Michael Coquerelle

Early dispersal of modern humans in Europe and implications for Neanderthal behaviour

The appearance of anatomically modern humans in Europe and the nature of the transition from the Middle to Upper Palaeolithic are matters of intense debate. Most researchers accept that before the arrival of anatomically modern humans, Neanderthals had adopted several ‘transitional’ technocomplexes. Two of these, the Uluzzian of southern Europe and the Châtelperronian of western Europe, are key to current interpretations regarding the timing of arrival of anatomically modern humans in the region and their potential interaction with Neanderthal populations. They are also central to current debates regarding the cognitive abilities of Neanderthals and the reasons behind their extinction. However, the actual fossil evidence associated with these assemblages is scant and fragmentary, and recent work has questioned the attribution of the Châtelperronian to Neanderthals on the basis of taphonomic mixing and lithic analysis. Here we reanalyse the deciduous molars from the Grotta del Cavallo (southern Italy), associated with the Uluzzian and originally classified as Neanderthal. Using two independent morphometric methods based on microtomographic data, we show that the Cavallo specimens can be attributed to anatomically modern humans. The secure context of the teeth provides crucial evidence that the makers of the Uluzzian technocomplex were therefore not Neanderthals. In addition, new chronometric data for the Uluzzian layers of Grotta del Cavallo obtained from associated shell beads and included within a Bayesian age model show that the teeth must date to ∼45,000–43,000 calendar years before present. The Cavallo human remains are therefore the oldest known European anatomically modern humans, confirming a rapid dispersal of modern humans across the continent before the Aurignacian and the disappearance of Neanderthals.

Sexual dimorphism of the human mandible and its association with dental development

The present study investigates whether the human mandible is sexually dimorphic during early postnatal development and whether early dimorphic features persist during subsequent ontogeny. We also examine whether mandibular dimorphism is linked to dimorphism of dental development. Dense CT-derived mandibular meshes of 84 females and 75 males, ranging from birth to adulthood, were analyzed using geometric morphometric methods. On the basis of the specimens chronological ages and mineralization stages of the deciduous and permanent teeth, we compute dental age as proxy for dental development by the additive conjoint measurement method. By birth, males have, on average, more advanced age-specific shapes than females. However, sex differences decrease quickly as females catch up via a different association between shape and size. This leads to an almost complete reduction of sexual dimorphism between the ages of 4 and 14. From puberty to adulthood, males are characterized by allometric shape changes while the shape of the female mandible continues to change even after size has ceased to increase. Dimorphism of dental maturation becomes visible only at puberty. Sexual dimorphism, concentrated at the ramus and the mental region during the earliest ontogenetic stages and again at adulthood, is not associated with the development of the teeth. At puberty there is a simultaneous peak in size increase, shape development, and dental maturation likely controlled by the surge of sex hormones with a dimorphic onset age. We argue that the infant and adult dimorphism of the mental region may be associated with the development of supralaryngeal structures.

Comparison of dental measurement systems for taxonomic assignment of first molars

Morphometrics of the molar crown is based traditionally on diameter measurements but is nowadays more often based on 2D image analysis of crown outlines. An alternative approach involves measurements at the level of the cervical line. We compare the information content of the two options in a three-dimensional (3D) digital sample of lower and upper first molars (M(1) and M(1) ) of modern human and Neanderthal teeth. The cervical outline for each tooth was created by digitizing the cervical line and then sectioning the tooth with a best fit plane. The crown outline was projected onto this same plane. The curves were analyzed by direct extraction of diameters, diagonals, and area and also by principal component analysis either of the residuals obtained by regressing out these measurements from the radii (shape information) or directly by the radii (size and shape information). For M(1) , the crown and cervical outline radii allow us to discriminate between Neanderthals and modern humans with 90% and 95% accuracy, respectively. Fairly good discrimination between the groups (80-82.5%) was also obtained using cervical measurements. With respect to M(1) , general overlap of the two groups was obtained by both crown and cervical measurements; however, the two taxa were differentiable by crown outline residuals (90-97%). Accordingly, while crown diameters or crown radii should be used for taxonomic analysis of unworn or slightly worn M(1) s, the crown outline, after regressing out size information, could be promising for taxonomic assignment of lower M1s.

