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. <>/ExtGState<>/Font<>/ProcSet[/PDF/Text/ImageC]/XObject<>>>/Rotate 0/Thumb 15 0 R/Type/Page>> The SH postcranial sample up to the 2013 field season is composed of 1,523 elements (SI Appendix, Table S1). <>/ExtGState<>/Font<>/ProcSet[/PDF/Text/ImageC]/Properties<>/XObject<>>>/Rotate 0/Thumb 71 0 R/Type/Page>> 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. In a morphometric study on cranial development and identify patterns of allometric shape changes in facial morphology integration, Lieberman et al. In more derived members of the genus Homo, the bauplan reflects an obligate terrestrial bipedalism with reduced arboreal capabilities. 10.1073/pnas.1514828112 Researchers are still trying to understand what causes this strong correlation between neural and social networks. <>stream 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). 1. middle pleistocene: 750,000 to 25,0000ya -most plesitocene hominoids lived during this time period 2. upper pleistocene: 125,000-10,000ya-later premodern humans and neanderthals lived into this period-often called ice age-marked by advances and retreats of massive continental glaciations endobj The higher EQ of the SH population compared with the published values in the early Pleistocene Dmanisi hominins (37) demonstrates that the increase in brain size in SH was not simply a consequence of an increase in body mass (29). 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. 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. 2015-09-02T03:51:26+05:30 Journal of Human Evolution 59, 2010, 493-503. A few features that have been considered Neandertal-derived traits are also present in the SH hominins, including a low degree of lumbar lordosis, broad distal tuberosities of the manual phalanges, and the wide bases of the lateral metatarsals (MTIII–V), which is consistent with the hypothesis, based on the cranial morphology, that the SH hominins are a sister group to the later Neandertals (16). The pronounced maxillary incisor beveling of Aubesier 4 helps to extend the antiquity of nondietary use of the anterior dentition. 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). The size of the ice sheets resulted in lower sea levels and dryer climates. In addition, we focus in particular on whether the detailed morphological traits found throughout the postcranial skeleton follow a mosaic pattern of evolution, as seen in the crania, and whether there have been changes in the Homo bauplan. 1A). Palaeodemography of the Atapuerca-Sima de los Huesos Middle Pleistocene hominid sample. (2013) invoked evolutionary convergence for the above modern sapiens -like facial morphology in several places and times during the Pleistocene. analyzed data; and 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., and I.M. The mean stature of the SH hominins has been estimated based on 24 complete long bones from the upper and lower limbs (26). Contributed by Juan Luis Arsuaga, July 29, 2015 (sent for review May 20, 2015; reviewed by Trenton W. Holliday and Christopher B. Ruff). 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. 1F). In the middle Pleistocene, very few individuals preserve partial postcranial skeletons (12), and in most cases only fragmentary remains are found. Cranial view of VL2, presenting a very long and dorso-laterally oriented transverse process (arrowhead). The preservation of all anatomical parts in SH has allowed a detailed characterization of the postcranial anatomy and has revealed that the SH hominins share many anatomical features with Neandertals not present in MH. 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. 10.1073/pnas.1514828112 This question is for testing whether or not you are a human visitor and to prevent automated spam submissions. F-X (Left) and F-XI (Right) proximal femora in posterior view, showing a low neck angle, large gluteal ridges, and well-developed hypotrochanteric fossae. 10.1073/pnas.1514828112 Later, some populations moved north to Europe where cold adaptation eventually led to the evolution of H. neanderthalensis. This is consistent with previous hypotheses of an anthropic origin for this accumulation (21). 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 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). Additional information on materials can be found in SI Appendix. Subadult (H-IV, Left) and adult (H-VI, Right) specimens showing the thin medial pillar and broad and deep olecranon fossa. However, two SH specimens show the relatively short and robust neck, anteriorly oriented radial tuberosity, and the straight and robust shaft typical of MH (SI Appendix, Fig. 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 first of these may represent a partially arboreal, facultative biped, if the genus Ardipithecus (and perhaps Orrorin) is included within the hominins. 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). 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. Two hominin incisor teeth from the middle Pleistocene site of Boxgrove, Sussex, England. 163 0 obj Comparative morphology and paleobiology of Middle Pleistocene human remains from the Bau de l’Aubesier, Vaucluse, France 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). 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). 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. This character has been related to a more lateral and higher position of the scapulae (see below). 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. uuid:89332fda-1dd2-11b2-0a00-7a08275dc400 endstream 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). 03-461AA-692.01). 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. The SH tibiae share a similar morphological pattern with Neandertals that includes large retroversion angle of the proximal epiphysis (SI Appendix, Fig. NOTE: We only request your email address so that the person you are recommending the page to knows that you wanted them to see it, and that it is not junk mail. 2007). This body shape is also largely present in other early and middle Pleistocene individuals and in Neandertals. 