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Homo erectus

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Homo erectus
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{{short description|Extinct species of the genus Homo}}{{redirect|H. erectus|other uses|H. erectus (disambiguation)|and|Homo erectus (disambiguation)}}{{pp-move-indef}}{{Use dmy dates|date=March 2017}}{{italic title}}{{Speciesbox| taxon = Homo erectus| image = Homme de Tautavel 01-08.jpg| image_upright = 0.7| image_caption = Reconstructed skeleton of Tautavel Manbased on numerous fossil remains of H. erectus. Museum of Prehistory Tautavel, France (2008 photograph)
Eugène Dubois>Dubois, 1893)2|0.07}}Early Pleistocene – Late Pleistocene| synonyms = }}File:Homo erectus adult female - head model - Smithsonian Museum of Natural History - 2012-05-17.jpg|thumb|upright=0.9|right|Forensic reconstruction of an adult female Homo erectus.Reconstruction by John Gurche (2010), Smithsonian Museum of Natural History, based on KNM ER 3733 and (KNM ER 992|992]].Abigail Tucker, "A Closer Look at Evolutionary Faces", Smithsonian.com, 25 February 2010.)(File:Homo erectus new.JPG|thumb|upright=0.9|right|Forensic reconstruction of an adult male Homo erectus.Reconstruction by W. Schnaubelt & N. Kieser (Atelier Wild Life Art), 2006, Westfälisches Museum für Archäologie, Herne, Germany.)Homo erectus (meaning '(:wikt:erectus|upright) man') is a species of archaic humans that lived throughout most of the Pleistocene geological epoch.Its earliest fossil evidence dates to 1.8 million years ago (discovered 1991 in Dmanisi, Georgia).BOOK, Haviland, William A., Walrath, Dana, Prins, Harald E.L., Harald E. L. Prins, McBride, Bunny, 2007, Evolution and Prehistory: The Human Challenge,weblink 8th, Belmont, CA, Cengage Learning, Thomson Wadsworth, 162, 978-0-495-38190-7, A debate regarding the classification, ancestry, and progeny of H. erectus, especially in relation to Homo ergaster, is ongoing, with two major positions:
1) H. erectus is the same species as H. ergaster, and thereby H. erectus is a direct ancestor of the later hominins including Homo heidelbergensis, Homo antecessor, Homo neanderthalensis, Homo denisova, and Homo sapiens; or, 2) it is in fact an Asian species or subspecies distinct from African H. ergaster.Klein, R. (1999). The Human Career: Human Biological and Cultural Origins. Chicago: University of Chicago Press, {{ISBN|0226439631}}.
Some paleoanthropologists consider H. ergaster to be a variety, that is, the "African" variety, of H. erectus; the labels "Homo erectus sensu stricto" (strict sense) for the Asian species and "Homo erectus sensu lato" (broad sense) have been offered for the greater species comprising both Asian and African populations.JOURNAL, Antón, S.C., 2003, Natural history of Homo erectus, Am. J. Phys. Anthropol., 122, 126–170, 10.1002/ajpa.10399, 14666536, By the 1980s, the growing numbers of H. erectus specimens, particularly in Africa, led to the realization that Asian H. erectus (H. erectus sensu stricto), once thought so primitive, was in fact more derived than its African counterparts. These morphological differences were interpreted by some as evidence that more than one species might be included in H. erectus sensu lato (e.g., Stringer, 1984; Andrews, 1984; Tattersall, 1986; Wood, 1984, 1991a, b; Schwartz and Tattersall, 2000) ... Unlike the European lineage, in my opinion, the taxonomic issues surrounding Asian vs. African H. erectus are more intractable. The issue was most pointedly addressed with the naming of H. ergaster on the basis of the type mandible KNM-ER 992, but also including the partial skeleton and isolated teeth of KNM-ER 803 among other Koobi Fora remains (Groves and Mazak, 1975). Recently, this specific name was applied to most early African and Georgian H. erectus in recognition of the less-derived nature of these remains vis à vis conditions in Asian H. erectus (see Wood, 1991a, p. 268; Gabunia et al., 2000a). At least portions of the paratype of H. ergaster (e.g., KNM-ER 1805) are not included in most current conceptions of that taxon. The H. ergaster question remains famously unresolved (e.g., Stringer, 1984; Tattersall, 1986; Wood, 1991a, 1994; Rightmire, 1998b; Gabunia et al., 2000a; Schwartz and Tattersall, 2000), in no small part because the original diagnosis provided no comparison with the Asian fossil record, JOURNAL, Suwa G, Asfaw B, Haile-Selassie Y, White T, Katoh S, WoldeGabriel G, Hart W, Nakaya H, Beyene Y, 10.1537/ase.061203, Early Pleistocene Homo erectus fossils from Konso, southern Ethiopia, Anthropological Science, 115, 2, 133–151, 2007, H. erectus eventually became extinct throughout its range in Africa, Europe and Asia, but developed into derived species, notably Homo heidelbergensis.As a chronospecies, the time of its disappearance is thus a matter of contention. The species name proposed in 1950defines Java Man as the type specimen (now H. e. erectus). Since then, there has been a trend in palaeoanthropology of reducing the number of proposed species of Homo, to the point where H. erectus includes allearly (Lower Paleolithic) forms of Homo sufficiently derived from H. habilis anddistinct from early H. heidelbergensis (in Africa also known as H. rhodesiensis). In this wider sense, H. erectus had mostly been replaced by H. heidelbergensis by about 300,000 years ago, with possible late survival in Java as late as 70,000 years ago.Homo erectus soloensis, found in Java, is considered the latest known survival of H. erectus.Formerly dated to as late as 50,000 to 40,000 years ago, a 2011 study pushed back the date of its extinction of H. e. soloensis to 75,000 years ago at the latest.Indriati E, Swisher CC III, Lepre C, Quinn RL, Suriyanto RA, et al. 2011 The Age of the 20 Meter Solo River Terrace, Java, Indonesia and the Survival of Homo erectus in Asia.PLoS ONE 6(6): e21562. {{doi|10.1371/journal.pone.0021562}}.The discovery of the morphologically divergent Dmanisi skull 5 in 2013 has reinforced the trend of subsuming fossils formerly given separate species names under H. erectus considered as a wide-ranging, polymorphous species.Skull suggests three early human species were one : Nature News & Comment Thus, H. ergaster is now well within the accepted morphological range of H. erectus, and it has been suggested that even H. rudolfensis and H. habilis (alternatively suggested as late forms of Australopithecus rather than early Homo)should be considered early varieties of H. erectus.JOURNAL, A Complete Skull from Dmanisi, Georgia, and the Evolutionary Biology of Early Homo, David Lordkipanidze, Marcia S. Ponce de Leòn, Ann Margvelashvili, Yoel Rak, G. Philip Rightmire, Abesalom Vekua, Christoph P.E. Zollikofer, Science, 18 October 2013, 342, 6156, 326–331, 10.1126/science.1238484, 24136960, 2013Sci...342..326L, NEWS, Switek, Brian, 17 October 2013, Beautiful Skull Spurs Debate on Human History,weblink National Geographic, 22 September 2014,

