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middle pleistocene humans’ morphology

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endobj Postcranial skeleton 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. The site, The temporal bones from Sima de los Huesos Middle Pleistocene site (Sierra de Atapuerca, Spain). This was followed by a subsequent further increase in the EQ in Neandertals and MH. received financial support from Binghamton University (SUNY) and the American Museum of Natural History; E.P.-R. was supported by a Comunidad Autónoma de Madrid Grant S2010/BMD-2330; and L.R. This character has been related to a more lateral and higher position of the scapulae (see below). However, the dorsoventral size of the single complete first rib is longer than MH and Neandertals, and an incomplete second rib suggests that it was dorso-ventrally longer than that of Kebara 2. C. M. Fitzgerald and S. W. Hillson. 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. It is apparent from the suite of Neandertal lineage features present in the Aubesier Middle Pleistocene human remains that they provide further evidence for the gradual emergence during the second half of the Middle Pleistocene and the early Late Pleistocene of Europe of the derived (at least in frequency) Neandertal morphological pattern. Body mass (BM) can be reconstructed from hominin skeletal remains using both morphometric [stature and bi-iliac breadth (BIB)] (29) or mechanical approaches (joint surface size of weight-bearing skeletal elements) (30). Abstract. Although there are no known pelvic remains attributed to H. habilis, in our view, a ML relatively wide biotype was likely present (as the most parsimonious interpretation) in the earliest members of the genus Homo and was inherited from their early hominin ancestors. Thus, the full suite of Neandertal features did not arise all at once, and the evolution of the postcranial skeleton could be characterized as following a mosaic pattern. Britain was a peninsula and the Indonesian islands were connected to the Southeast Asian mainland, for example. The SH sample shows remarkably broad, tall, and AP-expanded pelvises. Regarding the thorax, the absence of complete midthoracic ribs makes it difficult to assess whether the size and shape of the SH costal skeleton is similar to that of Neandertals (32). The morphology of the proximal ulna has been shown to effectively differentiate archaic or premodern humans (such as Homo heidelbergensis and H. neanderthalensis) from modern humans (H. sapiens). Copyright © 2021 National Academy of Sciences. Later, some populations moved north to Europe where cold adaptation eventually led to the evolution of H. neanderthalensis. (A) Third lumbar vertebra (L3). Thus, the full suite of Neandertal features did not arise all at once, and the evolution of the postcranial skeleton could be characterized as following a mosaic pattern. 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). A team of scientists led by LIU Wu and WU Xiujie from the Institute of Vertebrate Paleontology and Paleoanthropology (IVPP) of the Chinese Academy of Sciences reported the first ever Middle Pleistocene human skull found in southeastern China, revealing the variation and continuity in early Asian humans. Subadult (H-IV, Left) and adult (H-VI, Right) specimens showing the thin medial pillar and broad and deep olecranon fossa. Although the evidence is still limited, a growing body of research suggests music may have beneficial effects for diseases such as Parkinson’s. In light of recent results, they’re not so sure. A.G.-O. 10.1073/pnas.1514828112 Reviewers: T.W.H., Tulane University; and C.B.R., The Johns Hopkins University School of Medicine. 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. The SH sample shows a dominant dorsal position (n = 8) of the axillary sulcus for the Musculus teres minor (on the axillary border), resembling the predominant condition in Neandertals. 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. 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 (). A phylogenetic approach, Clavicles, scapulae and humeri from the Sima de los Huesos site (Sierra de Atapuerca, Spain), Metric and morphological study of the upper cervical spine from the Sima de los Huesos site (Sierra de Atapuerca, Burgos, Spain), Middle Pleistocene lower back and pelvis from an aged human individual from the Sima de los Huesos site, Spain, Stature estimation from complete long bones in the Middle Pleistocene humans from the Sima de los Huesos, Sierra de Atapuerca (Spain), Human talus bones from the Middle Pleistocene site of Sima de los Huesos (Sierra de Atapuerca, Burgos, Spain), Human calcanei from the Middle Pleistocene site of Sima de los Huesos (Sierra de Atapuerca, Burgos, Spain), Body mass and encephalization in Pleistocene. 