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{{short description|Extinct species of archaic human}}
{{short description|Extinct species of archaic human}}
{{good article}}
{{redirect|H. erectus|other uses|H. erectus (disambiguation)|and|Homo erectus (disambiguation)}}
{{redirect|H. erectus|other uses|H. erectus (disambiguation)|and|Homo erectus (disambiguation)}}
 
{{redirect|Pithecanthropus erectus|the song and album by Charles Mingus|Pithecanthropus Erectus (album)}}
{{Use dmy dates|date=September 2020}}
{{pp-move}}
{{Use dmy dates|date=January 2025}}
{{Speciesbox
{{Speciesbox
| fossil_range = {{Fossil range|2|0.1}}<small>[[Early Pleistocene]] – [[Late Pleistocene]]<ref name=Rizal/></small>
| fossil_range = {{Fossil range|2|0.1}} [[Pleistocene]]<ref name=Rizal2020/>
| image = Peking Man Skull (replica) presented at Paleozoological Museum of China.jpg
| image = Java Man holotype.jpg
| image_caption = Replica of the skull of [[Peking Man]] at the [[Paleozoological Museum of China]]
| image_caption = [[Java Man]], the [[holotype]] of ''H. erectus''{{efn-num|The Java Man [[femur]] presents a noticeable [[osteocyte]], likely [[Paget's disease of bone]] and [[osteopetrosis]] (thickening of the bone), probably stemming from [[skeletal fluorosis]] caused by ingestion of food contaminated by [[fluoride toxicity|fluoride]]-filled [[volcanic ash#ingesting ash|volcanic ash]] (as the specimen was found in ash-filled [[stratum (geology)|strata]]). Livestock that graze on volcanic ash-ridden fields typically die of [[acute toxicity]] within a few days or weeks.<ref>{{cite journal |last=Soriano|first=M. |title=The fluoric origin of the bone lesion in the ''Pithecanthropus erectus'' femur |journal=American Journal of Physical Anthropology |volume=32 |issue=1 |pages=49–57 |year=1970 |pmid=4984453 |doi=10.1002/ajpa.1330320107 |bibcode=1970AJPA...32...49S }}</ref>}}
| extinct = yes
| taxon = Homo erectus
| taxon = Homo erectus
| authority = ([[Eugène Dubois|Dubois]], 1893)
| authority = ([[Eugène Dubois|Dubois]], 1893)
| extinct = yes
| synonyms = * ''[[Java Man|Anthropopithecus erectus]]'' <small>[[Eugène Dubois|Dubois]], 1893</small>
* ''[[Java Man|Pithecanthropus erectus]]'' <small>([[Eugène Dubois|Dubois]], 1893)</small>
* ''[[Peking Man|Sinanthropus pekinensis]]''
* ''[[Solo Man|Javanthropus soloensis]]''
* ''[[Tighennif|Atlanthropus mauritanicus]]''
* (?) ''[[Homo ergaster|Telanthropus capensis]]''
* (?) ''[[Homo georgicus]]''
* (?) ''[[Tautavel Man|Homo tautavelensis]]''
}}
}}


'''''Homo erectus''''' ({{IPAc-en|ˌ|h|oʊ|m|oʊ|_|ə|'|r|ɛ|k|t|ə|s}}; meaning "[[:wikt:erectus|upright]] man") is an extinct [[species]] of [[archaic human]] from the [[Pleistocene]], with its earliest occurrence about 2&nbsp;million years ago.<ref name=Herries>{{cite journal | vauthors = Herries AI, Martin JM, Leece AB, Adams JW, Boschian G, Joannes-Boyau R, Edwards TR, Mallett T, Massey J, Murszewski A, Neubauer S, Pickering R, Strait DS, Armstrong BJ, Baker S, Caruana MV, Denham T, Hellstrom J, Moggi-Cecchi J, Mokobane S, Penzo-Kajewski P, Rovinsky DS, Schwartz GT, Stammers RC, Wilson C, Woodhead J, Menter C | display-authors = 6 | title = Contemporaneity of ''Australopithecus'', ''Paranthropus'', and early ''Homo erectus'' in South Africa | journal = Science | volume = 368 | issue = 6486 | pages = eaaw7293 | date = April 2020 | pmid = 32241925 | doi = 10.1126/science.aaw7293 | doi-access = free }}</ref> Its specimens are among the first recognizable members of the genus ''[[Homo]]''.  
'''''Homo erectus''''' ({{IPAc-en|ˌ|h|oʊ|m|oʊ|_|ə|'|r|ɛ|k|t|ə|s}} {{lit|[[:wikt:erectus|upright]] man}}) is an [[extinction|extinct]] [[species]] of [[Homo|archaic human]] from the [[Pleistocene]], spanning nearly 2 million years. It is the first human species to evolve a humanlike [[body plan]] and [[human gait|gait]], to [[early expansions of hominins out of Africa|leave Africa]] and colonize Asia and Europe, and to [[Control of fire by early humans|wield fire]]. Some populations of ''H. erectus'' were ancestors of later human species, including ''[[Homo heidelbergensis|H. heidelbergensis]]'' — the [[last common ancestor]] of [[human|modern humans]], [[Neanderthal]]s, and [[Denisovan]]s. As such a widely distributed species both geographically and temporally, ''H. erectus'' anatomy varies considerably. Subspecies are sometimes recognized: ''[[Java Man|H. e. erectus]]'', ''[[Peking Man|H. e. pekinensis]]'', ''[[Solo Man|H. e. soloensis]]'', ''[[Homo ergaster|H. e. ergaster]]'', ''[[Dmanisi hominins|H. e. georgicus]]'', and ''[[Tautavel Man|H. e. tautavelensis]]''.


Several human species, such as ''[[H. heidelbergensis]]'' and ''[[H. antecessor]]'', appear to have evolved from ''H.&nbsp;erectus'', and [[Neanderthal]]s, [[Denisovan]]s, and [[modern humans]] are in turn generally considered to have evolved from ''H. heidelbergensis''.<ref>{{cite journal | vauthors = Dembo M, Radovčić D, Garvin HM, Laird MF, Schroeder L, Scott JE, Brophy J, Ackermann RR, Musiba CM, de Ruiter DJ, Mooers AØ, Collard M | display-authors = 6 | title = The evolutionary relationships and age of Homo naledi: An assessment using dated Bayesian phylogenetic methods | journal = Journal of Human Evolution | volume = 97 | pages = 17–26 | date = August 2016 | pmid = 27457542 | doi = 10.1016/j.jhevol.2016.04.008 | hdl = 2164/8796 | hdl-access = free }}</ref> ''H.&nbsp;erectus'' was the first human ancestor to spread throughout [[Eurasia]], with a continental range extending from the [[Iberian Peninsula]] to [[Java]]. Asian populations of ''H.&nbsp;erectus'' may be ancestral to ''[[H.&nbsp;floresiensis]]''<ref>{{cite journal | vauthors = van den Bergh GD, Kaifu Y, Kurniawan I, Kono RT, Brumm A, Setiyabudi E, Aziz F, Morwood MJ | display-authors = 6 | title = Homo floresiensis-like fossils from the early Middle Pleistocene of Flores | journal = Nature | volume = 534 | issue = 7606 | pages = 245–248 | date = June 2016 | pmid = 27279221 | doi = 10.1038/nature17999 | bibcode = 2016Natur.534..245V | s2cid = 205249218 | author5-link = Adam Brumm }}</ref> and possibly to ''[[Homo luzonensis|H.&nbsp;luzonensis]]''.<ref>{{cite journal | vauthors = Détroit F, Mijares AS, Corny J, Daver G, Zanolli C, Dizon E, Robles E, Grün R, Piper PJ | display-authors = 6 | title = A new species of Homo from the Late Pleistocene of the Philippines | journal = Nature | volume = 568 | issue = 7751 | pages = 181–186 | date = April 2019 | pmid = 30971845 | doi = 10.1038/s41586-019-1067-9 | s2cid = 106411053 | bibcode = 2019Natur.568..181D | url = https://hal.archives-ouvertes.fr/hal-02296712/file/Detroit_%26_al_2019_Nature_postprint.pdf }}</ref> The last known population of ''H.&nbsp;erectus'' is ''[[Solo Man|H. e. soloensis]]'' from Java, around 117,000–108,000 years ago.<ref name=Rizal>{{cite journal | vauthors = Rizal Y, Westaway KE, Zaim Y, van den Bergh GD, Bettis EA, Morwood MJ, Huffman OF, Grün R, Joannes-Boyau R, Bailey RM, Westaway MC, Kurniawan I, Moore MW, Storey M, Aziz F, Zhao JX, Sipola ME, Larick R, Zonneveld JP, Scott R, Putt S, Ciochon RL | display-authors = 6 | title = Last appearance of Homo erectus at Ngandong, Java, 117,000-108,000 years ago | journal = Nature | volume = 577 | issue = 7790 | pages = 381–385 | date = January 2020 | pmid = 31853068 | doi = 10.1038/s41586-019-1863-2 | s2cid = 209410644 }}</ref>
The species was first [[species description|described]] by [[Eugène Dubois]] in 1893 as "''[[Anthropopithecus|Pithecanthropus]] erectus''" using a skullcap, [[molar (tooth)|molar]], and [[femur]] from [[Java]], Indonesia. Further discoveries around East Asia were used to contend that humanity evolved out of Asia. Based on [[historical race concepts]], it was argued that local ''H. erectus'' populations evolved directly into local modern human populations ([[multiregional origin of modern humans|polycentrism]]) rather than all humanity sharing a single [[early modern human|anatomically modern]] ancestor ([[monogenism]]). As the fossil record improved over the mid-to-late 20th century, [[Early expansions of hominins out of Africa|"Out of Africa"]] theory and monogenism became the consensus.


''H. erectus'' had a more modern gait and body proportions, and was the first human species to have exhibited a flat face, prominent nose, and possibly sparse body hair coverage. Though the species' brain size certainly exceeds that of ancestor species, capacity varied widely depending on the population. In earlier populations, brain development seemed to cease early in childhood, suggesting that offspring were largely self-sufficient at birth, thus limiting cognitive development through life. ''H. erectus'' was an [[apex predator]];<ref name="Ben-Dor2021">{{cite journal | vauthors = Ben-Dor M, Sirtoli R, Barkai R | title = The evolution of the human trophic level during the Pleistocene | journal = American Journal of Physical Anthropology | volume = 175 | issue = Suppl 72 | pages = 27–56 | date = August 2021 | pmid = 33675083 | doi = 10.1002/ajpa.24247 | doi-access = free }}</ref> sites generally show consumption of medium to large animals, such as [[bovine]]s or [[elephant]]s, and suggest the development of predatory behavior and coordinated hunting. ''H. erectus'' is associated with the [[Acheulean]] stone tool [[industry (archaeology)|industry]], and is postulated to have been the earliest human ancestor capable of using fire,<ref>{{Cite journal |last1=Zohar |first1=Irit |last2=Alperson-Afil |first2=Nira |last3=Goren-Inbar |first3=Naama |last4=Prévost |first4=Marion |last5=Tütken |first5=Thomas |last6=Sisma-Ventura |first6=Guy |last7=Hershkovitz |first7=Israel |last8=Najorka |first8=Jens |date=2022-11-14 |title=Evidence for the cooking of fish 780,000 years ago at Gesher Benot Ya'aqov, Israel |url=https://www.nature.com/articles/s41559-022-01910-z |journal=Nature Ecology & Evolution |volume=6 |issue=12 |pages=2016–2028 |language=en |doi=10.1038/s41559-022-01910-z |pmid=36376603 |s2cid=253522354 |issn=2397-334X}}</ref> hunting and gathering in coordinated groups, caring for injured or sick group members, and possibly seafaring and art (though examples of art are controversial, and are otherwise rudimentary and few and far between).
The typical skull has a pronounced brow ridge, a protruding jaw, and large teeth. The bones are much thicker than in modern humans. East Asian populations normally have an even more robust skeleton and larger brain volume — averaging {{cvt|1000|cc}}. Western ''H. erectus'' brain volume could be as low as {{cvt|546|cc}} in ''H. e. georgicus''. ''H. erectus'' probably had a faster apelike [[human height|growth trajectory]], lacking the [[altriciality|extended childhood]] required for [[language acquisition]]. Reconstructed adult body dimensions range from {{cvt|141–167|cm|ftin|0}} in height and about {{cvt|50|kg}} in weight.


''H. erectus'' males and females may have been roughly the same size as each other (i.e. exhibited reduced [[sexual dimorphism]]), which could indicate [[monogamy]] in line with general trends exhibited in primates. Size, nonetheless, ranged widely from {{cvt|146–185|cm|ftin|sigfig=1}} in height and {{cvt|40–68|kg}} in weight. It is unclear if ''H. erectus'' was anatomically capable of speech, though it is postulated they communicated using some [[Origin of language|proto-language]].
''H. erectus'' invented the [[Acheulean]] [[industry (archaeology)|tool industry]], a major innovation of large, heavy-duty [[stone tools]]. These may have been used in butchery, vegetable processing, and woodworking of [[spear]]s and [[digging stick]]s. ''H. erectus'' was a major predator of large herbivores on the expanding savannas during the [[Quaternary glaciation]]. The species is usually characterized as the first [[hunter-gatherer]] and the first to practice [[sexual division of labor]]. Fire usage and cave habitation were probably not important aspects of daily life. Similarly, ''H. erectus'' may not have often ventured into colder regions or cooked meat. The last known occurrence of ''H. erectus'' is 108,000 to 117,000 years ago (''H. e. soloensis'') in Southeast Asia, until the last savannas in the region gave way to jungle.


==Taxonomy==
==Taxonomy==
===Naming===
{{See also|Human taxonomy}}
{{see|Java Man}}
===Research history===
[[File:Pithecanthropus erectus-PeterMaas Naturalis.jpg|thumb|left|[[Java Man]] at [[Naturalis Biodiversity Center|Naturalis]]]]
 
Contrary to the view [[Charles Darwin]] expressed in his 1871 book ''[[Descent of Man]]'', many late-19th century evolutionary naturalists postulated that Asia, not Africa, was the birthplace of humankind as it is midway between Europe and America, providing optimal dispersal routes throughout the world (the [[Out of Asia theory]]). Among these was German naturalist [[Ernst Haeckel]], who argued that the first human species evolved on the now-disproven hypothetical continent "[[Lemuria (continent)|Lemuria]]" in what is now Southeast Asia, from a species he termed "''[[Pithecanthropus]] alalus''" ("speechless apeman").{{sfn|Theunissen|2012|loc=p. 6}} "Lemuria" had supposedly sunk below the [[Indian Ocean]], so no fossils could be found to prove this. Nevertheless, Haeckel's model inspired Dutch scientist [[Eugène Dubois]] to journey to the [[Dutch East Indies]]. Because no directed expedition had ever discovered human fossils (the few known had all been discovered by accident), and the economy was strained by the [[Long Depression]], the Dutch government refused to fund Dubois. In 1887, he enlisted in the [[Royal Netherlands Indies Army|Dutch East India Army]] as a medical officer, and was able to secure a post in 1887 in the Indies to search for his "[[missing link (human evolution)|missing link]]" in his spare time.{{sfn|Theunissen|2012|loc=p. 33}} On [[Java]], he found a skullcap in 1891 and a [[femur]] in 1892 ([[Java Man]]) dating to the [[late Pliocene]] or [[early Pleistocene]] at the [[Trinil]] site along the [[Solo River]], which he named ''Pithecanthropus erectus'' ("upright apeman") in 1893. He attempted unsuccessfully to convince the European scientific community that he had found an upright-walking ape-man. Given few fossils of ancient humans had even been discovered at the time, they largely dismissed his findings as a malformed non-human ape.<ref name=HsiaoPei2014>{{cite journal | vauthors = Yen HP | title = Evolutionary Asiacentrism, Peking man, and the origins of sinocentric ethno-nationalism | journal = Journal of the History of Biology | volume = 47 | issue = 4 | pages = 585–625 | year = 2014 | pmid = 24771020 | doi = 10.1007/s10739-014-9381-4 | s2cid = 23308894 }}</ref>
[[File:Hypothetical sketch of the monophylitic origin and of the diffusion of the 12 varieties of men from Lemuria over the earth LCCN2014649358.jpg|thumb|left|upright=1.5|In 1868, [[Ernst Haeckel]] suggested early humans dispersed from the now-disproven hypothetical continent "[[Lemuria (continent)|Lemuria]]".<ref name=HsiaoPei2014/><ref>{{cite book|first=E.|last=Haeckel|author-link=Ernst Haeckel|orig-year=1868|year=1876|trans-title=The history of creation|title=Natürliche Schöpfungsgeschichte|translator=Lankester, E. R.|publisher=Henry S. King & Co.|url=https://archive.org/details/historyofcreatio76hist/page/360|page=361}}</ref>]]
 
While [[Charles Darwin]] had hypothesized in his 1871 ''[[Descent of Man]]'' that humans most likely evolved in Africa,{{efn-num|Darwin's work mainly aimed to demonstrate that his [[scientific theory|theory]] of [[common descent]] with modification by [[natural selection|natural]] and [[sexual selection in humans|sexual selection]] applied to humans, "The sole object of this work is to consider, firstly, whether man, like every other species, is descended from some pre-existing form; secondly, the manner of his development; and thirdly, the value of the [[social Darwinism|differences]] between the so-called [[historical race concepts|races of man]]."<ref>{{cite book|first=C.|last=Darwin|author-link=Charles Darwin|year=1871|url=https://www.gutenberg.org/cache/epub/34967/pg34967-images.html#Page_3|title=The Descent of Man and Selection in Relation to Sex|publisher=John Murray|page=3}}</ref>}} many late-19th century evolutionary naturalists postulated that Asia was the birthplace of humankind. Asia is midway between all continents via land routes or short sea crossings, providing optimal dispersal routes throughout the world. Among the major proponents of "Out of Asia" theory was [[Ernst Haeckel]], who argued that the first human species (which he speculatively named ''Homo primigenius'') evolved on a now-disproven hypothetical continent "[[Lemuria (continent)|Lemuria]]" from a species he termed ''[[Anthropopithecus|Pithecanthropus alalus]]'' (speechless ape-man). "Lemuria" had supposedly sunk below the [[Indian Ocean]], accounting for the lack of fossil evidence.<ref name=HsiaoPei2014/>


The significance of these fossils would not be realized until the 1927 discovery of what Canadian paleoanthropologist [[Davidson Black]] called "''Sinanthropus pekinensis''" (Peking Man) at the [[Zhoukoudian]] cave near [[Beijing]], China. Black lobbied across North America and Europe for funding to continue excavating the site,{{sfn|Sigmon|1981|loc=p. 64}} which has since become the most productive ''H. erectus'' site in the world.<ref name=Yang2014>{{cite book| vauthors = Yang L |year=2014|chapter=Zhoukoudian: Geography and Culture|title=Encyclopedia of Global Archaeology|pages=7961–7965|publisher=Springer Science+Business Media|isbn=978-1-4419-0466-9|doi=10.1007/978-1-4419-0465-2_1899}}</ref> Continued interest in Java led to further ''H. erectus'' fossil discoveries at Ngandong ([[Solo Man]]) in 1931, [[Mojokerto child|Mojokerto]] (Java Man) in 1936, and [[Sangiran]] (Java Man) in 1937. The Sangiran site yielded the best preserved Java Man skull.<ref>{{cite book| vauthors = Ciochon RL, Huffman OF |year=2014|chapter=Java Man|title=Encyclopedia of Global Archaeology|pages=4182–4188| veditors = Smith C |doi=10.1007/978-1-4419-0465-2_712|isbn=978-1-4419-0426-3|s2cid=241324984 }}</ref> German paleoanthropologist [[Franz Weidenreich]] provided much of the detailed description of the Chinese specimens in several monographs. The original specimens were lost during the [[Second Sino-Japanese War]] after an attempt to smuggle them out of China for safekeeping. Only [[plaster cast|casts]] remain.
Dutch scientist [[Eugène Dubois]] joined the [[Royal Netherlands East Indies Army]] to search for the "[[missing link (human evolution)|missing link]]" of human evolution in [[Java]].{{efn-num|Dubois was studying the anatomy and evolution of the [[larynx]] in [[vertebrate]]s in [[Amsterdam]] with German anatomist [[Max Fürbringer]], but fed up with both the research and Fürbringer, he decided to drop everything and board the [[SS Prinses Amalia]] to the Dutch East Indies on 29 October 1887. Dubois said his interest in human evolution began in secondary school (the State [[Hogere Burgerschool]]), where he heard of lectures given by [[Carl Vogt]] in 1868.{{sfn|Theunissen|1989|loc=pp. 27–30}}}} At the [[Trinil]] site, his team found a skullcap and [[molar (tooth)|molar]] in 1891, and a [[femur]] in 1892 ([[Java Man]]), which he named "''Pithecanthropus erectus''" in 1893.{{sfn|Theunissen|1989|loc=54–59}} He attempted to convince the European scientific community that he had found an upright-walking ape-man dating to the [[late Pliocene]] or [[Early Pleistocene]]; they dismissed his findings as some kind of non-human ape.<ref name=HsiaoPei2014>{{cite journal |first=Y. |last=Hsiao-Pei |year=2014 |title=Evolutionary Asiacentrism, Peking Man, and the Origins of Sinocentric Ethno-Nationalism |journal=Journal of the History of Biology |volume=47 |issue=4 |pages=585–625 |doi=10.1007/s10739-014-9381-4 |pmid=24771020 |s2cid=23308894}}</ref>


Similarities between Java Man and Peking Man led [[Ernst Mayr]] to rename both as ''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]] the most prominent among them—theorized that humans' earliest ancestors were African. Darwin had pointed out that chimpanzees and gorillas, humans' closest relatives, evolved and exist only in Africa.<ref>{{cite book | vauthors = Darwin CR |title=The Descent of Man and Selection in Relation to Sex|url= https://archive.org/details/descentmanandse03darwgoog |publisher=John Murray |year=1871 |isbn=978-0-8014-2085-6}}</ref> Darwin did not include orangutans among the great apes of the Old World, likely because he thought of orangutans as primitive humans rather than apes.<ref>{{Cite journal |last1=van Wyhe |first1=John |last2=Kjærgaard |first2=Peter C. |date=2015-06-01 |title=Going the whole orang: Darwin, Wallace and the natural history of orangutans |url=https://www.sciencedirect.com/science/article/pii/S1369848615000370 |journal=Studies in History and Philosophy of Science Part C: Studies in History and Philosophy of Biological and Biomedical Sciences |language=en |volume=51 |pages=53–63 |doi=10.1016/j.shpsc.2015.02.006 |pmid=25861859 |s2cid=20089470 |issn=1369-8486}}</ref> While Darwin considered Africa as the most probable birthplace of human ancestors, he also made the following statement about the geographic location of human origins in his book ''The Descent of Man, and Selection in Relation to Sex'': "... it is useless to speculate on this subject; for two or three anthropomorphous apes, one the Dryopithecus …, existed in Europe during the Miocene age; and since so remote a period the earth has certainly undergone many great revolutions, and there has been ample time for migration on the largest scale.” (1889, pp. 155-156).
{{multiple image|total_width = 300
|image1=Homo erectus Sangiran 17 IMG 5644 BMNH.jpg
|image2=Sinanthropus Skull XII.png
|footer=[[Franz Weidenreich]] and [[Gustav Heinrich Ralph von Koenigswald|Ralph von Koenigswald]] argued [[Java Man]] was an ancient human using fossils like [[Sangiran]] (left) and [[Peking Man]] (right)
}}


In 1949, the species was reported in [[Swartkrans]] Cave, South Africa, by South African paleoanthropologists [[Robert Broom]] and [[John Talbot Robinson]], who described it as "''Telanthropus capensis''".<ref>{{cite journal | vauthors = Curnoe D | title = A review of early Homo in southern Africa focusing on cranial, mandibular and dental remains, with the description of a new species (Homo gautengensis sp. nov.) | journal = Homo | volume = 61 | issue = 3 | pages = 151–177 | date = June 2010 | pmid = 20466364 | doi = 10.1016/j.jchb.2010.04.002 }}</ref> ''Homo'' fossils have also been reported from nearby caves, but their species designation has been a tumultuous discussion. A few North African sites have additionally yielded ''H. erectus'' remains, which at first were classified as "''Atlantanthropus mauritanicus''" in 1951.{{sfn|Sigmon|1981|loc=p. 231}} Beginning in the 1970s, propelled most notably by [[Richard Leakey]], more were being unearthed in East Africa predominantly at the [[Koobi Fora]] site, Kenya, and [[Olduvai Gorge]], Tanzania.{{sfn|Sigmon|1981|loc=p. 193}}
Dubois argued that "''P. erectus''" was a [[gibbon]]-like ape which was the precursor to a more familiar human body plan, but in the 1930s, German-American anatomist [[Franz Weidenreich]] noticed a striking similarity with ancient human remains recently being unearthed in China ([[Peking Man]], "''Sinanthropus pekinensis''").{{sfn|Theunissen|1989|loc=p. 156}}{{sfn|Boaz|Ciochon|2004|loc=pp. 59–60}} This characterization became better supported as German-Dutch palaeontologist [[Gustav Heinrich Ralph von Koenigswald]] discovered more Indonesian ancient human remains over the decade at [[Mojokerto child|Mojokerto]], [[Sangiran]], and [[Solo Man|Ngandong]].{{efn-num|Dubois never accepted that the Java Man was a type of human, and continued to fight Weidenreich and von Koenigswald until his death in 1940. Sir [[Arthur Keith]] described Dubois in an obituary notice as, "an idealist, his ideas being so firmly held that his mind tended to bend facts rather than alter his ideas to fit them."{{sfn|Theunissen|1989|loc=pp. 159–165}}}}{{sfn|Boaz|Ciochon|2004|loc=pp. 60–62}}<ref>{{cite journal |first1=G. H. R. |last1=von Koenigswald |author-link1=Gustav Heinrich Ralph von Koenigswald |first2=F. |last2=Weidenreich |author2-link=Franz Weidenreich |year=1939 |title=The Relationship between ''Pithecanthropus'' and ''Sinanthropus'' |journal=Nature |volume=144 |issue=3657 |pages=926–929 |doi=10.1038/144926a0|bibcode=1939Natur.144..926V }}</ref> Weidenreich believed that they were the direct ancestors of the local modern human [[human taxonomy#Homo sapiens subspecies|''Homo sapiens'' subspecies]], in accord with [[historical race concepts]] ([[multiregional origin of modern humans|polycentricism]]) — that is, Peking Man was the direct ancestor of specifically [[Chinese people]], and Java Man of [[Aboriginal Australian]]s.<ref name="Weidenreich1940">{{cite journal |first=F. |last=Weidenreich |author-link=Franz Weidenreich |year=1940 |title=Some problems dealing with ancient man |journal=American Anthropologist |volume=42 |issue=3 |pages=375–383 |doi=10.1525/aa.1940.42.3.02a00010}}</ref><ref name=Mayr1950/> As the significance of racial distinction diminished with the development of [[modern evolutionary synthesis]], many fossil human species and genera around Asia, Africa, and Europe (including "''Pithecanthropus''" and "''Sinanthropus''") were reclassified as subspecies of ''Homo erectus''.<ref name=Schwartz2010/>{{sfn|Boaz|Ciochon|2004|loc=pp. 70–73}}


Archaic human fossils unearthed across Europe used to be assigned to ''H. erectus'', but have since been separated as ''[[Homo heidelbergensis|H. heidelbergensis]]'' as a result of British physical anthropologist [[Chris Stringer]]'s work.<ref name=Lumley2015>{{cite journal| vauthors = de Lumley MA |year=2015|title=L'homme de Tautavel. Un ''Homo erectus'' européen évolué. ''Homo erectus tautavelensis''|trans-title=Tautavel Man. An evolved European ''Homo erectus''. ''Homo erectus tautavelensis''|language=fr|journal=L'Anthropologie|volume=119|issue=3|pages=342–344|doi=10.1016/j.anthro.2015.06.001}}</ref>
During the late 20th century, some of the oldest ''H. erectus'' fossils were discovered across Africa, the first being Kenyan archeologist [[Louis Leakey]]'s [[Chellean Man|Olduvai Hominin 9]] in 1960.{{sfn|Boaz|Ciochon|2004|loc=pp. 68–72}} As the human fossil record expanded, the "Out of Africa" theory and [[monogenism]] became the consensus: that all modern humans share a fully [[early modern human|anatomically modern]] common ancestor. ''H. erectus'' is now generally considered to be an African species which later dispersed across Eurasia, with later African populations giving rise to the modern human lineage.{{sfn|Boaz|Ciochon|2004|loc=pp. 151–153}}


===Evolution===
===Subspecies===
{{see|Early human expansions out of Africa}}
By the middle of the 20th century, [[human taxonomy]] was in turmoil, with many poorly defined species and genera described across Europe, Asia, and Africa, which exaggerated the differences among them.{{sfn|Boaz|Ciochon|2004|loc=p. 65}} In 1940, Weidenreich was the first to suggest reclassifying "''Sinanthropus pekinensis''" and "''Pithecanthropus erectus''" as subspecies of ''H. erectus''.<ref name="Weidenreich1940"/> In 1950, German-American evolutionary biologist [[Ernst Mayr]] entered this field. Surveying a "bewildering diversity of names" and many proposals for consolidation, he decided to reclassify human fossils into three species of ''[[Homo]]'': "''H. transvaalensis''" (the [[australopithecine]]s), ''H. erectus'' (including "''Sinanthropus''", "''Pithecanthropus''", and various other Asian, African, and European taxa), and ''H. sapiens'' (including anything younger than ''H. erectus'', such as modern humans and Neanderthals). Mayr defined these species as a sequential lineage, each evolving into the next ([[chronospecies]]).<ref name=Mayr1950>{{cite journal |last=Mayr |first=E. |author-link=Ernst Mayr |year=1950 |title=Taxonomic categories in fossil hominids |journal=Cold Spring Harbor Symposia on Quantitative Biology |volume=15 |pages=109–118 |doi=10.1101/SQB.1950.015.01.013|pmid=14942702 }}</ref> Though later Mayr changed his opinion on the australopithecines (recognizing ''[[Australopithecus]]''), his more [[lumpers and splitters|conservative]] view of [[archaic human]] diversity became widely adopted in the subsequent decades.<ref name=Schwartz2010>{{cite journal |last1=Schwartz |first1=J. H. |author1-link=Jeffrey H. Schwartz |last2=Tattersall |first2=I. |author2-link=Ian Tattersall |year=2010 |title=Fossil evidence for the origin of ''Homo sapiens'' |journal=American Journal of Physical Anthropology |volume=143 |issue=S51 |pages=96–98; 101–103 |doi=10.1002/ajpa.21443 |pmid=21086529 |bibcode=2010AJPA..143S..94S |doi-access=free}}</ref>
{{Human timeline}}
[[File:Carte hachereaux.jpg|thumb|upright=1.5|Map of the distribution of Middle Pleistocene ([[Acheulean]]) [[Cleaver (tool)|cleaver]] finds]]
It has been proposed that ''H. erectus'' evolved from ''[[Homo habilis|H. habilis]]'' about 2 Mya, though this has been called into question because they coexisted for at least a half a million years. Alternatively, a group of ''H. habilis'' may have been [[reproductively isolated]], and only this group developed into ''H. erectus'' ([[cladogenesis]]).<ref name=Spoor2007/>


