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.

Dental data challenge the ubiquitous presence of Homo in the Cradle of Humankind.

The origins of Homo, as well as the diversity and biogeographic distribution of early Homo species, remain critical outstanding issues in paleoanthropology. Debates about the recognition of early Homo, first appearance dates, and taxonomic diversity within Homo are particularly important for determining the role that southern African taxa may have played in the origins of the genus. The correct identification of Homo remains also has implications for reconstructing phylogenetic relationships between species of Australopithecus and Paranthropus, and the links between early Homo species and Homo erectus. We use microcomputed tomography and landmark-free deformation-based three-dimensional geometric morphometrics to extract taxonomically informative data from the internal structure of postcanine teeth attributed to Early Pleistocene Homo in the southern African hominin-bearing sites of Sterkfontein, Swartkrans, Drimolen, and Kromdraai B. Our results indicate that, from our sample of 23 specimens, only 4 are unambiguously attributed to Homo, 3 of them coming from Swartkrans member 1 (SK 27, SK 847, and SKX 21204) and 1 from Sterkfontein (Sts 9). Three other specimens from Sterkfontein (StW 80 and 81, SE 1508, and StW 669) approximate the Homo condition in terms of overall enamel-dentine junction shape, but retain Australopithecus-like dental traits, and their generic status remains unclear. The other specimens, including SK 15, present a dominant australopith dental signature. In light of these results, previous dietary and ecological interpretations can be reevaluated, showing that the geochemical signal of one tooth from Kromdraai (KB 5223) and two from Swartkrans (SK 96 and SKX 268) is consistent with that of australopiths.

Evidence for habitual climbing in a Pleistocene hominin in South Africa.

Bipedalism is a defining trait of the hominin lineage, associated with a transition from a more arboreal to a more terrestrial environment. While there is debate about when modern human-like bipedalism first appeared in hominins, all known South African hominins show morphological adaptations to bipedalism, suggesting that this was their predominant mode of locomotion. Here we present evidence that hominins preserved in the Sterkfontein Caves practiced two different locomotor repertoires. The trabecular structure of a proximal femur (StW 522) attributed to Australopithecus africanus exhibits a modern human-like bipedal locomotor pattern, while that of a geologically younger specimen (StW 311) attributed to either Homo sp. or Paranthropus robustus exhibits a pattern more similar to nonhuman apes, potentially suggesting regular bouts of both climbing and terrestrial bipedalism. Our results demonstrate distinct morphological differences, linked to behavioral differences between Australopithecus and later hominins in South Africa and contribute to the increasing evidence of locomotor diversity within the hominin clade.

A new absolute date from Swartkrans Cave for the oldest occurrences of Paranthropus robustus and Oldowan stone tools in South Africa.

The Early Pleistocene site of Swartkrans in South Africa's Cradle of Humankind World Heritage Site has been significant for our understanding of the evolution of both early Homo and Paranthropus, as well as the earliest archaeology of southern Africa. Previous attempts to improve a faunal age estimate of the earliest deposit, Member 1, had produced results obtained with uranium-lead dating (U-Pb) on flowstones and cosmogenic burial dating of quartz, which placed the entire member in the range of >1.7/1.8 Ma and <2.3 Ma. In 2014, two simple burial dates for the Lower Bank, the earliest unit within Member 1, narrowed its age to between ca. 1.8 Ma and 2.2 Ma. A new dating program using the isochron method for burial dating has established an absolute age of 2.22 ± 0.09 Ma for a large portion of the Lower Bank, which can now be identified as containing the earliest Oldowan stone tools and fossils of Paranthropus robustus in South Africa. This date agrees within one sigma with the U-Pb age of 2.25 ± 0.08 Ma previously published for the flowstone underlying the Lower Bank and confirms a relatively rapid rate of accumulation for a large portion of the talus.

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.

The long limb bones of the StW 573 Australopithecus skeleton from Sterkfontein Member 2: Descriptions and proportions.

