StW 573 Australopithecus prometheus: Its Significance for an Australopith Bauplan.

The StW 573 skeleton of Australopithecus prometheus from Sterkfontein Member 2 is some 93% complete and thus by far the most complete member of that genus yet found. Firmly dated at 3.67 Ma, it is one of the earliest specimens of its genus. A crucial aspect of interpretation of locomotor behaviour from fossil remains is an understanding of the palaeoenvironment in which the individual lived and the manner in which it would have used it. While the value of this ecomorphological approach is largely accepted, it has not been widely used as a stable framework on which to build evolutionary biomechanical interpretations. Here, we collate the available evidence on StW 573's anatomy in order, as far as currently possible, to reconstruct what might have been this individual's realized and potential niche. We explore the concept of a common Australopithecus "bauplan" by comparing the morphology and ecological context of StW 573 to that of paenocontemporaneous australopiths including Australopithecus anamensis and KSD-VP-1/1 Australopithecus afarensis. Each was probably substantially arboreal and woodland-dwelling, relying substantially on arboreal resources. We use a hypothesis-driven approach, tested by: virtual experiments, in the case of extinct species; biomechanical analyses of the locomotor behaviour of living great ape species; and analogical experiments with human subjects. From these, we conclude that the habitual locomotor mode of all australopiths was upright bipedalism, whether on the ground or on branches. Some later australopiths such as Australopithecus sediba undoubtedly became more terrestrial, allowing sacrifice of arboreal stability in favour of manual dexterity. Indeed, modern humans retain arboreal climbing skills but have further sacrificed arboreal effectiveness for enhanced ability to sustain striding terrestrial bipedalism over much greater distances. We compare StW 573's locomotor adaptations to those of living great apes and protohominins, and agree with those earlier observers who suggest that the common panin-hominin last common ancestor was postcranially more like Gorilla than Pan.

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.

Trabecular organization of the proximal femur in Paranthropus robustus: Implications for the assessment of its hip joint loading conditions.

Reconstruction of the locomotor repertoire of the australopiths (Australopithecus and Paranthropus) has progressively integrated information from the mechanosensitive internal structure of the appendicular skeleton. Recent investigations showed that the arrangement of the trabecular network at the femoral head center is biomechanically compatible with the pattern of cortical bone distribution across the neck, both suggesting a full commitment to bipedalism in australopiths, but associated with a slightly altered gait kinematics compared to Homo involving more lateral deviation of the body center of mass over the stance limb. To provide a global picture in Paranthropus robustus of the trabecular architecture of the proximal femur across the head, neck and greater trochanter compartments, we applied techniques of virtual imaging to the variably preserved Early Pleistocene specimens SK 82, SK 97, SK 3121, SKW 19 and SWT1/LB-2 from the cave site of Swartkrans, South Africa. We also assessed the coherence between the structural signals from the center of the head and those from the trabecular network of the inferolateral portion of the head and the inferior margin of the neck, sampling the so-called vertical bundle, which in humans represents the principal compressive system of the joint. Our analyses show a functionally related trabecular organization in Pa. robustus that closely resembles the extant human condition, but which also includes some specificities in local textural arrangement. The network of the inferolateral portion of the head shows a humanlike degree of anisotropy and a bone volume fraction intermediate between the extant human and the African ape patterns. These results suggest slight differences in gait kinematics between Pa. robustus and extant humans. The neck portion of the vertical bundle revealed a less biomechanically sensitive signal. Future investigations on the australopith hip joint loading environment should more carefully investigate the trabecular structure of the trochanteric region and possible structural covariation between cortical bone distribution across the neck and site-specific trabecular properties of the arcuate bundle.

Cortical bone distribution in the femoral neck of Paranthropus robustus.

