Nobody's land? The oldest evidence of early Upper Paleolithic settlements in inland Iberia.

The Iberian Peninsula is a key region for unraveling human settlement histories of Eurasia during the period spanning the decline of Neandertals and the emergence of anatomically modern humans (AMH). There is no evidence of human occupation in central Iberia after the disappearance of Neandertals ~42,000 years ago until approximately 26,000 years ago, rendering the region "nobody's land" during the Aurignacian period. The Abrigo de la Malia provides irrefutable evidence of human settlements dating back to 36,200 to 31,760 calibrated years before the present (cal B.P.) This site also records additional levels of occupation around 32,420 to 26,260 cal B.P., suggesting repeated settlement of this territory. Our multiproxy examination identifies a change in climate trending toward colder and more arid conditions. However, this climatic deterioration does not appear to have affected AMH subsistence strategies or their capacity to inhabit this region. These findings reveal the ability of AMH groups to colonize regions hitherto considered uninhabitable, reopening the debate on early Upper Paleolithic population dynamics of southwestern Europe.

Sexual dimorphism in the first rib of Homo sapiens.

This work aimed to study sexual dimorphism in the first rib of modern humans, with a special focus on whether differences in shape are due to divergent allometric growth in males and females. Also, we compare the accuracy of sex classification using different approaches based on two methodologies, traditional morphometry based on linear measurements and geometric morphometric analysis based on 2D landmark coordinates. The sample studied here comprised 121 right and left first ribs from 65 female and male adult recent Euro-American Homo sapiens individuals. For traditional morphometrics, 12 metric variables were collected from each rib using a digital caliper, and for geometric morphometrics, six landmarks and 31 semilandmarks were captured from photographs using digital software. Both geometric morphometric and metric data were analyzed to calculate the index of sexual dimorphism, variation related to lateral asymmetry, variation in size and shape, and allometric trends between males and females. Finally, a linear discriminant analysis (LDA) was performed comparing both methodologies to test the best approach for sex classification. Results indicated that there are significant sex differences in the size and shape of the first ribs of recent Euro-American Homo sapiens. Regression analysis revealed different allometric patterns for males and females, and this could partially explain shape differences between sexes. Additionally, traditional morphometrics showed that all characteristics analyzed are significantly dimorphic, with the midshaft minimum craniocaudal diameter, the sternal end minimum diameter, and the neck minimum craniocaudal diameter displaying the most dimorphic scores. Similarly, geometric morphometrics results indicated that males have more curved and interno-exteriorly wider first ribs. Finally, analysis of sex classification using LDA yielded slightly better accuracy for traditional morphometry (83.8%) than the geometric morphometrics approach (81.3%) based on form Procrustes coordinates. This study demonstrates the usefulness of applying two different morphometric approaches to obtain more comprehensive results.

Covariation between the cranium and the cervical vertebrae in hominids.

The analysis of patterns of integration is crucial for the reconstruction and understanding of how morphological changes occur in a taxonomic group throughout evolution. These patterns are relatively constant; however, both patterns and the magnitudes of integration may vary across species. These differences may indicate morphological diversification, in some cases related to functional adaptations to the biomechanics of organisms. In this study, we analyze patterns of integration between two functional and developmental structures, the cranium and the cervical spine in hominids, and we quantify the amount of divergence of each anatomical element through phylogeny. We applied these methods to three-dimensional data from 168 adult hominid individuals, summing a total of more than 1000 cervical vertebrae. We found the atlas (C1) and axis (C2) display the lowest covariation with the cranium in hominids (Homo sapiens, Pan troglodytes, Pan paniscus, Gorilla gorilla, Gorilla beringei, Pongo pygmaeus). H. sapiens show a relatively different pattern of craniocervical correlation compared with chimpanzees and gorillas, especially in variables implicated in maintaining the balance of the head. Finally, the atlas and axis show lower magnitude of shape change during evolution than the rest of the cervical vertebrae, especially those located in the middle of the subaxial cervical spine. Overall, results suggest that differences in the pattern of craniocervical correlation between humans and gorillas and chimpanzees could reflect the postural differences between these groups. Also, the stronger craniocervical integration and larger magnitude of shape change during evolution shown by the middle cervical vertebrae suggests that they have been selected to play an active role in maintaining head balance.

The human remains from Axlor (Dima, Biscay, northern Iberian Peninsula).

