1 Stringer, C. Modern human origins: progress and prospects. Phil. Trans. R. Soc. B 357, 563–579 (2002)

2 Bräuer, G. The origin of modern anatomy: by speciation or intraspecific evolution? Evol. Anthropol. 17, 22–37 (2008)

3 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)

4 Ennouchi, E. Le deuxième crâne de l’homme d’Irhoud. Annales de Paléontologie (Vértébrés) LIV, 117–128 (1968)

5 Hublin, J.-J. & Tillier, A. M. in Aspects of Human Evolution Vol. 21 Symposia of the Society for the study of Human Biology, Volume XXI (ed. Stringer, C. B. ) 167–186 (Taylor & Francis, 1981)

6 Hublin, J.-J., Tillier, A. M. & Tixier, J. L’humerus d’enfant moustérien (Homo 4) du Jebel Irhoud (Maroc) dans son contexte archéologique. Bull. Mem. Soc. Anthropol. Paris 4, 115–141 (1987)

7 Tixier, J ., Brugal, J.-P ., Tillier, A.-M ., Bruzeket, J . & Hublin, J.-J. in Actes des 1ères Journées Nationales d’Archéologie et du Patrimoine. Préhistoire Vol. 1 149–153 (Société Marocaine d’Archéologie et du Patrimoine, 2001)

8 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. Comptes Rendus à l'Académie des Sciences de Paris 316, 847–852 (1993)

9 Ennouchi, E. Un Néanderthalien: l’homme du Jebel Irhoud (Maroc). Anthropologie 66, 279–299 (1962)

10 Stringer, C. B in Recent advances in Primatology. (eds Chivers, D. J . & Joysey, K. A. ) 395–418 (Academic, 1978)

11 Hublin, J.-J. Recent human evolution in northwestern Africa. Phil. Trans. R. Soc. B 337, 185–191 (1992)

12 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. Quat. Res. 80, 552–561 (2013)

13 Smith, T. M. et al. Earliest evidence of modern human life history in North African early Homo sapiens. Proc. Natl Acad. Sci. USA 104, 6128–6133 (2007)

14 Gonder, M. K., Mortensen, H. M., Reed, F. A., de Sousa, A. & Tishkoff, S. A. Whole-mtDNA genome sequence analysis of ancient African lineages. Mol. Biol. Evol. 24, 757–768 (2007)

15 McDougall, I., Brown, F. H. & Fleagle, J. G. Stratigraphic placement and age of modern humans from Kibish, Ethiopia. Nature 433, 733–736 (2005)

16 White, T. D. et al. Pleistocene Homo sapiens from Middle Awash, Ethiopia. Nature 423, 742–747 (2003)

17 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)

18 Bruner, E. & Pearson, O. Neurocranial evolution in modern humans: the case of Jebel Irhoud 1. Anthropol. Sci. 121, 31–41 (2013)

19 Bermúdez de Castro, J. M. & Martinón-Torres, M. Evolutionary interpretation of the modern human-like facial morphology of the Atapuerca Gran Dolina-TD6 hominins. Anthropol. Sci. 122, 149–155 (2014)

20 Schwartz, J. H. & Tattersall, I. The human chin revisited: what is it and who has it? J. Hum. Evol. 38, 367–409 (2000)

21 Trinkaus, E. Dental remains from the Shanidar adult Neanderthals. J. Hum. Evol. 7, 369–382 (1978)

22 Gunz, P. et al. A uniquely modern human pattern of endocranial development. Insights from a new cranial reconstruction of the Neandertal newborn from Mezmaiskaya. J. Hum. Evol. 62, 300–313 (2012)

23 Klein, R. G. The Human Career: Human Biological and Cultural Origins 3rd edn (Chicago Univ. Press, 2009)

24 Grün, R. et al. Direct dating of Florisbad hominid. Nature 382, 500–501 (1996)

25 Stringer, C. The Origin of Our Species. (Allen Lane, Penguin, 2011)

26 Harvati, K . et al. The later Stone Age calvaria from Iwo Eleru, Nigeria: morphology and chronology. PLoS ONE 6, e24024 (2011)

27 Gunz, P. et al. Early modern human diversity suggests subdivided population structure and a complex out-of-Africa scenario. Proc. Natl Acad. Sci. USA 106, 6094–6098 (2009)

28 Arsuaga, J. L. et al. Neandertal roots: cranial and chronological evidence from Sima de los Huesos. Science 344, 1358–1363 (2014)

