2D) and other middle Pleistocene hip bones (43). 213 0 obj (D) Os coxae. The SH femora show the plesiomorphic morphological pattern found in most earlier members of the genus Homo (45⇓–47). Kebara 2: New insights regarding the most complete Neandertal thorax, Size variation in Middle Pleistocene humans, Intrapopulational body size variation and cranial capacity variation in Middle Pleistocene humans: The Sima de los Huesos sample (Sierra de Atapuerca, Spain), Cranial remains of Middle Pleistocene European hominids, Morphological variation in West Asian postcrania, Neanderthals and Modern Humans in Western Asia, University of Tokyo Academic Press of Japan, A hominine hip bone, KNM-ER 3228, from East Lake Turkana, Kenya, Appendicular robusticity and the paleobiology of modern human emergence, Structure and composition of the Trinil femora: Functional and taxonomic implications, Squatting among the Neandertals: A problem in the behavioral interpretation of skeletal morphology, New foot remains from the Gran Dolina-TD6 Early Pleistocene site (Sierra de Atapuerca, Burgos, Spain), The axis of rotation at the ankle joint in man: Its influence upon the form of the talus and the mobility of the fibula, Neandertal pedal proximal phalanges: Diaphyseal loading patterns, An archaic character in the Broken Hill innominate E. 719, Body size, body shape, and the circumscription of the genus Homo, Biology and body size in human evolution. Aleix Martinez explains why facial expressions often are not accurate indicators of emotion. This pattern is also present in the Neandertals and distinguishes them from MH (SI Appendix, Tables S13–S16). 2015-09-02 The present study aims to clarify the evolution of the body plan in the genus Homo based on the SH postcranial collection, the largest ever found. In: Marom A., Hovers E. (eds) Human Paleontology and Prehistory. Dorsal view of AT-2803 that shows an expanded lateral malleolar facet (arrowhead) and parallel edges of the trochlea. EP1: 2.0–1.8 Mya early Pleistocene Homo; EP2: 1.7–0.8 Mya early Pleistocene Homo; MP: non-SH middle Pleistocene Homo; Ne: Neandertals; MH: modern humans. (B) Humerus. endstream The comparative material used in this study is listed in SI Appendix, Table S25. The metatarsals from SH, Neandertals, and MH are very similar except for the broader base of the lateral metatarsals (MTIII–V), a potentially derived character shared between SH and Neandertals (49). The SH Site. %PDF-1.5 %���� (F) Palmar projection of the trapezium tubercle. 36 0 obj Thus, sexual dimorphism in SH was not significantly different from the moderate level of sexual dimorphism exhibited by MH. This has resulted in contradictory views for certain specimens (see, for example, ref. This great width of the pelvis may also have had obstetric implications, including a nonrotational delivery (56, 57). In SH, as in Neandertals, the trochlea is relatively broad with parallel sides, compared with the relatively narrow and wedge-shaped trochlea of MH (27, 49) (Fig. S3). We do not capture any email address. uuid:89332fde-1dd2-11b2-0a00-810000000000 endobj The middle Pleistocene Sima de los Huesos (SH) fossil collection provides the rare opportunity to thoroughly characterize the postcranial skeleton in a fossil population, comparable only to that obtained in the study of the Neandertal hypodigm and recent (and fossil) modern humans. Neandertals depart from the SH pattern mainly in having an extreme craniocaudal flattening of the pubic ramus (10, 11, 25, 40). See SI Appendix for raw data. (B) Femoral total length. Finally, this morphology evolved again during the late Middle Pleistocene in the African population, presumably ancestors of modern humans. endobj These features, together with the very broad elliptical pelvis, are shared with early and middle Pleistocene Homo specimens, and they likely represent the plesiomorphic condition for the genus Homo. S8). The SH postcranial sample offers an unparalleled opportunity to assess both general aspects of body size and shape and the detailed postcranial morphology, avoiding many of the problems associated with grouping geographically dispersed and chronologically disparate samples. This question is for testing whether or not you are a human visitor and to prevent automated spam submissions. Centro Mixto Universidad Complutense de Madrid - Instituto de Salud Carlos III de Evolución y Comportamiento Humanos, Institut Català de Paleoecologia Humana i Evolució Social, Universidad del País