Recent Modern Humans Research Articles

Cross-sectional properties of the lower limb long bones in the Middle Pleistocene Sima de los Huesos sample (Sierra de Atapuerca, Spain)

The recovery to date of three complete and five partial femora, seven complete tibiae, and four complete fibulae from the Atapuerca Sima de los Huesos site provides an opportunity to analyze the biomechanical cross-sectional properties in this Middle Pleistocene population and to compare them with those of other fossil hominins and recent modern humans.We have performed direct comparisons of the cross-sectional geometric parameters and reduced major axis (RMA) regression lines among different samples. We have determined that Atapuerca Sima de los Huesos (SH) fossils have significantly thicker cortices than those of recent modern humans for the three leg bones at all diaphyseal levels, except that of the femur at 35% of biomechanical length. The SH bones are similar to those of Neandertals and Middle Pleistocene humans and different from Homo sapiens in their diaphyseal cross-sectional shape and strength parameters.When standardized by estimated body size, both the SH and Neandertal leg bones have in general greater strength than those of H. sapiens from the early modern (EMH), Upper Paleolithic (UP), and recent populations (RH). The Sima de los Huesos human leg bones have, in general terms, an ancestral pattern similar to that of Pleistocene humans and differing from H. sapiens.

Cranial measures and ancient DNA both show greater similarity of Neandertals to recent modern Eurasians than to recent modern sub-Saharan Africans.

Ancient DNA analysis has shown that present-day humans of Eurasian ancestry are more similar to Neandertals than are present-day humans of sub-Saharan African ancestry, reflecting interbreeding after modern humans first left Africa. We use craniometric data to test the hypothesis that the crania of recent modern humans show the same pattern. We computed Mahalanobis squared distances between a published Neandertal centroid based on 37 craniometric traits and each of 2,413 recent modern humans from the Howells global data set (N = 373 sub-Saharan Africans, N = 2,040 individuals of Eurasian descent). The average distance to the Neandertal centroid is significantly lower for Eurasian crania than for sub-Saharan African crania as expected from the findings of ancient DNA (p < 0.001). This result holds when examining distances for separate geographic regions of humans of Eurasian descent (Europeans, Asians, Australasians, Native Americans, and Pacific Islanders). Most of these results are also seen when examining distances partitioning size and shape variation. Our results show that the genetic difference in Neandertal ancestry seen in the DNA of present-day sub-Saharan Africans and Eurasians is also found in patterns of recent modern human craniometric variation.

Morphology and structure of Homo erectus humeri from Zhoukoudian, Locality 1.

BackgroundRegional diversity in the morphology of the H. erectus postcranium is not broadly documented, in part, because of the paucity of Asian sites preserving postcranial fossils. Yet, such an understanding of the initial hominin taxon to spread throughout multiple regions of the world is fundamental to documenting the adaptive responses to selective forces operating during this period of human evolution.MethodsThe current study reports the first humeral rigidity and strength properties of East Asian H. erectus and places its diaphyseal robusticity into broader regional and temporal contexts. We estimate true cross-sectional properties of Zhoukoudian Humerus II and quantify new diaphyseal properties of Humerus III using high resolution computed tomography. Comparative data for African H. erectus and Eurasian Late Pleistocene H. sapiens were assembled, and new data were generated from two modern Chinese populations.ResultsDifferences between East Asian and African H. erectus were inconsistently expressed in humeral cortical thickness. In contrast, East Asian H. erectus appears to exhibit greater humeral robusticity compared to African H. erectus when standardizing diaphyseal properties by the product of estimated body mass and humeral length. East Asian H. erectus humeri typically differed less in standardized properties from those of side-matched Late Pleistocene hominins (e.g., Neanderthals and more recent Upper Paleolithic modern humans) than did African H. erectus, and often fell in the lower range of Late Pleistocene humeral rigidity or strength properties.DiscussionQuantitative comparisons indicate that regional variability in humeral midshaft robusticity may characterize H. erectus to a greater extent than presently recognized. This may suggest a temporal difference within H. erectus, or possibly different ecogeographical trends and/or upper limb loading patterns across the taxon. Both discovery and analysis of more adult H. erectus humeri are critical to further evaluating and potentially distinguishing between these possibilities.