Because the earliest remains of ''H. erectus'' are found in both Africa and East Asia (in China as early as 2.1 Mya,<ref name = "Zhu_2018">{{cite journal | vauthors = Zhu Z, Dennell R, Huang W, Wu Y, Qiu S, Yang S, Rao Z, Hou Y, Xie J, Han J, Ouyang T | display-authors = 6 | title = Hominin occupation of the Chinese Loess Plateau since about 2.1 million years ago | journal = Nature | volume = 559 | issue = 7715 | pages = 608–612 | date = July 2018 | pmid = 29995848 | doi = 10.1038/s41586-018-0299-4 | s2cid = 49670311 | author10 = Han Jiangwei ({{lang|zh-Hans|韩江伟}}) | author11 = Ouyang Tingping ({{lang|zh-Hans|欧阳婷萍}}) | author4 = Wu Yi ({{lang|zh-Hans|吴翼}}) | author5 = Qiu Shifan ({{lang|zh-Hans|邱世藩}}) | author6 = Yang Shixia ({{lang|zh-Hans|杨石霞}}) | author7 = Rao Zhiguo ({{lang|zh-Hans|饶志国}}) | author3 = Huang Weiwen ({{lang|zh-Hans|黄慰文}}) | bibcode = 2018Natur.559..608Z | author8 = Hou Yamei ({{lang|zh-Hans|侯亚梅}}) | author9 = Xie Jiubing ({{lang|zh-Hans|谢久兵}}) | name-list-style = vanc }}</ref><ref>{{Cite journal| vauthors = Barras C |year=2018|title=Tools from China are oldest hint of human lineage outside Africa|url=https://www.nature.com/articles/d41586-018-05696-8 |journal=Nature|doi=10.1038/d41586-018-05696-8|s2cid=188286436|issn=0028-0836}}</ref><ref>{{cite journal | vauthors = Hao L, Chao Rong L, Kuman K | year = 2017 | title = Longgudong, an Early Pleistocene site in Jianshi, South China, with stratigraphic association of human teeth and lithics | journal = Science China Earth | volume = 60 | issue = 3| pages = 452–462 | doi = 10.1007/s11430-016-0181-1 | bibcode = 2017ScChD..60..452L | s2cid = 132479732 }}</ref> in South Africa 2.04 Mya<ref name=Herries/><ref>{{cite web |title=Our direct human ancestor Homo erectus is older than we thought |url=https://www.eurekalert.org/pub_releases/2020-04/uoj-odh040120.php |website=EurekAlert |publisher=[[American Association for the Advancement of Science|AAAS]]}}</ref>), it is debated where ''H. erectus'' evolved. A 2011 study suggested that it was ''H. habilis'' who reached West Asia from Africa, that early ''H. erectus'' developed there, and that 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]]), eventually evolving into modern humans in Africa.<ref>{{cite journal | vauthors = Ferring R, Oms O, Agustí J, Berna F, Nioradze M, Shelia T, Tappen M, Vekua A, Zhvania D, Lordkipanidze D | display-authors = 6 | title = Earliest human occupations at Dmanisi (Georgian Caucasus) dated to 1.85-1.78 Ma | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 108 | issue = 26 | pages = 10432–10436 | date = June 2011 | pmid = 21646521 | pmc = 3127884 | doi = 10.1073/pnas.1106638108 | doi-access = free | bibcode = 2011PNAS..10810432F }}</ref><ref>{{cite journal |doi=10.1016/j.quascirev.2010.04.012|date=June 2011 | vauthors = Augusti J, Lordkipanidze D |title=How "African" was the early human dispersal out of Africa?|volume=30|issue=11–12 |pages=1338–1342 |journal=Quaternary Science Reviews|bibcode=2011QSRv...30.1338A }}</ref> Others have suggested that ''H. erectus''/''H. ergaster'' developed in Africa, where it eventually evolved into modern humans.<ref>{{cite journal| vauthors = Rightmire GP |title= Human Evolution in the Middle Pleistocene: The Role of ''Homo heidelbergensis'' |year=1998 |journal=Evolutionary Anthropology|doi=10.1002/(sici)1520-6505(1998)6:6<218::aid-evan4>3.0.co;2-6 |volume=6 |issue=6 |pages=218–227|s2cid= 26701026 }}</ref><ref name="Asfawpmid11907576">{{cite journal | vauthors = Asfaw B, Gilbert WH, Beyene Y, Hart WK, Renne PR, WoldeGabriel G, Vrba ES, White TD | display-authors = 6 | title = Remains of Homo erectus from Bouri, Middle Awash, Ethiopia | journal = Nature | volume = 416 | issue = 6878 | pages = 317–320 | date = March 2002 | pmid = 11907576 | doi = 10.1038/416317a | s2cid = 4432263 | bibcode = 2002Natur.416..317A }}</ref>
{{blockquote|text=...never more than one species of man existed on the earth at any one time... If fossils of [[African pygmies|Congo pygmies]] and of [[Tutsi|Watusi]] were to be found in the same deposit by a paleontologist, a million years hence, he might well think that they belonged to two different species.|author=[[Ernst Mayr]], 1950<ref name=Mayr1950/>}}


''H. erectus'' had reached [[Sangiran]], Java, by 1.8 Mya,<ref>{{Cite journal |last1=Husson |first1=Laurent |last2=Salles |first2=Tristan |last3=Lebatard |first3=Anne-Elisabeth |last4=Zerathe |first4=Swann |last5=Braucher |first5=Régis |last6=Noerwidi |first6=Sofwan |last7=Aribowo |first7=Sonny |last8=Mallard |first8=Claire |last9=Carcaillet |first9=Julien |last10=Natawidjaja |first10=Danny H. |last11=Bourlès |first11=Didier |last12=ASTER team |last13=Aumaitre |first13=Georges |last14=Bourlès |first14=Didier |last15=Keddadouche |first15=Karim |date=2022-11-08 |title=Javanese Homo erectus on the move in SE Asia circa 1.8 Ma |journal=Scientific Reports |language=en |volume=12 |issue=1 |pages=19012 |doi=10.1038/s41598-022-23206-9 |issn=2045-2322 |pmc=9643487 |pmid=36347897|bibcode=2022NatSR..1219012H }}</ref> and a second and distinct wave of ''H. erectus'' had colonized [[Zhoukoudian]], China, about 780 kya. Early teeth from Sangiran are bigger and more similar to those of basal (ancestral) Western ''H. erectus'' and ''H. habilis'' than to those of the derived Zhoukoudian ''H. erectus''. However, later Sangiran teeth seem to reduce in size, which could indicate a secondary colonization event of Java by the Zhoukoudian or some closely related population.<ref>{{cite journal | vauthors = Zaim Y, Ciochon RL, Polanski JM, Grine FE, Bettis EA, Rizal Y, Franciscus RG, Larick RR, Heizler M, Eaves KL, Marsh HE | display-authors = 6 | title = New 1.5 million-year-old Homo erectus maxilla from Sangiran (Central Java, Indonesia) | journal = Journal of Human Evolution | volume = 61 | issue = 4 | pages = 363–376 | date = October 2011 | pmid = 21783226 | doi = 10.1016/j.jhevol.2011.04.009 }}</ref>
In the 1970s, as [[population genetics]] was being formulated, the anatomical variation of ''H. erectus'' across its wide geographic and temporal range (the basis for the subspecies distinctions) became better understood as [[cline (biology)|clines]] — different populations which attained some anatomical regionality but were not reproductively isolated.{{sfn|Boaz|Ciochon|2004|loc=pp. 151–153}} In general, subspecies names for ''H. erectus'' are now used for convenience to indicate time and region rather than specific anatomical trends.<ref name=Anton2002>{{cite journal |first=S. C. |last=Antón |year=2002 |title=Evolutionary significance of cranial variation in Asian ''Homo erectus'' |journal=American Journal of Physical Anthropology |volume=118 |issue=4 |page=302 |doi=10.1002/ajpa.10091|pmid=12124912 |bibcode=2002AJPA..118..301A }}</ref>


===Subspecies===
{{blockquote|text=...to paleontologists in general, subspecies are epiphenomena which do not merit the attention paid to species... The pursuit of subspecies in the fossil record is at best fraught with difficulty, and is more probably futile.|author=[[Ian Tattersall]], 1986<ref>{{cite journal |first=I. |last=Tattersall |author-link=Ian Tattersall |year=1986 |title=Species recognition in human paleontology |journal=Journal of Human Evolution |volume=15 |issue=3 |page=168 |doi=10.1016/S0047-2484(86)80043-4|bibcode=1986JHumE..15..165T }}</ref>}}
"[[Wushan Man]]" was proposed as ''Homo erectus wushanensis'', but is now thought to be based upon fossilized fragments of an extinct non-hominin ape.<ref name="Ciochon">{{cite journal | vauthors = Ciochon RL | title = The mystery ape of Pleistocene Asia | journal = Nature | volume = 459 | issue = 7249 | pages = 910–911 | date = June 2009 | pmid = 19536242 | doi = 10.1038/459910a | s2cid = 205047272 | doi-access = free | bibcode = 2009Natur.459..910C }}</ref>
 
{{multiple image
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| image1 = Ergaster Skull.png
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| image2 = Erectus Skull.png
| caption2 =  
| footer = Reconstructions of ''[[Homo ergaster|H. e. ergaster]]'' ([[KNM ER 3733]]) left and ''[[Peking Man|H. e. pekinensis]]'' right
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The more commonly used subspecies (if any are used)<!-- many times subspecies are not used --> are:{{sfn|Antón|2003|loc=p. 153}}
*''[[Java Man|H. e. erectus]]'' for earlier Indonesian fossils
*''[[Peking Man|H. e. pekinensis]]'' for Chinese fossils
*''[[Solo Man|H. e. soloensis]]'' for the latest-surviving Indonesian fossils
*''[[Homo ergaster|H. e. ergaster]]'' for African fossils
*''[[Dmanisi hominins|H. e. georgicus]]'' for an early group of fossils from [[Prehistoric Georgia|Georgia]]<ref name=Rightmire2006/>
*''[[Tautavel Man|H. e. tautavelensis]]'' for Western European fossils<ref name=Lumley2015>{{cite journal |first=M.-A. |last=de Lumley |year=2015 |title=L'homme de Tautavel. Un ''Homo erectus'' européen évolué. ''Homo erectus tautavelensis'' |trans-title=Tautavel Man. An evolved European ''Homo erectus''. ''Homo erectus tautavelensis'' |language=fr |journal=L'Anthropologie |volume=119 |issue=3 |pages=342–344 |doi=10.1016/j.anthro.2015.06.001}}</ref>


Since its discovery in 1893 ([[Java Man]]), there has been a trend in paleoanthropology of reducing the number of proposed species of ''Homo'', to the point where ''H. erectus'' includes all early ([[Lower Paleolithic]]) forms of ''Homo'' sufficiently derived from ''[[Homo habilis|H. habilis]]'' and
The ancient Georgia fossils have variably been classified as ''H. e. ergaster'' (or quadrinomial ''H. e. ergaster georgicus''),<ref>{{cite journal |last1=Zollikofer |first1=C. P. E. |last2=Ponce de León |first2=M. S. |last3=Margvelashvili |first3=A. |last4=Rightmire |first4=G. Philip |last5=Lordkipanidze |first5=D. |s2cid=206554612 |year=2014 |title=Response to Comment on "A Complete Skull from Dmanisi, Georgia, and the Evolutionary Biology of Early ''Homo''" |url=https://www.science.org/doi/10.1126/science.1250056 |journal=Science |volume=344 |issue=6182 |pages=360–b |doi=10.1126/science.1250081 |pmid=24763573 |bibcode=2014Sci...344..360Z|url-access=subscription }}</ref> as their own subspecies as ''H. e. georgicus'', or as their own species ''H. georgicus''.<ref name=Rightmire2006>{{Cite journal |last1=Rightmire |first1=G. Philip |last2=Lordkipanidze |first2=David |last3=Vekua |first3=Abesalom |year=2006 |title=Anatomical descriptions, comparative studies and evolutionary significance of the hominin skulls from Dmanisi, Republic of Georgia |url=https://www.sciencedirect.com/science/article/pii/S0047248405001624 |journal=Journal of Human Evolution |volume=50 |issue=2 |page=140 |doi=10.1016/j.jhevol.2005.07.009 |pmid=16271745|bibcode=2006JHumE..50..115R |url-access=subscription }}</ref> Some authors may also elevate ''H. ergaster'',<ref>{{Cite journal |last1=Rightmire |first1=G. Philip |last2=Ponce de León |first2=Marcia S. |last3=Lordkipanidze |first3=David |last4=Margvelashvili |first4=Ann |last5=Zollikofer |first5=Christoph P. E. |year=2017 |title=Skull 5 from Dmanisi: Descriptive anatomy, comparative studies, and evolutionary significance |journal=Journal of Human Evolution |volume=104 |pages=51 |doi=10.1016/j.jhevol.2017.01.005 |pmid=28317556 |doi-access=free|bibcode=2017JHumE.104...50R }}</ref> ''H. soloensis'',<ref>{{cite journal |first1=V. |last1=Zeitoun |first2=F. |last2=Détroit |first3=D. |last3=Grimaud-Hervé |first4=H. |last4=Widianto |year=2010 |title=Solo man in question: Convergent views to split Indonesian ''Homo erectus'' in two categories |journal=[[Quaternary International]] |volume=223–224 |page=288 |doi=10.1016/j.quaint.2010.01.018 |bibcode=2010QuInt.223..281Z |s2cid=84764668 |url=https://hal.archives-ouvertes.fr/hal-02977578/file/Solo.pdf |access-date=22 May 2021 |archive-date=29 April 2021 |archive-url=https://web.archive.org/web/20210429071324/https://hal.archives-ouvertes.fr/hal-02977578/file/Solo.pdf |url-status=live}}</ref> and ''H. pekinensis'' to species level.<ref>{{cite journal |first=B. |last=Marwick |year=2009 |title=Biogeography of Middle Pleistocene hominins in mainland Southeast Asia: A review of current evidence |journal=Quaternary International |volume=202 |issue=1–2 |page=53 |doi=10.1016/j.quaint.2008.01.012|bibcode=2009QuInt.202...51M }}</ref> Fossils relegated to ''H. e. tautavelensis'' are traditionally assigned to ''[[Homo heidelbergensis|H. heidelbergensis]]''.<ref name=Lumley2015/>
distinct from early ''[[Homo heidelbergensis|H. heidelbergensis]]'' (in Africa also known as ''[[Homo rhodesiensis|H. rhodesiensis]]'').<ref name=Kaifu2005/> It is sometimes considered as a wide-ranging, polymorphous species.<ref>{{Cite journal | doi=10.1038/nature.2013.13972| title=Skull suggests three early human species were one| year=2013| vauthors = Perkins S | journal=Nature| s2cid=88314849}}</ref>


Due to such a wide range of variation, it has been suggested that the ancient ''[[Homo rudolfensis|H. rudolfensis]]'' and ''[[Homo habilis|H. habilis]]'' should be considered early varieties of ''H. erectus''.<ref name=dmanisiskull5>{{cite journal | vauthors = Lordkipanidze D, Ponce de León MS, Margvelashvili A, Rak Y, Rightmire GP, Vekua A, Zollikofer CP | title = A complete skull from Dmanisi, Georgia, and the evolutionary biology of early Homo | journal = Science | volume = 342 | issue = 6156 | pages = 326–331 | date = October 2013 | pmid = 24136960 | doi = 10.1126/science.1238484 | s2cid = 20435482 | bibcode = 2013Sci...342..326L }}</ref><ref name=National_Geographic>{{cite news | vauthors = Black R |date=17 October 2013 |title= Beautiful Skull Spurs Debate on Human History |url=https://www.nationalgeographic.com/culture/article/131017-skull-human-origins-dmanisi-georgia-erectus/ |work=[[National Geographic (magazine)|National Geographic]] |access-date=6 June 2021}}</ref> The primitive ''H. e. georgicus'' from [[Dmanisi skulls|Dmanisi]], Georgia has the smallest brain capacity of any known Pleistocene hominin (about 600 cc), and its inclusion in the species would greatly expand the range of variation of ''H. erectus'' to perhaps include species as ''H. rudolfensis'', ''[[Homo gautengensis|H. gautengensis]]'', ''[[Homo ergaster|H. ergaster]]'', and perhaps ''H. habilis''.<ref>{{cite news |title= Skull of Homo erectus throws story of human evolution into disarray |url= https://www.theguardian.com/science/2013/oct/17/skull-homo-erectus-human-evolution | vauthors = Sample I  |work= The Guardian |date= 17 October 2013 }}</ref> However, a 2015 study suggested that ''H. georgicus'' represents an earlier, more primitive species of ''Homo'' derived from an older dispersal of hominins from Africa, with ''H. ergaster/erectus'' possibly deriving from a later dispersal.<ref name="Guimares">{{cite journal |vauthors=Giumares SW, Merino CL |title=Dmanisi hominin fossils and the problem of multiple species in the early Homo genus |journal=Nexus: The Canadian Student Journal of Anthropology |volume=23 |date=September 2015 |s2cid=73528018 |url=https://pdfs.semanticscholar.org/0f43/cf9c1c394c7af01cf10160f164cdec6dd77b.pdf |archive-url=https://web.archive.org/web/20200114162011/https://pdfs.semanticscholar.org/0f43/cf9c1c394c7af01cf10160f164cdec6dd77b.pdf |archive-date=2020-01-14 }}</ref> ''H. georgicus'' is sometimes not even regarded as ''H. erectus''.<ref>{{cite journal | vauthors = Argue D, Groves CP, Lee MS, Jungers WL | title = The affinities of Homo floresiensis based on phylogenetic analyses of cranial, dental, and postcranial characters | journal = Journal of Human Evolution | volume = 107 | pages = 107–133 | date = June 2017 | pmid = 28438318 | doi = 10.1016/j.jhevol.2017.02.006 }}</ref><ref>{{Cite encyclopedia| vauthors = Lordkipanidze D |chapter=Dmanisi|date=2018-10-04|encyclopedia=The International Encyclopedia of Biological Anthropology|pages=1–4| veditors = Trevathan W, Cartmill M, Dufour D, Larsen C |publisher=John Wiley & Sons, Inc.|language=en|doi=10.1002/9781118584538.ieba0139|isbn=9781118584422|s2cid=240090147}}</ref>
===Evolution and dispersal===
{{See also|Human evolution|Early expansions of hominins out of Africa}}
[[File:Dmanisi fossils D 2282 + D 211 (Replika).JPG|left|thumb|upright=0.8|''[[Dmanisi hominins|H. e. georgicus]]'' (above) represents one of the earliest dispersals out of Africa about 1.8 million years ago.<ref name=Ferring2011/>]]
''H. erectus'' evolved in Africa from a population of ''[[Homo habilis|H. habilis]]''{{sfn|Boaz|Ciochon|2004|loc=pp. 143–144}}<ref name=Zaim2011/> and they coexisted for about half a million years.<ref>{{cite journal|first1=F.|last1=Spoor|first2=M. G.|last2=Leakey|first3=P. N.|last3=Gathogo|first4=F. H.|last4=Brown|title=Implications of new early ''Homo'' fossils from Ileret, east of Lake Turkana, Kenya|journal=Nature|year=2007|issn=0028-0836|page=689|volume=448|issue=7154|doi=10.1038/nature05986|first5=S. C.|last5=Antón|first6=I.|last6=McDougall|first7=C.|last7=Kiarie|first8=F. K.|last8=Manthi|first9=L. N.|last9=Leakey |pmid=17687323 |bibcode=2007Natur.448..688S }}</ref> During this time interval, ''H. erectus'' populations could display a mix of more "classically ''erectus''" or "classically ''habilis''" cranial anatomy and a wide range of brain volumes.<ref name=Baab2025>{{cite journal|first1=Karen L.|last1=Baab|first2=Yousuke|last2=Kaifu|first3=Sarah E.|last3=Freidline|first4=Michael J.|last4=Rogers|title=New reconstruction of DAN5 cranium (Gona, Ethiopia) supports complex emergence of Homo erectus|journal=Nature Communications|year=2025|issn=2041-1723|volume=16|issue=1|pmid=41402278|pmc=12708782|doi=10.1038/s41467-025-66381-9|first5=Sileshi|last5=Semaw|doi-access=free}}</ref> The oldest identified ''H. erectus'' specimen is a 2.04 million year old skull, DNH 134, from [[Drimolen]], South Africa, coexisting with the australopithecine ''[[Paranthropus robustus]]''.<ref>{{Cite journal |last1=Herries |first1=Andy I. R. |last2=Martin |first2=Jesse M. |year=2020 |title=Contemporaneity of ''Australopithecus'', ''Paranthropus'', and early ''Homo erectus'' in South Africa  |journal=Science |volume=368 |issue=6486 |article-number=eaaw7293 |doi=10.1126/science.aaw7293 |pmid=32241925 |bibcode=2020Sci...368w7293H |ref=CITEREFHerriesMartinLeeceAdams2020 |hdl=11568/1040368 |s2cid=214763272 |hdl-access=free}}</ref> ''H. erectus'' dispersed out of Africa soon after evolution, the earliest recorded instances being ''H. e. georgicus'' 1.78 to 1.85 million years ago in Georgia,<ref name=Ferring2011>{{Cite journal |last1=Ferring |first1=Reid |last2=Oms |first2=Oriel |last3=Agustí |first3=Jordi |last4=Berna |first4=Francesco |last5=Nioradze |first5=Medea |last6=Shelia |first6=Teona |last7=Tappen |first7=Martha |last8=Vekua |first8=Abesalom |last9=Zhvania |first9=David|last10=Lordkipanidze|first10=David |year=2011 |title=Earliest human occupations at Dmanisi (Georgian Caucasus) dated to 1.85–1.78 Ma |journal=Proceedings of the National Academy of Sciences of the United States of America |volume=108 |issue=26 |article-number=10432–6 |doi=10.1073/pnas.1106638108 |pmid=21646521 |pmc=3127884 |bibcode=2011PNAS..10810432F |doi-access=free}}</ref> the Indonesian Mojokerto and Sangiran sites 1.6 to 1.8 million years ago,{{sfn|Antón|2003|loc=p. 142}}{{sfn|Boaz|Ciochon|2004|loc=p. 162}} and the Chinese [[Yunxian Man]] 1.77 million years ago.<ref>{{cite journal|first1=Hua|last1=Tu|first2=Xiaobo|last2=Feng|first3=Lan|last3=Luo|first4=Zhongping|last4=Lai|title=The oldest in situ ''Homo erectus'' crania in eastern Asia: The Yunxian site dates to ~1.77 Ma|url=https://www.science.org/doi/10.1126/sciadv.ady2270|journal=Science Advances|year=2026|issn=2375-2548|volume=12|issue=8|doi=10.1126/sciadv.ady2270|first5=Darryl|last5=Granger|first6=Christopher|last6=Bae|first7=Guanjun|last7=Shen}}</ref> Populations may have pushed into northwestern Europe at around the same time.<ref>{{cite journal |first1=J. W. F. |last1=Reumer |first2=N. |last2=Peters |first3=J. |last3=de Vos |year=2021 |title=Did ''Homo erectus'' sensu lato live in western Europe during the Early Pleistocene? An ecological approach to a vexing question |journal=Cainozoic Research |volume=21 |issue=1 |pages=97–109 |url=https://natuurtijdschriften.nl/pub/1019587/}}</ref> While ''H. erectus'' is usually considered the first [[hominin]] to leave Africa, stone tools that may date to as far back as 2.48 and 2.1 million years ago (from [[Zarqa River|Zarqa Valley]], Jordan, and [[Shangchen]], China, respectively) could indicate that an earlier hominin species left Africa.<ref>{{cite journal|first1=Giancarlo|last1=Scardia|first2=Fabio|last2=Parenti|first3=Daniel P.|last3=Miggins|first4=Axel|last4=Gerdes|title=Chronologic constraints on hominin dispersal outside Africa since 2.48 Ma from the Zarqa Valley, Jordan|url=https://linkinghub.elsevier.com/retrieve/pii/S0277379119302847|journal=Quaternary Science Reviews|year= 2019|issn=0277-3791|pages=1–19|volume=219|doi=10.1016/j.quascirev.2019.06.007|first5=Astolfo G. M.|last5=Araujo|first6=Walter A.|last6=Neves|bibcode=2019QSRv..219....1S |url-access=subscription|hdl=11449/187832|hdl-access=free}}</ref> Since ''H. erectus'' was first defined in East Asia, those populations are sometimes distinguished as ''H. erectus sensu stricto'' ("in the strict sense"), and African and West Eurasian populations as ''H. erectus sensu lato'' ("in the broad sense"), but this may not reflect how these populations are actually related to each other.{{efn-num|There may have been two different ''H. erectus'' dispersals into East Asia: an early one that led to the Sangiran [[Java Man]] (specifically the older material), and a later one which led to the northern Chinese [[Peking Man]]. That is, the older Sangiran material may be more closely related to ''H. erectus sensu lato'' than to Peking Man. The younger Sangiran material, though, possibly descended from or interbred with Chinese ''H. erectus''.<ref name=Zaim2011>{{cite journal |last1=Zaim|first1=Y.|last2=Ciochon |first2=R. L.|author2-link=Russell Ciochon|last3=Polanski|first3=J. M.|last4=Grine|first4=F. E.|last5=Bettis|first5=E. A.|last6=Rizal |first6=Y.|last7=Franciscus|first7=R. G.|last8=Larick|first8=R. R.|last9=Heizler|first9=M.|last10=Eaves|first10=K. L.|last11=Marsh|first11=H. E.|title=New 1.5 million-year-old ''Homo erectus'' maxilla from Sangiran (Central Java, Indonesia) |journal=Journal of Human Evolution |volume=61 |issue=4 |pages=363–376 |year=2011 |pmid=21783226 |doi=10.1016/j.jhevol.2011.04.009 |bibcode=2011JHumE..61..363Z}}</ref>}}<ref name=Anton2002/><ref name=Zaim2011/>


It is debated whether the African ''H. e. ergaster'' is a separate species (and that ''H. erectus'' evolved in Asia, then migrated to Africa),<ref name="Baab">{{cite journal |vauthors=Baab K |title=Defining Homo erectus |journal=Handbook of Paleoanthropology |edition=2 |pages=2189–2219 |url=https://www.researchgate.net/publication/283477977 |date=December 2015 |doi=10.1007/978-3-642-39979-4_73 |isbn=978-3-642-39978-7 }}</ref> or is the African form (''[[sensu lato]]'') of ''H. erectus ([[sensu stricto]])''. In the latter, ''H. ergaster'' has also been suggested to represent the immediate ancestor of ''H. erectus''.<ref>{{cite book| vauthors = Tattersall I, Schwartz J |title=Extinct Humans|year=2001|isbn=978-0-8133-3482-0|place=Boulder, Colorado |publisher=Westview/Perseus |url=https://archive.org/details/extincthumans00tatt}}{{page needed|date=December 2019}}</ref> It has also been suggested that ''H. ergaster'' instead of ''H. erectus'', or some hybrid between the two, was the immediate ancestor of other archaic humans and modern humans.{{citation needed|date=April 2020}} It has been proposed that Asian ''H. erectus'' have several unique characteristics from non-Asian populations ([[autapomorphies]]), but there is no clear consensus on what these characteristics are or if they are indeed limited to only Asia. Based on supposed derived characteristics, the 120 kya Javan ''H. e. soloensis'' has been proposed to have speciated from ''H. erectus'', as ''H. soloensis'', but this has been challenged because most of the basic cranial features are maintained.<ref>{{cite journal | vauthors = Kaifu Y, Aziz F, Indriati E, Jacob T, Kurniawan I, Baba H | title = Cranial morphology of Javanese Homo erectus: new evidence for continuous evolution, specialization, and terminal extinction | journal = Journal of Human Evolution | volume = 55 | issue = 4 | pages = 551–580 | date = October 2008 | pmid = 18635247 | doi = 10.1016/j.jhevol.2008.05.002 }}</ref> In a wider sense, ''H. erectus'' had mostly been replaced by ''H. heidelbergensis'' by about 300 kya, with possible late survival of ''[[Homo erectus soloensis|H. erectus soloensis]]'' in Java an estimated 117-108 kya.<ref name="Rizal" />
Once established around the [[Old World]], ''H. erectus'' evolved into other later species in the genus ''Homo'', including: ''H. heidelbergensis'', ''[[Homo antecessor|H. antecessor]]'',<ref name=Stringer2012/> ''[[Homo floresiensis|H. floresiensis]]'',<ref>{{cite journal |last1=Berger |first1=L. R. |author-link=Lee Rogers Berger |last2=Churchill |first2=S. E. |display-authors=et al. |year=2008 |title=Small-Bodied Humans from Palau, Micronesia |journal=PLOS ONE |volume=3 |issue=3 |article-number=e1780 |bibcode=2008PLoSO...3.1780B |doi=10.1371/journal.pone.0001780 |pmc=2268239 |pmid=18347737 |doi-access=free}}</ref> and ''[[Homo luzonensis|H. luzonensis]]''.<ref>{{cite journal |first=N. |last=Fleming |year=2019 |title=Unknown human relative discovered in Philippine cave |journal=Nature News |doi=10.1038/d41586-019-01152-3 |pmid=32269371 |s2cid=146786512}}</ref> ''H. heidelbergensis'', in turn, is usually placed as the [[last common ancestor]] of [[Neanderthal]]s (''H. neanderthalensis''), [[Denisovan]]s, and modern humans.<ref name=Stringer2012>{{cite journal |last=Stringer |first=C. |year=2012 |title=What makes a modern human |journal=[[Nature (journal)|Nature]] |volume=485 |number=7396 |pages=33–35 |doi=10.1038/485033a |pmid=22552077 |bibcode=2012Natur.485...33S |s2cid=4420496 |doi-access=free}}</ref> ''H. erectus'' is thus a non-natural, [[paraphyletic]] grouping of fossils and does not include all the descendants of a last common ancestor.<ref name="Ni2021">{{Cite journal |last1=Ni |first1=Xijun |last2=Ji |first2=Qiang |last3=Wu |first3=Wensheng |last4=Shao |first4=Qingfeng |last5=Ji |first5=Yannan |last6=Zhang |first6=Chi |last7=Liang |first7=Lei |last8=Ge |first8=Junyi |last9=Guo |first9=Zhen |last10=Li |first10=Jinhua |last11=Li |first11=Qiang |year=2021 |title=Massive cranium from Harbin in northeastern China establishes a new Middle Pleistocene human lineage |journal=The Innovation |language=en |volume=2 |issue=3 |doi=10.1016/j.xinn.2021.100130 |issn=2666-6758 |pmc=8454562 |pmid=34557770 |last13=Stringer |first13=Chris |last12=Grün |first12=Rainer |article-number=100130 |bibcode=2021Innov...200130N}}</ref> Despite being designated as a different species, ''H. erectus'' may have [[interbreeding between archaic and modern humans|interbred]] with some of its descendant species, namely the common ancestor of Neanderthals and Denisovans ("Neandersovans").<ref>{{cite journal |first=P. J. |last=Waddell |year=2013 |title=Happy New Year ''Homo erectus''? More evidence for interbreeding with archaics predating the modern human/Neanderthal split |journal=Quantitative Biology |pages=2–3 |arxiv=1312.7749}}</ref>