OBJECTIVES: We provide the description and comparative analysis of all the human fossil remains found at Axlor during the excavations carried out by J. M. de Barandiarán from 1967 to 1974: a cranial vault fragment and seven teeth, five of which likely belonged to the same individual, although two are currently lost. Our goal is to describe in detail all these human remains and discuss both their taxonomic attribution and their stratigraphic context. MATERIALS AND METHODS: We describe external and internal anatomy, and use classic and geometric morphometrics. The teeth from Axlor are compared to Neandertals, Upper Paleolithic, and recent modern humans. RESULTS: Two teeth (a left dm2 , a left di1 ) and the parietal fragment show morphological features consistent with a Neandertal classification, and were found in an undisturbed Mousterian context. The remaining three teeth (plus the two lost ones), initially classified as Neandertals, show morphological features and a general size that are more compatible with their classification as modern humans. DISCUSSION: A left parietal fragment (Level VIII) from a single probably adult Neandertal individual was recovered during the old excavations performed by Barandiarán. Additionally, two different Neandertal children lost deciduous teeth during the formations of levels V (left di1 ) and IV (right dm2 ). In addition, a modern human individual is represented by five remains (two currently lost) from a complex stratigraphic setting. Some of the morphological features of these remains suggest that they may represent one of the scarce examples of Upper Paleolithic modern human remains in the northern Iberian Peninsula, which should be confirmed by direct dating.

Evolutionary selection and morphological integration in the vertebral column of modern humans.

OBJECTIVES: The main objective is to quantify integration, modularity, and response to selection in the presacral vertebral column of modern humans. MATERIALS AND METHODS: Seventeen linear variables on each presacral vertebra were collected in 108 modern humans producing a total of ~39,000 measurements. Then, we studied patterns and magnitudes of integration at regional, vertebral, and intra-vertebral levels. Additionally, we calculated the ability of vertebrae to respond to selection by quantifying differences in evolvability, flexibility, and constraint throughout the spine. RESULTS: The results indicate that caudal vertebrae are more evolvable than those located more cranially in the presacral vertebral column, following an increasing pattern of evolvability from the cervical to the lumbar region. Additionally, the atlas and fifth lumbar vertebra show the lowest values of integration, while central thoracic vertebrae display the highest magnitudes of integration. DISCUSSION: These results could be related to three main factors: body plan organization expressed by the Hox genes, the strong developmental constraints that determine the number of mammalian vertebrae, and, finally, the functional requirements of an adaptation to bipedal locomotion in the human lineage.

3D virtual reconstruction of the Kebara 2 Neandertal thorax.

The size and shape of the Neandertal thorax has been debated since the first discovery of Neandertal ribs more than 150 years ago, with workers proposing different interpretations ranging from a Neandertal thoracic morphology that is indistinguishable from modern humans, to one that was significantly different from them. Here, we provide a virtual 3D reconstruction of the thorax of the adult male Kebara 2 Neandertal. Our analyses reveal that the Kebara 2 thorax is significantly different but not larger from that of modern humans, wider in its lower segment, which parallels his wide bi-iliac breadth, and with a more invaginated vertebral column. Kinematic analyses show that rib cages that are wider in their lower segment produce greater overall size increments (respiratory capacity) during inspiration. We hypothesize that Neandertals may have had a subtle, but somewhat different breathing mechanism compared to modern humans.

First data of Neandertal bird and carnivore exploitation in the Cantabrian Region (Axlor; Barandiaran excavations; Dima, Biscay, Northern Iberian Peninsula).

Neandertals were top predators who basically relied on middle- to large-sized ungulates for dietary purposes, but there is growing evidence that supports their consumption of plants, leporids, tortoises, marine resources, carnivores and birds. The Iberian Peninsula has provided the most abundant record of bird exploitation for meat in Europe, starting in the Middle Pleistocene. However, the bird and carnivore exploitation record was hitherto limited to the Mediterranean area of the Iberian Peninsula. Here we present the first evidence of bird and carnivore exploitation by Neandertals in the Cantabrian region. We have found cut-marks in two golden eagles, one raven, one wolf and one lynx remain from the Mousterian levels of Axlor. The obtaining of meat was likely the primary purpose of the cut-marks on the golden eagle and lynx remains. Corvids, raptors, felids and canids in Axlor could have likely acted as commensals of the Neandertals, scavenging upon the carcasses left behind by these hunter-gatherers. This could have brought them closer to Neandertal groups who could have preyed upon them. These new results provide additional information on their dietary scope and indicate a more complex interaction between Neandertals and their environment.

