The Recently Extinct Plants and Animals Database

Homo erectus Dubois, 1892

Upright man



Taxonomy & Nomenclature

Synonym/s: Homo ergaster; Pithecanthropus erectus Dubois, 1892


Conservation Status

Last Record: 117kya-108kya (Rizal et al., 2019)















NB: The bibliography below includes all African members of H. ergaster as members of H. erectus following (Baab, 2008). If they really do constitute different taxa then the bibliography for H. erectus would necessarily be much shorter.


Abdelhady, A. A., Elewa, A. M. T. and El-Dawy, M. H. (2019). The position of Neandertal and Homo erectus within the hominid clade based on craniodental morphology and whole mtDNA genomes. Homo: Internationale Zeitschrift fur die Vergleichende Forschung am Menschen. DOI: 10.1127/homo/2019/1119 [Abstract]

Ackerman, Sandra J. (2017). Neanderthals Reenvisioned. American Scientist; Research Triangle Park 105(1): 6-7. [Abstract]

Aguirre E. (1994). Homo erectus and Homo sapiens: one or more species? Courier Forsch Inst Senckenberg 171: 333-339.

Aiello, L. C. and Key, C. (2002). Energetic consequences of being a Homo erectus female. American Journal of Human Biology 14(5): 551-565.

An, Z., Gao W, Zhu Y, Kan X, Wang J, Sun J, Wei M. (1990). Magnetostratigrpahic dates of Lantian Homo erectus. Acta Anthropol Sin 9: 1-7.

Andrews, P. (1984). An alternative explanation of the characters used to define Homo erectus. Courier Forschungsinstitut Senckenberg 69: 167-178.

Antón, S. C. (1997). Developmental age and taxonomic affinity of the Mojokerto child, Java, Indonesia. American Journal of Physical Anthropology 102(4): 497-514. [Abstract]

Antón, S. C. (1999). Cranial growth in H. erectus: how credible are the Ngandong juveniles? Am. J. Phys. Anthropol. 108: 223-236.

Antón, S.C., 2001. Cranial growth in Homo erectus. In: Minugh-Purvis, N., McNamara, K.J. (Eds.), Human Evolution through Developmental Change. Johns Hopkins University Press, Baltimore, pp. 349e380.

Antón, SC. (2002a). Evolutionary significance of cranial variation in Asian Homo erectus. Am J Phys Anthropol 118(4): 301-323.

Antón, S. C. (2002b). Cranial growth in Homo erectus, pp. 349-380. In: Minugh Purvis, N. and McNamara, K. J. (eds.). Human evolution through developmental change. Baltimore: Johns Hopkins University Press.

Antón, Susan C. (2003). Natural history of Homo erectus. American Journal of Physical Anthropology 122(S37): 126–170.

Antón, Susan C. (2013). Homo Erectus and Related Taxa, pp. 497-516. In: Begun, David R. (ed.). A Companion to Paleoanthropology. Wiley-Blackwell. 648 pp. [Abstract]

Antón, S. C. and Leigh, S. R. (2003). Growth and life history in Homo erectus. In: Thompson J, Krovitz G, Nelson A. editors. Patterns of growth and development in the genus Homo Cambridge: Cambridge University Press. [incomplete citation]

Antón, Susan C. and Swisher, C. C. III. (2001). Evolution and variation of cranial capacity in Asian Homo erectus, pp. 25-39. In: Indriati E. (ed.). A scientific life: papers in honor of Professor Dr. Teuku Jacob. Yogyakarta, Indonesia: Bigraf Publishing.

Antón, Susan C., Taboada, Hannah G. et al. (2016). Morphological variation in [i]Homo erectus[/i] and the origins of developmental plasticity. Phil. Trans. R. Soc. B 371(1698). DOI: 10.1098/rstb.2015.0236

Arif, J., Baba, H., Kaifu, Y., 2005. The skull IX (Tjf-1993.05) as one of the variant examples of Asian Homo erectusfrom Indonesia. PaleoAnthropology Suppl. A49.

Arif J, Baba H, Suparka ME, Zaim Y, Setoguchi T. (2001). Preliminary study of Homo erectus Skull IX (Tjg-1993.05) from Sangiran, Central Java, Indonesia. Bull Natl Sci Mus Tokyo Ser D 27:1–17.

Úlfur Árnason and Björn Hallström. (2020). The reversal of human phylogeny: Homo left Africa as erectus, came back as sapiens sapiens. Hereditas 157: 51.

Ascenzi, A., Biddittu, I., Cassoli, P. F., Segre, A. G., & Segre-Naldini, E. (1996). A calvarium of late [i]Homo erectus[/i] from Ceprano, Italy. Journal of Human Evolution 31(5): 409-423.

