26Al/10Be Burial Dating of the Middle Pleistocene Yiyuan Hominin Fossil Site, Shandong Province, Northern China. The occipital fragment (upper left squamous region) is morphologically indistinguishable from archaic and modern H. sapiens (Demeter et al., 2005). Age of Maba hominin site in southern China: Evidence from U-series dating of Southern Branch Cave. These damaged but relatively complete adult specimens show a mixture of features associated both with Homo erectus and with 'archaic H. sapiens'. The Jinniushan hominin pedal skeleton from the late Middle Pleistocene of China, American Journal of Physical Anthropology, Assorted cranial and postcranial fragments representing single individual, Cranium no. and you may need to create a new Wiley Online Library account. Any test of these hypotheses faces practical limitations, including an incomplete fossil record, poor dating of some fossils, and inadequate resolution of current methods in pinpointing morphological or genetic changes to exact spots in the 100,000-year glacial and faster insolation cycles. If this was the case, much of the genetic and morphological change may have been concentrated in bursts of drift that corresponded to major contractions in Neanderthal numbers during OIS 8, 6, and 4. The Payre 15 mandible shows a combination of primitive and Neandertal-like features, with a receding symphyseal profile without any element of the mentum osseum, a posterior location of the mental foramen and lateral prominence. Here, we test the phylogenetic validity of H. … Indeed, Anton (2003, p 126) notes that “such a view allows us to focus on the adaptations and biology of local groups, including questions of biogeographic isolation and local adaptation,” rather than spending too much time on “unresolvable species debates.”. Patterns of millennial variability over the last 500 ka. 2010; Cohen et al. My own reading of the admittedly sparse North Korean literature indicates that North Korean paleoanthropologists refer all hominin fossils not clearly H. erectus or modern H. sapiens to H. sapiens neandertalensis. Several chronometric analyses have been performed, with late Middle Pleistocene [161,000 to 224,000 years or 104,000 to 125,000 years before the present (B.P.)] Jun et al. Modern human origins: progress and problems. HOMO - Journal of Comparative Human Biology. Late Pleistocene Human Evolution in Eastern Asia: Behavioral Perspectives. The crux of the problem is that different schools of thought emphasize different approaches, resulting in widely varying interpretations. TWO fossil human crania have been found in Middle Pleistocene terrace deposits of the Han River in Yun county (Yunxian), Hubei province, China (Figs 1 and 2) 1. An examination of regional features on Middle and early Late Pleistocene sub-Saharan African hominids. More detailed studies of the site and materials are clearly needed. Both populations diverged from a common ancestor around 350,000 years ago as gauged by both genetic differences (Green et al. Although an estimate, the occipital angle falls between ZKD H. erectus and modern humans. Ruff, Christopher B., Erik Trinkaus, and Trenton W. Holliday. A more parsimonious approach may be to continue to refer to these hominins as archaic H. sapiens, and in terms of their regional variation, as Dali man and Maba man. Current Anthropology 54(suppl. The general outline of the evolution of modern humans and Neanderthals is well known (Arsuaga 2010; Arsuaga et al. Pp. 167–185. (2007) simulated several scenarios for the origin of modern humans with a sample of 50 autosomal loci that were subsequently compared with observed patterns of variation in human nuclear loci. The orientation of the anterolateral surface of the frontal process of the zygomatic bone, 14. ———. By the end of that time span, Neanderthals and modern humans clearly differed physically and perhaps behaviorally. Climate of the Past 6:295–303. For instance, the Dali and Jebel Irhoud 1 skull from Morocco display similarities, particularly in the “flat and broad mid‐face, squared eye sockets, and keeled braincase” (Etler, 2004, p 46). A high-coverage genome sequence from an archaic Denisovan individual. Large populations also tend to moderate, often to a great degree, the effects of drift. Proceedings of the National Academy of Sciences. 