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Defining Homo erectus

  • Karen L. Baab
Living reference work entry

Abstract

Pithecanthropus (now Homo) erectus was first recognized as a species by Eugène Dubois in the 1890s from fossils at the Indonesian site of Trinil. Additional finds from Indonesia and then China expanded the morphological, geographic, and temporal bounds of this species, but it was not until 1960 that H. erectus was recognized in Africa. Since that time, H. erectus has become among the best sampled species in human evolution and has also come to include fossils from Eurasia and possibly South Africa. These fossils are united by a shared neurocranial shape and the presence of a large number of discrete traits. Yet, there is considerable variation among and within fossil assemblages, which has been interpreted within the framework of a long-lived and polytypic species by some, and as indicative of multiple species by others. This chapter reviews the morphological features of the skull that serve to define H. erectus, as well as the extent and implications of variation across the fossil hypodigm. Sometimes the smaller and less derived African and Georgian fossils have been assigned to a separate species, H. ergaster. Yet, geometric morphometric analysis indicates that the degree of cranial shape variation for the entire sample (including African, Georgian, Indonesian, and Chinese fossils) is within the range of many single primate species, and the variation between Indonesian and Chinese fossils is of the same magnitude as that between African/Georgian and Asian fossils. Therefore, the single-species model for H. erectus cannot be rejected on the basis of cranial shape. However, not every fossil assigned to this species is a perfect fit, and some of the most recent additions to H. erectus expand the range of variation in directions that are unexpected based on established patterns of intraspecific variation. The analysis of shape has further confirmed that much of the cranial shape variation present within the species is partitioned among circumscribed temporo-geographic groups. Additional work is necessary to better understand the utility of discrete traits for systematic research and their distribution in the expanded Pleistocene Homo fossil record.

Keywords

Cranial Vault Cranial Capacity Fossil Sample Early Homo Cranial Shape 
These keywords were added by machine and not by the authors. This process is experimental and the keywords may be updated as the learning algorithm improves.

Notes

Acknowledgement

I gratefully acknowledge the Editors, Winfried Henke and Ian Tattersall, for the invitation to participate in this edition of the Handbook of Paleoanthropology. Thanks also to the following individuals and institutions for permission to study their collections: Ian Tattersall, Ken Mowbray, and Gisselle Garcia, American Museum of Natural History; Marta Lahr and Maggie Bellati, University of Cambridge; Teuku Jacob and Etty Indriati, Gadja Mada University; Dominique Grimaud-Hervé, Institut de Paleontologie Humaine; Philippe Mennecier, Musée de L’Homme; Mamitu Yilma, National Museums of Ethiopia; Emma Mbua and Idle Omar Farah, National Museums of Kenya; Rob Kruszynski and Louise Humphrey, Natural History Museum; Michele Morgan, Peabody Museum, Harvard University; Alan Morris, University of Cape Town. Grant support was provided by NSF (BCS 04-24262, DGE 03-33415, and DBI 96-02234), and the Leakey and Sigma Xi Foundations; this is NYCEP Morphometrics contribution 90. Would like to propose CR: Gerhard Weber Visual Anthropology and Biomechanics

References

  1. An Z, Ho CK (1989) New magnetostratigraphic dates of Lantian Homo erectus. Quat Res 32:213–221Google Scholar
  2. An Z, Gao W, Zhu Y, Kan X, Wang J, Sun J, Wei M (1990) Magnetostratigraphic dates of Lantian Homo erectus. Acta Anthropol Sin 9:1–7 (in Chinese, with English abstract)Google Scholar
  3. Andrews P (1984) An alternative interpretation of the characters used to define Homo erectus. Cour Forsch-Inst Senckenberg 69:167–175Google Scholar
  4. Antón SC (2002) Evolutionary significance of cranial variation in Asian Homo erectus. Am J Phys Anthropol 118:801–828Google Scholar
