Taxonomic differences in deciduous lower first molar crown outlines of Homo sapiens and Homo neanderthalensis.

Recent studies have demonstrated that the outline shapes of deciduous upper and lower second molars and the deciduous upper first molar are useful for diagnosing hominin taxa-especially Homo neanderthalensis and Homo sapiens. Building on these studies, we use geometric morphometric methods to assess the taxonomic significance of the crown outline of the lower first deciduous molar (dm1). We test whether the crown shape of the dm1 distinguishes H. neanderthalensis from H. sapiens and explore whether dm1 crown shape can be used to accurately assign individuals to taxa. Our fossil sample includes 3 early H. sapiens, 7 Upper Paleolithic H. sapiens, and 13 H. neanderthalensis individuals. Our recent human sample includes 103 individuals from Africa, Australia, Europe, South America, and South Asia. Our results indicate that H. neanderthalensis dm1s cluster fairly tightly and separate well from those of Upper Paleolithic H. sapiens. However, we also found that the range of shapes in the recent human sample completely overlaps the ranges of all fossil samples. Consequently, results of the quadratic discriminant analysis based on the first 8 principal components (PCs) representing more than 90% of the variation were mixed. Lower dm1s were correctly classified in 87.3% of the individuals; the combined H. sapiens sample had greater success (90.2%) in assigning individuals than did the H. neanderthalensis sample (61.5%). When the analysis was run removing the highly variable recent human sample, accuracy increased to 84.6% for H. neanderthalensis, and 57.1% of Upper Paleolithic H. sapiens were classified correctly by using the first 4 PCs (70.3%). We conclude that caution is warranted when assigning isolated dm1 crowns to taxa; while an assignment to H. neanderthalensis has a high probability of being correct, assignment to Upper Paleolithic H. sapiens is less certain.

The Châtelperronian conundrum: Blade and bladelet lithic technologies from Quinçay, France.

The discovery of an almost complete Neanderthal skeleton in a Châtelperronian context at Saint-Césaire 35 years ago changed our perspective on the beginning of the Upper Paleolithic in western Europe. Since then, the Châtelperronian has generally been considered a "transitional" industry rather than an Upper or a Middle Paleolithic industry because of its chronological position, and the association of Neanderthal remains with blades, bone tools and personal ornaments. Several competing hypotheses have been proposed to explain the association between Neanderthals and these types of artefacts including post-depositional mixing, acculturation from anatomically modern human populations, or an independent technological evolution by local Neanderthal populations. Quinçay Cave is the only Châtelperronian site where personal ornaments have been found that does not contain an overlying Upper Paleolithic layer. This means that the post-depositional mixing of later elements into the Châtelperronian may not be used as an explanation for the presence of these materials. We report here on a detailed technological analysis of lithic artefacts from the three Châtelperronian layers at Quinçay Cave. We compare our results with the technology of Mousterian blade industries dating to OIS (oxygen isotope stage) 5, the Mousterian of Acheulian Tradition type B, and the Proto-Aurignacian. We show that the Châtelperronian is sufficiently divergent from the Middle Paleolithic to be classified as a fully Upper Paleolithic industry, with a focus on blade and bladelet production. We also show that the Quinçay Châtelperronian includes retouched bladelets that resemble those found in the Proto-Aurignacian, but were produced in a different manner. We argue that a technological convergence cannot account for these behaviors, since the specific type of retouched bladelet associated with the Châtelperronian was also regularly used by Proto-Aurignacian of neighboring regions. We suggest that the idea of retouched bladelets may have diffused from the northern Proto-Aurignacian to the Quinçay Châtelperronian and that the transmission of the morphology of this desired end-product without the transmission of its manufacturing process may point toward a low degree of social intimacy between these groups. We conclude that the apparent paradox of the Châtelperronian is the result of the complexity of interaction between Neanderthal and anatomically modern human groups in western Europe between 45,000 and 40,000 years ago.

Allometry, merism, and tooth shape of the lower second deciduous molar and first permanent molar.