Discovery and type specimen

{{see|Trinil 2}}The Dutch anatomist Eugène Dubois, inspired by Darwin's theory of evolution as it applied to humanity, set out in 1886 for Asia (despite Darwin's theory of African origin) to find a human ancestor.In 1891–92, his team discovered first a tooth, then a skullcap, and finally a femur of a human fossil on the island of Java, Dutch East Indies (now Indonesia). Excavated from the bank of the Solo River at Trinil, in East Java, he first (1893) allocated the material to a genus of fossil chimpanzees as Anthropopithecus erectus, then the following year assigned his species to a new genus as Pithecanthropus erectus (the genus name had been coined by Ernst Haeckel in 1868 for the hypothetical link between humans and fossil Apes)—from the Greek πίθηκος (píthēkos, "ape") and ἄνθρωπος (ánthrōpos, "human")—based on the proposal that the femur suggested that the creature had been bipedal, like Homo sapiens.Dubois' 1891 find was the first fossil of a Homo-species (or any hominin species) found as result of a directed expedition and search (the first recognized human fossil had been the circumstantial discovery of Homo neanderthalensis in 1856; see List of human evolution fossils).The Java fossil from Indonesia aroused much public interest. It was dubbed by the popular press as Java Man; but few scientists accepted Dubois' argument that his fossil was the transitional form—the so-called "missing link"—between humans and nonhuman apes.{{Sfn|Swisher|Curtis|Lewin|2000|p=70}}File:Ficha del homo georgicus. Museo Arqueológico Nacional de España.jpg|thumb|left| Poster of homo georgicus. National Archaeological Museum of SpainNational Archaeological Museum of SpainMost of the spectacular discoveries of H. erectus next took place at the Zhoukoudian Project, now known as the Peking Man site, in Zhoukoudian, China. This site was first discovered by Johan Gunnar Andersson in 1921NEWS, The First Knock at the Door, Peking Man Site Museum, In the summer of 1921, Dr. J.G. Andersson and his companions discovered this richly fossiliferous deposit through the local quarry men's guide. During examination he was surprised to notice some fragments of white quartz in tabus, a mineral normally foreign in that locality. The significance of this occurrence immediately suggested itself to him and turning to his companions, he exclaimed dramatically "Here is primitive man, now all we have to do is find him!", and was first excavated in 1921, and produced two human teeth.NEWS, The First Knock at the Door, Peking Man Site Museum, For some weeks in this summer and a longer period in 1923 Dr. Otto Zdansky carried on excavations of this cave site. He accumulated an extensive collection of fossil material, including two Homo erectus teeth that were recognized in 1926. So, the cave home of Peking Man was opened to the world., Davidson Black's initial description (1921) of a lower molar as belonging to a previously unknown species (which he named Sinanthropus pekinensis)from sino-, a combining form of the Greek Σίνα "China", and the Latinate pekinensis, "of Peking" prompted widely publicized interest. Extensive excavations followed, which altogether uncovered 200 human fossils from more than 40 individuals including five nearly complete skullcaps.NEWS, Review of the History, Peking Man Site Museum, During 1927–1937, abundant human and animal fossils as well as artefact were found at Peking Man Site, it made the site to be the most productive one of the Homo erectus sites of the same age all over the world. Other localities in the vicinity were also excavated almost at the same time., Franz Weidenreich provided much of the detailed description of this material in several monographs published in the journal Palaeontologica Sinica (Series D).Nearly all of the original specimens were lost during World War II during an attempt to smuggle them out of China for safekeeping; however, authentic casts were made by Weidenreich, which exist at the American Museum of Natural History in New York City and at the Institute of Vertebrate Paleontology and Paleoanthropology in Beijing, and are considered to be reliable evidence.Similarities between Java Man and Peking Man led Ernst Mayr to rename both Homo erectus in 1950.Throughout much of the 20th century, anthropologists debated the role of H. erectus in human evolution. Early in the century, due in part to the discoveries at Java and Zhoukoudian, the belief that modern humans first evolved in Asia was widely accepted.A few naturalists—Charles Darwin most prominent among them—theorized that humans' earliest ancestors were African: Darwin pointed out that chimpanzees and gorillas, humans' closest relatives, evolved and exist only in Africa.BOOK, Darwin, Charles R., The Descent of Man and Selection in Relation to Sex, John Murray, 1871, 978-0-8014-2085-6,