  5. Antón SC (2003) Natural history of Homo erectus. Yearb Phys Anthropol 46:126–170Google Scholar
  6. Antón SC (2004) The face of Olduvai Hominid 12. J Hum Evol 46(3):335–345Google Scholar
  7. Arambourg C (1954) L’hominien fossile de Ternifine (Algérie). CR Hebd Acad Sci 239:893–895Google Scholar
  8. Arif J, Baba H, Kaifu, Y (2005) The skull IX (Tjf-1993.05) as one of the variant examples of Asian Homo erectus from Indonesia. PaleoAnthropology 2005:A49(Abst)Google Scholar
  9. Asfaw B, Gilbert WH, Beyene Y, Hart WK, Renne PR, WoldeGabriel G, Vrba ES, White TD (2002) Remains of Homo erectus from Bouri, Middle Awash, Ethiopia. Nature 416:317–320PubMedGoogle Scholar
  10. Baab KL (2007) Cranial shape variation in Homo erectus. City University of New York, New YorkGoogle Scholar
  11. Baab KL (2008a) A re-evaluation of the taxonomic affinities of the early Homo cranium KNM-ER 42700. J Hum Evol 55:741–746PubMedGoogle Scholar
  12. Baab KL (2008b) The taxonomic implications of cranial shape variation in Homo erectus. J Hum Evol 54(6):827–847PubMedGoogle Scholar
  13. Baab KL (2010) Cranial shape in Asian Homo erectus: geographic, anagenetic, and size-related variation. In: Norton CJ, Braun DR (eds) Asian paleoanthropology: from Africa to China and Beyond. Springer, New York, pp 57–79Google Scholar
  14. Baab KL, Freidline SE, Wang SL, Hanson T (2010) Relationship of cranial robusticity to cranial form, geography and climate in Homo sapiens. Am J Phys Anthropol 141:97–115PubMedGoogle Scholar
  15. Baab KL, McNulty KP, Harvati K (2013) Homo floresiensis contextualized: a geometric morphometric comparative analysis of fossil and pathological human samples. PLoS ONE 8(7):e69119PubMedCentralPubMedGoogle Scholar
  16. Baba H, Aziz F, Kaifu Y, Suwa G, Kono RT, Jacob T (2003a) Homo erectus calvarium from the Pleistocene of Java. Science 299:1384–1388PubMedGoogle Scholar
  17. Baba H, Aziz F, Kaifu Y, Suwa G, Kono RT, Jacob T (2003b) Homo erectus calvarium from the Pleistocene of Java. Science 299(5611):1384–1388PubMedGoogle Scholar
  18. Balzeau A (2006) Are thickened cranial bones and equal participation of the three structural bone layers autapomorphic traits of Homo erectus? Bulletins et Mémoires de la Société d’Anthropologie de Paris 18(3–4):145–163Google Scholar
  19. Balzeau A (2013) Thickened cranial vault and parasagittal keeling: correlated traits and autapomorphies of Homo erectus? J Hum Evol 64:631–644PubMedGoogle Scholar
  20. Bermúdez de Castro JM, Arsuaga JL, Carbonell E, Rosas A, Martı́nez I, Mosquera M (1997) A hominid from the lower Pleistocene of Atapuerca, Spain: possible ancestor to Neandertals and modern humans. Science 276(5317):1392–1395PubMedGoogle Scholar
  21. Bermúdez de Castro JM, Rosas A, Nicolás ME (1999) Dental remains from Atapuerca-TD6 (Gran Dolina site, Burgos, Spain). J Hum Evol 37(3):523–566PubMedGoogle Scholar
  22. Bilsborough A (2000) Chronology, variability and evolution in Homo erectus. Var Evol 8:5–30Google Scholar
  23. Brain CK (1993) Swartkrans: a cave’s chronicle of early man, vol 8. Transvaal Museum, PretoriaGoogle Scholar
  24. Bräuer G (1990) The occurrence of some controversial Homo erectus cranial features in the Zhoukoudian and East African hominids. Acta Anthropol Sinica 9:352–358Google Scholar
  25. Bräuer G (1994) How different are Asian and African Homo erectus? Cour Forsch-Inst Senckenberg 171:301–318Google Scholar
  26. Bräuer G, Mbua E (1992) Homo erectus features used in cladistics and their variability in Asian and African hominids. J Hum Evol 22:79–108Google Scholar
  27. Brown P (1994) Cranial vault thickness in Asian Homo erectus and Homo sapiens. Cour Forsch-Inst Senckenberg 171:33–46Google Scholar
  28. Brown P (2001) Chinese Middle Pleistocene hominids and modern human origins. In: Barham L, Brown KR (eds) Human roots – Africa and Asia in the Middle Pleistocene. Western Academic and Specialist Press, Bristol, pp 135–145Google Scholar
  29. Brown P, Maeda T (2009) Liang Bua Homo floresiensis mandibles and mandibular teeth: a contribution to the comparative morphology of a new hominin species. J Hum Evol 57(5):571–596PubMedGoogle Scholar
