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New fossils from Jebel Irhoud, Morocco and the pan-African origin of Homo sapiens | Nature
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The exact place and time that our species emerged remains obscure because the fossil record is limited and the chronological age of many key specimens remains uncertain. Previous fossil evidence has placed the emergence of modern human biology in eastern Africa around 200,000 years ago. In this issue of Nature, Jean-Jaques Hublin and colleagues report new human fossils from Jebel Irhoud, Morocco; their work is accompanied by a separate report on the dating of the fossils by Shannon McPherron and colleagues. Together they report remains dating back 300,000–350,000 years. They identify numerous features, including a facial, mandibular and dental morphology, that align the material with early or recent modern humans. They also identified more primitive neurocranial and endocranial morphology. Collectively, the researchers believe that this mosaic of features displayed by the Jebel Irhoud hominins assigns them to the earliest evolutionary phase of Homo sapiens. 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All rights reserved."/> <meta name="dc.rights" content="2017 Macmillan Publishers Limited, part of Springer Nature. All rights reserved."/> <meta name="dc.rightsAgent" content="journalpermissions@springernature.com"/> <meta name="dc.description" content="New human fossils from Jebel Irhoud (Morocco) document the earliest evolutionary stage of Homo sapiens and display modern conditions of the face and mandible combined with more primative features of the neurocranium. The exact place and time that our species emerged remains obscure because the fossil record is limited and the chronological age of many key specimens remains uncertain. Previous fossil evidence has placed the emergence of modern human biology in eastern Africa around 200,000 years ago. In this issue of Nature, Jean-Jaques Hublin and colleagues report new human fossils from Jebel Irhoud, Morocco; their work is accompanied by a separate report on the dating of the fossils by Shannon McPherron and colleagues. Together they report remains dating back 300,000–350,000 years. They identify numerous features, including a facial, mandibular and dental morphology, that align the material with early or recent modern humans. They also identified more primitive neurocranial and endocranial morphology. Collectively, the researchers believe that this mosaic of features displayed by the Jebel Irhoud hominins assigns them to the earliest evolutionary phase of Homo sapiens. Both papers suggest that the evolutionary processes behind the emergence of modern humans were not confined to sub-Saharan Africa. Fossil evidence points to an African origin of Homo sapiens from a group called either H. heidelbergensis or H. rhodesiensis. However, the exact place and time of emergence of H. sapiens remain obscure because the fossil record is scarce and the chronological age of many key specimens remains uncertain. In particular, it is unclear whether the present day ‘modern’ morphology rapidly emerged approximately 200 thousand years ago (ka) among earlier representatives of H. sapiens1 or evolved gradually over the last 400 thousand years2. Here we report newly discovered human fossils from Jebel Irhoud, Morocco, and interpret the affinities of the hominins from this site with other archaic and recent human groups. We identified a mosaic of features including facial, mandibular and dental morphology that aligns the Jebel Irhoud material with early or recent anatomically modern humans and more primitive neurocranial and endocranial morphology. In combination with an age of 315 ± 34 thousand years (as determined by thermoluminescence dating)3, this evidence makes Jebel Irhoud the oldest and richest African Middle Stone Age hominin site that documents early stages of the H. sapiens clade in which key features of modern morphology were established. 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Paris; citation_title=L’humerus d’enfant moustérien (Homo 4) du Jebel Irhoud (Maroc) dans son contexte archéologique; citation_author=J-J Hublin, AM Tillier, J Tixier; citation_volume=4; citation_publication_date=1987; citation_pages=115-141; citation_id=CR6"/> <meta name="citation_reference" content="citation_journal_title=Actes des 1ères Journées Nationales d’Archéologie et du Patrimoine. Préhistoire Vol.; citation_author=J Tixier, J-P Brugal, A-M Tillier, J Bruzeket, J-J Hublin; citation_volume=1; citation_publication_date=2001; citation_pages=149-153; citation_id=CR7"/> <meta name="citation_reference" content="citation_journal_title=Comptes Rendus à l'Académie des Sciences de Paris; citation_title=Le gisement moustérien du Djebel Irhoud, Maroc: précisions sur la faune et la biochronologie, et description d’un nouveau reste humain; citation_author=F Amani, D Geraads; citation_volume=316; citation_publication_date=1993; citation_pages=847-852; citation_id=CR8"/> <meta name="citation_reference" content="citation_journal_title=Anthropologie; citation_title=Un Néanderthalien: l’homme du Jebel Irhoud (Maroc); citation_author=E Ennouchi; citation_volume=66; citation_publication_date=1962; citation_pages=279-299; citation_id=CR9"/> <meta name="citation_reference" content="Stringer, C. B in Recent advances in Primatology. (eds Chivers, D. J . & Joysey, K. A. ) 395–418 (Academic, 1978)"/> <meta name="citation_reference" content="citation_journal_title=Phil. Trans. R. Soc. B; citation_title=Recent human evolution in northwestern Africa; citation_author=J-J Hublin; citation_volume=337; citation_publication_date=1992; citation_pages=185-191; citation_id=CR11"/> <meta name="citation_reference" content="citation_journal_title=Quat. Res.; citation_title=The rodents from the late middle Pleistocene hominid-bearing site of J’bel Irhoud, Morocco, and their chronological and paleoenvironmental implications; citation_author=D Geraads; citation_volume=80; citation_publication_date=2013; citation_pages=552-561; citation_id=CR12"/> <meta name="citation_reference" content="citation_journal_title=Proc. Natl Acad. Sci. USA; citation_title=Earliest evidence of modern human life history in North African early Homo sapiens; citation_author=TM Smith; citation_volume=104; citation_publication_date=2007; citation_pages=6128-6133; citation_id=CR13"/> <meta name="citation_reference" content="citation_journal_title=Mol. Biol. Evol.; citation_title=Whole-mtDNA genome sequence analysis of ancient African lineages; citation_author=MK Gonder, HM Mortensen, FA Reed, A de Sousa, SA Tishkoff; citation_volume=24; citation_publication_date=2007; citation_pages=757-768; citation_id=CR14"/> <meta name="citation_reference" content="citation_journal_title=Nature; citation_title=Stratigraphic placement and age of modern humans from Kibish, Ethiopia; citation_author=I McDougall, FH Brown, JG Fleagle; citation_volume=433; citation_publication_date=2005; citation_pages=733-736; citation_id=CR15"/> <meta name="citation_reference" content="citation_journal_title=Nature; citation_title=Pleistocene Homo sapiens from Middle Awash, Ethiopia; citation_author=TD White; citation_volume=423; citation_publication_date=2003; citation_pages=742-747; citation_id=CR16"/> <meta name="citation_reference" content="Smith, F. H. in Continuity or Replacement, Controversies in Homo sapiens evolution (eds Bräuer, G . & Smith, F. H. ) 145–156 (A. A. Balkema, Zagreb, 1992)"/> <meta name="citation_reference" content="citation_journal_title=Anthropol. Sci.; citation_title=Neurocranial evolution in modern humans: the case of Jebel Irhoud 1; citation_author=E Bruner, O Pearson; citation_volume=121; citation_publication_date=2013; citation_pages=31-41; citation_id=CR18"/> <meta name="citation_reference" content="citation_journal_title=Anthropol. Sci.; citation_title=Evolutionary interpretation of the modern human-like facial morphology of the Atapuerca Gran Dolina-TD6 hominins; citation_author=JM Bermúdez de Castro, M Martinón-Torres; citation_volume=122; citation_publication_date=2014; citation_pages=149-155; citation_id=CR19"/> <meta name="citation_reference" content="citation_journal_title=J. Hum. Evol.; citation_title=The human chin revisited: what is it and who has it?; citation_author=JH Schwartz, I Tattersall; citation_volume=38; citation_publication_date=2000; citation_pages=367-409; citation_id=CR20"/> <meta name="citation_reference" content="citation_journal_title=J. Hum. Evol.; citation_title=Dental remains from the Shanidar adult Neanderthals; citation_author=E Trinkaus; citation_volume=7; citation_publication_date=1978; citation_pages=369-382; citation_id=CR21"/> <meta name="citation_reference" content="citation_journal_title=J. Hum. Evol.; citation_title=A uniquely modern human pattern of endocranial development. Insights from a new cranial reconstruction of the Neandertal newborn from Mezmaiskaya; citation_author=P Gunz; citation_volume=62; citation_publication_date=2012; citation_pages=300-313; citation_id=CR22"/> <meta name="citation_reference" content="Klein, R. G. The Human Career: Human Biological and Cultural Origins 3rd edn (Chicago Univ. Press, 2009)"/> <meta name="citation_reference" content="citation_journal_title=Nature; citation_title=Direct dating of Florisbad hominid; citation_author=R Grün; citation_volume=382; citation_publication_date=1996; citation_pages=500-501; citation_id=CR24"/> <meta name="citation_reference" content="Stringer, C. The Origin of Our Species. (Allen Lane, Penguin, 2011)"/> <meta name="citation_reference" content="citation_journal_title=PLoS ONE; citation_title=The later Stone Age calvaria from Iwo Eleru, Nigeria: morphology and chronology; citation_author=K Harvati; citation_volume=6; citation_publication_date=2011; citation_pages=e24024; citation_id=CR26"/> <meta name="citation_reference" content="citation_journal_title=Proc. Natl Acad. Sci. USA; citation_title=Early modern human diversity suggests subdivided population structure and a complex out-of-Africa scenario; citation_author=P Gunz; citation_volume=106; citation_publication_date=2009; citation_pages=6094-6098; citation_id=CR27"/> <meta name="citation_reference" content="citation_journal_title=Science; citation_title=Neandertal roots: cranial and chronological evidence from Sima de los Huesos; citation_author=JL Arsuaga; citation_volume=344; citation_publication_date=2014; citation_pages=1358-1363; citation_id=CR28"/> <meta name="citation_reference" content="citation_journal_title=Science; citation_title=A high-coverage genome sequence from an archaic Denisovan individual; citation_author=M Meyer; citation_volume=338; citation_publication_date=2012; citation_pages=222-226; citation_id=CR29"/> <meta name="citation_reference" content="citation_journal_title=J. Hum. Evol.; citation_title=Did a discrete event 200,000–100,000 years ago produce modern humans?; citation_author=TD Weaver; citation_volume=63; citation_publication_date=2012; citation_pages=121-126; citation_id=CR30"/> <meta name="citation_reference" content="citation_journal_title=Source Code Biol. Med.; citation_title=MIA - a free and open source software for gray scale medical image analysis; citation_author=G Wollny; citation_volume=8; citation_publication_date=2013; citation_pages=20; citation_id=CR31"/> <meta name="citation_reference" content="citation_journal_title=Med. Image Anal.; citation_title=Landmark methods for forms without landmarks: morphometrics of group differences in outline shape; citation_author=FL Bookstein; citation_volume=1; citation_publication_date=1997; citation_pages=225-243; citation_id=CR32"/> <meta name="citation_reference" content="Gunz, P., Mitteroecker, P. & Bookstein, F. L. in Modern Morphometrics in Physical Anthropology. (ed. Slice, D.E. ) 73–98 (Kluwer Academic/Plenum Publishers, 2005)"/> <meta name="citation_reference" content="citation_journal_title=Hystrix; citation_title=Semilandmarks: a method for quantifying curves and surfaces; citation_author=P Gunz, P Mitteroecker; citation_volume=24; citation_publication_date=2013; citation_pages=103-109; citation_id=CR34"/> <meta name="citation_reference" content="citation_journal_title=16th IEEE Visualization Conference (VIS 2005); citation_author=DF Wiley; citation_volume=55; citation_publication_date=2005; citation_pages=431-438; citation_id=CR35"/> <meta name="citation_reference" content="citation_journal_title=J. Hum. Evol.; citation_title=Middle Pleistocene human facial morphology in an evolutionary and developmental context; citation_author=SE Freidline, P Gunz, K Harvati, J-J Hublin; citation_volume=63; citation_publication_date=2012; citation_pages=723-740; citation_id=CR36"/> <meta name="citation_reference" content="citation_journal_title=J. Anat.; citation_title=The pattern of endocranial ontogenetic shape changes in humans; citation_author=S Neubauer, P Gunz, J-J Hublin; citation_volume=215; citation_publication_date=2009; citation_pages=240-255; citation_id=CR37"/> <meta name="citation_reference" content="citation_journal_title=J. Hum. Evol.; citation_title=Cioclovina (Romania): affinities of an early modern European; citation_author=K Harvati, P Gunz, D Grigorescu; citation_volume=53; citation_publication_date=2007; citation_pages=732-746; citation_id=CR38"/> <meta name="citation_reference" content="citation_journal_title=Nature; citation_title=Early dispersal of modern humans in Europe and implications for Neanderthal behaviour; citation_author=S Benazzi; citation_volume=479; citation_publication_date=2011; citation_pages=525-528; citation_id=CR39"/> <meta name="citation_reference" content="citation_journal_title=Am. J. Phys. Anthropol.; citation_title=Allometry, merism, and tooth shape of the upper deciduous M2 and permanent M1; citation_author=SE Bailey, S Benazzi, J-J Hublin; citation_volume=154; citation_publication_date=2014; citation_pages=104-114; citation_id=CR40"/> <meta name="citation_reference" content="citation_journal_title=Am. J. Phys. Anthropol.; citation_title=Cervical and crown outline analysis of worn Neanderthal and modern human lower second deciduous molars; citation_author=S Benazzi; citation_volume=149; citation_publication_date=2012; citation_pages=537-546; citation_id=CR41"/> <meta name="citation_reference" content="citation_journal_title=J. Hum. Evol.; citation_title=Enamel–dentine junction (EDJ) morphology distinguishes the lower molars of Australopithecus africanus and Paranthropus robustus; citation_author=MM Skinner, P Gunz, BA Wood, J-J Hublin; citation_volume=55; citation_publication_date=2008; citation_pages=979-988; citation_id=CR42"/> <meta name="citation_reference" content="citation_journal_title=Am. J. Phys. Anthropol.; citation_title=Discrimination of extant Pan species and subspecies using the enamel–dentine junction morphology of lower molars; citation_author=MM Skinner, P Gunz, BA Wood, C Boesch, J-J Hublin; citation_volume=140; citation_publication_date=2009; citation_pages=234-243; citation_id=CR43"/> <meta name="citation_reference" content="citation_journal_title=Nature; citation_title=Reconstructed Homo habilis type OH 7 suggests deep-rooted species diversity in early Homo; citation_author=F Spoor; citation_volume=519; citation_publication_date=2015; citation_pages=83-86; citation_id=CR44"/> <meta name="citation_reference" content="Beucher, S . & Lantuéjoul, C. in International Workshop on Image Processing: Real-Time Edge and Motion Detection 2.1–2.12 (Rennes, France, 1979)"/> <meta name="citation_reference" content="citation_journal_title=J. Hum. Evol.; citation_title=Anterior tooth root morphology and size in Neanderthals: taxonomic and functional implications; citation_author=A Le Cabec, P Gunz, K Kupczik, J Braga, J-J Hublin; citation_volume=64; citation_publication_date=2013; citation_pages=169-193; citation_id=CR46"/> <meta name="citation_reference" content="citation_journal_title=Evol. Biol.; citation_title=Advances in geometric morphometrics; citation_author=P Mitteroecker, P Gunz; citation_volume=36; citation_publication_date=2009; citation_pages=235-247; citation_id=CR47"/> <meta name="citation_reference" content="citation_journal_title=J. Hum. Evol.; citation_title=Principles for the virtual reconstruction of hominin crania; citation_author=P Gunz, P Mitteroecker, S Neubauer, GW Weber, FL Bookstein; citation_volume=57; citation_publication_date=2009; citation_pages=48-62; citation_id=CR48"/> <meta name="citation_reference" content="citation_journal_title=Syst. Zool.; citation_title=Extensions of the procrustes method for the optimal superimposition of landmarks; citation_author=FJ Rohlf, D Slice; citation_volume=39; citation_publication_date=1990; citation_pages=40-59; citation_id=CR49"/> <meta name="citation_reference" content="citation_journal_title=Evol. Biol.; citation_title=Linear discrimination, ordination, and the visualization of selection gradients in modern morphometrics; citation_author=P Mitteroecker, F Bookstein; citation_volume=38; citation_publication_date=2011; citation_pages=100-114; citation_id=CR50"/> <meta name="citation_reference" content="R Development Core Team. R: a language and environment for statistical computing http://www.r-project.org (R Foundation for Statistical Computing, Vienna, Austria, 2012)"/> <meta name="citation_reference" content="citation_journal_title=Am. J. Phys. Anthropol.; citation_title=The fossil human remains from the Paleolithic site of Sidi Abderrahman (Morocco); citation_author=C Arambourg, P Biberson; citation_volume=14; citation_publication_date=1956; citation_pages=467-489; citation_id=CR52"/> <meta name="citation_reference" content="citation_journal_title=Anthropologie; citation_title=La mandibule atlanthropienne de la carrière Thomas I (Casablanca); citation_author=F Sausse; citation_volume=79; citation_publication_date=1975; citation_pages=81-112; citation_id=CR53"/> <meta name="citation_reference" content="citation_journal_title=J. Hum. Evol.; citation_title=Comparative studies of Late Pleistocene human remains from Klasies River Mouth, South Africa; citation_author=GP Rightmire; citation_volume=20; citation_publication_date=1991; citation_pages=131-156; citation_id=CR54"/> <meta name="citation_reference" content="Stewart, T. D. in Annual Report of the Smithsonian Institution 521–533 (US Government Printing Office, Washington, 