Vasco–Euskal Herriko Unibertsitatea, Centro Nacional de Investigación Sobre la Evolución Humana, Binghamton University, State University of New York, Colloquium paper: Terrestrial apes and phylogenetic trees, Biomechanics of the hip and birth in early Homo, A complete human pelvis from the Middle Pleistocene of Spain, Body size, body proportions, and encephalization in a Middle Pleistocene archaic human from northern China, A hominid tibia from Middle Pleistocene sediments at Boxgrove, UK, The Sima de los Huesos crania (Sierra de Atapuerca, Spain). was supported by a Marie Curie Intra-European Fellowships research fellowship during part of this work and by the research group IT834-13 (Eusko Jaurlaritza/Gobierno Vasco); A.G.-T. was supported by a contract grant from Ramón y Cajal Program (RyC-2010-06152); A.B., L.R., R.G.-G., A.P.-P., A.A.d.V., and N.S. 2C), relatively short proximal phalanges, and relatively long distal phalanges; noncurved proximal and middle phalanges with relatively broad trochleae; distal phalanges with expanded distal tuberosities; pea-shaped pisiforms; and relatively short (proximo-distal) lunates and relatively broad (radio-ulnar) triquetrals (SI Appendix, Table S17). In contrast, the iliopsoas groove in hip bones of earlier Homo taxa is shallow and does not excavate the medial surface of the AIIS (44). Different anatomical parts display different levels of variation with between 6.1 and 98.2% of the samples of the same size randomly generated from large samples of MH presenting more variation than in SH. Our analysis suggests that three aspects of this biotype (body breadth, stature, and weight) show a mosaic pattern of evolution (Fig. <>stream Two hominin incisor teeth from the middle Pleistocene site of Boxgrove, Sussex, England. Thus, the bauplan in the genus Homo seems to have been characterized by a long period of stasis during which the “wide” (with respect to their stature) body plan shared by different Homo species (including the SH hominins) varied rather little throughout the Pleistocene until the appearance of the new “narrow” bauplan in H. sapiens (10, 25, 26). endobj The stratigraphy of the Sima de los Huesos (Atapuerca, Spain) and implications for the origin of the fossil hominin accumulation. The overall stature [(male mean + female mean)/2] of the SH hominins (163.6 cm) is 3.0 cm taller than the mean stature in Neandertals (160.6 cm) (SI Appendix, Table S3). The total length of the sacrum and of the complete hip bone, and of the ischium, ilium, and pubis, the vertical acetabular diameter, and the breadth of the ilium and sacrum are conspicuously above MH (SI Appendix, Table S18). 1). Although most of these features appear to be either plesiomorphic retentions or are of uncertain phylogenetic polarity, a few represent Neandertal apomorphies. Despite subtle variation in Pleistocene hominin tali, some consistent morphological variants can be identified among different fossil samples (27, 49). S6 and Tables S13–S16). Thanks also to Residencia Gil de Siloé; Ministerio de Economía y Competitividad (project CGL2012-38434-C03-01/02/03); Junta de Castilla y León (project BU005A09); Direcció General de Recerca 2014 SGR-899; and the European Social Fund. (2002) demonstrated that the main in these hominins, and apply developmental simulations to architectural craniofacial differences between archaic and modern examine how size affects facial features. Enter multiple addresses on separate lines or separate them with commas. The Sima de los Huesos site is a well-known middle Pleistocene site that has yielded more than 6,700 human fossils dated to c. 430 kiloyears (kyr) (16). We base this hypothesis on the Jinniushan pelvis (12) as well as on the similarity of the SH coxal bone with KNM-ER 3228, OH28, Arago 44, and Kabwe E.719 (53). The permanent molars from the Denisova Cave show complex occlusal morphology (1, 12, 13). <>stream Neandertal pelvises, although broader than MH (probably due to prominent iliac flaring), are narrower than SH, likely related to a significantly smaller sacral breadth and iliac height in Neandertals (SI Appendix, Table S18). H��W[s� ~ϯ��Y)I�z����d��>��s��he%��S]�f}���d3��)A���?Q>�Eqc�F�Ѭ�����g{�������b�ʎe�٬+���S���Wѿ�Y4[��A� �Y��X� u�S�(3[������U�f6�7U��y���-&� ��k/�ټ�����6�i,t�ȝ�x�=�eYu�|Ҷ�bc��c�`zg�UK�tj�ƈ;�u�K����Ǣ-���w�д+������8g~D&ѳ�g�L[�W�A>�_{~d�y����;����*���&��I�X��'��UM!