  30. Brown FH, McDougall I (1993) Geologic setting and age. In: Walker A, Leakey R (eds) The Nariokotome Homo erectus Skeleton. Harvard University Press, Cambridge, pp 9–20Google Scholar
  31. Brown F, Harris J, Leakey R, Walker A (1985) Early Homo erectus skeleton from west Lake Turkana, Kenya. Nature 316(6031):788–792PubMedGoogle Scholar
  32. 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(16):5674–5678PubMedCentralPubMedGoogle Scholar
  33. Chen T-M, Yang Q, Hu Y-Q, Bao W-B, Li T-Y (1997) ESR dating of tooth enamel from Yunxian Homo erectus site, China. Quat Sci Rev 16(3–5):455–458Google Scholar
  34. Curnoe D (2006) Odontometric systematic assessment of the Swartkrans SK 15 mandible. HOMO 57(4):263–294Google Scholar
  35. de Lumley M-A, Gabounia L, Vekua A, Lordkipandize D (2006) Les restes humains du Pliocène final et du dèbut du Pléistocène inféfieur de Dmanissi, Géorgie (1991–2000). I – Les crânes, D 2280, D 2282, D 2700. Anthropologie 110:1–110Google Scholar
  36. de Lumley H, Khatib S, Mestour B, Gattacceca J, Faure O, Rochette P, Vadeboin F, Feng X (2008) Étude du paléomagnétisme du site de l’Homme de Yunxian. In: de Lumley H, Li T (eds) Le site de l’Homme de Yunxian. Quyuanhekou, Qingqu, Yunxian, Province du Hubei. CNRS éditions et éditions Recherche sur les Civilisations, Paris, pp 185–236Google Scholar
  37. Delson E, Harvati K, Reddy D, Marcus LF, Mowbray K, Sawyer GJ, Jacob T, Marquez S (2001) The Sambungmacan 3 Homo erectus calvaria: a comparative morphometric and morphological analysis. Anat Rec 262(4):380–397PubMedGoogle Scholar
  38. Dennell R, Roebroeks W (2005) An Asian perspective on early human dispersal from Africa. Nature 438(7071):1099–1104PubMedGoogle Scholar
  39. Dubois E (1894) Pithecanthropus erectus, Eine menschenähnliche Übergangsform aus Java. Landesdruckerei, BataviaGoogle Scholar
  40. Dubois E (1924) On the principal characters of the cranium and the brain, the mandible and the teeth of Pithecanthropus erectus. Proc Koninkl Akad Wetensch Amsterdam 27:265–278Google Scholar
  41. Etler DA (1996) The fossil evidence for human evolution in Asia. Annu Rev Anthropol 25:275–301Google Scholar
  42. Etler DA (2004) Homo erectus in East Asia: human ancestor or evolutionary dead-end? Athena Rev 4:37–50Google Scholar
  43. Feibel CS, Brown FH, McDougall I (1989) Stratigraphic context of fossil hominids from the Omo Group deposits, northern Turkana Basin, Kenya and Ethiopia. Am J Phys Anthropol 78:595–622PubMedGoogle Scholar
  44. Ferring R, Oms O, Agustí J, Berna F, Nioradze M, Shelia T, Tappen M, Vekua A, Zhvania D, Lordkipanidze D (2011) Earliest human occupations at Dmanisi (Georgian Caucasus) dated to 1.85–1.78 Ma. Proc Natl Acad Sci 108(26):10432–10436PubMedCentralPubMedGoogle Scholar
  45. Gabounia L, de Lumley M-A, Vekua A, Lordkipandize D, de Lumley H (2002) Découverte d’un nouvel hominidé à Dmanisśi (Transcaucasie, Georgie). CR Palevol 1:243–253Google Scholar
  46. Gabunia L (1992) Der Menschliche Unterkiefer von Dmanisi. Jahrbuch RGZM 39:185–208Google Scholar
  47. Gabunia L, Vekua AK (1995) A Plio-Pleistocene hominid from Dmanisi, East Georgia, Caucasus. Nature 373:509–512PubMedGoogle Scholar
  48. Gabunia L, Vekua A, Lordkipanidze D (1999) A hominid metatarsal from Dmanisi (Eastern Georgia). Anthropos (Brno) 37(2):163–166Google Scholar
  49. Gabunia L, Vekua A, Lordkipanidze D, Swisher CC III, Ferring R, Justus A, Nioradze M, Tvalchrelidze M, Antón SC, Bosinski G (2000) Earliest Pleistocene hominid cranial remains from Dmanisi, Republic of Georgia: taxonomy, geological setting, and age. Science 288(5468):1019–1025PubMedGoogle Scholar
  50. Gower JC (1975) Generalized Procrustes analysis. Psychometrika 40:33–51Google Scholar
  51. Grimaud-Hervé D, Widianto H, Détroit F, Jacob T (2005) The new skull of Bukuran from Sangiran dome, Java, Indonesia. PaleoAnthropology (2005):A46(Abst)Google Scholar
  52. Grimaud-Hervé D, Widianto H, Détroit F, Sémah F (2012) Comparative morphological and morphometric description of the hominin calvaria from Bukuran (Sangiran, Central Java, Indonesia). J Hum Evol 63(5):637–652PubMedGoogle Scholar