1962)"/> <meta name="citation_reference" content="citation_journal_title=Le Squelette moustérien de Kébara; citation_author=A-M Tillier; citation_volume=2; citation_publication_date=1991; citation_pages=97-112; citation_id=CR56"/> <meta name="citation_reference" content="citation_journal_title=Proc. Am. Phil. Soc.; citation_title=The Neanderthal skeletal remains from Shanidar Cave, Iraq: a summary of findings to date.; citation_author=TD Stewart; citation_volume=121; citation_publication_date=1977; citation_pages=121-165; citation_id=CR57"/> <meta name="citation_reference" content="Trinkaus, E. The Shanidar Neanderthals (Academic, 1983)"/> <meta name="citation_reference" content="Leroi-Gourhan, A. Étude des Restes Humains Fossiles provenant des Grottes d’Arcy-sur-Cure (Masson et Cie, 1958)"/> <meta name="citation_reference" content="citation_journal_title=J. Hum. Evol.; citation_title=A Neandertal mandible from the Cova del Gegant (Sitges, Barcelona, Spain); citation_author=J Daura; citation_volume=49; citation_publication_date=2005; citation_pages=56-70; citation_id=CR60"/> <meta name="citation_reference" content="citation_journal_title=Rev. Antropol.; citation_title=Les caracteres simiens de la machoire de la Naulette.; citation_author=P Topinard; citation_volume=15; citation_publication_date=1886; citation_pages=385-431; citation_id=CR61"/> <meta name="citation_reference" content="citation_journal_title=Anthropol. Rev.; citation_title=On a human jaw from the cave of La Naulette, near Dinant, Belgium; citation_author=CC Blake; citation_volume=5; citation_publication_date=1867; citation_pages=294-303; citation_id=CR62"/> <meta name="citation_reference" content="Leguebe, A. & Toussaint, M. La Mandibule et les Cubitus de la Naulette: Morphologie et Morphométrie 15 (Editions du Centre National de la Recherche Scientifique, 1988)"/> <meta name="citation_reference" content="citation_journal_title=Anthropologie; citation_title=Les hommes fossiles de La Ferrassie (Dordogne) et le problem de la definition des Neandertaliens classiques. III. Squelette céphalique; citation_author=JL Heim; citation_volume=78; citation_publication_date=1974; citation_pages=321-378; citation_id=CR64"/> <meta name="citation_reference" content="citation_journal_title=Etudes Quaternaires; citation_title=Les Néandertaliens de la grotte de l’Hortus.; citation_author=M-A de Lumley; citation_volume=1; citation_publication_date=1972; citation_pages=375-385; citation_id=CR65"/> <meta name="citation_reference" content="citation_journal_title=PLoS ONE; citation_title=Possible interbreeding in late Italian Neanderthals? New data from the Mezzena jaw (Monti Lessini, Verona, Italy); citation_author=S Condemi; citation_volume=8; citation_publication_date=2013; citation_pages=e59781; citation_id=CR66"/> <meta name="citation_reference" content="citation_journal_title=Memorie del Museo civico di Storia naturale di Verona; citation_title=Resti scheletrici umani del ‘Riparo Mezzena’; citation_author=C Corrain; citation_volume=16; citation_publication_date=1968; citation_pages=97-101; citation_id=CR67"/> <meta name="citation_reference" content="citation_journal_title=Am. J. Phys. Anthropol.; citation_title=Neandertal mandibles from the Sima de las Palomas del Cabezo Gordo, Murcia, southeastern Spain; citation_author=MJ Walker, AV Lombardi, J Zapata, E Trinkaus; citation_volume=142; citation_publication_date=2010; citation_pages=261-272; citation_id=CR68"/> <meta name="citation_reference" content="citation_journal_title=L’homme préhistorique; citation_title=Machoire humaine moustérienne trouvée dans la station de La Quina; citation_author=H Martin; citation_volume=13; citation_publication_date=1926; citation_pages=3-21; citation_id=CR69"/> <meta name="citation_reference" content="citation_journal_title=Bull. Soc. Préhistorique; citation_title=Position stratigraphique des Ossements humains recueillis dans le Moustérien de La Quina de 1908 à 1912; citation_author=H Martin; citation_volume=9; citation_publication_date=1912; citation_pages=700-709; citation_id=CR70"/> <meta name="citation_reference" content="Martin, H. L’Homme fossile de la Quina. (Libraire Octave Doin, 1923)"/> <meta name="citation_reference" content="citation_journal_title=Ann. Hist. Nat. Mus. Natl. Hung.; citation_title=The Subalyuk Neanderthal remains (Hungary): a re-examination; citation_author=I Pap, AM Tillier, B Arensburg, M Chech; citation_volume=88; citation_publication_date=1996; citation_pages=233-270; citation_id=CR72"/> <meta name="citation_reference" content="citation_journal_title=Hum. Evol.; citation_title=Comparative biometrical study of the Mousterian mandible from Cueva del Boquete de Zafarraya (Málaga, Spain); citation_author=F Sanchez; citation_volume=14; citation_publication_date=1999; citation_pages=125-138; citation_id=CR73"/> <meta name="citation_reference" content="Vlcˇek, E. Fossile Menschenfunde von Weimar-Ehringsdorf, Weimarer Monographien zur Ur- und Frühgeschichte Vol. 30 (Landesamt für Archäologische Denkmalpflege, 1993)"/> <meta name="citation_reference" content="Condemi, S. Les néandertaliens de La Chaise: abri Bourgeois-Delaunay. Comité des travaux historiques et scientifiques (CTHS, 2001)"/> <meta name="citation_reference" content="Bar-Yosef, O . & Vandermeersch, B . (eds) Le squelette moustérien de Kébara 2. (Editions du Centre National de la Recherche Scientifique, 1991)"/> <meta name="citation_reference" content="Arambourg, C . & Hoffstetter, R. Le gisement de Ternifine Vol. 1. (Masson, 1963)"/> <meta name="citation_reference" content="citation_journal_title=The ancient Palestinian: Skhu¯ l V reconstruction.; citation_author=EA Hooton, HO Hencken, ChE Snow; citation_volume=17; citation_publication_date=1953; citation_pages=5-10; citation_id=CR78"/> <meta name="citation_reference" content="citation_journal_title=J. R. Anthropol. Inst.; citation_title=The Chancelade skull; citation_author=WJ Sollas; citation_volume=57; citation_publication_date=1927; citation_pages=89-122; citation_id=CR79"/> <meta name="citation_reference" content="citation_journal_title=Bull. Mem. Soc. Anthropol. Paris; citation_title=Caractères des squelettes humains quaternaires de la vallée du Roc (Charente); citation_author=H Martin; citation_volume=8; citation_publication_date=1927; citation_pages=103-129; citation_id=CR80"/> <meta name="citation_reference" content="citation_journal_title=J. Anthropol. Sci.; citation_title=The Late Upper Paleolithic skeleton Villabruna 1 (Italy): a source of data on biology and behavior of a 14,000 year-old hunter; citation_author=G Vercellotti, G Alciati, MP Richards, V Formicola; citation_volume=86; citation_publication_date=2008; citation_pages=143-163; citation_id=CR81"/> <meta name="citation_reference" content="citation_journal_title=Rev. Antropol.; citation_title=Una mandibola umana dal deposito dell’Epigravettiano finale delle Arene Candite (scavi 197O).; citation_author=V Formicola; citation_volume=64; citation_publication_date=1986; citation_pages=271-278; citation_id=CR82"/> <meta name="citation_reference" content="citation_journal_title=Bull. d’Archéologie Marocaine; citation_title=Le gisement préhistorique de Dar-es-Soltane 2. Champ de tir de El Menzeh à Rabat (Maroc). Note préliminaire. 2-Étude anthropologique des restes post-atériens; citation_author=AM Odano, R Riquet; citation_volume=11; citation_publication_date=1978; citation_pages=25-63; citation_id=CR83"/> <meta name="citation_reference" content="citation_journal_title=CR Acad. Sci. Paris; citation_title=Nouveaux restes humains atériens du Maroc.; citation_author=A Debénath; citation_volume=290; citation_publication_date=1980; citation_pages=851-852; citation_id=CR84"/> <meta name="citation_reference" content="citation_journal_title=Paéorient; citation_title=Les Natoufiens du Nahal Oren (Ouadi Fallah) Etude anthropologique; citation_author=E Crognier, M Dupouy-Madre; citation_volume=2; citation_publication_date=1974; citation_pages=103-121; citation_id=CR85"/> <meta name="citation_reference" content="citation_journal_title=Z. Morphol. Anthropol.; citation_title=Vergleichend-morphologische Kennzeichnung der Jungpaläolithiker von Oberkassel bei Bonn.; citation_author=W Henke; citation_volume=75; citation_publication_date=1984; citation_pages=27-44; citation_id=CR86"/> <meta name="citation_reference" content="citation_journal_title=Am. J. Phys. Anthropol.; citation_title=Ohalo II H2: a 19,000-year-old skeleton from a water-logged site at the Sea of Galilee, Israel; citation_author=I Hershkovitz; citation_volume=96; citation_publication_date=1995; citation_pages=215-234; citation_id=CR87"/> <meta name="citation_reference" content="citation_journal_title=Proc. Natl Acad. Sci. USA; citation_title=Early modern humans from the Pestera Muierii, Baia de Fier, Romania; citation_author=A Soficaru, A Dobos, E Trinkaus; citation_volume=103; citation_publication_date=2006; citation_pages=17196-17201; citation_id=CR88"/> <meta name="citation_reference" content="Crevecoeur, I. Etude anthropologique des restes humains de Nazlet Khater (Paléolithique Superieur, Egypte). PhD Thesis, Université Sciences et Technologies Bordeaux I (2006)"/> <meta name="citation_reference" content="citation_journal_title=J. Hum. Evol.; citation_title=Morphology and affinities of the Nazlet Khater man; citation_author=A Thoma; citation_volume=13; citation_publication_date=1984; citation_pages=287-296; citation_id=CR90"/> <meta name="citation_reference" content="citation_journal_title=The Prehistory of Nubia; citation_author=JE Anderson, F Wendorf; citation_volume=2; citation_publication_date=1968; citation_pages=996-1040; citation_id=CR91"/> <meta name="citation_reference" content="citation_journal_title=Anthropologie; citation_title=From the Nile to the Danube: a comparison of the Nazlet Khater 2 and Oase 1 early modern human mandibles; citation_author=I Crevecoeur; citation_volume=42; citation_publication_date=2004; citation_pages=203-213; citation_id=CR92"/> <meta name="citation_reference" content="citation_journal_title=Proc. Natl Acad. Sci. USA; citation_title=An early modern human from the Pes¸ tera cu Oase, Romania; citation_author=E Trinkaus; citation_volume=100; citation_publication_date=2003; citation_pages=11231-11236; citation_id=CR93"/> <meta name="citation_reference" content="Trinkaus, E. & Svoboda, J. (eds) Early Modern Human Evolution in Central Europe. The People of Dolní Veˇ stonice and Pavlov. The Dolni Vestonice Studies Vol. 12 (Oxford Univ. Press, 2006)"/> <meta name="citation_reference" content="citation_journal_title=The Dolni Vestonice Studies; citation_title=The People of the Pavlovian. Skeletal Catalogue and Osteometrics of the Gravettian Fossil Hominids from Dolni Vestonice and Pavlov.; citation_author=V Sládek, E Trinkaus, SW Hillson, TW Holliday; citation_volume=5; citation_publication_date=2000; citation_pages=1-244; citation_id=CR95"/> <meta name="citation_reference" content="citation_journal_title=Archeologické rozhledy; citation_title=The evaluation of a rediscovered fragment of human lower jaw, No 21 from Prˇedmostí u Prˇerova.; citation_author=E Drozdová; citation_volume=53; citation_publication_date=2001; citation_pages=452-460; citation_id=CR96"/> <meta name="citation_reference" content="citation_journal_title=Travaux du Laboratoire d’Anthropologie et de Préhistoire des Pays de la Méditerranée Occidentale; citation_title=Palimpseste paléoanthropologique sur l’«Homme fossile d’Asselar» (Sahara); citation_author=O Dutour; citation_volume=1; citation_publication_date=1992; citation_pages=73-83; citation_id=CR97"/> <meta name="citation_reference" content="citation_journal_title=Antiquités nationales; citation_title=Vestiges humains du gisement d’Isturitz (Pyrénées—Atlantiques): étude anthropologique et analyse des traces d’action humaine intentionelle; citation_author=D Gambier; citation_volume=22–23; citation_publication_date=1990; citation_pages=9-26; citation_id=CR98"/> <meta name="citation_reference" content="citation_journal_title=Am. J. Phys. Anthropol.; citation_title=Hominid molars from a middle Stone Age level at the Mumba Rock Shelter, Tanzania; citation_author=G Bräuer, MJ Mehlman; citation_volume=75; citation_publication_date=1988; citation_pages=69-76; citation_id=CR99"/> <meta name="citation_reference" content="Protsch, R. R. R. Di Archäologischen und Anthropologischen Ergebnisse der Kohl-Larsen-Expeditionen in Nord-Tanzania 1933–1939. (Institut für Urgeschichte der Universität Tübingen, 1981)"/> <meta name="citation_reference" content="citation_journal_title=Am. J. Phys. Anthropol.; citation_title=Preliminary observations on the BK 8518 mandible from Baringo, Kenya; citation_author=BA Wood, FL Van Noten; citation_volume=69; citation_publication_date=1986; citation_pages=117-127; citation_id=CR101"/> <meta name="citation_reference" content="citation_journal_title=Am. J. Phys. Anthropol.; citation_title=Middle Pleistocene hominids from Olduvai Gorge, northern Tanzania; citation_author=GP Rightmire; citation_volume=53; citation_publication_date=1980; citation_pages=225-241; citation_id=CR102"/> <meta name="citation_reference" content="citation_journal_title=J. Hum. Evol.; citation_title=Dental remains from Atapuerca (Spain) I. Metrics.; citation_author=JM Bermúdez de Castro; citation_volume=15; citation_publication_date=1986; citation_pages=265-287; citation_id=CR103"/> <meta name="citation_reference" content="citation_journal_title=Courier Forschungsinstitut Senckenberg; citation_title=Fossil hominid teeth from the Sangiran Dome (Java, Indonesia); citation_author=FE Grine, JL Franzen; citation_volume=171; citation_publication_date=1994; citation_pages=75-103; citation_id=CR104"/> <meta name="citation_reference" content="citation_journal_title=J. Hum. Evol.; citation_title=Anterior tooth root morphology and size in Neanderthals: taxonomic and functional implications; citation_author=A Le Cabec, P Gunz, K Kupczik, J Braga, J-J Hublin; citation_volume=64; citation_publication_date=2013; citation_pages=169-193; citation_id=CR105"/> <meta name="citation_reference" content="Turner, C. G. II, Nichol, C. R. & Scott, G. R. Advances in Dental Anthropology (eds Kelley, M., & Larsen, C. ) 13–31 (Wiley Liss, 1991)"/> <meta name="citation_reference" content="Bailey, S. E. Neandertal Dental Morphology: Implications for modern human origins. PhD thesis, Arizona State Univ. (2002)"/> <meta name="citation_author" content="Hublin, Jean-Jacques"/> <meta name="citation_author_institution" content="Department of Human Evolution, Max Planck Institute for Evolutionary Anthropology, Leipzig, Germany"/> <meta name="citation_author_institution" content="Chaire Internationale de Paléoanthropologie, Collège de France, Paris, France"/> <meta name="citation_author" content="Ben-Ncer, Abdelouahed"/> <meta name="citation_author_institution" content="Institut National des Sciences de l’Archéologie et du Patrimoine, Rabat"/> <meta name="citation_author" content="Bailey, Shara E."/> <meta name="citation_author_institution" content="Department of Anthropology, Center for the Study of Human Origins, New York University, New York, USA"/> <meta name="citation_author" content="Freidline, Sarah E."/> <meta name="citation_author_institution" content="Department of Human Evolution, Max Planck Institute for Evolutionary Anthropology, Leipzig, Germany"/> <meta name="citation_author" content="Neubauer, Simon"/> <meta name="citation_author_institution" content="Department of Human Evolution, Max Planck Institute for Evolutionary Anthropology, Leipzig, Germany"/> <meta name="citation_author" content="Skinner, Matthew M."/> <meta name="citation_author_institution" content="School of Anthropology and Conservation, University of Kent, Canterbury, UK"/> <meta name="citation_author" content="Bergmann, Inga"/> <meta name="citation_author_institution" content="Department of Human Evolution, Max Planck Institute for Evolutionary Anthropology, Leipzig, Germany"/> <meta name="citation_author" content="Le Cabec, Adeline"/> <meta name="citation_author_institution" content="Department of Human Evolution, Max Planck Institute for Evolutionary Anthropology, Leipzig, Germany"/> <meta name="citation_author" content="Benazzi, Stefano"/> <meta name="citation_author_institution" content="Department of Cultural Heritage, University of Bologna, Ravenna, Italy"/> <meta name="citation_author" content="Harvati, Katerina"/> <meta name="citation_author_institution" content="Paleoanthropology, Senckenberg Center for Human Evolution and Paleoenvironment, and DFG Center for Advanced Studies: “Words, Bones, Genes, Tools”, Eberhard Karls Universität, Tübingen, Germany"/> <meta name="citation_author" content="Gunz, Philipp"/> <meta name="citation_author_institution" content="Department of Human Evolution, Max Planck Institute for Evolutionary Anthropology, Leipzig, Germany"/> <meta name="access_endpoint" content="https://www.nature.com/platform/readcube-access"/> <meta name="twitter:site" content="@nature"/> <meta name="twitter:card" content="summary_large_image"/> <meta name="twitter:image:alt" content="Content cover image"/> <meta name="twitter:title" content="New fossils from Jebel Irhoud, Morocco and the pan-African origin of Homo sapiens"/> <meta name="twitter:description" content="Nature - The exact place and time that our species emerged remains obscure because the fossil record is limited and the chronological age of many key specimens remains uncertain. 