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.

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.

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.

Earth's volatile contents established by melting and vaporization.

The silicate Earth is strongly depleted in moderately volatile elements (such as lead, zinc, indium and alkali elements) relative to CI chondrites, the meteorites that compositionally most closely resemble the Sun. This depletion may be explained qualitatively by accretion of 10 to 20 per cent of a volatile-rich body to a reduced volatile-free proto-Earth, followed by partial extraction of some elements to the core. However, there are several unanswered questions regarding the sources of Earth's volatiles, notably the overabundance of indium in the silicate Earth. Here we examine the melting processes that occurred during accretion on Earth and precursor bodies and report vaporization experiments under conditions of fixed temperature and oxygen fugacity. We find that the pattern of volatile element depletion in the silicate Earth is consistent with partial melting and vaporization rather than with simple accretion of a volatile-rich chondrite-like body. We argue that melting and vaporization on precursor bodies and possibly during the giant Moon-forming impact were responsible for establishing the observed abundances of moderately volatile elements in Earth.

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.

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.

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.

A Mercury-like component of early Earth yields uranium in the core and high mantle (142)Nd.

Recent (142)Nd isotope data indicate that the silicate Earth (its crust plus the mantle) has a samarium to neodymium elemental ratio (Sm/Nd) that is greater than that of the supposed chondritic building blocks of the planet. This elevated Sm/Nd has been ascribed either to a 'hidden' reservoir in the Earth or to loss of an early-formed terrestrial crust by impact ablation. Since removal of crust by ablation would also remove the heat-producing elements--potassium, uranium and thorium--such removal would make it extremely difficult to balance terrestrial heat production with the observed heat flow. In the 'hidden' reservoir alternative, a complementary low-Sm/Nd layer is usually considered to reside unobserved in the silicate lower mantle. We have previously shown, however, that the core is a likely reservoir for some lithophile elements such as niobium. We therefore address the question of whether core formation could have fractionated Nd from Sm and also acted as a sink for heat-producing elements. We show here that addition of a reduced Mercury-like body (or, alternatively, an enstatite-chondrite-like body) rich in sulfur to the early Earth would generate a superchondritic Sm/Nd in the mantle and an (142)Nd/(144)Nd anomaly of approximately +14 parts per million relative to chondrite. In addition, the sulfur-rich core would partition uranium strongly and thorium slightly, supplying a substantial part of the 'missing' heat source for the geodynamo.

Statistical estimates of hominin origination and extinction dates: A case study examining the Australopithecus anamensis-afarensis lineage.

Reliable estimates of when hominin taxa originated and went extinct are central to addressing many paleoanthropological questions, including those relating to macroevolutionary patterns. The timing of hominin temporal ranges can be used to test chronological predictions generated from phylogenetic hypotheses. For example, hypotheses of phyletic ancestor-descendant relationships, based on morphological data, predict no temporal range overlap between the two taxa. However, a fossil taxon's observed temporal range is almost certainly underestimated due to the incompleteness of both the fossil record itself and its sampling, and this decreases the likelihood of observing temporal overlap. Here, we focus on a well-known and widely accepted early hominin lineage, Australopithecus anamensis-afarensis, and place 95% confidence intervals (CIs) on its origination and extinction dates. We do so to assess whether its temporal range is consistent with it being a phyletic descendant of Ardipithecus ramidus and/or a direct ancestor to the earliest claimed representative of Homo (i.e., Ledi-Geraru). We find that the last appearance of Ar. ramidus falls within the origination CI of Au. anamensis-afarensis, whereas the claimed first appearance of Homo postdates the extinction CI. These results are consistent with Homo evolving from Au. anamensis-afarensis, but temporal overlap between Ar. ramidus and Au. anamensis-afarensis cannot be rejected at this time. Though additional samples are needed, future research should extend our initial analyses to incorporate the uncertainties surrounding the range endpoints of Ar. ramidus and earliest Homo. Overall, our findings demonstrate the need for quantifying the uncertainty surrounding the appearances and disappearances of hominin taxa in order to better understand the timing of evolutionary events in our clade's history.

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.

Birth of primate comparative anatomy.

In 1698, a creature with a perplexing mix of human and "ape" features died in London. Brought back to England by merchants who had acquired it during a trading mission to West Africa, it attracted the attention of the Royal Society, and after the death of what we now know was a juvenile chimpanzee, Edward Tyson, a distinguished physician/anatomist, was commissioned to undertake its dissection. Tyson, who was assisted by William Cowper, prepared a detailed written and graphic description of their meticulous dissection, and this forms the major part of his 1699 publication Orang-outang sive Homo sylvestris: or The Anatomy of a Pygmie compared with that of a Monkey, an Ape, and a Man. Tyson records the many ways his "pygmie" resembled, and differed from, modern humans, including acute assessments of its brain and pelvic anatomy. Tyson's monograph is a text-book example of the comparative method. He, and it, deserve more recognition.

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.

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.

Brain enlargement and dental reduction were not linked in hominin evolution.

The large brain and small postcanine teeth of modern humans are among our most distinctive features, and trends in their evolution are well studied within the hominin clade. Classic accounts hypothesize that larger brains and smaller teeth coevolved because behavioral changes associated with increased brain size allowed a subsequent dental reduction. However, recent studies have found mismatches between trends in brain enlargement and posterior tooth size reduction in some hominin species. We use a multiple-variance Brownian motion approach in association with evolutionary simulations to measure the tempo and mode of the evolution of endocranial and dental size and shape within the hominin clade. We show that hominin postcanine teeth have evolved at a relatively consistent neutral rate, whereas brain size evolved at comparatively more heterogeneous rates that cannot be explained by a neutral model, with rapid pulses in the branches leading to later Homo species. Brain reorganization shows evidence of elevated rates only much later in hominin evolution, suggesting that fast-evolving traits such as the acquisition of a globular shape may be the result of direct or indirect selection for functional or structural traits typical of modern humans.

Pattern and process in hominin brain size evolution are scale-dependent.

A large brain is a defining feature of modern humans, yet there is no consensus regarding the patterns, rates and processes involved in hominin brain size evolution. We use a reliable proxy for brain size in fossils, endocranial volume (ECV), to better understand how brain size evolved at both clade- and lineage-level scales. For the hominin clade overall, the dominant signal is consistent with a gradual increase in brain size. This gradual trend appears to have been generated primarily by processes operating within hypothesized lineages-64% or 88% depending on whether one uses a more or less speciose taxonomy, respectively. These processes were supplemented by the appearance in the fossil record of larger-brained Homo species and the subsequent disappearance of smaller-brained Australopithecus and Paranthropus taxa. When the estimated rate of within-lineage ECV increase is compared to an exponential model that operationalizes generation-scale evolutionary processes, it suggests that the observed data were the result of episodes of directional selection interspersed with periods of stasis and/or drift; all of this occurs on too fine a timescale to be resolved by the current human fossil record, thus producing apparent gradual trends within lineages. Our findings provide a quantitative basis for developing and testing scale-explicit hypotheses about the factors that led brain size to increase during hominin evolution.