29 Meyer, M. et al. A high-coverage genome sequence from an archaic Denisovan individual. Science 338, 222–226 (2012)

30 Weaver, T. D. Did a discrete event 200,000–100,000 years ago produce modern humans? J. Hum. Evol. 63, 121–126 (2012)

31 Wollny, G. et al. MIA - a free and open source software for gray scale medical image analysis. Source Code Biol. Med. 8, 20 (2013)

32 Bookstein, F. L. Landmark methods for forms without landmarks: morphometrics of group differences in outline shape. Med. Image Anal. 1, 225–243 (1997)

33 Gunz, P., Mitteroecker, P. & Bookstein, F. L. in Modern Morphometrics in Physical Anthropology. (ed. Slice, D.E. ) 73–98 (Kluwer Academic/Plenum Publishers, 2005)

34 Gunz, P. & Mitteroecker, P. Semilandmarks: a method for quantifying curves and surfaces. Hystrix 24, 103–109 (2013)

35 Wiley, D. F . et al. Evolutionary Morphing. In 16th IEEE Visualization Conference (VIS 2005) 55, 431–438 (American Journal of Physical Anthropology, 2005)

36 Freidline, S. E., Gunz, P., Harvati, K. & Hublin, J.-J. Middle Pleistocene human facial morphology in an evolutionary and developmental context. J. Hum. Evol. 63, 723–740 (2012)

37 Neubauer, S., Gunz, P. & Hublin, J.-J. The pattern of endocranial ontogenetic shape changes in humans. J. Anat. 215, 240–255 (2009)

38 Harvati, K., Gunz, P. & Grigorescu, D. Cioclovina (Romania): affinities of an early modern European. J. Hum. Evol. 53, 732–746 (2007)

39 Benazzi, S. et al. Early dispersal of modern humans in Europe and implications for Neanderthal behaviour. Nature 479, 525–528 (2011)

40 Bailey, S. E., Benazzi, S. & Hublin, J.-J. Allometry, merism, and tooth shape of the upper deciduous M2 and permanent M1. Am. J. Phys. Anthropol. 154, 104–114 (2014)

41 Benazzi, S. et al. Cervical and crown outline analysis of worn Neanderthal and modern human lower second deciduous molars. Am. J. Phys. Anthropol. 149, 537–546 (2012)

42 Skinner, M. M., Gunz, P., Wood, B. A. & Hublin, J.-J. Enamel–dentine junction (EDJ) morphology distinguishes the lower molars of Australopithecus africanus and Paranthropus robustus. J. Hum. Evol. 55, 979–988 (2008)

43 Skinner, M. M., Gunz, P., Wood, B. A., Boesch, C. & Hublin, J.-J. Discrimination of extant Pan species and subspecies using the enamel–dentine junction morphology of lower molars. Am. J. Phys. Anthropol. 140, 234–243 (2009)

44 Spoor, F. et al. Reconstructed Homo habilis type OH 7 suggests deep-rooted species diversity in early Homo. Nature 519, 83–86 (2015)

45 Beucher, S . & Lantuéjoul, C. in International Workshop on Image Processing: Real-Time Edge and Motion Detection 2.1–2.12 (Rennes, France, 1979)

46 Le Cabec, A., Gunz, P., Kupczik, K., Braga, J. & Hublin, J.-J. Anterior tooth root morphology and size in Neanderthals: taxonomic and functional implications. J. Hum. Evol. 64, 169–193 (2013)

47 Mitteroecker, P. & Gunz, P. Advances in geometric morphometrics. Evol. Biol. 36, 235–247 (2009)

48 Gunz, P., Mitteroecker, P., Neubauer, S., Weber, G. W. & Bookstein, F. L. Principles for the virtual reconstruction of hominin crania. J. Hum. Evol. 57, 48–62 (2009)

49 Rohlf, F. J. & Slice, D. Extensions of the procrustes method for the optimal superimposition of landmarks. Syst. Zool. 39, 40–59 (1990)

50 Mitteroecker, P. & Bookstein, F. Linear discrimination, ordination, and the visualization of selection gradients in modern morphometrics. Evol. Biol. 38, 100–114 (2011)

51 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)

52 Arambourg, C. & Biberson, P. The fossil human remains from the Paleolithic site of Sidi Abderrahman (Morocco). Am. J. Phys. Anthropol. 14, 467–489 (1956)