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class="c-article-header"> <header> <ul class="c-article-identifiers" data-test="article-identifier"> <li class="c-article-identifiers__item" data-test="article-category">Letter</li> <li class="c-article-identifiers__item">Published: <time datetime="2017-06-08">08 June 2017</time></li> </ul> <h1 class="c-article-title" data-test="article-title" data-article-title="">New fossils from Jebel Irhoud, Morocco and the pan-African origin of <i>Homo sapiens</i></h1> <ul class="c-article-author-list c-article-author-list--short" data-test="authors-list" data-component-authors-activator="authors-list"><li class="c-article-author-list__item"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Jean_Jacques-Hublin-Aff1-Aff2" data-author-popup="auth-Jean_Jacques-Hublin-Aff1-Aff2" data-author-search="Hublin, Jean-Jacques" data-corresp-id="c1">Jean-Jacques Hublin<svg width="16" height="16" focusable="false" role="img" aria-hidden="true" class="u-icon"><use xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="#icon-eds-i-mail-medium"></use></svg></a><sup class="u-js-hide"><a href="#Aff1">1</a>,<a href="#Aff2">2</a></sup>, </li><li class="c-article-author-list__item"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Abdelouahed-Ben_Ncer-Aff3" data-author-popup="auth-Abdelouahed-Ben_Ncer-Aff3" data-author-search="Ben-Ncer, Abdelouahed">Abdelouahed Ben-Ncer</a><sup class="u-js-hide"><a href="#Aff3">3</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Shara_E_-Bailey-Aff4" data-author-popup="auth-Shara_E_-Bailey-Aff4" data-author-search="Bailey, Shara E.">Shara E. Bailey</a><sup class="u-js-hide"><a href="#Aff4">4</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Sarah_E_-Freidline-Aff1" data-author-popup="auth-Sarah_E_-Freidline-Aff1" data-author-search="Freidline, Sarah E.">Sarah E. Freidline</a><sup class="u-js-hide"><a href="#Aff1">1</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Simon-Neubauer-Aff1" data-author-popup="auth-Simon-Neubauer-Aff1" data-author-search="Neubauer, Simon">Simon Neubauer</a><sup class="u-js-hide"><a href="#Aff1">1</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Matthew_M_-Skinner-Aff5" data-author-popup="auth-Matthew_M_-Skinner-Aff5" data-author-search="Skinner, Matthew M.">Matthew M. Skinner</a><sup class="u-js-hide"><a href="#Aff5">5</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Inga-Bergmann-Aff1" data-author-popup="auth-Inga-Bergmann-Aff1" data-author-search="Bergmann, Inga">Inga Bergmann</a><sup class="u-js-hide"><a href="#Aff1">1</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Adeline-Le_Cabec-Aff1" data-author-popup="auth-Adeline-Le_Cabec-Aff1" data-author-search="Le Cabec, Adeline">Adeline Le Cabec</a><sup class="u-js-hide"><a href="#Aff1">1</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" 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Philipp" data-corresp-id="c2">Philipp Gunz<svg width="16" height="16" focusable="false" role="img" aria-hidden="true" class="u-icon"><use xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="#icon-eds-i-mail-medium"></use></svg></a><sup class="u-js-hide"><a href="#Aff1">1</a></sup> </li></ul><button aria-expanded="false" class="c-article-author-list__button"><svg width="16" height="16" focusable="false" role="img" aria-hidden="true" class="u-icon"><use xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="#icon-eds-i-chevron-down-medium"></use></svg><span>Show authors</span></button> <p class="c-article-info-details" data-container-section="info"> <a data-test="journal-link" href="/" data-track="click" data-track-action="journal homepage" data-track-category="article body" data-track-label="link"><i data-test="journal-title">Nature</i></a> <b data-test="journal-volume"><span class="u-visually-hidden">volume</span> 546</b>, <span class="u-visually-hidden">pages </span>289–292 (<span 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</header> <div class="u-js-hide" data-component="article-subject-links"> <h3 class="c-article__sub-heading">Subjects</h3> <ul class="c-article-subject-list"> <li class="c-article-subject-list__subject"><a href="/subjects/archaeology" data-track="click" data-track-action="view subject" data-track-label="link">Archaeology</a></li><li class="c-article-subject-list__subject"><a href="/subjects/biological-anthropology" data-track="click" data-track-action="view subject" data-track-label="link">Biological anthropology</a></li> </ul> </div> <div class="u-mb-8 c-status-message c-status-message--boxed c-status-message--info"> <span class="c-status-message__icon"> <svg class="u-icon" width="18" height="18" aria-hidden="true" focusable="false"> <use xlink:href="#icon-eds-i-info-filled-medium"></use> </svg> </span> <p class="u-mt-0">A <a href="https://doi.org/10.1038/s41586-018-0166-3" class="relation-link" data-track="click" data-track-action="view linked article" data-track-label="link">Correction</a> to this article was published on 13 June 2018</p> </div> <div class="u-mb-8 c-status-message c-status-message--boxed c-status-message--info"> <span class="c-status-message__icon"> <svg class="u-icon" width="18" height="18" aria-hidden="true" focusable="false"> <use xlink:href="#icon-eds-i-info-filled-medium"></use> </svg> </span> <p class="u-mt-0">This article has been <a href="#change-history">updated</a></p> </div> </div> <div class="c-article-body"> <section aria-labelledby="Abs3" data-title="Abstract" lang="en"><div class="c-article-section" id="Abs3-section"><h2 class="c-article-section__title js-section-title js-c-reading-companion-sections-item" id="Abs3">Abstract</h2><div class="c-article-section__content" id="Abs3-content"><p>Fossil evidence points to an African origin of <i>Homo sapiens</i> from a group called either <i>H. heidelbergensis</i> or <i>H. rhodesiensis</i>. However, the exact place and time of emergence of <i>H. sapiens</i> remain obscure because the fossil record is scarce and the chronological age of many key specimens remains uncertain. In particular, it is unclear whether the present day ‘modern’ morphology rapidly emerged approximately 200 thousand years ago (ka) among earlier representatives of <i>H. sapiens</i><sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 1" title="Stringer, C. Modern human origins: progress and prospects. Phil. Trans. R. Soc. B 357, 563–579 (2002)" href="/articles/nature22336#ref-CR1" id="ref-link-section-d136325645e573">1</a></sup> or evolved gradually over the last 400 thousand years<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2" title="Bräuer, G. The origin of modern anatomy: by speciation or intraspecific evolution? Evol. Anthropol. 17, 22–37 (2008)" href="/articles/nature22336#ref-CR2" id="ref-link-section-d136325645e578">2</a></sup>. Here we report newly discovered human fossils from Jebel Irhoud, Morocco, and interpret the affinities of the hominins from this site with other archaic and recent human groups. We identified a mosaic of features including facial, mandibular and dental morphology that aligns the Jebel Irhoud material with early or recent anatomically modern humans and more primitive neurocranial and endocranial morphology. In combination with an age of 315 ± 34 thousand years (as determined by thermoluminescence dating)<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 3" title="Richter, D. et al. The age of the Jebel Irhoud (Morocco) hominins and the origins of the Middle Stone Age. Nature 
 http://dx.doi.org/10.1038/nature22335
 
 (2017)" href="/articles/nature22336#ref-CR3" id="ref-link-section-d136325645e582">3</a></sup>, this evidence makes Jebel Irhoud the oldest and richest African Middle Stone Age hominin site that documents early stages of the <i>H. sapiens</i> clade in which key features of modern morphology were established. 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js-c-reading-companion-sections-item" id="change-history">Change history</h2><div class="c-article-section__content" id="change-history-content"><ul class="c-article-change-list"><li class="c-article-change-list__item u-mb-24" id="chg1"><ins datetime="2018-06-13"><h3 class="c-article-change-list__heading u-h3 u-pr-8 u-display-inline">13 June 2018</h3><div class="c-article-change-list__details"><p>In the originally published version of this Letter, the PC1 and PC2 values in Fig. 3a were incorrect; in addition, the enumeration of Qafzeh samples should have been 'Qafzeh (9, 25)', and Extended Data Fig. 1c and Extended Data Table 2 have been altered accordingly. This has been corrected online.</p></div></ins></li></ul></div></div></section><div id="MagazineFulltextArticleBodySuffix"><section aria-labelledby="Bib1" data-title="References"><div class="c-article-section" id="Bib1-section"><h2 class="c-article-section__title js-section-title js-c-reading-companion-sections-item" id="Bib1">References</h2><div class="c-article-section__content" id="Bib1-content"><div data-container-section="references"><ol class="c-article-references" data-track-component="outbound reference" data-track-context="references section"><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="1"><p class="c-article-references__text" id="ref-CR1">Stringer, C. Modern human origins: progress and prospects. <i>Phil. Trans. R. Soc. B</i> <b>357</b>, 563–579 (2002)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed reference" data-track-action="pubmed reference" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=12028792" aria-label="PubMed reference 1">PubMed</a> <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed central reference" data-track-action="pubmed central reference" href="http://www.ncbi.nlm.nih.gov/pmc/articles/PMC1692961" aria-label="PubMed Central reference 1">PubMed Central</a> <a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 1" href="http://scholar.google.com/scholar_lookup?&title=Modern%20human%20origins%3A%20progress%20and%20prospects&journal=Phil.%20Trans.%20R.%20Soc.%20B&volume=357&pages=563-579&publication_year=2002&author=Stringer%2CC"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="2"><p class="c-article-references__text" id="ref-CR2">Bräuer, G. The origin of modern anatomy: by speciation or intraspecific evolution? <i>Evol. Anthropol.</i> <b>17</b>, 22–37 (2008)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 2" href="http://scholar.google.com/scholar_lookup?&title=The%20origin%20of%20modern%20anatomy%3A%20by%20speciation%20or%20intraspecific%20evolution%3F&journal=Evol.%20Anthropol.&volume=17&pages=22-37&publication_year=2008&author=Br%C3%A4uer%2CG"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="3"><p class="c-article-references__text" id="ref-CR3">Richter, D. et al. The age of the Jebel Irhoud (Morocco) hominins and the origins of the Middle Stone Age. <i>Nature</i> <a href="https://doi.org/10.1038/nature22335" data-track="click_references" data-track-action="external reference" data-track-value="external reference" data-track-label="10.1038/nature22335">http://dx.doi.org/10.1038/nature22335</a> (2017)</p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="4"><p class="c-article-references__text" id="ref-CR4">Ennouchi, E. Le deuxième crâne de l’homme d’Irhoud. <i>Annales de Paléontologie (Vértébrés)</i> <b>LIV</b>, 117–128 (1968)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 4" href="http://scholar.google.com/scholar_lookup?&title=Le%20deuxi%C3%A8me%20cr%C3%A2ne%20de%20l%E2%80%99homme%20d%E2%80%99Irhoud&journal=Annales%20de%20Pal%C3%A9ontologie%20%28V%C3%A9rt%C3%A9br%C3%A9s%29&volume=LIV&pages=117-128&publication_year=1968&author=Ennouchi%2CE"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="5"><p class="c-article-references__text" id="ref-CR5">Hublin, J.-J. & Tillier, A. M. in <i>Aspects of Human Evolution</i> Vol. <b>21</b> Symposia of the Society for the study of Human Biology, Volume XXI (ed. Stringer, C. B. ) 167–186 (Taylor & Francis, 1981)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 5" href="http://scholar.google.com/scholar_lookup?&title=Symposia%20of%20the%20Society%20for%20the%20study%20of%20Human%20Biology%2C%20Volume%20XXI&journal=Aspects%20of%20Human%20Evolution&volume=21&pages=167-186&publication_year=1981&author=Hublin%2CJ-J&author=Tillier%2CAM&author=Stringer%2CCB"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="6"><p class="c-article-references__text" id="ref-CR6">Hublin, J.-J., Tillier, A. M. & Tixier, J. L’humerus d’enfant moustérien (Homo 4) du Jebel Irhoud (Maroc) dans son contexte archéologique. <i>Bull. Mem. Soc. Anthropol. Paris</i> <b>4</b>, 115–141 (1987)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 6" href="http://scholar.google.com/scholar_lookup?&title=L%E2%80%99humerus%20d%E2%80%99enfant%20moust%C3%A9rien%20%28Homo%204%29%20du%20Jebel%20Irhoud%20%28Maroc%29%20dans%20son%20contexte%20arch%C3%A9ologique&journal=Bull.%20Mem.%20Soc.%20Anthropol.%20Paris&volume=4&pages=115-141&publication_year=1987&author=Hublin%2CJ-J&author=Tillier%2CAM&author=Tixier%2CJ"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="7"><p class="c-article-references__text" id="ref-CR7">Tixier, J ., Brugal, J.-P ., Tillier, A.-M ., Bruzeket, J . & Hublin, J.-J. in <i>Actes des 1ères Journées Nationales d’Archéologie et du Patrimoine. Préhistoire Vol.</i> <b>1</b> 149–153 (Société Marocaine d’Archéologie et du Patrimoine, 2001)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 7" href="http://scholar.google.com/scholar_lookup?&title=&journal=Actes%20des%201%C3%A8res%20Journ%C3%A9es%20Nationales%20d%E2%80%99Arch%C3%A9ologie%20et%20du%20Patrimoine.%20Pr%C3%A9histoire%20Vol.&volume=1&pages=149-153&publication_year=2001&author=Tixier%2CJ&author=Brugal%2CJ-P&author=Tillier%2CA-M&author=Bruzeket%2CJ&author=Hublin%2CJ-J"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="8"><p class="c-article-references__text" id="ref-CR8">Amani, F. & Geraads, D. Le gisement moustérien du Djebel Irhoud, Maroc: précisions sur la faune et la biochronologie, et description d’un nouveau reste humain. <i>Comptes Rendus à l'Académie des Sciences de Paris</i> <b>316</b>, 847–852 (1993)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 8" href="http://scholar.google.com/scholar_lookup?&title=Le%20gisement%20moust%C3%A9rien%20du%20Djebel%20Irhoud%2C%20Maroc%3A%20pr%C3%A9cisions%20sur%20la%20faune%20et%20la%20biochronologie%2C%20et%20description%20d%E2%80%99un%20nouveau%20reste%20humain&journal=Comptes%20Rendus%20%C3%A0%20l%27Acad%C3%A9mie%20des%20Sciences%20de%20Paris&volume=316&pages=847-852&publication_year=1993&author=Amani%2CF&author=Geraads%2CD"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="9"><p class="c-article-references__text" id="ref-CR9">Ennouchi, E. Un Néanderthalien: l’homme du Jebel Irhoud (Maroc). <i>Anthropologie</i> <b>66</b>, 279–299 (1962)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 9" href="http://scholar.google.com/scholar_lookup?&title=Un%20N%C3%A9anderthalien%3A%20l%E2%80%99homme%20du%20Jebel%20Irhoud%20%28Maroc%29&journal=Anthropologie&volume=66&pages=279-299&publication_year=1962&author=Ennouchi%2CE"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="10"><p class="c-article-references__text" id="ref-CR10">Stringer, C. B in <i>Recent advances in Primatology.</i> (eds Chivers, D. J . & Joysey, K. A. ) 395–418 (Academic, 1978)</p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="11"><p class="c-article-references__text" id="ref-CR11">Hublin, J.-J. Recent human evolution in northwestern Africa. <i>Phil. Trans. R. Soc. B</i> <b>337</b>, 185–191 (1992)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="ads reference" data-track-action="ads reference" href="http://adsabs.harvard.edu/cgi-bin/nph-data_query?link_type=ABSTRACT&bibcode=1992RSPTB.337..185H" aria-label="ADS reference 11">ADS</a> <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="cas reference" data-track-action="cas reference" href="/articles/cas-redirect/1:STN:280:DyaK3s%2FivVOksQ%3D%3D" aria-label="CAS reference 11">CAS</a> <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed reference" data-track-action="pubmed reference" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=1357693" aria-label="PubMed reference 11">PubMed</a> <a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 11" href="http://scholar.google.com/scholar_lookup?&title=Recent%20human%20evolution%20in%20northwestern%20Africa&journal=Phil.%20Trans.%20R.%20Soc.%20B&volume=337&pages=185-191&publication_year=1992&author=Hublin%2CJ-J"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="12"><p class="c-article-references__text" id="ref-CR12">Geraads, D. et al. The rodents from the late middle Pleistocene hominid-bearing site of J’bel Irhoud, Morocco, and their chronological and paleoenvironmental implications. <i>Quat. Res.</i> <b>80</b>, 552–561 (2013)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 12" href="http://scholar.google.com/scholar_lookup?&title=The%20rodents%20from%20the%20late%20middle%20Pleistocene%20hominid-bearing%20site%20of%20J%E2%80%99bel%20Irhoud%2C%20Morocco%2C%20and%20their%20chronological%20and%20paleoenvironmental%20implications&journal=Quat.%20Res.&volume=80&pages=552-561&publication_year=2013&author=Geraads%2CD"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="13"><p class="c-article-references__text" id="ref-CR13">Smith, T. M. et al. Earliest evidence of modern human life history in North African early <i>Homo sapiens</i>. <i>Proc. Natl Acad. Sci. USA</i> <b>104</b>, 6128–6133 (2007)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="ads reference" data-track-action="ads reference" href="http://adsabs.harvard.edu/cgi-bin/nph-data_query?link_type=ABSTRACT&bibcode=2007PNAS..104.6128S" aria-label="ADS reference 13">ADS</a> <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="cas reference" data-track-action="cas reference" href="/articles/cas-redirect/1:CAS:528:DC%2BD2sXks1artLk%3D" aria-label="CAS reference 13">CAS</a> <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed reference" data-track-action="pubmed reference" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=17372199" aria-label="PubMed reference 13">PubMed</a> <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed central reference" data-track-action="pubmed central reference" href="http://www.ncbi.nlm.nih.gov/pmc/articles/PMC1828706" aria-label="PubMed Central reference 13">PubMed Central</a> <a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 13" href="http://scholar.google.com/scholar_lookup?&title=Earliest%20evidence%20of%20modern%20human%20life%20history%20in%20North%20African%20early%20Homo%20sapiens&journal=Proc.%20Natl%20Acad.%20Sci.%20USA&volume=104&pages=6128-6133&publication_year=2007&author=Smith%2CTM"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="14"><p class="c-article-references__text" id="ref-CR14">Gonder, M. K., Mortensen, H. M., Reed, F. A., de Sousa, A. & Tishkoff, S. A. Whole-mtDNA genome sequence analysis of ancient African lineages. <i>Mol. Biol. Evol.</i> <b>24</b>, 757–768 (2007)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="cas reference" data-track-action="cas reference" href="/articles/cas-redirect/1:CAS:528:DC%2BD2sXjtlers7w%3D" aria-label="CAS reference 14">CAS</a> <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed reference" data-track-action="pubmed reference" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=17194802" aria-label="PubMed reference 14">PubMed</a> <a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 14" href="http://scholar.google.com/scholar_lookup?