The affinities of Homo antecessor – a review of craniofacial features and their taxonomic validity

Open access


The phylogenetic affinities of Homo antecessor, a hominin dating from the early Middle Pleistocene of Europe, are still unclear. In this study we conducted a comprehensive review of the TD6 hypodigm within the context of the historical development of paleoanthropological issues concerning this species. H. antecessor, based on all available craniofacial features to date, displays a midfacial morphology very similar to specimens attributed to Classic Homo erectus, suggesting that H. antecessor is the geographical European variant of Classic H. erectus.

If the inline PDF is not rendering correctly, you can download the PDF file here.

  • Antón S. 2003. Natural history of Homo erectus. Am J Phys Anthropol 46:126–70.

  • Antón SC. 2012. Early Homo: Who When and Where. Current Anthropology 53 Suppl 6:S278-98.

  • Antón C Potts R Aiello LC. 2014. Evolution of early Homo: An integrated biological perspective. Science 345:1236828.

  • Antón SC Spoor F Fellmann CD Swisher CC III. 2007. Defining Homo erectus: Size considered. In: Henke W Tattersall I editors. Handbook of Paleoanthropology. 1st ed. Berlin Heidelberg New York: Springer-Verlag.

  • Antón SC Swisher CC 3rd. 2004. Early dispersal of Homo from Africa. Ann Rev Anthropol 33:271–96.

  • Antón SC. 2004. The face of Olduvai Hominid 12. J Hum Evol 46:337–47.

  • Arsuaga JL Martínez I Carretero JM Gracia A Lorenzo C Bermúdez de Castro JM. 1996. Evolución humana en Europa: regis-tro y debates. Revista Española de Paleontología Nº Extraordinario:269-77.

  • Arsuaga JL Martínez I Lorenzo C Gracia A Muñoz A Alonso O Gallego J. 1999. The human cranial remains from Gran Dolina Lower Pleistocene site (Sierra de Atapuerca Spain). J Hum Evol 37:431-57.

  • Asfaw B Gilbert WH Beyene Y Hart WK Rennek PR WoldeGabriel G Vrba ES White TD. 2002. Remains of Homo erectus from Bouri Middle Awash Ethiopia. Nature 416:317–20.

  • Bailey SE. 2002. A closer look at Neanderthal postcanine dental morphology: The mandibular dentition. Anat Rec 269:148–56.

  • Bermúdez de Castro JM Arsuaga JL Carbonell E Rosas A Martínez I Mosquera M. 1997. A hominid from the Lower Pleistocene of Atapuerca Spain: Possible ancestor to neandertals and modern humans. Science 276:1392–5.

  • Bermúdez de Castro JM Martinón-Torres M Martín-Francés L Modesto-Mata M Martínez-de-Pinillos M García C Carbonell E. 2017. Homo antecessor: The state of the art eighteen years later. Quaternary International 433:22–31.

  • Blumenschine RJ Peters CR Masao FT Clarke RJ Deino AL Hay RL Swisher CC Stanistreet IG Ashley GM McHenry LJ Sikes NE Van Der Merwe NJ Tactikos JC Cushing AE Deocampo DM Njau JK Ebert JI. 2003. Late Pliocene Homo and Hominid Land Use from Western Olduvai Gorge Tanzania. Science 299:1217–21.

  • Braüer G Schultz M. 1996. The morphological affinities of the Plio-Pleistocene mandible from Dmanisi Georgia. J Hum Evol 30:445–481.

  • Bromage TG. 1989. Ontogeny of the early hominid face. J Hum Evol 18:751-73..

  • Carbonell E Bermúdez de Castro JM Arsuaga JL Allue E Bastir M Benito A Cáceres I Canals T Díez JC Van der Made J Mosquera M Ollé A Pérez-González A Rodríguez J Rodríguez XP Rosas A Rosell J Sala R Vallverdú J Vergés JM. 2005. An Early Pleistocene hominin mandible from Atapuerca-TD6 Spain. Proc Natl Acad Sci USA 102:5674–8.

  • Carbonell E Bermúdez de Castro JM Arsuaga JL Díez JC Rosas A Cuenca-Bescós G Sala R Mosquera M Rodríguez XP. 1995. Lower Pleistocene Hominids and Artifacts from Atapuerca-TD6 (Spain). Science 269:826–30.