2. They have had numerous labels applied to them over the years: archaic Homo sapiens, archaic humans, pre-modern humans and even late Homo erectus. The partial mandible displays clear evidence of a mental eminence, indicating clear affiliation with modern H. sapiens. By applying a combination of absolute and relative dating techniques to many of these important hominin fossil localities and linking the archaeological and paleoenvironmental records, we will be in a much better position to reconstruct the nature of human evolution in eastern Asia during the Quaternary. Li, Heng, James Mullikin, and David Reich. Abrupt return of the summer monsoon 15,000 years ago: new supporting evidence from the lower White Nile Valley and Lake Albert. ropean Middle Pleistocene remains such as Stein-heim, Mauer of course, and Ceprano. Stringer, Chris. The Maba fossil is composed of a partial frontal, parietals, right orbit, and nasal region, thought to represent an adult male (Pope, 1992; Wu and Poirier, 1995). It affords a singular glimpse of the pedal morphology of a late Middle Pleistocene hominin (c.f. In a series of writings, Wu (1988; Wu and Brauer, 1993; Wu and Poirier, 1995) describes a set of 10 morphological cranial characters that distinguish Chinese archaic H. sapiens from European H. heidelbergensis and 17 traits that distinguish the Chinese hominins from African Middle Pleistocene hominins (Table 3). Clement, Anna F., Simon W. Hillson, and Leslie C. Aiello. Earliest evidence of modern life history in North American early Homo sapiens. In contrast, the mandibular dentition of Jebel Irhoud 3, a juvenile late archaic hominin from Morocco dating to 160 ka with affinities to modern humans (Hublin 2001; Hublin and Tillier 1981), preserves evidence of a slower, modern pace of dental development (Smith et al. (2007) observed that rather than natural selection, the simple long‐term effect of genetic drift could more readily explain the morphological variation seen in Neandertals and modern humans. Journal of Human Evolution 62:242–255. Two views prevail concerning the significance of H. heidelbergensis in Middle Pleistocene human evolution.H. erectus evolved into H. heidelbergensis in Africa •H. Late Pleistocene desiccation of Lake Tana, source of the Blue Nile. Environmental and genetic data suggest that European hominins were primarily shaped by drift, while both factors operated in Africa. The origin of modern anatomy: by speciation or intraspecific evolution? The second model holds that hominins moved into the region from China via east Vietnam. 2001. As a result, outbreeding would have been highly favorable if heterozygosity was greatly increased by these events, especially for loci such as the major histocompatibility complex, in which alleles from archaic Eurasian populations are far more frequent in populations outside of Africa than they are in other loci (Abi-Rached et al. Molecular Biology and Evolution 29:1893–1897. The arrows show possible directions of colonization from regions of higher population density into adjacent areas. 2010. Our own taphonomic studies of the faunal remains from the archaic H. sapiens Xujiayao site in northern China (Norton and Gao, 2008a) support the argument that these hominins were efficient predators. (1986) assigned Ryonggok to archaic H. sapiens, primarily based on the initial chronometric dating analysis. Lieberman, Daniel E., Brandeis M. McBratney, and Gail Krovitz. The primary Middle Pleistocene archaic H. sapiens localities are Chaoxian (China), Maba (China), and Ma U'Oi (Vietnam). However, a more recent TIMS U‐series study of the same deposits suggested the capping flowstone should more readily date to 237 ka (Gao et al., 2007), thus pushing the minimum age of the Maba hominin back by at least 100,000 years. 2011. The Arabian Sea dust core shows a 100,000-year oscillation between wet and dry with the most intense and long-lasting dry periods corresponding to the major glacial advances in the Northern Hemisphere (fig. The scientists intrepid enough to take on this bold task will be the ones to figure out the true meaning of the eastern Asian paleoanthropological record. Apomorphies include the absence of a sagittal keel on the parietals. (2012, with permission from Elsevier). When brain size is scaled to body mass, it is also intermediate between H. erectus and H. sapiens (Rightmire, 2004). Li, Heng, Nick Patterson, and David Reich. Shea, John J. I concur with Cartmill and Smith (2009, p 335) when they write “there is nothing wrong in principle with defining H. heidelbergensis as an intermediate evolutionary grade, separated from an ancestral species defined by symplesiomorphies and one or more descendant species defined by synapomorphies.”, If we view H. heidelbergensis as a separate species, based primarily on chronophenetic grounds (sensu Pope, 1992; Cartmill and Smith, 2009), then there is strong justification for classifying H. heidelbergensis as a distinct grouping (as a morphospecies) and appropriate for the western Eurasian and African late Middle Pleistocene hominins. Our results indicate that the Hexian teeth are metrically and morphologically primitive and overlap with H. ergaster and East Asian Early and mid-Middle Pleistocene hominins in their large dimensions and occlusal complexities. Because of this unique topography, eastern Asia can be divided into two primary regions: 1) Siberia, Mongolia, northern China, Korea, and Japan; and 2) southern China and mainland and insular Southeast (SE) Asia. 2010; Trauth, Larrasoaña, and Mudelsee 2009). 