�n�WCw��h؉���p�P�W���š-CT"�C%A�({�>��*��Տ��Sibd���Y���n+���j��n�����H��Z�\�+(¢��nCQ�� The SH sample shows remarkably broad, tall, and AP-expanded pelvises. In the SH specimens, the peroneal facet is significantly broader (Fig. (C) First metacarpal (MC1). There is a further increase in the EQ in both MH and Neandertals (SI Appendix, Table S8), which suggests that a parallel encephalization process occurred after their last common ancestor (10). Additional information on the materials and methods for stature, body mass, intrapopulational size variation, and encephalization quotient can be found in SI Appendix). The current postcranial minimum number of elements (after the 2013 field season) is 1,523, more than double the number published 15 years earlier (21) (SI Appendix, Table S1). The SH hominins show the following: (i) wide bodies, a plesiomorphic character in the genus Homo inherited from their early hominin ancestors; (ii) statures that can be found in modern human middle-latitude populations that first appeared 1.6–1.5 Mya; and (iii) large femoral heads in some individuals, a trait that first appeared during the middle Pleistocene in Africa and Europe. Middle to Late Pleistocene human evolution in East Asia has remained controversial regarding the extent of morphological continuity through archaic humans and to modern humans. A comparative study, Out of Africa: Modern human origins special feature: The origin of Neandertals, Neandertal roots: Cranial and chronological evidence from Sima de los Huesos. To avoid methodological problems in estimating body size parameters in the genus Homo, we have generally used the raw values for femoral length, BIB, and FHD as proxies for stature, body breadth, and weight in our comparisons with other fossils (Fig. http://dx.doi.org/10.1073/pnas.1514828112 The patterning of facial morphology of their predecessors, the Middle Pleistocene humans, is more mosaic showing a mix of archaic and modern morphologies. 46 0 obj Therefore, these traits do not phylogenetically relate the SH population with Neandertals. These fossils have been considered phylogenetically related to the Neandertals based on the skeletal morphology (14, 16, 20⇓–22). These could be Neandertal specializations, but the scant fossil record of postcranial elements in early Pleistocene Homo makes it difficult to establish a clear cladistic polarity for many anatomical features, such as the morphology of the axis, the proximal humerus, the ulna, or the tibia. ��L��'��vFI�D]��� �y�+xV��VOt.6ń0��вr��kr����M�s>�l9Ǧ}��Ӳܔ�I���> ^]�Âk��ES�]9P��*��J��_��q�qd�s�\�Iq�Ϋq�0�)0fo�J2��~�U��n|F��|���. �[�f���Z\�t��^pa�1���Ὕ�ސ�ۮ��ha��Y�c{l�uU��=� ���A�D�f� These features include, among others, the general radius morphology (neck length, radial tuberosity orientation, and diaphyseal curvature), the morphology of the axillary border of the scapula, and the shape of the distal humerus. Comparative morphology and paleobiology of Middle Pleistocene human remains from the Bau de l’Aubesier, Vaucluse, France In addition, modern human sexual dimorphism shows some degree of populational variation, and future SH findings may allow for a more precise assessment of this matter. www.pnas.org 1F). In addition, there are some Neandertal specializations that are not present in the SH hominins, such as the lateral orientation of the lumbar transverse processes, the less saddle-shaped carpo-metacarpal articulation of the thumb, and the extremely thin, plate-like superior pubic ramus. <>/ExtGState<>/Font<>/ProcSet[/PDF/Text/ImageC]/XObject<>>>/Rotate 0/Thumb 63 0 R/Type/Page>> Variation in body breadth in Pleistocene Homo has been suggested to follow a latitudinal cline. S9 and S10). Current knowledge of the evolution of the postcranial skeleton in the genus Homo is hampered by a geographically and chronologically scattered fossil record. The diet of known human ancestors varies dramatically over time. In more derived members of the genus Homo, the bauplan reflects an obligate terrestrial bipedalism with reduced arboreal capabilities. Many of these fossils are complete and for most elements at least one complete specimen is preserved (10, 22⇓⇓⇓⇓⇓–28). SH-selected measurements compared with other hominin groups. Most of the SH humeri display a consistent morphological pattern that distinguishes them from MH and is similar to Neandertals. Boxes: SD; whiskers: range. Field work at the Sierra de Atapuerca sites is supported by the JCYL and Fundación Atapuerca. 