The costal remains of the El Sidrón Neanderthal site (Asturias, northern Spain) and their importance for understanding Neanderthal thorax morphology

The study of the Neanderthal thorax has attracted the attention of the scientific community for more than a century. It is agreed that Neanderthals have a more capacious thorax than modern humans, but whether this was caused by a medio-lateral or an antero-posterior expansion of the thorax is still debated, and is key to understanding breathing biomechanics and body shape in Neanderthals. The fragile nature of ribs, the metameric structure of the thorax and difficulties in quantifying thorax morphology all contribute to uncertainty regarding precise aspects of Neanderthal thoracic shape. The El Sidrón site has yielded costal remains from the upper to the lower thorax, as well as several proximal rib ends (frequently missing in the Neanderthal record), which help to shed light on Neanderthal thorax shape. We compared the El Sidrón costal elements with ribs from recent modern humans as well as with fossil modern humans and other Neanderthals through traditional morphometric methods and 3D geometric morphometrics, combined with missing data estimation and virtual reconstruction (at the 1st, 5th and 11th costal levels). Our results show that Neanderthals have larger rib heads and articular tubercles than their modern human counterparts. Neanderthal 1st ribs are smaller than in modern humans, whereas 5th and 11th ribs are considerably larger. When we articulated mean ribs (size and shape) with their corresponding vertebral elements, we observed that compared to modern humans the Neanderthal thorax is medio-laterally expanded at every level, especially at T5 and T11. Therefore, in the light of evidence from the El Sidrón costal remains, we hypothesize that the volumetric expansion of the Neanderthal thorax proposed by previous authors would mainly be produced by a medio-lateral expansion of the thorax.

Estimation of cranial capacity of healthy adult students of a medical university in eastern part of Nepal

The Late Middle Pleistocene hominin fossils from the Xujiayao site in northern China have been closely studied in light of their morphological variability. However, all previous studies have focused on separated cranial fragments. Here, we report the first reconstruction of a fairly complete posterior cranium, Xujiayao 6 (XJY 6), confidently dated to ∼200–160 ka, which facilitated an assessment of its overall cranial size. XJY 6 was reconstructed from three of the original fragments—the PA1486 (No.7/XJY 6a) occipital bone, PA1490 (No.10/XJY 6b) right parietal bone, and PA1498 (No.17/XJY 15) left temporal bone—which originated from the same young adult individual. The XJY 6 endocranial capacity, estimated by measuring endocranial volume, was estimated using multiple regression formulae derived from ectocranial and endocranial measurements on select samples of Pleistocene hominins and recent modern humans. The results indicate that the larger pooled sample of both Pleistocene and recent modern humans was more robust for the endocranial capacity estimate. Based on the pooled sample using the ectocranial and endocranial measurements, we conservatively estimate the XJY 6 endocranial volume to be ∼1700 cm3 with a 95% confidence interval of 1555–1781 cm3. This is close to Xuchang 1, which dates to 125–105 ka and whose endocranial volume is ∼1800 cm3. Thus, XJY 6 provides the earliest evidence of a brain size that falls in the upper range of Neanderthals and modern Homo sapiens. XJY 6, together with Xuchang 1, Homo floresiensis, Homo luzonensis, and Homo naledi, challenge the general pattern that brain size gradually increases over geological time. This study also finds that hominin brain size expansion occurred at different rates across time and space.

The Late Neandertal permanent lower left third premolar from Walou Cave (Trooz, Belgium) and its context.