OBJECTIVES: This study investigates the effect of allometry on the shape of lower dm2 (dm2) and lower M1 (M1) crown outlines and examines whether the trajectory and magnitude of allometric scaling are shared between Neandertals and Homo sapiens. METHODS: Our sample included 164 specimens: 57 recent H. sapiens, 44 Upper Paleolithic H. sapiens, 17 early H. sapiens, and 46 Neandertals. Of these, 59 represent dm2/M1 pairs from the same individuals. Occlusal photographs were used to obtain crown shapes of dm2s and M1s. Principal components analysis (PCA) of the matrix of shape coordinates was used to explore the pattern of morphological variation across the dm2 and M1 samples. Allometry was investigated by means of the Pearson product-moment correlation coefficient. Two-block partial least squares (2B-PLS) analysis was used to explore patterns of covariation between dm2 and M1 crown outlines of matched individual pairs. RESULTS: The PCA confirmed significant differences between Neandertal and H. sapiens dm2 and M1 shapes. Allometry accounted for a small but statistically significant proportion of the total morphological variance. The magnitude of the allometric contribution to crown shape was stronger among Neandertals than among H. sapiens. However, we could not reject the null hypothesis that the two species share the same allometric trajectory. The 2B-PLS analysis of the pooled sample of paired individuals revealed a significant correlation in crown shape between dm2 and M1. While Procrustes distances differed significantly between dm2 and M1 in Neandertals, it did not among H. sapiens groups. CONCLUSIONS: Our results confirm several of the results obtained by a similar study of upper dm2/M1 (dm(2)/M(1)), but there are differences as well. Neandertal dm2/M1 shapes are less derived than those of the dm(2)/M(1). Such differences may support previous studies, which have suggested that different developmental and/or epigenetic factors affect the upper and lower dentitions.

A comparison of proximal humeral cancellous bone of great apes and humans.

The shoulder is the most mobile joint in the primate body, and is involved in both locomotor and manipulative activities. The presumed functional sensibility of trabecular bone can offer a way of decoding the activities to which the forelimbs of fossil primates were subjected. We examine the proximal humeral trabecular architecture in a relatively closely related group of similarly sized hominids (Pongo pygmaeus, Pan troglodytes, and Homo sapiens), in order to evaluate the effect of diverging habitual motion behaviors of the shoulder complex in a coherent phylogenetic group. In order to characterize and compare the humeral trabecular architectures of the three species, we imaged a large sample by high-resolution computed tomography (HrCT) and quantified their trabecular architectures by standard bone 3D morphometric parameters. Univariate statistical analysis was performed, showing significant differences among the species. However, univariate statistics could not highlight the structural particularity in the cancellous bone of each species. A principal component analysis also showed clear separation of the three taxa and enabled a structural characterization of the humeral trabecular bone of each species. We conclude that the differences in the architectural setup of the three hominids likely reflect multiple differences in their habitual activity patterns of their shoulder joint, although individual structural features are difficult to relate to specific loading conditions.

A comprehensive morphometric analysis of the frontal and zygomatic bone of the Zuttiyeh fossil from Israel.

The Zuttiyeh hominin craniofacial fossil was discovered in Israel in 1925. Radiometric dates and the archaeological context (Acheulo-Yabrudian) bracket the associated cave layers to between 200 and 500 ka (thousands of years ago), making it one of the earliest cranial fossils discovered in the Near East thus far. Its geographic position, at the corridor between Africa and Eurasia, in combination with its probable Middle Pleistocene date make it a crucial specimen for interpreting later human evolution. Since its discovery, qualitative descriptive and traditional morphometric methods have variously suggested affinities to Homo erectus (Zhoukoudian), Homo neanderthalensis (Tabun), and early Homo sapiens (Skhul and Qafzeh). To better determine the taxonomic affinities of the Zuttiyeh fossil, this study uses 3D semilandmark geometric morphometric techniques and multivariate statistical analyses to quantify the frontal and zygomatic region and compare it with other Middle to Late Pleistocene African and Eurasian hominins. Our results show that the frontal and zygomatic morphology of Zuttiyeh is most similar to Shanidar 5, a Near East Neanderthal, Arago 21, a European Middle Pleistocene hominin, and Skhul 5, an early H. sapiens. The shape differences between archaic hominins (i.e., Homo heidelbergensis and H. neanderthalensis) in this anatomical region are very subtle. We conclude that Zuttiyeh exhibits a generalized frontal and zygomatic morphology, possibly indicative of the population that gave rise to modern humans and Neanderthals. However, given that it most likely postdates the split between these two lineages, Zuttiyeh might also be an early representative of the Neanderthal lineage. Neanderthals largely retained this generalized overall morphology, whereas recent modern humans depart from this presumably ancestral morphology.