  • Cartmill M Smith FH. 2009. The human lineage. Hoboken New Jersey: Wiley-Blackwell.

  • Clarke RJ. 2012. A Homo habilis maxilla and other newly-discovered hominid fossils from Olduvai Gorge Tanzania. J Hum Evol 63:418–28.

  • De Lumley H Li TY directors. 2008. Le site de l’homme de Yunxian: Quyuanhekou Quingqu Yunxian province du Hubei. Paris: CNRS Editions.

  • Delson E. 1997. One skull does not a specie make. Nature 389:445–6.

  • Enlow DH Wang S. 1965. Growth and remodelling of the human maxilla. Am J Orthodontics 51:446–64.

  • Enlow DH. 1966. A morphogenetic analysis of facial growth. Am J Orthodontics 52:283–99.

  • Etler DA. 1994. The Chinese Hominidae: New Finds New Interpretations. PhD thesis. Berkeley: University of California.

  • Etler DA. 2004. Homo erectus in East Asia: human ancestor or evolutionary dead-end? Athena Review 4:37–49.

  • Falk D. 1986. Evolution of Cranial Blood Drainage in Hominids - Enlarged Occipital Marginal Sinuses and Emissary Foramina. Am J Phys Anthropol 70:311–324.

  • Feibel CS Lepre CJ Quinn RL. 2009. Stratigraphy correlation and age estimates for fossils from Area 123 Koobi Fora. J Hum Evol 57:112–22.

  • Feng X. 2008. Stratégie de débitage et mode de façonnage des industries du Paléolithique inférieur en Chine et en Europe entre 1 Ma et 400 000 ans Ressemblances et différences de la culture de l’homme de Yunxian et Acheuléen européen. L’Anthropologie 112:423–47.

  • Feng XB Lu CQ Wang H. 2011. New advance in the study of the Homo erectus site in Yunxian County in Hubei. Jianghan Archaeology 120:57–64.

  • Franciscus RG Trinkaus E. 1988. Nasal morphology and the emergence of Homo erectus. Am J Phys Anthropol 75:517–27.

  • Franciscus RG. 2003. Internal nasal floor configuration in Homo with special reference to the evolution of Neandertal facial form. J Hum Evol 44:701–29.

  • Freidline SE Gunz P Harvati K Hublin JJ. 2013. Evaluating developmental shape changes in Homo antecessor subadult facial morphology. J Hum Evol 65:404–23.

  • Gabunia L de Lumley M-A Vekua A Lordkipanidze D de Lumley H. 2002. Découverte d’un nouvel hominidé à Dmanissi (Transcaucasie Géorgie). C R Paleovol 1:243–53.

  • Gabunia L Vekua A. 1995. A Plio-Pleistocene hominid from Dmanisi East Georgia Caucasus. Nature 373:509–12.

  • Gabunia L Vekua A Lordkipanidze D Swisher CC 3rd Ferring R Justus A Nioradze M Tvalchrelidze M Antón SC Bosinski G Jöris O Lumley MA Majsuradze G Mouskhelishvili A. 2000. Earliest Pleistocene hominid cranial remains from Dmanisi Republic of Georgia: taxonomy geological setting and age. Science 288:1019–25.

  • Gower CG. 1923. A contribution to the morphology of the apertura piriformis. Am J Phys Anthropol 6:27–36.

  • Hay RL. 1976. The setting of the gorge and history of the geological investigations. In: Hay RL editor. The Geology of the Olduvai Gorge. Berkeley: University of California Press.

  • Henke W. 1995. Qualitative and quantitative analysis of the Dmanisi mandible. In: Radlanski RJ Renz H editors. Proceedings of the 10th international symposium on dental morphology. Berlin: Christine und Michael Brünae GbA.

  • Johanson DC Masao FT Eck GG White TD Walter RC Kimbel WH Asfaw B Manega P Ndessokia P Suwa G. 1987. New partial skeleton of Homo habilis from Olduvai Gorge Tanzania. Nature 327:205–9.