  53. Grine FE, Jungers WL, Schultz J (1996) Phenetic affinities among early Homo crania from East and South Africa. J Hum Evol 30(3):189–225Google Scholar
  54. Groves CP (1989) A theory of human and primate evolution. Clarendon Press Oxford, New YorkGoogle Scholar
  55. Grün R, Huang P-H, Huang W, McDermott F, Thorne A, Stringer CB, Yan G (1998) ESR and U-series analyses of teeth from the palaeoanthropological site of Hexian, Anhui Province, China. J Hum Evol 34(6):555–564PubMedGoogle Scholar
  56. Harrison T (1993) Cladistic concepts and the species problem in hominoid evolution. In: Kimbel WH, Martin LB (eds) Species, species concepts, and primate evolution. Plenum Press, New York, pp 346–371Google Scholar
  57. Hawks J, Oh S, Hunley K, Dobson S, Cabana G, Dayalu P, Wolpoff M (2000) An Australasian test of the recent African origin theory using the WLH-50 calvarium. J Hum Evol 39:1–22PubMedGoogle Scholar
  58. Hay RL (1990) Olduvai Gorge: a case history in the interpretation of hominid paleoenvironments in East Africa, establishment of a geologic framework for paleoanthropology. Geological Society of America, BoulderGoogle Scholar
  59. Howell FC (1978) Hominidae. In: Maglio VJ, Cooke HBS (eds) Evolution of African mammals. Harvard University Press, Cambridge, pp 154–248Google Scholar
  60. Howells WW (1980) Homo erectus – who, when and where: a survey. Yearb Phys Anthropol 23:1–23Google Scholar
  61. Huang PH, Grün R (1998) Study on burying ages of fossil teeth from Yuanmou Man site, Yunnan province, China. Acta Anthropol Sin 17:165–170 (in Chinese with English abstract)Google Scholar
  62. Hublin J-J (1986) Some comments on the diagnostic features of Homo erectus. Anthropos (Brno) 23:175–187Google Scholar
  63. Hublin J-J (1987) Les Charactéres dérivés d’Homo erectus: Rélation avec l’augmentation de la masse squelettique. In: Giacobini G (ed) Hominidae: proceedings of the second international congress of human paleontology. Jaca, Torino, pp 199–204Google Scholar
  64. Hublin J-J (2001) Northwestern African Middle Pleistocene hominids and their bearing on the emergence of Homo sapiens. In: Barnham L, Robson-Brown K (eds) Human roots: Africa and Asia in the Middle Pleistocene. Western Academic and Specialist Press Limited, Bristol, pp 99–121Google Scholar
  65. Huffman OF (2001) Geologic context and age of the Perning/Mojokerto Homo erectus, East Java. J Hum Evol 40:353–362PubMedGoogle Scholar
  66. Huffman OF, Shipman P, Hertler C, de Vos J, Aziz F (2005) Historical evidence of the 1936 Mojokerto skull discovery, East Java. J Hum Evol 48:321–363PubMedGoogle Scholar
  67. Huffman OF, Zaim Y, Kappelman J, Ruez DR Jr, de Vos J, Rizal Y, Aziz F, Hertler C (2006) Relocation of the 1936 Mojokerto skull discovery site near Perning, East Java. J Hum Evol 50:431–451PubMedGoogle Scholar
  68. Hyodo M, Nakaya H, Urabe A, Saegusa H, Shunrong X, Jiyun Y, Xuepin J (2002) Paleomagnetic dates of hominid remains from Yuanmou, China, and other Asian sites. J Hum Evol 43(1):27–41PubMedGoogle Scholar
  69. Indriati E, Antón SC (2008) Earliest Indonesian facial and dental remains from Sangiran, Java: a description of Sangiran 27. Anthropol Sci 116(3):219–229Google Scholar
  70. Indriati E, Swisher CC III, Lepre C, Quinn RL, Suriyanto RA, Hascaryo AT, Grün R, Feibel CS, Pobiner BL, Aubert M, Lees W, Antón SC (2011) The age of the 20 meter Solo River terrace, Java, Indonesia and the survival of Homo erectus in Asia. PLoS ONE 6(6):e21562PubMedCentralPubMedGoogle Scholar
  71. Jacob T (1980) The Pithecanthropus of Indonesia: phenotype, genetics and ecology. In: Königson LK (ed) Current argument on Early Man. Pergamon, Oxford, pp 170–179Google Scholar
  72. Jelínek J (1982) The east and southeast Asian way of regional evolution. Anthropos (Brno) 21:195–212Google Scholar
  73. Jolly CJ (2001) A proper study for mankind: analogies from the papionin monkeys and their implications for human evolution. Yearb Phys Anthropol 44:177–204Google Scholar
  74. Jungers WL (2009) Interlimb proportions in humans and fossil hominins: variability and scaling. In: Grine FE, Fleagle JG, Leakey RE (eds) The first humans: origin and early evolution of the genus Homo. Springer, New York, pp 93–98Google Scholar