Out of Africa: modern human origins special feature: the origin of Neandertals.

Western Eurasia yielded a rich Middle (MP) and Late Pleistocene (LP) fossil record documenting the evolution of the Neandertals that can be analyzed in light of recently acquired paleogenetical data, an abundance of archeological evidence, and a well-known environmental context. Their origin likely relates to an episode of recolonization of Western Eurasia by hominins of African origin carrying the Acheulean technology into Europe around 600 ka. An enhancement of both glacial and interglacial phases may have played a crucial role in this event, as well as in the subsequent evolutionary history of the Western Eurasian populations. In addition to climatic adaptations and an increase in encephalization, genetic drift seems to have played a major role in their evolution. To date, a clear speciation event is not documented, and the most likely scenario for the fixation of Neandertal characteristics seems to be an accretion of features along the second half of the MP. Although a separation time for the African and Eurasian populations is difficult to determine, it certainly predates OIS 11 as phenotypic Neandertal features are documented as far back as and possibly before this time. It is proposed to use the term "Homo rhodesiensis" to designate the large-brained hominins ancestral to H. sapiens in Africa and at the root of the Neandertals in Europe, and to use the term "Homo neanderthalensis" to designate all of the specimens carrying derived metrical or non-metrical features used in the definition of the LP Neandertals.

Dental development of the Taï Forest chimpanzees revisited.

Developmental studies consistently suggest that teeth are more buffered from the environment than other skeletal elements. The surprising finding of late tooth eruption in wild chimpanzees (Zihlman et al., 2004) warrants reassessment in a broader study of crown and root formation. Here we re-examine the skeletal collection of Taï Forest juvenile chimpanzees using radiography and physical examination. Several new individuals are included, along with genetic and histological assessments of questionable identities. Only half of the Taï juveniles employed by Zihlman et al. (2004) have age of death known with accuracy sufficient for precise comparisons with captive chimpanzees. One key individual in the former study, misidentified during field recovery as Xindra (age 8.3), is re-identified as Goshu (age 6.4). For crown formation we find that onset and duration greatly overlap captive chimpanzees, whereas root development may be more susceptible to acceleration in captive individuals. Kuykendall's (1996) equation relating captive tooth formation stage to age gives reasonable estimates of young wild subjects' true ages. Direct comparisons of tooth eruption ages are limited. A key 3.76 year-old individual likely possessed an emerging mandibular M1 at death (previously estimated from the maxillary molar as occurring at 4.1 years). Wild individuals appear to fall near the middle or latter half of captive eruption ranges. While minor developmental differences are apparent in some comparisons, our reanalysis does not show an "unambiguous pattern" of slower tooth formation in this wild environment. These data do not undermine recent developmental studies of the comparative life histories of fossil hominins.

Early brain growth in Homo erectus and implications for cognitive ability.