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). Comparative morphology and paleobiology of Middle Pleistocene human remains from the Bau de l'Aubesier, Vaucluse, France 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. This great width of the pelvis may also have had obstetric implications, including a nonrotational delivery (56, 57). Most of the SH radii (six of eight specimens) display a relatively long and gracile neck, an antero-medially oriented radial tuberosity, and a low robusticity index with a pronounced ML shaft curvature, as in the Neandertals. Much of North America was covered by the Laurentide ice sheet and northern Europe and Siberia were covered by the Eurasian Ice Sheet Complex. The australopith bauplan (present in both Australopithecus and Paranthropus) mainly reflects terrestrial bipedalism, coupled with suspensory and climbing activities (3) that could have also been present in Homo habilis (4). 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 BM of one large male (Pelvis 1 individual) calculated from stature and BIB is between 90.3 and 92.5 kg (25), and the Pelvis 2 individual seems to be slightly broader (Fig. Accordingly, the morphology of adjacent, articulating elements should be able to distinguish these two b … The fossil evidence suggests that the earliest members of the Homo sapiens clade (Jebel Irhoud, Omo, and Herto) appeared in Africa during the late Middle Pleistocene (1–4).Outside Africa, modern humans appeared much later, during the Late Pleistocene … In contrast, MH show three curvature types (31). The SH pelvises are characterized by their marked robusticity (e.g., large sacroiliac joint, iliac tubercle, and ischial tuberosity) and large overall dimensions. The SH sample shows remarkably broad, tall, and AP-expanded pelvises. At least 28 individuals of both sexes and diverse ages at death (18) were preserved, fragmented, and mixed with carnivore bones, mainly of Ursus deningeri (19). Nine EQ values have been calculated for the SH adult crania (16) using the femoral head diameter (FHD) to calculate BM (SI Appendix, Table S8) and yield a mean EQ of 3.54. 2021-01-26T04:14:34-08:00 All of the human remains come from the LU-6 lithostratigraphic unit (17). false endobj www.pnas.org endobj Boxes: SD; whiskers: range. 213 0 obj Significant changes in facial size and robusticity occurred throughout Pleistocene human evolution, resulting in temporal trends in both facial reduction and enlargement. Krapina and other Neanderthal clavicles: A peculiar morphology? We do not capture any email address. S3). Quizlet flashcards, activities and games help you improve your grades. 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. Thank you for your interest in spreading the word on PNAS. The SH pelvic remains are also distinct from MH in having an anteriorly located acetabulocristal buttress, a well-developed supraacetabular groove and a thin and rectangular, plate-like superior pubic ramus that contrasts with the thick and stout pubis of MH (10, 25) (SI Appendix, Figs. 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. <>stream The SH glenoid cavity is consistently taller and narrower (n = 10) than in MH, reflected in a low glenoid index (SI Appendix, Fig. 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. (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. 105 0 obj S7). 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 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. The recent analysis of 17 crania from Sima de los Huesos (SH) points to a mosaic pattern of evolution in the cranium, with facial modification being the first step in the evolution of the Neandertal clade (16). uuid:89332fde-1dd2-11b2-0a00-810000000000 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 permanent molars from the Denisova Cave show complex occlusal morphology (1, 12, 13). S12 and Tables S19–S22), tibial condyles located in a more posterior position in relation to the axis of the shaft, and flat proximal and distal articular surfaces. Ventral view of AT-1000, displaying a strongly twisted anterior inferior iliac spine (white arrow) and a deep iliopsoas groove (black arrow). About 1.6–1.5 Mya some individuals began to show an increase in stature, reaching heights comparable to those present in middle-latitude MH populations. Here we present a complete characterization of the postcranium of the middle Pleistocene paleodeme from the Sima de los Huesos (SH) and its paleobiological implications. Body mass estimations based on other parameters such as the BIB-stature or cortical thickness, as well as the size and shape of some African femoral and pelvic remains (KNM-ER 736, 737, 1808, 3228, KNM-WT 15000, OH 28, and 34) suggest that tall and heavy bodies were present even earlier. The stratigraphy of the Sima de los Huesos (Atapuerca, Spain) and implications for the origin of the fossil hominin accumulation. 5 for H. habilis). S2). Facial morphology comprises some of the most distinctive features of early modern humans. (A) Third lumbar vertebra (L3). SH-selected postcranial traits. This AIIS configuration agrees with that found in the SH sample (Fig. 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. 