  75. Jungers WL, Harcourt-Smith WEH, Wunderlich RE, Tocheri MW, Larson SG, Sutikna T, Due RA, Morwood MJ (2009) The foot of Homo floresiensis. Nature 459(7243):81–84PubMedGoogle Scholar
  76. Kaifu Y, Baba H, Aziz F, Indriati E, Schrenk F, Jacob T (2005) Taxonomic affinities and evolutionary history of the early Pleistocene hominids of Java: dentognathic evidence. Am J Phys Anthropol 128(4):709–726PubMedGoogle Scholar
  77. Kaifu Y, Aziz F, Indriati E, Jacob T, Kurniawan I, Baba H (2008) Cranial morphology of Javanese Homo erectus: new evidence for continuous evolution, specialization, and terminal extinction. J Hum Evol 55(4):551–580PubMedGoogle Scholar
  78. Kaifu Y, Zaim Y, Baba H, Kurniawan I, Kubo D, Rizal Y, Arif J, Aziz F (2011) New reconstruction and morphological description of a Homo erectus cranium: skull IX (Tjg-1993.05) from Sangiran, Central Java. J Hum Evol 61(3):270–294PubMedGoogle Scholar
  79. Kappelman J, Alçiçek MC, Kazancı N, Schultz M, Özkul M, Şen Ş (2008) First Homo erectus from Turkey and implications for migrations into temperate Eurasia. Am J Phys Anthropol 135(1):110–116PubMedGoogle Scholar
  80. Kelley J, Plavcan JM (1998) A simulation test of hominoid species number at Lufeng, China: implications for the use of the coefficient of variation in paleotaxonomy. J Hum Evol 35(6):577–596PubMedGoogle Scholar
  81. Kennedy G (1991) On the autapomorphic traits of Homo erectus. J Hum Evol 20:375–412Google Scholar
  82. Kidder JH, Durband AC (2004a) A re-evaluation of the metric diversity within Homo erectus. J Hum Evol 46(3):297–313Google Scholar
  83. Kidder JH, Durband AC (2004b) A re-evaluation of the metric diversity within Homo erectus. J Hum Evol 46(3):299–315PubMedGoogle Scholar
  84. Kramer A (1993) Human taxonomic diversity in the Pleistocene: does Homo erectus represent multiple hominid species? Am J Phys Anthropol 91(2):161–171PubMedGoogle Scholar
  85. Kramer A, Djubiantono T, Aziz F, Bogard JS, Weeks RA, Weinand DC, Hames WE, Michael Elam J, Durband AC (2005) The first hominid fossil recovered from West Java, Indonesia. J Hum Evol 48(6):661–667PubMedGoogle Scholar
  86. Lahr MM, Wright RSV (1996) The question of robusticity and the relationship between cranial size and shape in Homo sapiens. J Hum Evol 31:157–191Google Scholar
  87. Larick R, Ciochon RL, Zaim Y, Suminto S, Rizal Y, Reagan M, Heizler M (2001) Early Pleistocene 40Ar/39Ar ages for Bapang Formation hominins, Central Java, Indonesia. Proc Natl Acad Sci 98:4866–4871PubMedCentralPubMedGoogle Scholar
  88. Le Gros Clark WE (1964) The fossil evidence for human evolution. University of Chicago Press, ChicagoGoogle Scholar
  89. Leakey LSB (1961) New finds at Olduvai Gorge. Nature 189:649–650PubMedGoogle Scholar
  90. Leakey REF (1976) New hominid fossils from the Koobi Fora formation in Northern Kenya. Nature 261:574–576PubMedGoogle Scholar
  91. Leakey REF, Walker AC (1985) Further hominids from the Plio-Pleistocene of Koobi Fora, Kenya. Am J Phys Anthropol 67(2):135–163PubMedGoogle Scholar
  92. Leakey MG, Spoor F, Brown FH, Gathogo PN, Leakey LN (2003) A new hominin calvaria from Ileret (Kenya). Am J Phys Anthropol 36:136, SupplGoogle Scholar
  93. Lee SH (2005) Is variation in the cranial capacity of the Dmanisi sample too high to be from a single species? Am J Phys Anthropol 127:263–266PubMedGoogle Scholar
  94. Lee S-H, Wolpoff MH (2009) The pattern of evolution in Pleistocene human brain size. Paleobiology 29:186–196Google Scholar
  95. Leigh SR (1992) Cranial capacity evolution in Homo erectus and early Homo sapiens. Am J Phys Anthropol 87(1):1–13PubMedGoogle Scholar
  96. Li T, Etler DA (1992) New Middle Pleistocene hominid crania from Yunxian in China. Nature 357:404–407PubMedGoogle Scholar
  97. Li HY, Mei Y (1983) The upper age of Hexian Homo erectus. Chinese Sci Bull 28:703Google Scholar
  98. Li P, Qian F, Ma XH, Pu QY, Na LS, Ju SQ (1976) A preliminary chronological study on fossils of Homo erectus yuanmouensis using a paleomagnetic technique. Sci Sinica 6:579–591Google Scholar
