids and selected homininans. The StW 573 pectoral girdle combines features intermediate between those of humans and other apes: a long and curved clavicle, suggesting a relatively dorsally positioned scapula; an enlarged and uniquely proportioned supraspinous fossa; a relatively cranially oriented glenoid fossa; and ape-like reinforcement of the axillary margin by a stout ventral bar. StW 573 scapulae are as follows: smaller than those of some homininans (i.e., KSD-VP-1/1 and KNM-ER 47000A), larger than others (i.e., A.L. 288-1, Sts 7, and MH2), and most similar in size to another australopith from Sterkfontein, StW 431. Moreover, StW 573 and StW 431 exhibit similar structural features along their axillary margins and inferior angles. As the StW 573 pectoral girdle (e.g., scapular configuration) has a greater affinity to that of apes-Gorilla in particular-rather than modern humans, we suggest that the StW 573 morphological pattern appears to reflect adaptations to arboreal behaviors, especially those with the hand positioned above the head, more than human-like manipulatory capabilities. When compared with less complete pectoral girdles from middle/late Miocene apes and that of the penecontemporaneous KSD-VP-1/1 (Australopithecus afarensis), and mindful of consensus views on the adaptiveness of arboreal positional behaviors soliciting abducted glenohumeral joints in early Pliocene taxa, we propose that the StW 573 pectoral girdle is a reasonable model for hypothesizing pectoral girdle configuration of the crown hominin last common ancestor.

Calcaneal shape variation in humans, nonhuman primates, and early hominins.

The foot has played a prominent role in evaluating early hominin locomotion. The calcaneus, in particular, plays an important role in weight-bearing. Although the calcanei of early hominins have been previously scrutinized, a three-dimensional analysis of the entire calcaneal shape has not been conducted. Here, we investigate the relationship between external calcaneal shape and locomotion in modern Homo sapiens (n = 130), Gorilla (n = 86), Pan (n = 112), Pongo (n = 31), Papio (n = 28), and hylobatids (Hylobates, Symphalangus; n = 32). We use these results to place the calcanei attributed to Australopithecus sediba, A. africanus, A. afarensis, H. naledi, and Homo habilis/Paranthropus boisei into a locomotor context. Calcanei were scanned using either surface scanning or micro-CT and their external shape analyzed using a three-dimensional geometric morphometric sliding semilandmark analysis. Blomberg's K statistic was used to estimate phylogenetic signal in the shape data. Shape variation was summarized using a principal components analysis. Procrustes distances between all taxa as well as distances between each fossil and the average of each taxon were calculated. Blomberg's K statistic was small (K = 0.1651), indicating weak phylogenetic effects, suggesting variation is driven by factors other than phylogeny (e.g., locomotion or body size). Modern humans have a large calcaneus relative to body size and display a uniquely convex cuboid facet, facilitating a rigid midfoot for bipedalism. More arboreal great apes display relatively deeper cuboid facet pivot regions for increased midfoot mobility. Australopithecus afarensis demonstrates the most human-like calcaneus, consistent with obligate bipedalism. Homo naledi is primarily modern human-like, but with some intermediate traits, suggesting a different form of bipedalism than modern humans. Australopithecus africanus, A. sediba, and H. habilis/P. boisei calcanei all possess unique combinations of human and nonhuman ape-like morphologies, suggesting a combination of bipedal and arboreal behaviors.

The DNH 7 skull of Australopithecus robustus from Drimolen (Main Quarry), South Africa.

Although the early hominin species Australopithecus robustus has been known for more than eight decades and is represented by hundreds of fossils from sites in South Africa, a complete, well-preserved skull has been elusive. DNH 7, an adult cranium and mandible from the Drimolen site, was identified, on the basis of its small size, as a presumptive female of A. robustus. Here, we provide a detailed comparative description of the specimen. In cranial, facial, and dental size, DNH 7 is confirmed to lie at the extreme small end of the A. robustus range of variation, along with a few fragmentary maxillofacial specimens from Swartkrans. In addition, relative to the classically derived craniofacial features of the Swartkrans+Kromdraai portions of the A. robustus hypodigm, primitive anatomy pervades the DNH 7 face, braincase, and cranial base. Taken together, these pieces of evidence place DNH 7 in a previously unfilled position on the robust Australopithecus morphocline, where the specimen highlights the morphological distinctions between southern and eastern African species of this group and epitomizes the anatomy expected of the group's last common ancestor.

New Pliocene hominin remains from the Leado Dido'a area of Woranso-Mille, Ethiopia.

Fossiliferous deposits at Woranso-Mille span the period when Australopithecus anamensis gave rise to Australopithecus afarensis (3.8-3.6 Ma) and encompass the core of the A. afarensis range (ca. 3.5-3.2 Ma). Within the latter period, fossils described to date include the intriguing but taxonomically unattributed Burtele foot, dentognathic fossils attributed to Australopithecus deyiremeda, and one specimen securely attributed to A. afarensis (the Nefuraytu mandible). These fossils suggest that at least one additional hominin lineage lived alongside A. afarensis in the Afar Depression. Here we describe a collection of hominin fossils from a new locality in the Leado Dido'a area of Woranso-Mille (LDD-VP-1). The strata in this area are correlated to the same chron as those in the Burtele area (C2An.3n; 3.59-3.33 Ma), and similar in age to the Maka Sands and the Basal through lower Sidi Hakoma Members of the Hadar Formation. We attribute all but one of the LDD hominin specimens to A. afarensis, based on diagnostic morphology of the mandible, maxilla, canines, and premolars. The LDD specimens generally fall within the range of variation previously documented for A. afarensis but increase the frequency of some rare morphological variants. However, one isolated M3 is extremely small, and its taxonomic affinity is currently unknown. The new observations support previous work on temporal trends in A. afarensis and demonstrate that the large range of variation accepted for this species is present even within a limited spatiotemporal range. The value added with this sample lies in its contribution to controlling for spatiotemporal differences among site samples in the A. afarensis hypodigm and its contemporaneity with non-A. afarensis specimens at Woranso-Mille.