Origin and dispersal

{{see|Early human expansions out of Africa|Acheulean}}File:Carte hachereaux.jpg|thumb|upright=1.5|Map of the distribution of Middle Pleistocene (Acheulean) cleaver finds]]The derivation of the genus Homo from Australopithecina took place in East Africa after 3 million years ago.The inclusion of species dated to just before 2 million years ago, Homo habilis and Homo rudolfensis, into Homo is somewhat contentious.MAGAZINE,weblink Oldest known member of human family found in Ethiopia, New Scientist, 4 March 2015, 2015-03-07, Especially as H. habilis appears to have coexisted with H. ergaster/erectus for a substantial period after 2 Mya, it has been proposed that ergaster may not be directly derived from habilis.JOURNAL, Implications of new early Homo fossils from Ileret, east of Lake Turkana, Kenya, F. Spoor, M.G. Leakey, P.N. Gathogo, F.H. Brown, S.C. Antón, I. McDougall, C. Kiarie, F.K. Manthi, L.N. Leakey, Nature, 7154, 688–691, 9 August 2007, 10.1038/nature05986, 448, 17687323, 2007Natur.448..688S, "A partial maxilla assigned to H. habilis reliably demonstrates that this species survived until later than previously recognized, making an anagenetic relationship with H. erectus unlikely. [...] these two early taxa were living broadly sympatrically in the same lake basin for almost half a million years."Homo erectus emerged about 2 million years ago. Fossils dated close to 1.8 million years ago have been found both in Africa and in West Asia, so it is unclear whether H. erectus emerged in Africa or in Asia. Ferring et al. (2011) suggest that it was still H. habilis which reached West Asia, and that early H. erectus developed there. Early H. erectus would then have dispersed from West Asia, to East Asia (Peking Man) Southeast Asia (Java Man), back to Africa (Homo ergaster), and to Europe (Tautavel Man).JOURNAL, Ferring, R., Oms, O., Agusti, J., Berna, F., Nioradze, M., Shelia, T., Tappen, M., Vekua, A., Zhvania, D., Lordkipanidze, D., 10.1073/pnas.1106638108, Earliest human occupations at Dmanisi (Georgian Caucasus) dated to 1.85-1.78 Ma, Proceedings of the National Academy of Sciences, 108, 26, 10432–10436, 2011, 21646521, 3127884, 2011PNAS..10810432F, JOURNAL, 10.1016/j.quascirev.2010.04.012, June 2011, Augusti, Jordi, Lordkipanidze, David, How "African" was the early human dispersal out of Africa?, 30, 11–12, 1338–1342, Quaternary Science Reviews, 2011QSRv...30.1338A,

Africa{{anchor|Homo_ergaster}}

File:Homo ergaster.jpg|thumb|upright| KNM-ER 3733 (1.6 Mya, discovered 1975 at Koobi Fora, KenyaKenyaIn the 1950s, archaeologists John T. Robinson and Robert Broom named Telanthropus capensis;JOURNAL, Robinson JT, The nature of Telanthropus capensis, Nature, 171, 4340, 33, January 1953, 13025468, 10.1038/171033a0, 1953Natur.171...33R, Robinson had discovered a jaw fragment in 1949 in Swartkrans, South Africa. Later,{{when|date=April 2018}} Simonetta{{who|date=April 2018}} proposed to re-designate it to Homo erectus, and Robinson agreed.BOOK, The First Humans: Origin and Early Evolution of the Genus Homo, Frederick E. Grine, John G. Fleagle, Richard E. Leakey, 1 Jun 2009, Springer, 7, Chapter 2: Homo habilis – A Premature Discovery: Remember by One of Its Founding Fathers, 42 Years Later, From the 1950s forward, numerous finds in East Africa suggested sympatric coexistence for H. ergaster and H. habilis for several hundred millennia, which tends to confirm the hypothesis that they represent separate lineages from a common ancestor; that is, the ancestral relationship between them was not anagenetic, but was cladogenetic, which here suggests that a subgroup population of H. habilis—or of a common ancestor of H. habilis and H. ergaster/erectus—became reproductively isolated from the main-group population, eventually evolving into the new species Homo ergaster (Homo erectus sensu lato).JOURNAL, Implications of new early Homo fossils from Ileret, east of Lake Turkana, Kenya, F. Spoor, M.G. Leakey, P.N. Gathogo, F.H. Brown, S.C. Antón, I. McDougall, C. Kiarie, F.K. Manthi, L.N. Leakey, Nature, 7154, 688–691, 2007-08-09, 10.1038/nature05986, 448, 17687323, 2007Natur.448..688S, "A partial maxilla assigned to H. habilis reliably demonstrates that this species survived until later than previously recognized, making an anagenetic relationship with H. erectus unlikely"In 1961, Yves Coppens discovered a skull in northern Chad. He coined the name Tchadanthropus uxoris for what he considered the earliest fossil human discovered in north Africa.BOOK, Kalb, Jon E, Adventures in the Bone Trade: The Race to Discover Human Ancestors in Ethiopia's Afar Depression, Springer, 2001, 76, 978-0-387-98742-2,weblink 2010-12-02, Although once considered to be a specimen of H. habilis,BOOK, Cornevin, Robert, Histoire de l'Afrique, Payotte, 1967, 440, 978-2-228-11470-7, T. uxoris has been subsumed into H. erectus but it is no longer considered a valid taxon.WEB, Mikko's Phylogeny Archive,weblinkweblink" title="web.archive.org/web/20070106024346weblink">weblink 2007-01-06, Finnish Museum of Natural History, University of Helsinki, It was reported that the fossil "had been so eroded by wind-blown sand that it mimicked the appearance of an australopith, a primitive type of hominid".JOURNAL, 11 July 2002, Wood, Bernard, Palaeoanthropology: Hominid revelations from Chad, Nature (journal), Nature, 418, 6894, 133–135, 10.1038/418133a, 12110870,weblinkweblink" title="web.archive.org/web/20110717090936weblink">weblink yes, 17 July 2011, 2 December 2010, 2002Natur.418..133W, It is probably only 10,000 years old according to stratigraphy, paleontology and C14 dating presented in Michel Servant's PhD as early as 1973.Servant 1983, pp. 462–464.{{BSN|date=April 2019}}{{clarify|date=April 2019|reason=it's an erectus skull that's only 10,000 years old? This seems completely wrong}}