Due to its completeness, the A.L. 288-1 ('Lucy') skeleton has long served as the archetypal bipedal Australopithecus. However, there remains considerable debate about its limb proportions. There are three competing, but not necessarily mutually exclusive, explanations for the high humerofemoral index of A.L. 288-1: (1) a retention of proportions from an Ardipithecus-like chimp/human last common ancestor (CLCA); (2) indication of some degree of climbing ability; (3) allometry. Recent discoveries of other partial skeletons of Australopithecus, such as those of Australopithecus sediba (MH1 and MH2) and Australopithecus afarensis (KSD-VP-1/1 and DIK-1/1), have provided new opportunities to test hypotheses of early hominin body size and limb proportions. Yet, no early hominin is as complete (>90%), as is the ∼3.67 Ma 'Little Foot' (StW 573) skeleton from Sterkfontein Member 2. Here, we provide the first descriptions of its upper and lower long limb bones, as well as a comparative context of its limb proportions. We found that StW 573 possesses absolutely longer limb lengths than A.L. 288-1, but both skeletons show similar limb proportions. This finding seems to argue against a purely allometric explanation for A.L. 288-1 limb proportions. In fact, our multivariate allometric analysis suggests that limb lengths of Australopithecus, as represented by StW 573 and A.L. 288-1, exhibit a significantly different (p < 0.001) allometric pattern than that which typifies modern humans and African apes. Like some previous analyses, our results also suggest that hominin limb evolution occurred in two stages with: first, a modest increase in lower limb length and a concurrent shortening of the antebrachium between Ardipithecus and Australopithecus, followed by a considerable lengthening of the lower limb along with a decrease of both upper limb elements occurring between Australopithecus and Homo sapiens.

A multiscale stratigraphic investigation of the context of StW 573 'Little Foot' and Member 2, Sterkfontein Caves, South Africa.

The Sterkfontein Caves is currently the world's richest Australopithecus-bearing site. Included in Sterkfontein's hominin assemblage is StW 573 ('Little Foot'), a near-complete Australopithecus skeleton discovered in Member 2 in the Silberberg Grotto. Because of its importance to the fossil hominin record, the geological age of StW 573 has been the subject of significant debate. Three main hypotheses have been proposed regarding the formation and age of Member 2 and by association StW 573. The first proposes that Member 2 (as originally defined in the type section in the Silberberg Grotto) started to accumulate at around 2.58 Ma and that the unit is contained within the Silberberg Grotto. The second proposes that Member 2 started forming before 3.67 ± 0.16 Ma and that the deposit extends into the Milner Hall and close to the base of the cave system. The third proposes a 'two-stage burial scenario', in which some sediments and StW 573 represent a secondary and mixed-age accumulation reworked from a higher cave. The stratigraphic and sedimentological implications of these hypotheses are tested here through the application of a multiscale investigation of Member 2, with reference to the taphonomy of the StW 573 skeleton. The complete infilling sequence of Member 2 is described across all exposures of the deposit in the Silberberg Grotto and into the Milner Hall. Sediments are generally stratified and conformably deposited in a sequence of silty sands eroded from well-developed lateritic soils on the landscape surface. Voids, clasts and bioclasts are organized consistently across and through Member 2 conforming with the underlying deposit geometry, indicating gradual deposit accretion with no distinct collapse facies evident and only localized intra-unit postdepositional modification. The stratigraphy and sedimentology of Member 2 support a simple single-stage accumulation process of Member 2 and a primary association between the sediments of Member 2 and the StW 573 'Little Foot' skeleton.

The bony labyrinth of StW 573 ("Little Foot"): Implications for early hominin evolution and paleobiology.

Because of its exceptional degree of preservation and its geological age of ∼3.67 Ma, StW 573 makes an invaluable contribution to our understanding of early hominin evolution and paleobiology. The morphology of the bony labyrinth has the potential to provide information about extinct primate taxonomic diversity, phylogenetic relationships and locomotor behaviour. In this context, we virtually reconstruct and comparatively assess the bony labyrinth morphology in StW 573. As comparative material, we investigate 17 southern African hominin specimens from Sterkfontein, Swartkrans and Makapansgat (plus published data from two specimens from Kromdraai B), attributed to Australopithecus, early Homo or Paranthropus, as well as 10 extant human and 10 extant chimpanzee specimens. We apply a landmark-based geometric morphometric method for quantitatively assessing labyrinthine morphology. Morphology of the inner ear in StW 573 most closely resembles that of another Australopithecus individual from Sterkfontein, StW 578, recovered from the Jacovec Cavern. Within the limits of our sample, we observe a certain degree of morphological variation in the Australopithecus assemblage of Sterkfontein Member 4. Cochlear morphology in StW 573 is similar to that of other Australopithecus as well as to Paranthropus specimens included in this study, but it is substantially different from early Homo. Interestingly, the configuration of semicircular canals in Paranthropus specimens from Swartkrans differs from other fossil hominins, including StW 573. Given the role of the cochlea in the sensory-driven interactions with the surrounding environment, our results offer new perspectives for interpreting early hominin behaviour and ecology. Finally, our study provides additional evidence for discussing the phylogenetic polarity of labyrinthine traits in southern African hominins.