* ''[[Bilzingsleben (Paleolithic site)|H. e. bilzingslebenensis]]'' (Vlček 1978): Originally described from a series of skulls from Bilzingsleben, with the individual of [[Samu (fossil)|Vertesszöllös]] being referred. <ref>{{Cite journal |last=Vlček |first=Emanuel |date=1978-03-01 |title=A new discovery of Homo erectus in central Europe |url=https://www.sciencedirect.com/science/article/pii/S0047248478801158 |journal=Journal of Human Evolution |language=en |volume=7 |issue=3 |pages=239–251 |doi=10.1016/S0047-2484(78)80115-8 |issn=0047-2484}}</ref> The material historically referred to this taxon are now affiliated with [[Neanderthal|Neanderthals]] and the hominins at [[Sima de los huesos|Sima de los Huesos]].<ref>{{Cite journal |last=Arsuaga |first=Juan-Luis |last2=Martínez |first2=Ignacio |last3=Gracia |first3=Ana |last4=Carretero |first4=José-Miguel |last5=Carbonell |first5=Eudald |date=1993 |title=Three new human skulls from the Sima de los Huesos Middle Pleistocene site in Sierra de Atapuerca, Spain |url=https://www.nature.com/articles/362534a0 |journal=Nature |language=en |volume=362 |issue=6420 |pages=534–537 |doi=10.1038/362534a0 |issn=1476-4687}}</ref>
[[File:Spreading homo sapiens la.svg|thumb|upright=2.1|Successive dispersals of {{color box|#e8e22c}} ''Homo erectus'' (yellow), {{color box|#e4ca30}} ''[[Neanderthal|Homo neanderthalensis]]'' (ochre) and {{color box|#e9252c}} ''[[Human|Homo sapiens]]'' (red, ''[[Out of Africa II]]'')]]
* ''[[Boskop Man|H. e. capensis]]'' (Broom 1917): A variant of "''[[Boskop Man|Homo capensis]]''",<ref>{{Cite journal |last=Coon |first=Carleton S. |date=1966 |title=Review of The Nomenclature of the Hominidae, including a Definitive List of Hominid Taxa |url=https://www.jstor.org/stable/41449277 |journal=Human Biology |volume=38 |issue=3 |pages=344–347 |issn=0018-7143}}</ref> a taxon erected from a skull from South Africa formally classified as a type of "[[Race (human categorization)|race]]" but is now considered a representative of the [[Khoisan]].<ref>{{Cite journal |last=Singer |first=Ronald |date=1958 |title=232. The Boskop 'Race' Problem |url=https://www.jstor.org/stable/2795854 |journal=Man |volume=58 |pages=173–178 |doi=10.2307/2795854 |issn=0025-1496}}</ref>
The dispersal of ''H. erectus'' is generally ascribed to the evolution of obligate [[biped]]alism, better technology, and adoption of a carnivorous diet.<ref name=Carotenuto2016/> However, the sudden adoption of carnivory could be [[sampling bias]], with earlier species consuming the same amount of meat.<ref>{{cite journal |first1=W. A. |last1=Barr |first2=B. |last2=Pobiner |first3=J. |last3=Rowan |first4=J. T. |last4=Faith |year=2022 |title=No sustained increase in zooarchaeological evidence for carnivory after the appearance of ''Homo erectus'' |journal=Proceedings of the National Academy of Sciences |volume=119 |issue=5 |article-number=e2115540119 |doi=10.1073/pnas.2115540119|doi-access=free |pmid=35074877 |pmc=8812535 |bibcode=2022PNAS..11915540B }}</ref> Populations spread out via open grassland and woodland [[savanna]]s, which were expanding due to a global [[aridification]] trend at the onset of the [[Quaternary glaciation]].<ref name=Carotenuto2016/> ''H. erectus'' is usually thought to have occupied the [[Sahara]] and West Asia during [[African humid period|humid periods]], but populations may have persisted into desert periods.<ref>{{Cite journal |last1=Mercader |first1=Julio |last2=Akuku |first2=Pamela |last3=Boivin |first3=Nicole |last4=Camacho |first4=Alfredo |last5=Carter |first5=Tristan |last6=Clarke |first6=Siobhán |last7=Cueva Temprana |first7=Arturo |last8=Favreau |first8=Julien |last9=Galloway |first9=Jennifer |last10=Hernando |first10=Raquel |last11=Huang |first11=Haiping |last12=Hubbard |first12=Stephen |last13=Kaplan |first13=Jed O. |last14=Larter |first14=Steve |last15=Magohe |first15=Stephen |year=2025 |title=''Homo erectus'' adapted to steppe-desert climate extremes one million years ago |journal=Communications Earth & Environment |language=en |volume=6 |issue=1 |article-number=1 |doi=10.1038/s43247-024-01919-1 |issn=2662-4435|doi-access=free |pmid=39830897 |pmc=11738993 |bibcode=2025ComEE...6....1M }}</ref>
* ''[[Lantian Man|H. e. chenchiawoensis]]'': A name utilized in a 2007 review of Chinese archeology; the text suggests that it and ''gongwanglingensis'' are contenders in taxonomy<ref name=":03">{{Cite book |last=李学勤 |url=https://books.google.ca/books?id=KtB7iumTYcsC&dq |title=20世纪中国学术大典: 考古学, 博物馆学 |date=2007 |publisher=福建教育出版社 |isbn=978-7-5334-3641-4 |language=zh}}</ref> (despite this name not appearing in the literature).
* ''[[Java Man|H. e. erectus]]'' (Dubois 1891):<ref>{{Cite journal |last=E |first=Dubois |date=1891 |title=Palaeontologische onderzoekingen op Java |url=https://cir.nii.ac.jp/crid/1573387448916597760 |journal=Verslag van het mijnwezen 3^e/4^e kwartaal |pages=12–15}}</ref> The Javanese specimens of ''H. erectus'' were once classified as a distinct subspecies in the 1970s. The [[Java Man|cranium]] from [[Trinil]] is the holotype.<ref>{{Cite book |last=Tuttle |first=Russell H. |url=https://books.google.com/books?id=ejsyIZMsC9oC&pg=PA327 |title=Paleoanthropology: Morphology and Paleoecology |date=2011-05-12 |publisher=Walter de Gruyter |isbn=978-3-11-081069-1 |language=en}}</ref>
* ''[[Homo ergaster|H. e. ergaster]]'' (Groves and Mazák 1975): Antón and Middleton (2023) suggested that ''ergaster'' should be disused based on poor diagnoses.<ref name=":22">{{Cite journal |last=Antón |first=Susan C. |last2=Middleton |first2=Emily R. |date=2023-06-01 |title=Making meaning from fragmentary fossils: Early Homo in the Early to early Middle Pleistocene |url=https://www.sciencedirect.com/science/article/pii/S0047248422001671 |journal=Journal of Human Evolution |language=en |volume=179 |pages=103307 |doi=10.1016/j.jhevol.2022.103307 |issn=0047-2484|doi-access=free }}</ref> The name ''Homo erectus ergaster georgicus'' was created to classify the [[Dmanisi]] population as a subspecies of ''H. e. ergaster'', but [[Binomial nomenclature|quadrinomials]] are not supported by the [[ICZN]].<ref>{{Cite journal |last=Schwartz |first=Jeffrey H. |last2=Tattersall |first2=Ian |last3=Chi |first3=Zhang |date=2014-04-25 |title=Comment on “A Complete Skull from Dmanisi, Georgia, and the Evolutionary Biology of Early Homo ” |url=https://www.science.org/doi/10.1126/science.1250056 |journal=Science |language=en |volume=344 |issue=6182 |pages=360–360 |doi=10.1126/science.1250056 |issn=0036-8075}}</ref>
* ''[[Dmanisi hominins|H. e. georgicus]]'' (Gabounia 1991):<ref>{{Cite journal |last=Gabounia |first=Léo |last2=de Lumley |first2=Marie-Antoinette |last3=Vekua |first3=Abesalom |last4=Lordkipanidze |first4=David |last5=de Lumley |first5=Henry |date=2002-09-01 |title=Découverte d’un nouvel hominidé à Dmanissi (Transcaucasie, Géorgie) |url=https://www.sciencedirect.com/science/article/pii/S1631068302000325 |journal=Comptes Rendus Palevol |language=fr |volume=1 |issue=4 |pages=243–253 |doi=10.1016/S1631-0683(02)00032-5 |issn=1631-0683}}</ref> This hypothetical subspecific designation unites the D2600 cranium with the remainder of the Dmanisi sample, a connection that was, at the time, controversial and was only suggested if the single-species hypothesis could be proven true.<ref>{{Cite journal |last=Rightmire |first=G. Philip |last2=Lordkipanidze |first2=David |last3=Vekua |first3=Abesalom |date=2006-02-01 |title=Anatomical descriptions, comparative studies and evolutionary significance of the hominin skulls from Dmanisi, Republic of Georgia |url=https://www.sciencedirect.com/science/article/pii/S0047248405001624 |journal=Journal of Human Evolution |language=en |volume=50 |issue=2 |pages=115–141 |doi=10.1016/j.jhevol.2005.07.009 |issn=0047-2484}}</ref>
* ''[[Lantian Man|H. e. gongwanglingensis]]'':  A name utilized in a 2007 review of Chinese archeology; the text suggests that it and ''chenchiawoensis'' are contenders in taxonomy.<ref name=":03" /> Rukang (1992) notes that this taxon was born in a "subspecies fever".<ref>{{Cite journal |last=Rukang |first=Wu |date=1992-06-15 |title=On the classification of subspecies of Homo |url=http://www.anthropol.ac.cn/EN/ |journal=Acta Anthropologica Sinica |language=en |volume=11 |issue=02 |pages=109 |issn=1000-3193}}</ref>
* ''[[Homo habilis|H. e. habilis]]'' (Leakey, Tobias, and Napier 1964): D.R. Hughes believed that the [[Olduvai Gorge|Olduvai]] specimens were not distinct enough to be assigned to ''[[Australopithecus]]'', so he created this taxon, as an early variation of ''H. erectus''.<ref>{{Cite journal |last=Tobias |first=Phillip V. |date=1991 |title=The species Homo habilis: example of a premature discovery |url=https://www.jstor.org/stable/23735461 |journal=Annales Zoologici Fennici |volume=28 |issue=3/4 |pages=371–380 |issn=0003-455X}}</ref>
* ''[[Homo heidelbergensis|H. e. heidelbergensis]]'' (Schoetensack 1908): This taxon was used as an alternative to standard ''H. heidelbergensis'' during the middle 20th century, and it was used as a Eurocentric devision of the wider Middle Pleistocene hominin morph.<ref>{{Cite journal |last=Mounier |first=A. |last2=Caparros |first2=M. |date=2015-10-01 |title=Le statut phylogénétique d’Homo heidelbergensis – étude cladistique des homininés du Pléistocène moyen |url=https://doi.org/10.1007/s13219-015-0127-4 |journal=BMSAP |language=fr |volume=27 |issue=3 |pages=110–134 |doi=10.1007/s13219-015-0127-4 |issn=1777-5469}}</ref>
* ''[[Hexian Man|H. e. hexianensis]]'' (Huang 1982): Established based on the Hexian cranium.<ref>{{Cite journal |last=W |first=Huang |date=1982 |title=Preliminary study on the fossil hominid skull and fauna of Hexian, Anhui |url=https://cir.nii.ac.jp/crid/1572543025128829184 |journal=Vertebrata PalAsiatica |volume=20 |pages=plate 1}}</ref>
* ''[[Samu (fossil)|H. e. hungaricus]]'' (Naddeo 2023): A Hungarian paper submitted to a [[Academic conference|conference]] lists this subspecies as an alternate name for the Vertesszöllös remains.<ref>{{Cite journal |last=Naddeo |first=Michelangelo |date=2023 |title=Az ősi magyar jelképrendszer keresése |url=https://epa.oszk.hu/01400/01445/00066/pdf/EPA01445_acta_hungarica_2023_1_0460-0492.pdf |journal=Acta Historica Hungaricus |volume=38}}</ref>
* ''[[Lantian Man|H. e. lantianensis]]'' (Ju-Kang 1964): Based on hominin fossils discovered in [[Lantian County|Lantian]], originally named as a species of ''[[Sinanthropus]]'' and then reclassified as a subspecies.<ref name="Woo1964b">{{cite journal |last=Woo |first=J.-K. |year=1964 |title=The Hominid Skull of Lantian, Shenshi |url=http://www.ivpp.cas.cn/cbw/gjzdwxb/xbwzxz/200912/P020091214552793457098.pdf |journal=Vertebrata PalAsiatica |volume=10 |issue=1}}</ref>
* ''[[Olduvai Hominid 9|H. e. leakeyi]]'' (Heberer 1963): A conditional name and thus unavailable for [[Taxonomy|taxonomic]] use, once used to describe [[Olduvai Hominid 9|OH 9]]. The replacement name is ''louisleakeyi''.<ref>{{Cite journal |last=Groves |first=Colin P. |date=1999-12-01 |title=Nomenclature of African Plio-Pleistocene hominins |url=http://www.sciencedirect.com/science/article/pii/S0047248499903664 |journal=Journal of Human Evolution |language=en |volume=37 |issue=6 |pages=869–872 |doi=10.1006/jhev.1999.0366 |issn=0047-2484 |pmid=10600324}}</ref> It received limited use as a subspecies.<ref name=":12">{{Cite journal |last=Coon |first=Carleton S. |date=1966 |title=Review of The Nomenclature of the Hominidae, including a Definitive List of Hominid Taxa |url=https://www.jstor.org/stable/41449277 |journal=Human Biology |volume=38 |issue=3 |pages=344–347 |issn=0018-7143}}</ref>
* ''[[Maba Man|H. e. mapaensis]]'' (Kurth 1965): A name that was proposed for the [[Maba Man|Maba cranium]], although the use of the word 'perhaps' was interpreted by the [[Paleo Core]] database to be a conditional proposal and thus not available for valid reuse under the ICZN. Groves (1989) classified it as a subspecies of [[Human|''H. sapiens'']], and Howell (1999) did not assign the species to a genus.<ref>{{Cite web |date=2023 |title=Homo erectus mapaensis Kurth, 1965 |url=https://paleocore.org/origins/nomina/detail/218/ |access-date=2023-08-07 |website=Paleo Core |language=en}}</ref>
* ''[[Tighennif|H. e. mauritanicus]]'' (Arambourg 1954): A subspecies that received limited use as a descriptor for the cranial and mandibular material discovered at [[Tighennif|Tighenif]].<ref name=":12" />
* ''[[Narmada cranium|H. e. narmadensis]]'' (Sonakia 1984): The name given to the [[Narmada cranium]].<ref>{{Cite journal |last=Sonakia |first=Arun |last2=Kennedy |first2=Kenneth A. R. |date=1985 |title=Skull Cap of an Early Man from the Narmada Valley Alluvium (Pleistocene) of Central India |url=https://www.jstor.org/stable/678879 |journal=American Anthropologist |volume=87 |issue=3 |pages=612–616 |issn=0002-7294}}</ref>
* ''[[Sambungmacan crania|H. e. newyorkensis]]'' (Laitman and Tattersall 2001): A name based on the Sambungmacan 3 cranium.<ref>{{Cite journal |last=Laitman |first=Jeffrey T. |last2=Tattersall |first2=Ian |date=2001-04-01 |title=Homo erectus newyorkensis: An Indonesian fossil rediscovered in Manhattan sheds light on the middle phase of human evolution |url=https://onlinelibrary.wiley.com/doi/10.1002/ar.1042 |journal=The Anatomical Record |language=en |volume=262 |issue=4 |pages=341–343 |doi=10.1002/ar.1042 |issn=0003-276X}}</ref>
* [[Solo Man|''H. e. ngandongensis'']] (Sartono 1976): A name that was used in the process of splitting Pithecanthropus into many subspecies.<ref>{{Cite book |last=SARTONO R |url=https://pascal-francis.inist.fr/vibad/index.php?action=getRecordDetail&idt=12954719 |title=The Javanese Pleistocene Hominids: A Re-Appraisal |date=1976}}</ref>
* ''[[Olduvai Hominid 9|H. e. olduvaiensis]]'': A subspecies that described the OH 9 cranium, compared to the Bilzingsleben cranial fragments.<ref>{{Cite journal |last=Vlček |first=Emanuel |last2=Mania |first2=Dietrich |date=1977 |title=Ein Neuer Fund Von Homo Erectus in Europa: Bilzingsleben (ddr) |url=https://www.jstor.org/stable/26294532 |journal=Anthropologie (1962-) |volume=15 |issue=2/3 |pages=159–169 |issn=0323-1119}}</ref>
* ''[[Peking Man|H. e. pekinensis]]'' (Black and Zdansky 1927): Originally assigned the type of ''Sinanthropus'' based on a single molar.<ref>{{Cite journal |last=D |first=Black |date=1927 |title=On a lower molar hominid tooth from the Chou Kou Tien deposit |url=https://cir.nii.ac.jp/crid/1571698599180803456 |journal=Palaeont.Sinica, Ser.D |volume=7 |pages=1–29}}</ref> Antón and Middleton (2023) suggested that [[Zhoukoudian Peking Man Site|Zhoukoudian]] and Nanjing may be referrable under this name if they exhibit enough discontinuity from ''H. erectus'' proper.<ref name=":22" />
* ''[[Reilingen cranium|H. e. reilingensis]]'' (Czarnetzki 1989): Referring to a single cranial fragment, this subspecies is now considered a member of the Neanderthal lineage.<ref>{{Cite journal |last=Dean |first=David |last2=Hublin |first2=Jean-Jacques |last3=Holloway |first3=Ralph |last4=Ziegler |first4=Reinhard |date=1998 |title=On the phylogenetic position of the pre-Neandertal specimen from Reilingen, Germany |url=https://linkinghub.elsevier.com/retrieve/pii/S0047248498902147 |journal=Journal of Human Evolution |language=en |volume=34 |issue=5 |pages=485–508 |doi=10.1006/jhev.1998.0214}}</ref>
* ''[[Solo Man|H. e. soloensis]]'' (Oppenoorth 1932): The original name devised by Oppenoorth for the Ngandong crania.<ref>{{Cite journal |last=Oppenoorth |first=W.F.F. |date=1932 |title=Homo (Javanthropus) soloensis. Ein plistocene mensch van Java |journal=Wetenschappelijke mededeelingen |volume=20 |pages=49-75}}</ref>
* ''[[Tautavel Man|H. e. tautavelensis]]'' (de Lumley and de Lumley 1971): Referring to the remains discovered at [[Caune de l'Arago|Arago]], with many preferring allocation to ''Homo heidelbergensis''.<ref>{{Cite thesis |title=Reconstitution et position phylétique des restes crâniens de l'Homme de Tautavel (Arago 21-47) et de Biache-Saint-Vaast 2. Apports de l'imagerie et de l'analyse tridimensionnelle. |url=https://theses.hal.science/tel-00653112 |publisher=Université Paul Cézanne - Aix-Marseille III |date=2005-11-30 |degree=phdthesis |language=fr |first=Gaspard |last=Guipert}}</ref> The remains were determined not to be ''H. erectus'' by Antón and Middleton (2023).<ref name=":22" />
* ''[[Java Man|H. e. trinilensis]]'' (Sartono 1976): A tentative classification scheme, thus making the name conditional and unable for use.<ref>{{Cite journal |last=Sartono |first=S. |date=1980 |title=On the Javanese Pleistocene hominids: A reappraisal |journal=Abstracts of the IUSPP Nice}}</ref>
* ''[[Wajak crania|H. e. wadjakensis]]'' (Dubois 1921): A species established by Eugene Dubois based on the Wajak skulls.<ref>{{Cite journal |last=Dubois |first=E. |date=1921-01-01 |title=The proto-Australian fossil man of Wadjak, Java |url=https://ui.adsabs.harvard.edu/abs/1921KNAB...23.1013D |journal=Koninklijke Nederlandse Akademie van Wetenschappen Proceedings Series B Physical Sciences |volume=23 |pages=1013–1051}}</ref> Pramujiono classified these materials as a subspecies, and incorrectly self-published the name as ''wajakensis''.<ref>{{Cite journal |last=Pramujiono |first=Agung |title=BERBAGAI PANDANGAN ASAL BANGSA DAN BAHASA INDONESIA: DARI KAJIAN LINGUSITIK HISTORIS KOMPARATIF SAMPAI ARKEOLINGUISTIK DAN PALEOLINGUISTIK |url=https://www.academia.edu/9091394/BERBAGAI_PANDANGAN_ASAL_BANGSA_DAN_BAHASA_INDONESIA_DARI_KAJIAN_LINGUSITIK_HISTORIS_KOMPARATIF_SAMPAI_ARKEOLINGUISTIK_DAN_PALEOLINGUISTIK}}</ref>
* ''[[Wushan Man|H. e. wushanensis]]'' (Huang and Fang 1991): Originally conceived as a hominin, the remains this taxon is founded on are more likely referred to [[Ponginae]].<ref>{{Cite journal |last=Wei |first=Guangbiao |last2=Huang |first2=Wanbo |last3=Chen |first3=Shaokun |last4=He |first4=Cunding |last5=Pang |first5=Libo |last6=Wu |first6=Yan |date=2014-12-15 |title=Paleolithic culture of Longgupo and its creators |url=https://www.sciencedirect.com/science/article/pii/S1040618214002110 |journal=Quaternary International |series=Multidisciplinary Perspectives on the Gigantopithecus Fauna and Quaternary Biostratigraphy in East Asia |language=en |volume=354 |pages=154–161 |doi=10.1016/j.quaint.2014.04.003 |issn=1040-6182}}</ref><ref>{{Cite journal |last=Zanolli |first=Clément |last2=Kullmer |first2=Ottmar |last3=Kelley |first3=Jay |last4=Bacon |first4=Anne-Marie |last5=Demeter |first5=Fabrice |last6=Dumoncel |first6=Jean |last7=Fiorenza |first7=Luca |last8=Grine |first8=Frederick E. |last9=Hublin |first9=Jean-Jacques |last10=Nguyen |first10=Anh Tuan |last11=Nguyen |first11=Thi Mai Huong |last12=Pan |first12=Lei |last13=Schillinger |first13=Burkhard |last14=Schrenk |first14=Friedemann |last15=Skinner |first15=Matthew M. |date=May 2019 |title=Evidence for increased hominid diversity in the Early to Middle Pleistocene of Indonesia |url=https://www.nature.com/articles/s41559-019-0860-z |journal=Nature Ecology & Evolution |language=en |volume=3 |issue=5 |pages=755–764 |doi=10.1038/s41559-019-0860-z |issn=2397-334X}}</ref>
* ''[[Yuanmou Man|H. e. yuanmouensis]]'' (Li ''et al.'' 1977): Based on hominin remains<ref>{{Cite journal |last=Chengzhi |first=Hu |date=1973 |title=Yunnan Yuanmou faxian de yuanren yachi huashi |journal=Dizhi xuebao |volume=1 |pages=65-71}}</ref> that Antón and Middleton (2023) suggest do not belong to the taxon H. erectus, although they do not provide an alternate classification.<ref name=":22" />


[[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 [[Georgia (country)|Georgian]] [[Transcaucasus]])]]
Most ''H. erectus sensu lato'' specimens date to 1 to 1.8 million years ago in the Early Pleistocene before giving way to descendant species.<ref name=Rizal2020/> The classification of [[Middle Pleistocene]] ''Homo'' has been a controversial topic, termed "the muddle in the middle".<ref name=Lumley2015/><ref>{{cite journal |first1=C. J. |last1=Bae |first2=X. |last2=Wu |year=2024 |title=Making sense of eastern Asian Late Quaternary hominin variability |journal=Nature Communications |volume=15 |issue=1 |article-number=9479 |doi=10.1038/s41467-024-53918-7 |doi-access=free |pmid=39488555 |pmc=11531466|bibcode=2024NatCo..15.9479B }}</ref> ''H. erectus sensu stricto'' persisted much longer than ''sensu lato'', with the youngest population (''H. e. soloensis'') dating to 108,000 to 117,000 years ago in [[Late Pleistocene]] Java.<ref name=Rizal2020>{{Cite journal |last1=Rizal |first1=Y. |last2=Westaway |first2=K. E. |last3=Zaim |first3=Y. |last4=van den Bergh |first5=E. A. |last5=Bettis III |first6=M. J. |last6=Morwood |first7=O. F. |last7=Huffman |first8=R. |last8=Grün |first9=R. |last9=Joannes-Boyau|first10=R. M.|last10=Bailey |author11=Sidarto |first12=M. C. |last12=Westaway |first13=I. |last13=Kurniawan |first14=M. W. |last14=Moore |first15=M. |last15=Storey |first16=F. |last16=Aziz |author17=Suminto |first18=J. |last18=Zhao |author19=Aswan|first20=M. E.|last20=Sipola |first21=R. |last21=Larick |first22=J.-P. |last22=Zonneveld |first23=R. |last23=Scott |first24=S. |last24=Putt |first25=R. L. |last25=Ciochon |author25-link=Russell Ciochon|year=2020 |title=Last appearance of ''Homo erectus'' at Ngandong, Java, 117,000–108,000 years ago |journal=[[Nature (journal)|Nature]] |volume=577 |issue=7790 |pages=381–385 |doi=10.1038/s41586-019-1863-2 |issn=0028-0836 |pmid=31853068 |s2cid=209410644|hdl=10072/397335 |hdl-access=free }}</ref> This population appears to have died out when the savannah corridors closed and tropical jungle took over.<ref>{{cite journal |first1=J. |last1=Louys |first2=P. |last2=Roberts |year=2020 |title=Environmental drivers of megafauna and hominin extinction in Southeast Asia |journal=[[Nature (journal)|Nature]] |volume=586 |issue=7829 |pages=402–406 |doi=10.1038/s41586-020-2810-y |pmid=33029012 |bibcode=2020Natur.586..402L |hdl=10072/402368 |s2cid=222217295 |hdl-access=free}}</ref>


===Descendants and synonyms===
A 2021 phylogeny of some ''H. erectus'' fossils using [[tip dating]]:<ref name="Ni2021"/>
{{More citations needed section|date=July 2021}}
''Homo erectus'' is the most long-lived species of ''Homo'', having survived for almost two million years. By contrast, ''[[Homo sapiens]]'' emerged about a third of a million years ago.