We describe a hominin permanent lower left third premolar unearthed in 1997 at Walou Cave (Belgium), found in direct association with a Mousterian lithic industry, in a layer directly dated to 40-38,000years BP. The taxonomical attribution of the tooth is addressed through comparative morphometric analyses, and stable isotope analyses aimed at determining the diet of the individual. The Walou P3 plots within the Neandertal range of variation and is significantly different from recent modern humans in all morphometric assessments. The isotope data showed that like other Neandertals, the Walou individual acquired its dietary proteins primarily from terrestrial food sources. We discuss the implications of the existence of a clearly Neandertal premolar dating to the period of the Middle to Upper Paleolithic transition in the Meuse river basin.

Brief communication: An analysis of dental development in Pleistocene Homo using skeletal growth and chronological age

This study takes a new approach to interpreting dental development in Pleistocene Homo in comparison with recent modern humans. As rates of dental development and skeletal growth are correlated given age in modern humans, using age and skeletal growth in tandem yields more accurate dental development estimates. Here, I apply these models to fossil Homo to obtain more individualized predictions and interpretations of their dental development relative to recent modern humans. Proportional odds logistic regression models based on three recent modern human samples (N = 181) were used to predict permanent mandibular tooth development scores in five Pleistocene subadults: Homo erectus/ergaster, Neanderthals, and anatomically modern humans (AMHs). Explanatory variables include a skeletal growth indicator (i.e., diaphyseal femoral length), and chronological age. AMHs Lagar Velho 1 and Qafzeh 10 share delayed incisor development, but exhibit considerable idiosyncratic variation within and across tooth types, relative to each other and to the reference samples. Neanderthals Dederiyeh 1 and Le Moustier 1 exhibit delayed incisor coupled with advanced molar development, but differences are reduced when femoral diaphysis length is considered. Dental development in KNM-WT 15,000 Homo erectus/ergaster, while advanced for his age, almost exactly matches the predictions once femoral length is included in the models. This study provides a new interpretation of dental development in KNM-WT 15000 as primarily reflecting his faster rates of skeletal growth. While the two AMH specimens exhibit considerable individual variation, the Neanderthals exhibit delayed incisor development early and advanced molar development later in ontogeny.

Manot 1 calvaria and Recent Modern Human Evolution: an Anthropological Perspective

The time range between 60 ka and 50 ka is one of the most dramatic phases in human biological evolution. In this period, the western part of Eurasia (Europe and the Near East) was populated by Neanderthals, whereas the eastern part (Central Asia and Siberia) was populated by Denisovans. However, by 30 ka, these two populations were replaced by anatomically modern humans (AMH). When did these newcomers arrive and from where? There is accumulating archaeological and genetic evidence suggesting that this demographic shift occurred at the end of MIS 4 [1–3]. Moreover, it is quite clear that a major dispersal of AMH out of Africa was the source of the new populations [4–7]. In this study, we examined specific morphological characteristics of Manot 1 (e.g., suprainiac fossa), and assessed their similarities to the corresponding traits found among Neanderthals. We will show that although the terminology is similar, the traits in each hominin group are of different entities. We also show that Manot 1 and Early Upper Palaeolithic skulls of Europe have many traits in common (e.g., suprainiac fossa, bunning), although Manot 1 is much more gracile. Finally, some of the archaic traits (e.g., suprainiac fossa) seen in Manot 1 can be traced to the Late Pleistocene Aduma skull (~79–105 ka) from Ethiopia or even Eyasi 1 (~200–400 ka) from Tanzania.