Eurasia

Caucasus{{anchor|Homo erectus georgicus}}

File:Homo Georgicus IMG 2921.JPG|thumb|upright|Dmanisi skull 3 (fossils skull D2700 and jaw D2735, two of several found in Dmanisi in the Georgian TranscaucasusTranscaucasusFile:MEH Homo georgicus 29-04-2012 11-35-22 2372x3863.JPG|thumb|upright|Reconstruction of Homo georgicus based on D2700, by Élisabeth Daynès, Museo de la Evolución HumanaMuseo de la Evolución HumanaHomo erectus georgicus is the subspecies name assigned to fossil skulls and jaws found in Dmanisi, Georgia. First proposed as a separate species, it is now classified within H. erectus.JOURNAL, 10.1126/science.1072953, 12098694, 2002, 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, A new skull of early Homo from Dmanisi, Georgia, 297, 5578, 85–89, Science, 2002Sci...297...85V, JOURNAL, Lordkipanidze D, Jashashvili T, Vekua A, Ponce de León MS, Zollikofer CP, 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, Postcranial evidence from early Homo from Dmanisi, Georgia, 10.1038/nature06134, Nature,weblink 449, 7160, 305–310, 2007, 17882214, 2007Natur.449..305L, LORDKIPANIDZE > FIRST1 = D.
FIRST2 = A. FIRST3 = R. FIRST4 = G.P. FIRST5 = J. FIRST6 = G. FIRST7 = A. FIRST8 = M. FIRST9 = M.S.P. FIRST10 = M. FIRST11 = C.P.E., 10.1038/434717b, Anthropology: The earliest toothless hominin skull, Nature, 434, 7034, 717–718, 2005, 15815618, 2005Natur.434..717L, The site was discovered in 1991 by Georgian scientist David Lordkipanidze. Five skulls were excavated from 1991 forward, including a "very complete" skull in 2005. Excavations at Dmanisi have yielded 73 stone tools for cutting and chopping and 34 bone fragments from unidentified fauna.JOURNAL, Ferring, R., Oms, O., Agusti, J., Berna, F., Nioradze, M., Shelia, T., Tappen, M., Vekua, A., Zhvania, D., Lordkipanidze, D., 10.1073/pnas.1106638108, Earliest human occupations at Dmanisi (Georgian Caucasus) dated to 1.85–1.78 Ma, Proceedings of the National Academy of Sciences, 108, 26, 10432–10436, 2011, 21646521, 3127884, 2011PNAS..10810432F, After their initial assessment, some scientists were persuaded to name the Dmanisi find as a new species, Homo georgicus, which they posited as a descendant of African Homo habilis and an ancestor to Asian Homo erectus. This classification, however, was not supported, and the fossil was instead designated a divergent subgroup of Homo erectus.JOURNAL,weblink A Shrunken Head for African Homo erectus, 10.1126/science.300.5621.893a, 2003, Gibbons, A., Science, 300, 5621, 893a–893, 12738831, JOURNAL, Tattersall, I., Schwartz, J.H., 10.1146/annurev.earth.031208.100202, Evolution of the GenusHomo, Annual Review of Earth and Planetary Sciences, 37, 1, 67–92, 2009, 2009AREPS..37...67T, RIGHTMIRE > FIRST1 = G.P. FIRST2 = D. FIRST3 = A., 10.1016/j.jhevol.2005.07.009, Anatomical descriptions, comparative studies and evolutionary significance of the hominin skulls from Dmanisi, Republic of Georgia, Journal of Human Evolution, 50, 2, 115–141, 2006, 16271745, JOURNAL, 10.1126/science.288.5468.1019 first1 = L. first2 = A. first3 = D. first4 = 3. first5 = R. first6 = A. first7 = M. first8 = M. first9 = S.C. first10 = G. first11 = O. first12 = M.A. first13 = G. first14 = A., Earliest Pleistocene hominid cranial remains from Dmanisi, Republic of Georgia: Taxonomy, geological setting, and age, Science, 288, 5468, 1019–1025, 2000, 10807567, 2000Sci...288.1019G, The fossil skeletons present a species primitive in its skull and upper body but with relatively advanced spine and lower limbs, implying greater mobility than the previous morphology.JOURNAL, 10.2307/4019325, Evolutionary back story: Thoroughly modern spine supported human ancestor, Bower, Bruce, Science News, 169, 18, 275–276, 3 May 2006, 4019325, It is now thought not to be a separate species, but to represent a stage soon after the transition between H. habilis to H. erectus; it has been dated at 1.8 Mya.NEWS, John Noble, Wilford, John Noble Wilford, 19 September 2007, New Fossils Offer Glimpse of Human Ancestors,weblink The New York Times, 9 September 2009, The assemblage includes one of the largest Pleistocene Homo mandibles (D2600), one of the smallest Lower Pleistocene mandibles (D211), a nearly complete sub-adult (D2735), and a toothless specimen D3444/D3900.JOURNAL, 10.1016/j.jhevol.2008.02.003, 18394678, 2008, Rightmire, G. Philip, Van Arsdale, Adam P., Lordkipanidze, David, David Lordkipanidze, Variation in the mandibles from Dmanisi, Georgia, 54, 6, 904–908, Journal of Human Evolution, Two of the skulls—D2700, with a brain volume of {{Convert|600|cc}}, and D4500 or Dmanisi Skull 5, with a brain volume of about 546 centimetres—present the two smallest and most primitive Hominina skulls from the Pleistocene period. The variation in these skulls were compared to variations in modern humans and within a sample group of chimpanzees. The researchers found that, despite appearances, the variations in the Dmanisi skulls were no greater than those seen among modern people and among chimpanzees. These findings suggest that previous fossil finds that were classified as different species on the basis of the large morphological variation among them—including Homo rudolfensis, Homo gautengensis, H. ergaster, and potentially even H. habilis—should perhaps be re-classified to the same lineage as Homo erectus.NEWS, Skull of Homo erectus throws story of human evolution into disarray,weblink Ian Sample, The Guardian, 17 October 2013,