Regarding many [[archaic humans]], there is no definite consensus as to whether they should be classified as [[Human subspecies|subspecies]] of ''H. erectus'' or ''H. sapiens'' or as separate species.
{{Clade|{{Clade
* African ''H. erectus'' candidates
  |1=''[[Homo habilis|H. habilis]]''
** ''[[Homo ergaster]]'' (or "African ''H. erectus''")
  |label2='''''H. erectus'''''
** ''[[Homo naledi]]''
  |2={{clade
* Eurasian ''H. erectus'' candidates:
    |1={{clade
** ''[[Homo antecessor]]''
      |1=[[Homo gautengensis|Stw 53]] (1.9 million years ago)
** ''[[Homo heidelbergensis]]''
    |2=[[Dmanisi hominins|Dmanisi]] (1.8 million years ago)
* ''[[Homo floresiensis]]''<ref>There was long-standing uncertainty whether ''H. floresiensis'' should be considered close to ''H. erectus'', close to ''H. sapiens'', or an altogether separate species.
    }}
In 2017, it was suggested on morphological grounds that ''H. floresiensis'' is a sister species to either ''H. habilis'' or to a minimally ''habilis''-''erectus''-''ergaster''-''sapiens'' [[clade]], and its line much more ancient than Homo erectus itself.
  |2={{clade
{{cite journal | vauthors = Argue D, Groves CP, Lee MS, Jungers WL | title = The affinities of Homo floresiensis based on phylogenetic analyses of cranial, dental, and postcranial characters | journal = Journal of Human Evolution | volume = 107 | pages = 107–133 | date = June 2017 | pmid = 28438318 | doi = 10.1016/j.jhevol.2017.02.006 }}</ref>
    |1=[[Turkana Boy|Turkana]] (1.7 million years ago)
* ''[[Homo rhodesiensis]]''
    |2={{clade
* the Narmada fossil, discovered in 1982 in [[Madhya Pradesh]], India, was at first suggested as ''H. erectus'' or ''Homo erectus narmadensis''.<ref>{{cite journal | vauthors = Kennedy KA, Sonakia A, Chiment J, Verma KK | title = Is the Narmada hominid an Indian Homo erectus? | journal = American Journal of Physical Anthropology | volume = 86 | issue = 4 | pages = 475–496 | date = December 1991 | pmid = 1776655 | doi = 10.1002/ajpa.1330860404 }}</ref>
      |1=[[Olduvai Hominid 9]] (1.5 million years ago)
      |2={{clade
        |1=[[Sangiran]] (1.4 million years ago)
        |2={{clade
        |1={{clade
          |1={{clade
          |1=[[Nanjing Man]] (0.6 million years ago)
          |2=[[Peking Man]] (0.5 million years ago)
          }}
          |2={{clade
            |1=[[He County#Archaeology|Hexian]] (0.5 million years ago)
            |2={{clade
              |1=[[Sambungmacan crania|Sambungmacan]] (0.2 million years ago)
              |2=[[Solo Man|Ngandong]] (0.1 million years ago)
            }}
            }}
        }}
        |2= {{clade
          |1=''[[Homo heidelbergensis|H. heidelbergensis]]''{{efn-num|''H. heidelbergensis'' is also a [[paraphyly|paraphyletic]] assemblage of fossils.<ref name="Ni2021"/>}}
          |2={{clade
            |1=''[[Neanderthal|H. neanderthalensis]]''
            |2=''[[human|H. sapiens]]''
            }}
          }}
        }}
      }}
    }}
  }}
  }}
}}|label1=''[[Homo]]'' (2.85 million years ago)|style=font-size:85%;line-height:75%}}


''[[Meganthropus]]'', based on fossils found in Java, dated to between 1.4 and 0.9 Mya, was tentatively grouped with ''H. erectus'' in contrast to earlier interpretations of it as a giant species of early human<ref name=Kaifu2005>{{cite journal | vauthors = Kaifu Y, Baba H, Aziz F, Indriati E, Schrenk F, Jacob T | title = Taxonomic affinities and evolutionary history of the Early Pleistocene hominids of Java: dentognathic evidence | journal = American Journal of Physical Anthropology | volume = 128 | issue = 4 | pages = 709–726 | date = December 2005 | pmid = 15761880 | doi = 10.1002/ajpa.10425 }}</ref> although older literature has placed the fossils outside of ''Homo'' altogether.<ref>Krantz, G.S. (1975). "An explanation for the diastema of Javan erectus Skull IV". In: Paleoanthropology, Morphology and Paleoecology. La Hague: Mouton, 361–372.</ref> However, Zanolli et al. (2019) judged ''Meganthropus'' to be a distinct genus of extinct ape.<ref>{{cite journal | vauthors = Zanolli C, Kullmer O, Kelley J, Bacon AM, Demeter F, Dumoncel J, Fiorenza L, Grine FE, Hublin JJ, Nguyen AT, Nguyen TM, Pan L, Schillinger B, Schrenk F, Skinner MM, Ji X, Macchiarelli R | display-authors = 6 | title = Evidence for increased hominid diversity in the Early to Middle Pleistocene of Indonesia | journal = Nature Ecology & Evolution | volume = 3 | issue = 5 | pages = 755–764 | date = May 2019 | pmid = 30962558 | doi = 10.1038/s41559-019-0860-z | s2cid = 102353734 | url = https://kar.kent.ac.uk/72814/1/01-Indonesian_hominid_paleobiodiversity_v2.pdf }}</ref>
==Biology==
As such a widely distributed species both across regions and through time, the anatomy of ''H erectus'' can vary considerably. Among living [[primate]]s, the degree of regionality{{clarify|date=March 2026}} achieved by ''H. erectus'' ([[phenotypic plasticity]]) is only observed in modern humans.<ref name=Anton2016/>


==Anatomy==
===Head===
===Head===
[[File:Skull pekingman.jpg|thumb|left|upright=0.8|Skull of ''H. e. pekinensis'' showing a flat face, pronounced brow ridge, and a [[sagittal keel]]]]
[[File:Solo Man reconstruction.png|thumb|[[Franz Weidenreich]]'s reconstruction of the ''[[Solo Man|H. e. soloensis]]'' skull]]
''Homo erectus'' featured a flat face compared to earlier hominins; pronounced brow ridge; and a low, flat skull.<ref>{{cite journal | vauthors = Baba H, Aziz F, Kaifu Y, Suwa G, Kono RT, Jacob T | title = Homo erectus calvarium from the Pleistocene of Java | journal = Science | volume = 299 | issue = 5611 | pages = 1384–1388 | date = February 2003 | pmid = 12610302 | doi = 10.1126/science.1081676 | s2cid = 20437090 }}</ref><ref name=Balzeau2006/> The presence of [[sagittal keel|sagittal]], [[frontal suture|frontal]], and [[coronal suture|coronal]] keels, which are small crests that run along these [[suture (anatomy)|suture]] lines, has been proposed to be evidence of significant thickening of the skull, specifically the [[cranial vault]]. [[CT scan]] analyses reveal this to not be the case. However, the [[squamous part of occipital bone]], particularly the [[internal occipital crest]], at the rear of the skull is notably thicker than that of modern humans, likely a [[basal (phylogenetics)|basal]] (ancestral) trait.<ref name=Balzeau2006>{{cite journal| vauthors = Balzeau A |year=2006|title=Are thickened cranial bones and equal participation of the three structural bone layers autapomorphic traits of ''Homo erectus''?|journal=Bulletins et mémoires de la Société d'Anthropologie de Paris|volume=18|issue=3–4|pages=145–163|doi=10.4000/bmsap.1528|url=https://journals.openedition.org/bmsap/1528}}</ref><ref>{{cite journal | vauthors = Copes LE, Kimbel WH | title = Cranial vault thickness in primates: Homo erectus does not have uniquely thick vault bones | journal = Journal of Human Evolution | volume = 90 | pages = 120–134 | date = January 2016 | pmid = 26767964 | doi = 10.1016/j.jhevol.2015.08.008 | doi-access = free }}</ref> The fossil record indicates that ''H. erectus'' was the first human species to have featured a projecting nose, which is generally thought to have evolved in response to breathing dry air in order to retain moisture.<ref>{{cite journal | vauthors = Franciscus RG, Trinkaus E | title = Nasal morphology and the emergence of Homo erectus | journal = American Journal of Physical Anthropology | volume = 75 | issue = 4 | pages = 517–527 | date = April 1988 | pmid = 3133950 | doi = 10.1002/ajpa.1330750409 | author2-link = Erik Trinkaus }}</ref> American psychologist Lucia Jacobs hypothesized that the projecting nose instead allowed for distinguishing the direction different smells come from (stereo olfaction) to facilitate navigation and long-distance migration.<ref>{{cite journal | vauthors = Jacobs LF | title = The navigational nose: a new hypothesis for the function of the human external pyramid | journal = The Journal of Experimental Biology | volume = 222 | issue = Pt Suppl 1 | page = jeb186924 | date = February 2019 | pmid = 30728230 | doi = 10.1242/jeb.186924 | doi-access = free }}</ref>
Dubois originally described the species using a skullcap, noting the traits of a low and thickened [[cranial vault]] and a continuous bar of bone forming the brow ridge (supraorbital torus).{{sfn|Antón|2003|loc=p. 132}} ''H. erectus'' fossils typically share these traits, but the Kenyan [[Koobi Fora]] skulls notably have thinner skulls and weaker supraorbital tori.{{sfn|Antón|2003|loc=p. 147}} He also used several other traits now considered more typical of ''H. erectus sensu stricto'', such as a [[sagittal keel]] running across the midline of the skullcap, a bar of bone across the back of the skull ([[occipital bone|occipital]] torus), and a strong crest on the [[mastoid part of the temporal bone]].{{sfn|Antón|2003|loc=p. 132}} These traits can be still be found, nonetheless, in a few ''H. erectus sensu lato'' specimens, namely the 1.47 million year old Olduvai Hominin 9.{{sfn|Antón|2003|loc=p. 136}}


The average brain size of Asian ''H. erectus'' is about {{cvt|1000|cc}}. However, markedly smaller specimens have been found in Dmanisi, Georgia (''H. e. georgicus''); [[Koobi Fora]] and [[Olorgesailie]], Kenya; and possibly [[Gona, Ethiopia|Gona]], Ethiopia. Overall, ''H. erectus'' brain size varies from {{cvt|546–1251|cc}},<ref name=Anton2016/> which is greater than the range of variation seen in modern humans and chimps, though less than that of gorillas.{{Citation needed|date=August 2020}}
Compared to ''H. erectus sensu lato'', the skullcap of ''sensu stricto'' narrows considerably at the front, the face is bigger and presumably more [[prognathism|prognathic]] (it juts out more, but the face is poorly documented), and the molars are larger particularly in Indonesian fossils.{{sfn|Antón|2003|loc=pp. 146–147}} ''H. erectus'' was the first human species with a fleshy [[human nose|nose]], which is generally thought to have evolved in response to breathing dry air in order to retain moisture.<ref>{{cite journal |last1=Franciscus|first1=R. G.|last2=Trinkaus|first2=E.|author2-link=Erik Trinkaus |title=Nasal morphology and the emergence of ''Homo erectus'' |journal=American Journal of Physical Anthropology |volume=75 |issue=4 |pages=517–527 |year=1988 |pmid=3133950 |doi=10.1002/ajpa.1330750409 |bibcode=1988AJPA...75..517F }}</ref> Compared to earlier ''Homo'', ''H. erectus'' has smaller teeth, thinner [[tooth enamel|enamel]], and weaker [[mandible]]s (jawbone), likely due to a greater reliance on tool use and food processing.<ref name=Ungar2006>{{cite journal |last1=Ungar|first1=P. S.|last2=Grine|first2=F. E. |year=2006 |title=Diet in Early ''Homo'': A Review of the Evidence and a New Model of Adaptive Versatility |journal=Annual Review of Anthropology |volume=35 |issue=1 |pages=208–228 |doi=10.1146/annurev.anthro.35.081705.123153 |bibcode=2006ARAnt..35..209U }}</ref>


[[File:Homo erectus reconstruction, Natural History Museum, London.jpg|thumb|upright=1.5|Homo erectus reconstruction, Natural History Museum, London.]]
The brain size of ''H. erectus'' varies considerably, but is generally smaller in ''H. erectus sensu lato'', as low as {{cvt|546|cc}} in [[Dmanisi skull 5]].<ref>{{Cite journal |last1=Lordkipanidze |first1=David |last2=Ponce de León |first2=Marcia S. |last3=Margvelashvili |first3=Ann |last4=Rak |first4=Yoel |last5=Rightmire |first5=G. Philip |last6=Vekua |first6=Abesalom |last7=Zollikofer |first7=Christoph P. E. |year=2013 |title=A Complete Skull from Dmanisi, Georgia, and the Evolutionary Biology of Early ''Homo'' |url=https://www.science.org/doi/10.1126/science.1238484 |journal=Science |volume=342 |issue=6156 |pages=326–331 |doi=10.1126/science.1238484 |pmid=24136960 |bibcode=2013Sci...342..326L |s2cid=20435482|url-access=subscription }}</ref> East Asian ''H.&nbsp;erectus'' overall are rather big-brained, averaging roughly 1,000 cc,<ref name=Anton2016>{{cite journal |first1=S. C. |last1=Antón |first2=H. G. |last2=Taboada |display-authors=et al. |year=2016 |title=Morphological variation in ''Homo erectus'' and the origins of developmental plasticity |journal=[[Philosophical Transactions of the Royal Society B]] |volume=371 |issue=1698 |article-number=20150236 |doi=10.1098/rstb.2015.0236 |pmc=4920293 |pmid=27298467}}</ref> staying within the range of variation for modern humans.<ref>{{Cite journal |doi=10.2478/anre-2021-0029 |title=Interpopulational variation in human brain size: Implications for hominin cognitive phylogeny |year=2022 |last1=Clark |first1=G. |last2=Henneberg |first2=M. |journal=Anthropological Review |volume=84 |issue=4 |pages=405–429 |doi-access=free}}</ref> The late-surviving ''H. e. soloensis'' has the biggest brain volume with one specimen measuring {{cvt|1251|cc}}.{{sfn|Antón|2003|loc=p. 136}}
In an article published in 2021 titled "Interpopulational variation in human brain size: Implications for hominin cognitive phylogeny," it was found that the brain size of Asian ''H. erectus'' over the last 600,000 years overlaps significantly with modern human populations. Significantly, some small brained modern populations showed greater affinity with ''H. erectus'' than they did with other large brained and large bodied modern populations. The paper points out methodological flaws in current understanding of brain size increase in human evolution, where species averages are compared with fossils, which overlooks interpopulational variation. It also overlooks the fact that some modern populations have not seen any dramatic brain size increase relative to ''H. erectus'' with most of the increase occurring in northern populations, which has the result of obscuring interpopulational variation. As the authors write '...the increase in the mean of ''H. sapiens'' cranial capacity is to a large extent due to an increase in the upper limit with a much less pronounced increase in the lower limit relative to our ''H. erectus'' sample. And this increase in the upper limit seems to be more pronounced in northern populations – which may be a result of correlated increases in body size  in  addition  to  climatic  factors'. Consequently, the authors argue that purely based on brain size similarities, Asian ''H. erectus'' could be re-classified as a subspecies of ''H. sapiens'', that is ''H. sapiens soloensis'' - as was suggested by earlier authors.<ref>{{Cite journal|doi = 10.2478/anre-2021-0029|title = Interpopulational variation in human brain size: Implications for hominin cognitive phylogeny|year = 2022|last1 = Clark|first1 = Gary|last2 = Henneberg|first2 = Maciej|journal = Anthropological Review|volume = 84|issue = 4|pages = 405–429|doi-access = free}}</ref>
 
Dentally, ''H. erectus'' have the thinnest [[tooth enamel|enamel]] of any Plio–Pleistocene hominin. Enamel prevents the tooth from breaking from hard foods, but impedes shearing through tough foods. The [[Mandible#Body|bodies of the mandibles]] of ''H. erectus'', and all early ''Homo'', are thicker than those of modern humans and all living apes. The mandibular body resists torsion from the [[bite force]] or chewing, meaning their jaws could produce unusually powerful stresses while eating, but the practical application of this is unclear. Nonetheless, the mandibular bodies of ''H. erectus'' are somewhat thinner than those of early ''Homo''. The premolars and molars also have a higher frequency of pits than ''H. habilis'', suggesting ''H. erectus'' ate more brittle foods (which cause pitting). These all indicate that the ''H. erectus'' mouth was less capable of processing hard foods and more at shearing through tougher foods, thus reducing the variety of foods it could process, likely as a response to tool use.<ref name=Ungar2006>{{cite journal| vauthors = Ungar PS, Grine FE |year=2006|title=Diet in Early ''Homo'': A Review of the Evidence and a New Model of Adaptive Versatility|journal=Annual Review of Anthropology|volume=35|pages=208–228|doi=10.1146/annurev.anthro.35.081705.123153}}</ref>


===Body===
===Body===
[[File:Turkana boy by Mauricio Antón.jpg|thumb|Skeleton and reconstruction of [[Turkana Boy]] by [[Mauricio Antón]]]]  
[[File:Neanderthal Museum 15.jpg|thumb|left|[[Turkana Boy]] at the [[Neanderthal Museum]]]]
Like modern humans, ''H. erectus'' varied widely in size, ranging from {{cvt|146–185|cm|ftin|sigfig=1}} in height and {{cvt|40–68|kg}} in weight, thought to be due to regional differences in climate, mortality rates, or nutrition.<ref>{{cite book
The rest of the body is primarily understood by three partial skeletons from the Kenyan [[Lake Turkana#Anthropology|Lake Turkana]] site, notably [[Turkana Boy]]. Other postcranial fossils (all bones aside from the skull) attributed to ''H. erectus'' are not associated with a skull, making attribution unverifiable. Though the body plan of earlier ''Homo'' is poorly understood, ''H. erectus'' is usually characterized as the first ''Homo'' species with a human body plan, distinct from non-human apes.{{sfn|Antón|2003|loc=pp. 147–152}}<ref name=Anton2016/><ref>{{cite journal |last1=Haeusler|first1=M.|last2=Schiess|first2=R.|last3=Boeni|first3=T.|title=New vertebral and rib material point to modern bauplan of the Nariokotome ''Homo erectus'' skeleton |journal=Journal of Human Evolution |volume=61 |issue=5 |pages=575–582 |year=2011 |pmid=21868059 |doi=10.1016/j.jhevol.2011.07.004 |bibcode=2011JHumE..61..575H |url=https://www.zora.uzh.ch/id/eprint/50126/6/Haeusler_New_vertebral_and_rib_material_point_to_modern_bauplan.pdf}}</ref> The chest may have been short and [[barrel chest|barrel-shaped]], like other archaic humans.<ref name=Bastir2020>{{cite journal|first1=M.|last1=Bastir|first2=D.|last2=García-Martínez|first3=N.|last3=Torres-Tamayo|first4=C. A.|last4=Palancar|first5=B.|last5=Beyer|first6=A.|last6=Barash|first7=C.|last7=Villa|first8=J. A.|last8=Sanchis-Gimeno|first9=A.|last9=Riesco-López|first10=S.|last10=Nalla|first11=I.|last11=Torres-Sánchez|first12=F.|last12=García-Río|first13=E.|last13=Been|first14=A.|last14=Gómez-Olivencia|first15=M.|last15=Haeusler|first16=S. A.|last16=Williams|first17=F.|last17=Spoor|year=2020|title=Rib cage anatomy in ''Homo erectus'' suggests a recent evolutionary origin of modern human body shape|journal=Nature Ecology & Evolution|volume=4|issue=9 |pages=1178–1187|doi=10.1038/s41559-020-1240-4|pmid=32632258 |bibcode=2020NatEE...4.1178B |hdl=2013/ULB-DIPOT:oai:dipot.ulb.ac.be:2013/311388|hdl-access=free}}</ref> [[Fossil track]]s near [[Ileret]], Kenya, suggest a [[human gait]]. This adaptation is implicated in the dispersal of ''H. erectus'' across the Old World.<ref name=Hatala>{{cite journal |last1=Hatala|first1=K. G.|last2=Roach|first2=N. T.|last3=Ostrofsky |first3=K. R.|last4=Wunderlich|first4=R. E.|last5=Dingwall|first5=H. L.|last6=Villmoare|first6=B. A.|last7=Green|first7=D. J.|last8=Harris |first8=J. W.|last9=Braun|first9=D. R.|last10=Richmond|first10=B. G. |title=Footprints reveal direct evidence of group behavior and locomotion in ''Homo erectus'' |journal=Scientific Reports |volume=6 |issue=28766 |article-number=28766 |year=2016 |pmid=27403790 |pmc=4941528 |doi=10.1038/srep28766 |doi-access=free |bibcode=2016NatSR...628766H}}</ref>
| isbn = 9780674600751
| url = https://books.google.com/books?id=RGGVsCPuv1cC&dq=homo+erectus+tall&pg=PA412
| title = The Nariokotome Homo erectus skeleton
| publisher = Harvard University Press
| date = 1993
| access-date = 2 October 2022
| page = 412
| author = Alan Walker, Richard Leakey
}}</ref><ref>{{cite journal| vauthors = Migliano AB, Guillon M |year=2012|title=The Effects of Mortality, Subsistence, and Ecology on Human Adult Height and Implications for ''Homo'' Evolution|journal=Current Anthropology|volume=53|issue=S6|pages=359–368|url=https://www.researchgate.net/publication/233905432|doi=10.1086/667694|s2cid=84442763}}</ref> Among primates, this marked of a response to environmental stressors ([[phenotypic plasticity]]) is only demonstrated in modern humans.<ref name=":0"/><ref>{{Cite journal |last1=Antón |first1=Susan C. |last2=Potts |first2=Richard |last3=Aiello |first3=Leslie C. |date=2014-07-04 |title=Evolution of early ''Homo'': An integrated biological perspective |url=https://www.science.org/doi/10.1126/science.1236828 |journal=Science |language=en |volume=345 |issue=6192 |pages=1236828 |doi=10.1126/science.1236828 |pmid=24994657 |s2cid=30188239 |issn=0036-8075}}</ref><ref name=":02">{{Cite journal |last1=Antón |first1=Susan C. |last2=Taboada |first2=Hannah G. |last3=Middleton |first3=Emily R. |last4=Rainwater |first4=Christopher W. |last5=Taylor |first5=Andrea B. |last6=Turner |first6=Trudy R. |last7=Turnquist |first7=Jean E. |last8=Weinstein |first8=Karen J. |last9=Williams |first9=Scott A. |date=2016-07-05 |title=Morphological variation in ''Homo erectus'' and the origins of developmental plasticity |journal=Philosophical Transactions of the Royal Society B: Biological Sciences |volume=371 |issue=1698 |pages=20150236 |doi=10.1098/rstb.2015.0236 |issn=0962-8436 |pmc=4920293 |pmid=27298467}}</ref>


Like modern humans and unlike other great apes, there does not seem to have been a great size disparity between ''H. erectus'' males and females (size-specific [[sexual dimorphism]]), though there is not much fossil data regarding this.<ref name=Simpson2008/> Brain size in two adults from [[Koobi Fora]] measured {{cvt|848|and|804|cc}},<ref name=Anton2016>{{cite journal | vauthors = Antón SC, Taboada HG, Middleton ER, Rainwater CW, Taylor AB, Turner TR, Turnquist JE, Weinstein KJ, Williams SA | display-authors = 6 | title = Morphological variation in Homo erectus and the origins of developmental plasticity | journal = Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences | volume = 371 | issue = 1698 | page = 20150236 | date = July 2016 | pmid = 27298467 | pmc = 4920293 | doi = 10.1098/rstb.2015.0236 }}</ref> and another significantly smaller adult measured {{cvt|691|cc}}, which could possibly indicate sexual dimorphism, though sex was undetermined.<ref name=Spoor2007>{{cite journal | vauthors = Spoor F, Leakey MG, Gathogo PN, Brown FH, Antón SC, McDougall I, Kiarie C, Manthi FK, Leakey LN | display-authors = 6 | title = Implications of new early Homo fossils from Ileret, east of Lake Turkana, Kenya | journal = Nature | volume = 448 | issue = 7154 | pages = 688–691 | date = August 2007 | pmid = 17687323 | doi = 10.1038/nature05986 | author2-link = Mary Leakey | s2cid = 35845 | bibcode = 2007Natur.448..688S }}</ref> Another case that depicts the difficulty of assigning sex to the fossil record is a few samples taken in Olduvai Gorge. In 1960, in Olduvai Gorge two skulls identified as OH12 and OH9, were found to be that of ''H. erectus'' with a cranial capacities of 1000 cc and 700 cc.<ref name=":5">{{Cite book |last=Leakey |first=Mary D |url=http://catalog.hathitrust.org/api/volumes/oclc/6131892.html |title=Olduvai Gorge: my search for early man |date=1979 |publisher=Collins |location=London |isbn=9780002116138 |language=English |oclc=647137093}}</ref> It is unclear if sexual dimorphism is at play here since the remains are fragmentary.<ref name=":5" /> If ''H. erectus'' did not exhibit sexual dimorphism, then it is possible that they were the first in the human line to do so, though the fragmentary fossil record for earlier species makes this unclear. If yes, then there was a substantial and sudden increase in female height.<ref name=Plavcan2012>{{cite journal| vauthors = Plavcan JM |year=2012|title=Body Size, Size Variation, and Sexual Size Dimorphism in Early ''Homo''|journal=Current Anthropology|volume=53|issue=S6|pages=309–423|doi=10.1086/667605|s2cid=84095311}}</ref> Certain features of sexual dimorphism are often identified in the possibility of determining sex such as lack of muscle marking.<ref name=":13">{{Cite web |title=Rightmire GP. The Evolution of Homo Erectus: Comparative Anatomical Studies of an Extinct Human Species. Cambridge University Press; 1990. |url=https://search.library.ucr.edu/permalink/01CDL_RIV_INST/14qc2ti/alma991029515979704706 |access-date=2022-05-05 |website=search.library.ucr.edu |language=en}}</ref>
It is unclear when human ancestors [[prehistory of nakedness and clothing|lost most of their body hair]]. Genetic analysis suggests that high activity in the [[melanocortin 1 receptor]], which produces dark skin, dates back to 1.2 million years ago. This could indicate the evolution of hairlessness around this time, as a lack of body hair would have left the skin exposed to harmful [[UV radiation]].<ref>{{cite journal |last1=Rogers|first1=A. R.|last2=Iltis|first2=D.|last3=Wooding|first3=S. |year=2004 |title=Genetic Variation at the MC1R Locus and the Time since Loss of Human Body Hair |journal=Current Anthropology |volume=45 |issue=1 |pages=105–108 |doi=10.1086/381006 |bibcode=2004CurrA..45..105R |s2cid=224795768}}</ref> It is possible that populations in higher latitudes developed lighter skin to prevent [[vitamin D deficiency]],<ref>{{cite journal |last=Jablonski|first=N. G. |title=Human skin pigmentation as an example of adaptive evolution |journal=Proceedings of the American Philosophical Society |volume=156 |issue=1 |pages=45–57 |year=2012 |pmid=23035389 |jstor=23558077}}</ref> though a 300,000 to 500,000 year old Turkish ''H. erectus'' specimen presents the earliest case of [[tuberculous meningitis]], which is typically exacerbated by vitamin D deficiency in dark-skinned people living in higher latitudes.<ref>{{cite journal |last1=Kappelman|first1=J.|last2=Alçiçek|first2=M. C.|last3=Kazanci|first3=N.|last4=Schultz|first4=M.|last5=Ozkul|first5=M.|last6=Sen|first6=S. |title=First ''Homo erectus'' from Turkey and implications for migrations into temperate Eurasia |journal=American Journal of Physical Anthropology |volume=135 |issue=1 |pages=110–116 |year=2008 |pmid=18067194 |doi=10.1002/ajpa.20739 |bibcode=2008AJPA..135..110K }}</ref> Hairlessness is generally thought to have facilitated sweating,<ref>{{cite journal |last1=Best|first1=A.|last2=Kamilar|first2=J. M. |title=The evolution of eccrine sweat glands in human and nonhuman primates |journal=Journal of Human Evolution |volume=117 |pages=33–43 |year=2018 |pmid=29544622 |doi=10.1016/j.jhevol.2017.12.003 |bibcode=2018JHumE.117...33B |s2cid=3921318}}</ref> but it may also have helped to reduce parasite load, and was possibly reinforced by [[sexual selection]].<ref>{{cite book |last1=Pagel|first1=M|last2=Bodmer|first2=W. |year=2004 |chapter=The Evolution of Human Hairlessness: Cultural Adaptations and the Ectoparasite Hypothesis |title=Evolutionary Theory and Processes: Modern Horizons |pages=329–335 |publisher=Springer, Dordrecht |doi=10.1007/978-94-017-0443-4_17 |isbn=978-94-017-0443-4}}</ref><ref>{{cite journal |last=Gile|first=J. |year=2010 |title=Naked Love: The Evolution of Human Hairlessness |journal=Biological Theory |volume=5 |issue=4 |pages=326–336 |doi=10.1162/BIOT_a_00062 |s2cid=84164968}}</ref>
[[File:Homo.erectus.adult.female.smithsonian.timevanson.flickr.jpg|thumb|Reconstruction of a female ''H. erectus'']]
''H. erectus'' had about the same limb configurations and proportions as modern humans, implying humanlike locomotion,<ref>{{cite journal | vauthors = Ruff C | title = Femoral/humeral strength in early African Homo erectus | journal = Journal of Human Evolution | volume = 54 | issue = 3 | pages = 383–390 | date = March 2008 | pmid = 17977577 | doi = 10.1016/j.jhevol.2007.09.001 }}</ref> the first in the ''Homo'' lineage.<ref name=":0">{{Cite journal |last1=Antón |first1=Susan C. |last2=Taboada |first2=Hannah G. |last3=Middleton |first3=Emily R. |last4=Rainwater |first4=Christopher W. |last5=Taylor |first5=Andrea B. |last6=Turner |first6=Trudy R. |last7=Turnquist |first7=Jean E. |last8=Weinstein |first8=Karen J. |last9=Williams |first9=Scott A. |date=2016-07-05 |title=Morphological variation in Homo erectus and the origins of developmental plasticity |journal=Philosophical Transactions of the Royal Society B: Biological Sciences |volume=371 |issue=1698 |pages=20150236 |doi=10.1098/rstb.2015.0236 |issn=0962-8436 |pmc=4920293 |pmid=27298467}}</ref> ''H. erectus'' tracks near [[Ileret]], Kenya, also indicate a [[human gait]].<ref name=Hatala/> A humanlike shoulder suggests an ability for high speed throwing.<ref name=":4">Roach, & Richmond. (2015). "Clavicle length, throwing performance and the reconstruction of the Homo erectus shoulder". ''Journal of Human Evolution'', 80(C), 107–113.</ref> It was once thought that Turkana boy had 6 [[lumbar vertebra]] instead of the 5 seen in modern humans and 11 instead of 12 [[thoracic vertebra]]e, but this has since been revised, and the specimen is now considered to have exhibited a humanlike curvature of the spine ([[lordosis]]) and the same number of respective vertebrae.<ref>{{cite journal | vauthors = Haeusler M, Schiess R, Boeni T | title = New vertebral and rib material point to modern bauplan of the Nariokotome Homo erectus skeleton | journal = Journal of Human Evolution | volume = 61 | issue = 5 | pages = 575–582 | date = November 2011 | pmid = 21868059 | doi = 10.1016/j.jhevol.2011.07.004 | url = https://www.zora.uzh.ch/id/eprint/50126/6/Haeusler_New_vertebral_and_rib_material_point_to_modern_bauplan.pdf }}</ref>