Ascenzi, A., Mallegni, F., Manzi, G., Segre, A. G. and Segre-Naldini, E. (2000). A re-appraisal of Ceprano calvaria affinities with
Homo erectus, after the new reconstruction. J. Human Evol. 39: 443-450.

Ascenzi, A. and Segre, A. G. (2000). The fossil calvaria of Homo erectus from Ceprano (central Italy): a new reconstruction, pp. 25-33. In: Aloisi, M., Battaglia, B., Carafoli, E. and Danieli, G. A. (eds.). The Origin of Humankind. Amsterdam: IOS.

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: 317-320. [Abstract]

Ayala, Francisco J. and Cela-Conde, Camilo J. (2017a). Processes in Human Evolution: The Journey from Early Hominins to Neandertals and Modern Humans (second edition). Oxford: Oxford University Press. [reprinted with corrections in 2018]

Ayala, Francisco J. and Cela-Conde, Camilo J. (2017b). Middle and Lower Pleistocene: the Homo radiation, pp. [?-?]. In: Ayala, Francisco J. and Cela-Conde, Camilo J. (eds.). Processes in Human Evolution: The Journey from Early Hominins to Neandertals and Modern Humans (second edition). Oxford: Oxford University Press. [reprinted with corrections in 2018]

Aziz, F., Baba, H. and Watanabe, N. (1996). Morphological study on the Javanese Homo erectus Sangiran 17 skull based upon the new reconstruction. Geol Res Dev Ctr Paleontol Ser 8: 11–25.

Baab, Karen L. (2007). Cranial Shape Variation in Homo erectus. Ph.D. Dissertation. City University of New York.

Baab, Karen L. (2008). The taxonomic implications of cranial shape variation in Homo erectus. Journal of Human Evolution 54(6): 827-847.

Baab, Karen L. (2010). Cranial shape in Asian Homo erectus: geographic, anagenetic, and size-related variation, pp. 57-80. In: Norton, C. J. and Braun, D. R. (eds.). Asian Paleoanthropology: From Africa to China and Beyond. Dordrecht: Springer Press.

Baab, Karen L. (2016). The role of neurocranial shape in defining the boundaries of an expanded Homo erectus hypodigm. Journal of Human Evolution 92: 1-21. [Abstract]

Baab, Karen L. (2021). Reconstructing cranial evolution in an extinct hominin. Proceedings of the Royal Society B 288(1943). doi: [Abstract] [supplementary information]

Baab, Karen L., Nesbitt, Allison, Hublin, Jean-Jacques and Neubaeur, Simon. (2021). Assessing the status of the KNM-ER 42700 fossil using Homo erectus neurocranial development. Journal of Human Evolution 154: 102980. [Abstract]

Baba H, Aziz F, Kaifu Y, Suwa G, Kono RT, Jacob T. (2003). Homo erectus calvarium from the Pleistocene of Java. Science 299: 1384-1388.

Bailey, Shara E. (2004). A morphometric analysis of maxillary molar crowns of Middle-Late Pleistocene hominins. Journal of Human Evolution, 47(3): 183-198.

Bailey, Shara E. et al. (In Press, 2014). Taxonomic differences in deciduous upper second molar crown outlines of Homo sapiens, Homo neanderthalensis and Homo erectus. Journal of Human Evolution. [Abstract]

Barash, Alon, Bastir, Markus and Been, Ella. (2015). 3D Morphometric Study of the Mandibular Fossa and Its Implication for Species Recognition in Homo erectus. Advances in Anthropology 5: 152-163.

Bartstra, G. J. (1987). Late Homo erectus or Ngandong man of Java. Palaeohist Groningen 29: 1-7.

Bauer, Catherine C. and Harvati, Katarina. (In Press, 2015). A virtual reconstruction and comparative analysis of the KNM-ER 42700 cranium. Anthropologischer Anzeiger. DOI: [Abstract]

Begun, D. & Walker, A. in The Nariokotome Homo erectus Skeleton (eds Walker, A. & Leakey, R.) 326–358 (Springer, Berlin, 1993).

Berghuis, H. W. K. et al. (2021). Hominin homelands of East Java: Revised stratigraphy and landscape reconstructions for Plio-Pleistocene Trinil. Quaternary Science Reviews 260: 106912.

Bilsborough, A., 2000. Chronology, variability and evolution in Homo erectus. Var. Evol. 8, 5e30

Boaz, Noel T. et al. (2001). The Scavenging of ‘Peking Man. Natural History 110: 46-52.

Boaz, Noel T. and Ciochon, Russell L. (2004). Dragon Bone Hill: An Ice-Age Saga of [i]Homo erectus[/i]. Oxford and New York: Oxford University Press. 264 pp.