Morphological integration in the gorilla, chimpanzee, and human neck.

OBJECTIVES: Although integration studies are important to understand the evolution of organisms' traits across phylogenies, vertebral integration in primates is still largely unexplored. Here we describe and quantify patterns of morphological integration and modularity in the subaxial cervical vertebrae (C3-C7) in extant hominines incorporating the potential influence of size. MATERIALS AND METHODS: Three-dimensional landmarks were digitized on 546 subaxial cervical vertebrae from 141 adult individuals of Gorilla gorilla, Pan troglodytes, and Homo sapiens. Integration and modularity, and the influence of size effects, were quantified using geometric morphometric approaches. RESULTS: All subaxial cervical vertebrae from the three species show a strong degree of integration. Gorillas show the highest degree of integration; conversely, humans have the lowest degree of integration. Analyses of allometric regression residuals show that size is an important factor promoting integration in gorillas, with lesser influence in chimpanzees and almost no effect in humans. DISCUSSION: Results point to a likely ancestral pattern of integration in non-human hominines, whereby the degree of integration decreases from cranial to caudal positions. Humans deviate from this pattern in the cranialmost (C3) and, to a lesser extent, in the caudalmost (C7) vertebrae, which are less integrated. These differences can be tentatively related to the emergence of bipedalism due to the presence of modern human-like C3 in australopiths, which still preserve a more chimpanzee-like C7.

The Neandertal vertebral column 2: The lumbar spine.

Here we provide the most extensive metric and morphological analysis performed to date on the Neandertal lumbar spine. Neandertal lumbar vertebrae show differences from modern humans in both the vertebral body and in the neural arch, although not all Neandertal lumbar vertebrae differ from modern humans in the same way. Differences in the vertebral foramen are restricted to the lowermost lumbar vertebrae (L4 and L5), differences in the orientation of the upper articular facets appear in the uppermost lumbar vertebrae (probably in L1 and L2-L3), and differences in the horizontal angle of the transverse process appear in L2-L4. Neandertals, when compared to modern humans, show a smaller degree of lumbar lordosis. Based on a still limited fossil sample, early hominins (australopiths and Homo erectus) had a lumbar lordosis that was similar to but below the mean of modern humans. Here, we hypothesize that from this ancestral degree of lumbar lordosis, the Neandertal lineage decreased their lumbar lordosis and Homo sapiens slightly increased theirs. From a postural point of view, the lower degree of lordosis is related to a more vertical position of the sacrum, which is also positioned more ventrally with respect to the dorsal end of the pelvis. This results in a spino-pelvic alignment that, though different from modern humans, maintained an economic postural equilibrium. Some features, such as a lower degree of lumbar lordosis, were already present in the middle Pleistocene populations ancestral to Neandertals. However, these middle Pleistocene populations do not show the full suite of Neandertal lumbar morphologies, which probably means that the characteristic features of the Neandertal lumbar spine did not arise all at once.

The role of allometry and posture in the evolution of the hominin subaxial cervical spine.

Cervical vertebrae not only protect the spinal cord but also are the insertion and origin points for muscles related to the movement of the head, upper limb, and trunk, among others, and are thus important elements in primate evolution. While previous work has been undertaken on the first two cervical vertebrae, there is a dearth of studies on the subaxial cervical spine in hominines. In this paper, we provide detailed morphological information on two important aspects of the subaxial cervical vertebrae (C3 - C7): mid-sagittal morphology and superior facet orientation. We studied large samples of African apes including modern humans and the most complete fossil hominin subaxial cervical vertebrae using both traditional and geometric morphometrics. There are significant differences between extant hominoids related to the relative length and orientation of the spinous process as well as to the orientation of the articular facets, which are related to size, locomotion, and neck posture. In fact, fossil hominins do not completely conform to any of the extant groups. Our assessment of mid-sagittal morphology and superior articular facet orientation shows that australopiths have more Homo-like upper subaxial cervical vertebrae coupled with more "primitive" lower cervical vertebrae. Based on these results, we hypothesize that those changes, maybe related to postural changes derived from bipedalism, did not affect the entire subaxial cervical spine at once. From a methodological point of view, the combination of traditional and geometric morphometric data provides a more integrative perspective of morphological change and evolution, which is certainly useful in human evolutionary studies.