53 Sausse, F. La mandibule atlanthropienne de la carrière Thomas I (Casablanca). Anthropologie 79, 81–112 (1975)

54 Rightmire, G. P. Comparative studies of Late Pleistocene human remains from Klasies River Mouth, South Africa. J. Hum. Evol. 20, 131–156 (1991)

55 Stewart, T. D. in Annual Report of the Smithsonian Institution 521–533 (US Government Printing Office, Washington, 1962)

56 Tillier, A.-M. in Le Squelette moustérien de Kébara 2 97–112 (Centre National de la Recherche Scientifique, Paris, 1991)

57 Stewart, T. D. The Neanderthal skeletal remains from Shanidar Cave, Iraq: a summary of findings to date. Proc. Am. Phil. Soc. 121, 121–165 (1977)

58 Trinkaus, E. The Shanidar Neanderthals (Academic, 1983)

59 Leroi-Gourhan, A. Étude des Restes Humains Fossiles provenant des Grottes d’Arcy-sur-Cure (Masson et Cie, 1958)

60 Daura, J. et al. A Neandertal mandible from the Cova del Gegant (Sitges, Barcelona, Spain). J. Hum. Evol. 49, 56–70 (2005)

61 Topinard, P. Les caracteres simiens de la machoire de la Naulette. Rev. Antropol. 15, 385–431 (1886)

62 Blake, C. C. On a human jaw from the cave of La Naulette, near Dinant, Belgium. Anthropol. Rev. 5, 294–303 (1867)

63 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)

64 Heim, J. L. Les hommes fossiles de La Ferrassie (Dordogne) et le problem de la definition des Neandertaliens classiques. III. Squelette céphalique. Anthropologie 78, 321–378 (1974)

65 de Lumley, M.-A. Les Néandertaliens de la grotte de l’Hortus. Etudes Quaternaires 1, 375–385 (1972)

66 Condemi, S. et al. Possible interbreeding in late Italian Neanderthals? New data from the Mezzena jaw (Monti Lessini, Verona, Italy). PLoS ONE 8, e59781 (2013)

67 Corrain, C. Resti scheletrici umani del ‘Riparo Mezzena’. Memorie del Museo civico di Storia naturale di Verona 16, 97–101 (1968)

68 Walker, M. J., Lombardi, A. V., Zapata, J. & Trinkaus, E. Neandertal mandibles from the Sima de las Palomas del Cabezo Gordo, Murcia, southeastern Spain. Am. J. Phys. Anthropol. 142, 261–272 (2010)

69 Martin, H. Machoire humaine moustérienne trouvée dans la station de La Quina. L’homme préhistorique 13, 3–21 (1926)

70 Martin, H. Position stratigraphique des Ossements humains recueillis dans le Moustérien de La Quina de 1908 à 1912. Bull. Soc. Préhistorique 9, 700–709 (1912)

71 Martin, H. L’Homme fossile de la Quina. (Libraire Octave Doin, 1923)

72 Pap, I., Tillier, A. M., Arensburg, B. & Chech, M. The Subalyuk Neanderthal remains (Hungary): a re-examination. Ann. Hist. Nat. Mus. Natl. Hung. 88, 233–270 (1996)

73 Sanchez, F. Comparative biometrical study of the Mousterian mandible from Cueva del Boquete de Zafarraya (Málaga, Spain). Hum. Evol. 14, 125–138 (1999)

74 Vlcˇek, E. Fossile Menschenfunde von Weimar-Ehringsdorf, Weimarer Monographien zur Ur- und Frühgeschichte Vol. 30 (Landesamt für Archäologische Denkmalpflege, 1993)

75 Condemi, S. Les néandertaliens de La Chaise: abri Bourgeois-Delaunay. Comité des travaux historiques et scientifiques (CTHS, 2001)

76 Bar-Yosef, O . & Vandermeersch, B . (eds) Le squelette moustérien de Kébara 2. (Editions du Centre National de la Recherche Scientifique, 1991)

77 Arambourg, C . & Hoffstetter, R. Le gisement de Ternifine Vol. 1. (Masson, 1963)

78 Hooton, E.A ., Hencken, H.O & Snow, Ch. E. The ancient Palestinian: Skhu¯ l V reconstruction. 17, 5–10 (American School of Prehistoric Research, 1953)

79 Sollas, W. J. The Chancelade skull. J. R. Anthropol. Inst. 57, 89–122 (1927)

80 Martin, H. Caractères des squelettes humains quaternaires de la vallée du Roc (Charente). Bull. Mem. Soc. Anthropol. Paris 8, 103–129 (1927)