Low trabecular bone density in recent sedentary modern humans

Research on a limited number of samples suggests that trabecular bone density (i.e., bone volume fraction, BVF) within specific articulations is lower among more sedentary Holocene agricultural populations compared with Holocene foragers, implying that activity levels have a significant effect on trabecular BVF. However, it is unclear to what extent BVF differs among groups with varying activity levels and how general this phenomenon is across multiple limb articulations. Here, we test two hypotheses that: (i) sedentary populations have lower BVF compared with active populations across limb articulations; and (ii) these declines are more uniform in the lower limb (because of its more direct relationship to mobility), and more variable in the upper limb. We estimated BVF in seven lower and upper limb articulations of five Holocene population samples with subsistence strategies spanning from foraging through horticultural to industrial using pQCT (peripheral Quantitative Computed Tomography). Both hypotheses are largely supported. First, the most active groups have significantly greater BVF in most limb elements compared with more sedentary groups. Second, all sedentary groups have relatively similar (and lower) BVF in the lower limb but show more variation in upper limb articulations. These results suggest that a decline in activity levels associated with the adoption of agriculture and industrialization significantly contributed to the reduction in BVF in recent modern humans, but specific behavioral changes, particularly in the upper limb, also affected these patterns.

The morphology of the enamel–dentine junction in Neanderthal molars: Gross morphology, non-metric traits, and temporal trends

This study explores the morphological differences between the enamel–dentine junction (EDJ) of maxillary and mandibular molars of Neanderthals (n = 150) and recent modern humans (n = 106), and between an earlier Neanderthal sample (consisting of Pre-Eemian and Eemian Neanderthals dating to before 115 ka) and a later Neanderthal sample (consisting of Post-Eemian Neanderthals dating to after 115 ka). The EDJ was visualised by segmenting microtomographic scans of each molar. A geometric morphometric methodology compared the positioning of the dentine horns, the shape of the marginal ridge between the dentine horns, and the shape of the cervix. We also examined the manifestation of non-metric traits at the EDJ including the crista obliqua, cusp 5, and post-paracone tubercle. Furthermore, we report on additional morphological features including centrally placed dentine horn tips and twinned dentine horns. Our results indicate that EDJ morphology can discriminate with a high degree of reliability between Neanderthals and recent modern humans at every molar position, and discriminate between the earlier and the later Neanderthal samples at every molar position, except for the M3 in shape space. The cervix in isolation can also discriminate between Neanderthals and recent modern humans, except at the M3 in form space, and is effective at discriminating between the earlier and the later Neanderthal samples, except at the M2/M2 in form space. In addition to demonstrating the taxonomic valence of the EDJ, our analysis reveals unique manifestations of dental traits in Neanderthals and expanded levels of trait variation that have implications for trait definitions and scoring.

The Qesem Cave hominin material (part 2): A morphometric analysis of dm2-QC2 deciduous lower second molar

The Qesem Cave Middle Pleistocene hominin site has yielded a well preserved lower second deciduous molar (dm2-QC2), among several other human dental remains. It has been previously described by Hershkovitz et al. using traditional methods. In this study, we used multiple approaches in order to characterize the outer and inner morphology of dm2-QC2, namely a descriptive investigation of the inner morphology, analysis of the dental tissues, and comparative 3D geometric morphometric investigation of various aspects of the dental crown based on data gathered from μCT images. Dm2-QC2 was compared to a sample of 44 specimens, including recent and fossil modern humans, Neanderthals, and Homo erectus.Our comprehensive quantitative investigation agrees with Hershkovitz et al. with regard to the mixed morphology of this specimen. Dm2-QC2 allies morphologically with Neanderthals and Skhul X for its squared cervical outline, but is intermediate between modern humans and Neanderthals for its mildly distally expanded crown outline. Dm2-QC2 falls within Neanderthal variability in having relatively high dentine horns with inwardly bent tips. It is peculiar for its mesio-distal elongation of the occlusal marginal ridge at the enamel–dentine junction. The relative enamel thickness delivers different results if measured at the mesial section (intermediate but closer to modern humans) or at the entire crown (close to the Neanderthal distribution). In terms of size, dm2-QC2 and Qafzeh 15 are among the largest specimens in our sample.Dm2-QC2 shows a mosaic of features, with a prevalence of those typical or at least frequent in Neanderthals. Among the latter, the mid-trigonid crest and taurodontism are only slightly expressed. On the basis of this isolated dental evidence, and considering the paucity of information available on human evolution in the Levantine Middle Pleistocene, we do not attempt a taxonomic classification of this single tooth. However, dm2-QC2 indicates that Neanderthal features were already present in the Middle Pleistocene Levant. We cannot rule out the possibility that the dm2-QC2 population contributed to later Levantine or other Eurasian populations, but this is too speculative given the information currently available.