It is largely unclear when human ancestors lost most of their body hair. Genetic analysis suggests that high activity in the [[melanocortin 1 receptor]], which would produce dark skin, dates back to 1.2 Mya. This could indicate the evolution of hairlessness around this time, as a lack of body hair would have left the skin exposed to harmful [[UV radiation]].<ref>{{cite journal| vauthors = Rogers AR, Iltis D, Wooding S |year=2004|title=Genetic Variation at the MC1R Locus and the Time since Loss of Human Body Hair|journal=Current Anthropology|volume=45|issue=1|pages=105–108|doi=10.1086/381006|s2cid=224795768}}</ref> It is possible that exposed skin only became maladaptive in the Pleistocene, because the increasing [[axial tilt|tilt]] of the Earth (which also caused the [[Quaternary glaciation|ice ages]]) would have increased solar radiation bombardment- which would suggest that hairlessness first emerged in the australopithecines.<ref name=Gilligan2010/> However, australopithecines seem to have lived at much higher, much colder elevations—typically {{cvt|1000–1600|m}} where the nighttime temperature can drop to {{cvt|10|or|5|C|F}}—so they may have required hair to stay warm, unlike early ''Homo'' which inhabited lower, hotter elevations.<ref>{{cite journal | vauthors = Dávid-Barrett T, Dunbar RI | title = Bipedality and hair loss in human evolution revisited: The impact of altitude and activity scheduling | journal = Journal of Human Evolution | volume = 94 | pages = 72–82 | date = May 2016 | pmid = 27178459 | pmc = 4874949 | doi = 10.1016/j.jhevol.2016.02.006 }}</ref> Populations in higher latitudes potentially developed lighter skin to prevent [[vitamin D deficiency]].<ref>{{cite journal | vauthors = Jablonski NG | title = Human skin pigmentation as an example of adaptive evolution | journal = Proceedings of the American Philosophical Society | volume = 156 | issue = 1 | pages = 45–57 | date = March 2012 | pmid = 23035389 | jstor = 23558077 }}</ref> A 500–300 kya ''H. erectus'' specimen from Turkey was diagnosed with the earliest known case of [[tuberculous meningitis]], which is typically exacerbated in dark-skinned people living in higher latitudes due to vitamin D deficiency.<ref name="First Homo erectus from Turkey and">{{cite journal | vauthors = Kappelman J, Alçiçek MC, Kazanci N, Schultz M, Ozkul M, Sen S | title = First Homo erectus from Turkey and implications for migrations into temperate Eurasia | journal = American Journal of Physical Anthropology | volume = 135 | issue = 1 | pages = 110–116 | date = January 2008 | pmid = 18067194 | doi = 10.1002/ajpa.20739 }}</ref> Hairlessness is generally thought to have facilitated sweating,<ref>{{cite journal | vauthors = Best A, Kamilar JM | title = The evolution of eccrine sweat glands in human and nonhuman primates | journal = Journal of Human Evolution | volume = 117 | pages = 33–43 | date = April 2018 | pmid = 29544622 | doi = 10.1016/j.jhevol.2017.12.003 }}</ref> but reduction of parasite load and [[sexual selection]] have also been proposed.<ref>{{cite book| vauthors = Pagel M, Bodmer W |year=2004|chapter=The Evolution of Human Hairlessness: Cultural Adaptations and the Ectoparasite Hypothesis|title=Evolutionary Theory and Processes: Modern Horizons|pages=329–335|publisher=Springer, Dordrecht|doi=10.1007/978-94-017-0443-4_17|isbn=978-94-017-0443-4}}</ref><ref>{{cite journal| vauthors = Gile J |year=2010|title=Naked Love: The Evolution of Human Hairlessness|journal=Biological Theory|volume=5|issue=4|pages=326–336|doi=10.1162/BIOT_a_00062|s2cid=84164968}}</ref>
===Size===
Height reconstructions range approximately {{cvt|141–167|cm|ftin|0}}, with tropical populations typically reconstructed as scoring on the higher end like modern human populations. Adult weight is harder to approximate, but about {{cvt|50|kg}} may have been normal. ''H. erectus'' is usually thought to be the first human species with little size-specific [[sexual dimorphism]], but the variability of postcranial material makes this unclear.<ref name=Anton2016/> A 2010 study estimates that the [[Turkana Boy]] would have reached a height of {{cvt|163|cm|ftin|0}} if he had reached adulthood.<ref>{{cite journal |first1=R. R. |last1=Graves |first2=A. C. |last2=Lupo |first3=R. C. |last3=McCarthy |first4=D. J. |last4=Wescott |first5=D. L. |last5=Cunningham |year=2010 |title=Just how strapping was KNM-WT 15000? |journal=Journal of Human Evolution |volume=59 |issue=5 |pages=542–554 |doi=10.1016/j.jhevol.2010.06.007|pmid=20846707 |bibcode=2010JHumE..59..542G }}</ref>


===Metabolism===
===Growth and development===
[[File:Pithecanthropus modjokertensis Tjokro Handojo.JPG|thumb|left|Front view of the [[Mojokerto child]] skull]]
The dimensions of a 1.8&nbsp;million years old adult female ''H. e. ergaster'' pelvis from [[Gona, Ethiopia|Gona]], Ethiopia, suggests that she would have been capable of birthing children with a maximum prenatal brain size of {{cvt|315|cc}}, about 30–50% of adult brain size, falling between [[chimpanzee]]s (~40%) and modern humans (28%).<ref name=Simpson2008/> Similarly, a 1.5 million year old infant skull from Mojokerto had a brain volume of about 72–84% the size of an adult, which suggests a brain growth trajectory more similar to that of non-human apes.<ref name=Coqueugniot2004>{{cite journal |last1=Coqueugniot |first1=H. |last2=Hublin |first2=J.-J. |display-authors=et al. |year=2004 |title=Early brain growth in ''Homo erectus'' and implications for cognitive ability |url=https://www.nature.com/articles/nature02852 |journal=Nature |volume=431 |issue=7006 |pages=299–302 |doi=10.1038/nature02852 |pmid=15372030 |bibcode=2004Natur.431..299C |s2cid=4428043|url-access=subscription }}</ref> This suggests that the childhood growth and development of ''H. erectus'' was intermediate between that of chimpanzees and modern humans,<ref name=Simpson2008>{{Cite journal |last1=Simpson |first1=Scott W. |last2=Quade |first2=Jay |last3=Levin |first3=Naomi E. |last4=Butler |first4=Robert |last5=Dupont-Nivet |first5=Guillaume |last6=Everett |first6=Melanie |last7=Semaw |first7=Sileshi |year=2008 |title=A Female ''Homo erectus'' Pelvis from Gona, Ethiopia |url=https://www.science.org/doi/10.1126/science.1163592 |journal=Science |volume=322 |issue=5904 |pages=1089–1092 |doi=10.1126/science.1163592 |pmid=19008443 |bibcode=2008Sci...322.1089S |citeseerx=10.1.1.710.7337 |s2cid=22191315}}</ref> and the faster development rate suggests that [[altriciality]] (an extended childhood) evolved at a later stage in human evolution.<ref name=Coqueugniot2004/> The faster development rate might also indicate a shorter expected lifespan compared to later ''Homo''.<ref>{{Cite journal |last1=Caspari |first1=Rachel |last2=Lee |first2=Sang-Hee |year=2004 |title=Older age becomes common late in human evolution |journal=PNAS |volume=101 |issue=30 |pages=10,895–10,900 |doi=10.1073/pnas.0402857101 |pmid=15252198 |pmc=503716 |doi-access=free}}</ref>
The 1.8 Ma [[Mojokerto child]] specimen from Java, who died at about 1 year of age, presented 72–84% of the average adult brain size, which is more similar to the faster brain growth trajectory of great apes than modern humans. This indicates that ''H. erectus'' was probably not cognitively comparable to modern humans, and that [[altriciality|secondary altriciality]]—an extended childhood and long period of dependency due to the great amount of time required for brain maturation—evolved much later in human evolution, perhaps in the modern human/Neanderthal last common ancestor.<ref name=Coqueugniot2004>{{cite journal | vauthors = Coqueugniot H, Hublin JJ, Veillon F, Houët F, Jacob T | title = Early brain growth in Homo erectus and implications for cognitive ability | journal = Nature | volume = 431 | issue = 7006 | pages = 299–302 | date = September 2004 | pmid = 15372030 | doi = 10.1038/nature02852 | s2cid = 4428043 | bibcode = 2004Natur.431..299C }}</ref> It was previously believed that, based on the narrow pelvis of Turkana boy, ''H. erectus'' could only safely deliver a baby with a brain volume of about {{cvt|230|cc}}, equating to a similar brain growth rate as modern humans to achieve the average adult brain size of {{cvt|600–1067|cc}}. However, a 1.8 Ma female pelvis from Gona<!--[[Gona]] directs to a village in Papua New Guinea-->, Ethiopia, shows that ''H. erectus'' babies with a brain volume of {{cvt|310|cc}} could have been safely delivered, which is 34–36% the mean adult size, compared to 40% in chimps and 28% in modern humans. This more aligns with the conclusions drawn from the Mojokerto child.<ref name=Simpson2008>{{cite journal | vauthors = Simpson SW, Quade J, Levin NE, Butler R, Dupont-Nivet G, Everett M, Semaw S | title = A female Homo erectus pelvis from Gona, Ethiopia | journal = Science | volume = 322 | issue = 5904 | pages = 1089–1092 | date = November 2008 | pmid = 19008443 | doi = 10.1126/science.1163592 | s2cid = 22191315 | bibcode = 2008Sci...322.1089S }}</ref> A faster development rate could indicate a lower expected lifespan.<ref>{{cite journal | vauthors = Caspari R, Lee SH | title = Older age becomes common late in human evolution | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 101 | issue = 30 | pages = 10895–10900 | date = July 2004 | pmid = 15252198 | pmc = 503716 | doi = 10.1073/pnas.0402857101 | doi-access = free }}</ref>
 
Based on an average mass of {{cvt|63|kg}} for males and {{cvt|52.3|kg}} for females, the daily energy expenditure (DEE)—the amount of calories metabolized in one day—was estimated to be about 2271.8 and 1909.5 [[kcal]], respectively. This is similar to that of earlier ''Homo'', despite a marked increase in activity and migratory capacity, likely because the longer legs of ''H. erectus'' were more energy-efficient in long-distance movement. Nonetheless, the estimate for ''H. erectus'' females is 84% higher than that for ''Australopithecus'' females, possibly due to an increased body size and a decreased growth rate.<ref>{{cite journal | vauthors = Steudel-Numbers KL | title = Energetics in Homo erectus and other early hominins: the consequences of increased lower-limb length | journal = Journal of Human Evolution | volume = 51 | issue = 5 | pages = 445–453 | date = November 2006 | pmid = 16780923 | doi = 10.1016/j.jhevol.2006.05.001 }}</ref> A 2011 study, assuming high energy or dietary fat requirements based on the abundance of large game animals at ''H. erectus'' sites, calculated a DEE of 2,700–3,400 kcal of which 27–44% derived from fat, and 44–62% of the fat from animal sources. In comparison, modern humans with a similar activity level have a DEE of 2,450 calories, of which 33% derives from fat, and 49% of the fat from animals.<ref name=BenDor2011/>


===Bone thickness===
===Bone thickness===
[[File:Homo erectus humeri.png|thumb|Cross sections of Chinese ''H. erectus'' [[humerus|humeri]] (upper arm bones) showing extremely thickened [[cortical bone]]]]
[[File:Homo erectus skull cross-section.png|thumb|Cross sections of [[Peking Man]] Skulls III (A) and XII (B), and [[Java Man]] Skull II (C)]]
The [[cortical bone]] (the outer layer of the bone) is extraordinarily thickened, particularly in East Asian populations. The skullcaps have oftentimes been confused with fossil turtle [[carapace]]s,<ref name=Boaz2004/> and the [[medullary canal]] in the [[long bone]]s (where the [[bone marrow]] is stored, in the limbs) is extremely narrowed (medullary [[stenosis]]). This degree of thickening is usually exhibited in semi-aquatic animals which used their heavy ([[pachyosteosclerosis|pachyosteosclerotic]]) bones as ballasts to help them sink, induced by [[hypothyroidism]]. Male specimens have thicker cortical bone than females.<ref name=Kennedy1985/>
The bones are extraordinarily thickened, particularly in ''Homo erectus sensu stricto'', so much so that skull fragments have sometimes been confused for fossil turtle [[carapace]]s.{{sfn|Boaz|Ciochon|2004|loc=p. 74}} The [[medullary canal]] in the [[long bone]]s (where the [[bone marrow]] is stored, in the limbs) is extremely narrowed (medullary [[stenosis]]). This degree of thickening is usually exhibited in semi-aquatic animals which use their heavy ([[pachyosteosclerosis|pachyosteosclerotic]]) bones as ballasts to help them sink, induced by [[hypothyroidism]].<ref name=Kennedy1985>{{cite journal |last=Kennedy|first=G. E. |year=1985 |title=Bone thickness in ''Homo erectus'' |journal=Journal of Human Evolution |volume=14 |issue=8 |pages=699–708 |doi=10.1016/S0047-2484(85)80052-X |bibcode=1985JHumE..14..699K}}</ref>


It is largely unclear what function this could have served. All pathological inducers would leave scarring or some other indicator not normally exhibited in ''H. erectus''. Before more complete skeletons were discovered, Weidenreich suggested ''H. erectus'' was a gigantic species, thickened bone required to support the massive weight. It was hypothesized that intense physical activity could have induced bone thickening, but in 1970, human biologist [[Stanley Marion Garn]] demonstrated there is a low correlation between the two at least in modern humans. Garn instead noted different races have different average cortical bone thicknesses, and concluded it is genetic rather than environmental. It is unclear if the condition is caused by increased bone apposition (bone formation) or decreased [[bone resorption]], but Garn noted the stenosis is quite similar to the [[congenital]] condition in modern humans induced by hyper-apposition. In 1985, biological anthropologist Gail Kennedy argued for resorption as a result of [[hyperparathyroidism]] caused by [[hypocalcemia]] ([[calcium in biology|calcium]] deficiency), a consequence of a dietary shift to low-calcium meat. Kennedy could not explain why the [[calcium metabolism]] of ''H. erectus'' never adjusted.<ref name=Kennedy1985>{{cite journal | vauthors = Kennedy GE |year=1985|title=Bone thickness in ''Homo erectus''|journal=Journal of Human Evolution|volume=14|issue=8|pages=699–708|doi=10.1016/S0047-2484(85)80052-X}}</ref> In 1985, American paleoanthropologist Mary Doria Russell<!--not the novelist--> and colleagues argued the supraorbital torus is a response to withstanding major [[bending moment]] which localizes in that region when significant force is applied through the front teeth, such as while using the mouth as a third hand to carry objects.<ref>{{cite journal| vauthors = Russell MD, Brown T, Garn SM, Giris F, Turkel S, İşcan MY, Oyen OJ, Jacobshagen B, Pietrusewsky M, Rightmire GP, Smith FH | display-authors = 6 |year=1985|title=The Supraorbital Torus: 'A Most Remarkable Peculiarity'|journal=Current Anthropology|volume=26|issue=3|pages=337–350|doi=10.1086/203279|s2cid=146857927}}</ref>
It is unclear what function intense bone thickening could have served. Before more complete skeletons were discovered, Weidenreich suggested ''H. erectus'' was a gigantic species.<ref>{{cite book|first=F.|last=Weidenreich|authorlink=Franz Weidenreich|year=1946|chapter=Giants as Earliest Ancestors|title=Apes, Giants, and Man|chapter-url=https://archive.org/details/in.ernet.dli.2015.531106/page/n57|publisher=University of Chicago Press}}</ref> Other explanations include a far more violent and impact-prone lifestyle than other ''Homo'', or pathological nutrient deficiencies.{{sfn|Boaz|Ciochon|2004|loc=pp. 80–86}} The supraorbital torus thickens with age, and may be a response to [[bending]] stresses from habitual loading of the front teeth.<ref>{{cite journal |last1=Russell|first1=M. D.|last2=Brown|first2=T.|last3=Garn|first3=S. M.|last4=Giris|first4=F.|last5=Turkel|first5=S.|last6=İşcan|first6=M. Y.|last7=Oyen|first7=O. J.|last8=Jacobshagen|first8=B.|last9=Pietrusewsky|first9=M.|last10=Rightmire|first10=G. P.|last11=Smith|first11=F. H.|year=1985 |title=The Supraorbital Torus: 'A Most Remarkable Peculiarity' |journal=Current Anthropology |volume=26 |issue=3 |pages=337–350 |doi=10.1086/203279 |s2cid=146857927}}</ref>
 
In 2004, Noel Boaz and Russel Ciochon suggested it was a result of a cultural practice, wherein ''H. erectus'' would fight each other with fists, stones, or clubs to settle disputes or battle for mates, since the skull is reinforced in key areas. The mandible is quite robust, capable of absorbing heavy blows (no "glass jaw"); the heavy brow ridge protects the eyes, and transitions into a bar covering the ears, connecting all the way in the back of the skull, meaning blows to any of these regions can be effectively dissipated across the skull; and the sagittal keel protects the top of the braincase. Many skullcaps bear usually debilitating fractures, such as the Peking Man skull X, yet they can show signs of surviving and healing. Anthropologist [[Peter Brown (anthropologist)|Peter Brown]] suggested a similar reason for the unusual thickening of the modern [[Australian Aboriginal]] skull, a result of a ritual popular in central and southeast Australian tribes where adversaries would wack each other with [[waddy|waddies]] (sticks) until [[knockout]].<ref name=Boaz2004>{{cite journal| vauthors = Boaz N, Ciochon R |year=2004|title=Headstrong Hominids|url=https://www.researchgate.net/publication/295215638|journal=Natural History|volume=113|issue=1|pages=28–34}}</ref>


==Culture==
==Culture==
===Social structure===
===Subsistence===
[[File:Ileret trackways.jpg|thumb|upright=1.3|Diagram of fossil trackways from 2 sites near [[Ileret]], Kenya]]
{{See also|Hunting hypothesis}}
The only fossil evidence regarding ''H. erectus'' group composition comes from 4 sites outside of [[Ileret]], Kenya, where 97 footprints made 1.5 Mya were likely left by a group of at least 20 individuals. One of these trackways, based on the size of the footprints, may have been an entirely male group, which could indicate they were some specialised task group, such as a hunting or foraging party, or a border patrol. If correct, this would also indicate sexual division of labour, which distinguishes human societies from those of other great apes and social mammalian carnivores. In modern hunter gatherer societies who target large prey items, typically male parties are dispatched to bring down these high-risk animals, and, due to the low success rate, female parties focus on more predictable foods.<ref name=Hatala>{{cite journal | vauthors = Hatala KG, Roach NT, Ostrofsky KR, Wunderlich RE, Dingwall HL, Villmoare BA, Green DJ, Harris JW, Braun DR, Richmond BG | display-authors = 6 | title = Footprints reveal direct evidence of group behavior and locomotion in Homo erectus | journal = Scientific Reports | volume = 6 | issue = 28766 | pages = 28766 | date = July 2016 | pmid = 27403790 | pmc = 4941528 | doi = 10.1038/srep28766 | doi-access = free | bibcode = 2016NatSR...628766H }}</ref> Based on modern day savanna chimp and [[baboon]] group composition and behavior, ''H. erectus ergaster'' may have lived in large, multi-male groups in order to defend against large savanna predators in the open and exposed environment.<ref name=Willems2017/> However, dispersal patterns indicate that ''H. erectus'' generally avoided areas with high carnivore density.<ref name=Carotenuto2016/> It is possible that male–male bonding and male–female friendships were important societal aspects.<ref name=Willems2017>{{cite journal | vauthors = Willems EP, van Schaik CP | title = The social organization of Homo ergaster: Inferences from anti-predator responses in extant primates | journal = Journal of Human Evolution | volume = 109 | pages = 11–21 | date = August 2017 | pmid = 28688456 | doi = 10.1016/j.jhevol.2017.05.003 }}</ref>
[[File:Megalochelys AMNH.jpg|thumb|left|''H. erectus'' overhunting may have led to the extinction of ''[[Megalochelys]]'' (above).<ref name=Rhodin2015/>]]
''H. erectus'' was early-on portrayed as the earliest [[hunter-gatherer]] and a skilled predator of big game, relying on running. The few identified specimens of the ''H. e. ergaster'' torso and pelvis may indicate a body plan more conducive for power running, unlike modern humans better [[endurance running hypothesis|adapted for endurance running]].<ref name=Bastir2020/> The gradual shift to "[[apex predator|top predator]]" may have led to its dispersal throughout Afro-Eurasia.<ref name=Carotenuto2016/> Though scavenging may have instead played a bigger role at least in some populations, ''H. erectus'' fossils are often associated with the butchered remains of large herbivores,{{sfn|Boaz|Ciochon|2004|loc=p. 105}} especially [[elephant]]s, [[rhino]]s, [[hippo]]s, [[bovine]]s, and [[boar]]s. Tracking complex prey behaviors as well as the nutritional value of meat have been connected to brain volume growth.<ref name=BenDor2011/>


Because ''H. erectus'' children had faster brain growth rates, ''H. erectus'' likely did not exhibit the same degree of maternal investment or child-rearing behaviours as modern humans.<ref name=Simpson2008/>
''H. erectus'' is usually assumed to have practiced [[sexual division of labor]] much like recent hunter-gatherer societies, with men hunting and women gathering. This model is supported by a fossil trackway from Ileret, Kenya, made by a probably all-male band of over 20 ''H. erectus'' individuals, possibly a hunting party or (similar to chimpanzees) a border patrol group.<ref>{{Cite journal |last1=Hatala |first1=Kevin G. |last2=Roach |first2=Neil T. |last3=Ostrofsky |first3=Kelly R. |last4=Wunderlich |first4=Roshna E. |last5=Dingwall |first5=Heather L. |last6=Villmoare |first6=Brian A. |last7=Green |first7=David J. |last8=Harris |first8=John W. K. |last9=Braun |first9=David R.|last10=Richmond|first10=Brian G. |year=2016 |title=Footprints reveal direct evidence of group behavior and locomotion in ''Homo erectus'' |journal=Scientific Reports |volume=6 |issue=28766 |article-number=28766 |doi=10.1038/srep28766 |pmid=27403790 |pmc=4941528 |bibcode=2016NatSR...628766H}}</ref>  


Because ''H. erectus'' males and females are thought to have been about the same size compared to other great apes (exhibit less size-specific sexual dimorphism), it is generally hypothesised that they lived in a monogamous society, as reduced sexual dimorphism in primates is typically correlated with this mating system.<ref name=Plavcan2012/> However, it is unclear if ''H. erectus'' did in fact exhibit humanlike rates of sexual dimorphism.<ref name=Spoor2007/> If they did, then it would mean only female height increased from the ancestor species, which could have been caused by a shift in female fertility or diet, and/or reduced pressure on males for large size. This in turn could imply a shift in female behavior which made it difficult for males to maintain a harem.<ref>{{cite journal | vauthors = Plavcan JM |year=2012|title=Implications of Male and Female Contributions to Sexual Size Dimorphism for Inferring Behavior in the Hominin Fossil Record|journal=International Journal of Primatology|volume=33|issue=6|pages=1364–1381|doi=10.1007/s10764-012-9642-z|s2cid=17850676}}</ref>
Since common modern human tapeworms began to diverge from those of other predators roughly 1.7 million years ago (specifically the [[Taenia solium|pork tapeworm]], [[Taenia saginata|beef tapeworm]], and [[Taenia asiatica|Asian tapeworm]]), not only was ''H. erectus'' consuming meat regularly enough for speciation to occur in these parasites, but meat was probably consumed raw more often than not.{{sfn|Boaz|Ciochon|2004|loc=p. 105–107}} Some populations were collecting aquatic resources like fish, shellfish, and turtles at waterside sites, such as Lake Turkana<ref>{{cite journal |last=Steele|first=T. E. |title=A unique hominin menu dated to 1.95 million years ago |journal=[[Proceedings of the National Academy of Sciences of the United States of America]] |volume=107 |issue=24 |pages=10771–10772 |year=2010 |pmid=20534542 |pmc=2890732 |doi=10.1073/pnas.1005992107 |doi-access=free |bibcode=2010PNAS..10710771S}}</ref> and Trinil.<ref>{{cite journal |last1=Joordens|first1=J. C.|last2=Wesselingh|first2=F. P.|last3=de Vos|first3=J.|last4=Vonhof|first4=H. B.|last5=Kroon|first5=D. |title=Relevance of aquatic environments for hominins: a case study from Trinil (Java, Indonesia) |journal=[[Journal of Human Evolution]] |volume=57 |issue=6 |pages=656–671 |year=2009 |pmid=19683789 |doi=10.1016/j.jhevol.2009.06.003 |bibcode=2009JHumE..57..656J}}</ref> Underground [[storage organ]]s (roots, tubers, etc.) were likely also major dietary components, and traces of the edible plant ''[[Celtis]]'' have been documented at several ''H. erectus'' sites.<ref name=Hardy2018>{{cite journal |first=K. |last=Hardy |year=2018 |title=Plant use in the Lower and Middle Palaeolithic: Food, medicine and raw materials |journal=Quaternary Science Reviews |volume=191 |pages=393–398 |doi=10.1016/j.quascirev.2018.04.028|bibcode=2018QSRv..191..393H }}</ref>


===Food===
Possibly due to overhunting of the biggest game available, the dispersal of ''H. erectus'' and descendant species may be implicated in the extinctions of large herbivores and the gradual reduction of average herbivore size over the Pleistocene.<ref>{{cite journal |first1=J. |last1=Dembitzer |first2=R. |last2=Barkai |first3=M. |last3=Ben-Dor |first4=S. |last4=Meiri |year=2022 |title=Levantine overkill: 1.5 million years of hunting down the body size distribution |journal=Quaternary Science Reviews |volume=276 |article-number=107316 |doi=10.1016/j.quascirev.2021.107316|bibcode=2022QSRv..27607316D }}</ref> ''H. erectus'' overhunting has been blamed by some authors for the decline of [[proboscidea]]n species as well as competing carnivores,<ref name=BenDor2011>{{cite journal |last1=Ben-Dor|first1=M.|last2=Gopher|first2=A.|last3=Hershkovitz|first3=I|last4=Barkai|first4=R. |title=Man the fat hunter: the demise of ''Homo erectus'' and the emergence of a new hominin lineage in the Middle Pleistocene (ca. 400 kyr) Levant |journal=PLOS ONE |volume=6 |issue=12 |article-number=e28689 |year=2011 |pmid=22174868 |pmc=3235142 |doi=10.1371/journal.pone.0028689 |bibcode=2011PLoSO...628689B |doi-access=free}}</ref><ref>{{cite journal |first1=T. |last1=Surovell |first2=N. |last2=Waguespack |first3=P. J. |last3=Brantingham |year=2005 |title=Global archaeological evidence for proboscidean overkill |journal=Proceedings of the National Academy of Sciences |volume=102 |issue=17 |pages=6,231-6,236 |doi=10.1073/pnas.0501947102|doi-access=free |pmid=15829581 |pmc=1087946 |bibcode=2005PNAS..102.6231S }}</ref><ref name=Faith2020>{{cite journal|last1=Faith|first1=J. Tyler|last2=Rowan|first2=John|last3=Du|first3=Andrew|last4=Barr|first4=W. Andrew|year=2020|title=The uncertain case for human-driven extinctions prior to ''Homo sapiens''|journal=Quaternary Research|volume=96|pages=88–104|doi=10.1017/qua.2020.51 |bibcode=2020QuRes..96...88F }}</ref> but their decline may be better attributed to the spread of grasslands.<ref name=Faith2020/><ref>{{cite journal|last1=Saarinen|first1=Juha|last2=Lister|first2=Adrian M.|year=2023|title=Fluctuating climate and dietary innovation drove ratcheted evolution of proboscidean dental traits|journal=Nature Ecology & Evolution|volume=7|issue=9 |pages=1490–1502|doi=10.1038/s41559-023-02151-4|pmid=37580434 |pmc=10482678|bibcode=2023NatEE...7.1490S }}</ref> The giant tortoise ''[[Megalochelys]]'' may have been driven to extinction by ''H. erectus'' in [[Sundaland]] (what is now [[Island Southeast Asia]]), since species went extinct shortly after the arrival of ''H. erectus''.<ref name=Rhodin2015>{{Cite book |url=http://www.iucn-tftsg.org/cbftt/ |title=Conservation Biology of Freshwater Turtles and Tortoises |chapter=Turtles and Tortoises of the World During the Rise and Global Spread of Humanity: First Checklist and Review of Extinct Pleistocene and Holocene Chelonians |year=2015 |publisher=Chelonian Research Foundation |isbn=978-0-9653540-9-7 |editor-last=Rhodin |editor-first=A. |edition=First |series=Chelonian Research Monographs |volume=5 |doi=10.3854/crm.5.000e.fossil.checklist.v1.2015 |editor-last2=Pritchard |editor-first2=P. |editor-last3=van Dijk |editor-first3=P. P. |editor-last4=Saumure |editor-first4=R. |editor-last5=Buhlmann |editor-first5=K. |editor-last6=Iverson |editor-first6=J. |editor-last7=Mittermeier |editor-first7=R. |page=15}}</ref>
Increasing brain size is often directly associated with a meatier diet and resultant higher caloric intake. [[Human entomophagy]] and therefore an increase in protein consumption through insects has also been proposed as a possible cause. However, it is also possible that the energy-expensive guts decreased in size in ''H. erectus'', because the large ape gut is used to synthesize fat by fermenting plant matter which was replaced by dietary animal fat, allowing more energy to be diverted to brain growth. This would have increased brain size indirectly while maintaining the same caloric requirements of ancestor species. ''H. erectus'' may have also been the first to use a [[Hunter-gatherer|hunting and gathering]] food collecting strategy as a response to the increasing dependence on meat. With an emphasis on teamwork, division of labor, and food sharing, hunting and gathering was a dramatically different subsistence strategy from previous modes.<ref name=Ungar2006/><ref name=BenDor2011/>