3D Morphometric Analysis of Fossil Canid Skulls Contradicts the Suggested Domestication of Dogs During the Late Paleolithic

Whether dogs were domesticated during the Pleistocene, when humans were hunter-gatherers, or during the Neolithic, when humans began to form permanent settlements and engage in agriculture, remains controversial. Recently discovered Paleolithic fossil skulls, Goyet dated 31,680 +/- 250 YBP and Eliseevichi MAE 447/5298 dated 13,905 +/- 55 YBP, were previously identified as dogs. However, new genetic studies contradict the identification of these specimens as dogs, questioning the validity of traditional measurements used to morphologically identify canid fossil skulls. We employ 3D geometric morphometric analyses to compare the cranial morphology of Goyet and Eliseevichi MAE to that of ancient and modern dogs and wolves. We demonstrate that these Paleolithic canids are definitively wolves and not dogs. Compared to mesaticephalic (wolf-like breeds) dog skulls, Goyet and Eliseevichi MAE, do not have cranial flexion and the dorsal surface of their muzzles has no concavity near the orbits. Morphologically, these early fossil canids resemble wolves, and thus no longer support the establishment of dog domestication in the Paleolithic.

Differential Growth and Development of the Upper and Lower Human Thorax

The difficulties in quantifying the 3D form and spatial relationships of the skeletal components of the ribcage present a barrier to studies of the growth of the thoracic skeleton. Thus, most studies to date have relied on traditional measurements such as distances and indices from single or few ribs. It is currently known that adult-like thoracic shape is achieved early, by the end of the second postnatal year, with the circular cross-section of the newborn thorax transforming into the ovoid shape of adults; and that the ribs become inclined such that their anterior borders come to lie inferior to their posterior. Here we present a study that revisits growth changes using geometric morphometrics applied to extensive landmark data taken from the ribcage. We digitized 402 (semi) landmarks on 3D reconstructions to assess growth changes in 27 computed tomography-scanned modern humans representing newborns to adults of both sexes. Our analyses show a curved ontogenetic trajectory, resulting from different ontogenetic growth allometries of upper and lower thoracic units. Adult thoracic morphology is achieved later than predicted, by diverse modifications in different anatomical regions during different ontogenetic stages. Besides a marked increase in antero-posterior dimensions, there is an increase in medio-lateral dimensions of the upper thorax, relative to the lower thorax. This transforms the pyramidal infant thorax into the barrel-shaped one of adults. Rib descent is produced by complex changes in 3D curvature. Developmental differences between upper and lower thoracic regions relate to differential timings and rates of maturation of the respiratory and digestive systems, the spine and the locomotor system. Our findings are relevant to understanding how changes in the relative rates of growth of these systems and structures impacted on the development and evolution of modern human body shape.

Comparison of dental measurement systems for taxonomic assignment of Neanderthal and modern human lower second deciduous molars

Traditional morphometric approaches for taxonomic assignment of Neanderthal and modern human dental remains are mainly characterized by caliper measurements of tooth crowns. Several studies have recently described differences in dental tissue proportions and enamel thickness between Neanderthal and modern human teeth. At least for the lower second deciduous molar (dm(2)), a three-dimensional lateral relative enamel thickness index has been proposed for separating the two taxa. This index has the advantage over other measurements of being applicable to worn teeth because it ignores the occlusal aspect of the crown. Nevertheless, a comparative evaluation of traditional crown dimensions and lateral dental tissue proportion measurements for taxonomic assignment of Neanderthal and modern human dm(2)s has not yet been performed. In this study, we compare various parameters gathered from the lateral aspects of the crown. These parameters include crown diameters, height of the lateral wall of the crown (lateral crown height = LCH), lateral enamel thickness, and dentine volume of the lateral wall, including the volume of the coronal pulp chamber (lateral dentine plus pulp volume = LDPV), in a 3D digital sample of Neanderthal and modern human dm(2)s to evaluate their utility in separating the two taxa. The LDPV and the LCH allow us to discriminate between Neanderthals and modern humans with 88.5% and 92.3% accuracy, respectively. Though our results confirm that Neanderthal dm(2)s have lower relative enamel thickness (RET) index compared with modern humans (p = 0.005), only 70% of the specimens were correctly classified on the basis of the RET index. We also emphasize that results of the lateral enamel thickness method depend on the magnitude of the interproximal wear. Accordingly, we suggest using the LCH or the LDPV to discriminate between Neanderthal and modern human dm(2)s. These parameters are more independent of interproximal wear and loss of lateral enamel.