&title=Whole-mtDNA%20genome%20sequence%20analysis%20of%20ancient%20African%20lineages&journal=Mol.%20Biol.%20Evol.&volume=24&pages=757-768&publication_year=2007&author=Gonder%2CMK&author=Mortensen%2CHM&author=Reed%2CFA&author=de%20Sousa%2CA&author=Tishkoff%2CSA"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="15"><p class="c-article-references__text" id="ref-CR15">McDougall, I., Brown, F. H. & Fleagle, J. G. Stratigraphic placement and age of modern humans from Kibish, Ethiopia. <i>Nature</i> <b>433</b>, 733–736 (2005)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="ads reference" data-track-action="ads reference" href="http://adsabs.harvard.edu/cgi-bin/nph-data_query?link_type=ABSTRACT&bibcode=2005Natur.433..733M" aria-label="ADS reference 15">ADS</a> <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="cas reference" data-track-action="cas reference" href="/articles/cas-redirect/1:CAS:528:DC%2BD2MXhtleqs7k%3D" aria-label="CAS reference 15">CAS</a> <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed reference" data-track-action="pubmed reference" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=15716951" aria-label="PubMed reference 15">PubMed</a> <a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 15" href="http://scholar.google.com/scholar_lookup?&title=Stratigraphic%20placement%20and%20age%20of%20modern%20humans%20from%20Kibish%2C%20Ethiopia&journal=Nature&volume=433&pages=733-736&publication_year=2005&author=McDougall%2CI&author=Brown%2CFH&author=Fleagle%2CJG"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="16"><p class="c-article-references__text" id="ref-CR16">White, T. D. et al. Pleistocene <i>Homo sapiens</i> from Middle Awash, Ethiopia. <i>Nature</i> <b>423</b>, 742–747 (2003)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="ads reference" data-track-action="ads reference" href="http://adsabs.harvard.edu/cgi-bin/nph-data_query?link_type=ABSTRACT&bibcode=2003Natur.423..742W" aria-label="ADS reference 16">ADS</a> <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="cas reference" data-track-action="cas reference" href="/articles/cas-redirect/1:CAS:528:DC%2BD3sXksV2itrs%3D" aria-label="CAS reference 16">CAS</a> <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed reference" data-track-action="pubmed reference" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=12802332" aria-label="PubMed reference 16">PubMed</a> <a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 16" href="http://scholar.google.com/scholar_lookup?&title=Pleistocene%20Homo%20sapiens%20from%20Middle%20Awash%2C%20Ethiopia&journal=Nature&volume=423&pages=742-747&publication_year=2003&author=White%2CTD"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="17"><p class="c-article-references__text" id="ref-CR17">Smith, F. H. in <i>Continuity or Replacement, Controversies in Homo sapiens evolution</i> (eds Bräuer, G . & Smith, F. H. ) 145–156 (A. A. Balkema, Zagreb, 1992)</p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="18"><p class="c-article-references__text" id="ref-CR18">Bruner, E. & Pearson, O. Neurocranial evolution in modern humans: the case of Jebel Irhoud 1. <i>Anthropol. Sci.</i> <b>121</b>, 31–41 (2013)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 18" href="http://scholar.google.com/scholar_lookup?&title=Neurocranial%20evolution%20in%20modern%20humans%3A%20the%20case%20of%20Jebel%20Irhoud%201&journal=Anthropol.%20Sci.&volume=121&pages=31-41&publication_year=2013&author=Bruner%2CE&author=Pearson%2CO"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="19"><p class="c-article-references__text" id="ref-CR19">Bermúdez de Castro, J. M. & Martinón-Torres, M. Evolutionary interpretation of the modern human-like facial morphology of the Atapuerca Gran Dolina-TD6 hominins. <i>Anthropol. Sci.</i> <b>122</b>, 149–155 (2014)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 19" href="http://scholar.google.com/scholar_lookup?&title=Evolutionary%20interpretation%20of%20the%20modern%20human-like%20facial%20morphology%20of%20the%20Atapuerca%20Gran%20Dolina-TD6%20hominins&journal=Anthropol.%20Sci.&volume=122&pages=149-155&publication_year=2014&author=Berm%C3%BAdez%20de%20Castro%2CJM&author=Martin%C3%B3n-Torres%2CM"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="20"><p class="c-article-references__text" id="ref-CR20">Schwartz, J. H. & Tattersall, I. The human chin revisited: what is it and who has it? <i>J. Hum. Evol.</i> <b>38</b>, 367–409 (2000)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="cas reference" data-track-action="cas reference" href="/articles/cas-redirect/1:STN:280:DC%2BD3c7kvFamsw%3D%3D" aria-label="CAS reference 20">CAS</a> <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed reference" data-track-action="pubmed reference" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=10683306" aria-label="PubMed reference 20">PubMed</a> <a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 20" href="http://scholar.google.com/scholar_lookup?&title=The%20human%20chin%20revisited%3A%20what%20is%20it%20and%20who%20has%20it%3F&journal=J.%20Hum.%20Evol.&volume=38&pages=367-409&publication_year=2000&author=Schwartz%2CJH&author=Tattersall%2CI"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="21"><p class="c-article-references__text" id="ref-CR21">Trinkaus, E. Dental remains from the Shanidar adult Neanderthals. <i>J. Hum. Evol.</i> <b>7</b>, 369–382 (1978)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 21" href="http://scholar.google.com/scholar_lookup?&title=Dental%20remains%20from%20the%20Shanidar%20adult%20Neanderthals&journal=J.%20Hum.%20Evol.&volume=7&pages=369-382&publication_year=1978&author=Trinkaus%2CE"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="22"><p class="c-article-references__text" id="ref-CR22">Gunz, P. et al. A uniquely modern human pattern of endocranial development. Insights from a new cranial reconstruction of the Neandertal newborn from Mezmaiskaya. <i>J. Hum. Evol.</i> <b>62</b>, 300–313 (2012)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed reference" data-track-action="pubmed reference" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=22221766" aria-label="PubMed reference 22">PubMed</a> <a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 22" href="http://scholar.google.com/scholar_lookup?&title=A%20uniquely%20modern%20human%20pattern%20of%20endocranial%20development.%20Insights%20from%20a%20new%20cranial%20reconstruction%20of%20the%20Neandertal%20newborn%20from%20Mezmaiskaya&journal=J.%20Hum.%20Evol.&volume=62&pages=300-313&publication_year=2012&author=Gunz%2CP"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="23"><p class="c-article-references__text" id="ref-CR23">Klein, R. G. <i>The Human Career: Human Biological and Cultural Origins</i> 3rd edn (Chicago Univ. Press, 2009)</p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="24"><p class="c-article-references__text" id="ref-CR24">Grün, R. et al. Direct dating of Florisbad hominid. <i>Nature</i> <b>382</b>, 500–501 (1996)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="ads reference" data-track-action="ads reference" href="http://adsabs.harvard.edu/cgi-bin/nph-data_query?link_type=ABSTRACT&bibcode=1996Natur.382..500G" aria-label="ADS reference 24">ADS</a> <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed reference" data-track-action="pubmed reference" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=8700221" aria-label="PubMed reference 24">PubMed</a> <a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 24" href="http://scholar.google.com/scholar_lookup?&title=Direct%20dating%20of%20Florisbad%20hominid&journal=Nature&volume=382&pages=500-501&publication_year=1996&author=Gr%C3%BCn%2CR"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="25"><p class="c-article-references__text" id="ref-CR25">Stringer, C. <i>The Origin of Our Species.</i> (Allen Lane, Penguin, 2011)</p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="26"><p class="c-article-references__text" id="ref-CR26">Harvati, K . et al. The later Stone Age calvaria from Iwo Eleru, Nigeria: morphology and chronology. <i>PLoS ONE</i> <b>6</b>, e24024 (2011)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="ads reference" data-track-action="ads reference" href="http://adsabs.harvard.edu/cgi-bin/nph-data_query?link_type=ABSTRACT&bibcode=2011PLoSO...624024H" aria-label="ADS reference 26">ADS</a> <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="cas reference" data-track-action="cas reference" href="/articles/cas-redirect/1:CAS:528:DC%2BC3MXht1OmsrjO" aria-label="CAS reference 26">CAS</a> <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed reference" data-track-action="pubmed reference" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=21949689" aria-label="PubMed reference 26">PubMed</a> <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed central reference" data-track-action="pubmed central reference" href="http://www.ncbi.nlm.nih.gov/pmc/articles/PMC3174138" aria-label="PubMed Central reference 26">PubMed Central</a> <a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 26" href="http://scholar.google.com/scholar_lookup?&title=The%20later%20Stone%20Age%20calvaria%20from%20Iwo%20Eleru%2C%20Nigeria%3A%20morphology%20and%20chronology&journal=PLoS%20ONE&volume=6&publication_year=2011&author=Harvati%2CK"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="27"><p class="c-article-references__text" id="ref-CR27">Gunz, P. et al. Early modern human diversity suggests subdivided population structure and a complex out-of-Africa scenario. <i>Proc. Natl Acad. Sci. USA</i> <b>106</b>, 6094–6098 (2009)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="ads reference" data-track-action="ads reference" href="http://adsabs.harvard.edu/cgi-bin/nph-data_query?link_type=ABSTRACT&bibcode=2009PNAS..106.6094G" aria-label="ADS reference 27">ADS</a> <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="cas reference" data-track-action="cas reference" href="/articles/cas-redirect/1:CAS:528:DC%2BD1MXlsFCrsbY%3D" aria-label="CAS reference 27">CAS</a> <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed reference" data-track-action="pubmed reference" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=19307568" aria-label="PubMed reference 27">PubMed</a> <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed central reference" data-track-action="pubmed central reference" href="http://www.ncbi.nlm.nih.gov/pmc/articles/PMC2669363" aria-label="PubMed Central reference 27">PubMed Central</a> <a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 27" href="http://scholar.google.com/scholar_lookup?&title=Early%20modern%20human%20diversity%20suggests%20subdivided%20population%20structure%20and%20a%20complex%20out-of-Africa%20scenario&journal=Proc.%20Natl%20Acad.%20Sci.%20USA&volume=106&pages=6094-6098&publication_year=2009&author=Gunz%2CP"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="28"><p class="c-article-references__text" id="ref-CR28">Arsuaga, J. L. et al. Neandertal roots: cranial and chronological evidence from Sima de los Huesos. <i>Science</i> <b>344</b>, 1358–1363 (2014)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="ads reference" data-track-action="ads reference" href="http://adsabs.harvard.edu/cgi-bin/nph-data_query?link_type=ABSTRACT&bibcode=2014Sci...344.1358A" aria-label="ADS reference 28">ADS</a> <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="cas reference" data-track-action="cas reference" href="/articles/cas-redirect/1:CAS:528:DC%2BC2cXpslygsb4%3D" aria-label="CAS reference 28">CAS</a> <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed reference" data-track-action="pubmed reference" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=24948730" aria-label="PubMed reference 28">PubMed</a> <a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 28" href="http://scholar.google.com/scholar_lookup?&title=Neandertal%20roots%3A%20cranial%20and%20chronological%20evidence%20from%20Sima%20de%20los%20Huesos&journal=Science&volume=344&pages=1358-1363&publication_year=2014&author=Arsuaga%2CJL"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="29"><p class="c-article-references__text" id="ref-CR29">Meyer, M. et al. A high-coverage genome sequence from an archaic Denisovan individual. <i>Science</i> <b>338</b>, 222–226 (2012)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="ads reference" data-track-action="ads reference" href="http://adsabs.harvard.edu/cgi-bin/nph-data_query?link_type=ABSTRACT&bibcode=2012Sci...338..222M" aria-label="ADS reference 29">ADS</a> <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="cas reference" data-track-action="cas reference" href="/articles/cas-redirect/1:CAS:528:DC%2BC38XhsVKrtr3N" aria-label="CAS reference 29">CAS</a> <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed reference" data-track-action="pubmed reference" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=22936568" aria-label="PubMed reference 29">PubMed</a> <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed central reference" data-track-action="pubmed central reference" href="http://www.ncbi.nlm.nih.gov/pmc/articles/PMC3617501" aria-label="PubMed Central reference 29">PubMed Central</a> <a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 29" href="http://scholar.google.com/scholar_lookup?&title=A%20high-coverage%20genome%20sequence%20from%20an%20archaic%20Denisovan%20individual&journal=Science&volume=338&pages=222-226&publication_year=2012&author=Meyer%2CM"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="30"><p class="c-article-references__text" id="ref-CR30">Weaver, T. D. Did a discrete event 200,000–100,000 years ago produce modern humans? <i>J. Hum. Evol.</i> <b>63</b>, 121–126 (2012)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed reference" data-track-action="pubmed reference" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=22658331" aria-label="PubMed reference 30">PubMed</a> <a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 30" href="http://scholar.google.com/scholar_lookup?&title=Did%20a%20discrete%20event%20200%2C000%E2%80%93100%2C000%20years%20ago%20produce%20modern%20humans%3F&journal=J.%20Hum.%20Evol.&volume=63&pages=121-126&publication_year=2012&author=Weaver%2CTD"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="31"><p class="c-article-references__text" id="ref-CR31">Wollny, G. et al. MIA - a free and open source software for gray scale medical image analysis. <i>Source Code Biol. Med.</i> <b>8</b>, 20 (2013)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed reference" data-track-action="pubmed reference" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=24119305" aria-label="PubMed reference 31">PubMed</a> <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed central reference" data-track-action="pubmed central reference" href="http://www.ncbi.nlm.nih.gov/pmc/articles/PMC4015836" aria-label="PubMed Central reference 31">PubMed Central</a> <a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 31" href="http://scholar.google.com/scholar_lookup?&title=MIA%20-%20a%20free%20and%20open%20source%20software%20for%20gray%20scale%20medical%20image%20analysis&journal=Source%20Code%20Biol.%20Med.&volume=8&publication_year=2013&author=Wollny%2CG"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="32"><p class="c-article-references__text" id="ref-CR32">Bookstein, F. L. Landmark methods for forms without landmarks: morphometrics of group differences in outline shape. <i>Med. Image Anal.</i> <b>1</b>, 225–243 (1997)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="cas reference" data-track-action="cas reference" href="/articles/cas-redirect/1:STN:280:DyaK1M%2FptVGqtg%3D%3D" aria-label="CAS reference 32">CAS</a> <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed reference" data-track-action="pubmed reference" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=9873908" aria-label="PubMed reference 32">PubMed</a> <a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 32" href="http://scholar.google.com/scholar_lookup?&title=Landmark%20methods%20for%20forms%20without%20landmarks%3A%20morphometrics%20of%20group%20differences%20in%20outline%20shape&journal=Med.%20Image%20Anal.&volume=1&pages=225-243&publication_year=1997&author=Bookstein%2CFL"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="33"><p class="c-article-references__text" id="ref-CR33">Gunz, P., Mitteroecker, P. & Bookstein, F. L. in <i>Modern Morphometrics in Physical Anthropology.</i> (ed. Slice, D.E. ) 73–98 (Kluwer Academic/Plenum Publishers, 2005)</p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="34"><p class="c-article-references__text" id="ref-CR34">Gunz, P. & Mitteroecker, P. Semilandmarks: a method for quantifying curves and surfaces. <i>Hystrix</i> <b>24</b>, 103–109 (2013)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 34" href="http://scholar.google.com/scholar_lookup?&title=Semilandmarks%3A%20a%20method%20for%20quantifying%20curves%20and%20surfaces&journal=Hystrix&volume=24&pages=103-109&publication_year=2013&author=Gunz%2CP&author=Mitteroecker%2CP"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="35"><p class="c-article-references__text" id="ref-CR35">Wiley, D. F . et al. Evolutionary Morphing. In <i>16th IEEE Visualization Conference (VIS 2005)</i> <b>55</b>, 431–438 (American Journal of Physical Anthropology, 2005)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 35" href="http://scholar.google.com/scholar_lookup?&title=&journal=16th%20IEEE%20Visualization%20Conference%20%28VIS%202005%29&volume=55&pages=431-438&publication_year=2005&author=Wiley%2CDF"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="36"><p class="c-article-references__text" id="ref-CR36">Freidline, S. E., Gunz, P., Harvati, K. & Hublin, J.-J. Middle Pleistocene human facial morphology in an evolutionary and developmental context. <i>J. Hum. Evol.</i> <b>63</b>, 723–740 (2012)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed reference" data-track-action="pubmed reference" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=22981042" aria-label="PubMed reference 36">PubMed</a> <a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 36" href="http://scholar.google.com/scholar_lookup?&title=Middle%20Pleistocene%20human%20facial%20morphology%20in%20an%20evolutionary%20and%20developmental%20context&journal=J.