[[File:Elephasantiquus.jpg|thumb|upright=1|''H. erectus'' ate primarily large game, such as the [[straight-tusked elephant]] (above)]]
===Technology===
''H. erectus'' sites frequently are associated with assemblages of medium- to large-sized game, namely [[elephant]]s, [[rhino]]s, [[hippo]]s, [[bovine]], and [[boar]]. ''H. erectus'' would have had considerable leftovers, potentially pointing to food sharing or long-term [[food preservation]] (such as by drying) if most of the kill was indeed utilized. It is possible that ''H. erectus'' grew to become quite dependent on large-animal meat, and the disappearance of ''H. erectus'' from the [[Levant]] is correlated with the local extinction of the [[straight-tusked elephant]].<ref name=BenDor2011>{{cite journal | vauthors = Ben-Dor M, Gopher A, Hershkovitz I, Barkai R | title = Man the fat hunter: the demise of Homo erectus and the emergence of a new hominin lineage in the Middle Pleistocene (ca. 400 kyr) Levant | journal = PLOS ONE | volume = 6 | issue = 12 | pages = e28689 | year = 2011 | pmid = 22174868 | pmc = 3235142 | doi = 10.1371/journal.pone.0028689 | bibcode = 2011PLoSO...628689B | doi-access = free }}</ref> Nonetheless, ''H. erectus'' diet likely varied widely depending upon location. For example, at the 780 kya [[Daughters of Jacob Bridge|Gesher Benot Ya'aqov]] site, Israel, the inhabitants gathered and ate 55 different types of fruits, vegetables, seeds, nuts, and tubers, and it appears that they used fire to roast certain plant materials that otherwise would have been inedible; they also consumed amphibians, reptiles, birds, aquatic and terrestrial invertebrates, in addition to the usual large creatures such as elephant and [[fallow deer]].<ref>{{cite journal | vauthors = Melamed Y, Kislev ME, Geffen E, Lev-Yadun S, Goren-Inbar N | title = The plant component of an Acheulian diet at Gesher Benot Ya'aqov, Israel | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 113 | issue = 51 | pages = 14674–14679 | date = December 2016 | pmid = 27930293 | pmc = 5187744 | doi = 10.1073/pnas.1607872113 | bibcode = 2016PNAS..11314674M | doi-access = free }}</ref> At the 1.95 Mya FwJJ20 lakeside site in the [[East Turkana]] Basin, Kenya, the inhabitants ate (alongside the usual bovids, hippos, and rhinos) aquatic creatures such as [[turtle]]s, [[crocodile]]s, and [[catfish]]. The large animals were likely scavenged at this site, but the turtles and fish were possibly collected live.<ref>{{cite journal | vauthors = Steele TE | title = A unique hominin menu dated to 1.95 million years ago | journal = [[Proceedings of the National Academy of Sciences of the United States of America]] | volume = 107 | issue = 24 | pages = 10771–10772 | date = June 2010 | pmid = 20534542 | pmc = 2890732 | doi = 10.1073/pnas.1005992107 | doi-access = free | bibcode = 2010PNAS..10710771S }}</ref> In East Africa between 2.0 and 1.4 Mya, carcasses of {{C4}}-grazing ungulates, particularly [[Alcelaphini|alcelaphins]], featured increasingly prominently in the diet of these hominins.<ref>{{cite journal |last1=Patterson |first1=D. B. |last2=Braun |first2=D. R. |last3=Behrensmeyer |first3=A. K. |last4=Merritt |first4=S. |last5=Zliobaite |first5=I. |last6=Reeves |first6=J. S. |last7=Wood |first7=B. A. |last8=Fortelius |first8=M. |last9=Bobe |first9=R. |date=1 September 2017 |title=Ecosystem evolution and hominin paleobiology at East Turkana, northern Kenya between 2.0 and 1.4 Ma |url=https://www.sciencedirect.com/science/article/abs/pii/S0031018216307787 |journal=[[Palaeogeography, Palaeoclimatology, Palaeoecology]] |volume=481 |pages=1–13 |doi=10.1016/j.palaeo.2017.05.001 |bibcode=2017PPP...481....1P |access-date=22 January 2023|doi-access=free }}</ref> At the 1.5 Mya [[Trinil H. K. Fauna|Trinil H. K.]] site, Java, ''H. erectus'' likely gathered fish and shellfish.<ref>{{cite journal | vauthors = Joordens JC, Wesselingh FP, de Vos J, Vonhof HB, Kroon D | title = Relevance of aquatic environments for hominins: a case study from Trinil (Java, Indonesia) | journal = [[Journal of Human Evolution]] | volume = 57 | issue = 6 | pages = 656–671 | date = December 2009 | pmid = 19683789 | doi = 10.1016/j.jhevol.2009.06.003 }}</ref>
====Stone tools====
 
Dentally, ''H. erectus'' mouths were not as versatile as those of ancestor species, capable of processing a narrower range of foods. However, tools were likely used to process hard foods, thus affecting the chewing apparatus, and this combination may have instead increased dietary flexibility (though this does not equate to a highly varied diet). Such versatility may have permitted ''H. erectus'' to inhabit a range of different environments, and migrate beyond Africa.<ref name=Ungar2006/>


In 1999, British anthropologist [[Richard Wrangham]] proposed the "cooking hypothesis" which states that ''H. erectus'' speciated from the ancestral ''H. habilis'' because of fire usage and cooking 2 million years ago to explain the rapid doubling of brain size between these two species in only a 500,000 year timespan, and the sudden appearance of the typical human body plan. Cooking makes protein more easily digestible, speeds up nutrient absorption, and destroys food-borne pathogens, which would have increased the environment's natural carrying capacity, allowing group size to expand, causing selective pressure for sociality, requiring greater brain function.<ref name=Gowlett2016/><ref name=Gowlett2013/> However, the fossil record does not associate the emergence of ''H. erectus'' with fire usage nor with any technological breakthrough for that matter, and cooking likely did not become a common practice until after 400 kya.<ref name=Ungar2006/><ref name=BenDor2011/>
[[File:Pièce bifaciale Saint-Acheul La Garenne Archéologie Nationale 01042018.jpg|thumb|A [[handaxe]] from the [[Amiens|Saint-Acheul]] site at the [[National Archaeological Museum, France|Musée d'Archéologie nationale]], France ]]


Java Man's dispersal through Southeast Asia coincides with the [[extirpation]] of the giant turtle ''[[Megalochelys]]'', possibly due to overhunting as the turtle would have been an easy, slow-moving target which could have been stored for quite some time.<ref>{{Cite book|url=http://www.iucn-tftsg.org/cbftt/|title=Conservation Biology of Freshwater Turtles and Tortoises|year=2015|publisher=Chelonian Research Foundation|isbn=978-0-9653540-9-7| veditors = Rhodin A, Pritchard P, van Dijk PP, Saumure R, Buhlmann K, Iverson J, Mittermeier R |edition=First|series=Chelonian Research Monographs|volume=5|doi=10.3854/crm.5.000e.fossil.checklist.v1.2015|page=15}}</ref>
''H. erectus'' manufactured [[Lower Paleolithic]] technologies, and is credited with the invention of the [[Acheulean]] stone tool [[industry (archaeology)|industry]] at latest 1.95 million years ago.<ref>{{cite journal|first1=Margherita|last1=Mussi|first2=Matthew M.|last2=Skinner|first3=Rita T.|last3=Melis|first4=Joaquín|last4=Panera|title=Early ''Homo erectus'' lived at high altitudes and produced both Oldowan and Acheulean tools|url=https://www.science.org/doi/10.1126/science.add9115|journal=Science|year= 2023|pages=713–718|volume=382|issue=6671|doi=10.1126/science.add9115|first5=Susana|last5=Rubio-Jara|first6=Thomas W.|last6=Davies|first7=Denis|last7=Geraads|first8=Hervé|last8=Bocherens|first9=Giuseppe|last9=Briatico|first10=Adeline|last10=Le Cabec|first11=Jean-Jacques|last11=Hublin|first12=Agness|last12=Gidna|first13=Raymonde|last13=Bonnefille|first14=Luca|last14=Di Bianco|first15=Eduardo|last15=Méndez-Quintas|pmid=37824630 |bibcode=2023Sci...382..713M }}</ref> This was a major technological breakthrough featuring large, heavy-duty tools; most iconically, the [[handaxe]]. Over hundreds of thousands of years, the Acheulean eventually replaced its predecessor — the [[Oldowan]] (a [[chopper (archaeology)|chopper]] and [[lithic flake|flake]] industry) — in Africa, and spread out across Western Eurasia.<ref name=Torre2016/> This sudden innovation was typically explained as a response to environmental instability in order to process more types of food and broaden the diet combined with increasing brain size, which also allowed ''H. erectus'' to colonize Eurasia. Despite this characterization of the Acheulean, the small-brained ''H. e. georgicus'' was able to leave Africa despite only manufacturing Oldowan-style tools,<ref name=Carotenuto2016/><ref name=Torre2016>{{cite journal |last=de la Torre|first=I. |title=The origins of the Acheulean: past and present perspectives on a major transition in human evolution |journal=Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences |volume=371 |issue=1698 |article-number=20150245 |year=2016 |pmid=27298475 |pmc=4920301 |doi=10.1098/rstb.2015.0245}}</ref> the 1.6 million year old DAN5/P1 specimen from [[Gona, Ethiopia]] is associated with Acheulean style tools despite its low brain volume of {{cvt|598|cc}},<ref name=Baab2025/> and the handaxe does not seem to have been manufactured commonly in East Asia.<ref>{{cite journal |first1=H. |last1=Li |first2=C. |last2=Li |first3=K. |last3=Kuman |year=2014 |title=Rethinking the "Acheulean" in East Asia: Evidence from recent investigations in the Danjiangkou Reservoir Region, central China |journal=Quaternary International |volume=347 |pages=163–175 |doi=10.1016/j.quaint.2014.03.059|bibcode=2014QuInt.347..163L }}</ref> The lack of East Asian handaxes was first noted by American archaeologist [[Hallam L. Movius]] in 1948, who drew the "[[Movius Line]]", dividing the East into a "chopping-tool culture" and the West into a "hand axe culture".<ref name=Movius1948>{{cite journal |first=H. L. |last=Movius |author-link=Hallam L. Movius |year=1948 |title=The Lower Palaeolithic Cultures of Southern and Eastern Asia |journal=Transactions of the American Philosophical Society |volume=38 |issue=4 |pages=386–403 |doi=10.2307/1005632 |jstor=1005632}}</ref> Movius took this as evidence of inferiority of Far Eastern populations:


===Technology===
{{blockquote|text=...as early as [[Lower Paleolithic|Lower Palaeolithic]] times Southern and Eastern Asia as a whole was a region of cultural retardation...very primitive forms of Early Man apparently persisted there long after types at a comparable stage of physical evolution had become extinct elsewhere.|author=[[Hallam L. Movius]], 1948<ref name=Movius1948/>}}
====Tool production====
{{Multiple image|image1=Canto tallado 2-Guelmim-Es Semara.jpg|caption1=[[Oldowan]] choppers did not become completely replaced until about 1 Mya|image2=Bifaz cordiforme.jpg|caption2=An [[Acheulean]] [[cordiform axe]]|direction=vertical}}
''H. erectus'' is credited with inventing the [[Acheulean]] stone tool industry, succeeding the [[Oldowan]] industry,<ref>{{cite book | vauthors = Beck RB, Black L, Krieger LS, Naylor PC, Shabaka DI |title=World History: Patterns of Interaction |url=https://archive.org/details/mcdougallittellw00beck |url-access=registration |publisher=McDougal Littell |year=1999 |location=Evanston, IL |isbn=978-0-395-87274-1 }}{{page needed|date=December 2019}}</ref><ref>{{cite journal | vauthors = Richards MP | title = A brief review of the archaeological evidence for Palaeolithic and Neolithic subsistence | journal = European Journal of Clinical Nutrition | volume = 56 | issue = 12 | pages = 1270–1278 | date = December 2002 | pmid = 12494313 | doi = 10.1038/sj.ejcn.1601646 | doi-access = free }}</ref> and were the first to make [[lithic flake]]s bigger than {{cvt|10|cm}}, and [[hand axe]]s (which includes bifacial tools with only 2 sides, such as picks, knives, and [[cleaver (tool)|cleavers]]).<ref>{{cite journal | vauthors = de la Torre I | title = The origins of the Acheulean: past and present perspectives on a major transition in human evolution | journal = Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences | volume = 371 | issue = 1698 | pages = 20150245 | date = July 2016 | pmid = 27298475 | pmc = 4920301 | doi = 10.1098/rstb.2015.0245 }}</ref> Though larger and heavier, these hand axes had sharper, chiseled edges.<ref name=Lepre2011/> They were likely multi-purpose tools, used in variety of activities such as cutting meat, wood, or edible plants.<ref name=Nowell2009/> In 1979, American paleontologist Thomas Wynn stated that Acheulean technology required operational intelligence (foresight and planning), being markedly more complex than Oldowan technology which included lithics of unstandardized shape, cross-sections, and symmetry. Based on this, he concluded that there is not a significant disparity in intelligence between ''H. erectus'' and modern humans and that, for the last 300,000 years, increasing intelligence has not been a major influencer of cultural evolution.<ref>{{cite journal| vauthors = Wynn T |year=1979|title=The Intelligence of Later Acheulean Hominids|journal=Man|volume=14|issue=3|pages=371–391|doi=10.2307/2801865|jstor=2801865}}</ref> However, a 1 year old ''H. erectus'' specimen shows that this species lacked an extended childhood required for greater brain development, indicating lower cognitive capabilities.<ref name=Coqueugniot2004/> A few sites, likely due to occupation over several generations, features hand axes en masse, such as at [[Melka Kunture]], Ethiopia; [[Olorgesailie]], Kenya; [[Isimila]], Tanzania; and [[Kalambo Falls]], Zambia.<ref name=Nowell2009/>


The earliest record of Acheulean technology comes from [[West Turkana]], Kenya 1.76 Mya. Oldowan lithics are also known from the site, and the two seemed to coexist for some time. The earliest records of Acheulean technology outside of Africa date to no older than 1 Mya, indicating it only became widespread after some secondary ''H. erectus'' dispersal from Africa.<ref name=Lepre2011>{{cite journal | vauthors = Lepre CJ, Roche H, Kent DV, Harmand S, Quinn RL, Brugal JP, Texier PJ, Lenoble A, Feibel CS | display-authors = 6 | title = An earlier origin for the Acheulian | journal = Nature | volume = 477 | issue = 7362 | pages = 82–85 | date = August 2011 | pmid = 21886161 | doi = 10.1038/nature10372 | s2cid = 4419567 | bibcode = 2011Natur.477...82L }}</ref>
[[File:Homo-erectus Turkana-Boy (Ausschnitt) Fundort Nariokotome, Kenia, Rekonstruktion im Neanderthal Museum.jpg|thumb|left|upright=1.2|Reconstruction of [[Turkana Boy]] at the [[Neanderthal Museum]]]]
''H. erectus'' seems to have been using stone tools in butchery, vegetable processing, and woodworking (maybe manufacturing [[spear]]s and [[digging stick]]s).<ref name=Hardy2018/><ref name=Dominguez2001/> In Africa, Oldowan sites are typically found alongside major fossil assemblages, but Acheulean sites normally feature more stone tools than fossils, so ''H. erectus'' could have been using choppers and handaxes for different activities.<ref name=Dominguez2001>{{cite journal |first1=M. |last1=Dominguez-Rodrigo |first2=J. |last2=Serrallonga |first3=J. |last3=Juan-Tresserras |first4=L. |last4=Alcala |first5=L. |last5=Luque |year=2001 |title=Woodworking activities by early humans: a plant residue analysis on Acheulian stone tools from Peninj (Tanzania) |journal=Journal of Human Evolution |volume=40 |issue=4 |pages=289–299 |doi=10.1006/jhev.2000.0466|pmid=11312582 |bibcode=2001JHumE..40..289D }}</ref> These stone tools probably were not [[hafting|hafted]] onto spears; this innovation is associated with the transition to the [[Middle Paleolithic]] and the emergence of Neanderthals and modern humans.<ref>{{cite journal|first1=J.|last1=Blinkhorn|title=Examining the Origins of Hafting in South Asia|journal=Journal of Paleolithic Archaeology|year=2019|issn=2520-8217|pages=466–481|volume=2|issue=4|doi=10.1007/s41982-019-00034-4|doi-access=free}}</ref>


On Java, ''H. erectus'' produced tools from shells at [[Sangiran]]<ref>{{cite journal| vauthors = Choi K, Driwantoro D |year=2007|title=Shell tool use by early members of ''Homo erectus'' in Sangiran, central Java, Indonesia: cut mark evidence|journal=Journal of Archaeological Science|volume=34|issue=1|pages=48–58|doi=10.1016/j.jas.2006.03.013|bibcode=2007JArSc..34...48C }}</ref> and Trinil.<ref name=":1"/> Spherical stones, measuring {{cvt|6–12|cm}} in diameter, are frequently found in African and Chinese Lower Paleolithic sites, and were potentially used as [[bolas]]; if correct, this would indicate string and cordage technology.<ref>{{cite book|url={{google books|plainurl=yes|id=ODtqDQAAQBAJ|page=6}}| vauthors = Turner J |year=1996|title=History and Science of Knots|publisher=World Scientific|pages=6–8|isbn=9789810224691}}</ref>
Materials for stone tools were normally sourced locally, and it seems [[lithic reduction|blanks]] were usually chosen based on size rather than material quality.<ref name=Torre2016/> ''H. erectus'' also produced tools from shells at Sangiran<ref>{{cite journal |last1=Choi|first1=K.|last2=Driwantoro|first2=D. |year=2007 |title=Shell tool use by early members of ''Homo erectus'' in Sangiran, central Java, Indonesia: cut mark evidence |journal=Journal of Archaeological Science |volume=34 |issue=1 |pages=48–58 |doi=10.1016/j.jas.2006.03.013 |bibcode=2007JArSc..34...48C}}</ref> and Trinil.<ref name=Joordens2015/>
 
{{anchor|Fire}}


====Fire====
====Fire====
{{See also|Control of fire by early humans}}
{{Main|Control of fire by early humans}}
''H. erectus'' is credited as the first human ancestor to have used fire, though the timing of this invention is debated mainly because campfires very rarely and very poorly preserve over long periods of time, let alone thousands or millions of years. The earliest claimed fire sites are in Kenya, FxJj20 at [[Koobi Fora]]<ref>{{cite journal| vauthors = Hlubik S, Berna F, Feibel C, Braun D |year=2017|title=Researching the Nature of Fire at 1.5 Mya on the Site of FxJj20 AB, Koobi Fora, Kenya, Using High-Resolution Spatial Analysis and FTIR Spectrometry|journal=Current Anthropology|volume=58|pages=S243–S257|doi=10.1086/692530|s2cid=148948219}}</ref><ref name=Gowlett2016/><ref name=Roebroeks2011/> and GnJi 1/6E in the [[Chemoigut Formation]], as far back as 1.5 Mya,<ref name=Gowlett2016/><ref name=Roebroeks2011/> and in South Africa, [[Wonderwerk Cave]], 1.7 Mya.<ref name=Beaumont2011>{{cite journal | vauthors = Beaumont PB |year=2011|title=The Edge: More on Fire-Making by about 1.7 Million Years Ago at Wonderwerk Cave in South Africa|journal=Current Anthropology|volume=52|issue=4|pages=585–595|doi=10.1086/660919|s2cid=144176681}}</ref> The first firekeepers are thought to have simply transported to caves and maintained naturally occurring fires for extended periods of time or only sporadically when the opportunity arose. Maintaining fires would require firekeepers to have knowledge on slow-burning materials such as dung.<ref name=Gowlett2016>{{cite journal | vauthors = Gowlett JA | title = The discovery of fire by humans: a long and convoluted process | journal = Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences | volume = 371 | issue = 1696 | pages = 20150164 | date = June 2016 | pmid = 27216521 | pmc = 4874402 | doi = 10.1098/rstb.2015.0164 }}</ref> Fire becomes markedly more abundant in the wider archaeological record after 400,000–300,000 years ago, which can be explained as some advancement in fire management techniques took place at this time<ref name=Gowlett2016/> or human ancestors only opportunistically used fire until this time.<ref name=Roebroeks2011>{{cite journal | vauthors = Roebroeks W, Villa P | title = On the earliest evidence for habitual use of fire in Europe | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 108 | issue = 13 | pages = 5209–5214 | date = March 2011 | pmid = 21402905 | pmc = 3069174 | doi = 10.1073/pnas.1018116108 | doi-access = free | bibcode = 2011PNAS..108.5209R }}</ref><ref name=Sandgathe2017/><ref name=Ungar2006/><ref name=BenDor2011/> It is possible that firestarting was invented and lost and reinvented multiple times and independently by different communities rather than being invented in one place and spreading throughout the world.<ref name=Sandgathe2017>{{cite journal| vauthors = Sandgathe D |year=2017|title=Identifying and Describing Pattern and Process in the Evolution of Hominin Use of Fire|journal=Current Anthropology|volume=58|pages=S360–S370|doi=10.1086/691459|url=https://www.researchgate.net/publication/317042963|hdl=11858/00-001M-0000-002C-0141-3|s2cid=165025762|hdl-access=free}}</ref> The earliest evidence of hearths comes from Gesher Benot Ya'aqov, Israel, over 700,000 years ago, where fire is recorded in multiple layers in an area close to water, both uncharacteristic of natural fires.<ref name=Gowlett2013/>
''H. erectus'' is credited as the first human species to wield fire. The earliest claimed fire site is [[Wonderwerk Cave]], South Africa, at 1.7 million years old.<ref name=Beaumont2011>{{cite journal |last=Beaumont|first=P. B. |year=2011 |title=The Edge: More on Fire-Making by about 1.7 Million Years Ago at Wonderwerk Cave in South Africa |journal=Current Anthropology |volume=52 |issue=4 |pages=585–595 |doi=10.1086/660919 |s2cid=144176681}}</ref> While the species' dispersal far out of Africa has often been attributed to fire and cave dwelling, fire does not become common in the archaeological record until 300,000 to 400,000 years ago,<ref name=Roebroeks2011>{{cite journal |last1=Roebroeks|first1=W.|last2=Villa|first2=P. |title=On the earliest evidence for habitual use of fire in Europe |journal=Proceedings of the National Academy of Sciences of the United States of America |volume=108 |issue=13 |pages=5209–5214 |year=2011 |pmid=21402905 |pmc=3069174 |doi=10.1073/pnas.1018116108 |doi-access=free |bibcode=2011PNAS..108.5209R}}</ref> and cave-dwelling about 600,000 years ago.<ref>{{cite journal |url=https://www.researchgate.net/publication/303107882 |last1=Ullman|first1=M.|last2=Hovers|first2=E.|last3=Goren-Inbar|first3=N.|last4=Frumkin|first4=A. |year=2013 |title=Levantine cave dwellers: geographic and environmental aspects of early humans use of caves, case study from Wadi Amud, northern Israel |journal=International Congress of Speleology |volume=1 |page=169}}</ref> Therefore, ''H. erectus'' may have only been scavenging fire opportunistically. Similarly, ''H. erectus'' sites usually stay within warmer tropical or subtropical latitudes.<ref name=Carotenuto2016>{{cite journal |last1=Carotenuto|first1=F.|last2=Tsikaridze|first2=N.|last3=Rook|first3=L.|last4=Lordkipanidze|first4=D.|last5=Longo|first5=L.|last6=Condemi|first6=S.|last7=Raia|first7=P. |title=Venturing out safely: The biogeography of ''Homo erectus'' dispersal out of Africa |journal=Journal of Human Evolution |volume=95 |pages=1–12 |year=2016 |pmid=27260171 |doi=10.1016/j.jhevol.2016.02.005 |bibcode=2016JHumE..95....1C |hdl=10356/82274 |hdl-access=free}}</ref>


Artificial lighting may have led to increased waking hours—modern humans have about a 16-hour waking period, whereas other apes are generally awake from only sunup to sundown—and these additional hours were probably used for socializing. Because of this, fire usage is probably also linked to the [[origin of language]].<ref name=Gowlett2016/><ref name=Gowlett2013/> Artificial lighting may have also made sleeping on the ground instead of the trees possible by keeping terrestrial predators at bay.<ref name=Gowlett2013/>
The dating of northerly populations (namely Peking Man) could suggest that they were retreating to warmer refugia during [[glacial period]]s, but the precise age of the Peking Man fossils is poorly resolved.{{sfn|Antón|2003|loc=p. 132}}<ref name="auto"/> There have been claims of manmade hearths and "clear-cut evidence for intentional fire use",<ref>{{cite journal |first1=X. |last1=Gao |first2=S. |last2=Zhang |first3=Y. |last3=Zhang |first4=F. |last4=Chen |year=2017 |title=Evidence of Hominin Use and Maintenance of Fire at Zhoukoudian |journal=Current Anthropology |volume=58 |issue=S16 |pages=S267–S277 |doi=10.1086/692501 |s2cid=164616520 |url=https://www.researchgate.net/publication/317987720}}</ref> ostensibly as far back as 770,000 years ago in the supposed cave home of Peking Man.<ref name="auto">{{cite journal |last1=Zhong |first1=M. |last2=Shi |first2=C. |display-authors=et al. |year=2013 |title=On the possible use of fire by ''Homo erectus'' at Zhoukoudian, China |journal=Chinese Science Bulletin |volume=59 |issue=3 |pages=335–343 |doi=10.1007/s11434-013-0061-0 |s2cid=93590269}}</ref> At the French Caune de L'Arago, Tautavel Man does not seem to have been using fire at all, even though occupation sequences span two cold periods.<ref name=Lumley2015/>


Migration into the frigid climate of Ice Age Europe may have only been possible because of fire, but evidence of fire usage in Europe until about 400–300,000 years ago is notably absent.<ref name=Roebroeks2011/> If these early European ''H. erectus'' did not have fire, it is largely unclear how they stayed warm, avoided predators, and prepared animal fat and meat for consumption. There was also a lower likelihood of naturally occurring fires due to lightning being less common in areas further north. It is possible that they only knew how to maintain fires in certain settings in the landscapes and prepared food some distance away from home, meaning evidence of fire and evidence of hominin activity are spaced far apart.<ref name=Gowlett2013>{{cite journal| vauthors = Gowlett JA, Wrangham RW |year=2013|title=Earliest fire in Africa: Towards the convergence of archaeological evidence and the cooking hypothesis|journal=Azania: Archaeological Research in Africa|volume=48|issue=1|pages=5–30|doi=10.1080/0067270X.2012.756754|s2cid=163033909|url=https://www.researchgate.net/publication/271530765}}</ref> Alternatively, ''H. erectus'' may have only pushed farther north during warmer [[interglacial]] periods—thus not requiring fire, food storage, or clothing technology—<ref>{{cite journal | vauthors = Antón SC | title = Natural history of Homo erectus | journal = American Journal of Physical Anthropology | volume = 122 | issue = S37 | pages = 126–170 | year = 2003 | pmid = 14666536 | doi = 10.1002/ajpa.10399 }}</ref> and their dispersal patterns indicate they generally stayed in warmer lower-to-middle latitudes.<ref name=Carotenuto2016>{{cite journal | vauthors = Carotenuto F, Tsikaridze N, Rook L, Lordkipanidze D, Longo L, Condemi S, Raia P | title = Venturing out safely: The biogeography of Homo erectus dispersal out of Africa | journal = Journal of Human Evolution | volume = 95 | pages = 1–12 | date = June 2016 | pmid = 27260171 | doi = 10.1016/j.jhevol.2016.02.005 | hdl = 10356/82274 | hdl-access = free }}</ref> It is debated if the ''H. e. pekinensis'' inhabitants of [[Zhoukoudian]], Northern China, were capable of controlling fires as early as 770 kya to stay warm in what may have been a relatively cold climate.<ref>{{cite journal| vauthors = Zhong M, Shi C, Gao X, Wu X, Chen F, Zhang S, Zhang X, Olsen JW |year=2013|title=On the possible use of fire by ''Homo erectus'' at Zhoukoudian, China|journal=Chinese Science Bulletin|volume=59|issue=3|pages=335–343|doi=10.1007/s11434-013-0061-0|s2cid=93590269|url=https://www.researchgate.net/publication/272016716}}</ref>
====Healthcare====
 
{{See also|Prehistoric medicine}}
==== Construction ====
[[File:Dmanisi fossils D 3444 + D 3900 (Replika).jpg|thumb|upright=0.8|The single-toothed ''[[Dmanisi hominins|H. e. georgicus]]'' specimen (above) is the earliest probable example of human group care.<ref name=Spikins2019/>]]
[[File:Terra-Amata-Hut.gif|thumb|Reconstruction of a [[Terra Amata (archaeological site)|Terra Amata]] dwelling<ref>{{cite web|url=https://www.musee-terra-amata.org/musee/le-site-acheuleen-de-terra-amata/ |title=Le site acheuléen de Terra Amata |language=fr |trans-title=The Acheulean site of Terra Amata |author=Musée de Préhistoire Terra Amata |website=Musée de Préhistoire Terra Amata |access-date=10 June 2022}}</ref>]]
Like other primates, ''H. erectus'' probably used [[medicinal plant]]s<ref name=Hardy2018/> and infirmed sick group members. The earliest probable example of human group care is a 1.77 million year old ''H. e. georgicus'' specimen who had lost all but one tooth due to age or [[gum disease]] (the earliest example of severe chewing impairment) yet still survived for several years afterwards.<ref name=Spikins2019>{{cite journal |last1=Spikins|first1=P.|last2=Needham|first2=A.|last3=Wright|first3=B|last4=Dytham|first4=C|last5=Gatta |first5=M.|last6=Hitchens|first6=G. |year=2019 |title=Living to fight another day: The ecological and evolutionary significance of Neanderthal healthcare |journal=Quaternary Science Reviews |volume=217 |pages=98–118 |doi=10.1016/j.quascirev.2018.08.011 |bibcode=2019QSRv..217...98S |doi-access=free}}</ref>
In 1962, a {{cvt|12x14x1|ft|cm|order=flip}} circle made with volcanic rocks was discovered in [[Olduvai Gorge]]. At {{cvt|2–2.5|ft|cm|order=flip|adj=on}} intervals, rocks were piled up to {{cvt|6–9|in|cm|order=flip}} high. British palaeoanthropologist [[Mary Leakey]] suggested the rock piles were used to support poles stuck into the ground, possibly to support a [[windbreak]] or a rough hut. Some modern day nomadic tribes build similar low-lying rock walls to build temporary shelters upon, bending upright branches as poles and using grasses or animal hide as a screen.<ref>{{cite book|url={{google books|plainurl=yes|id=eepULHufmF8C|page=24}} |vauthors=Leakey MD |author-link=Mary Leakey |year=1971 |title=Olduvai Gorge: Volume 3, Excavations in Beds I and II, 1960-1963 |publisher=[[Cambridge University Press]] |page=24 |isbn=9780521077231}}</ref> Dating to 1.75 Mya, it is the oldest claimed evidence of architecture.<ref>{{cite book|vauthors=Ingold T |year=2000 |title=The Perception of the Environment: Essays on Livelihood, Dwelling and Skill |chapter=Building, dwelling, living: how animals and people make themselves at home in the world |publisher=Psychology Press |page=184 |url={{google books|plainurl=yes|id=S3GakE5OT-kC|page=184}} |isbn=9780415228329}}</ref>
 