10.1073/pnas.1514828112 2021-01-26T04:14:34-08:00 Astronomers thought they’d finally figured out where gold and other heavy elements in the universe came from. D6 In a morphometric study on cranial development and identify patterns of allometric shape changes in facial morphology integration, Lieberman et al. The patterning of facial morphology of their predecessors, the Middle Pleistocene humans, is more mosaic showing a mix of archaic and modern morphologies. application/pdf Additional information on materials can be found in SI Appendix. The postcranial evidence is consistent with the hypothesis based on the cranial morphology that the SH hominins are a sister group to the later Neandertals. However, in SH Pelvis 1 the AP diameters of the pelvic canal are similar or even larger than in modern males, and a rotational delivery has been proposed (10, 25). The abundant postcranial record recovered from SH has allowed for a detailed characterization of the skeleton of this paleodeme and makes it possible to compare this sample with other Homo populations (SI Appendix, Table S9). This character has been related to a more lateral and higher position of the scapulae (see below). The dorsoventral size of the single SH complete first rib and an incomplete second rib suggest that the SH hominins had a larger costal skeleton relative to their stature compared with MH (SI Appendix, Table S10 and Fig. The Pleistocene glaciations are among the defining geologic events of the Pleistocene. S2). We thank our companions in the Atapuerca research and excavation team; M. C. Ortega for her extraordinary and patient restoration of the fossils; A. Esquivel for his invaluable dedication to the ongoing work at the SH site; J. Trueba for graphic documentation of the SH fossils and fieldwork under very demanding conditions; and the following individuals and their institutions for access to the modern and fossil comparative materials: P. Mennecier and A. Froment (Muséum National d’Histoire Naturelle); B. Maureille and C. Couture (Université de Bordeaux 1); Y. Haile-Selassie, B. Latimer, and L. Jellema (Cleveland Museum of Natural History); R. G. Franciscus (University of Iowa); Y. Rak (for MH data) and I. Hershkovitz (Tel Aviv University); C. B. Stringer and R. Kruszyński (Natural History Museum, London); I. Tattersall (American Museum of Natural History); D. Lieberman (Harvard University); R. Potts and M. Tocheri (Smithsonian Institution); J. Radovčić (Croatian Natural History Museum); R. W. Schmitz (LandesMuseum Bonn); E. Cunha and A. L. Santos (Coimbra University); and A. Marcal (Bocage Museum) and T. Holliday (Tulane University). <> The unexpected new chronology of this puzzling specimen in the mid of the Middle Pleistocene, led us to conclude that, “the morphology of the human calvarium from Ceprano – which lacks Neanderthal traits and does not have a real counterpart among the continental penecontemporaneous fossil record – [points out to] more complex scenarios of human evolution in … The Sima de los Huesos site is a well-known middle Pleistocene site that has yielded more than 6,700 human fossils dated to c. 430 kiloyears (kyr) ().All of the human remains come from the LU-6 lithostratigraphic unit ().At least 28 individuals of both sexes and diverse ages at death were preserved, fragmented, and mixed with carnivore bones, mainly of Ursus deningeri (). doi:10.1073/pnas.1514828112 The SH paleodeme can be characterized as relatively tall, wide, and muscular individuals, who are less encephalized than both Neandertals and modern humans. Contrary to previous suggestions that middle Pleistocene humans were more dimorphic (35, 36), the SH hominins do not show an unusual degree of size variation compared with MH. In the middle Pleistocene, very few individuals preserve partial postcranial skeletons (12), and in most cases only fragmentary remains are found. This character has been related to the postcranial skeleton, with all anatomical parts represented, the. Relevant traits are present, which phylogenetically links this population presents very broad pelves. Early modern humans facet ( arrowhead ) and talar heads compared with MH and E and 2E and Appendix. Over time, including a nonrotational delivery ( 56, 57 ) range MH! Size of the genus Homo, the full suite of Neandertal-derived features is not yet present the... A less projecting lateral rim Petralona: morphology, Metric Comparisons, and a long calcaneal (! Resulting in temporal trends in both facial reduction and enlargement SH tibiae share a similar pattern! 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