Humans differ from other primates in their significantly lengthened growth period. The persistence of a fetal pattern of brain growth after birth is another important feature of human development. Here we present the results of an analysis of the 1.8-million-year-old Mojokerto child (Perning 1, Java), the only well preserved skull of a Homo erectus infant, by computed tomography. Comparison with a large series of extant humans and chimpanzees indicates that this individual was about 1 yr (0-1.5 yr) old at death and had an endocranial capacity at 72-84% of an average adult H. erectus. This pattern of relative brain growth resembles that of living apes, but differs from that seen in extant humans. It implies that major differences in the development of cognitive capabilities existed between H. erectus and anatomically modern humans.

Brief communication: dental development and enamel thickness in the Lakonis Neanderthal molar.

Developmental and structural affinities between modern human and Neanderthal dental remains continue to be a subject of debate as well as their utility for informing assessments of life history and taxonomy. Excavation of the Middle Paleolithic cave site Lakonis in southern Greece has yielded a lower third molar (LKH 1). Here, we detail the crown development and enamel thickness of the distal cusps of the LKH 1 specimen, which has been classified as a Neanderthal based on the presence of an anterior fovea and mid-trigonid crest. Crown formation was determined using standard histological techniques, and enamel thickness was measured from a virtual plane of section. Developmental differences include thinner cuspal enamel and a lower periodicity than modern humans. Crown formation in the LKH 1 hypoconid is estimated to be 2.6-2.7 years, which is shorter than modern human times. The LKH 1 hypoconid also shows a more rapid overall crown extension rate than modern humans. Relative enamel thickness was approximately half that of a modern human sample mean; enamel on the distal cusps of modern human third molars is extremely thick in absolute and relative terms. These findings are consistent with recent studies that demonstrate differences in crown development, tissue proportions, and enamel thickness between Neanderthals and modern humans. Although overlap in some developmental variables may be found, the results of this and other studies suggest that Neanderthal molars formed in shorter periods of time than modern humans, due in part to thinner enamel and faster crown extension rates.

Pretreatment and gaseous radiocarbon dating of 40-100 mg archaeological bone.

Radiocarbon dating archaeological bone typically requires 300-1000 mg material using standard protocols. We report the results of reducing sample size at both the pretreatment and 14C measurement stages for eight archaeological bones spanning the radiocarbon timescale at different levels of preservation. We adapted our standard collagen extraction protocol specifically for <100 mg bone material. Collagen was extracted at least twice (from 37-100 mg material) from each bone. Collagen aliquots containing <100 µg carbon were measured in replicate using the gas ion source of the AixMICADAS. The effect of sample size reduction in the EA-GIS-AMS system was explored by measuring 14C of collagen containing either ca. 30 µg carbon or ca. 90 µg carbon. The gas dates were compared to standard-sized graphite dates extracted from large amounts (500-700 mg) of bone material pretreated with our standard protocol. The results reported here demonstrate that we are able to reproduce accurate radiocarbon dates from <100 mg archaeological bone material back to 40,000 BP.

Molar enamel thickness and dentine horn height in Gigantopithecus blacki.

Absolutely thick molar enamel is consistent with large body size estimates and dietary inferences about Gigantopithecus blacki, which focus on tough or fibrous vegetation. In this study, 10 G. blacki molars demonstrating various stages of attrition were imaged using high-resolution microtomography. Three-dimensional average enamel thickness and relative enamel thickness measurements were recorded on the least worn molars within the sample (n = 2). Seven molars were also virtually sectioned through the mesial cusps and two-dimensional enamel thickness and dentine horn height measurements were recorded. Gigantopithecus has the thickest enamel of any fossil or extant primate in terms of absolute thickness. Relative (size-scaled) measures of enamel thickness, however, support a thick characterization (i.e., not "hyper-thick"); G. blacki relative enamel thickness overlaps slightly with Pongo and completely with Homo. Gigantopithecus blacki dentine horns are relatively short, similar to (but shorter than) those of Pongo, which in turn are shorter than those of humans and African apes. Gigantopithecus blacki molar enamel (and to a lesser extent, that of Pongo pygmaeus) is distributed relatively evenly across the occlusal surface compared with the more complex distribution of enamel thickness in Homo sapiens. The combination of evenly distributed occlusal enamel and relatively short dentine horns in G. blacki results in a flat and low-cusped occlusal surface suitable to grinding tough or fibrous food objects. This suite of molar morphologies is also found to varying degrees in Pongo and Sivapithecus, but not in African apes and humans, and may be diagnostic of subfamily Ponginae.