East and Southeast Asia

H. erectus is attested with certainty in East and Southeast Asia from about 0.7 Ma, with possible early presence before 1 Ma; stone tools from Shangchen discovered in 2018 were even claimed to be older than 2 Mya.JOURNAL, Zhu Zhaoyu (), Dennell, Robin, Huang Weiwen (), Wu Yi (), Qiu Shifan (), Yang Shixia (), Rao Zhiguo (), Hou Yamei (), Xie Jiubing (), Han Jiangwei (), Ouyang Tingping (), 2018, Hominin occupation of the Chinese Loess Plateau since about 2.1 million years ago, Nature, 559, 7715, 608–612, 10.1038/s41586-018-0299-4, 29995848, 0028-0836, 2018Natur.559..608Z, JOURNAL, Barras, Colin, 2018, Tools from China are oldest hint of human lineage outside Africa,weblink Nature, 10.1038/d41586-018-05696-8, 0028-0836, Meganthropus refers to a group of fossils found in Java, dated to between 1.4 and 0.9 Mya, which are tentatively grouped with H. erectus, at least in the wider sense of the term in which "all earlier Homo populations that are sufficiently derived from African early Homo belong to H. erectus",Kaifu, Y., et al. (2005). "Taxonomic affinities and evolutionary history of the Early Pleistocene hominids of Java: dentognathic evidence". Am. J. Phys. Anthropol. 128: 709–726. although older literature has placed the fossils outside of Homo altogether.Krantz, G.S. (1975). "An explanation for the diastema of Javan erectus Skull IV". In: Paleoanthropology, Morphology and Paleoecology. La Hague: Mouton, 361–372.Java Man (H. e. erectus, the type specimen for H. erectus), discovered on the island of Java in 1891/2, is dated to 1.0–0.7 Mya. Lantian Man (H. e. lantianensis), discovered in 1963 in Lantian County, Shaanxi province, China, is roughly contemporary with Java Man.Peking Man (H. e. pekinensis), discovered in 1923–27 at Zhoukoudian (Chou K'ou-tien) near Beijing, China, dates to about 0.75 Mya.NEWS, 'Peking Man' older than thought, BBC News, Paul Rincon, 11 March 2009,weblink 22 May 2010, and a new 26Al/10Be dating suggests they are in the range of 680,000–780,000 years old.JOURNAL, 10.1038/nature07741, March 2009, Shen, G, Gao, X, Gao, B, Granger, De, Age of Zhoukoudian Homo erectus determined with (26)Al/(10)Be burial dating, 458, 7235, 198–200, 0028-0836, 19279636, Nature, 2009Natur.458..198S, NEWS,weblink BBC News, 'Peking Man' older than thought, 11 March 2009, 22 May 2010, Yuanmou Man (H. e. yuanmouensis), discovered in Yuanmou County in Yunnan, China, in 1965, is likely of similar age as Peking Man (but with dates proposed as early as 1.7 Mya).Qian F, Li Q, Wu P, Yuan S, Xing R, Chen H, and Zhang H (1991). Lower Pleistocene, Yuanmou Formation: Quaternary Geology and Paleoanthropology of Yuanmou, Yunnan, China. Beijing: Science Press, pp. 17–50Nanjing Man (H. e. nankinensis), discovered in 1993 in the Hulu cave on the Tangshan hills near Nanjing, dates to about 0.6 Mya.JOURNAL, W. Rukang, L. Xingxue,weblink Homo erectus from Nanjing, PaleoAnthropology, 2003, JOURNAL, J. Zhao, K. Hu, K.D. Collerson, H. Xu,weblink Thermal ionisation mass spectrometry U-series dating of a hominid site near Nanjing, China,weblink 2017-09-08, Geology, 29, 27, 2001, 10.1130/0091-7613(2001)0292.0.CO;2, Solo Man (H. e. soloensis), discovered between 1931/1933 along the Solo River, Java, is of uncertain age, dated between 0.25 and 0.075 Mya (the younger date would qualify Solo Man as the latest fossil still classified as H. erectus, even though it shows some derived features, notably larger cranial capacity).WEB,weblink Finding showing human ancestor older than previously thought offers new insights into evolution, 5 July 2011, Terra Daily, 2019-04-22,