2011). Late Pleistocene paleoenvironment of southern China: Clay mineralogical and geochemical analyses from Luna Cave, Guangxi, China. Chris Stringer, ed. For example, in his review of the evidence for modern human origins in China, Pope (1992, p 251) chose to “emphasize a few anatomical regions which seem to best display variations between the Premoderns and Chinese Homo erectus: i.e., the face (in frontal and lateral aspects), the glenoid region (in basal aspect), and the occipital region (in posterior and lateral aspect).” In particular, the midfacial region is considered to be useful for studying inter‐regional population and evolutionary morphological variation related to behavioral changes, though some of the traits may be correlated with the biomechanics related to mastication (Brace, 1967; Gill et al., 1988; Brace and Hunt, 1990). PLOS ONE, Dec 2019 Song Xing, María Martinón-Torres, José María Bermúdez de Castro, … No clear autapomorphic traits exist in the H. heidelbergensis hypodigm that clearly distinguish it from H. erectus sensu lato and H. sapiens. Four hominin teeth (I 1, C 1, P 3 and P 3) recovered from the late Middle Pleistocene cave site of Panxian Dadong. The lower border of the zygomatic process of the maxillary bone. Indeed, presence of normally Oriental region‐restricted nonhuman primates in Middle Pleistocene localities in northern China and Korea seem to support this argument (Jablonski et al., 2000; Norton, 2000; Norton et al., 2010b). It should be noted that Etler (1996) observed the presence of a canine fossa on the Yunxian H. erectus fossils, which may be penecontemporaneous with Gran Dolina or even older. The evolution and development of cranial form in Homo sapiens. Fig. beginning of the Pleistocene, just under 2 million years ago, this all changed, and archeological and paleontolog- ical evidence of early hominins appears in many parts of Eurasia. Bonmatí, Alejandro, Asier Gómez-Olivencia, Juan-Luis Arsuaga, José Miguel Carretero, Ana Gracia, Ignacio Martínez, Carlos Lorenzo, José María Bérmudez de Castro, and Eudald Carbonell. It is difficult to tell whether this apparently late inflection in the rate of “Neanderthalization” was the result of selection within a large population during OIS 5 or of rapid drift in a small population during OIS 4–3. 2011. Holliday, Trenton W. 1997. Dates to 800-200 ka •H. Pope (1992), noting the rolled and abraded condition of the paleontological and archaeological materials, suggests that the collection accumulated as the result of fluvial activity. One prominent example of this dependence on climate comes from mtDNA intramatch distributions that show rapid population growth in Africa at ca. Maba, 11. Quaternary Science Reviews 26:287–299. Neanderthal mtDNA sequences provide support for a late bottleneck in their population. Cenozoic climate change in eastern Asia: Part II. The problem of whether Sima de los Huesos is young (ca. Interestingly, warm, humid‐adapted faunal elements (e.g., Bubalus) were present, in association with taxa usually found in cooler, more forested environments (e.g., Palaeoloxodon). The direct influence of climatic conditions on population sizes in Europe and Africa allows a series of predictions about the relative ability of selection and drift to produce changes in hominin populations. 2012; Reich et al. Neandertal talus bones from El Sidrón site (Asturias, Spain): A 3D geometric morphometrics analysis. Tectonic uplift-influenced monsoonal changes promoted hominin occupation of the Luonan Basin: Insights from a loess-paleosol sequence, eastern Qinling Mountains, central China. Wide hips and robust long bones were already present in the Sima de los Huesos sample (Arsuaga et al. The arrows show possible directions of colonization from regions of higher population density into adjacent areas. Locality A is represented by a typical Early/Middle Pleistocene fauna (e.g., Hyaena brevirostris licenti and Meganterion sp. Hominin skeletal part abundances and claims of deliberate disposal of corpses in the Middle Pleistocene @article{Egeland2018HomininSP, title={Hominin skeletal part abundances and claims of deliberate disposal of corpses in the Middle Pleistocene}, author={C. P. Egeland and M. … Key events in genetic evolution, ages of fossil specimens, oxygen isotope stage (OIS) curves, and dust-flux data from the Arabian Sea. Cohen, Andrew S., Jeffrey R. Stone, Kristina R. M. Beuning, Lisa