Studies of the australopith (Australopithecus and Paranthropus) proximal femur have increasingly integrated information from the local arrangement of the cortical and cancellous bone to allow functional-biomechanical inferences on the locomotor behavioral patterns. In Australopithecus africanus and Paranthropus robustus, the cancellous bone organization at the center of the femoral head shows principal strut orientation similar to that of fossil and recent humans, which indicates that australopiths were human-like in many aspects of their bipedalism. However, by combining outer morphology with superoinferior asymmetry in cortical bone thickness at the base of neck and mid-neck, it has been suggested that, while adapted for terrestrial bipedality, australopiths displayed a slightly altered gait kinematics compared to Homo. We used techniques of 2D and 3D virtual imaging applied to an X-ray microtomographic record to assess cortical bone distribution along the entire femoral neck compartment in four upper femora from Swartkrans, South Africa (SK 82, SK 97, SK 3121, and SWT1/LB-2) and compared the results to the extant human and chimpanzee conditions. Our results support and extend previous evidence for more symmetric superior and inferior femoral neck cortical thicknesses in P. robustus than in modern humans and show that the differences are even greater than previously reported. However, P. robustus and humans still share a trend of lateral-to-medial decrease in asymmetry of the superior/inferior cortical thickness ratio, while this pattern is reversed in chimpanzees. We also identified two features uniquely characterizing P. robustus: an accentuated contrast between the relatively thicker anterior and the thinner posterior walls, and a more marked lateral-to-medial thinning of both cortices compared to extant humans and chimpanzees, which indicate wider interspecific differences among hominids in structural organization of the proximal femur than previously reported. It remains to be ascertained if, and to what extent, these features also characterize the femoral neck of Australopithecus.

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.

Chimpanzee extractive foraging with excavating tools: Experimental modeling of the origins of human technology.

It is hypothesized that tool-assisted excavation of plant underground storage organs (USOs) played an adaptive role in hominin evolution and was also once considered a uniquely human behavior. Recent data indicate that savanna chimpanzees also use tools to excavate edible USOs. However, those chimpanzees remain largely unhabituated and we lack direct observations of this behavior in the wild. To fill this gap in our knowledge of hominoid USO extractive foraging, we conducted tool-mediated excavation experiments with captive chimpanzees naïve to this behavior. We presented the chimpanzees with the opportunity to use tools in order to excavate artificially-placed underground foods in their naturally forested outdoor enclosure. No guidance or demonstration was given to the chimpanzees at any time. The chimpanzees used tools spontaneously in order to excavate the underground foods. They exhibited six different tool use behaviors in the context of excavation: probe, perforate, dig, pound, enlarge and shovel. However, they still excavated manually more often than they did with tools. Chimpanzees were selective in their choice of tools that we provided, preferring longer tools for excavation. They also obtained their own tools mainly from naturally occurring vegetation and transported them to the excavation site. They reused some tools throughout the study. Our new data provide a direction for the study of variables relevant to modeling USO extractive foraging by early hominins.

Ages-at-death distribution of the early Pleistocene hominin fossil assemblage from Drimolen (South Africa).

OBJECTIVES: A prevailing hypothesis in paleoanthropology is that early Pleistocene hominin bones were accumulated in South African caves by carnivores, which used those shelters, and the trees surrounding them, as refuge and feeding sites. We tested this hypothesis at the site of Drimolen, by comparing its hominin age-at-death distribution to that of the nearby and roughly contemporaneous site of Swartkrans. MATERIALS AND METHODS: We employed standard dental aging systems in order to categorize the Drimolen hominin teeth into age classes of 5 years each. We then compared the age-at-death distribution for Drimolen with the published data available for the Swartkrans hominins. RESULTS: Age-at-death distributions indicate that the age category "young adults" is the best represented age category at Swartkrans and the most poorly represented one at Drimolen. Moreover, Drimolen has a preponderance of infant specimens. Both sites have a low frequency of old adult specimens. CONCLUSIONS: Differences observed in frequencies of the age-at-death categories suggest different mechanisms of hominin skeletal accumulation at Drimolen and Swartkrans. Swartkrans' frequency curve reflects mortality in a population subjected to predation and is thus consistent with the carnivore-accumulating hypothesis. In contrast, the Drimolen curve is similar to that of wild populations of living apes. Living primates have been observed exploiting caves as sleeping shelters, for nutritional, security, drinking, and thermoregulatory purposes. We suggest that similar cave use by Pleistocene hominins can explain, in large part, the accumulation of hominin bones at Drimolen. Such a conclusion is another illustration of the growing awareness that a "one-size-fits-all" taphonomic model for South African early Pleistocene hominin sites is probably insufficient.

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.

The Oldowan industry from Swartkrans cave, South Africa, and its relevance for the African Oldowan.