Modern human molar enamel thickness and enamel-dentine junction shape.

This study examines cross-sections of molar crowns in a diverse modern human sample to quantify variation in enamel thickness and enamel-dentine junction (EDJ) shape. Histological sections were generated from molars sectioned buccolingually across mesial cusps. Enamel cap area, dentine area, EDJ length, and bi-cervical diameter were measured on micrographs using a digitizing tablet. Nine landmarks along the EDJ were defined, and X and Y coordinates were digitized in order to quantify EDJ shape. Upper molars show greater values for the components of enamel thickness, leading to significantly greater average enamel thickness than in lower molars. Average enamel thickness increased significantly from M1 to M3 in both molar rows, due to significantly increasing enamel cap area in upper molars, and decreasing dentine area in lower molars. Differences in EDJ shape were found among maxillary molars in combined and individual populations. Sex differences were also found; males showed significantly greater dentine area, EDJ length, and bi-cervical diameters in certain tooth types, which resulted in females having significantly thicker average enamel. Differences in enamel thickness and EDJ shape within molars were also found among populations, although few consistent trends were evident. This study demonstrates that enamel thickness and EDJ shape vary among molars, between sexes, and among populations; these factors must be considered in the categorization and comparison of ape and human molars, particularly when isolated teeth or fossil taxa are included. Human relative enamel thickness encompasses most values reported for fossil apes and humans, suggesting limited taxonomic value when considered alone.

Archaeology. The makers of the Protoaurignacian and implications for Neandertal extinction.

The Protoaurignacian culture is pivotal to the debate about the timing of the arrival of modern humans in western Europe and the demise of Neandertals. However, which group is responsible for this culture remains uncertain. We investigated dental remains associated with the Protoaurignacian. The lower deciduous incisor from Riparo Bombrini is modern human, based on its morphology. The upper deciduous incisor from Grotta di Fumane contains ancient mitochondrial DNA of a modern human type. These teeth are the oldest human remains in an Aurignacian-related archaeological context, confirming that by 41,000 calendar years before the present, modern humans bearing Protoaurignacian culture spread into southern Europe. Because the last Neandertals date to 41,030 to 39,260 calendar years before the present, we suggest that the Protoaurignacian triggered the demise of Neandertals in this area.

Three-dimensional molar enamel distribution and thickness in Australopithecus and Paranthropus.

Thick molar enamel is among the few diagnostic characters of hominins which are measurable in fossil specimens. Despite a long history of study and characterization of Paranthropus molars as relatively 'hyper-thick', only a few tooth fragments and controlled planes of section (designed to be proxies of whole-crown thickness) have been measured. Here, we measure molar enamel thickness in Australopithecus africanus and Paranthropus robustus using accurate microtomographic methods, recording the whole-crown distribution of enamel. Both taxa have relatively thick enamel, but are thinner than previously characterized based on two-dimensional measurements. Three-dimensional measurements show that P. robustus enamel is not hyper-thick, and A. africanus enamel is relatively thinner than that of recent humans. Interspecific differences in the whole-crown distribution of enamel thickness influence cross-sectional measurements such that enamel thickness is exaggerated in two-dimensional sections of A. africanus and P. robustus molars. As such, two-dimensional enamel thickness measurements in australopiths are not reliable proxies for the three-dimensional data they are meant to represent. The three-dimensional distribution of enamel thickness shows different patterns among species, and is more useful for the interpretation of functional adaptations than single summary measures of enamel thickness.

Isotopic dietary analysis of a Neanderthal and associated fauna from the site of Jonzac (Charente-Maritime), France.