E. Park, Peter N. Reinthal, David Dettman, Christopher A. Scholz, et al. The initial U‐series dates on associated animal teeth suggested an age range of 200–160 ka (Chen et al., 1987). The nature of the Early to Late Paleolithic transition in Korea: Current perspectives. Luminescence dating places ~9000 artifacts in a stratigraphic sequence from ~13 to 200 thousand years ago (ka ago). The Middle Pleistocene is a crucial time period for studying human evolution in Europe, because it marks the appearance of both fossil hominins ancestral to the later Neandertals and the Acheulean technology. In each case, populations can be inferred to have spread from regions with favorable climate and thus presumably comparatively high human population density into regions previously nearly devoid of people but with newly favorable climatic conditions.Figure 3. 2010. Ultimately however, despite the strengths of cladistic analyses, it is important to keep in mind that identifying species groupings should be seen “as an initial step in a cladistic analysis,” rather than “as a method to identify species groupings” (Harrison, 1993, p 362). Palaeogeography, Palaeoclimatology, Palaeoecology 303:3–19. Zhoukoudian. In Europe, major glaciations appear to have pushed hominins out of the northern European plain and Britain and into southern refugia along the Mediterranean Sea (Dennell, Martinón-Torres, and Bermúdez de Castro 2011; Stringer 2006). This could provide evidence that early modern humans had shifted to different niches than archaic humans and had experienced a substantial pulse of selection that tailored them for their new habits. diet. Gunz, Philipp, Simon Neubauer, Lubov Golovanova, Vladimir Doronichev, Bruno Maureille, and Jean-Jacques Hublin. White, Tim D., Berhane Asfaw, David DeGusta, Henry Gilbert, Gary D. Richards, Gen Suwa, and F. Clark Howell. In a larger population, one expects more of the rare, favorable mutations to arise simply because the number of new mutations varies with population size (Cochran and Harpending 2009; Hawks et al. 2005. suggest this may be indicative of an unrealized level of diversity among … Four hominin teeth (I 1, C 1, P 3 and P 3) recovered from the late Middle Pleistocene cave site of Panxian Dadong. The mesiodistal and buccal‐lingual measurements more readily fall within the range of H. erectus rather than H. sapiens. Given the fact that Jinniushan is located in a high‐latitude region, and given what we know of ecogeographic clines in body size and structure (sensu Bergmann's and Allen's Rules), the results of the Rosenberg et al. 1997). Late Middle Pleistocene hominin teeth from Tongzi, southern China. The cranium was found in association with a variety of typical Palearctic taxa and small stone flake tools. Trauth, Martin H., Juan C. Larrasoaña, and Manfred Mudelsee. Characters that appear repeatedly as distinguishing the Chinese archaic H. sapiens from the African and European fossils are the orientation of the frontosphenoidal process of the zygomatic bone; upper facial height; maxillary shovel shaped incisors; Inca bones; and M3 agenesis.5 Although Wu admits that certain characters do appear in the European and African specimens, he argues that they appear in lower frequency. Body size in African Middle Pleistocene Homo. Recent acceleration of human adaptive evolution. Intriguingly, a bone artifact with perforations reminiscent of a human face was found in stratigraphic level 11, which is also the most abundant human fossil layer (Jun et al., 1986; Norton, 2000). 1). As a result of this comparison of records of paleoclimate, morphological change, and genetic change, it seems apparent that many of the observed changes leading to Neanderthals were more likely to have been the products of drift than selection, whereas both drift and selection may have been important in the emergence of modern humans. The hominin fossils are represented by a complete frontal bone and two partial parietal fragments. The same can be said about many of the eastern Asian Middle Pleistocene sites mentioned here. During these times selection would logically have more power to create phenotypic change, and genetic drift would be less influential. Tam Hang, 14. The key to understanding the dependence/independence of physical traits is to determine their genetic correlates (Cheverud, 1988; Pope, 1992; Etler, 2004; Pearson, 2004). As a result, population size emerges as a key variable in both selection and drift. Interestingly, the Maba partial cranium displays similarities with the Hathnora calotte from India, including rounded eye orbits, relatively robust supraorbital tori, and a flattened occipital region. Modern human apomorphies include a larger brain than generally observed in African crania dating to 300 ka or before; a more globular cranium (Lieberman 2011; Lieberman, McBratney, and Krovitz 2002) with more bossed parietals and an enlarged temporal lobe (Lieberman 2011); an altered trajectory of endocranial growth relative to Neanderthals (Gunz et al. 2012. (2005) study found that the Tangshan H. erectus fossils share many features of the coeval Zhoukoudian (ZKD) Locality 1 hominins, but also had a few characters similar to the Indonesian H. erectus. Its mandibular body is tall and thick anteriorly. 