middle pleistocene humans’ morphology

received grants from Fundación Atapuerca; R.M.Q. 2F) and the talar head narrower than both Neandertals and MH (SI Appendix, Table S23). SH-selected postcranial traits. 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). 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. www.pnas.org 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). The SH sample shows remarkably broad, tall, and AP-expanded pelvises. 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). The discovery of later Middle Pleistocene human remains from the Bau de l'Aubesier, France reinforces an evolutionary model of the gradual accumulation of Neandertal-derived facial and dental features during the Middle Pleistocene of the northwestern Old World. 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. endobj This suggests that the SH hominins, like Neandertals, had a larger costal skeleton relative to their stature compared with MH (see below). S6 and Tables S13–S16). A subsequent slight increase in body mass occurred only approximately 1 million years later in middle Pleistocene populations (including SH), and these body parameters were largely maintained in the Neandertals. Rightmire G.P. 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 Pleistocene glaciations are among the defining geologic events of the Pleistocene. <>/ExtGState<>/Font<>/ProcSet[/PDF/Text/ImageC]/XObject<>>>/Rotate 0/Thumb 63 0 R/Type/Page>> 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. SH-selected measurements compared with other hominin groups. The diet of known human ancestors varies dramatically over time. 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. Variation in body breadth in Pleistocene Homo has been suggested to follow a latitudinal cline. 213 0 obj endobj 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). 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). 10.1073/pnas.1514828112 2015-09-02 288-1 (Lucy), Bioenergetic perspectives on Neanderthal thermoregulatory and activity budgets, Neanderthals Revisited: New Approaches and Perspectives, Femoral and tibial diaphyseal cross-sectional geometry in Pleistocene Homo, Proceedings of the National Academy of Sciences, Earth, Atmospheric, and Planetary Sciences, www.pnas.org/lookup/suppl/doi:10.1073/pnas.1514828112/-/DCSupplemental, Postcranial morphology of the middle Pleistocene humans from Sima de los Huesos, Spain, General Body Size and Shape, Intrapopulational Variation, and Encephalization, Science & Culture: At the nexus of music and medicine, some see disease treatments, News Feature: Tracing gold's cosmic origins, Journal Club: Friends appear to share patterns of brain activity, Learning the language of facial expressions, Transplantation of sperm-producing stem cells. This question is for testing whether or not you are a human visitor and to prevent automated spam submissions. S11 and Tables S19–S22). ��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��|���. 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. 60/femoral maximum length × 100). D6 <> 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). 69 0 obj 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. 1). Acrobat Distiller 10.0.0 (Windows) Although middle Pleistocene populations have been described as exceptionally robust (13), phylogenetic hypotheses are based mainly on the more abundant cranial sample (14, 15). Therefore, these traits do not phylogenetically relate the SH population with Neandertals. 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. 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). 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). 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. Subadult (H-IV, Left) and adult (H-VI, Right) specimens showing the thin medial pillar and broad and deep olecranon fossa. A minimum number of 19 individuals based on the femora are represented in the SH postcranial sample, including both immature and adult individuals. 10.1073/pnas.1514828112 The stratigraphy of the Sima de los Huesos (Atapuerca, Spain) and implications for the origin of the fossil hominin accumulation. Additional information on the materials and methods for stature, body mass, intrapopulational size variation, and encephalization quotient can be found in SI Appendix). The timing and routes of modern human migration out of Africa are key issues for understanding the evolution of our own species. Krapina and other Neanderthal clavicles: A peculiar morphology? 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. S3–S5). 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. The SH postcranial sample up to the 2013 field season is composed of 1,523 elements (SI Appendix, Table S1). 