Europe

{{see|Homo antecessor|Homo heidelbergensis}}It is conventional to label European archaic humans contemporary with late H. erectus under the separate species name of Homo heidelbergensis, the immediate predecessor of Homo neanderthalensis.H. heidelbergensis fossils are recorded from 600 ka (Mauer 1 mandible), the oldest complete skulls are"Tautavel Man" (Homo erectus tautavelensis), c. 450 ka, and the Atapuerca skull ("Miguelón"), c. 430 ka.The oldest known human fossils found in Europe is a molar from the Sima del Elefante site, Atapuerca Mountains, Spain, dated c. 1.2 Mya. This is associated with the skull fragments of the "Boy of Gran Dolina" found nearby, dated 0.9 Mya and classified as Homo antecessor by the discoverers. The relationship of these fossils to H. erectus is open to debate, as no complete skull has been found.H. e. bilzingslebenensis (Vlcek 1978) refers to skull fragments found at the Bilzingsleben site, Thuringia, Germany.There is, however, indirect evidence of human presence in Europe as early as 1.6 Mya, in the form of stone tools discovered at Lézignan-la-Cèbe, France, in 2008.WEB, Jones, Tim,weblink Lithic Assemblage Dated to 1.57 Million Years Found at Lézignan-la-Cébe, Southern France «, Anthropology.net, 21 June 2012, 2009-12-17, Other stone tools found in France and thought to predate 1 Mya are from Chilhac, Haute-Loire) and from the Grotte du Vallonnet, near Menton.Human presence in Great Britain close to 1 Mya is established by stone tools and fossilized footprints found near Happisburgh, Norfolk.NEWS, Moore, Matthew, Norfolk earliest known settlement in northern Europe,weblink 8 July 2010, The Daily Telegraph, 8 July 2010, London, NEWS, Ghosh, Pallab, Humans' early arrival in Britain,weblink BBC, 8 July 2010, 7 July 2010,