We report here on the isotopic analysis (carbon and nitrogen) of collagen extracted from a Neanderthal tooth and animal bone from the late Mousterian site of Jonzac (Charente-Maritime, France). This study was undertaken to test whether the isotopic evidence indicates that animal protein was the main source of dietary protein for this relatively late Neanderthal, as suggested by previous studies. This was of particular interest here because this is the first isotopic study of a relatively late Neanderthal associated with Mousterian of Acheulian Tradition (MTA, dating to approximately 55,000 to 40,000 BP) technology. We found that the Jonzac Neanderthal had isotopic values consistent with a diet in which the main protein sources were large herbivores, particularly bovids and horses. We also found evidence of different dietary niches between the Neanderthal and a hyena at the site, with the hyena consuming mainly reindeer.

Recent human evolution in northwestern Africa.

The first modern humans in the Maghreb are said to be associated with the Aterian industries which appeared at least 40 ka BP in the northwest. Their predecessors are mainly represented by the Jebel Irhoud (Morocco) specimens. Palaeontological evidence, as well as electron spin resonance (ESR) dating, suggests that this series is older than previously published, and should belong to oxygen isotope stage 5 or even 6. There is no evidence of any Neanderthal apomorphy in this group which can no longer be considered as 'African Nanderthals'. Clear synapomorphies with modern man combined with some plesiomorphic retentions indicate a slightly more primitive (and older?) grade than the Qafzeh-Skhul sample in southwestern Asia. The Northwestern evidence demonstrates that the mediterranean sea was a major biological barrier during the upper Middle and lower Upper Pleistocene and that the rise of anatomically modern features cannot be restricted to a sub-Saharan of eastern African area.

On the phylogenetic position of the pre-Neandertal specimen from Reilingen, Germany.

This paper describes the morphology and phylogenetic significance of a well-preserved fossil human cranium found near Reilingen, Germany in 1978. The specimen consists of two complete parietals, most of the right temporal, and 70% of the occipital. The specimen displays several features characteristic of early "archaic" Homo sapiens: maximum breadth at the supramastoid crests, roughly pentagonal occipital view, an angular torus, superoinferiorly elongate/ anteroposterior shortened squamous temporal, strong mastoid, and separation of the tympanic and mastoid temporal portions. Other features align the Reilingen cranium with the "classic" Neandertals: developed suprainiac fossa, bilaterally protuberant occipital torus, strong juxtamastoid eminence, a highly convex occipital plane, and lambdoid flattening. This morphology and our cephalometric analysis support the assignment of this specimen to a Stage 2 "pre-Neandertal" group (e.g., Steinheim, Swanscombe, and Atapuerca [SH site]). Following an "Accretion Model" of European hominid evolution, Stage 2 succeeds Stage 1 European "early-pre-Neandertals" (e.g., Mauer and Petralona), and precedes both the Saale-Eem Stage 3 "early Neandertals" (e.g., Biache 1), which present greater platycephaly, a reduced mastoid process, an enlarged juxtamastoid eminence, and true "en-bombe" (parieto-occipital form), and Weichsel Stage 4 "Neandertals" (sensu stricto, i.e., "classic" Neandertals).

A late Neanderthal associated with Upper Palaeolithic artefacts.

The French site of Arcy-sur-Cure is a key locality in documenting the Middle-Upper Palaeolithic transition in Europe. Reliable attribution of the fragmentary hominid fossils associated with its early Upper Palaeolithic Châtelperronian industry has not been possible. Here we report the first conclusive identification of one of these fossils as Neanderthal on the basis of newly discovered derived features of the bony labyrinth. Dated at about thirty-four thousand years (34 kyr) ago, the fossil is representative of the youngest known Neanderthal populations, and its archaeological context indicates that these hominids used a rich bone industry as well as personal ornaments. The evidence supports the hypothesis of a long term coexistence with technocultural interactions between the first modern humans and the last Neanderthals in Europe. However, the complete absence of the derived Neanderthal traits in labyrinths of modern Upper Palaeolithic specimens from western Europe argues against phylogenetic continuity between the two populations in this region.