Spatial sampling bias influences our understanding of early hominin evolution in eastern Africa.

The eastern branch of the Eastern African Rift System (EARS) is the source of a large proportion of the early hominin fossil record, but it covers a tiny fraction (ca. 1%) of the continent. Here we investigate how this mismatch between where fossils are preserved and where hominins probably lived may influence our ability to understand early hominin evolution, using extant mammals as analogues. We show that the eastern branch of the EARS is not an environmentally representative sample of the full species range for nearly all extant rift-dwelling mammals. Likewise, when we investigate published morphometric datasets for extant cercopithecine primates, evidence from the eastern branch alone fails to capture major portions of continental-scale cercopithecine cranial morphospace. We suggest that extant rift-dwelling species should be used as analogues to place confidence intervals on hominin habitat reconstructions. Furthermore, given the north-south orientation of the eastern branch of the EARS, morphoclines that are not aligned along this major north-south axis are likely to be poorly sampled by sites in the eastern branch. There is a pressing need for research on the geography of early hominin morphoclines to estimate how morphologically representative the hominin fossil sample from the eastern branch may be.

What we know and do not know after the first decade of Homo naledi.

It has been just over 10 years since the first fossils attributed to Homo naledi were recovered from the Rising Star Cave system in South Africa's Cradle of Humankind. The hominin fossil evidence for H. naledi displays a distinctive combination of primitive and derived morphology, yet for a time-averaged fossil sample it is remarkable for its relatively low level of variation. Thus-unusually for palaeoanthropology-there has been little pushback against the decision to recognize a single novel taxon for all of the material recovered from the Rising Star Cave system. However, almost everything else claimed about H. naledi-its age, burial context and behaviour-has been controversial. Here we examine the strength of the evidence for these claims.

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.

On the scientific credibility of paleoanthropology: Reply to Villmoare and Kimbel.

Smith and Wood reply to Villmoare and Kimbel regarding the scientific credibility of problems in paleoanthropology that require causal explanations for unique historical events.

Behavioral and phylogenetic correlates of limb length proportions in extant apes and monkeys: Implications for interpreting hominin fossils.

The body proportions of extant animals help inform inferences about the behaviors of their extinct relatives, but relationships between body proportions, behavior, and phylogeny in extant primates remain unclear. Advances in behavioral data, molecular phylogenies, and multivariate analytical tools make it an opportune time to perform comprehensive comparative analyses of primate traditional limb length proportions (e.g., intermembral, humerofemoral, brachial, and crural indices), body size-adjusted long bone proportions, and principal components. In this study we used a mix of newly-collected and published data to investigate whether and how the limb length proportions of a diverse sample of primates, including monkeys, apes, and modern humans, are influenced by behavior and phylogeny. We reconfirm that the intermembral index, followed by the first principal component of traditional limb length proportions, is the single most effective variable distinguishing hominoids and other anthropoids. Combined limb length proportions and positional behaviors are strongly correlated in extant anthropoid groups, but phylogeny is a better predictor of limb length proportion variation than of behavior. We confirm convergences between members of the Atelidae and extant apes (especially Pan), members of the Hylobatidae and Pongo, and a potential divergence of Presbytis limb proportions from some other cercopithecoids, which correlate with adaptations for forelimb-dominated behaviors in some colobines. Collectively, these results substantiate hypotheses indicating that extinct hominins and other hominoid taxa can be distinguished by analyzing combinations of their limb length proportions at different taxonomic levels. From these results, we hypothesize that fossil skeletons characterized by notably disparate limb length proportions are unlikely to have exhibited similar behavioral patterns.

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.

Hominin fossils from Kromdraai and Drimolen inform Paranthropus robustus craniofacial ontogeny.

Ontogeny provides critical information about the evolutionary history of early hominin adult morphology. We describe fossils from the southern African sites of Kromdraai and Drimolen that provide insights into early craniofacial development in the Pleistocene robust australopith Paranthropus robustus. We show that while most distinctive robust craniofacial features appear relatively late in ontogeny, a few do not. We also find unexpected evidence of independence in the growth of the premaxillary and maxillary regions. Differential growth results in a proportionately larger and more postero-inferiorly rotated cerebral fossa in P. robustus infants than in the developmentally older Australopithecus africanus juvenile from Taung. The accumulated evidence from these fossils suggests that the iconic SK 54 juvenile calvaria is more likely early Homo than Paranthropus. It is also consistent with the hypothesis that P. robustus is more closely related to Homo than to A. africanus.

Underappreciated pioneers.

This contribution focuses on a 1928 multiauthor paper reporting the discovery of a child's skull at Devil's Tower cave on the Rock of Gibraltar. It was ground-breaking. Two of the lead authors, Dorothy Garrod and Dorothea Bate, were women, and it was one of the earliest reports of a fossil hominin to incorporate and integrate detailed information about its stratigraphic and environmental context.

Distinguishing primate taxa with enamel incremental variables.

Enamel has long been of interest for its functional and phylogenetic significance among fossil hominins and other primates. Previous studies demonstrated that enamel incremental features distinguish among hominin fossil taxa, suggesting utility for highlighting taxonomy. However, not all features appear to be useful in mixed samples of fossils, living humans, and apes. Here we tested enamel incremental data from closely related primate taxa to determine which features, if any, distinguish among them. Enamel incremental variables were measured from the M2 of 40 living primate taxa, and we tested our variables using discriminant function analysis at the taxonomic ranks of parvorder, family, tribe, and genus. We then included enamel incremental data from Australopithecus afarensis, Australopithecus africanus, Paranthropus aethiopicus, Paranthropus boisei, and Paranthropus robustus to determine if these features distinguished fossil taxa from living humans and apes. Our initial results show that enamel incremental variables distinguish among primate taxa, but with low classification rates. Further testing with jackknifing methods shows overlap between groups at all taxonomic ranks, suggesting enamel incremental variables are unreliable for taxonomy. The addition of many common enamel incremental growth variables also resulted in multicollinearity in our multivariate analysis. As the dentition and isolated teeth remain a significant portion of the hominin fossil record, verifying enamel incremental features as a useful taxonomic tool is fundamentally important for hominin paleobiology.