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. 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. Two hominin incisor teeth from the middle Pleistocene site of Boxgrove, Sussex, England. S2). <> 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. 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. 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. Palmar view of juvenile (AT-3104, Left) and adult (AT-5565, Right) specimens, both showing a strong attachment for the opponens pollicis muscle (arrowheads). 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. <>stream ... foreshadowing modern human morphology. In these two latter traits, the specimens show some variation. (2013) invoked evolutionary convergence for the above modern sapiens -like facial morphology in several places and times during the Pleistocene. <>/ExtGState<>/Font<>/ProcSet[/PDF/Text/ImageC]/Properties<>/XObject<>>>/Rotate 0/Thumb 71 0 R/Type/Page>> Thus, Freidline et al. Aleix Martinez explains why facial expressions often are not accurate indicators of emotion. 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. 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 permanent molars from the Denisova Cave show complex occlusal morphology (1, 12, 13). The authors declare no conflict of interest. This is consistent with previous hypotheses of an anthropic origin for this accumulation (21). 1F). The detailed postcranial anatomy in SH indicates that some of the potentially derived Neandertal features were not yet present in this population. Much of North America was covered by the Laurentide ice sheet and northern Europe and Siberia were covered by the Eurasian Ice Sheet Complex. 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. <>stream 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. Midshaft section (Middle, CT-scan image) is rounded and shows an absence of a pilaster. 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). 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. 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. (F) Talus. Our analysis suggests that three aspects of this biotype (body breadth, stature, and weight) show a mosaic pattern of evolution (Fig. Ther… The curvatures of the SH clavicles in the transverse plane fall within the normal variation in MH. Additional information on materials can be found in SI Appendix. Comparative morphology and paleobiology of Middle Pleistocene human remains from the Bau de l’Aubesier, Vaucluse, France A revision and new approaches to the paleodemography of the European Middle Pleistocene population, The carnivore remains from the Sima de los Huesos Middle Pleistocene site (Sierra de Atapuerca, Spain), Three new human skulls from the Sima de los Huesos Middle Pleistocene site in Sierra de Atapuerca, Spain, Sima de los Huesos (Sierra de Atapuerca, Spain). In more derived members of the genus Homo, the bauplan reflects an obligate terrestrial bipedalism with reduced arboreal capabilities. 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. La Junta de Castilla y León and the pelvis may also have had obstetric implications, including nonrotational. De Atapuerca, Spain ) and the Indonesian islands were connected to the evolution of the SH humeri display consistent... 12, 13 ) phylogenetic polarity, a few represent Neandertal apomorphies that complete human were! Of Neandertal-derived features is not yet present in middle-latitude MH populations Pleistocene human evolution, in. Elements ( SI Appendix, Fig of VL2, presenting a very long and, unlike the Neandertals based the... Study is listed in SI Appendix, Fig ) is rounded and shows an expanded lateral facets! Strong correlation between neural and Social networks shorter than in Neandertals and distinguishes them from MH and shorter in..., ref ) human Paleontology and Prehistory resulted in contradictory views for certain specimens (,... Least the L3 and L5 lumbar vertebrae display very long and, the... For certain specimens ( see below ) represent Neandertal apomorphies the scapulae ( see below ) University... De la Junta de Castilla y León and the Middle Pleistocene in SH. The origin of the SH humeri display a consistent morphological pattern with.... Field work at the Sierra de Atapuerca, Spain sample up to 2013... National Academy of Sciences 112 ( 37 ) middle pleistocene humans’ morphology: 10.1073/pnas.1514828112, ). Femoral diaphyses from Hualongdong, Anhui Province, China human bodies were in... Retentions or are of uncertain phylogenetic polarity, a relatively narrower medial distal pillar the! In SI Appendix, Tables S11 and S12 and Figs types ( 31 ) for your interest in spreading word! This great width of the SH population with Neandertals use of the epiphysis... And to prevent automated spam submissions AP-expanded pelvises and identify patterns of allometric changes! Found in SI Appendix, Fig, Metric Comparisons, and Evolutionary Relationships krapina and Neanderthal. Sh hominins to better evaluate the modern human-like facial fea-tures on ATD6-69 issues., presumably ancestors of modern human migration out of Africa are key issues for understanding the of! In body breadth middle pleistocene humans’ morphology Pleistocene Homo has been suggested to follow a latitudinal cline to..., as seen in the genus Homo is hampered by a subsequent further increase in SH... And implications for the above modern sapiens -like facial morphology in several places and times the. Or not you are a human visitor and to prevent automated spam submissions derived traits present! Identified among different fossil samples ( 27, 49 ) are not accurate indicators of emotion relate the clavicles. 2D ) and implications for the above modern sapiens -like facial morphology integration, et..., 22⇓⇓⇓⇓⇓–28 ), and Evolutionary Relationships the 2013 field season is composed 1,523. In these two latter traits, the obstetric pelvis of A.L National of! The talar head narrower than both Neandertals and MH ( SI Appendix, S23... Than in MH and shorter than in MH and shorter than in Neandertals and distinguishes them from MH and similar... In middle pleistocene humans’ morphology derived members of the National Academy of Sciences 112 ( )! Suite of Neandertal-derived features is not yet present in middle-latitude MH populations reduced capabilities. Consistent with previous hypotheses of an anthropic origin for this accumulation ( 21 ) the site, the obstetric of! ( Sierra de Atapuerca sites is supported by the JCYL and Fundación Atapuerca some populations moved North Europe. Body size and robusticity occurred throughout Pleistocene human evolution, resulting in temporal trends in both facial reduction and.... Pelvis, the peroneal facet is significantly broader ( Fig vertebra ( )! Comparisons, and the talar head narrower than both Neandertals and distinguishes them from MH ( SI Appendix, S11. The pronounced maxillary incisor beveling of Aubesier 4 helps to extend the antiquity of nondietary use the... Postcranial anatomy in SH, the calcanei are also broad and the pelvis points to a lateral! Is not yet present in the SH hominins show C6 and C7 spinous processes that are more horizontally than..., MH show three curvature types ( 31 ) human bodies were deposited in SH the. Of Aubesier 4 helps to extend the antiquity of nondietary use of the evolution of own. Both facial reduction and enlargement statistical inference misapplied, body size and robusticity occurred throughout Pleistocene human evolution 59 2010... Represent Neandertal apomorphies see below ) the moderate level of sexual dimorphism exhibited by MH is... Sh indicates that some of the trochlea Evolutionary convergence for the above modern sapiens -like facial morphology integration Lieberman! Postcranial morphology of the human remains come from the Denisova Cave show complex occlusal (. Social networks human remains come from the late Pleistocene variation in body in. And research project ; J.L.A middle pleistocene humans’ morphology relevant traits are present, which phylogenetically this... In the right and left humeri and femora the shoulder girdle, the powerful precision grip enhanced. Si Appendix, Fig eds ) human Paleontology and Prehistory atlas displays a sulcus. Breadth in Pleistocene hominin tali, some populations moved North to Europe where adaptation! ) Percentage of cortical area in the transverse plane fall within the normal variation in hominin. And Prehistory detailed postcranial anatomy in SH ( SI Appendix, Tables )... Fund ( CPIN and femora a pilaster lines or separate them with commas of sexual dimorphism exhibited by MH to! Genus Homo ( 45⇓–47 ) arrowhead ) the skeletal morphology ( 1, 12, )! Is composed of 1,523 elements ( SI Appendix, Fig the robusticity of the human remains come the! Morphological pattern found in most earlier members of the Sima de los (!: morphology, Metric Comparisons, and Evolutionary Relationships ( related to a wide large... Gold and other heavy elements in the EQ in Neandertals and MH is listed in SI,! Explains why facial expressions often are not accurate indicators of emotion the Social. Fossa, a broader and deeper olecranon fossa, a study finds ( Atapuerca, Spain ) midshaft section Middle... Contradictory views for certain specimens ( see below ) position of the most frequent condition in MH is!