%20Hum.%20Evol.&volume=63&pages=723-740&publication_year=2012&author=Freidline%2CSE&author=Gunz%2CP&author=Harvati%2CK&author=Hublin%2CJ-J"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="37"><p class="c-article-references__text" id="ref-CR37">Neubauer, S., Gunz, P. & Hublin, J.-J. The pattern of endocranial ontogenetic shape changes in humans. <i>J. Anat.</i> <b>215</b>, 240–255 (2009)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed reference" data-track-action="pubmed reference" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=19531085" aria-label="PubMed reference 37">PubMed</a> <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed central reference" data-track-action="pubmed central reference" href="http://www.ncbi.nlm.nih.gov/pmc/articles/PMC2750758" aria-label="PubMed Central reference 37">PubMed Central</a> <a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 37" href="http://scholar.google.com/scholar_lookup?&title=The%20pattern%20of%20endocranial%20ontogenetic%20shape%20changes%20in%20humans&journal=J.%20Anat.&volume=215&pages=240-255&publication_year=2009&author=Neubauer%2CS&author=Gunz%2CP&author=Hublin%2CJ-J"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="38"><p class="c-article-references__text" id="ref-CR38">Harvati, K., Gunz, P. & Grigorescu, D. Cioclovina (Romania): affinities of an early modern European. <i>J. Hum. Evol.</i> <b>53</b>, 732–746 (2007)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed reference" data-track-action="pubmed reference" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=18001819" aria-label="PubMed reference 38">PubMed</a> <a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 38" href="http://scholar.google.com/scholar_lookup?&title=Cioclovina%20%28Romania%29%3A%20affinities%20of%20an%20early%20modern%20European&journal=J.%20Hum.%20Evol.&volume=53&pages=732-746&publication_year=2007&author=Harvati%2CK&author=Gunz%2CP&author=Grigorescu%2CD"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="39"><p class="c-article-references__text" id="ref-CR39">Benazzi, S. et al. Early dispersal of modern humans in Europe and implications for Neanderthal behaviour. <i>Nature</i> <b>479</b>, 525–528 (2011)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="ads reference" data-track-action="ads reference" href="http://adsabs.harvard.edu/cgi-bin/nph-data_query?link_type=ABSTRACT&bibcode=2011Natur.479..525B" aria-label="ADS reference 39">ADS</a> <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="cas reference" data-track-action="cas reference" href="/articles/cas-redirect/1:CAS:528:DC%2BC3MXhsVSktbbP" aria-label="CAS reference 39">CAS</a> <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed reference" data-track-action="pubmed reference" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=22048311" aria-label="PubMed reference 39">PubMed</a> <a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 39" href="http://scholar.google.com/scholar_lookup?&title=Early%20dispersal%20of%20modern%20humans%20in%20Europe%20and%20implications%20for%20Neanderthal%20behaviour&journal=Nature&volume=479&pages=525-528&publication_year=2011&author=Benazzi%2CS"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="40"><p class="c-article-references__text" id="ref-CR40">Bailey, S. E., Benazzi, S. & Hublin, J.-J. Allometry, merism, and tooth shape of the upper deciduous M2 and permanent M1. <i>Am. J. Phys. Anthropol.</i> <b>154</b>, 104–114 (2014)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed reference" data-track-action="pubmed reference" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=24482249" aria-label="PubMed reference 40">PubMed</a> <a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 40" href="http://scholar.google.com/scholar_lookup?&title=Allometry%2C%20merism%2C%20and%20tooth%20shape%20of%20the%20upper%20deciduous%20M2%20and%20permanent%20M1&journal=Am.%20J.%20Phys.%20Anthropol.&volume=154&pages=104-114&publication_year=2014&author=Bailey%2CSE&author=Benazzi%2CS&author=Hublin%2CJ-J"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="41"><p class="c-article-references__text" id="ref-CR41">Benazzi, S. et al. Cervical and crown outline analysis of worn Neanderthal and modern human lower second deciduous molars. <i>Am. J. Phys. Anthropol.</i> <b>149</b>, 537–546 (2012)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed reference" data-track-action="pubmed reference" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=23077004" aria-label="PubMed reference 41">PubMed</a> <a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 41" href="http://scholar.google.com/scholar_lookup?&title=Cervical%20and%20crown%20outline%20analysis%20of%20worn%20Neanderthal%20and%20modern%20human%20lower%20second%20deciduous%20molars&journal=Am.%20J.%20Phys.%20Anthropol.&volume=149&pages=537-546&publication_year=2012&author=Benazzi%2CS"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="42"><p class="c-article-references__text" id="ref-CR42">Skinner, M. M., Gunz, P., Wood, B. A. & Hublin, J.-J. Enamel–dentine junction (EDJ) morphology distinguishes the lower molars of <i>Australopithecus africanus</i> and <i>Paranthropus robustus</i>. <i>J. Hum. Evol.</i> <b>55</b>, 979–988 (2008)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed reference" data-track-action="pubmed reference" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=18824253" aria-label="PubMed reference 42">PubMed</a> <a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 42" href="http://scholar.google.com/scholar_lookup?&title=Enamel%E2%80%93dentine%20junction%20%28EDJ%29%20morphology%20distinguishes%20the%20lower%20molars%20of%20Australopithecus%20africanus%20and%20Paranthropus%20robustus&journal=J.%20Hum.%20Evol.&volume=55&pages=979-988&publication_year=2008&author=Skinner%2CMM&author=Gunz%2CP&author=Wood%2CBA&author=Hublin%2CJ-J"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="43"><p class="c-article-references__text" id="ref-CR43">Skinner, M. M., Gunz, P., Wood, B. A., Boesch, C. & Hublin, J.-J. Discrimination of extant <i>Pan</i> species and subspecies using the enamel–dentine junction morphology of lower molars. <i>Am. J. Phys. Anthropol.</i> <b>140</b>, 234–243 (2009)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed reference" data-track-action="pubmed reference" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=19382140" aria-label="PubMed reference 43">PubMed</a> <a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 43" href="http://scholar.google.com/scholar_lookup?&title=Discrimination%20of%20extant%20Pan%20species%20and%20subspecies%20using%20the%20enamel%E2%80%93dentine%20junction%20morphology%20of%20lower%20molars&journal=Am.%20J.%20Phys.%20Anthropol.&volume=140&pages=234-243&publication_year=2009&author=Skinner%2CMM&author=Gunz%2CP&author=Wood%2CBA&author=Boesch%2CC&author=Hublin%2CJ-J"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="44"><p class="c-article-references__text" id="ref-CR44">Spoor, F. et al. Reconstructed <i>Homo habilis</i> type OH 7 suggests deep-rooted species diversity in early <i>Homo</i>. <i>Nature</i> <b>519</b>, 83–86 (2015)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="ads reference" data-track-action="ads reference" href="http://adsabs.harvard.edu/cgi-bin/nph-data_query?link_type=ABSTRACT&bibcode=2015Natur.519...83S" aria-label="ADS reference 44">ADS</a> <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="cas reference" data-track-action="cas reference" href="/articles/cas-redirect/1:CAS:528:DC%2BC2MXjvVGjsrs%3D" aria-label="CAS reference 44">CAS</a> <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed reference" data-track-action="pubmed reference" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=25739632" aria-label="PubMed reference 44">PubMed</a> <a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 44" href="http://scholar.google.com/scholar_lookup?&title=Reconstructed%20Homo%20habilis%20type%20OH%207%20suggests%20deep-rooted%20species%20diversity%20in%20early%20Homo&journal=Nature&volume=519&pages=83-86&publication_year=2015&author=Spoor%2CF"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="45"><p class="c-article-references__text" id="ref-CR45">Beucher, S . & Lantuéjoul, C. in <i>International Workshop on Image Processing: Real-Time Edge and Motion Detection</i> 2.1–2.12 (Rennes, France, 1979)</p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="46"><p class="c-article-references__text" id="ref-CR46">Le Cabec, A., Gunz, P., Kupczik, K., Braga, J. & Hublin, J.-J. Anterior tooth root morphology and size in Neanderthals: taxonomic and functional implications. <i>J. Hum. Evol.</i> <b>64</b>, 169–193 (2013)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed reference" data-track-action="pubmed reference" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=23266488" aria-label="PubMed reference 46">PubMed</a> <a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 46" href="http://scholar.google.com/scholar_lookup?&title=Anterior%20tooth%20root%20morphology%20and%20size%20in%20Neanderthals%3A%20taxonomic%20and%20functional%20implications&journal=J.%20Hum.%20Evol.&volume=64&pages=169-193&publication_year=2013&author=Le%20Cabec%2CA&author=Gunz%2CP&author=Kupczik%2CK&author=Braga%2CJ&author=Hublin%2CJ-J"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="47"><p class="c-article-references__text" id="ref-CR47">Mitteroecker, P. & Gunz, P. Advances in geometric morphometrics. <i>Evol. Biol.</i> <b>36</b>, 235–247 (2009)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 47" href="http://scholar.google.com/scholar_lookup?&title=Advances%20in%20geometric%20morphometrics&journal=Evol.%20Biol.&volume=36&pages=235-247&publication_year=2009&author=Mitteroecker%2CP&author=Gunz%2CP"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="48"><p class="c-article-references__text" id="ref-CR48">Gunz, P., Mitteroecker, P., Neubauer, S., Weber, G. W. & Bookstein, F. L. Principles for the virtual reconstruction of hominin crania. <i>J. Hum. Evol.</i> <b>57</b>, 48–62 (2009)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed reference" data-track-action="pubmed reference" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=19482335" aria-label="PubMed reference 48">PubMed</a> <a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 48" href="http://scholar.google.com/scholar_lookup?&title=Principles%20for%20the%20virtual%20reconstruction%20of%20hominin%20crania&journal=J.%20Hum.%20Evol.&volume=57&pages=48-62&publication_year=2009&author=Gunz%2CP&author=Mitteroecker%2CP&author=Neubauer%2CS&author=Weber%2CGW&author=Bookstein%2CFL"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="49"><p class="c-article-references__text" id="ref-CR49">Rohlf, F. J. & Slice, D. Extensions of the procrustes method for the optimal superimposition of landmarks. <i>Syst. Zool.</i> <b>39</b>, 40–59 (1990)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 49" href="http://scholar.google.com/scholar_lookup?&title=Extensions%20of%20the%20procrustes%20method%20for%20the%20optimal%20superimposition%20of%20landmarks&journal=Syst.%20Zool.&volume=39&pages=40-59&publication_year=1990&author=Rohlf%2CFJ&author=Slice%2CD"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="50"><p class="c-article-references__text" id="ref-CR50">Mitteroecker, P. & Bookstein, F. Linear discrimination, ordination, and the visualization of selection gradients in modern morphometrics. <i>Evol. Biol.</i> <b>38</b>, 100–114 (2011)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 50" href="http://scholar.google.com/scholar_lookup?&title=Linear%20discrimination%2C%20ordination%2C%20and%20the%20visualization%20of%20selection%20gradients%20in%20modern%20morphometrics&journal=Evol.%20Biol.&volume=38&pages=100-114&publication_year=2011&author=Mitteroecker%2CP&author=Bookstein%2CF"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="51"><p class="c-article-references__text" id="ref-CR51">R Development Core Team. R: a language and environment for statistical computing <a href="http://www.r-project.org" data-track="click_references" data-track-action="external reference" data-track-value="external reference" data-track-label="http://www.r-project.org">http://www.r-project.org</a>(R Foundation for Statistical Computing, Vienna, Austria, 2012)</p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="52"><p class="c-article-references__text" id="ref-CR52">Arambourg, C. & Biberson, P. The fossil human remains from the Paleolithic site of Sidi Abderrahman (Morocco). <i>Am. J. Phys. Anthropol.</i> <b>14</b>, 467–489 (1956)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="cas reference" data-track-action="cas reference" href="/articles/cas-redirect/1:STN:280:DyaG2s%2Fisl2quw%3D%3D" aria-label="CAS reference 52">CAS</a> <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed reference" data-track-action="pubmed reference" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=13381832" aria-label="PubMed reference 52">PubMed</a> <a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 52" href="http://scholar.google.com/scholar_lookup?&title=The%20fossil%20human%20remains%20from%20the%20Paleolithic%20site%20of%20Sidi%20Abderrahman%20%28Morocco%29&journal=Am.%20J.%20Phys.%20Anthropol.&volume=14&pages=467-489&publication_year=1956&author=Arambourg%2CC&author=Biberson%2CP"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="53"><p class="c-article-references__text" id="ref-CR53">Sausse, F. La mandibule atlanthropienne de la carrière Thomas I (Casablanca). <i>Anthropologie</i> <b>79</b>, 81–112 (1975)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 53" href="http://scholar.google.com/scholar_lookup?&title=La%20mandibule%20atlanthropienne%20de%20la%20carri%C3%A8re%20Thomas%20I%20%28Casablanca%29&journal=Anthropologie&volume=79&pages=81-112&publication_year=1975&author=Sausse%2CF"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="54"><p class="c-article-references__text" id="ref-CR54">Rightmire, G. P. Comparative studies of Late Pleistocene human remains from Klasies River Mouth, South Africa. <i>J. Hum. Evol.</i> <b>20</b>, 131–156 (1991)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 54" href="http://scholar.google.com/scholar_lookup?&title=Comparative%20studies%20of%20Late%20Pleistocene%20human%20remains%20from%20Klasies%20River%20Mouth%2C%20South%20Africa&journal=J.%20Hum.%20Evol.&volume=20&pages=131-156&publication_year=1991&author=Rightmire%2CGP"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="55"><p class="c-article-references__text" id="ref-CR55">Stewart, T. D. in <i>Annual Report of the Smithsonian Institution</i> 521–533 (US Government Printing Office, Washington, 1962)</p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="56"><p class="c-article-references__text" id="ref-CR56">Tillier, A.-M. in <i>Le Squelette moustérien de Kébara</i> <b>2</b> 97–112 (Centre National de la Recherche Scientifique, Paris, 1991)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 56" href="http://scholar.google.com/scholar_lookup?&title=&journal=Le%20Squelette%20moust%C3%A9rien%20de%20K%C3%A9bara&volume=2&pages=97-112&publication_year=1991&author=Tillier%2CA-M"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="57"><p class="c-article-references__text" id="ref-CR57">Stewart, T. D. The Neanderthal skeletal remains from Shanidar Cave, Iraq: a summary of findings to date. <i>Proc. Am. Phil. Soc.</i> <b>121</b>, 121–165 (1977)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 57" href="http://scholar.google.com/scholar_lookup?&title=The%20Neanderthal%20skeletal%20remains%20from%20Shanidar%20Cave%2C%20Iraq%3A%20a%20summary%20of%20findings%20to%20date.&journal=Proc.%20Am.%20Phil.%20Soc.&volume=121&pages=121-165&publication_year=1977&author=Stewart%2CTD"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="58"><p class="c-article-references__text" id="ref-CR58">Trinkaus, E. <i>The Shanidar Neanderthals</i> (Academic, 1983)</p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="59"><p class="c-article-references__text" id="ref-CR59">Leroi-Gourhan, A. <i>Étude des Restes Humains Fossiles provenant des Grottes d’Arcy-sur-Cure</i> (Masson et Cie, 1958)</p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="60"><p class="c-article-references__text" id="ref-CR60">Daura, J. et al. A Neandertal mandible from the Cova del Gegant (Sitges, Barcelona, Spain). <i>J. Hum. Evol.</i> <b>49</b>, 56–70 (2005)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="cas reference" data-track-action="cas reference" href="/articles/cas-redirect/1:STN:280:DC%2BD2MzjvFehsQ%3D%3D" aria-label="CAS reference 60">CAS</a> <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed reference" data-track-action="pubmed reference" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=15896823" aria-label="PubMed reference 60">PubMed</a> <a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 60" href="http://scholar.google.com/scholar_lookup?&title=A%20Neandertal%20mandible%20from%20the%20Cova%20del%20Gegant%20%28Sitges%2C%20Barcelona%2C%20Spain%29&journal=J.%20Hum.%20Evol.&volume=49&pages=56-70&publication_year=2005&author=Daura%2CJ"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="61"><p class="c-article-references__text" id="ref-CR61">Topinard, P. Les caracteres simiens de la machoire de la Naulette. <i>Rev. Antropol.</i> <b>15</b>, 385–431 (1886)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 61" href="http://scholar.google.com/scholar_lookup?&title=Les%20caracteres%20simiens%20de%20la%20machoire%20de%20la%20Naulette.&journal=Rev.%20Antropol.&volume=15&pages=385-431&publication_year=1886&author=Topinard%2CP"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="62"><p class="c-article-references__text" id="ref-CR62">Blake, C. C. On a human jaw from the cave of La Naulette, near Dinant, Belgium. <i>Anthropol. Rev.</i> <b>5</b>, 294–303 (1867)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 62" href="http://scholar.google.com/scholar_lookup?&title=On%20a%20human%20jaw%20from%20the%20cave%20of%20La%20Naulette%2C%20near%20Dinant%2C%20Belgium&journal=Anthropol.%20Rev.