Taxonomy

{{see|Homo|Human taxonomy}}{{multiple image| align = right| direction =horizontal| header=Evolutionary models| total_width=500| image1 = Homo-Stammbaum, Version Stringer-en.svg
Homo over the last 2 million years (vertical axis) based on Stringer (2012).STRINGER TITLE=WHAT MAKES A MODERN HUMAN YEAR=2012 ISSUE=7396 DOI=10.1038/485033A BIBCODE=2012NATUR.485...33S, | image2 = Human evolution chart-en.svg"Figure 5. Temporal and Geographical Distribution of Hominid Populations Redrawn from Stringer (2003)" (edited from source), in REED LAST2=SMITH LAST3=HAMMOND LAST4=ROGERS LAST5=CLAYTON DATE=NOVEMBER 2004 JOURNAL=PLOS BIOLOGY ISSUE=11 DOI=10.1371/JOURNAL.PBIO.0020340 PMC=521174 DISPLAY-AUTHORS=3, Note the depiction of Homo ergaster as an ancestor of Homo erectus.| footer=}}Paleoanthropologists continue to debate the classification of Homo erectus and Homo ergaster as separate species. One school of thought calls H. ergaster the direct African ancestor of H. erectus, proposing that erectus emigrated out of Africa to Asia while branching into a distinct species.BOOK, Tattersall, Ian and Jeffrey Schwartz, Extinct Humans, 2001, 978-0-8133-3482-0, Boulder, Colorado, Westview/Perseus, Some scholars dispense with the species name ergaster, making no distinction between such fossils as the Turkana Boy and Peking Man.{{Citation needed|date=January 2010}} Still, "Homo ergaster" has gained some acceptance as a valid taxon, and the two species are still usually defined as distinct African and Asian populations of the greater species H. erectus, that is, "Homo erectus sensu lato".One of the features distinguishing H. erectus from H. sapiens is the size difference in teeth. H. erectus has large teeth while H. sapiens have smaller teeth.Homo erectus. (2018). Britannica Online Academic Edition, Encyclopædia Britannica, Inc. One theory for the reason of H. erectus having larger teeth is because of the requirements for eating raw meat instead of cooked meat like H. sapiens.Some have insisted that Ernst Mayr's biological species definition cannot be used to test the above hypotheses—that is, that the two species might be considered the same. Alternatively, the amount of variation of cranial morphology between known specimens of H. erectus and H. ergaster can be compared to the same variation within an appropriate population of living primates (that is, one of similar geographical distribution or close evolutionary relationship), such that: if the amount of variation between H. erectus and H. ergaster is greater than that within an appropriately selected population, for example, say, macaques, then H. erectus and H. ergaster may be considered as two different species.{{multiple image| align = right| direction =horizontal| header=Comparative morphology| total_width=300| image1 = Homo erectus skull.jpg| caption1 = The skull of Homo erectus; note the large teeth.| image2 = Homo sapiens-MGL on loan from MAHL-P5030047-white.jpg| caption2 = The skull of Homo sapiens, with its smaller teeth.| footer=}}Finding an extant (i.e., living) model suitable for field study, analysis, and comparison is very important; and selecting a living sample population of an appropriate species can be difficult. (For example, the morphological variation among the global population of H. sapiens is small, so our own species diversity may not be a trustworthy comparison. Fossils found in Dmanisi, Georgia were originally designated as a separate (but closely related) species; but subsequent specimens showed their variation to be within the range of Homo erectus. and they are now classified as Homo erectus georgicus.) New foot tracks found in 2009 in Kenya and reported in Science by Matthew Bennett of Bournemouth University in Britain and his colleagues, confirmed that the gait of Homo erectus was heel-to-toe, walking as a modern human does, rather than with the australopithecine-like method of its own ancestors.JOURNAL, A footprint in the sands of time,weblink The Economist, 22 December 2017, 2009-02-26, H. erectus fossils show a cranial capacity greater than that of Homo habilis (although the Dmanisi specimens have distinctively small crania): the earliest fossils show a cranial capacity of 850 cm³, while later Javan specimens measure up to 1100 cm³,Swisher, Carl Celso III; Curtis, Garniss H. and Lewin, Roger (2002) Java Man, Abacus, {{ISBN|0-349-11473-0}}. overlapping that of H. sapiens.; the frontal bone is less sloped and the dental arcade smaller than that of the australopithecines; the face is more orthognatic (less protrusive) than either the australopithecines or H. habilis, with large brow-ridges and less prominent zygomata (cheekbones). The early hominins stood about {{height|m=1.79|precision=0}}BOOK, Bryson, Bill, A Short History of Nearly Everything: Special Illustrated Edition, Doubleday Canada, Toronto, 2005, 978-0-385-66198-0, —only 17 percent of modern male humans are tallerBOOK, Khanna, Dev Raj, Human Evolution, Discovery Publishing House, 2004, 195, 978-8171417759,weblink African H. erectus, with a mean stature of 170 cm, would be in the tallest 17 percent of modern populations, even if we make comparisons only with males, 30 March 2013, —and were extraordinarily slender, with long arms and legs.NEWS, A Kid Tall For His Age, Roylance, Frank D. Roylance,weblink Baltimore Sun, Clearly this population of early people were tall, and fit. Their long bones were very strong. We believe their activity level was much higher than we can imagine today. We can hardly find Olympic athletes with the stature of these people, 6 February 1994, 30 March 2013, (File:Human arm bones diagram.svg|thumb|Diagram of modern human shoulder bones.)Throwing performance may have been an important mode for early hunting and defense in the genus Homo. Throwing performance in the genus has previously been linked to several anatomical shifts in the upper body during the evolution of Homo. Different fossils and skeletal measures used in reconstructing the Homo erectus shoulder make it possible for either an anteriorly facing shoulder configuration or a lateral orientation that is similar to modern humans.Roach, & Richmond. (2015). "Clavicle length, throwing performance and the reconstruction of the Homo erectus shoulder". Journal of Human Evolution, 80(C), 107–113. These two different orient the throwing ability and hunting behavior of early Homo species. However, it has been found that a commonly used metric for clavicle length (the claviculohumeral ratio) does not predict shoulder position on the torso accurately. Also, no connection between clavicle length and throwing performance was found. This new evidence supports the conclusion that Homo erectus fossil clavicles are similar to modern human variations, with Homo erectus having a shoulder construction that was lateral facing. This suggests that the ability for high speed throwing can be dated back to nearly two million years ago.Sexual dimorphism in H. erectus—males are about 25% larger than females—is slightly greater than seen in the later H. sapiens, but less than that of the earlier genus Australopithecus. Regarding evolution of human physiology, the discovery of the skeleton of "Turkana boy" (Homo ergaster) near Lake Turkana, Kenya, by Richard Leakey and Kamoya Kimeu in 1984—one of the most complete hominin skeletons ever discovered—has contributed greatly to the interpretation.Stringer (2003, 2012) and Reed, et al. (2004) and others have produced schematic graph-models for interpreting the evolution of Homo sapiens from earlier species of Homo, including Homo erectus and/or Homo ergaster, see graphs at right. Blue areas denote the existence of one or more hominin species at a given time and place (that is, region). These and other interpretations differ mainly in the taxonomy and geographical distribution of species.Stringer (see upper graph-model) depicts the presence of H. erectus as dominating the temporal and geographic development of human evolution; and as persisting broadly throughout Africa and Eurasia for nearly 2 million years, eventually evolving into H. heidelbergensis / H. rhodesiensis, which in turn evolved into H. sapiens. Reed, et al. shows Homo ergaster as the ancestor of Homo erectus; then it is ergaster, or a variety of ergaster, or perhaps a hybrid of ergaster and erectus, which develops into species that evolve into archaic and then modern humans and then out of Africa.Both models show the Asian variety of Homo erectus going extinct recently{{when|date=August 2018}}. And both models indicate species admixture: early modern humans spread from Africa across different regions of the globe and interbred with earlier descendants of H. heidelbergensis / H. rhodesiensis, namely the Neanderthals, Denisovans, as well as unknown archaic African hominins. See admixture; and see Neanderthal admixture theory.JOURNAL, Lovers not fighters, Scientific American, 298, 3, 20–21, Whitfield, John, 18 February 2008, 2008SciAm.298c..20W, 10.1038/scientificamerican0308-20,

Habitat

There has been evidence of H. erectus inhabiting a cave in Zhoukoudian, China. This evidence consisted of remains, stones, charred animal bones, collections of seeds, and possibly ancient hearths and charcoal. Although this does not prove that H. erectus lived in caves, it does show that H. erectus spent periods of time in caves of Zhoukoudian. Remains of H. erectus have more consistently been found in lake beds and in stream sediments. This suggest that H. erectus also lived in open encampments near streams and lakes.