  99. Liu D, Bada JL, Kminek G (2002) Amino acid racemization dating of fossil mammal teeth. In: Wu R, Li X, Wu X, Mu X (eds) Homo erectus from Nanjing. Jiangsu Science and Technology Publishing House, Nanjing, p 310Google Scholar
  100. 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(3):253–262PubMedGoogle Scholar
  101. Lordkipanidze D, Vekua A, Ferring R, Rightmire GP, Agusti J, Kiladze G, Mouskhelishvili A, Nioradze M, Ponce de León MS, Tappen M, Zollikofer CPE (2005) The earliest toothless hominin skull. Nature 434:717PubMedGoogle Scholar
  102. Lordkipanidze D, Jashashvili T, Vekua A, de Leon MSP, Zollikofer CPE, Rightmire GP, Pontzer H, Ferring R, Oms O, Tappen M, Bukhsianidze M, Agusti J, Kahlke R, Kiladze G, Martinez-Navarro B, Mouskhelishvili A, Nioradze M, Rook L (2007) Postcranial evidence from early Homo from Dmanisi, Georgia. Nature 449(7160):305–310PubMedGoogle Scholar
  103. 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 342:326–331PubMedGoogle Scholar
  104. Lu Z, Huang Y, Li P, Meng Z (1989) Yiyuan fossil man. Acta Anthropol Sinica 8:301–313Google Scholar
  105. Macaluso PJ, Hopwood DE, Kioukis G, Malit N, Nevgloski AJ, McCausland-Gaines JB, Lordkipanidze D (2004) Cranial variation among the Plio-Pleistocene hominins from Dmanisi, Georgia. Am J Phys Anthropol 123:139, SupplGoogle Scholar
  106. Maier W, Nkini A (1984) Olduvai Hominid 9: new results of investigation. Cour Forsch-Inst Senckenberg 69:123–130Google Scholar
  107. Márquez S, Mowbray K, Sawyer G, Jacob T, Silvers A (2001) New fossil hominid calvaria from Indonesia – Sambungmacan 3. Anat Rec 262(4):344–368PubMedGoogle Scholar
  108. Martinón-Torres M, Bermúdez de Castro JM, Gómez-Robles A, Margvelashvili A, Prado L, Lordkipanidze D, Vekua A (2008) Dental remains from Dmanisi (Republic of Georgia): morphological analysis and comparative study. J Hum Evol 55(2):249–273PubMedGoogle Scholar
  109. Mayr E (1950) Taxonomic categories in fossil hominids. Cold Spring Harb Symp Quant Biol 15:109–118PubMedGoogle Scholar
  110. Potts R, Behrensmeyer AK, Deino A, Ditchfield P, Clark J (2004) Small Mid-Pleistocene hominin associated with East African Acheulean technology. Science 305:75–78PubMedGoogle Scholar
  111. Qian F, Zhou GX (1991) Quaternary geology and paleoanthropology of Yuanmou, Yunnan, China. Science Press, BeijingGoogle Scholar
  112. Rightmire GP (1984) Comparisons of Homo erectus from Africa and Southeast Asia. Cour Forsch-Inst Senckenberg 69:83–98Google Scholar
  113. Rightmire GP (1986) Species recognition and Homo erectus. J Hum Evol 15:823–826Google Scholar
  114. Rightmire GP (1990) The evolution of Homo erectus: comparative anatomical studies of an extinct human species. Cambridge University Press, CambridgeGoogle Scholar
  115. Rightmire GP (1998a) Evidence from facial morphology for similarity of Asian and African representatives of Homo erectus. Am J Phys Anthropol 106(1):61–85PubMedGoogle Scholar
  116. Rightmire GP (1998b) Human evolution in the Middle Pleistocene: the role of Homo heidelbergensis. Evol Anthropol 6(6):218–227Google Scholar
  117. Rightmire GP (2004) Brain size and encephalization in early to Mid-Pleistocene Homo. Am J phys Anthrop 124:109–123PubMedGoogle Scholar
  118. Rightmire GP (2008) Homo in the Middle Pleistocene: hypodigms, variation, and species recognition. Evol Anthropol 17(1):8–21Google Scholar
  119. Rightmire GP (2013) Homo erectus and Middle Pleistocene hominins: brain size, skull form, and species recognition. J Hum Evol 65(3):223–252PubMedGoogle Scholar
  120. Rightmire GP, Lordkipanidze D (2009) Comparisons of early Pleistocene skulls from East Africa and the Georgian Caucasus: evidence bearing on the origin and systematics of genus Homo. In: Grine FE, Fleagle JG, Leakey RE (eds) The first humans–origin and early evolution of the genus Homo. Springer, New York, pp 39–48Google Scholar
  121. 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(2):115–141PubMedGoogle Scholar
  122. Robinson JT (1961) The australopithecines and their bearing on the origin of Man and of stone tool making. S Afr J Sci 57:3–16Google Scholar
  123. Rohlf FJ, Slice D (1990) Extensions of the Procrustes method for the optimal superimposition of landmarks. Syst Zool 39:40–59Google Scholar