The oldest recognized artifacts at the Swartkrans cave hominid-bearing site in South Africa have long been known to occur in the Lower Bank of Member 1, now dated with the cosmogenic nuclide burial method to ca. 1.8-2.19 Ma. However, the affinities of this industry have been debated due to small sample size. In this paper we present newly excavated material from the Lower Bank retrieved since 2005 in the Swartkrans Paleoanthropological Research Project. The sample is now large enough to confirm its affinity with the Oldowan industrial complex. The assemblage is highly expedient and core reduction strategies are largely casual. Although freehand flaking is present, the bipolar technique is most significant, even in non-quartz raw materials. The Swartkrans assemblage shows some significant contrasts with the Sterkfontein Oldowan, ca. 2.18 Ma, which can be explained by its closer proximity to raw material sources, its somewhat different geographic context, and its more expedient nature. The Swartkrans Oldowan now provides us with the first good indication of Oldowan variability in southern Africa, where only two sizeable assemblages have thus far been discovered. Comparisons are made with other sites across Africa that help to place this variability within our overall understanding of the Oldowan industrial complex.

Hominin skeletal part abundances and claims of deliberate disposal of corpses in the Middle Pleistocene.

Humans are set apart from other organisms by the realization of their own mortality. Thus, determining the prehistoric emergence of this capacity is of significant interest to understanding the uniqueness of the human animal. Tracing that capacity chronologically is possible through archaeological investigations that focus on physical markers that reflect "mortality salience." Among these markers is the deliberate and culturally mediated disposal of corpses. Some Neandertal bone assemblages are among the earliest reasonable claims for the deliberate disposal of hominins, but even these are vigorously debated. More dramatic assertions center on the Middle Pleistocene sites of Sima de los Huesos (SH, Spain) and the Dinaledi Chamber (DC, South Africa), where the remains of multiple hominin individuals were found in deep caves, and under reported taphonomic circumstances that seem to discount the possibility that nonhominin actors and processes contributed to their formation. These claims, with significant implications for charting the evolution of the "human condition," deserve scrutiny. We test these assertions through machine-learning analyses of hominin skeletal part representation in the SH and DC assemblages. Our results indicate that nonanthropogenic agents and abiotic processes cannot yet be ruled out as significant contributors to the ultimate condition of both collections. This finding does not falsify hypotheses of deliberate disposal for the SH and DC corpses, but does indicate that the data also support partially or completely nonanthropogenic formational histories.

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.

New early Pleistocene hominin teeth from the Swartkrans Formation, South Africa.

We describe 14 hominin teeth and tooth fragments excavated recently from Swartkrans Cave (South Africa). The fossils derive from Members 1 (Lower Bank) and 3, from the Member 2/3 interface and from two deposits not yet assigned to member (the "Talus Cone Deposit" and the "Underground North Excavation" [UNE]) of the Swartkrans Formation, and include the first hominin fossil from the UNE, the two smallest Paranthropus robustus deciduous maxillary second molars in the entire hominin fossil record, and one of the smallest P. robustus permanent maxillary second molars from Swartkrans. The small permanent molar is accompanied by another tooth from a different individual but from the same stratigraphic level of the Swartkrans Formation; this second tooth is among, if not, the largest P. robustus permanent maxillary first molars known from anywhere-lending credence to assertions that degrees of body size sexual dimorphism previously ascribed to this species may be underestimated. It is more equivocal whether this evidence also supports hypotheses proposing that P. robustus assemblages from Swartkrans (as well as those from other South African cave sites) formed through the taphonomically biasing actions of large carnivores.

Earliest modern human-like hand bone from a new >1.84-million-year-old site at Olduvai in Tanzania.

Modern humans are characterized by specialized hand morphology that is associated with advanced manipulative skills. Thus, there is important debate in paleoanthropology about the possible cause-effect relationship of this modern human-like (MHL) hand anatomy, its associated grips and the invention and use of stone tools by early hominins. Here we describe and analyse Olduvai Hominin (OH) 86, a manual proximal phalanx from the recently discovered >1.84-million-year-old (Ma) Philip Tobias Korongo (PTK) site at Olduvai Gorge (Tanzania). OH 86 represents the earliest MHL hand bone in the fossil record, of a size and shape that differs not only from all australopiths, but also from the phalangeal bones of the penecontemporaneous and geographically proximate OH 7 partial hand skeleton (part of the Homo habilis holotype). The discovery of OH 86 suggests that a hominin with a more MHL postcranium co-existed with Paranthropus boisei and Homo habilis at Olduvai during Bed I times.