On some aspects of Neandertal zygomatic morphology

Neandertals are characterized by a series of well-documented facial characteristics, including midfacial prognathism, large nasal and orbital areas, and a marked supraorbital torus. We provide a comparative morphometric study of another part of this facial complex, the frontal process of the zygomatic. We find that European Neandertals have a distinctly columnar form of the frontal process not found in recent modern humans and most Pleistocene modern humans. Some purportedly modern specimens and specimens pre-dating Neandertals exhibit the same pattern as European Neandertals, while others exhibit the modern human pattern. The columnar form is likely a retention of the ancestral state in Neandertals and the other late Pleistocene specimens that exhibit it, but variation in the pattern seen in early modern humans reveals possible insights into late Pleistocene human evolution.

Unconstrained cranial evolution in Neandertals and modern humans compared to common chimpanzees.

A variety of lines of evidence support the idea that neutral evolutionary processes (genetic drift, mutation) have been important in generating cranial differences between Neandertals and modern humans. But how do Neandertals and modern humans compare with other species? And how do these comparisons illuminate the evolutionary processes underlying cranial diversification? To address these questions, we used 27 standard cranial measurements collected on 2524 recent modern humans, 20 Neandertals and 237 common chimpanzees to estimate split times between Neandertals and modern humans, and between Pan troglodytes verus and two other subspecies of common chimpanzee. Consistent with a neutral divergence, the Neandertal versus modern human split-time estimates based on cranial measurements are similar to those based on DNA sequences. By contrast, the common chimpanzee cranial estimates are much lower than DNA-sequence estimates. Apparently, cranial evolution has been unconstrained in Neandertals and modern humans compared with common chimpanzees. Based on these and additional analyses, it appears that cranial differentiation in common chimpanzees has been restricted by stabilizing natural selection. Alternatively, this restriction could be due to genetic and/or developmental constraints on the amount of within-group variance (relative to effective population size) available for genetic drift to act on.

Perikymata distribution in Homo with special reference to the Xujiayao juvenile.

This study investigates where the Xujiayao juvenile (I(1) and C(1) ) fits into the array of perikymata distribution patterns found within the genus Homo. In addition to the I(1) and the C(1) of the Xujiayao juvenile, this study includes samples of early Homo (H. rudolfensis and H. erectus), Neandertals, early modern humans (Qafzeh), and recent modern humans from Southern Africa, Newcastle (UK), and North America (Inupiaq, AK). Three sets of analyses were undertaken, including a comparison of percentage of perikymata in the cervical half of the crown, repeated measures analysis of the percentage of total perikymata in each decile, and canonical variates analysis using both total perikymata number and the percentage of perikymata in the cervical half of the crown. The I(1) and C(1) of early Homo and Neandertals have a lower percentage of perikymata in the cervical half of the crown than modern human samples. Repeated measures analysis reveals clear distinctions in the distribution of perikymata between the modern human and fossil samples. Canonical variates analysis suggests greater differences between modern humans and the fossil samples than within the fossil samples, and classifies the Xujiayao teeth among modern humans. The present study further clarifies variation of perikymata distribution patterns within the genus Homo. The perikymata distribution of the Xujiayao juvenile tends to be more similar to that of modern humans than to either early Homo or Neandertals.

The endostructural pattern of a middle pleistocene human femoral diaphysis from the Karain E site (Southern Anatolia, Turkey).