&volume=5&pages=294-303&publication_year=1867&author=Blake%2CCC"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="63"><p class="c-article-references__text" id="ref-CR63">Leguebe, A. & Toussaint, M. <i>La Mandibule et les Cubitus de la Naulette: Morphologie et Morphométrie</i> 15 (Editions du Centre National de la Recherche Scientifique, 1988)</p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="64"><p class="c-article-references__text" id="ref-CR64">Heim, J. L. Les hommes fossiles de La Ferrassie (Dordogne) et le problem de la definition des Neandertaliens classiques. III. Squelette céphalique. <i>Anthropologie</i> <b>78</b>, 321–378 (1974)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 64" href="http://scholar.google.com/scholar_lookup?&title=Les%20hommes%20fossiles%20de%20La%20Ferrassie%20%28Dordogne%29%20et%20le%20problem%20de%20la%20definition%20des%20Neandertaliens%20classiques.%20III.%20Squelette%20c%C3%A9phalique&journal=Anthropologie&volume=78&pages=321-378&publication_year=1974&author=Heim%2CJL"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="65"><p class="c-article-references__text" id="ref-CR65">de Lumley, M.-A. Les Néandertaliens de la grotte de l’Hortus. <i>Etudes Quaternaires</i> <b>1</b>, 375–385 (1972)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 65" href="http://scholar.google.com/scholar_lookup?&title=Les%20N%C3%A9andertaliens%20de%20la%20grotte%20de%20l%E2%80%99Hortus.&journal=Etudes%20Quaternaires&volume=1&pages=375-385&publication_year=1972&author=de%20Lumley%2CM-A"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="66"><p class="c-article-references__text" id="ref-CR66">Condemi, S. et al. Possible interbreeding in late Italian Neanderthals? New data from the Mezzena jaw (Monti Lessini, Verona, Italy). <i>PLoS ONE</i> <b>8</b>, e59781 (2013)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="ads reference" data-track-action="ads reference" href="http://adsabs.harvard.edu/cgi-bin/nph-data_query?link_type=ABSTRACT&bibcode=2013PLoSO...859781C" aria-label="ADS reference 66">ADS</a> <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="cas reference" data-track-action="cas reference" href="/articles/cas-redirect/1:CAS:528:DC%2BC3sXlslCmtrg%3D" aria-label="CAS reference 66">CAS</a> <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed reference" data-track-action="pubmed reference" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=23544098" aria-label="PubMed reference 66">PubMed</a> <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed central reference" data-track-action="pubmed central reference" href="http://www.ncbi.nlm.nih.gov/pmc/articles/PMC3609795" aria-label="PubMed Central reference 66">PubMed Central</a> <a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 66" href="http://scholar.google.com/scholar_lookup?&title=Possible%20interbreeding%20in%20late%20Italian%20Neanderthals%3F%20New%20data%20from%20the%20Mezzena%20jaw%20%28Monti%20Lessini%2C%20Verona%2C%20Italy%29&journal=PLoS%20ONE&volume=8&publication_year=2013&author=Condemi%2CS"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="67"><p class="c-article-references__text" id="ref-CR67">Corrain, C. Resti scheletrici umani del ‘Riparo Mezzena’. <i>Memorie del Museo civico di Storia naturale di Verona</i> <b>16</b>, 97–101 (1968)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 67" href="http://scholar.google.com/scholar_lookup?&title=Resti%20scheletrici%20umani%20del%20%E2%80%98Riparo%20Mezzena%E2%80%99&journal=Memorie%20del%20Museo%20civico%20di%20Storia%20naturale%20di%20Verona&volume=16&pages=97-101&publication_year=1968&author=Corrain%2CC"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="68"><p class="c-article-references__text" id="ref-CR68">Walker, M. J., Lombardi, A. V., Zapata, J. & Trinkaus, E. Neandertal mandibles from the Sima de las Palomas del Cabezo Gordo, Murcia, southeastern Spain. <i>Am. J. Phys. Anthropol.</i> <b>142</b>, 261–272 (2010)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed reference" data-track-action="pubmed reference" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=20014182" aria-label="PubMed reference 68">PubMed</a> <a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 68" href="http://scholar.google.com/scholar_lookup?&title=Neandertal%20mandibles%20from%20the%20Sima%20de%20las%20Palomas%20del%20Cabezo%20Gordo%2C%20Murcia%2C%20southeastern%20Spain&journal=Am.%20J.%20Phys.%20Anthropol.&volume=142&pages=261-272&publication_year=2010&author=Walker%2CMJ&author=Lombardi%2CAV&author=Zapata%2CJ&author=Trinkaus%2CE"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="69"><p class="c-article-references__text" id="ref-CR69">Martin, H. Machoire humaine moustérienne trouvée dans la station de La Quina. <i>L’homme préhistorique</i> <b>13</b>, 3–21 (1926)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 69" href="http://scholar.google.com/scholar_lookup?&title=Machoire%20humaine%20moust%C3%A9rienne%20trouv%C3%A9e%20dans%20la%20station%20de%20La%20Quina&journal=L%E2%80%99homme%20pr%C3%A9historique&volume=13&pages=3-21&publication_year=1926&author=Martin%2CH"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="70"><p class="c-article-references__text" id="ref-CR70">Martin, H. Position stratigraphique des Ossements humains recueillis dans le Moustérien de La Quina de 1908 à 1912. <i>Bull. Soc. Préhistorique</i> <b>9</b>, 700–709 (1912)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 70" href="http://scholar.google.com/scholar_lookup?&title=Position%20stratigraphique%20des%20Ossements%20humains%20recueillis%20dans%20le%20Moust%C3%A9rien%20de%20La%20Quina%20de%201908%20%C3%A0%201912&journal=Bull.%20Soc.%20Pr%C3%A9historique&volume=9&pages=700-709&publication_year=1912&author=Martin%2CH"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="71"><p class="c-article-references__text" id="ref-CR71">Martin, H. <i>L’Homme fossile de la Quina.</i> (Libraire Octave Doin, 1923)</p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="72"><p class="c-article-references__text" id="ref-CR72">Pap, I., Tillier, A. M., Arensburg, B. & Chech, M. The Subalyuk Neanderthal remains (Hungary): a re-examination. <i>Ann. Hist. Nat. Mus. Natl. Hung.</i> <b>88</b>, 233–270 (1996)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 72" href="http://scholar.google.com/scholar_lookup?&title=The%20Subalyuk%20Neanderthal%20remains%20%28Hungary%29%3A%20a%20re-examination&journal=Ann.%20Hist.%20Nat.%20Mus.%20Natl.%20Hung.&volume=88&pages=233-270&publication_year=1996&author=Pap%2CI&author=Tillier%2CAM&author=Arensburg%2CB&author=Chech%2CM"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="73"><p class="c-article-references__text" id="ref-CR73">Sanchez, F. Comparative biometrical study of the Mousterian mandible from Cueva del Boquete de Zafarraya (Málaga, Spain). <i>Hum. Evol.</i> <b>14</b>, 125–138 (1999)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 73" href="http://scholar.google.com/scholar_lookup?&title=Comparative%20biometrical%20study%20of%20the%20Mousterian%20mandible%20from%20Cueva%20del%20Boquete%20de%20Zafarraya%20%28M%C3%A1laga%2C%20Spain%29&journal=Hum.%20Evol.&volume=14&pages=125-138&publication_year=1999&author=Sanchez%2CF"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="74"><p class="c-article-references__text" id="ref-CR74">Vlcˇek, E. <i>Fossile Menschenfunde von Weimar-Ehringsdorf, Weimarer Monographien zur Ur- und Frühgeschichte</i> Vol. <b>30</b> (Landesamt für Archäologische Denkmalpflege, 1993)</p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="75"><p class="c-article-references__text" id="ref-CR75">Condemi, S. <i>Les néandertaliens de La Chaise: abri Bourgeois-Delaunay. Comité des travaux historiques et scientifiques</i> (CTHS, 2001)</p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="76"><p class="c-article-references__text" id="ref-CR76">Bar-Yosef, O . & Vandermeersch, B . (eds) <i>Le squelette moustérien de Kébara</i> <b>2</b>. (Editions du Centre National de la Recherche Scientifique, 1991)</p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="77"><p class="c-article-references__text" id="ref-CR77">Arambourg, C . & Hoffstetter, R. <i>Le gisement de Ternifine</i> Vol. <b>1</b>. (Masson, 1963)</p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="78"><p class="c-article-references__text" id="ref-CR78">Hooton, E.A ., Hencken, H.O & Snow, Ch. E. <i>The ancient Palestinian: Skhu¯ l V reconstruction.</i> <b>17</b>, 5–10 (American School of Prehistoric Research, 1953)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 78" href="http://scholar.google.com/scholar_lookup?&title=&journal=The%20ancient%20Palestinian%3A%20Skhu%C2%AF%E2%80%89l%20V%20reconstruction.&volume=17&pages=5-10&publication_year=1953&author=Hooton%2CEA&author=Hencken%2CHO&author=Snow%2CChE"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="79"><p class="c-article-references__text" id="ref-CR79">Sollas, W. J. The Chancelade skull. <i>J. R. Anthropol. Inst.</i> <b>57</b>, 89–122 (1927)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 79" href="http://scholar.google.com/scholar_lookup?&title=The%20Chancelade%20skull&journal=J.%20R.%20Anthropol.%20Inst.&volume=57&pages=89-122&publication_year=1927&author=Sollas%2CWJ"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="80"><p class="c-article-references__text" id="ref-CR80">Martin, H. Caractères des squelettes humains quaternaires de la vallée du Roc (Charente). <i>Bull. Mem. Soc. Anthropol. Paris</i> <b>8</b>, 103–129 (1927)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 80" href="http://scholar.google.com/scholar_lookup?&title=Caract%C3%A8res%20des%20squelettes%20humains%20quaternaires%20de%20la%20vall%C3%A9e%20du%20Roc%20%28Charente%29&journal=Bull.%20Mem.%20Soc.%20Anthropol.%20Paris&volume=8&pages=103-129&publication_year=1927&author=Martin%2CH"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="81"><p class="c-article-references__text" id="ref-CR81">Vercellotti, G., Alciati, G., Richards, M. P. & Formicola, V. The Late Upper Paleolithic skeleton Villabruna 1 (Italy): a source of data on biology and behavior of a 14,000 year-old hunter. <i>J. Anthropol. Sci.</i> <b>86</b>, 143–163 (2008)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed reference" data-track-action="pubmed reference" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=19934473" aria-label="PubMed reference 81">PubMed</a> <a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 81" href="http://scholar.google.com/scholar_lookup?&title=The%20Late%20Upper%20Paleolithic%20skeleton%20Villabruna%201%20%28Italy%29%3A%20a%20source%20of%20data%20on%20biology%20and%20behavior%20of%20a%2014%2C000%20year-old%20hunter&journal=J.%20Anthropol.%20Sci.&volume=86&pages=143-163&publication_year=2008&author=Vercellotti%2CG&author=Alciati%2CG&author=Richards%2CMP&author=Formicola%2CV"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="82"><p class="c-article-references__text" id="ref-CR82">Formicola, V. Una mandibola umana dal deposito dell’Epigravettiano finale delle Arene Candite (scavi 197O). <i>Rev. Antropol.</i> <b>64</b>, 271–278 (1986)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 82" href="http://scholar.google.com/scholar_lookup?&title=Una%20mandibola%20umana%20dal%20deposito%20dell%E2%80%99Epigravettiano%20finale%20delle%20Arene%20Candite%20%28scavi%20197O%29.&journal=Rev.%20Antropol.&volume=64&pages=271-278&publication_year=1986&author=Formicola%2CV"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="83"><p class="c-article-references__text" id="ref-CR83">Odano, A. M. & Riquet, R. Le gisement préhistorique de Dar-es-Soltane 2. Champ de tir de El Menzeh à Rabat (Maroc). Note préliminaire. 2-Étude anthropologique des restes post-atériens. <i>Bull. d’Archéologie Marocaine</i> <b>11</b>, 25–63 (1978)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 83" href="http://scholar.google.com/scholar_lookup?&title=Le%20gisement%20pr%C3%A9historique%20de%20Dar-es-Soltane%202.%20Champ%20de%20tir%20de%20El%20Menzeh%20%C3%A0%20Rabat%20%28Maroc%29.%20Note%20pr%C3%A9liminaire.%202-%C3%89tude%20anthropologique%20des%20restes%20post-at%C3%A9riens&journal=Bull.%20d%E2%80%99Arch%C3%A9ologie%20Marocaine&volume=11&pages=25-63&publication_year=1978&author=Odano%2CAM&author=Riquet%2CR"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="84"><p class="c-article-references__text" id="ref-CR84">Debénath, A. Nouveaux restes humains atériens du Maroc. <i>CR Acad. Sci. Paris</i> <b>290</b>, 851–852 (1980)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 84" href="http://scholar.google.com/scholar_lookup?&title=Nouveaux%20restes%20humains%20at%C3%A9riens%20du%20Maroc.&journal=CR%20Acad.%20Sci.%20Paris&volume=290&pages=851-852&publication_year=1980&author=Deb%C3%A9nath%2CA"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="85"><p class="c-article-references__text" id="ref-CR85">Crognier, E. & Dupouy-Madre, M. Les Natoufiens du Nahal Oren (Ouadi Fallah) Etude anthropologique. <i>Paéorient</i> <b>2</b>, 103–121 (1974)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 85" href="http://scholar.google.com/scholar_lookup?&title=Les%20Natoufiens%20du%20Nahal%20Oren%20%28Ouadi%20Fallah%29%20Etude%20anthropologique&journal=Pa%C3%A9orient&volume=2&pages=103-121&publication_year=1974&author=Crognier%2CE&author=Dupouy-Madre%2CM"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="86"><p class="c-article-references__text" id="ref-CR86">Henke, W. Vergleichend-morphologische Kennzeichnung der Jungpaläolithiker von Oberkassel bei Bonn. <i>Z. Morphol. Anthropol.</i> <b>75</b>, 27–44 (1984)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="cas reference" data-track-action="cas reference" href="/articles/cas-redirect/1:STN:280:DyaL2M%2Fpt1yqsw%3D%3D" aria-label="CAS reference 86">CAS</a> <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed reference" data-track-action="pubmed reference" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=6516528" aria-label="PubMed reference 86">PubMed</a> <a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 86" href="http://scholar.google.com/scholar_lookup?&title=Vergleichend-morphologische%20Kennzeichnung%20der%20Jungpal%C3%A4olithiker%20von%20Oberkassel%20bei%20Bonn.&journal=Z.%20Morphol.%20Anthropol.&volume=75&pages=27-44&publication_year=1984&author=Henke%2CW"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="87"><p class="c-article-references__text" id="ref-CR87">Hershkovitz, I. et al. Ohalo II H2: a 19,000-year-old skeleton from a water-logged site at the Sea of Galilee, Israel. <i>Am. J. Phys. Anthropol.</i> <b>96</b>, 215–234 (1995)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="cas reference" data-track-action="cas reference" href="/articles/cas-redirect/1:STN:280:DyaK2MzgsFamtQ%3D%3D" aria-label="CAS reference 87">CAS</a> <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed reference" data-track-action="pubmed reference" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=7785722" aria-label="PubMed reference 87">PubMed</a> <a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 87" href="http://scholar.google.com/scholar_lookup?&title=Ohalo%20II%20H2%3A%20a%2019%2C000-year-old%20skeleton%20from%20a%20water-logged%20site%20at%20the%20Sea%20of%20Galilee%2C%20Israel&journal=Am.%20J.%20Phys.%20Anthropol.&volume=96&pages=215-234&publication_year=1995&author=Hershkovitz%2CI"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="88"><p class="c-article-references__text" id="ref-CR88">Soficaru, A., Dobos, A. & Trinkaus, E. Early modern humans from the Pestera Muierii, Baia de Fier, Romania. <i>Proc. Natl Acad. Sci. USA</i> <b>103</b>, 17196–17201 (2006)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="ads reference" data-track-action="ads reference" href="http://adsabs.harvard.edu/cgi-bin/nph-data_query?link_type=ABSTRACT&bibcode=2006PNAS..10317196S" aria-label="ADS reference 88">ADS</a> <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="cas reference" data-track-action="cas reference" href="/articles/cas-redirect/1:CAS:528:DC%2BD28Xht1Kqs7vE" aria-label="CAS reference 88">CAS</a> <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed reference" data-track-action="pubmed reference" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=17085588" aria-label="PubMed reference 88">PubMed</a> <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed central reference" data-track-action="pubmed central reference" href="http://www.ncbi.nlm.nih.gov/pmc/articles/PMC1859909" aria-label="PubMed Central reference 88">PubMed Central</a> <a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 88" href="http://scholar.google.com/scholar_lookup?&title=Early%20modern%20humans%20from%20the%20Pestera%20Muierii%2C%20Baia%20de%20Fier%2C%20Romania&journal=Proc.%20Natl%20Acad.%20Sci.%20USA&volume=103&pages=17196-17201&publication_year=2006&author=Soficaru%2CA&author=Dobos%2CA&author=Trinkaus%2CE"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="89"><p class="c-article-references__text" id="ref-CR89">Crevecoeur, I. Etude anthropologique des restes humains de Nazlet Khater (Paléolithique Superieur, Egypte). <i>PhD Thesis, Université Sciences et Technologies Bordeaux I</i> (2006)</p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="90"><p class="c-article-references__text" id="ref-CR90">Thoma, A. Morphology and affinities of the Nazlet Khater man. <i>J. Hum. Evol.</i> <b>13</b>, 287–296 (1984)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 90" href="http://scholar.google.com/scholar_lookup?&title=Morphology%20and%20affinities%20of%20the%20Nazlet%20Khater%20man&journal=J.%20Hum.%20Evol.&volume=13&pages=287-296&publication_year=1984&author=Thoma%2CA"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="91"><p class="c-article-references__text" id="ref-CR91">Anderson, J. E. In <i>The Prehistory of Nubia</i> Vol. <b>2</b> (ed. Wendorf, F. ) 996–1040 (Southern Methodist Univ. Press,1968)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 91" href="http://scholar.google.com/scholar_lookup?&title=&journal=The%20Prehistory%20of%20Nubia&volume=2&pages=996-1040&publication_year=1968&author=Anderson%2CJE&author=Wendorf%2CF"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="92"><p class="c-article-references__text" id="ref-CR92">Crevecoeur, I. From the Nile to the Danube: a comparison of the Nazlet Khater 2 and Oase 1 early modern human mandibles. <i>Anthropologie</i> <b>42</b>, 203–213 (2004)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 92" href="http://scholar.google.com/scholar_lookup?