  • Joordens J Dupont-Nivet G Feibel C Spoor F Sier MJ van der Lubbe JH Nielsen TK Knul MV Davies GR Vonhof HB. 2013. Improved age control on early Homo fossils from the upper Burgi Member at Koobi Fora Kenya. J Hum Evol 65:731–45.

  • Kaifu Y Indriati E Aziz F Kurniawan I Baba H. 2010. Cranial Morphology and Variation of the Earliest Indonesian Hominids. In: Norton CJ Braun DR; editors. Asian Paleoanthropology: From Africa to China and Beyond. Springer.

  • Kimbel WH Rak Y Johanson DC. 2004. The skull of Australopithecus afarensis. Oxford University Press.

  • Kimbel WH. 2009. The origin of Homo. In: Grine FE Fleagle JG Leakey RE editors. The first humans. Origin and early evolution of the genus Homo. Springer.

  • Kimbel WH Johanson DC Rak Y. 1997. Systematic Assessment of a Maxilla of Homo From Hadar Ethiopia. Am J Phys Anthropol 103:235–62.

  • Kimbel WH White TD Johanson DC. 1984. Cranial Morphology of Australopithecus afarensis: A Comparative Study Based on a Composite Reconstruction of the Adult Skull. Am J Phys Anthropol 64:337–388.

  • Koesbardiati T. 2000. On the relevance of the regional features of the face in East Asia. PhD Thesis. Hamburg: Universität Hamburg.

  • Kurihara S Enlow DH Rangel RD. 1980. Remodelling reversals in anterior parts of the human mandible and maxilla. Angle Orthod 50:98–106.

  • Lacruz RS Bermúdez de Castro JM Martinón-Torres M O’Higgins P Paine ML Carbonell E Arsuaga JL Bromage TG. 2013. Facial morphogenesis of the earliest Europeans. PLoS One 8: e65199.

  • Lacruz RS Bromage TG O’Higgins P Toro-Ibacache V Warshaw J Berger LR. 2015. Distinct growth of the nasomaxillary complex in Au. sediba. Sci Rep 5:15175.

  • Lahr MM. 1994. The Multiregional Model of Modern Human Origins: A Reassessment of its Morphological Basis. J Hum Evol 26:23–56.

  • Larick R Ciochon RL Zaim Y Sudijono Suminto Rizal Y Aziz F Reagan M Heizler M. 2001. Early Pleistocene 40Ar/39Ar ages for Bapang Formation hominins Central Java Indonesia. Proc Natl Acad Sci USA 989:4866–71.

  • Leakey MG Spoor F Dean MC Feibel GS Antón SC Kiarie C Leakey LN. 2012. New fossils from Koobi Fora in northern Kenya confirm taxonomic diversity in early Homo. Nature 488:201–4.

  • Leakey REF Walker AC. 1985. Further hominids from the Plio-Pleistocene of Koobi-Fora Kenya. Am J Phys Anthropol 67:135–63.

  • Li T Etler DA. 1992. New middle Pleistocene hominid crania from Yunxian in China. Nature 357: 404–7.

  • Liu W Zhang Y Wu X. 2005. Middle Pleistocene human cranium from Tangshan (Nanjing) Southeast China: A new reconstruction and comparisons with Homo erectus from Eurasia and Africa. Am J Phys Anthropol 127:253–62.

  • Lordkipanidze D Ponce de León MS Margvelashvili A Rak Y Rightmire GP Vekua A Zollikofer CPE. 2013. A Complete Skull from Dmanisi Georgia and the Evolutionary Biology of Early Homo. Science 342:326–31.

  • Maddux SD Franciscus RG. 2009. Allometric scaling of infraorbital surface topography in Homo. J Hum Evol 56:161–74.

  • Martinón-Torres M Bermúdez de Castro JM Gómez-Robles A Arsuaga JL Carbonell E Lordkipanidze D Manzi G Margvelashvili A. 2007. Dental evidence on the hominin dispersals during the Pleistocene. Proc Natl Acad Sci USA 104:13279–82.

  • Maureille B Houeët F. 1997. Nouvelles données sur charactèristiques derivées du masif facial superieur des Neandertaliens. Anthropol Prehistorie 108 :89–98.

  • McCollum MA Grine FE Ward SC Kimbel WH. 1993. Subnasal morphological variation in extant hominoids and fossil hominids. J Hum Evol 24:87–111.