Behaviour

Tool use

File:Canto_tallado_2-Guelmim-Es_Semara.jpg|thumb|Early Homo erectus appears to have inherited Oldowan technology and refined it into the AcheuleanAcheuleanFile:Bifaz_cordiforme.jpg|thumb|A cordiform biface as commonly found in the Acheulean, associated with Homo erectus and derived species such as Homo heidelbergensisHomo heidelbergensisThe Paleolithic Age (Old Stone Age) of prehistoric human history and industry is dated from 2.6 million years ago to about 10,000 years ago;Toth, Nicholas; Schick, Kathy (2007). BOOK, Handbook of Paleoanthropology, 10.1007/978-3-540-33761-4_64, Handbook of Paleoanthropology, 1943–1963, 2007, Toth, Nicholas, Schick, Kathy, 978-3-540-32474-4, In Henke, H.C. Winfried; Hardt, Thorolf; Tatersall, Ian. Handbook of Paleoanthropology. Volume 3. Berlin; Heidelberg; New York: Springer-Verlag. p. 1944. (Print: {{ISBN|978-3-540-32474-4}} Online: {{ISBN|978-3-540-33761-4}}) thus it closely coincides with the Pleistocene epoch of geologic time, which is 2.58 million to 11,700 years ago.WEB,weblink The Pleistocene Epoch, University of California Museum of Paleontology, 22 August 2014, The beginning of early human evolution reaches back to the earliest innovations of primitive technology and tool culture. H. erectus were the first to use fire to cook and to make hand axes out of stone.{{citation needed|date=March 2017}}Homo ergaster used more diverse and sophisticated stone tools than its predecessors, where early Homo erectus used comparatively primitive tools. This is probably because H. ergaster inherited, used, and created tools first of Oldowan technology and later advanced the technology to the Acheulean.BOOK, Beck, Roger B., Black, Linda, Krieger, Larry S., Naylor, Phillip C., Shabaka, Dahia Ibo, World History: Patterns of Interaction, McDougal Littell, 1999, Evanston, IL, 978-0-395-87274-1, Because the use of Acheulean tools began ca. 1.8 million years ago,The Earth Institute. (2011-09-01). Humans Shaped Stone Axes 1.8 Million Years Ago, Study Says. Columbia University. Accessed 5 January 2012. and the line of H. erectus diverged some 200,000 years before the general innovation of Acheulean industry in Africa, then it is plausible that the Asian migratory descendants of H. erectus made no use of Acheulean technology. It has been suggested that the Asian H. erectus may have been the first humans to use rafts to travel over bodies of water, including oceans.JOURNAL, Paleoanthropology: Ancient Island Tools Suggest Homo erectus Was a Seafarer, Science, 279, 5357, 1635–1637, 13 March 1998, Gibbons, Ann, 10.1126/science.279.5357.1635, And the oldest stone tool found in Turkey reveals that hominins passed through the Anatolian gateway from western Asia to Europe approximately 1.2 million years ago—much earlier than previously thought. Oldest stone tool ever found in Turkey discovered by the University of Royal Holloway London and published in ScienceDaily on December 23, 2014

Use of fire

East African sites, such as Chesowanja near Lake Baringo, Koobi Fora, and Olorgesailie in Kenya, show potential evidence that fire was utilized by early humans. At Chesowanja, archaeologists found fire-hardened clay fragments, dated to 1.42 M.Y.A.JOURNAL, James, Steven R., February 1989, Hominid Use of Fire in the Lower and Middle Pleistocene: A Review of the Evidence, Current Anthropology,weblink 30, 1, 1–26, 10.1086/203705, 2012-04-04,weblink" title="web.archive.org/web/20151212084645weblink">weblink 12 December 2015, yes, dmy-all, Analysis showed that, in order to harden it, the clay must have been heated to about {{convert|400|C}}. At Koobi Fora, two sites show evidence of control of fire by Homo erectus at about 1.5 M.Y.A., with reddening of sediment associated with heating the material to {{convert|200|-|400|C|F|abbr=off}}. At a "hearth-like depression" at a site in Olorgesailie, Kenya, some microscopic charcoal was found—but that could have resulted from natural brush fires.In Gadeb, Ethiopia, fragments of welded tuff that appeared to have been burned, or scorched, were found alongside H. erectus–created Acheulean artifacts; but such re-firing of the rocks may have been caused by local volcanic activity. In the Middle Awash River Valley, cone-shaped depressions of reddish clay were found that could have been created only by temperatures of {{convert|200|C|abbr=on}} or greater. These features are thought to be burnt tree stumps such that the fire was likely away from a habitation site. Burnt stones are found in the Awash Valley, but naturally burnt (volcanic) welded tuff is also found in the area.A site at Bnot Ya'akov Bridge, Israel is reported to show evidence that H. erectus or H. ergaster controlled fire there between 790,000 and 690,000 years ago;NEWS, Paul, Rincon,weblink Early human fire skills revealed, BBC News, 29 April 2004, 2007-11-12, to date this claim has been widely accepted. Some evidence is found that H. erectus was controlling fire less than 250,000 years ago. Evidence also exists that H. erectus were cooking their food as early as 500,000 years ago.BOOK, Worlds Together, Worlds Apart, Pollard, Elizabeth, Norton, 2015, 978-0-393-92207-3, New York, 13, Re-analysis of burnt bone fragments and plant ashes from the Wonderwerk Cave, South Africa, has been dubbed evidence supporting human control of fire there by 1 M.Y.A.{{citation|date=2 April 2012 |author=Pringle, Heather |title=Quest for Fire Began Earlier Than Thought |journal=ScienceNOW |url=http://news.sciencemag.org/sciencenow/2012/04/quest-for-fire-began-earlier-tha.html?ref=em |archive-url=https://archive.is/20130415203914weblink |dead-url=yes |archive-date=15 April 2013 |accessdate=2012-04-04 }}There is archaeological evidence that Homo erectus cooked their food.

Engravings and religion

File:Homo Erectus shell with geometric incisions circa 500,000 BP, Naturalis Biodiversity Center, Netherlands (with detail).jpg|thumb|upright=2|Homo Erectus shell with geometric incisions, circa 500,000 BP, has been claimed as the first known work of art. From Trinil, Java. Now in the Naturalis Biodiversity Center, Netherlands.JOURNAL, Callaway, Ewen, Homo erectus made world's oldest doodle 500,000 years ago, Nature News, 10.1038/nature.2014.16477,weblink en, 2014,


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