In Europe, evidence of constructed dwelling structures dating to or following the [[Holstein Interglacial]] (which began 424 kya) has been claimed in Bilzingsleben, Germany; [[Terra Amata (archaeological site)|Terra Amata]], France; and [[Fermanville]] and [[Saint-Germain-des-Vaux]] in [[Normandy]]. The oldest evidence of a dwelling (and a campfire) in Europe comes from [[Přezletice]], Czech Republic, 700 kya during the [[Cromerian Interglacial]]. This dwelling's base measured about {{cvt|3x4|m}} on the exterior and {{cvt|3x2|m}} on the interior, and is considered to have been a firm surface hut, probably with a vaulted roof made of thick branches or thin poles, supported by a foundation of big rocks and earth, and likely functioned as a winter base camp.<ref>{{cite journal|vauthors=Sklenář K |year=1987|title=The Lower Paleolithic Dwelling Structure at Přezletice and its Significance |journal=Anthropologie |volume=25 |issue=2 |pages=101–103 |jstor=26294864}}</ref>
 
The earliest evidence of cave habitation is Wonderwerk Cave, South Africa, about 1.6 Mya, but evidence of cave use globally is sporadic until about 600 kya.<ref>{{cite journal|url=https://www.researchgate.net/publication/303107882 |vauthors=Ullman M, Hovers E, Goren-Inbar N, Frumkin A |year=2013 |title=Levantine cave dwellers: geographic and environmental aspects of early humans use of caves, case study from Wadi Amud, northern Israel |journal=International Congress of Speleology |volume=1}}</ref>
 
====Clothing====
[[File:Homo-erectus Turkana-Boy (Ausschnitt) Fundort Nariokotome, Kenia, Rekonstruktion im Neanderthal Museum.jpg|thumb|upright=1.3|left|Reconstruction of Turkana boy with light clothing by Adrie and Alfons Kennis at the [[Neanderthal Museum]]]]
It is largely unclear when clothing was invented, with the earliest estimate stretching as far back as 3 Mya to compensate for a lack of insulating body hair.<ref name=Gilligan2010>{{cite journal| vauthors = Gilligan I |year=2010|title=The Prehistoric Development of Clothing: Archaeological Implications of a Thermal Model|journal=Journal of Archaeological Method and Theory |volume=15|pages=15–80|doi=10.1007/s10816-009-9076-x|s2cid=143004288}}</ref> It is known that [[head lice]] and [[body lice]] (the latter can only inhabit clothed individuals) for modern humans diverged about 170 kya, well before modern humans left Africa, meaning clothes were already well in use before encountering cold climates. One of the first uses of animal hide is thought to have been for clothing, and the oldest hide scrapers date to about 780 kya, though this is not indicative of clothing.<ref>{{cite journal | vauthors = Toups MA, Kitchen A, Light JE, Reed DL | title = Origin of clothing lice indicates early clothing use by anatomically modern humans in Africa | journal = Molecular Biology and Evolution | volume = 28 | issue = 1 | pages = 29–32 | date = January 2011 | pmid = 20823373 | pmc = 3002236 | doi = 10.1093/molbev/msq234 }}</ref>


====Seafaring====
====Seafaring====
Acheulean artifacts discovered on isolated islands that were never connected to land in the Pleistocene may show seafaring by ''H. erectus'' as early as 1 Mya in Indonesia. They had arrived on the islands of [[Flores]], [[Timor]], and [[Rote Island|Roti]], which would have necessitated crossing the [[Lombok Strait]] (the [[Wallace Line]]), at least before 800 kya. It is also possible they were the first European mariners as well and crossed the [[Strait of Gibraltar]] between North Africa and Spain. A 2021 genetic analysis of these island populations of ''H. erectus'' found no evidence of interbreeding with modern humans.<ref>{{Cite web|date=23 March 2021|title=New evidence in search for the mysterious Denisovans|url=https://www.sciencedaily.com/releases/2021/03/210323084732.htm|access-date=30 March 2021|website=ScienceDaily|language=en}}</ref> Seafaring capability would show ''H. erectus'' had a great capacity for planning, likely months in advance of the trip.<ref name=Bednarik1999>{{cite journal| vauthors = Bednarik RG |year=1999 |title= Pleistocene seafaring in the Mediterranean|journal=Anthropologie|volume=37|issue=3|pages=275–282|jstor=26294895}}</ref><ref>{{cite journal| vauthors = Bednarik RG |year=1998|title=An experiment in Pleistocene seafaring|journal=The International Journal of Nautical Archaeology|volume=27|issue=2|pages=139–149|doi=10.1111/j.1095-9270.1998.tb00797.x|url=http://www.ifrao.com/wp-content/uploads/2015/03/98Nautuical.pdf}}</ref>
''H. erectus'' made long sea crossings to arrive on the islands of [[Flores]], [[Luzon]],<ref name="Détroit2019">{{cite journal |last1=Détroit|first1=F.|last2=Mijares|first2=A. S.|last3=Corny|first3=J.|last4=Daver|first4=G.|last5=Zanolli|first5=C.|last6=Dizon|first6=E.|last7=Robles|first7=E.|last8=Grün|first8=R.|last9=Piper|first9=P. J. |title=A new species of ''Homo'' from the Late Pleistocene of the Philippines |journal=Nature |volume=568 |issue=7751 |pages=181–186 |year=2019 |pmid=30971845 |doi=10.1038/s41586-019-1067-9 |name-list-style=amp |s2cid=106411053 |bibcode=2019Natur.568..181D |hdl=10072/386785 |url=https://hal.archives-ouvertes.fr/hal-02296712/file/Detroit_%26_al_2019_Nature_postprint.pdf}}</ref> and some [[Mediterranean Sea|Mediterranean]] islands. Some authors have asserted that ''H. erectus'' intentionally made these crossings by inventing watercrafts and seafaring so early in time, speaking to advanced cognition and language skills. These populations could have also been founded by [[oceanic dispersal|natural rafting events]] instead.<ref name=":6">{{Cite journal |last=Botha |first=Rudolf |year=2024 |title=Did ''Homo erectus'' Have Language? The Seafaring Inference |journal=Cambridge Archaeological Journal |volume=35 |language=en |pages=21–37 |doi=10.1017/S0959774324000118 |issn=0959-7743 |doi-access=free}}</ref>
 
Similarly, ''[[Homo luzonensis]]'' is dated between 771,000 and 631,000 years ago. Because Luzon has always been an island in the Quaternary, the ancestors of ''H. luzonensis'' would have had to have made a substantial sea crossing and crossed the [[Huxley Line]].<ref name="Détroit2019">{{cite journal | vauthors = Détroit F, Mijares AS, Corny J, Daver G, Zanolli C, Dizon E, Robles E, Grün R, Piper PJ | display-authors = 6 | title = A new species of Homo from the Late Pleistocene of the Philippines | journal = Nature | volume = 568 | issue = 7751 | pages = 181–186 | date = April 2019 | pmid = 30971845 | doi = 10.1038/s41586-019-1067-9 | name-list-style = amp | s2cid = 106411053 | bibcode = 2019Natur.568..181D | url = https://hal.archives-ouvertes.fr/hal-02296712/file/Detroit_%26_al_2019_Nature_postprint.pdf }}</ref>
 
====Healthcare====
[[File:Dmanisi fossils D 3444 + D 3900 (Replika).jpg|thumb|upright|Skull of a toothless ''H. e. georgicus'']]
The earliest probable example of infirming sick group members is a 1.77 Mya ''H. e. georgicus'' specimen who had lost all but one tooth due to age or [[gum disease]], the earliest example of severe chewing impairment, yet still survived for several years afterwards. However, it is possible australopithecines were capable of caring for debilitated group members.<ref>{{cite journal| vauthors = Spikins P, Needham A, Wright B, Dytham C, Gatta M, Hitchens G |year=2019 |title= Living to fight another day: The ecological and evolutionary significance of Neanderthal healthcare|journal=Quaternary Science Reviews|volume=217|pages=98–118|doi=10.1016/j.quascirev.2018.08.011|bibcode=2019QSRv..217...98S|doi-access=free}}</ref> Unable to chew, this ''H. e. georgicus'' individual probably ate soft plant or animal foods possibly with assistance from other group members. High-latitude groups are thought to have been predominantly carnivorous, eating soft tissue such as [[bone marrow]] or brains, which may have increased survival rates for toothless individuals.<ref>{{cite journal | vauthors = Lordkipanidze D, Vekua A, Ferring R, Rightmire GP, Agusti J, Kiladze G, Mouskhelishvili A, Nioradze M, Ponce de León MS, Tappen M, Zollikofer CP | display-authors = 6 | title = Anthropology: the earliest toothless hominin skull | journal = Nature | volume = 434 | issue = 7034 | pages = 717–718 | date = April 2005 | pmid = 15815618 | doi = 10.1038/434717b | bibcode = 2005Natur.434..717L | s2cid = 52800194 }}</ref>
 
The 1.5 Mya Turkana boy was diagnosed with juvenile [[spinal disc herniation]], and, because this specimen was still growing, this caused some [[scoliosis]] (abnormal curving of the spine). These usually cause recurrent lower back pain and [[sciatica]] (pain running down the leg), and likely restricted Turkana boy in walking, bending, and other daily activities. The specimen appears to have survived into adolescence, which evidences advanced group care.<ref>{{cite journal | vauthors = Haeusler M, Schiess R, Boeni T | title = Evidence for juvenile disc herniation in a homo erectus boy skeleton | journal = Spine | volume = 38 | issue = 3 | pages = E123–E128 | date = February 2013 | pmid = 23154836 | doi = 10.1097/BRS.0b013e31827cd245 | s2cid = 11534863 | url = https://www.zora.uzh.ch/id/eprint/76396/1/Haeusler_et_al_Evidence_for_juvenile_disc_herniation.pdf }}</ref>
 
The 1,000–700 kya Java man specimen presents a noticeable [[osteocyte]] on the femur, likely [[Paget's disease of bone]], and [[osteopetrosis]], thickening of the bone, likely resulting from [[skeletal fluorosis]] caused by ingestion of food contaminated by fluorine-filled volcanic ash (as the specimen was found in ash-filled [[stratum (geology)|strata]]). Livestock that grazes on volcanic ash ridden fields typically die of acute intoxication within a few days or weeks.<ref>{{cite journal | vauthors = Soriano M | title = The fluoric origin of the bone lesion in the Pithecanthropus erectus femur | journal = American Journal of Physical Anthropology | volume = 32 | issue = 1 | pages = 49–57 | date = January 1970 | pmid = 4984453 | doi = 10.1002/ajpa.1330320107 }}</ref>


=== Art and rituals ===
=== Art and rituals ===
{{See also|Prehistoric art}}
{{See also|Prehistoric art}}
{{Multiple image|width=300|direction=vertical|image1=Homo Erectus shell with geometric incisions circa 500,000 BP, Naturalis Biodiversity Center, Netherlands (with detail).jpg|caption1=Engraved [[Pseudodon shell DUB1006-fL]] from [[Trinil]], [[Java]]|image2=Museo de la Evolucion Humana Burgos - Tan Tan and Berekhat Ram Pebbles.jpg|caption2=Replicas of the "[[Venus of Tan-Tan]]" (left) and "[[Venus of Berekhat Ram]]" (right)}}
An engraved [[Pseudodon shell DUB1006-fL]] with geometric markings could possibly be evidence of the earliest art-making, dating back to 546–436 kya. Art-making capabilities could be considered evidence of symbolic thinking, which is associated with modern cognition and behavior.<ref name=":1">{{cite journal | vauthors = Joordens JC, d'Errico F, Wesselingh FP, Munro S, de Vos J, Wallinga J, Ankjærgaard C, Reimann T, Wijbrans JR, Kuiper KF, Mücher HJ, Coqueugniot H, Prié V, Joosten I, van Os B, Schulp AS, Panuel M, van der Haas V, Lustenhouwer W, Reijmer JJ, Roebroeks W | display-authors = 6 | title = Homo erectus at Trinil on Java used shells for tool production and engraving | journal = Nature | volume = 518 | issue = 7538 | pages = 228–231 | date = February 2015 | pmid = 25470048 | doi = 10.1038/nature13962 | s2cid = 4461751 | bibcode = 2015Natur.518..228J }}</ref><ref>{{cite journal | vauthors = Henshilwood CS, d'Errico F, Watts I | title = Engraved ochres from the Middle Stone Age levels at Blombos Cave, South Africa | journal = Journal of Human Evolution | volume = 57 | issue = 1 | pages = 27–47 | date = July 2009 | pmid = 19487016 | doi = 10.1016/j.jhevol.2009.01.005 }}</ref><ref>{{cite journal | vauthors = d'Errico F, Moreno RG, Rifkin RF | year = 2012 | title = Technological, elemental and colorimetric analysis of an engraved ochre fragment from the Middle Stone Age levels of Klasies River Cave 1, South Africa | journal = J. Archaeol. Sci. | volume = 39 | issue = 4| pages = 942–952 | doi = 10.1016/j.jas.2011.10.032 | bibcode = 2012JArSc..39..942D }}</ref><ref name="Nature Article">{{cite journal | vauthors = Callaway E |title=''Homo erectus'' made world's oldest doodle 500,000 years ago |journal=Nature News |doi=10.1038/nature.2014.16477 |url=https://www.nature.com/news/homo-erectus-made-world-s-oldest-doodle-500-000-years-ago-1.16477|year=2014 |s2cid=164153158 }}</ref> In 1976, American archeologist [[Alexander Marshack]] asserted that engraved lines on an ox rib, associated with Acheulean lithics, from [[Pech de l'Azé]], France, are similar to a [[meander (art)|meander design]] found in modern human Upper Paleolithic cave art.<ref name=Dickson1992>{{cite book| vauthors = Dickson DB |year=1992|title=The Dawn of Belief: Religion in the Upper Paleolithic of Southwestern Europe|publisher=University of Arizona Press|pages=40–46|url={{google books|plainurl=yes|id=DNr5YIygjMMC|page=40}}|isbn=978-0-8165-1336-9}}</ref> Three [[ostrich eggshell beads]] associated with Achuelian lithics were found in northwestern Africa, the earliest disc beads ever found, and Acheulian disc beads have also been found in France and Israel.<ref name=Bednarik1999/> The Middle Pleistocene "[[Venus of Tan-Tan]]" and "[[Venus of Berekhat Ram]]" are postulated to been crafted by ''H. erectus'' to resemble a human form. They were mostly formed by natural weathering, but slightly modified to emphasize certain grooves to suggest hairline, limbs, and eyes.<ref name=Morriss2009/><ref>{{cite journal| vauthors = d'Errico F, Nowell A |year=2000|title=A New Look at the Berekhat Ram Figurine: Implications for the Origins of Symbolism|journal=Cambridge Archaeological Journal|volume=10|issue=1|pages=123–167|doi=10.1017/S0959774300000056|s2cid=163138037}}</ref> The former has traces of pigments on the front side, possibly indicating it was colored.<ref name=Morriss2009>{{cite journal | vauthors = Morriss-Kay GM | title = The evolution of human artistic creativity | journal = Journal of Anatomy | volume = 216 | issue = 2 | pages = 158–176 | date = February 2010 | pmid = 19900185 | pmc = 2815939 | doi = 10.1111/j.1469-7580.2009.01160.x }}</ref>


''H. erectus'' was also the earliest human to have intentionally collected red-colored pigments, namely [[ochre]], recorded as early as the Middle Pleistocene. Ochre lumps at [[Olduvai Gorge]], Tanzania—associated with the 1.4 Ma [[Olduvai Hominid 9]]—and [[Ambrona]], Spain—which dates to 424–374 kya—were suggested to have been struck by a hammerstone and purposefully shaped and trimmed.<ref name=Watts2014/><ref name=Dickson1992/> At Terra Amata, France—which dates to 425–400 or 355–325 kya—red, yellow, and brown ochres were recovered in association with pole structures; ochre was probably heated to achieve such a wide color range.<ref name=Watts2014>{{cite book| vauthors = Watts I |year=2014|chapter=The red thread: pigment use and the evolution of collective ritual|title=The Social Origins of Language|publisher=Oxford University Press|pages=222–223|url={{google books|plainurl=yes|id=0HtYCwAAQBAJ|page=222}}|isbn=978-0-19-966533-4}}</ref><ref>{{cite book| vauthors = de Lumley H, Boone Y |year=1976|chapter=Les structures d'habitat au Paléolithique moyen|trans-chapter=Housing structures from the lower Paleolithic|title=La Préhistoire française: Les civilisations paléolithiques et mésolithiques de la France|trans-title=French prehistory: the Paleolithic and Mesolithic civilizations of France| veditors = de Lumley H, Guilaine J |publisher=Éditions du Centre national de la recherche scientifique|isbn=978-2-222-01968-8}}</ref> As it is unclear if ''H. erectus'' could have used ochre for any practical application, ochre collection might indicate that ''H. erectus'' was the earliest human to have exhibited a sense of [[aesthetics]] and to think beyond simply survival. Later human species are postulated to have used ochre as body paint, but in the case of ''H. erectus'', it is contested if body paint was used so early in time. Further, it is unclear if these few examples are not simply isolated incidents of ochre use, as ochre is much more prevalent in Middle and Upper Paleolithic sites attributed to Neanderthals and ''H. sapiens''.<ref name=Wreschner1980>{{cite journal|url=http://sites.utexas.edu/butzer/files/2017/03/Wreschner_Butzer-1980-RedOchre.pdf| vauthors = Wreschner EE, Bolton R, Butzer KW, Delporte H, Häusler A, Heinrich A, Jacobson-Widding A, Malinowski T, Masset C, Miller SF, Ronen A | display-authors = 6 |year=1980|title=Red Ochre and Human Evolution: A Case for Discussion|journal=Current Anthropology|volume=21|issue=5|pages=632–633<!--only citing these pages-->|doi=10.1086/202541|jstor=2741829|s2cid=88099778}}</ref><ref name=Dickson1992/>
In East Asia, ''H. erectus'' is usually represented only by skullcaps, which used to be interpreted as widespread [[human cannibalism|cannibalism]] and ritual [[headhunting]]. This had been reinforced by the historic practice of headhunting and cannibalism in some recent Indonesian, Australian, and Polynesian cultures, which were formerly believed to have directly descended from these ''H. erectus'' populations. The lack of the rest of the skeleton is now normally explained by natural phenomena.<ref>{{cite journal |first=T. |last=Jacob |author-link=Teuku Jacob |year=1972 |title=The Problem of Head-Hunting and Brain-Eating among Pleistocene Men in Indonesia |journal=Archaeology and Physical Anthropology in Oceania |volume=7 |issue=2 |pages=86–89 |jstor=40386169}}</ref>


In 1935, Jewish-German anthropologist [[Franz Weidenreich]] speculated that the inhabitants of the Chinese [[Zhoukoudian|Zhoukoudian Peking Man site]] were members of some Lower Paleolithic Skull Cult because the skulls all showed fatal blows to the head, breaking in of the [[foramen magnum]] at the base of the skull, by-and-large lack of preserved facial aspects, an apparently consistent pattern of breaking on the mandible, and a lack of post-cranial remains (elements that are not the skull). He believed that the inhabitants were [[headhunting|headhunters]], and smashed open the skulls and ate the brains of their victims.<ref>{{cite journal| vauthors = Weidenreich F |author-link=Franz Weidenreich|year=1935|title=The ''Sinanthropus'' Population of Choukoutien (Locality 1) with a Preliminary Report on New Discoveries|journal=Bulletin of the Geological Society of China|volume=14|issue=4|pages=427–468|doi=10.1111/j.1755-6724.1935.mp14004001.x}}</ref><ref name=Dickson1992/> However, scavenging animals and natural forces such as flooding can also inflict the same kind of damage to skulls,<ref name=Dickson1992/> and there is not enough evidence to suggest manhunting or cannibalism.<ref>{{cite journal| vauthors = Binford LR, Ho CK |author-link=Lewis Binford |year=1985|title=Taphonomy at a Distance: Zhoukoudian, 'The Cave Home of Beijing Man'?|journal=Current Anthropology|volume=26|issue=4|pages=413–442|doi=10.1086/203303|jstor=2742759|s2cid=147164100}}</ref>
[[File:Homo Erectus shell with geometric incisions circa 500,000 BP, Naturalis Biodiversity Center, Netherlands (with detail).jpg|thumb|upright=1.5|Engraved ''[[Pseudodon]]'' shell [[Pseudodon shell DUB1006-fL|DUB1006-fL]] from [[Trinil]], [[Java]]]]


In 1999, British science writers [[Marek Kohn]] and [[Steven Mithen]] said that many hand axes exhibit no wear and were produced en masse, and concluded that these symmetrical, tear-drop shaped lithics functioned primarily as [[display (zoology)|display]] tools so males could prove their fitness to females in some courting ritual, and were discarded afterwards.<ref>{{cite journal| vauthors = Kohn M, Mithen S |author-link=Marek Kohn|author2-link=Steven Mithen|year=1999|title=Handaxes: products of sexual selection?|journal=Antiquity|volume=73|issue=281|pages=518–526|doi=10.1017/S0003598X00065078|s2cid=162903453}}</ref> However, an apparent lack of reported wearing is likely due to a lack of use-wear studies, and only a few sites yield an exorbitant sum of hand axes likely due to gradual accumulation over generations instead of mass production.<ref name=Nowell2009>{{cite journal| vauthors = Nowell A, Chang ML |year=2009|title=The Case Against Sexual Selection as an Explanation of Handaxe Morphology |journal=PaleoAnthropology |pages=77–88|url=http://paleoanthro.reedd.webfactional.com/static/journal/content/PA20090077.pdf}}</ref>
Art-making could be evidence of symbolic thinking. An engraved ''[[Pseudodon]]'' shell [[Pseudodon shell DUB1006-fL|DUB1006-fL]] from Trinil, Java, with geometric markings could possibly be the earliest example of art-making, dating to 436,000 to 546,000 years ago.<ref name=Joordens2015>{{cite journal |last1=Joordens|first1=J. C.|last2=d'Errico|first2=F.|last3=Wesselingh|first3=F. P.|last4=Munro|first4=S.|last5=de Vos|first5=J|last6=Wallinga|first6=J.|last7=Ankjærgaard|first7=C.|last8=Reimann|first8=T.|last9=Wijbrans|first9=J. R.|last10=Kuiper|first10=K. F.|last11=Mücher|first11=H. J.|last12=Coqueugniot|first12=H.|last13=Prié|first13=V|last14=Joosten|first14=I.|last15=van Os|first15=B.|last16=Schulp|first16=A. S.|last17=Panuel|first17=M.|last18=van der Haas|first18=V.|last19=Lustenhouwer|first19=W.|last20=Reijmer|first20=J. J.|last21=Roebroeks|first21=W. |title=''Homo erectus'' at Trinil on Java used shells for tool production and engraving |journal=Nature |volume=518 |issue=7538 |pages=228–231 |year=2015 |pmid=25470048 |doi=10.1038/nature13962 |s2cid=4461751 |bibcode=2015Natur.518..228J}}</ref><ref>{{cite journal |last=Callaway|first=E. |title=''Homo erectus'' made world's oldest doodle 500,000 years ago |journal=Nature News |doi=10.1038/nature.2014.16477 |url=https://www.nature.com/news/homo-erectus-made-world-s-oldest-doodle-500-000-years-ago-1.16477 |year=2014 |s2cid=164153158|url-access=subscription }}</ref><ref name=Dickson1992>{{cite book |last=Dickson|first=D. B. |year=1992 |title=The Dawn of Belief: Religion in the Upper Paleolithic of Southwestern Europe |publisher=University of Arizona Press |pages=40–46 |url={{google books |plainurl=yes |id=DNr5YIygjMMC |page=40}} |isbn=978-0-8165-1336-9}}</ref> ''H. erectus'' was also the earliest human to collect red-colored pigments, namely [[ochre]]. Ochre lumps at Olduvai Gorge, Tanzania, associated with the 1.4 million year old Olduvai Hominid 9 may have been purposefully shaped and trimmed by a [[hammerstone]]. Red ochre is normally recognized as bearing symbolic value when associated with modern humans.<ref name=Dickson1992/>


===Language===
===Language===
In 1984, the vertebral column of the 1.6 Mya adolescent [[Turkana boy]] indicated that this individual did not have properly developed respiratory muscles in order to produce speech. In 2001, American anthropologists Bruce Latimer and James Ohman concluded that Turkana boy was afflicted by [[skeletal dysplasia]] and [[scoliosis]].<ref>{{cite journal| vauthors = Latimer B, Ohman J |year=2001|title=Axial dysplasia in ''Homo erectus''|journal=Journal of Human Evolution|volume=40}}</ref> In 2006, American anthropologist [[Marc Meyer]] and colleagues described a 1.8 Mya ''H. e. georgicus'' specimen as having a spine within the range of variation of modern human spines, contending that Turkana boy had [[spinal stenosis]] and was thus not representative of the species. Also, because he considered ''H. e. georgicus'' ancestral to all non-African ''H. erectus'', Meyer concluded that the respiratory muscles of all ''H. erectus'' (at least non-''H. ergaster'') would not have impeded vocalisation or speech production.<ref>{{cite conference | vauthors = Meyer M, Lordkipanidze D, Vekua A | title = Language and empathy in Homo erectus: Behaviors suggested by a modern spinal cord from Dmanisi, but not Nariokotome. | conference = Annual meeting of the Paleoanthroplogy Society | location = San Juan, Puerto Rico | date = 2006 | url = https://www.researchgate.net/publication/263198625 }}</ref> However, in 2013 and 2014, anthropologist Regula Schiess and colleagues concluded that there is no evidence of any congenital defects in Turkana boy, and considered the specimen representative of the species.<ref>{{cite journal | vauthors = Schiess R, Haeusler M | title = No skeletal dysplasia in the Nariokotome boy KNM-WT 15000 (Homo erectus)--a reassessment of congenital pathologies of the vertebral column | journal = American Journal of Physical Anthropology | volume = 150 | issue = 3 | pages = 365–374 | date = March 2013 | pmid = 23283736 | doi = 10.1002/ajpa.22211 }}</ref><ref>{{cite journal | vauthors = Schiess R, Boeni T, Rühli F, Haeusler M | title = Revisiting scoliosis in the KNM-WT 15000 Homo erectus skeleton | journal = Journal of Human Evolution | volume = 67 | issue = 48–59 | pages = 48–59 | date = February 2014 | pmid = 24491377 | doi = 10.1016/j.jhevol.2013.12.009 | url = https://www.zora.uzh.ch/id/eprint/92705/1/Schiess_et_al._-_Scoliosis_in_WT15000_2013-10-24.pdf }}</ref>
{{See also|Origin of language|Origin of speech}}
 
The [[spinal column]] of the 1.6 million year old Turkana boy would not have supported properly developed respiratory muscles required to produce speech;<ref>{{cite journal |last1=Latimer|first1=B.|last2=Ohman|first2=J. |year=2001 |title=Axial dysplasia in ''Homo erectus'' |journal=Journal of Human Evolution |volume=40}}</ref><ref>{{cite journal |last1=Schiess|first1=R.|last2=Haeusler|first2=M. |title=No skeletal dysplasia in the Nariokotome boy KNM-WT 15000 (''Homo erectus'')--a reassessment of congenital pathologies of the vertebral column |journal=American Journal of Physical Anthropology |volume=150 |issue=3 |pages=365–374 |year=2013 |pmid=23283736 |doi=10.1002/ajpa.22211 |bibcode=2013AJPA..150..365S }}</ref> and a 1.5 million year old infant ''H. erectus'' skull from Mojokerto, Java, shows that this population did not have an extended childhood, which is a prerequisite for [[language acquisition]].<ref name=Coqueugniot2004/> On the other hand, despite the [[cochlea]]r (ear) anatomy of Sangiran 2 and 4 retaining several traits reminiscent of australopithecines, the hearing range may have included the higher frequencies used to discern speech.<ref>{{cite journal |first1=A. |last1=Urciuoli |first2=J. |last2=Kubat |first3=L. |last3=Schisanowski |first4=F. |last4=Schrenk |first5=B. |last5=Zipfel |first6=M. |last6=Tawane |first7=L. |last7=Bam |first8=D. M. |last8=Alba |first9=O. |last9=Kullmer |year=2022 |title=Cochlear morphology of Indonesian ''Homo erectus'' from Sangiran |journal=Journal of Human Evolution |volume=165 |article-number=103163 |doi=10.1016/j.jhevol.2022.103163|doi-access=free |pmid=35299091 |bibcode=2022JHumE.16503163U }}</ref>
Neurologically, all ''Homo'' have similarly configured brains, and, likewise, the [[Broca's area|Broca's]] and [[Wernicke's area|Wernicke's]] areas (in charge of sentence formulation and speech production in modern humans) of ''H. erectus'' were comparable to those of modern humans. However, this is not indicative of anything in terms of speech capability as even large chimpanzees can have similarly expanded Broca's area, and it is unclear if these areas served as language centers in archaic humans.<ref>{{cite book| vauthors = Luef EM |year=2018|chapter=Tracing the human brain's classical language areas in extant and extinct hominids|title=The talking species: Perspectives on the evolutionary, neuronal and cultural foundations of language|publisher=Uni-Press Graz|url=https://www.researchgate.net/publication/327285824|isbn=978-3-902666-52-9}}</ref> A 1-year-old ''H. erectus'' specimen shows that an extended childhood to allow for brain growth, which is a prerequisite in language acquisition, was not exhibited in this species.<ref name=Coqueugniot2004/>
 