Boaz, N. T., Ciochon, R. L., Xu, Q, & Lui, J. (2004). ‘Mapping and taphonomic analysis of teh Homo erectus loci at Locality 1 Zhoukoudian, China’. Journal of Human Evolution. 46 (5). pp.519-549.

Bonde, N. (1989). Erectus and neanderthalensis as species or subspecies of Homo with a model of speciation in hominids. In (G. Giacobini, Ed.) Hominidae, pp. 205–208. Milan: Jaca Book

Boyle, Eve K. and Desilva, Jeremy M. (2015). A Large Homo erectus Talus from Koobi Fora, Kenya (KNM-ER 5428), and Pleistocene Hominin Talar Evolution. PaleoAnthropology 2015: 1-13.

Brain CK, Sillen A. 1988. Evidence from the Swarkrans cave for the earliest use of fire. Nature 336: 464-466.

Bräuer, G. 1990. The occurrence of some controversial Homo erectus cranial features in the Zhoukoudian and East African hominids. Acta Anthropol Sin 9: 350-358.

Bräuer, G. (1994a). How different are Asian and African Homo erectus? Courier Forschungsinsitut Senckenburg 171: 301-318.

Bräuer, G. (1994b). How different are Asian and African Homo erectus?, pp. 38-46. In: Franzen, J. (ed.). 100 years of Pithecanthropus: The Homo erectus problem. Courier Forschungsinsitut Senckenburg.

Bräuer, G. and Mbua, E. (1992). Homo erectus features used in cladistics and their variability in Asian and African hominids. Journal of Human Evolution 22(2): 79-108.

Bräuer, G. and Schultz, M. (1996). The morphological affnities of the Plio-Pleistocene mandible from Dmanisi, Georgia. J. hum. Evol. 30: 445-481. [Abstract]

D.C. Broadfield , R.L. Holloway , K. Mowbray , A. Silvers , M.S. Yuan & S. Márquez . 2001. Endocast of Sambungmacan 3 (Sm 3): a new Homo erectus from Indonesia. The Anatomical Record 262: 369-379.

Brown, Frank et al. (1985). Early Homo erectus skeleton from west Lake Turkana, Kenya. Nature 316(6031): 788-792.

Brown, F. H. and McDougall, I. (1993). Geological setting and age, pp. 9-20. In: Walker, A. and Leakey, Richard (eds.). The Nariokotome [i]Homo erectus[/i] skeleton. Cambridge, MA: Harvard University Press.

Brown, P. (1994). Cranial vault thickness in Asian Homo erectus and Homo sapiens. Courier Forschungsinsitut Senckenburg 171: 33-46.

Cachel S, Harris JWK. 1998. The lifeways of Homo erectus inferred from archaeology and evolutionary ecology: a perspective from East Africa. In: Petraglia MD, Korisettar R, editors. Early human behaviour in global context: the rise and diversity of the Lower Paleolithic record. New York: Routledge. p 108-132.

Carotenuto, F. et al. (2016). Venturing out safely: The biogeography of [i]Homo erectus[/i] dispersal out of Africa. Journal of Human Evolution 95: 1-12. [Abstract]

Chamyal L, Dambricourt Malassé A, Maurya DM, Raj R, Bandhari S, Pant SK, Gaillard C. (2011). Discovery of a robust Homo sapiens in India (Orsang, Lower Narmada Basin, Gujrat). Possible continuity with Indian Homo erectus. Acta Anthropologia Sinica 2: 167-191.

Chen T, Wu E, Yang Q, Hu Y. 1997. The ESR dating of the Nanjing Homo erectus stratigraphy. Nuclear Tech 20:732–734 [in Chinese with English abstract and tables].

Choi, K. & Driwantoro, D. Shell tool use by early members of Homo erectus in Sangiran, central Java, Indonesia: cut mark evidence. J. Archaeol. Sci. 34, 48–58 (2007)

Ciochon, Russell L. (2010), "Divorcing Hominins from the Stegodon–Ailuropoda Fauna: New Views on the Antiquity of Hominins in Asia", in John G. Fleagle et al. (eds), Out of Africa I: The First Hominin Colonization of Eurasia , Vertebrate Paleobiology and Paleoanthropology Series, Dordrecht: Springer, pp. 111-126.