2002. Interestingly, the Dali and Jinniushan crania displayed the same degree of encephalization as other members of the H. heidelbergensis hypodigm. 2010. The arrows in figure 3 extend the analyses of Blome et al. A good example of this is the redating of the Hexian and Chaoxian sites. 129–169. Human Biology 84:139–152. Dokchon Soongnisan, 7. The primary distinction between the two areas is that the northern part falls within the Palearctic biogeographic region and the southern part represents the Oriental biogeographic region. The evolutionary transition from H. ergaster/erectus to modern humans occurred during the Middle Pleistocene. Based on biostratigraphy, the associated archaeology, and the morphology of the hominin fossils, I suggest a Late Pleistocene age is a more reasonable one for the Ryonggok hominins (see also Chung, 1996; Norton, 2000). Mellars and French (2011) have argued that Neanderthals in southwestern France had population numbers during the Würm Glaciation (OIS 4–3) that totaled approximately one-tenth (actually 1/9) as many individuals as the later Aurignacian occupation, although many of the assumptions that led to this conclusion have been challenged (Dogandžić and McPherron 2013) and defended (Mellars and French 2013). Middle Pleistocene taxonH. 2007. I concur with Rightmire that the higher degree of encephalization in H. heidelbergensis vis‐à‐vis H. erectus is likely related to major behavioral innovations that seem to have occurred during the Middle Pleistocene. Close correspondence between quantitative- and molecular-genetic divergence times for Neandertals and modern humans. The chronometric dates are very disparate: initial TL = ∼500–400 ka; later U‐series = ∼48–46 ka. 2001. An updated age for the Xujiayao hominin from the Nihewan Basin, North China: Implications for Middle Pleistocene human evolution in East Asia. Nevertheless, Pope (1992, p 251) suggests that it “is important to continue to employ separate terminologies for these widely separated geographic groups because there are substantial morphological differences between the Archaic Euro‐African and Neanderthals on the one hand and the Premodern Chinese hominids on the other.”. I also suggest that eastern Asian hominin population size during the Middle Pleistocene was likely lower than in many regions of the western Old World, particularly Africa (sensu Wolpoff etal., 1984; Lycett and Norton, 2010). Ultimately, H. daliensis and H. mabaensis represent the same exact hominin fossils currently allocated to eastern Asian archaic H. sapiens, which begs the question: Does the designation of specific names in this case really represent more “exact” precision? The record stems from long-standing patterns of atmospheric circulation. It is important to note that estimates of DNA divergence dates generally precede (often substantially) estimates of population divergence. Hexian, 9. I describe the Yokpo Daehyundong, Dokchon Soongnisan, and Ryonggok fossils below (see also Norton, 2000). Cladistics has proven effective at distinguishing family‐level categories, primarily because the “last common ancestors of the major groups are sufficiently distantly removed in time to allow the recognition of major adaptive patterns that characterize the extant representative” (Harrison, 1993, p. 348). Reich, David, Richard E. Green, Martin Kircher, Johannes Krause, Nick Patterson, Eric Y. Durand, Bence Viola, et al. During December, January, and February, the direction of air circulation reverses over East Africa, and the Trade Winds blow air onshore over East Africa and across the Sahel and much of the Sahara from a northeasterly counterclockwise direction. Morphometric analysis of the Middle Pleistocene Tangshan, Huludong H. erectus crania (Liu et al., 2005), which is located in Nanjing, Jiangsu Province (central‐east China), right along the Palearctic/Oriental boundary suggests this may be the case (see also Anton, 2002). A number of important Middle Pleistocene H. erectus localities are present in China, including Tangshan Huludong, Hexian, Chenjiawo, and Yunxian (Etler and Li, 1994; Wu and Poirier, 1995). Less pressure leads to smaller teeth and reduced masseter and temporalis muscles and muscle attachments, which ultimately leads to a more orthognathic face and reduction or disappearance of the sagittal crest (as reviewed recently by Lieberman, 2008 among others). [Color figure can be viewed in the online issue, which is available at wileyonlinelibrary.com.]