The human femur from Karain E Cave (Turkey) exhumed from a Mousterian level provided the opportunity to make an incursion into the structural morphology of a late adolescent, or a young adult, femoral shaft from the late Middle Pleistocene of Anatolia. Considering the chrono-ecogeographical context, this study focuses particularly on the endostructural morphological similarities between Karain and Neanderthal fossils. Comparative analysis shows that some femoral features of the Karain specimen are frequently observed in Neanderthals, in comparison to some Middle Pleistocene Homo and Middle/Upper Paleolithic modern humans. In particular, we note a high degree of circularity and a strong midshaft posteromedial reinforcement of cortical thickness on the medial side. According to the mapping of cortical thickness, this latter feature can be related to the medial spiral distribution pattern of cortical thickness in the mid-proximal shaft, which is present at Karain and in all Neanderthals available for this study. This spiral distribution was not identified in recent modern humans and may be absent from ancient Homo with femoral pilaster. The endostructural signature of Karain could indicate a similar biomechanical strain system to that of Neanderthals that could be linked to body shape. However, the presence of posteromedial reinforcement in Berg Aukas may point to an ancestral feature and may be independent of latitude. A larger comparative sample should further clarify the taxonomical, biomechanical, and chrono-ecogeographical origins of the structural femoral features observed in an evolutionary Neanderthal context from MIS 7-9 in Karain.

Enamel thickness trends in Plio-Pleistocene hominin mandibular molars

Enamel thickness continues to be an important morphological character in hominin systematics and is frequently invoked in dietary reconstructions of Plio-Pleistocene hominin taxa. However, to date, the majority of published data on molar enamel thickness of Pliocene and early Pleistocene hominins derive from naturally fractured random surfaces of a small number of specimens. In this study we systematically analyze enamel thickness in a large sample of Plio-Pleistocene fossil hominins (n = 99), extant hominoids (n = 57), and modern humans (n = 30). Based on analysis of 2D mesial planes of section derived from microtomography, we examine both average and relative enamel thickness, and the distribution of enamel across buccal, occlusal, and lingual components of mandibular molars. Our results confirm the trend of increasing enamel thickness during the Pliocene that culminates in the thick enamel of the robust Australopithecus species, and then decreases from early Homo to recent modern humans. All hominin taxa share a regional average enamel thickness pattern of thick occlusal enamel and greater buccal than lingual enamel thickness. Pan is unique in exhibiting the thinnest average enamel thickness in the occlusal basin. Statistical analysis indicates that among Pliocene hominins enamel thickness is a weak taxonomic discriminator. The data underlying these results are included in a table in the Supplementary Online Material.

Recent origin of low trabecular bone density in modern humans

Humans are unique, compared with our closest living relatives (chimpanzees) and early fossil hominins, in having an enlarged body size and lower limb joint surfaces in combination with a relatively gracile skeleton (i.e., lower bone mass for our body size). Some analyses have observed that in at least a few anatomical regions modern humans today appear to have relatively low trabecular density, but little is known about how that density varies throughout the human skeleton and across species or how and when the present trabecular patterns emerged over the course of human evolution. Here, we test the hypotheses that (i) recent modern humans have low trabecular density throughout the upper and lower limbs compared with other primate taxa and (ii) the reduction in trabecular density first occurred in early Homo erectus, consistent with the shift toward a modern human locomotor anatomy, or more recently in concert with diaphyseal gracilization in Holocene humans. We used peripheral quantitative CT and microtomography to measure trabecular bone of limb epiphyses (long bone articular ends) in modern humans and chimpanzees and in fossil hominins attributed to Australopithecus africanus, Paranthropus robustus/early Homo from Swartkrans, Homo neanderthalensis, and early Homo sapiens. Results show that only recent modern humans have low trabecular density throughout the limb joints. Extinct hominins, including pre-Holocene Homo sapiens, retain the high levels seen in nonhuman primates. Thus, the low trabecular density of the recent modern human skeleton evolved late in our evolutionary history, potentially resulting from increased sedentism and reliance on technological and cultural innovations.