  124. Santa Luca A (1980) The Ngandong fossil hominids: a comparative study of a far eastern Homo erectus group. Department of Anthropology, Yale University, New HavenGoogle Scholar
  125. Sartono S (1982) Characteristics and chronology of early men in Java. In: L'Homo erectus et la place de l'homme de Tautavel Parmi les hominidés fossiles: Congrès International de Paléontologie Humaine, 1er Congrès, Nice, France, Prétirage du Colloque, pp 492–533Google Scholar
  126. Schwartz JH, Tattersall I (2003) The human fossil record. Volume two: Craniodental morphology of the genus Homo (Africa and Asia). Wiley, New YorkGoogle Scholar
  127. Schwartz JH, Tattersall I (2005) The human fossil record. vol 4: Craniodental morphology of Early Hominids (Genera Australopithecus, Paranthropus, Orrorin) and overview. Wiley-Liss, New YorkGoogle Scholar
  128. Shen G, Ku T, Cheng H, Edwards R, Yuan Z, Wang Q (2001) High-precision U-series dating of Locality 1 at Zhoukoudian, China. J Hum Evol 41:679–688PubMedGoogle Scholar
  129. Shen G, Gao X, Gao B, Granger DE (2009) Age of Zhoukoudian Homo erectus determined with 26Al/10Be burial dating. Nature 458(7235):198–200PubMedGoogle Scholar
  130. Simonetta A (1957) Catalogo e sinonimia annotata degli ominidi fossili ed attuali, 1758–1955. Atti della Società toscana di scienze naturali 64:53–112Google Scholar
  131. Simpson SW, Quade J, Levin NE, Butler R, Dupont-Nivet G, Everett M, Semaw S (2008) A female Homo erectus pelvis from Gona, Ethiopia. Science 322(5904):1089–1092PubMedGoogle Scholar
  132. Skinner MM, Gordon AD, Collard NJ (2006) Mandibular size and shape variation in the hominins at Dmanisi, Republic of Georgia. J Hum Evol 51:36–49PubMedGoogle Scholar
  133. 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 448:688–691PubMedGoogle Scholar
  134. Spoor F, Leakey MG, Antón SC, Leakey LN (2008) The taxonomic status of KNM-ER 42700: a reply to Baab (2008a). J Hum Evol 55(4):747–750Google Scholar
  135. Stringer CB (1984) The definition of Homo erectus and the existence of the species in Africa and Europe. Cour Forsch-Inst Senckenberg 69:131–143Google Scholar
  136. Stringer C (2012) The status of Homo heidelbergensis (Schoetensack 1908). Evol Anthropol Issues News Rev 21(3):101–107Google Scholar
  137. Swisher CCI, Curtis GH, Jacob T, Getty AG, Suprijo A, Widiasmoro (1994) The age of the earliest hominids in Java, Indonesia. Science 263:1118–1121PubMedGoogle Scholar
  138. Tamrat E, Thouveny N, Taieb M, Opdyke ND (1995) Revised magnetostratigraphy of the Plio-Pleistocene sedimentary sequence of the Olduvai Formation (Tanzania). Palaeogeogr Palaeoclimatol Palaeoecol 1114:273–283Google Scholar
  139. Tattersall I (1986) Species recognition in human paleontology. J Hum Evol 15(3):165–175Google Scholar
  140. Tattersall I (1992) Species concepts and species identification in human evolution. J Hum Evol 22(4):341–349Google Scholar
  141. Terhune CE, Kimbel WH, Lockwood CA (2007) Variation and diversity in Homo erectus: a 3D geometric morphometric analysis of the temporal bone. J Hum Evol 53:41–60PubMedGoogle Scholar
  142. Thorne AG, Wolpoff MH (1981) Regional continuity in Australasian Pleistocene hominid evolution. Am J Phys Anthropol 55:337–349PubMedGoogle Scholar
  143. Turner A, Chamberlain A (1989) Speciation, morphological change and the status of African Homo erectus. J Hum Evol 18:115–130Google Scholar
  144. Vekua A, Lordkipanidze D, Rightmire GP, Agusti J, Ferring R, Maisuradze G, Mouskhelishvili A, Nioradze M, De Leon MP, Tappen M, Tvalchrelidze M, Zollikofer C (2002) A new skull of early Homo from Dmanisi, Georgia. Science 297(5578):85–89PubMedGoogle Scholar
  145. Vialet A, Guipert G, Jianing H, Xiaobo F, Zune L, Youping W, Tianyuan L, de Lumley M-A, de Lumley H (2010) Homo erectus from the Yunxian and Nankin Chinese sites: anthropological insights using 3D virtual imaging techniques. CR Palevol 9(6):331–339Google Scholar
  146. Vialet A, Guipert G, Cihat Alçiçek M (2012) Homo erectus found still further west: reconstruction of the Kocabaş cranium (Denizli, Turkey). CR Palevol 11(2–3):89–95Google Scholar