2015-09-02T03:51:26+05:30 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. The intrapopulational size variation in SH shows that the level of dimorphism was similar to modern humans (MH), but the SH hominins were less encephalized than Neandertals. 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. August 2015; Proceedings of the National Academy of Sciences 112(37) DOI: 10.1073/pnas.1514828112. designed research; J.L.A., J.-M.C., C.L., A.G.-O., A.P., L.R., R.G.-G., A.B., R.M.Q., A.P.-P., I.M., A.A., A.G.-T., E.P.-R., N.S., N.G., A.A.d.V., G.C.-B., J.M.B.d.C., and E.C. Most of the SH humeri display a consistent morphological pattern that distinguishes them from MH and is similar to Neandertals. (D) Os coxae. Unfortunately, our understanding of the evolution and variation of body size and shape in Pleistocene Homo before the Neandertals is still quite limited due to a fragmentary and geographically and chronologically scattered fossil record. 69 0 obj <> www.pnas.org In the SH hand, like in Neandertals, the powerful precision grip is enhanced by the thumb robusticity and well-developed flexor musculature. In these two latter traits, the specimens show some variation. 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. “On the basis of preserved morphology, BH-1 differs significantly from Middle Pleistocene European hominins generally grouped under Homo heidelbergensis. Evidence from the shoulder girdle, the thorax, and the pelvis points to a wide and large body type in the SH hominins. 60/femoral maximum length × 100). Ther… 1F). (C) First metacarpal (MC1). Newly found ∼300,000-y-old human remains from Hualongdong (HLD), China, … S2). The SH hand morphology indicates a powerful precision grip and fine precision grasping capabilities that are similar to what has been described in Neandertals (39) and MH. S6 and Tables S13–S16). In general, the body plan in the genus Homo has been largely characterized by stasis since ∼1.6 Mya until the appearance of MH (2). 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). Despite large periods of morphological stasis in the general body plan, the anatomical details of the postcranial skeleton, as revealed in the SH sample, offer the best evidence for a pattern of mosaic evolution in the postcranium within the Neandertal lineage. These fossils have been considered phylogenetically related to the Neandertals based on the skeletal morphology (14, 16, 20⇓–22). The Neandertal talus displays broader lateral malleolar facets (50) and talar heads compared with MH. 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). �[�f���Z\�t��^pa�1���Ὕ�ސ�ۮ��ha��Y�c{l�uU��=� ���A�D�f� The paleontological description and comparative analysis using discrete morphology, morphometrics (linear and geometric) and cross‐sectional geometry of three femoral diaphyseal sections from the Middle Pleistocene site of Hualongdong, China. This suggests that the SH hominins, like Neandertals, had a larger costal skeleton relative to their stature compared with MH (see below). At least 28 individuals of both sexes and diverse ages at death (18) were preserved, fragmented, 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. Although the use of the FHD rather than the BIB (see above) yields lower BM values and, consequently, higher EQ values, the EQ from the SH sample is still significantly lower than that of Neandertals (P < 0.003) and MH (P < 0.006). (E) Femur. The timing and routes of modern human migration out of Africa are key issues for understanding the evolution of our own species. 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). Our analysis suggests that three aspects of this biotype (body breadth, stature, and weight) show a mosaic pattern of evolution (Fig. The appearance of this “narrow” bauplan has energetic implications, which have been invoked as one of the reasons for the success of our species (58), although the major change in relative skeletal strength (lower-limb diaphyseal cross-sectional geometry) within Homo may have taken place after, not at, the origin of H. sapiens (59). endobj In: Marom A., Hovers E. (eds) Human Paleontology and Prehistory. 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. See SI Appendix for raw data. Field work at the Sierra de Atapuerca sites is supported by the JCYL and Fundación Atapuerca. In all of the anatomical details of the upper and lower limb, the SH immature individuals follow the same pattern as in the adult specimens (SI Appendix, Tables S14 and S20). The curvatures of the SH clavicles in the transverse plane fall within the normal variation in MH. The patterning of facial morphology of their predecessors, the Middle Pleistocene humans, is more mosaic showing a mix of archaic and modern morphologies. 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). <>/ExtGState<>/Font<>/ProcSet[/PDF/Text/ImageC]/XObject<>>>/Rotate 0/Thumb 15 0 R/Type/Page>> This body shape is also largely present in other early and middle Pleistocene individuals and in Neandertals. 