The endocast of StW 573 ("Little Foot") and hominin brain evolution.

One of the most crucial debates in human paleoneurology concerns the timing and mode of the emergence of the derived cerebral features in the hominin fossil record. Given its exceptional degree of preservation and geological age (i.e., 3.67 Ma), StW 573 ('Little Foot') has the potential to shed new light on hominin brain evolution. Here we present the first detailed comparative description of the external neuroanatomy of StW 573. The endocast was virtually reconstructed and compared to ten southern African hominin specimens from Makapansgat, Malapa, Sterkfontein and Swartkrans attributed to Australopithecus and Paranthropus. We apply an automatic method for the detection of sulcal and vascular imprints. The endocranial surface of StW 573 is crushed and plastically deformed in a number of locations. The uncorrected and therefore minimum cranial capacity estimate is 408 cm3 and plots at the lower end of Australopithecus variation. The endocast of StW 573 approximates the rostrocaudally elongated and dorsoventrally flattened endocranial shape seen in Australopithecus and displays a distinct left occipital petalia. StW 573 and the comparative early hominin specimens share a similar sulcal pattern in the inferior region of the frontal lobes that also resembles the pattern observed in extant chimpanzees. The presumed lunate sulcus in StW 573 is located above the sigmoid sinus, as in extant chimpanzees, while it is more caudally positioned in SK 1585 and StW 505. The middle branch of the middle meningeal vessels derives from the anterior branch, as in MH 1, MLD 37/38, StW 578. Overall, the cortical anatomy of StW 573 displays a less derived condition compared to the late Pliocene/early Pleistocene southern African hominins (e.g., StW 505, SK 1585).

Hominin vertebrae and upper limb bone fossils from Sterkfontein Caves, South Africa (1998-2003 excavations).

OBJECTIVES: We provide descriptions and functional interpretations of 11 >2.0 Ma hominin vertebral and upper limb fossils from Sterkfontein. MATERIALS AND METHODS: We employed taphonomic methods to describe postmortem damage observed on the fossils. We used osteometric tools and measurements to generate quantitative descriptions, which were added to qualitative descriptions of the fossils. These observations were then interpreted using published data on the same skeletal elements from extant and extinct hominoid taxa. RESULTS: Six of the fossils carry carnivore tooth marks. Two vertebrae show morphologies that are consistent with fully developed lordosis of the lumbar spine, but which are not completely consistent with bipedal loading of the same intensity and/or frequency as reflected in the lumbars of modern humans. A clavicle shows a combination of humanlike and apelike features, the latter of which would have endowed its hominin with good climbing abilities. When combined, analyses of fragmentary radius and ulna fossils yield more ambiguous results. DISCUSSION: The new fossil collection presents a mix of bipedal and climbing features. It is unclear whether this mix indicates that all Sterkfontein hominins of >2.0 Ma were terrestrial bipeds who retained adaptations for climbing or whether the collection samples two differently adapted, coeval hominins, Australopithecus africanus and Australopithecus prometheus, both of which are represented at Sterkfontein by skull remains. Regardless, the significant frequency of tooth-marked fossils in the sample might indicate that predation was a selection pressure that maintained climbing adaptations in at least some Sterkfontein hominins of this period.

Hominin hand bone fossils from Sterkfontein Caves, South Africa (1998-2003 excavations).

We describe eleven hominin metacarpals and phalanges recovered from Jacovec Cavern and Member 4 of the Sterkfontein Formation between 1998 and 2003. Collectively, the fossils date in excess of 2.0 Ma, and are probably attributable to Australopithecus africanus and/or Australopithecus prometheus. When combined with results of previous studies on Australopithecus postcranial functional morphology, the new data presented here suggest that at least some late Pliocene and/or early Pleistocene hominins from Sterkfontein were arboreally adept. This finding accords with the reconstruction of the site's >2.0 Ma catchment area as well-vegetated and containing significant woody components. In addition, most of the new specimens described here evince morphologies that indicate the hands from which they derived lacked complete modern humanlike manual dexterity, which is integral to the manufacture and use of intentionally shaped stone tools. The absence of lithic artifacts from both stratigraphic units from which the fossils were excavated is consistent with this conclusion.

RETRACTED: The revised stratigraphy of the hominin-bearing site of Kromdraai (Gauteng, South Africa) and associated perspectives.