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.

Birth of Australopithecus.

The announcement of a fossilized child's skull discovered in a quarry in 1924 sub-Saharan Africa might not have seemed destined to be a classic paper. This contribution focuses on anatomist Raymond Dart's 1925 paper in which he designated the Taungs skull the type specimen of Australopithecus africanus. We combine an account of Dart's training and experience, with a telling of the fossil's discovery, analysis, the initial response of a mostly skeptical community, and a review of subsequent discoveries that consolidated the case Dart made for a hitherto unknown human close relative. Dart's paper presented evidence that confirmed the prescience of Charles Darwin's prediction that Africa was the birthplace of modern humans. The Taungs skull's unique mix of great ape and human attributes eventually led to a paradigm shift in our understanding of human evolution.

Rethinking the ecological drivers of hominin evolution.

A central goal of paleoanthropology is understanding the role of ecological change in hominin evolution. Over the past several decades researchers have expanded the hominin fossil record and assembled detailed late Cenozoic paleoclimatic, paleoenvironmental, and paleoecological archives. However, effective use of these data is precluded by the limitations of pattern-matching strategies for inferring causal relationships between ecological and evolutionary change. We examine several obstacles that have hindered progress, and highlight recent research that is addressing them by (i) confronting an incomplete fossil record, (ii) contending with datasets spanning varied spatiotemporal scales, and (iii) using theoretical frameworks to build stronger inferences. Expanding on this work promises to transform challenges into opportunities and set the stage for a new phase of paleoanthropological research.

Birth of Homo erectus.

Eugène Dubois was the pioneer of human origins research in South-East Asia, specifically on two of the islands, Sumatra and Java, now included in Indonesia. Dubois was a polymath, whose research interests embraced encephalization and hydrology as well as paleoanthropology. His interpretations of the hominin fossil evidence he collected, which he eventually assigned to Pithecanthropus erectus, changed over the years, and he evidently felt defensive about those interpretations, but in his 1894 paper he presents cogent reasons for his decision. The taxon he introduced is still recognized, and while it is no longer seen as "the" link between fossil apes and modern humans, it is currently one of the longest surviving hominin taxa.

Nature and relationships of Sahelanthropus tchadensis.

A partial left femur (TM 266-01-063) was recovered in July 2001 at Toros-Menalla, Chad, at the same fossiliferous location as the late Miocene holotype of Sahelanthropus tchadensis (the cranium TM 266-01-060-1). It was recognized as a probable primate femur in 2004 when one of the authors was undertaking a taphonomic survey of the fossil assemblages from Toros-Menalla. We are confident the TM 266 femoral shaft belongs to a hominid. It could sample a hominid hitherto unrepresented at Toros-Menalla, but a more parsimonious working hypothesis is that it belongs to S. tchadensis. The differences between TM 266 and the late Miocene Orrorin tugenensis partial femur BAR 1002'00, from Kenya, are consistent with maintaining at least a species-level distinction between S. tchadensis and O. tugenensis. The results of our preliminary functional analysis suggest the TM 266 femoral shaft belongs to an individual that was not habitually bipedal, something that should be taken into account when considering the relationships of S. tchadensis. The circumstances of its discovery should encourage researchers to check to see whether there is more postcranial evidence of S. tchadensis among the fossils recovered from Toros-Menalla.

Mandibular corpus shape is a taxonomic indicator in extant hominids.

OBJECTIVES: The aim of this study is to understand whether the shape of three sub-regions of the mandibular corpus (the alveolar arch, corpus at M1 and posterior symphysis) are useful for making taxonomic assessments at the genus and species levels in extant hominids. MATERIALS AND METHODS: We use data taken from 3D surface scans of the mandibular corpus of seven extant hominid taxa: Gorilla gorilla gorilla, Gorilla beringei graueri, Homo sapiens, Pan paniscus, Pan troglodytes schweinfurthii, Pongo abelii, and Pongo pygmaeus pygmaeus to generate four shape variables: alveolar arch shape (AAS), corpus shape at M1 (CSM1 ), posterior symphysis shape at the midline (PSSM), and posterior symphysis shape (PSS). To ascertain how reliable each mandibular shape variable is for assessing taxonomy, we ran canonical discriminant and discriminant function analysis, reporting cross-validated results. RESULTS: Using a combination of three mandibular corpus shape variables, 99% of specimens were classified correctly for genus-level analyses. A maximum of 100% of Pan specimens, 94% of Gorilla specimens and 96% of Pongo specimens were classified correctly at the species level when up to three mandibular shape variables were included in the analyses. When mandibular corpus variables were considered in isolation, posterior symphysis shape yielded the highest overall correct classification results. DISCUSSION: The high taxonomic classification rates at both the genus and species level, using 3D surface data and advanced quantification techniques, show that the shape of the alveolar arch, corpus at M1 and symphysis can distinguish extant hominid taxa. These findings have implications for assessing the taxonomy of extinct hominid specimens which preserve these mandibular sub-regions.