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. Although most of these features appear to be either plesiomorphic retentions or are of uncertain phylogenetic polarity, a few represent Neandertal apomorphies. The SH hominins could be included within the “wide Homo” bauplan due to their absolutely and relatively large and ML-wide biotype consisting of a large thorax with broad shoulders and pelvises, above-medium-height body, thick bones, and great musculature and body mass. Although middle Pleistocene populations have been described as exceptionally robust (13), phylogenetic hypotheses are based mainly on the more abundant cranial sample (14, 15). Ventral view of AT-1000, displaying a strongly twisted anterior inferior iliac spine (white arrow) and a deep iliopsoas groove (black arrow). We will characterize the general body size and shape [stature, body breadth, body mass, and encephalization quotient (EQ)] in the SH paleodeme within the context of postcranial evolution in the genus Homo. codirected the Atapuerca excavations and research project; J.L.A. The SH glenoid cavity is consistently taller and narrower (n = 10) than in MH, reflected in a low glenoid index (SI Appendix, Fig. performed research; J.L.A., J.-M.C., C.L., A.G.-O., A.P., L.R., R.G.-G., A.B., R.M.Q., A.P.-P., I.M., A.A., A.G.-T., E.P.-R., N.S., N.G., A.A.d.V., and G.C.-B. Rightmire G.P. (2013) invoked evolutionary convergence for the above modern sapiens -like facial morphology in several places and times during the Pleistocene. 10.1073/pnas.1514828112 Acrobat Distiller 10.0.0 (Windows) 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). The SH femora show the plesiomorphic morphological pattern found in most earlier members of the genus Homo (45⇓–47). http://dx.doi.org/10.1073/pnas.1514828112 All of the human remains come from the LU-6 lithostratigraphic unit (17). The pronounced maxillary incisor beveling of Aubesier 4 helps to extend the antiquity of nondietary use of the anterior dentition. The authors declare no conflict of interest. Boxes: SD; whiskers: range. 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). Although there appears to be a somewhat elevated level of intrapopulational variation and sexual dimorphism in the SH sample when we compare the BM values to that of modern humans (SI Appendix, Table S4), this result is based on the still relatively small sample of femoral head estimates (n = 5) in SH. Additional information on the materials and methods for stature, body mass, intrapopulational size variation, and encephalization quotient can be found in SI Appendix). Unlike MH, the anterior inferior iliac spine (AIIS) of the Neandertals is medially twisted relative to the anterior margin of the iliac blade and is bordered by a deep iliopsoas groove that excavates the medial surface of the AIIS (41, 42). In order to better evaluate the modern human-like facial fea-tures on ATD6-69 several issues need to be clarified. This pattern includes a transversely oval humeral head, a projected and massive lesser trochanter, a narrower deltoid tuberosity with two muscular crests, thick cortical diaphyses (Fig. Thus, the rare Denisovan human remains identified to date show affinity to Middle Pleistocene hominins (2, 12, 13), particularly to those from China and, to a lesser extent, to the Neanderthal lineage . endobj 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. Am J Phys Anthropol. This pattern is also present in the Neandertals and distinguishes them from MH (SI Appendix, Tables S13–S16). (F) Palmar projection of the trapezium tubercle. The comparative material used in this study is listed in SI Appendix, Table S25. A mosaic pattern was also documented in the SH cranium (16) although, in this case, the Neandertal suite of derived features forms a single functional complex. The permanent molars from the Denisova Cave show complex occlusal morphology (1, 12, 13). Further studies (8, 9) and the discovery of additional fossil evidence (10, 11) support the idea that the original reconstruction of the pelvis of KNM WT-15000, and thus a number of the interpretations based on it, need to be reconsidered. Therefore, these traits do not phylogenetically relate the SH population with Neandertals. 1A) characterized by very wide sacra, pronounced lateral iliac flaring, and long pubic rami that clearly separate it from the MH pelvic configuration (see below). endobj Using a randomization method, relying on bootstrapping, the size variation in the SH hominins was studied as a proxy for their level of sexual dimorphism (33, 34), including additional anatomical parts that were previously underrepresented (SI Appendix, Tables S5–S7). 105 0 obj Thus, sexual dimorphism in SH was not significantly different from the moderate level of sexual dimorphism exhibited by MH. 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