&title=From%20the%20Nile%20to%20the%20Danube%3A%20a%20comparison%20of%20the%20Nazlet%20Khater%202%20and%20Oase%201%20early%20modern%20human%20mandibles&journal=Anthropologie&volume=42&pages=203-213&publication_year=2004&author=Crevecoeur%2CI"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="93"><p class="c-article-references__text" id="ref-CR93">Trinkaus, E. et al. An early modern human from the Pes¸ tera cu Oase, Romania. <i>Proc. Natl Acad. Sci. USA</i> <b>100</b>, 11231–11236 (2003)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="ads reference" data-track-action="ads reference" href="http://adsabs.harvard.edu/cgi-bin/nph-data_query?link_type=ABSTRACT&bibcode=2003PNAS..10011231T" aria-label="ADS reference 93">ADS</a> <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="cas reference" data-track-action="cas reference" href="/articles/cas-redirect/1:CAS:528:DC%2BD3sXotFKnu78%3D" aria-label="CAS reference 93">CAS</a> <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed reference" data-track-action="pubmed reference" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=14504393" aria-label="PubMed reference 93">PubMed</a> <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed central reference" data-track-action="pubmed central reference" href="http://www.ncbi.nlm.nih.gov/pmc/articles/PMC208740" aria-label="PubMed Central reference 93">PubMed Central</a> <a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 93" href="http://scholar.google.com/scholar_lookup?&title=An%20early%20modern%20human%20from%20the%20Pes%C2%B8%E2%80%89tera%20cu%20Oase%2C%20Romania&journal=Proc.%20Natl%20Acad.%20Sci.%20USA&volume=100&pages=11231-11236&publication_year=2003&author=Trinkaus%2CE"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="94"><p class="c-article-references__text" id="ref-CR94">Trinkaus, E. & Svoboda, J. (eds) <i>Early Modern Human Evolution in Central Europe. The People of Dolní Veˇ stonice and Pavlov. The Dolni Vestonice Studies</i> Vol. <b>12</b> (Oxford Univ. Press, 2006)</p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="95"><p class="c-article-references__text" id="ref-CR95">Sládek, V ., Trinkaus, E ., Hillson, S. W . & Holliday, T. W. The People of the Pavlovian. Skeletal Catalogue and Osteometrics of the Gravettian Fossil Hominids from Dolni Vestonice and Pavlov. <i>The Dolni Vestonice Studies</i> Vol. <b>5</b> 1–244 (BRNO, 2000)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 95" href="http://scholar.google.com/scholar_lookup?&title=The%20People%20of%20the%20Pavlovian.%20Skeletal%20Catalogue%20and%20Osteometrics%20of%20the%20Gravettian%20Fossil%20Hominids%20from%20Dolni%20Vestonice%20and%20Pavlov.&journal=The%20Dolni%20Vestonice%20Studies&volume=5&pages=1-244&publication_year=2000&author=Sl%C3%A1dek%2CV&author=Trinkaus%2CE&author=Hillson%2CSW&author=Holliday%2CTW"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="96"><p class="c-article-references__text" id="ref-CR96">Drozdová, E. The evaluation of a rediscovered fragment of human lower jaw, No 21 from Prˇedmostí u Prˇerova. <i>Archeologické rozhledy</i> <b>53</b>, 452–460 (2001)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 96" href="http://scholar.google.com/scholar_lookup?&title=The%20evaluation%20of%20a%20rediscovered%20fragment%20of%20human%20lower%20jaw%2C%20No%2021%20from%20Pr%CB%87edmost%C3%AD%20u%20Pr%CB%87erova.&journal=Archeologick%C3%A9%20rozhledy&volume=53&pages=452-460&publication_year=2001&author=Drozdov%C3%A1%2CE"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="97"><p class="c-article-references__text" id="ref-CR97">Dutour, O. Palimpseste paléoanthropologique sur l’«Homme fossile d’Asselar» (Sahara). <i>Travaux du Laboratoire d’Anthropologie et de Préhistoire des Pays de la Méditerranée Occidentale</i> <b>1</b>, 73–83 (1992)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 97" href="http://scholar.google.com/scholar_lookup?&title=Palimpseste%20pal%C3%A9oanthropologique%20sur%20l%E2%80%99%C2%ABHomme%20fossile%20d%E2%80%99Asselar%C2%BB%20%28Sahara%29&journal=Travaux%20du%20Laboratoire%20d%E2%80%99Anthropologie%20et%20de%20Pr%C3%A9histoire%20des%20Pays%20de%20la%20M%C3%A9diterran%C3%A9e%20Occidentale&volume=1&pages=73-83&publication_year=1992&author=Dutour%2CO"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="98"><p class="c-article-references__text" id="ref-CR98">Gambier, D. Vestiges humains du gisement d’Isturitz (Pyrénées—Atlantiques): étude anthropologique et analyse des traces d’action humaine intentionelle. <i>Antiquités nationales</i> <b>22–23</b>, 9–26 (1990)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 98" href="http://scholar.google.com/scholar_lookup?&title=Vestiges%20humains%20du%20gisement%20d%E2%80%99Isturitz%20%28Pyr%C3%A9n%C3%A9es%E2%80%94Atlantiques%29%3A%20%C3%A9tude%20anthropologique%20et%20analyse%20des%20traces%20d%E2%80%99action%20humaine%20intentionelle&journal=Antiquit%C3%A9s%20nationales&volume=22%E2%80%9323&pages=9-26&publication_year=1990&author=Gambier%2CD"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="99"><p class="c-article-references__text" id="ref-CR99">Bräuer, G. & Mehlman, M. J. Hominid molars from a middle Stone Age level at the Mumba Rock Shelter, Tanzania. <i>Am. J. Phys. Anthropol.</i> <b>75</b>, 69–76 (1988)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed reference" data-track-action="pubmed reference" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=3124633" aria-label="PubMed reference 99">PubMed</a> <a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 99" href="http://scholar.google.com/scholar_lookup?&title=Hominid%20molars%20from%20a%20middle%20Stone%20Age%20level%20at%20the%20Mumba%20Rock%20Shelter%2C%20Tanzania&journal=Am.%20J.%20Phys.%20Anthropol.&volume=75&pages=69-76&publication_year=1988&author=Br%C3%A4uer%2CG&author=Mehlman%2CMJ"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="100"><p class="c-article-references__text" id="ref-CR100">Protsch, R. R. R. <i>Di Archäologischen und Anthropologischen Ergebnisse der Kohl-Larsen-Expeditionen in Nord-Tanzania</i> 1933–1939. (Institut für Urgeschichte der Universität Tübingen, 1981)</p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="101"><p class="c-article-references__text" id="ref-CR101">Wood, B. A. & Van Noten, F. L. Preliminary observations on the BK 8518 mandible from Baringo, Kenya. <i>Am. J. Phys. Anthropol.</i> <b>69</b>, 117–127 (1986)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="cas reference" data-track-action="cas reference" href="/articles/cas-redirect/1:STN:280:DyaL287islKktg%3D%3D" aria-label="CAS reference 101">CAS</a> <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed reference" data-track-action="pubmed reference" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=3080897" aria-label="PubMed reference 101">PubMed</a> <a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 101" href="http://scholar.google.com/scholar_lookup?&title=Preliminary%20observations%20on%20the%20BK%208518%20mandible%20from%20Baringo%2C%20Kenya&journal=Am.%20J.%20Phys.%20Anthropol.&volume=69&pages=117-127&publication_year=1986&author=Wood%2CBA&author=Van%20Noten%2CFL"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="102"><p class="c-article-references__text" id="ref-CR102">Rightmire, G. P. Middle Pleistocene hominids from Olduvai Gorge, northern Tanzania. <i>Am. J. Phys. Anthropol.</i> <b>53</b>, 225–241 (1980)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="cas reference" data-track-action="cas reference" href="/articles/cas-redirect/1:STN:280:DyaL3M%2FhvFaltg%3D%3D" aria-label="CAS reference 102">CAS</a> <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed reference" data-track-action="pubmed reference" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=6774618" aria-label="PubMed reference 102">PubMed</a> <a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 102" href="http://scholar.google.com/scholar_lookup?&title=Middle%20Pleistocene%20hominids%20from%20Olduvai%20Gorge%2C%20northern%20Tanzania&journal=Am.%20J.%20Phys.%20Anthropol.&volume=53&pages=225-241&publication_year=1980&author=Rightmire%2CGP"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="103"><p class="c-article-references__text" id="ref-CR103">Bermúdez de Castro, J. M. Dental remains from Atapuerca (Spain) I. Metrics. <i>J. Hum. Evol.</i> <b>15</b>, 265–287 (1986)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 103" href="http://scholar.google.com/scholar_lookup?&title=Dental%20remains%20from%20Atapuerca%20%28Spain%29%20I.%20Metrics.&journal=J.%20Hum.%20Evol.&volume=15&pages=265-287&publication_year=1986&author=Berm%C3%BAdez%20de%20Castro%2CJM"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="104"><p class="c-article-references__text" id="ref-CR104">Grine, F. E. & Franzen, J. L. Fossil hominid teeth from the Sangiran Dome (Java, Indonesia). <i>Courier Forschungsinstitut Senckenberg</i> <b>171</b>, 75–103 (1994)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 104" href="http://scholar.google.com/scholar_lookup?&title=Fossil%20hominid%20teeth%20from%20the%20Sangiran%20Dome%20%28Java%2C%20Indonesia%29&journal=Courier%20Forschungsinstitut%20Senckenberg&volume=171&pages=75-103&publication_year=1994&author=Grine%2CFE&author=Franzen%2CJL"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="105"><p class="c-article-references__text" id="ref-CR105">Le Cabec, A., Gunz, P., Kupczik, K., Braga, J. & Hublin, J.-J. Anterior tooth root morphology and size in Neanderthals: taxonomic and functional implications. <i>J. Hum. Evol.</i> <b>64</b>, 169–193 (2013)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed reference" data-track-action="pubmed reference" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=23266488" aria-label="PubMed reference 105">PubMed</a> <a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 105" href="http://scholar.google.com/scholar_lookup?&title=Anterior%20tooth%20root%20morphology%20and%20size%20in%20Neanderthals%3A%20taxonomic%20and%20functional%20implications&journal=J.%20Hum.%20Evol.&volume=64&pages=169-193&publication_year=2013&author=Le%20Cabec%2CA&author=Gunz%2CP&author=Kupczik%2CK&author=Braga%2CJ&author=Hublin%2CJ-J"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="106"><p class="c-article-references__text" id="ref-CR106">Turner, C. G. II, Nichol, C. R. & Scott, G. R. <i>Advances in Dental Anthropology</i> (eds Kelley, M., & Larsen, C. ) 13–31 (Wiley Liss, 1991)</p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="107"><p class="c-article-references__text" id="ref-CR107">Bailey, S. E. Neandertal Dental Morphology: Implications for modern human origins. <i>PhD thesis, Arizona State Univ.</i> (2002)</p></li></ol><p class="c-article-references__download u-hide-print"><a data-track="click" data-track-action="download citation references" data-track-label="link" rel="nofollow" href="https://citation-needed.springer.com/v2/references/10.1038/nature22336?format=refman&flavour=references">Download references<svg width="16" height="16" focusable="false" role="img" aria-hidden="true" class="u-icon"><use xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="#icon-eds-i-download-medium"></use></svg></a></p></div></div></div></section></div><section data-title="Acknowledgements"><div class="c-article-section" id="Ack1-section"><h2 class="c-article-section__title js-section-title js-c-reading-companion-sections-item" id="Ack1">Acknowledgements</h2><div class="c-article-section__content" id="Ack1-content"><p>The research program at Jebel Irhoud is jointly conducted and supported by the Moroccan Institut National des Sciences de l’Archéologie et du Patrimoine and the Department of Human Evolution of the Max Planck Institute for Evolutionary Anthropology. We are grateful to the many curators and colleagues who, over the years, gave us access to recent and fossil hominin specimens for computed tomography scanning or analysis, to E. Trinkaus for providing comparative data and to C. Kiarie, M. Lui, C. Piot, D. Plotzki, A. Buchenau and H. Temming for their technical assistance.</p></div></div></section><section aria-labelledby="author-information" data-title="Author information"><div class="c-article-section" id="author-information-section"><h2 class="c-article-section__title js-section-title js-c-reading-companion-sections-item" id="author-information">Author information</h2><div class="c-article-section__content" id="author-information-content"><h3 class="c-article__sub-heading" id="affiliations">Authors and Affiliations</h3><ol class="c-article-author-affiliation__list"><li id="Aff1"><p class="c-article-author-affiliation__address">Department of Human Evolution, Max Planck Institute for Evolutionary Anthropology, Deutscher Platz 6, Leipzig, 04103, Germany</p><p class="c-article-author-affiliation__authors-list">Jean-Jacques Hublin, Sarah E. Freidline, Simon Neubauer, Inga Bergmann, Adeline Le Cabec & Philipp Gunz</p></li><li id="Aff2"><p class="c-article-author-affiliation__address">Chaire Internationale de Paléoanthropologie, Collège de France, Paris, France</p><p class="c-article-author-affiliation__authors-list">Jean-Jacques Hublin</p></li><li id="Aff3"><p class="c-article-author-affiliation__address">Institut National des Sciences de l’Archéologie et du Patrimoine, Rabat, Morocco</p><p class="c-article-author-affiliation__authors-list">Abdelouahed Ben-Ncer</p></li><li id="Aff4"><p class="c-article-author-affiliation__address">Department of Anthropology, Center for the Study of Human Origins, New York University, New York, 10003, New York, USA</p><p class="c-article-author-affiliation__authors-list">Shara E. Bailey</p></li><li id="Aff5"><p class="c-article-author-affiliation__address">School of Anthropology and Conservation, University of Kent, Canterbury, CT2 7NR, UK</p><p class="c-article-author-affiliation__authors-list">Matthew M. Skinner</p></li><li id="Aff6"><p class="c-article-author-affiliation__address">Department of Cultural Heritage, University of Bologna, Ravenna, 48121, Italy</p><p class="c-article-author-affiliation__authors-list">Stefano Benazzi</p></li><li id="Aff7"><p class="c-article-author-affiliation__address">Paleoanthropology, Senckenberg Center for Human Evolution and Paleoenvironment, and DFG Center for Advanced Studies: “Words, Bones, Genes, Tools”, Eberhard Karls Universität, Tübingen, Germany</p><p class="c-article-author-affiliation__authors-list">Katerina Harvati</p></li></ol><div class="u-js-hide u-hide-print" data-test="author-info"><span class="c-article__sub-heading">Authors</span><ol class="c-article-authors-search u-list-reset"><li id="auth-Jean_Jacques-Hublin-Aff1-Aff2"><span class="c-article-authors-search__title u-h3 js-search-name">Jean-Jacques Hublin</span><div class="c-article-authors-search__list"><div class="c-article-authors-search__item c-article-authors-search__list-item--left"><a href="/search?author=Jean-Jacques%20Hublin" class="c-article-button" data-track="click" data-track-action="author link - publication" data-track-label="link" rel="nofollow">View author publications</a></div><div class="c-article-authors-search__item c-article-authors-search__list-item--right"><p class="search-in-title-js c-article-authors-search__text">You can also search for this author in <span class="c-article-identifiers"><a class="c-article-identifiers__item" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=search&term=Jean-Jacques%20Hublin" data-track="click" data-track-action="author link - pubmed" data-track-label="link" rel="nofollow">PubMed</a><span class="u-hide"> </span><a class="c-article-identifiers__item" href="http://scholar.google.co.uk/scholar?as_q=&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_sauthors=%22Jean-Jacques%20Hublin%22&as_publication=&as_ylo=&as_yhi=&as_allsubj=all&hl=en" data-track="click" data-track-action="author link - scholar" data-track-label="link" rel="nofollow">Google Scholar</a></span></p></div></div></li><li id="auth-Abdelouahed-Ben_Ncer-Aff3"><span class="c-article-authors-search__title u-h3 js-search-name">Abdelouahed Ben-Ncer</span><div class="c-article-authors-search__list"><div class="c-article-authors-search__item c-article-authors-search__list-item--left"><a href="/search?author=Abdelouahed%20Ben-Ncer" class="c-article-button" data-track="click" data-track-action="author link - publication" data-track-label="link" rel="nofollow">View author publications</a></div><div class="c-article-authors-search__item c-article-authors-search__list-item--right"><p class="search-in-title-js c-article-authors-search__text">You can also search for this author in <span class="c-article-identifiers"><a class="c-article-identifiers__item" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=search&term=Abdelouahed%20Ben-Ncer" data-track="click" data-track-action="author link - pubmed" data-track-label="link" rel="nofollow">PubMed</a><span class="u-hide"> </span><a class="c-article-identifiers__item" href="http://scholar.google.co.uk/scholar?as_q=&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_sauthors=%22Abdelouahed%20Ben-Ncer%22&as_publication=&as_ylo=&as_yhi=&as_allsubj=all&hl=en" data-track="click" data-track-action="author link - scholar" data-track-label="link" rel="nofollow">Google Scholar</a></span></p></div></div></li><li id="auth-Shara_E_-Bailey-Aff4"><span class="c-article-authors-search__title u-h3 js-search-name">Shara E. 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scholar" data-track-label="link" rel="nofollow">Google Scholar</a></span></p></div></div></li></ol></div><h3 class="c-article__sub-heading" id="contributions">Contributions</h3><p>The study was conceived by J.-J.H., A.B.-N. and P.G. Cranial metrical and non-metrical data were compiled and analysed by J.-J.H., A.B.-N., S.E.F., S.N., K.H. and P.G. Mandibular metrical and non-metrical data were compiled and analysed by J.-J.H. and I.B. Dental metrical and non-metrical data were compiled and analysed by S.E.B., M.M.S., A.L.C. and S.B. J.-J.H. and P.G. wrote the manuscript with contributions from all other authors.</p><h3 class="c-article__sub-heading" id="corresponding-author">Corresponding authors</h3><p id="corresponding-author-list">Correspondence to <a id="corresp-c1" href="mailto:hublin@eva.mpg.de">Jean-Jacques Hublin</a> or <a id="corresp-c2" href="mailto:gunz@eva.mpg.de">Philipp Gunz</a>.</p></div></div></section><section data-title="Ethics declarations"><div class="c-article-section" id="ethics-section"><h2 class="c-article-section__title js-section-title js-c-reading-companion-sections-item" id="ethics">Ethics declarations</h2><div class="c-article-section__content" id="ethics-content"> <h3 class="c-article__sub-heading">Competing interests</h3> <p>The authors declare no competing financial interests.