The "bear" essentials: actualistic research on Ursus arctos arctos in the Spanish Pyrenees and its implications for paleontology and archaeology.

Neotaphonomic studies of large carnivores are used to create models in order to explain the formation of terrestrial vertebrate fossil faunas. The research reported here adds to the growing body of knowledge on the taphonomic consequences of large carnivore behavior in temperate habitats and has important implications for paleontology and archaeology. Using photo- and videotrap data, we were able to describe the consumption of 17 ungulate carcasses by wild brown bears (Ursus arctos arctos) ranging the Spanish Pyrenees. Further, we analyzed the taphonomic impact of these feeding bouts on the bones recovered from those carcasses. The general sequence of consumption that we charted starts with separation of a carcass's trunk; viscera are generally eaten first, followed by musculature of the humerus and femur. Long limb bones are not broken open for marrow extraction. Bears did not transport carcasses or carcass parts from points of feeding and did not disperse bones appreciably (if at all) from their anatomical positions. The general pattern of damage that resulted from bear feeding includes fracturing, peeling, crenulation, tooth pitting and scoring of axial and girdle elements and furrowing of the upper long limb bones. As predicted from observational data, the taphonomic consequences of bear feeding resemble those of other non-durophagus carnivores, such as felids, and are distinct from those of durophagus carnivores, such as hyenids. Our results have paleontological and archaeological relevance. Specifically, they may prove useful in building analogical models for interpreting the formation of fossil faunas for which bears are suspected bone accumulators and/or modifiers. More generally, our comparative statistical analyses draw precise quantitative distinctions between bone damage patterns imparted respectively by durophagus (modelled here primarily by spotted hyenas [Crocuta crocuta] and wolves [Canis lupus]) and non-durophagus (modelled here by brown bears and lions [Panthera leo]) carnivorans.

First partial skeleton of a 1.34-million-year-old Paranthropus boisei from Bed II, Olduvai Gorge, Tanzania.

Recent excavations in Level 4 at BK (Bed II, Olduvai Gorge, Tanzania) have yielded nine hominin teeth, a distal humerus fragment, a proximal radius with much of its shaft, a femur shaft, and a tibia shaft fragment (cataloged collectively as OH 80). Those elements identified more specifically than to simply Hominidae gen. et sp. indet are attributed to Paranthropus boisei. Before this study, incontrovertible P. boisei partial skeletons, for which postcranial remains occurred in association with taxonomically diagnostic craniodental remains, were unknown. Thus, OH 80 stands as the first unambiguous, dentally associated Paranthropus partial skeleton from East Africa. The morphology and size of its constituent parts suggest that the fossils derived from an extremely robust individual who, at 1.338±0.024 Ma (1 sigma), represents one of the most recent occurrences of Paranthropus before its extinction in East Africa.

First records of talon cusps on baboon maxillary incisors argue for standardizing terminology and prompt a hypothesis of their formation.

Dental characters can provide vital clues for understanding intra- and intertaxonomic morphological variation and its underlying genetic and environmental components. However, the unambiguous identification of particular traits and their comparative study is often confounded by lack of consistent terminology in the relevant literature. This difficulty is exacerbated when the etiologies are not completely understood, as is the case with talon cusps. To date, research on talon cusps has focused on modern humans. In many instances, descriptions of talon cusps appear in clinical case studies focusing on their treatment and removal. What is lacking in those discussions, though, is a comparative framework, in which the occurrence of talon cusps in nonhuman primates, and possibly other mammals, is established and understood. Here, we report on a taloned upper central incisor of a wild baboon (Papio hamadryas ursinus) from South Africa. The anomalous incisor of this individual includes an exaggerated accessory cusp diagnosed as a Type II talon. Microcomputed tomographic and radiographic analyses show that the taloned cusp possesses enamel, dentin, and pulp. In addition, we identified an unclassifiable talon cusp on a central maxillary incisor of a baboon skull housed in the Smithsonian Institution's Natural History Museum collection. Our observations of talon cusps on baboon incisors demonstrate that, with regard to this phenomenon, systematic study of nonhuman primates is much needed, along with a consistent use of terminology in the anatomical and anthropological literature. Finally, we present a hypothesis of the formation of talon cusps on mammalian incisors.