  • McCollum MA. 2000. Subnasal morphological variation in fossil hominids: a reassessment based on new observations and recent developmental findings. Am J Phys Anthropol 112:275–83.

  • McCollum MA. 2008. Nasomaxillary remodelling and facial form in robust Australopithecus: a reassessment. J Hum Evol 54:2–14.

  • Mellinger W. 1940. The canine fossa. Arch Otolaryngol 31:930-7.

  • Parés JM Arnold L Duval M Demuro M Pérez-González A Bermúdez de Castro JM Carbonell E Arsuaga JL. 2013. Reassessing the age of Atapuerca-TD6 (Spain): new paleomagnetic results. Journal of Archaeological Science 40:4586–95.

  • Pope GG. 1991. Evolution of the zygomaticomaxillary region in the genus Homo and its relevance to the origin of modern humans. J Hum Evol 21:189–213.

  • Potts R Beherensmeyer AK Deino A Ditch-field P Clark J. 2004. Small Mid-Pleistocene Hominin Associated with East African Acheulean Technology. Science 305:75–78.

  • Prat S. 2002. Anatomical study of the skull of the Kenyan specimen KNM-ER 1805: a re-evaluation o its taxonomic allocation? C. R. Paleoevol 1:27–33.

  • Prat S. 2004. Les premiers représentants du genre Homo en quête d’une identité. Apports de l’étude morphologique et de l’analyse cladistique. Bulletins et Mémoires de la Société d’Anthropologie de Paris 16:17–35.

  • Rabadà i Vives D. Los homínidos de Atapuerca: entre la sinonimia y la paradoja (Sierra de Atapuerca Burgos España). Batalleria 12:143–8.

  • Rak Y. 1983. The Australopithecine Face. New York: Academic Press.

  • Rak Y. 1986. The Neanderthal: A New Look at an Old Face. J Hum Evol 15:151–64.

  • Ribot F García Bartual M Wang Q. 2018. A comparative study of the craniofacial features defining ‘Homo antecessor’. Ahead Acta Anthropologica Sinica (in press).

  • Ribot F Gibert J García Bartual M. 2007. La evolución facial de los homínidos: un modelo de evolución darvinista. In: Braga JC Checa A Company M editors. XXIII Jornadas de la Sociedad Española de Pale-ontología. Libro de resúmenes. Instituto Geológico y Minero de España y Universidad de Granada 2007 195. [Abstract]. (Caravaca de la Cruz 3-6 de Octubre de 2007).

  • Ribot F Gibert J Sánchez F Iglesias A Gibert L. 2006. Homo antecessor: un nouveau taxon? XV UISPP Congress: World Congress of the International Union for Prehistoric and Protohistoric Sciences (UISPP). S03-29 Lisbon Portugal 4-9 September 2006.

  • Ribot Trafí F García Bartual M. 2016. Evolución del esqueleto mediofacial en Homo. Implicaciones para un modelo de evolución. In: Ribot F coordinador. Homenaje al Dr. José Gibert Clols. Una vida dedicada a la ciencia y al conocimiento de los primer-os europeos. Granada Spain: Publicaciones Diputación de Granada.

  • Rightmire GP Lordkipanidze D Vekua A. 2006. Anatomical descriptions comparative studies and evolutionary significance of the hominin skulls from Dmanisi Republic of Georgia. J Hum Evol 50:115–41.

  • Rightmire GP Ponce de León MS Lordkipanidze D Margvelashvili A Zollikofer CP. 2017. Skull 5 from Dmanisi: Descriptive anatomy comparative studies and evolutionary significance. J Hum Evol 104:50–79.

  • Rightmire GP. 1993. Variation among early Homo crania from Olduvai Gorge and the Koobi Fora region. Am J Phys Anthropol 90:1–33.

  • Rightmire GP. 1998a. Human Evolution in the Middle Pleistocene: The role of Homo heidelbergensis. Evolutionary Anthropology 6:218–27.

  • Rightmire GP. 1998b. Evidence from facial morphology for similarity of Asian and African representatives of Homo erectus. Am J Phys Anthropol 106:61–85.

  • Rightmire GP. 2001. Patterns of hominid evolution and dispersal in the Middle Pleistocene. Quaternary International 75:77–84.