Ciochon, R.L., Long, V.T., Larick, R., González, L., Grün, R., de Vos, J., Yonge, C., Taylor, L., Yoshida, H., Reagan, M. (1996). Dated co-occurrence of [i]Homo erectus[/i] and [i]Gigantopithecus[/i] from Tham Khuyen Cave, Vietnam. Proceedings of the National Academy of Sciences 93: 3016-3020. [Abstract]

J. D. Clark, J. de Heinzelin, K. D. Schick, W. K. Hart, T. D. White, G. WoldeGabriel, R. C. Walter, G. Suwa, B. Asfaw, E. Vrba and Y. H.-Selassie (1994). African Homo erectus: old radiometric ages and young Oldowan assemblages in the Middle Awash Valley, Ethiopia. Science 264(5167): 1907-1910.

Clarke RJ. 1994. On the significance of the Swartkrans Homo to the Homo erectus problem. Cour Forsch Inst Senckenberg 171: 185–193.

Clarke RJ. 2000. A corrected reconstruction and interpretation of the Homo erectus skull from Ceprano, Italy. J Hum Evol 39: 433– 442.

Clarke RJ, Howell FC. (1972). Affinities of the Swartkrans SK847 hominid crania. Am J Phys Anthropol 37:319-336.

Clegg, M. and Aiello, L. C. (1999). A comparison of the Nariokotome Homo erectus with juveniles from a modern human population. American Journal of Physical Anthropology 110(1): 81-93 (-94?).

Cofran, Z., DeSilva, J.M. 2015. A neonatal perspective on Homo erectus brain growth. Journal of Human Evolution. 81: 41-17.

Collard, Mark and Cross, Alan. (2017). Thermoregulation in [i]Homo erectus[/i] and the Neanderthals: A Reassessment Using a Segmented Model, pp. 161-174. In: Marom, Assaf and Hovers, Erella (eds.). Human Paleontology and Prehistory: Contributions in Honor of Yoel Rak. Springer International Publishing. [Abstract]

Copes, Lynn E. and Kimbel, William H. (2016). Cranial vault thickness in primates: [i]Homo erectus[/i] does not have uniquely thick vault bones. Journal of Human Evolution 90: 120-134. [Abstract]

Coppens, Y., 2004. The hominids of Melka Kunture. Some general reflections. In: Chavaillon, J., Piperno, M. (Eds.), Studies on the Early Paleolithic site of Melka Kunture, Ethiopia. Origines, Istituto Italiano di Preistoria e Protostoria, Firenze, pp. 685-686.

Coqueugniot, H., Hublin, J. J., Veillon, F., Houët, F., & Jacob, T. (2004). Early brain growth in Homo erectus and implications for cognitive ability. Nature 431(7006): 299-302.

Corvinus, G. (2004). Homo erectus in East and Southeast Asia, and the questions of the age of the species and its association with stone artifacts, with special attention to hand-axe-like tools. Quaternary International 117: 141-151.

Cunningham, Deborah L. et al. (In Press, 2018). The effect of ontogeny on estimates of KNM-WT 15000's adult body size. Journal of Human Evolution. [Abstract]

Day, M. H. (1971). Postcranial remains of Homo erectus from bed IV, Olduvai Gorge, Tanzania. Nature 232: 383-387. [Abstract]

Day MH 1984. Postcranial remains of Homo erectus from Africa, Asia and possibly Europe. Cour Forsch Inst Senckenberg 169: 113–121

Day, M. H. and Stringer, C. (1982). A reconsideration of the Omo Kibish remains and the erectus-sapiens transition, pp. 814-846. Proceedings of the 2nd International Congress of Human Palaeontology, Nice, France, 1982.

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de Vos, John; Sartono, S.; Hardjasasmita, S.; Sondaar, P. Y. (1982), "The fauna from Trinil, type locality of Homo erectus : a reinterpretation", Geologie en Mijnbouw , 61: 207–11.

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Dean C, Leakey MG, Reid D, Schrenk F, Schwartz GT, Stringer C, Walker A (2001) Growth processes in teeth distinguish modern humans from Homo erectus and earlier hominins. Nature 414:628–631

Dean, M. C. (2007). Dental development and life history in primates and a comparison of cuspal enamel growth trajectories in a specimen of Homo erectus from Java (Sangiran S7-37), a Neanderthal (Tabun C1), and an early Homo sapiens specimen (Skhul II), from Israel. in: M. Faerman, L.K. Horwitz, T. Kahana, U. Zilberman (Eds.), Faces from the Past: Diachronic Patterns in the Biology of Human Populations from the Eastern Mediterranean, Archaeopress, BAR International Series, 1603 (2007), pp. 21–27.

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Dean, C., Leakey, M. G., Reid, D., Schrenk, F., Schwartz, G. T., Stringer, C., & Walker, A. (2001). Growth processes in teeth distinguish modern humans from Homo erectus and earlier hominins. Nature 414(6864): 628-631. [Abstract]

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