81 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. J. Anthropol. Sci. 86, 143–163 (2008)

82 Formicola, V. Una mandibola umana dal deposito dell’Epigravettiano finale delle Arene Candite (scavi 197O). Rev. Antropol. 64, 271–278 (1986)

83 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. Bull. d’Archéologie Marocaine 11, 25–63 (1978)

84 Debénath, A. Nouveaux restes humains atériens du Maroc. CR Acad. Sci. Paris 290, 851–852 (1980)

85 Crognier, E. & Dupouy-Madre, M. Les Natoufiens du Nahal Oren (Ouadi Fallah) Etude anthropologique. Paéorient 2, 103–121 (1974)

86 Henke, W. Vergleichend-morphologische Kennzeichnung der Jungpaläolithiker von Oberkassel bei Bonn. Z. Morphol. Anthropol. 75, 27–44 (1984)

87 Hershkovitz, I. et al. Ohalo II H2: a 19,000-year-old skeleton from a water-logged site at the Sea of Galilee, Israel. Am. J. Phys. Anthropol. 96, 215–234 (1995)

88 Soficaru, A., Dobos, A. & Trinkaus, E. Early modern humans from the Pestera Muierii, Baia de Fier, Romania. Proc. Natl Acad. Sci. USA 103, 17196–17201 (2006)

89 Crevecoeur, I. Etude anthropologique des restes humains de Nazlet Khater (Paléolithique Superieur, Egypte). PhD Thesis, Université Sciences et Technologies Bordeaux I (2006)

90 Thoma, A. Morphology and affinities of the Nazlet Khater man. J. Hum. Evol. 13, 287–296 (1984)

91 Anderson, J. E. In The Prehistory of Nubia Vol. 2 (ed. Wendorf, F. ) 996–1040 (Southern Methodist Univ. Press,1968)

92 Crevecoeur, I. From the Nile to the Danube: a comparison of the Nazlet Khater 2 and Oase 1 early modern human mandibles. Anthropologie 42, 203–213 (2004)

93 Trinkaus, E. et al. An early modern human from the Pes¸ tera cu Oase, Romania. Proc. Natl Acad. Sci. USA 100, 11231–11236 (2003)

94 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)

95 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. The Dolni Vestonice Studies Vol. 5 1–244 (BRNO, 2000)

96 Drozdová, E. The evaluation of a rediscovered fragment of human lower jaw, No 21 from Prˇedmostí u Prˇerova. Archeologické rozhledy 53, 452–460 (2001)

97 Dutour, O. Palimpseste paléoanthropologique sur l’«Homme fossile d’Asselar» (Sahara). Travaux du Laboratoire d’Anthropologie et de Préhistoire des Pays de la Méditerranée Occidentale 1, 73–83 (1992)

98 Gambier, D. Vestiges humains du gisement d’Isturitz (Pyrénées—Atlantiques): étude anthropologique et analyse des traces d’action humaine intentionelle. Antiquités nationales 22–23, 9–26 (1990)

99 Bräuer, G. & Mehlman, M. J. Hominid molars from a middle Stone Age level at the Mumba Rock Shelter, Tanzania. Am. J. Phys. Anthropol. 75, 69–76 (1988)

100 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)

101 Wood, B. A. & Van Noten, F. L. Preliminary observations on the BK 8518 mandible from Baringo, Kenya. Am. J. Phys. Anthropol. 69, 117–127 (1986)

102 Rightmire, G. P. Middle Pleistocene hominids from Olduvai Gorge, northern Tanzania. Am. J. Phys. Anthropol. 53, 225–241 (1980)

103 Bermúdez de Castro, J. M. Dental remains from Atapuerca (Spain) I. Metrics. J. Hum. Evol. 15, 265–287 (1986)

104 Grine, F. E. & Franzen, J. L. Fossil hominid teeth from the Sangiran Dome (Java, Indonesia). Courier Forschungsinstitut Senckenberg 171, 75–103 (1994)

105 Le Cabec, A., Gunz, P., Kupczik, K., Braga, J. & Hublin, J.-J. Anterior tooth root morphology and size in Neanderthals: taxonomic and functional implications. J. Hum. Evol. 64, 169–193 (2013)

106 Turner, C. G. II, Nichol, C. R. & Scott, G. R. Advances in Dental Anthropology (eds Kelley, M., & Larsen, C. ) 13–31 (Wiley Liss, 1991)