</p> </div></div></section><section data-title="Additional information"><div class="c-article-section" id="additional-information-section"><h2 class="c-article-section__title js-section-title js-c-reading-companion-sections-item" id="additional-information">Additional information</h2><div class="c-article-section__content" id="additional-information-content"><p><b>Reviewer Information</b> <i>Nature</i> thanks R. G. Klein, C. Stringer and the other anonymous reviewer(s) for their contribution to the peer review of this work.</p><p>Publisher's note: Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.</p></div></div></section><section data-title="Extended data figures and tables"><div class="c-article-section" id="Sec14-section"><h2 class="c-article-section__title js-section-title js-c-reading-companion-sections-item" id="Sec14">Extended data figures and tables</h2><div class="c-article-section__content" id="Sec14-content"><div data-test="supplementary-info"><div id="figshareContainer" class="c-article-figshare-container" data-test="figshare-container"></div><div class="c-article-supplementary__item js-c-reading-companion-figures-item" data-test="supp-item" id="Fig4"><h3 class="c-article-supplementary__title u-h3"><a class="print-link" data-track="click" data-track-action="view supplementary info" data-test="supp-info-link" data-track-label="extended data figure 1 mandibular morphology." href="/articles/nature22336/figures/4" data-supp-info-image="//media.springernature.com/lw685/springer-static/image/art%3A10.1038%2Fnature22336/MediaObjects/41586_2017_Article_BFnature22336_Fig4_ESM.jpg">Extended Data Figure 1 Mandibular morphology.</a></h3><div class="c-article-supplementary__description" data-component="thumbnail-container"><p><b>a</b>, Symphyseal section of the Irhoud 11 mandible showing the mental angle. <b>b</b>, Mental area of Irhoud 11 before virtual reconstruction (top) and Irhoud 3 (bottom). Both images are surface models generated from micro-computed tomography data. <b>c</b>, Bivariate plot of mandibular corpus breadth versus height at the mental foramen. Irhoud 11 (pink star) falls within the EMH distribution and has one of the largest corpus heights among Middle to Late Pleistocene hominins. Values are in mm. <i>n</i> indicates sample size. Data sources and sample compositions can be found in the Methods. Scale bar, 20 mm.</p></div></div><div class="c-article-supplementary__item js-c-reading-companion-figures-item" data-test="supp-item" id="Fig5"><h3 class="c-article-supplementary__title u-h3"><a class="print-link" data-track="click" data-track-action="view supplementary info" data-test="supp-info-link" data-track-label="extended data figure 2 dental morphology." href="/articles/nature22336/figures/5" data-supp-info-image="//media.springernature.com/lw685/springer-static/image/art%3A10.1038%2Fnature22336/MediaObjects/41586_2017_Article_BFnature22336_Fig5_ESM.jpg">Extended Data Figure 2 Dental morphology.</a></h3><div class="c-article-supplementary__description" data-component="thumbnail-container"><p><b>a</b>, Shape–space PCA plot of Late Early and Middle Pleistocene archaic <i>Homo</i>, Neanderthals and RMH M<sup>1</sup> crown outlines. The deformed mean crown outlines in the four directions of the PCs are drawn at the extremity of each axis. Sample compositions and abbreviations can be found in the Methods. <b>b</b>, EDJ morphology of the M<sub>2</sub> and P<sub>4</sub>. Top left, the PCA analysis of the EDJ shape of the M<sub>2</sub> places Irhoud 11 intermediate between <i>H. erectus</i> and RMH (along with other north Africa fossil humans) and distinct from Neanderthals. Surface models illustrate EDJ shape changes along PC1 (bottom left) and PC2 (top right); the former separating <i>H. erectus</i> from RMH, Neanderthals and north African EMH and the latter separating Neanderthals from RMH and north African EMH. Bottom right, a PCA analysis of the EDJ shape of the P<sub>4</sub> groups Irhoud 11 with modern and fossil humans.</p></div></div><div class="c-article-supplementary__item js-c-reading-companion-figures-item" data-test="supp-item" id="Fig6"><h3 class="c-article-supplementary__title u-h3"><a class="print-link" data-track="click" data-track-action="view supplementary info" data-test="supp-info-link" data-track-label="extended data figure 3 shape analysis of i2 roots." href="/articles/nature22336/figures/6" data-supp-info-image="//media.springernature.com/lw685/springer-static/image/art%3A10.1038%2Fnature22336/MediaObjects/41586_2017_Article_BFnature22336_Fig6_ESM.jpg">Extended Data Figure 3 Shape analysis of I<sub>2</sub> roots.</a></h3><div class="c-article-supplementary__description" data-component="thumbnail-container"><p>A between-group PCA shows a complete separation between Neanderthals and a worldwide sample of recent modern humans based on subtle shape differences. Irhoud 11 (pink star) plots at the fringes of RMH, close to the EMH from Contrebandiers 1 (Tem). Colour-coded Procrustes group mean shapes are plotted in the same orientation as the I<sub>2</sub> root surface of Irhoud 11. Although Irhoud 11 is more similar, overall, to Neanderthals in terms of root size, its root shape is clearly modern. The <i>H. erectus</i> specimen KNM-WT 15000 and hypothetical EMH Tabun C2 have incisor root shapes similar to Neanderthals, suggesting that roots that are labially more convex than in RMH represent a conserved primitive condition with limited taxonomical value. Sample compositions and abbreviations can be found in the Methods.</p></div></div><div class="c-article-supplementary__item js-c-reading-companion-figures-item" data-test="supp-item" id="Fig7"><h3 class="c-article-supplementary__title u-h3"><a class="print-link" data-track="click" data-track-action="view supplementary info" data-test="supp-info-link" data-track-label="extended data figure 4 shape analysis of the exter" href="/articles/nature22336/figures/7" data-supp-info-image="//media.springernature.com/lw685/springer-static/image/art%3A10.1038%2Fnature22336/MediaObjects/41586_2017_Article_BFnature22336_Fig7_ESM.jpg">Extended Data Figure 4 Shape analysis of the external vault.</a></h3><div class="c-article-supplementary__description" data-component="thumbnail-container"><p><b>a</b>, PC scores of PC1 and PC2 of external braincase shape in <i>H. erectus</i>, archaic Middle Pleistocene <i>Homo</i>, geographically diverse RMH and Neanderthals. Results are consistent with the analysis of endocranial shape (<a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/nature22336#Fig3">Fig. 3a</a>). However, several EMH and Upper Palaeolithic specimens fall outside the RMH variation. This is probably owing to the projecting supraorbital tori in these specimens. <b>b</b>, Shape changes associated with PC1 (two standard deviations in either direction) shown as thin-plate spline deformation grids in lateral and oblique view. PC1 captures a contrast between elongated braincases with projecting supraorbital tori (low scores, in black) and a more globular braincase with gracile supraorbital tori (high scores, in red). Sample compositions and abbreviations can be found in the Methods.</p></div></div><div class="c-article-supplementary__item js-c-reading-companion-figures-item" data-test="supp-item" id="Fig8"><h3 class="c-article-supplementary__title u-h3"><a class="print-link" data-track="click" data-track-action="view supplementary info" data-test="supp-info-link" data-track-label="extended data figure 5 facial and endocranial shap" href="/articles/nature22336/figures/8" data-supp-info-image="//media.springernature.com/lw685/springer-static/image/art%3A10.1038%2Fnature22336/MediaObjects/41586_2017_Article_BFnature22336_Fig8_ESM.jpg">Extended Data Figure 5 Facial and endocranial shape differences among <i>Homo</i> groups.</a></h3><div class="c-article-supplementary__description" data-component="thumbnail-container"><p>Visualizations of GMM shape analyses in <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/nature22336#Fig3">Fig. 3</a>. <b>a</b>, Average endocranial shape differences between <i>H. erectus</i>, recent <i>H. sapiens</i> and Neanderthals. Thin-plate spline grids are exaggerated. <b>b</b>, Visualization of shape changes along PC1 in <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/nature22336#Fig3">Fig. 3b</a> in frontal, lateral and superior view; two standard deviations in either direction from the mean shape (grey, negative; black, positive). <b>c</b>, Shape changes along PC2. All recent and fossil modern humans (low scores along PC2) share smaller, orthognathic faces, that differ from the larger, robust and prognathic faces of the Middle Pleistocene humans and Neanderthals (high scores along PC1). Arrow length is colour-coded (short, blue; long, red). As these visualizations are affected by the Procrustes superimposition, we also show grids for the maxilla and the supraorbital area. The arrow points to the plane of the maxillary thin-plate spline (red) in the template configuration.</p></div></div><div class="c-article-supplementary__item" data-test="supp-item"><div class="c-article-table" data-test="inline-table" data-container-section="table" id="table-1"><figure><figcaption class="c-article-table__figcaption"><b id="Tab1" data-test="table-caption">Extended Data Table 1 List of hominin specimens</b></figcaption><div class="u-text-right u-hide-print"><a class="c-article__pill-button" data-test="table-link" data-track="click" data-track-action="view table" data-track-label="button" rel="nofollow" href="/articles/nature22336/tables/1" aria-label="Full size table 1"><span>Full size table</span><svg width="16" height="16" focusable="false" role="img" aria-hidden="true" class="u-icon"><use xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="#icon-eds-i-chevron-right-small"></use></svg></a></div></figure></div></div><div class="c-article-supplementary__item" data-test="supp-item"><div class="c-article-table" data-test="inline-table" data-container-section="table" id="table-2"><figure><figcaption class="c-article-table__figcaption"><b id="Tab2" data-test="table-caption">Extended Data Table 2 Measurements of the Irhoud 11 mandible after reconstruction</b></figcaption><div class="u-text-right u-hide-print"><a class="c-article__pill-button" data-test="table-link" data-track="click" data-track-action="view table" data-track-label="button" rel="nofollow" href="/articles/nature22336/tables/2" aria-label="Full size table 2"><span>Full size table</span><svg width="16" height="16" focusable="false" role="img" aria-hidden="true" class="u-icon"><use xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="#icon-eds-i-chevron-right-small"></use></svg></a></div></figure></div></div><div class="c-article-supplementary__item" data-test="supp-item"><div class="c-article-table" data-test="inline-table" data-container-section="table" id="table-3"><figure><figcaption class="c-article-table__figcaption"><b id="Tab3" data-test="table-caption">Extended Data Table 3 Dental measurements (upper dentition)</b></figcaption><div class="u-text-right u-hide-print"><a class="c-article__pill-button" data-test="table-link" data-track="click" data-track-action="view table" data-track-label="button" rel="nofollow" href="/articles/nature22336/tables/3" aria-label="Full size table 3"><span>Full size table</span><svg width="16" height="16" focusable="false" role="img" aria-hidden="true" class="u-icon"><use xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="#icon-eds-i-chevron-right-small"></use></svg></a></div></figure></div></div><div class="c-article-supplementary__item" data-test="supp-item"><div class="c-article-table" data-test="inline-table" data-container-section="table" id="table-4"><figure><figcaption class="c-article-table__figcaption"><b id="Tab4" data-test="table-caption">Extended Data Table 4 Dental measurements (lower dentition)</b></figcaption><div class="u-text-right u-hide-print"><a class="c-article__pill-button" data-test="table-link" data-track="click" data-track-action="view table" data-track-label="button" rel="nofollow" href="/articles/nature22336/tables/4" aria-label="Full size table 4"><span>Full size table</span><svg width="16" height="16" focusable="false" role="img" aria-hidden="true" class="u-icon"><use xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="#icon-eds-i-chevron-right-small"></use></svg></a></div></figure></div></div><div class="c-article-supplementary__item" data-test="supp-item"><div class="c-article-table" data-test="inline-table" data-container-section="table" id="table-5"><figure><figcaption class="c-article-table__figcaption"><b id="Tab5" data-test="table-caption">Extended Data Table 5 Morphological dental trait comparison</b></figcaption><div class="u-text-right u-hide-print"><a class="c-article__pill-button" data-test="table-link" data-track="click" data-track-action="view table" data-track-label="button" rel="nofollow" href="/articles/nature22336/tables/5" aria-label="Full size table 5"><span>Full size table</span><svg width="16" height="16" focusable="false" role="img" aria-hidden="true" class="u-icon"><use xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="#icon-eds-i-chevron-right-small"></use></svg></a></div></figure></div></div></div></div></div></section><section data-title="Related audio"><div class="c-article-section" id="Sec15-section"><h2 class="c-article-section__title js-section-title js-c-reading-companion-sections-item" id="Sec15">Related audio</h2><div class="c-article-section__content" id="Sec15-content"><div data-test="supplementary-info"><div class="c-article-supplementary__item" data-test="supp-item" id="MOESM4"><h3 class="c-article-supplementary__title u-h3"><a class="print-link" data-track="click" data-track-action="view supplementary info" data-test="supp-info-link" data-track-label="kerri smith finds out how a cave in morocco is cha" href="https://static-content.springer.com/esm/art%3A10.1038%2Fnature22336/MediaObjects/41586_2017_BFnature22336_MOESM4_ESM.mp3" data-supp-info-image="">Kerri Smith finds out how a cave in Morocco is changing our understanding of human history</a></h3></div></div></div></div></section><section data-title="PowerPoint slides"><div class="c-article-section" id="Sec16-section"><h2 class="c-article-section__title js-section-title js-c-reading-companion-sections-item" id="Sec16">PowerPoint slides</h2><div class="c-article-section__content" id="Sec16-content"><div data-test="supplementary-info"><div class="c-article-supplementary__item" data-test="supp-item" id="MOESM1"><h3 class="c-article-supplementary__title u-h3"><a class="print-link" data-track="click" data-track-action="view supplementary info" data-test="supp-info-link" data-track-label="powerpoint slide for fig. 1" 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id="citeas">Cite this article</h3><p class="c-bibliographic-information__citation">Hublin, JJ., Ben-Ncer, A., Bailey, S. <i>et al.</i> New fossils from Jebel Irhoud, Morocco and the pan-African origin of <i>Homo sapiens</i>. <i>Nature</i> <b>546</b>, 289–292 (2017). https://doi.org/10.1038/nature22336</p><p class="c-bibliographic-information__download-citation u-hide-print"><a data-test="citation-link" data-track="click" data-track-action="download article citation" data-track-label="link" data-track-external="" rel="nofollow" href="https://citation-needed.springer.com/v2/references/10.1038/nature22336?format=refman&flavour=citation">Download citation<svg width="16" height="16" focusable="false" role="img" aria-hidden="true" class="u-icon"><use xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="#icon-eds-i-download-medium"></use></svg></a></p><ul class="c-bibliographic-information__list" data-test="publication-history"><li 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Previous fossil evidence has placed the emergence of modern human biology in eastern Africa around 200,000 years ago. In this issue of <i>Nature</i>, Jean-Jaques Hublin and colleagues report new human fossils from Jebel Irhoud, Morocco; their work is accompanied by a separate report on the dating of the fossils by Shannon McPherron and colleagues. Together they report remains dating back 300,000–350,000 years. They identify numerous features, including a facial, mandibular and dental morphology, that align the material with early or recent modern humans. They also identified more primitive neurocranial and endocranial morphology. Collectively, the researchers believe that this mosaic of features displayed by the Jebel Irhoud hominins assigns them to the earliest evolutionary phase of <i>Homo sapiens</i>. Both papers suggest that the evolutionary processes behind the emergence of modern humans were not confined to sub-Saharan Africa.</p></div><button type="button" id="show-button" data-track="click" data-track-label="button" data-track-action="editorial summary show less" class="c-article-editorial-summary__button">show all</button></section></div> <div class="c-article-associated-content__container"> <section> <h2 class="c-article-associated-content__title u-mb-24">Associated content</h2> <div class="c-article-associated-content__collection collection u-mb-24"> <section> <p class="c-article-associated-content__collection-label u-sans-serif u-text-bold u-mb-8">Collection</p> <h3 class="c-article-associated-content__collection-title u-h3 u-mb-8"> <a href="https://www.nature.com/collections/ffjxdjnlcq" class="u-link-inherit" data-track="click" data-track-action="view collection" data-track-category="associated content" data-track-label="collection" data-test="collection-link">Human Evolution</a> </h3> </section> </div> <div class="u-full-height u-mb-24"> <article class="u-full-height c-card c-card--flush"> <div class="c-card__layout u-full-height"> <div class="c-card__body"> <h3 class="c-card__title"> <a href="https://www.nature.com/articles/nature22335" class="c-card__link u-link-inherit" data-track="click" data-track-action="view article" data-track-category="associated content" data-track-label="research">The age of the hominin fossils from Jebel Irhoud, Morocco, and the origins of the Middle Stone Age</a> </h3> <ul data-test="author-list" class="c-author-list c-author-list--compact c-author-list--truncated"> <li>Daniel Richter</li><li>Rainer Grün</li><li>Shannon P. 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