This article has been retracted: please see Elsevier Policy on Article Withdrawal (http://www.elsevier.com/locate/withdrawalpolicy). This article has been retracted at the request of the co-Editors-in-Chief and the authors. The Results and Discussion of this article duplicate significant parts of book chapter "A revised stratigraphy of Kromdraai", published by L.B., R.M., R.C., F.T. and J.B. in Braga, J. and Thackeray, J.F. (Eds.), "Kromdraai. A Birthplace of Paranthropus in the Cradle of Humankind" (2016, SUN MeDIA MeTRO, pp. 31-47), https://doi.org/10.18820/9781928355076. One of the conditions of submission of a paper to Journal of Human Evolution is that authors declare explicitly that that their work is original and has not been published previously. Reuse of any data should be appropriately cited. As such this article represents an abuse of the scientific publishing system. The scientific community takes a very strong view on this matter and apologies are offered to readers of the journal that this was not detected during the submission process.

Cranial vault thickness variation and inner structural organization in the StW 578 hominin cranium from Jacovec Cavern, South Africa.

The Sterkfontein Caves site is one of the richest early hominin fossil localities in Africa. More specifically, the fossiliferous deposits within the lower-lying Jacovec Cavern have yielded valuable hominin remains; prominent among them is the Australopithecus partial cranium StW 578. Due to the fragmentary nature of the braincase, the specimen has not yet been formally assigned to a species. In this context, we employ microtomography to quantify cranial thickness and composition of StW 578 in order to assess its taxonomic affinity. As comparative material, we investigate 10 South African hominin cranial specimens from Sterkfontein (StW 505, Sts 5, Sts 25, Sts 71), Swartkrans (SK 46, SK 48, SK 49) and Makapansgat (MLD 1, MLD 10, MLD 37/38), attributed to either Australopithecus or Paranthropus, as well as 10 extant human and 10 extant chimpanzee crania. Thickness variation in and structural arrangement of the inner and outer cortical tables and the diploë are automatically assessed at regular intervals along one parasagittal and one coronal section. Additionally, topographic cranial vault thickness distribution is visualized using color maps. Comparisons highlight an absolutely and relatively thickened condition of the StW 578 cranial vault versus those of other South African Plio-Pleistocene hominins. Moreover, in StW 578, as well as in the Australopithecus specimens Sts 5 and Sts 71 from Sterkfontein, the diploic layer contributes substantially to cumulative vault thickness (i.e., >60%). Within the comparative sample investigated here, StW 505 and Sts 71 from Sterkfontein Member 4, both attributed to Australopithecus, most closely resemble StW 578 in terms of cranial vault thickness values, tissue proportions, and two- and three-dimensional distributions. Including additional Plio-Pleistocene Australopithecus and Paranthropus crania from South and East Africa in future studies would further help establish morphological variability in these hominin taxa.

Stratigraphic analysis of the Sterkfontein StW 573 Australopithecus skeleton and implications for its age.

StW 573, Little Foot, is the most complete Australopithecus skeleton yet discovered, with many of its bones found in their correct anatomical position. Since the discovery of the in situ skeleton in the Silberberg Grotto in 1997, several teams have attempted to date the fossil. This appeared a simple process because several flowstones are inter-bedded in the breccia above and below StW 573. Dating of these flowstones, using U-Pb (uranium-lead) isotope decay techniques, gave younger results than expected from the fauna and stratigraphic position, around 2.2 Ma (millions of years). Our recent stratigraphic, micromorphological and geochemical studies revealed that the stratigraphy is much more complicated than was previously thought, with localized post-depositional processes leading to the creation of voids within the breccia around the skeleton. These voids were then filled by multiple generations of flowstone growth. The research we present here demonstrates that the proposed dates based on the flowstone deposition can give only a minimum age for StW 573 and that the flowstone formation came after, and probably long after, the breccia deposition. If one takes account of the long evolution of these karst fillings, StW 573 appears to be significantly older than 2.2 Ma.

Frontal sinuses and human evolution.

The frontal sinuses are cavities inside the frontal bone located at the junction between the face and the cranial vault and close to the brain. Despite a long history of study, understanding of their origin and variation through evolution is limited. This work compares most hominin species' holotypes and other key individuals with extant hominids. It provides a unique and valuable perspective of the variation in sinuses position, shape, and dimensions based on a simple and reproducible methodology. We also observed a covariation between the size and shape of the sinuses and the underlying frontal lobes in hominin species from at least the appearance of Homo erectus. Our results additionally undermine hypotheses stating that hominin frontal sinuses were directly affected by biomechanical constraints resulting from either chewing or adaptation to climate. Last, we demonstrate their substantial potential for discussions of the evolutionary relationships between hominin species.