Fetal and infant growth patterns of the mandibular symphysis in modern humans and chimpanzees (Pan troglodytes)

Comparison of the early development of the mandibular symphysis between primates and modern humans is of particular interest in human palaeontology. Using geometric morphometric methods, we explored and compared the ontogenetic shape changes of 14 chimpanzee mandibles (Pan troglodytes) against 66 human CT‐scanned mandibles over the age range from fetal life to the complete emergence of the deciduous dentition in a visualization incorporating the deciduous tooth arrangement. The results reveal that the symphysis is anteriorly inclined in the youngest chimpanzee fetuses but develops an increasingly vertical orientation up until birth. At the same time, the anterior teeth reorient before a vertical emergence, and a symphyseal tuber appears on the labial side. When the deciduous canine emerges, the symphysis inclines anteriorly again, exhibiting the adult characteristic slope. These two phases are characterized by a repositioning of the simian shelf. Unlike chimpanzees, the human symphysis remains vertical throughout fetal development. However, the combination of morphological changes observed in chimpanzee fetuses is similar to that of modern humans after birth, as the mental region projects forward. By elongating the alveolar process, the inclination of the chimpanzee symphysis could be a key event for emergence of the deciduous canine, as space is lacking at the alveolar ridge in a vertical symphysis once the deciduous incisors and molars have emerged. The repositioning of the simian shelf suggests that the suprahyoid muscles have a significant influence on the anterior growth of the symphysis. The anteroposterior positioning of the basal symphysis in both species may be related to hyoid bone position during ontogeny.

A mass grave from the catacomb of Saints Peter and Marcellinus in Rome, second-third century AD

Investigations in a catacomb revealed an early mass grave, in which the bodies were bound tightly with plaster and textile in a type of mummification. Over 100 individuals, mostly young adults, including women, were stacked in rows apparently following a communal fatal incident, perhaps an epidemic. The presence of traces of gold, silver and probably amber with many of the bodies, and their burial in an imperial property suggests a group of some status being interred in the early years of the catacomb, at the end of the second century AD or beginning of the third.

Virtual Reconstruction of Very Large Skull Defects Featuring Partly and Completely Missing Midsagittal Planes

Despite the development of computer‐based methods, cranial reconstruction of very large skull defects remains a challenge particularly if the damage affects the midsagittal region hampering the usage of mirror imaging techniques. This pilot study aims to deliver a new method that goes beyond mirror imaging, giving the possibility to reconstruct crania characterized by large missing areas, which might be useful in the fields of paleoanthropology, bioarcheology, and forensics. We test the accuracy of digital reconstructions in cases where two‐thirds or more of a human cranium were missing. A three‐dimensional (3D) virtual model of a human cranium was virtually damaged twice to compare two destruction‐reconstruction scenarios. In the first case, a small fraction of the midsagittal region was still preserved, allowing the application of mirror imaging techniques. In the second case, the damage affected the complete midsagittal region, which demands a new approach to estimate the position of the midsagittal plane. Reconstructions were carried out using CT scans from a sample of modern humans (12 males and 13 females), to which 3D digital modeling techniques and geometric morphometric methods were applied. As expected, the second simulation showed a larger variability than the first one, which underlines the fact that the individual midsagittal plane is of course preferable in order to minimize the reconstruction error. However, in both simulations the Procrustes mean shape was an effective reference for the reconstruction of the entire cranium, producing models that showed a remarkably low error of about 3 mm, given the extent of missing data. Anat Rec, 296:745–758, 2013.

Morphological Integration and Variation in Facial Orientation in Pongo pygmaeus pygmaeus: A Geometric Morphometric Approach via Partial Least Squares

The orientation of the face relative to the cranial base is an important aspect of intraspecific and interspecific variation in hominoids. In contrast to African apes, Pongo is characterized by a structural condition known as airorhynchy, an upwardly rotated face relative to the basicranium. In this study we investigated patterns of covariation between facial orientation and basicranial and orbital shape. The sample comprised 43 adult specimens of Pongo pygmaeus pygmaeus. The shape of the craniofacial complex is represented by 105 three-dimensional landmarks and semilandmarks digitized on images obtained with computed tomography (CT). We investigated covariation patterns and facial orientation using partial least squares analysis of basicranial and orbital shape vis-à-vis maxillary shape. We found strong integration between the shape of the cranial base, orbits, and the shape and orientation of the palate and premaxilla. Facial orientation in our sample is also highly variable. As a dorsally rotated palate is associated with a ventral rotation of the orbits but not with a reduced basicranial angle, the “facial block” hypothesis cannot explain the variation in facial orientation in Pongo. Our results also indicate that mature males have a more ventrally oriented lower face that may be due to a reconfiguration of the craniofacial complex during the extended male growth phase. These findings demonstrate that there is strong integration between the cranial base and face and a high degree of variation in facial orientation in Pongo.