  147. Villmoare B (2005) Metric and non-metric randomization methods, geographic variation, and the single-species hypothesis for Asian and African Homo erectus. J Hum Evol 49(6):680–701PubMedGoogle Scholar
  148. von Koenigswald GHR, Weidenreich F (1939) The relationship between Pithecanthropus and Sinanthropus. Nature 144:926–929Google Scholar
  149. Walker A, Leakey RE (1978) The hominids of East Turkana. Sci Am 239(2):54PubMedGoogle Scholar
  150. Wang Y, Wu J, Xu H (2002) 230Th ages dated by thermal ionization and mass spectrometry (TIMS) technique. In: Wu R, Li X, Wu X, Mu X (eds) Homo erectus from Nanjing. Jiangsu Science and Technology Publishing House, Nanjing, p 309Google Scholar
  151. Weidenreich F (1940) Some problems dealing with ancient man. Am Anthropol 42(3):375–383Google Scholar
  152. Weidenreich F (1943) The skull of Sinanthropus pekinensis: a comparative study on a primitive hominid skull. Paleont Sinica Ser D 10:1–298Google Scholar
  153. Weidenreich F (1951) Morphology of solo man. Anthropol Pap Am Mus Nat Hist 43:222–288Google Scholar
  154. White T (1995) African omnivores: global climatic change and Plio-Pleistocene hominids and suids. In: Vrba ES, Denton GH, Partridge TC, Burckle LH (eds) Paleoclimate and evolution, with emphasis on human origins. Yale University Press, New Haven, pp 369–384Google Scholar
  155. Widianto H, Grimaud-Herve D (2000) Un nouveau crâne humain fossile dans le dôme de Sangiran (Java, Indonésie). CR Acad Sci Ser IIa Sci Terre Planets 330(12):883–888Google Scholar
  156. Wolpoff MH (1984) Evolution in Homo erectus: the question of stasis. Paleobiology 10:389–406Google Scholar
  157. Wolpoff MH, Xinzhi W, Thorne AG (1984) Modern Homo sapiens origins: a general theory of hominid evolution involving the fossil evidence from east Asia. In: Smith FH, Spencer F (eds) The origins of modern humans. Liss, New YorkGoogle Scholar
  158. Wolpoff MH, Thorne AG, Jelínek J, Zhang Y (1994) The case for sinking Homo erectus. 100 years of Pithecanthropus is enough! Cour Forsch-Inst Senckenberg 171:341–361Google Scholar
  159. Wood BA (1984) The origin of Homo erectus. Cour Forsch-Inst Senckenberg 69:111Google Scholar
  160. Wood BA (1991) Koobi Fora research project: hominid cranial remains, vol 4. Oxford University Press, New YorkGoogle Scholar
  161. Wood B (1992) Early hominid species and speciation. J Hum Evol 22:351–365Google Scholar
  162. Wood BA (1994) Taxonomy and evolutionary relationships of Homo erectus. Cour Forsch-Inst Senckenberg 171:159–165Google Scholar
  163. Wu X, Poirier FE (1995) Human evolution in China: a metric description of the fossils and a review of the sites. Oxford University Press, OxfordGoogle Scholar
  164. Wu R, Wu X, Zhang S (1989) Early humankind in China. Science Press, BeijingGoogle Scholar
  165. Yan GL (1993) Preliminary study on the magnetic stratigraphy of Yunxian Man Site, Hubei Province. Geosci Bull Chi Geol Univ 18:221–226, In ChineseGoogle Scholar
  166. Zaim Y, Ciochon RL, Polanski JM, Grine FE, Bettis EA III, Rizal Y, Franciscus RG, Larick RR, Heizler M, Aswan EKL, Marsh HE (2011) New 1.5 million-year-old Homo erectus maxilla from Sangiran (Central Java, Indonesia). J Hum Evol 61(4):363–376PubMedGoogle Scholar
  167. Zhang Y (1998) Fossil human crania from Yunxian, China: morphological comparison with Homo erectus crania from Zhoukoudian. Hum Evol 13:45–48Google Scholar
  168. Zhao SS, Pei JX, Guo SL, Liu SS, Qian F, Chou SH, Li XG (1985) Study of chronology of Peking Man site. In: Wu RK, Ren ME, Zhu XM et al (eds) Multi-disciplinary study of the Peking Man site at Zhoukoudian. Science Press, Beijing, pp 239–240Google Scholar
  169. Zhao J, Hu K, Collerson KD, Xu H (2001) Thermal ionization mass spectrometry U-series dating of a hominid site near Nanjing, China. Geology 29:27–30Google Scholar
  170. Zhu R, An Z, Potts R, Hoffman KA (2003) Magnetostratigraphic dating of early humans in China. Earth Sci Rev 61(3–4):341–359Google Scholar

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© Springer-Verlag Berlin Heidelberg 2013

Authors and Affiliations

  1. 1.Department of AnthropologyStony Brook UniversityStony BrookUSA

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