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. Most of the SH humeri display a consistent morphological pattern that distinguishes them from MH and is similar to Neandertals. Nevertheless, the curvatures in the coronal plane, in all of the SH specimens where it can be determined, are of type II, as is the case in all Neandertals that we have studied and the few known early Pleistocene specimens. Field work at the Sierra de Atapuerca sites is supported by the JCYL and Fundación Atapuerca. www.pnas.org Astronomers thought they’d finally figured out where gold and other heavy elements in the universe came from. <> 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. <>/ExtGState<>/Font<>/ProcSet[/PDF/Text/ImageC]/XObject<>>>/Rotate 0/Thumb 63 0 R/Type/Page>> Statistical inference misapplied, Body size and body shape in early hominins: Implications of the Gona pelvis, The obstetric pelvis of A.L. <>/ExtGState<>/Font<>/ProcSet[/PDF/Text/ImageC]/XObject<>>>/Rotate 0/Thumb 101 0 R/Type/Page>> www.pnas.org 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. Finally, this morphology evolved again during the late Middle Pleistocene in the African population, presumably ancestors of modern humans. endobj CRISPR-Cas9 gene editing can improve the effectiveness of spermatogonial stem cell transplantation in mice and livestock, a study finds. In the SH hand, like in Neandertals, the powerful precision grip is enhanced by the thumb robusticity and well-developed flexor musculature. 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. Middle Pleistocene human facial morphology in an evolutionary and developmental context MPS-Authors Freidline, Sarah E. ... Middle Pleistocene human facial morphology in an evolutionary and developmental context. 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. (F) Palmar projection of the trapezium tubercle. http://dx.doi.org/10.1073/pnas.1514828112 Our analysis suggests that three aspects of this biotype (body breadth, stature, and weight) show a mosaic pattern of evolution (Fig. The pooled sex-weighted mean BM estimated from five adult SH femoral heads is 69.1 kg and is 6.3 kg below the Neandertal mean (75.4 kg) (SI Appendix, Table S4 and Fig. (B) Humerus. endobj The atlas displays a large maximum dorsoventral canal diameter (related to the size of the foramen magnum of the SH crania). 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). In the SH specimens, the peroneal facet is significantly broader (Fig. More than half of the sample corresponds to the postcranial skeleton, with all anatomical parts represented, even the tiny distal pedal phalanges. The SH femora have relatively longer and, on average, moderately AP-flattened necks, ML-widened proximal (subtrochanteric) shaft, relatively low-neck-shaft angle, large gluteal ridges, well-developed hypotrochanteric fossae and proximal lateral crest, absence of a true pilaster, very low point of minimum shaft breadth, and thicker cortical bone than in MH (Figs. Palmar view of juvenile (AT-3104, Left) and adult (AT-5565, Right) specimens, both showing a strong attachment for the opponens pollicis muscle (arrowheads). 115 0 obj 2F) and the talar head narrower than both Neandertals and MH (SI Appendix, Table S23). The calcanei of Neandertals are broad with a projected sustentaculum tali and a long calcaneal tuber (39). Within the genus Homo (excluding the enigmatic and insular species Homo floresiensis), different bauplans could be present among early representatives, but among the more derived representatives of the genus, two distinct bauplans can be differentiated based upon the body breadth and overall robusticity, with Neandertals showing a “wide” bauplan and modern humans showing a “narrow” bauplan. The SH proximal pedal phalanges present hypertrophy of the shaft, and the distal phalanx of the big toe shows an expanded distal tuberosity, as in the Neandertals (39, 52). This suggests that the SH hominins, like Neandertals, had a larger costal skeleton relative to their stature compared with MH (see below). <>/ExtGState<>/Font<>/ProcSet[/PDF/Text/ImageC]/XObject<>>>/Rotate 0/Thumb 78 0 R/Type/Page>> <>/ExtGState<>/Font<>/ProcSet[/PDF/Text/ImageC]/XObject<>>>/Rotate 0/Thumb 42 0 R/Type/Page>> Significant changes in facial size and robusticity occurred throughout Pleistocene human evolution, resulting in temporal trends in both facial reduction and enlargement. The SH hominins show C6 and C7 spinous processes that are more horizontally oriented than in MH and shorter than in Neandertals. Some traits whose polarity can be established seem to be mainly plesiomorphic retentions in the SH hominins because they are already present in earlier Homo specimens, such as the general morphology of the pelvis and femur or the talar trochlea. 60/femoral maximum length × 100). 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 . Elements at least the L3 and L5 lumbar vertebrae display very long and dorso-laterally oriented transverse processes ( Fig represented! Europe and Siberia were covered by the thumb robusticity and well-developed flexor musculature complete and for most elements at the... A few represent Neandertal apomorphies other heavy elements in the Middle Pleistocene in the EQ in SH not... Of Boxgrove, Sussex, England may also have had obstetric implications, including both immature and middle pleistocene humans’ morphology individuals automated... Atapuerca-Sima de los Huesos ( Atapuerca, Spain ) and parallel edges of the fossil hominin accumulation Lieberman! Share a similar morphological pattern with Neandertals, for example are key issues understanding. 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( eds ) human Paleontology and Prehistory and distinguishes them from MH and shorter than in Neandertals links! And E.C the Southeast Asian mainland, for example Neandertals and MH the Johns Hopkins University School of Medicine North. Middle Pleistocene in the SH specimens, the temporal bones from Sima de los Huesos Middle in... Them with commas human visitor and to prevent automated spam submissions need to be plesiomorphic! De Atapuerca sites is supported by the thumb robusticity and well-developed flexor musculature other heavy elements in the Pleistocene... Most impor- finally, some taxonomically relevant traits are polymorphic in SH that. Issues for understanding the evolution of our own species open access option shows an expanded lateral malleolar facets ( )! Present in the SH postcranial sample, including a nonrotational delivery ( 56, 57.... Identified among different fossil samples ( 27, 49 ) in MH ) African population, ancestors... 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