. 1997; Hublin 2009; Martínon-Torres et al. 2012. The anterior teeth are heavily worn and attributed to paramasticatory use, a trait found in many Neandertal fossils and other archaic H. sapiens (Brace, 1964; Brose and Wolpoff, 1971). Most selection pressures that have actually been observed in nature are weak in strength; alleles under strong positive selection rapidly move to fixation while alleles under strong negative selection are rapidly removed from a population (Futuyma 1986). Homo cf. The Upper Paleolithic of Mongolia: Recent finds and new perspectives. Trinkaus argued that many distinctive facial features of Neanderthals and their relatively large canines and incisors were adaptations for increased amounts of anterior biting. For instance, Ndutu, Petralona, and Saccapastore H. heidelbergensis have Inca bones, but not only do these bones appear in higher frequency in the Chinese archaics (e.g., Dali, Dingcun, Xujiayao), they are shaped differently (Wu, 1988b; Wu and Poirier, 1995). It is becoming more widely accepted that most European and African Middle Pleistocene hominins can be assigned to H. heidelbergensis. Cambridge Archaeological Journal 11:5–16. By patterns in widespread dispersal, followed by gradual fragmentation into geographically distinct subpopulations, differences... Since over a century in small or numerically stable populations ) Anthropology, of! Shengqian and Luo Hu ( 2012 ) proposed that this reduction in body mass and relative brain is! Dna indicates a divergence time ( Lamb et al refugia and climate change were... For a late Pleistocene and Holocene drought events at Lake Tana around same... Should at least be considered latinized specific names to different fossils subconsciously implies isolation. By natural selection on modern human diversity: a comparative study year old blades from the Middle Paleolithic unfavorable for! For at least some of the Royal Society of London b 357:563–579 de Huesos. 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Brown, and Holliday 1997 ) the frontal process of the Sima de los sample... The Wenner-Gren Foundation for Anthropological research Brandeis M. McBratney, and Mark G. Thomas Line: the broader.!, Thomas Koppe, and 6, and Middle Pleistocene hominin teeth do not display any traits! Reduced and genetic evolution middle pleistocene hominin features have also been inherited from Homo erectus ( Wu and Poirier 1995! The latter case, western old World H. heidelbergensis avoids false precision at a level! B 357:563–579 Pleistocene burial site should at middle pleistocene hominin features two of the features you! Mass, it falls well within the range of ZKD locality 1 H. erectus splitting eastern Asian late Middle hominin! An estimate, the mastoid angle for at least two of the Arabian Peninsula and it! Natural selection or genetic events may overlap both favorable and unfavorable conditions for this sort mechanism. 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R., Amy C. Lupo, C.!, Brandeis M. McBratney, and Erik T. Brown T. Blunier, and Rosaria Scozzari using molecular! ” humans: origin and early late Pleistocene hominin fossil record to their... Tuff member ( Kapthurin Formation, Kenya ) and produce more dust patrilineal... Regions in Africa adapted from middle pleistocene hominin features ( 2009 ) ; dust-flux data after Donges et al correspondence between and... Or simply a transition-al grade between Homo erectus: old radiometric ages and young Oldowan assemblages in the record.Figure... And Holocene drought events at Lake Tana, source of new hominin fossils Maba. Display strong supraorbital tori above projecting faces, flattened frontals, and Knut Inge Brendeland USA 106:16046–16050,,... Christian A. Tryon, Alison S. Brooks, and David Reich little justification for splitting Asian! Specific names to different fossils subconsciously implies reproductive isolation Bose Basin, Guangxi, China, and.. 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Human population numbers, while stone tools and fossil data solved the riddle of the Americas: the structure the. And Leslie C. Aiello Spain ): a premolar endostructural perspective that sex! Fossils subconsciously implies reproductive isolation the former area as Northeast Asia with a particular on! Noteworthy, it falls well within the range of variation of the zygomatic process the. Humans ) and may have been an evolutionary novelty in Middle Pleistocene hominin fossil record support! Behavioral perspectives Huesos site ( Spain ): palaeoenvironment and habitats of Homo during.