Functional Morphology of the Neandertal Scapular Glenoid Fossa

Neandertals and Homo sapiens are known to differ in scapular glenoid fossa morphology. Functional explanations may be appropriate for certain aspects of glenoid fossa morphology; however, other factors--e.g., allometry, evolutionary development--must be addressed before functional morphology is considered. Using three-dimensional geometric morphometrics, shape of the scapular glenoid fossa was compared among Neandertals, early and recent modern humans, chimpanzees, orangutans, Australopithecus afarensis, and Au. sediba. Permutation analysis revealed that side, sex, and lifestyle did not correlate with shape. Of the features we found to differ between groups, anterior glenoid rim morphology and fossa curvature did not correlate with the aforementioned shape variables; thus, a functional explanation is appropriate for these components of glenoid fossa shape. Shared morphology among recent humans and chimpanzees (to the exclusion of Neandertals and orangutans) suggests independent forces contributing to these morphological configurations. Potential explanations include adaptations to habitual behavior and locomotor adaptations in the scapulae of recent humans and chimpanzees; these explanations are supported by clinical and experimental literature. The absence of these morphological features in Neandertals may support the lack of these selective forces on their scapular glenoid fossa morphology.

Hominin teeth from the early Late Pleistocene site of Xujiayao, Northern China.

It is generally accepted that from the late Middle to the early Late Pleistocene (∼340-90 ka BP), Neanderthals were occupying Europe and Western Asia, whereas anatomically modern humans were present in the African continent. In contrast, the paucity of hominin fossil evidence from East Asia from this period impedes a complete evolutionary picture of the genus Homo, as well as assessment of the possible contribution of or interaction with Asian hominins in the evolution of Homo sapiens and Homo neanderthalensis. Here we present a comparative study of a hominin dental sample recovered from the Xujiayao site, in Northern China, attributed to the early Late Pleistocene (MIS 5 to 4). Our dental study reveals a mosaic of primitive and derived dental features for the Xujiayao hominins that can be summarized as follows: i) they are different from archaic and recent modern humans, ii) they present some features that are common but not exclusive to the Neanderthal lineage, and iii) they retain some primitive conformations classically found in East Asian Early and Middle Pleistocene hominins despite their young geological age. Thus, our study evinces the existence in China of a population of unclear taxonomic status with regard to other contemporary populations such as H. sapiens and H. neanderthalensis. The morphological and metric studies of the Xujiayao teeth expand the variability known for early Late Pleistocene hominin fossils and suggest the possibility that a primitive hominin lineage may have survived late into the Late Pleistocene in China.

Variations in size, shape and asymmetries of the third frontal convolution in hominids: Paleoneurological implications for hominin evolution and the origin of language

The study of brain structural asymmetries as anatomical substrates of functional asymmetries in extant humans, great apes, and fossil hominins is of major importance in understanding the structural basis of modern human cognition. We propose methods to quantify the variation in size, shape and bilateral asymmetries of the third frontal convolution (or posterior inferior frontal gyrus) among recent modern humans, bonobos and chimpanzees, and fossil hominins using actual and virtual endocasts. These methodological improvements are necessary to extend previous qualitative studies of these features. We demonstrate both an absolute and relative bilateral increase in the size of the third frontal convolution in width and length between Pan species, as well as in hominins. We also observed a global bilateral increase in the size of the third frontal convolution across all species during hominin evolution, but also non-allometric intra-group variations independent of brain size within the fossil samples. Finally, our results show that the commonly accepted leftward asymmetry of Broca's cap is biased by qualitative observation of individual specimens. The trend during hominin evolution seems to be a reduction in size on the left compared with the right side, and also a clearer definition of the area. The third frontal convolution considered as a whole projects more laterally and antero-posteriorly in the right hemisphere. As a result, the left ‘Broca's cap’ looks more globular and better defined. Our results also suggest that the pattern of brain asymmetries is similar between Pan paniscus and hominins, leaving the gradient of the degree of asymmetry as the only relevant structural parameter. As the anatomical substrate related to brain asymmetry has been present since the appearance of the hominin lineage, it is not possible to prove a direct relationship between the extent of variations in the size, shape, and asymmetries of the third frontal convolution and the origin of language in hominins.