  • Rightmire GP. 2007. Later Middle Pleistocene Homo. In: Henke W Tattersall I editors. Handbook of Paleoanthropology. 1st ed. Berlin Heidelberg New York: Springer-Verlag.

  • Rodrigues Freire A Bevilacqua Prado F Cláudia Rossi A Yoshito Noritomi P Haiter Neto F Ferreira Caria PH. 2014. Biomechanics of the human canine pillar based on its geometry using finite element analysis. Int J Morphol 32:214–20.

  • Rosas A. 2000. Human evolution in the last million years — The Atapuerca evidence. Acta Anthropol Sin 19 (Suppl.):8–17.

  • Sawyer GJ Tattersall I. 1995. A new reconstruction of the skull of Homo erectus from Zhoukoudian China. In: Gibert J Sánchez F Gibert L Ribot F editors. The Hominids and their environment during the lower and middle Pleistocene of Eurasia. Proceedings of the International Conference of Human Palaeontology (Orce). Granada: Museo de Prehistoria y Paleontología J. Gibert y Ayuntamiento de Orce.

  • Schrenk F Kullmer O Bromage T. 2015. The Earliest Putative Homo Fossils. In: Henke W Tattersall I editors. Handbook of Paleo-anthropology. 2nd ed. Berlin Heidelberg New York: Springer-Verlag.

  • Schwartz J Tattersall I editors. 2002. The Human Fossil Record Volume 1: Terminology and Craniodental Morphology of Genus Homo (Europe). New York: Wiley-Liss.

  • Schwartz JH Tattersall I. 2003. The human fossil record. Volume 2. Craniodental morphology of genus Homo (Africa and Asia). New York: Wiley-Liss.

  • Shen G Gao X Gao B Granger DE. 2009. Age of Zhoukoudian Homo erectus determined with 26Al/10Be burial dating. Nature 458:198-200.

  • Shen G Jin L. 1991. Restudy of the upper age limit of Beijing Man cave. Acta Anthrop Sinica 10:273–277.

  • Shen G. 2001. Current status and prospects for dating fossil hominid sites in China. In: Tobias PV Rath MA Moggi-Cecchi J Doyle GA editors. Humanity from African naissance to coming millennia. Florence Italy: Firenze Univ. Press.

  • Simpson SW. 2015. Early Pleistocene Homo. In: Muehlenbein MP editor. Basics in Human Evolution. Elsevier.

  • Spoor F Leakey MG Gathogo PN Brown FH Antón SC McDougall I Kiarie C Manthi FK Leakey LN. 2007. Implications of new early Homo fossils from Ileret east of Lake Turkana Kenya. Nature 448:688–91.

  • Stan MC Bordei P Dina C Iliescu DM. 2013. Characteristics of some anatomical landmarks on the anterior face of the maxilla. ARS Medica Tomitana 2:79–83.

  • Strait DS Grine FE Moniz MA. 1997. J Hum Evol 32: 17–82.

  • Stringer C. 2002. Modern human origins: progress and prospects. Philos Trans R Soc Lond B Biol Sci 357:563–79.

  • Swisher CC 3rd Curtis GH Jacob T Getty AG Suprijo A Widiasmoro. 1994. Age of the earliest known hominids in Java Indonesia. Science 263:1118–21.

  • Tamrat E Tohveny N Taieb M Opdyke N. 1995. Revised magnetostratigraphy of Plio-Pleistocene sedimentary sequence of the Olduvai Formation (Tanzania). Palaeogeogr Palaeoclimatol Palaeoecol 114:273–83.

  • Tattersall I. 2015. Homo ergaster and Its Contemporaries. In: Henke W Tattersall I editors. Handbook of Paleoanthropology. 2nd ed. Berlin Heidelberg New York: Springer-Verlag.

  • Tattersall I Sawyer GJ. 1996. The skull of “Sinanthropus” from Zhoukoudian China: a new reconstruction. J Hum Evol 31:311–4.

  • Tobias PV. 1967. The Cranium and Maxillary Dentition of Australopithecus (Zinjanthropusy) boisei En: Olduvai Gorge: Volume 2. Cambridge: Cambridge University Press.

  • Tobias PV. 1980. The natural history of the helicoidal occlusal plane and its evolution in early Homo. Am J Phys Anthropol. 53:173–187.

  • Tobias PV. 1991. The Skulls Endocasts and Teeth of Horno habilis In: Olduvai Gorge: Volume 4. Cambridge: Cambridge University Press.

  • Vekua A Lordkipanidze D Rightmire GP Agustı´ J Ferring R Majsuradze G Mouskhelishvili A Noradze M Ponce de León M Tappen M Tvalchrelidze M Zollikofer C. 2002. A new skull of early Homo from Dmanisi Georgia. Science 297:85–9.

  • Walker AC Leakey MD. 1993. The Nariokotome H. erectus skeleton. Cambridge: Harvard University Press.

  • Wang Q Sun L. 2000. Eightieth year of Peking Man: Current status of Peking Man and the Zhoukoudian site. Anthropol Rev 63:19–30.

  • Wang Q Tobias PV. 2000a. Review of the phylogenetic position of Chinese Homo erectus in light of midfacial morphology. Acta Anthropol Sin 19 (Suppl.):23–33.

  • Wang Q Tobias PV. 2000b. Recent advances in Chinese palaeo-anthropology. South African Journal of Science 96:463–6.

  • Wang Q Tobias PV. 2001. An old species and a new frontier: Some thoughts on the taxonomy of Homo erectus. Przeglad Antropologiczny - Anthropological Review 64:9–20.

  • Wang Q Wright BW Smith A Chalk J Byron GD. 2010. Mechanical impact of incisor loading on the primate midfacial skeleton and its relevance to human evolution. The Anatomical Record 293:607–17.

  • Wang Q. 1998. Comparison of Homo heidelbergensis and Chinese Homo erectus. Ph.D. Dissertation. Institute of Vertebrate Pale-ontology and Paleoanthropology Chinese Academy of Sciences Beijing.

  • Weidenreich F. 1937a. Reconstruction of the entire skull of an adult female individual of Sinanthropus pekinensis. Nature 140:1010–1.

  • Weidenreich F. 1937b. The dentition of Sinanthropus pekinensis. Palaeont Sin New Ser D1:1–180.

  • Weidenreich F. 1943. The skull of Sinanthropus pekinensis: A comparative study on a primitive hominid skull. Palaeont Sin New Ser D10:1–484.

  • Williams FL Schroeder L Ackermann RR. 2012. The mid-face of lower Pleistocene hominins and its bearing on the attribution of SK 847 and StW 53. HOMO - Journal of Comparative Human Biology 63:245–57.

  • Woo J-K. 1966. The hominid skull of Lantian Shensi. Vertebrata PalAsiatica 10:1–16.

  • Wood B. 1991. Koobi Fora Research Project. Vol. 4: Hominid cranial remains. Oxford: Clarendon Press.

  • Wood BA. 1992. Origin and evolution of the genus Homo. Nature 355:783–90.

  • Wood B Richmond BG. 2000. Human evolution: taxonomy and paleobiology. J Anat 196:19–60.

  • Wu X Poirier FE. 1995. Human Evolution in China: A Metric Description of the Fossils and a Review of the Sites. London: Oxford University Press.

  • Yan GL. 1993. A preliminary study on magnetic stratigraphy of the geological section with the fossil bed of Yunxian Homo of Hubei. Earth Science-Journal of China University of Geosciences 18:221–6.

  • Zhao J Hu K Collerson KD Xu H. 2001. Thermal ionization mass spectrometry U-series dating of a hominid site near Nanjing China. Geology 29:27–30.

  • Zhu ZY Dennell R Huang WW Wu Y Rao ZG Qiu SF Xie JB Liu W Fu SQ Han JW Zhou HY Ou Yang TP Li HM. 2015. New dating of the Homo erectus cranium from Lantian (Gongwangling) China. J Hum Evol 78:144–57.

Journal information
Impact Factor

CiteScore 2018: 0.55

SCImago Journal Rank (SJR) 2018: 0.22
Source Normalized Impact per Paper (SNIP) 2018: 0.472

All Time Past Year Past 30 Days
Abstract Views 0 0 0
Full Text Views 393 301 25
PDF Downloads 294 214 23