The [[hyoid bone]] supports the tongue and makes possible modulation of the [[vocal tract]] to control pitch and volume. A 400 kya ''H. erectus'' hyoid bone from [[Castel di Guido]], Italy, is bar-shaped—more similar to that of other ''Homo'' than to that of non-human apes and ''Australopithecus''—but is devoid of muscle impressions, has a shield-shaped body, and is implied to have had reduced greater horns, meaning ''H. erectus'' lacked a humanlike vocal apparatus and thus anatomical prerequisites for a modern human level of speech.<ref>{{cite journal | vauthors = Capasso L, Michetti E, D'Anastasio R | title = A Homo erectus hyoid bone: possible implications for the origin of the human capability for speech | journal = Collegium Antropologicum | volume = 32 | issue = 4 | pages = 1007–1011 | date = December 2008 | pmid = 19149203 }}</ref> Increasing brain size and cultural complexity in tandem with technological refinement, and the hypothesis that articulate Neanderthals and modern humans may have inherited speech capabilities from the last common ancestor, could possibly indicate that ''H. erectus'' used some [[Origin of language|proto-language]] and built the basic framework which fully fledged languages would eventually be built around.<ref>{{cite journal | vauthors = Hillert DG | title = On the Evolving Biology of Language | journal = Frontiers in Psychology | volume = 6 | pages = 1796 | year = 2015 | pmid = 26635694 | pmc = 4656830 | doi = 10.3389/fpsyg.2015.01796 | doi-access = free }}</ref> However, this ancestor may have instead been ''H. heidelbergensis'', as a hyoid bone of a 530 kya ''H. heidelbergensis'' specimen from the Spanish [[Sima de los Huesos]] Cave is like that of modern humans,<ref>{{cite journal | vauthors = Martínez I, Arsuaga JL, Quam R, Carretero JM, Gracia A, Rodríguez L | title = Human hyoid bones from the middle Pleistocene site of the Sima de los Huesos (Sierra de Atapuerca, Spain) | journal = Journal of Human Evolution | volume = 54 | issue = 1 | pages = 118–124 | date = January 2008 | pmid = 17804038 | doi = 10.1016/j.jhevol.2007.07.006 | url = https://eprints.ucm.es/26853/1/1-s2.0-S0047_1.pdf }}</ref> and another specimen from the same area shows an auditory capacity sensitive enough to pick up human speech.<ref>{{cite journal | vauthors = Martínez I, Rosa M, Arsuaga JL, Jarabo P, Quam R, Lorenzo C, Gracia A, Carretero JM, Bermúdez de Castro JM, Carbonell E | display-authors = 6 | title = Auditory capacities in Middle Pleistocene humans from the Sierra de Atapuerca in Spain | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 101 | issue = 27 | pages = 9976–9981 | date = July 2004 | pmid = 15213327 | pmc = 454200 | doi = 10.1073/pnas.0403595101 | doi-access = free | bibcode = 2004PNAS..101.9976M }}</ref>
 
==Extinction==
{{Expand section|date=December 2020}}
The last known occurrence of ''Homo erectus'' is 117,000–108,000 years ago in [[Ngandong]], [[Java (island)|Java]] according to a study published in 2019.<ref name=Rizal/>
 
In 2020 researchers reported that ''Homo erectus'' and ''[[Homo heidelbergensis]]'' lost more than half of their climate [[Ecological niche|niche]] – climate they were adapted to – with no corresponding reduction in physical range, just before [[extinction]] and that climate change played a substantial role in extinctions of past ''Homo'' species.<ref>{{cite news | vauthors = Padmanaban D |title=Climate Change May Have Been a Major Driver of Ancient Hominin Extinctions |url=https://www.sapiens.org/archaeology/hominin-extinctions/ |website=SAPIENS |access-date=9 November 2020 |date=6 November 2020}}</ref><ref>{{cite news |title=Climate change likely drove early human species to extinction, modeling study suggests |url=https://phys.org/news/2020-10-climate-drove-early-human-species.html |access-date=9 November 2020 |work=phys.org |language=en}}</ref><ref>{{cite journal | vauthors = Raia P, Mondanaro A, Melchionna M, Di Febbraro M, Diniz-Filho JA, Rangel TF, Holden PB, Carotenuto F, Edwards NR, Lima-Ribeiro MS, Profico A, Maiorano L, Castiglione S, Serio C, Rook L | display-authors = 6 |title=Past Extinctions of Homo Species Coincided with Increased Vulnerability to Climatic Change |journal=One Earth |date=23 October 2020 |volume=3 |issue=4 |pages=480–490 |doi=10.1016/j.oneear.2020.09.007 | bibcode = 2020OEart...3..480R |hdl=2158/1211341 |language=en |issn=2590-3330|doi-access=free }}</ref>
 
==Fossils==
[[File:Homo erectus KNM ER 3733.jpg|thumb|Homo erectus KNM ER 3733 actual skull]]
The lower cave of China's Zhoukoudian Cave is one of the most important archaeological sites worldwide.<ref name=":2">Zanolli, Clément, et al. "Inner Tooth Morphology of Homo Erectus from Zhoukoudian. New Evidence from an Old Collection Housed at Uppsala University, Sweden." ''Journal of Human Evolution'', vol. 116, Mar. 2018, pp. 1–13.</ref> There have been remains of 45 ''Homo erectus'' individuals found and thousands of tools recovered.<ref name=":2" /> Most of these remains were lost during World War 2, with the exception of two postcranial elements that were rediscovered in China in 1951 and four human teeth from 'Dragon Bone Hill'.<ref name=":2" />
 
New evidence has shown that ''Homo erectus'' does not have uniquely thick vault bones, as was previously thought.<ref name=":3">Copes, Lynn E., and William H. Kimbel. "Cranial Vault Thickness in Primates: Homo Erectus Does Not Have Uniquely Thick Vault Bones." ''Journal of Human Evolution'', vol. 90, Jan. 2016, pp. 120–134.</ref> Testing showed that neither Asian nor African ''Homo erectus'' had uniquely large vault bones.<ref name=":3" />
 
=== Individual fossils ===
Some of the major ''Homo erectus'' fossils:
* Indonesia (island of Java): [[Trinil 2]] ([[holotype]]), [[Sangiran]] collection, Sambungmachan collection,<ref>{{cite journal | vauthors = Delson E, Harvati K, Reddy D, Marcus LF, Mowbray K, Sawyer GJ, Jacob T, Márquez S | display-authors = 6 | title = The Sambungmacan 3 Homo erectus calvaria: a comparative morphometric and morphological analysis | journal = The Anatomical Record | volume = 262 | issue = 4 | pages = 380–397 | date = April 2001 | pmid = 11275970 | doi = 10.1002/ar.1048 | s2cid = 25438682 | doi-access = free }}</ref> [[Solo Man|Ngandong collection]]
* China ("[[Peking Man]]"): Lantian (Gongwangling and Chenjiawo), Yunxian, [[Zhoukoudian]], Nanjing, [[Hexian]]
* Kenya: [[KNM ER 3883]], [[KNM ER 3733]]
* Vietnam: Northern, [[Tham Khuyen]],<ref>{{cite journal | vauthors = Ciochon R, Long VT, Larick R, González L, Grün R, de Vos J, Yonge C, Taylor L, Yoshida H, Reagan M | display-authors = 6 | title = Dated co-occurrence of Homo erectus and Gigantopithecus from Tham Khuyen Cave, Vietnam | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 93 | issue = 7 | pages = 3016–3020 | date = April 1996 | pmid = 8610161 | pmc = 39753 | doi = 10.1073/pnas.93.7.3016 | doi-access = free | bibcode = 1996PNAS...93.3016C }}</ref> Hoa Binh{{citation needed|date=October 2013}}
* Republic of Georgia: Dmanisi collection ("''[[Homo erectus georgicus]]''")
* Ethiopia: [[Daka skull|Daka calvaria]]
* Eritrea: Buia cranium (possibly ''H. ergaster'')<ref>{{cite web|url= http://archive.archaeology.org/9809/newsbriefs/eritrea.html| title= New Skull from Eritrea| volume= 51 |number= 5| date= September–October 1998 | vauthors = Schuster AM | work= Archaeology| access-date= 3 October 2015}}</ref>
* [[Denizli Province]], Turkey: Kocabas fossil<ref name="First Homo erectus from Turkey and"/>
* [[Drimolen]], South Africa: DNH 134<ref>{{cite journal | vauthors = Herries AI, Martin JM, Leece AB, Adams JW, Boschian G, Joannes-Boyau R, Edwards TR, Mallett T, Massey J, Murszewski A, Neubauer S, Pickering R, Strait DS, Armstrong BJ, Baker S, Caruana MV, Denham T, Hellstrom J, Moggi-Cecchi J, Mokobane S, Penzo-Kajewski P, Rovinsky DS, Schwartz GT, Stammers RC, Wilson C, Woodhead J, Menter C | display-authors = 6 | title = Contemporaneity of ''Australopithecus'', ''Paranthropus'', and early ''Homo erectus'' in South Africa | journal = Science | volume = 368 | issue = 6486 | pages = eaaw7293 | date = April 2020 | pmid = 32241925 | doi = 10.1126/science.aaw7293 | hdl-access = free | s2cid = 214763272 | hdl = 11568/1040368 }}</ref>
 
== Phylogeny ==
A cladogram of ''Homo erectus'' is as follows.<ref>{{Cite journal |last1=Ni |first1=Xijun |last2=Ji |first2=Qiang |last3=Wu |first3=Wensheng |last4=Shao |first4=Qingfeng |last5=Ji |first5=Yannan |last6=Zhang |first6=Chi |last7=Liang |first7=Lei |last8=Ge |first8=Junyi |last9=Guo |first9=Zhen |last10=Li |first10=Jinhua |last11=Li |first11=Qiang |date=2021-08-28 |title=Massive cranium from Harbin in northeastern China establishes a new Middle Pleistocene human lineage |journal=The Innovation |language=English |volume=2 |issue=3 |doi=10.1016/j.xinn.2021.100130 |issn=2666-6758 |pmc=8454562 |pmid=34557770 |last13=Stringer |first13=Chris |last12=Grün |first12=Rainer|page=100130 |bibcode=2021Innov...200130N }}</ref> It is indicated how many million years ago the clades diverged.
 
{{Clade|{{Clade
  |1=''[[Homo habilis]]'' (†1.7 Mya)
  |label2='''''Homo erectus'' s.l. (2.3)'''
  |2={{clade
    |label1=(2.1)
    |1={{clade
      |1=''[[H. gautengensis]]'' (†1.9)
    |2=[[Dmanisi hominins|Dmanisi]] (†1.8)
    }}
  |label2='''(2.0)'''
  |2={{clade
    |1=[[Turkana Boy|Turkana]] (†1.7)
    |label2='''(1.8)'''
    |2={{clade
      |1=[[Olduvai Hominid 9|Olduvai Hominids]] (†1.5)
      |label2='''Asian ''H. e.'' (1.6)'''
      |2={{clade
        |1=[[Sangiran]] (†1.4)
        |label2='''(1.4)'''
        |2={{clade
          |label1=(1.1)
        |1={{clade
          |label1=(0.8)
          |1={{clade
          |1=[[Nanjing Man]] (†0.6)
          |2=[[Peking Man]] (†0.5)
          }}
        |label2=(0.9)
          |2={{clade
            |1=[[Hexian]] (†0.5)
            |label2=(0.6)
            |2={{clade
              |1=[[Solo Man|Sambungmacan]] (†0.2)
              |2=[[Ngandong]] (†0.1)
            }}
            }}
        }}
        |2='''''[[Homo heidelbergensis|Homo rhodesiensis/heidelbergensis]]''''' (incl. ''[[Homo sapiens]]''))
}}
        }}
      }}
  }}
  }}
}}|label1='''[[Homo]] (2.85)'''|style=font-size:75%;line-height:75%}}
 
''Homo erectus'' was originally African. The extant ''Homo heidelbergensis'' ([[Cladistics|cladistically]] granting ''[[Homo sapiens]]''), which was originally African, emerged within the Asian ''Homo erectus''. Contemporary groups appear to have been interbreeding, so any phylogeny like this only gives a coarse impression of the evolution of ''Homo'', and extinct lineage may have partially continued in other groupings. Not included are other contemporary groups such as ''[[Homo floresiensis]], [[Homo naledi]]'', ''[[Homo luzonensis]]'', ''[[Homo rudolfensis]]'', ''[[Australopithecus sediba]], [[Australopithecus africanus]], and [[Paranthropus]].''


==Gallery==
Given expanding brain size and technological innovation, ''H. erectus'' may have been using some basic proto-language in combination with gesturing, and built the basic framework around which fully-fledged languages would eventually be formed.<ref>{{cite journal |last=Hillert|first=D. G. |title=On the Evolving Biology of Language |journal=Frontiers in Psychology |volume=6 |pages=1796 |year=2015 |pmid=26635694 |pmc=4656830 |doi=10.3389/fpsyg.2015.01796 |doi-access=free}}</ref>
<gallery mode="packed-hover" heights="120" class="center">
File:Homo erectus tautavelensis.jpg|''[[Homo erectus tautavelensis]]'' skull.
File:Tautavel UK 2.JPG|Replica of lower jaws of ''Homo erectus'' from [[Tautavel]], [[France]].
File:Calvaria Sangiran II (A).jpg|[[Calvaria (skull)|Calvaria]] "[[Sangiran]] II" original, collection [[Gustav Heinrich Ralph von Koenigswald|Koenigswald]], [[Senckenberg Museum]].
File:Homo erectus hand axe Daka Ethiopia.jpg|A reconstruction based on evidence from the [[Daka skull|Daka]] Member, Ethiopia
File:Pithecanthropus-erectus.jpg|Original fossils of ''Pithecanthropus erectus'' (now ''Homo erectus'') found in [[Java]] in 1891.
</gallery>


== See also ==
==See also==
* ''[[Anthropopithecus]]''
*[[Cro-Magnon]]
* [[Kozarnika, Dimovo Municipality]]
*[[Behavioral modernity]]
* [[Multiregional origin of modern humans]]
*[[Evolution of human intelligence]]
*''[[Homo naledi]]''


'''General:'''
==Notes==
* [[List of fossil sites]] ''(with link directory)''
{{notelist-num}}
* [[List of human evolution fossils]] ''(with images)''


== References ==
==References==
{{Reflist}}
{{reflist}}


== Further reading ==
===Bibliography===
* {{cite magazine|title=Homo Erectus Unearthed| vauthors = Leakey R, Walker A |magazine=[[National Geographic (magazine)|National Geographic]]|pages=624–629|volume=168|issue=5|date=November 1985|issn=0027-9358|oclc=643483454}}
* {{cite book |last1=Boaz |first1=N. T. |last2=Ciochon |first2=R. |author2-link=Russell Ciochon |year=2004 |title=Dragon Bone Hill: An Ice-Age Saga of Homo erectus |publisher=Oxford University Press |isbn=978-0-19-803488-9}}
* {{cite book| vauthors = Sigmon BA, Cybulski JS |year=1981|title=Homo erectus: Papers in Honor of Davidson Black|publisher=University of Toronto Press|jstor=10.3138/j.ctvcj2jdw.11|ref={{harvid|Sigmon|1981}}}}
* {{cite journal |first=S. C. |last=Antón |year=2003 |title=Natural history of ''Homo erectus''<sup>†</sup> |journal=American Journal of Biological Anthropology |volume=122 |issue=S37 |pages=126–170 |doi=10.1002/ajpa.10399 |doi-access=free|pmid=14666536 |bibcode=2003AJPA..122S.126A }}
* {{cite book| vauthors = Theunissen B, Theunissen LT |year=2012|title=Eugène Dubois and the Ape-Man from Java|publisher=Springer Netherlands|isbn=9789400922099|ref={{harvid|Theunissen|2012}}}}
* {{cite book |last=Theunissen |first=B. |year=1989 |title=Eugène Dubois and the Ape-Man from Java |publisher=Kluwer Academic Publishers |isbn=978-1-55608-081-4}}


== External links ==
== External links ==
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[[Category:Early species of Homo]]
[[Category:Early species of Homo]]
[[Category:Fossil taxa described in 1893]]
[[Category:Fossil taxa described in 1893]]
[[Category:Extinct animals of Indonesia]]
[[Category:Apex predators]]

Latest revision as of 21:59, 15 March 2026


Homo erectus
Java Man, the holotype of H. erectusTemplate:Efn-num
Scientific classification edit
: [[Template:Taxonomy/Homo]]
Species:
H. erectus
Binomial name
Homo erectus
(Dubois, 1893)

Homo erectus (/ˌhm əˈrɛktəs/ lit. 'upright man') is an extinct species of archaic human from the Pleistocene, spanning nearly 2 million years. It is the first human species to evolve a humanlike body plan and gait, to leave Africa and colonize Asia and Europe, and to wield fire. Some populations of H. erectus were ancestors of later human species, including H. heidelbergensis — the last common ancestor of modern humans, Neanderthals, and Denisovans. As such a widely distributed species both geographically and temporally, H. erectus anatomy varies considerably. Subspecies are sometimes recognized: H. e. erectus, H. e. pekinensis, H. e. soloensis, H. e. ergaster, H. e. georgicus, and H. e. tautavelensis.

The species was first described by Eugène Dubois in 1893 as "Pithecanthropus erectus" using a skullcap, molar, and femur from Java, Indonesia. Further discoveries around East Asia were used to contend that humanity evolved out of Asia. Based on historical race concepts, it was argued that local H. erectus populations evolved directly into local modern human populations (polycentrism) rather than all humanity sharing a single anatomically modern ancestor (monogenism). As the fossil record improved over the mid-to-late 20th century, "Out of Africa" theory and monogenism became the consensus.

The typical skull has a pronounced brow ridge, a protruding jaw, and large teeth. The bones are much thicker than in modern humans. East Asian populations normally have an even more robust skeleton and larger brain volume — averaging 1,000 cc (61 cu in). Western H. erectus brain volume could be as low as 546 cc (33.3 cu in) in H. e. georgicus. H. erectus probably had a faster apelike growth trajectory, lacking the extended childhood required for language acquisition. Reconstructed adult body dimensions range from 141–167 cm (4 ft 8 in – 5 ft 6 in) in height and about 50 kg (110 lb) in weight.

H. erectus invented the Acheulean tool industry, a major innovation of large, heavy-duty stone tools. These may have been used in butchery, vegetable processing, and woodworking of spears and digging sticks. H. erectus was a major predator of large herbivores on the expanding savannas during the Quaternary glaciation. The species is usually characterized as the first hunter-gatherer and the first to practice sexual division of labor. Fire usage and cave habitation were probably not important aspects of daily life. Similarly, H. erectus may not have often ventured into colder regions or cooked meat. The last known occurrence of H. erectus is 108,000 to 117,000 years ago (H. e. soloensis) in Southeast Asia, until the last savannas in the region gave way to jungle.

Taxonomy[edit | edit source]

Research history[edit | edit source]

In 1868, Ernst Haeckel suggested early humans dispersed from the now-disproven hypothetical continent "Lemuria".[2][3]

While Charles Darwin had hypothesized in his 1871 Descent of Man that humans most likely evolved in Africa,Template:Efn-num many late-19th century evolutionary naturalists postulated that Asia was the birthplace of humankind. Asia is midway between all continents via land routes or short sea crossings, providing optimal dispersal routes throughout the world. Among the major proponents of "Out of Asia" theory was Ernst Haeckel, who argued that the first human species (which he speculatively named Homo primigenius) evolved on a now-disproven hypothetical continent "Lemuria" from a species he termed Pithecanthropus alalus (speechless ape-man). "Lemuria" had supposedly sunk below the Indian Ocean, accounting for the lack of fossil evidence.[2]

Dutch scientist Eugène Dubois joined the Royal Netherlands East Indies Army to search for the "missing link" of human evolution in Java.Template:Efn-num At the Trinil site, his team found a skullcap and molar in 1891, and a femur in 1892 (Java Man), which he named "Pithecanthropus erectus" in 1893.[4] He attempted to convince the European scientific community that he had found an upright-walking ape-man dating to the late Pliocene or Early Pleistocene; they dismissed his findings as some kind of non-human ape.[2]

Franz Weidenreich and Ralph von Koenigswald argued Java Man was an ancient human using fossils like Sangiran (left) and Peking Man (right)

Dubois argued that "P. erectus" was a gibbon-like ape which was the precursor to a more familiar human body plan, but in the 1930s, German-American anatomist Franz Weidenreich noticed a striking similarity with ancient human remains recently being unearthed in China (Peking Man, "Sinanthropus pekinensis").[5][6] This characterization became better supported as German-Dutch palaeontologist Gustav Heinrich Ralph von Koenigswald discovered more Indonesian ancient human remains over the decade at Mojokerto, Sangiran, and Ngandong.Template:Efn-num[7][8] Weidenreich believed that they were the direct ancestors of the local modern human Homo sapiens subspecies, in accord with historical race concepts (polycentricism) — that is, Peking Man was the direct ancestor of specifically Chinese people, and Java Man of Aboriginal Australians.[9][10] As the significance of racial distinction diminished with the development of modern evolutionary synthesis, many fossil human species and genera around Asia, Africa, and Europe (including "Pithecanthropus" and "Sinanthropus") were reclassified as subspecies of Homo erectus.[11][12]

During the late 20th century, some of the oldest H. erectus fossils were discovered across Africa, the first being Kenyan archeologist Louis Leakey's Olduvai Hominin 9 in 1960.[13] As the human fossil record expanded, the "Out of Africa" theory and monogenism became the consensus: that all modern humans share a fully anatomically modern common ancestor. H. erectus is now generally considered to be an African species which later dispersed across Eurasia, with later African populations giving rise to the modern human lineage.[14]

Subspecies[edit | edit source]

By the middle of the 20th century, human taxonomy was in turmoil, with many poorly defined species and genera described across Europe, Asia, and Africa, which exaggerated the differences among them.[15] In 1940, Weidenreich was the first to suggest reclassifying "Sinanthropus pekinensis" and "Pithecanthropus erectus" as subspecies of H. erectus.[9] In 1950, German-American evolutionary biologist Ernst Mayr entered this field. Surveying a "bewildering diversity of names" and many proposals for consolidation, he decided to reclassify human fossils into three species of Homo: "H. transvaalensis" (the australopithecines), H. erectus (including "Sinanthropus", "Pithecanthropus", and various other Asian, African, and European taxa), and H. sapiens (including anything younger than H. erectus, such as modern humans and Neanderthals). Mayr defined these species as a sequential lineage, each evolving into the next (chronospecies).[10] Though later Mayr changed his opinion on the australopithecines (recognizing Australopithecus), his more conservative view of archaic human diversity became widely adopted in the subsequent decades.[11]

...never more than one species of man existed on the earth at any one time... If fossils of Congo pygmies and of Watusi were to be found in the same deposit by a paleontologist, a million years hence, he might well think that they belonged to two different species.

— Ernst Mayr, 1950[10]

In the 1970s, as population genetics was being formulated, the anatomical variation of H. erectus across its wide geographic and temporal range (the basis for the subspecies distinctions) became better understood as clines — different populations which attained some anatomical regionality but were not reproductively isolated.[14] In general, subspecies names for H. erectus are now used for convenience to indicate time and region rather than specific anatomical trends.[16]

...to paleontologists in general, subspecies are epiphenomena which do not merit the attention paid to species... The pursuit of subspecies in the fossil record is at best fraught with difficulty, and is more probably futile.

— Ian Tattersall, 1986[17]
Reconstructions of H. e. ergaster (KNM ER 3733) left and H. e. pekinensis right

The more commonly used subspecies (if any are used) are:[18]

The ancient Georgia fossils have variably been classified as H. e. ergaster (or quadrinomial H. e. ergaster georgicus),[21] as their own subspecies as H. e. georgicus, or as their own species H. georgicus.[19] Some authors may also elevate H. ergaster,[22] H. soloensis,[23] and H. pekinensis to species level.[24] Fossils relegated to H. e. tautavelensis are traditionally assigned to H. heidelbergensis.[20]

Evolution and dispersal[edit | edit source]

H. e. georgicus (above) represents one of the earliest dispersals out of Africa about 1.8 million years ago.[25]

H. erectus evolved in Africa from a population of H. habilis[26][27] and they coexisted for about half a million years.[28] During this time interval, H. erectus populations could display a mix of more "classically erectus" or "classically habilis" cranial anatomy and a wide range of brain volumes.[29] The oldest identified H. erectus specimen is a 2.04 million year old skull, DNH 134, from Drimolen, South Africa, coexisting with the australopithecine Paranthropus robustus.[30] H. erectus dispersed out of Africa soon after evolution, the earliest recorded instances being H. e. georgicus 1.78 to 1.85 million years ago in Georgia,[25] the Indonesian Mojokerto and Sangiran sites 1.6 to 1.8 million years ago,[31][32] and the Chinese Yunxian Man 1.77 million years ago.[33] Populations may have pushed into northwestern Europe at around the same time.[34] While H. erectus is usually considered the first hominin to leave Africa, stone tools that may date to as far back as 2.48 and 2.1 million years ago (from Zarqa Valley, Jordan, and Shangchen, China, respectively) could indicate that an earlier hominin species left Africa.[35] Since H. erectus was first defined in East Asia, those populations are sometimes distinguished as H. erectus sensu stricto ("in the strict sense"), and African and West Eurasian populations as H. erectus sensu lato ("in the broad sense"), but this may not reflect how these populations are actually related to each other.Template:Efn-num[16][27]

Once established around the Old World, H. erectus evolved into other later species in the genus Homo, including: H. heidelbergensis, H. antecessor,[36] H. floresiensis,[37] and H. luzonensis.[38] H. heidelbergensis, in turn, is usually placed as the last common ancestor of Neanderthals (H. neanderthalensis), Denisovans, and modern humans.[36] H. erectus is thus a non-natural, paraphyletic grouping of fossils and does not include all the descendants of a last common ancestor.[39] Despite being designated as a different species, H. erectus may have interbred with some of its descendant species, namely the common ancestor of Neanderthals and Denisovans ("Neandersovans").[40]

Successive dispersals of   Homo erectus (yellow),   Homo neanderthalensis (ochre) and   Homo sapiens (red, Out of Africa II)

The dispersal of H. erectus is generally ascribed to the evolution of obligate bipedalism, better technology, and adoption of a carnivorous diet.[41] However, the sudden adoption of carnivory could be sampling bias, with earlier species consuming the same amount of meat.[42] Populations spread out via open grassland and woodland savannas, which were expanding due to a global aridification trend at the onset of the Quaternary glaciation.[41] H. erectus is usually thought to have occupied the Sahara and West Asia during humid periods, but populations may have persisted into desert periods.[43]

Most H. erectus sensu lato specimens date to 1 to 1.8 million years ago in the Early Pleistocene before giving way to descendant species.[1] The classification of Middle Pleistocene Homo has been a controversial topic, termed "the muddle in the middle".[20][44] H. erectus sensu stricto persisted much longer than sensu lato, with the youngest population (H. e. soloensis) dating to 108,000 to 117,000 years ago in Late Pleistocene Java.[1] This population appears to have died out when the savannah corridors closed and tropical jungle took over.[45]

A 2021 phylogeny of some H. erectus fossils using tip dating:[39]

Template:Clade

Biology[edit | edit source]

As such a widely distributed species both across regions and through time, the anatomy of H erectus can vary considerably. Among living primates, the degree of regionality[clarification needed] achieved by H. erectus (phenotypic plasticity) is only observed in modern humans.[46]

Head[edit | edit source]

Franz Weidenreich's reconstruction of the H. e. soloensis skull

Dubois originally described the species using a skullcap, noting the traits of a low and thickened cranial vault and a continuous bar of bone forming the brow ridge (supraorbital torus).[47] H. erectus fossils typically share these traits, but the Kenyan Koobi Fora skulls notably have thinner skulls and weaker supraorbital tori.[48] He also used several other traits now considered more typical of H. erectus sensu stricto, such as a sagittal keel running across the midline of the skullcap, a bar of bone across the back of the skull (occipital torus), and a strong crest on the mastoid part of the temporal bone.[47] These traits can be still be found, nonetheless, in a few H. erectus sensu lato specimens, namely the 1.47 million year old Olduvai Hominin 9.[49]

Compared to H. erectus sensu lato, the skullcap of sensu stricto narrows considerably at the front, the face is bigger and presumably more prognathic (it juts out more, but the face is poorly documented), and the molars are larger particularly in Indonesian fossils.[50] H. erectus was the first human species with a fleshy nose, which is generally thought to have evolved in response to breathing dry air in order to retain moisture.[51] Compared to earlier Homo, H. erectus has smaller teeth, thinner enamel, and weaker mandibles (jawbone), likely due to a greater reliance on tool use and food processing.[52]

The brain size of H. erectus varies considerably, but is generally smaller in H. erectus sensu lato, as low as 546 cc (33.3 cu in) in Dmanisi skull 5.[53] East Asian H. erectus overall are rather big-brained, averaging roughly 1,000 cc,[46] staying within the range of variation for modern humans.[54] The late-surviving H. e. soloensis has the biggest brain volume with one specimen measuring 1,251 cc (76.3 cu in).[49]

Body[edit | edit source]

Turkana Boy at the Neanderthal Museum

The rest of the body is primarily understood by three partial skeletons from the Kenyan Lake Turkana site, notably Turkana Boy. Other postcranial fossils (all bones aside from the skull) attributed to H. erectus are not associated with a skull, making attribution unverifiable. Though the body plan of earlier Homo is poorly understood, H. erectus is usually characterized as the first Homo species with a human body plan, distinct from non-human apes.[55][46][56] The chest may have been short and barrel-shaped, like other archaic humans.[57] Fossil tracks near Ileret, Kenya, suggest a human gait. This adaptation is implicated in the dispersal of H. erectus across the Old World.[58]

It is unclear when human ancestors lost most of their body hair. Genetic analysis suggests that high activity in the melanocortin 1 receptor, which produces dark skin, dates back to 1.2 million years ago. This could indicate the evolution of hairlessness around this time, as a lack of body hair would have left the skin exposed to harmful UV radiation.[59] It is possible that populations in higher latitudes developed lighter skin to prevent vitamin D deficiency,[60] though a 300,000 to 500,000 year old Turkish H. erectus specimen presents the earliest case of tuberculous meningitis, which is typically exacerbated by vitamin D deficiency in dark-skinned people living in higher latitudes.[61] Hairlessness is generally thought to have facilitated sweating,[62] but it may also have helped to reduce parasite load, and was possibly reinforced by sexual selection.[63][64]

Size[edit | edit source]

Height reconstructions range approximately 141–167 cm (4 ft 8 in – 5 ft 6 in), with tropical populations typically reconstructed as scoring on the higher end like modern human populations. Adult weight is harder to approximate, but about 50 kg (110 lb) may have been normal. H. erectus is usually thought to be the first human species with little size-specific sexual dimorphism, but the variability of postcranial material makes this unclear.[46] A 2010 study estimates that the Turkana Boy would have reached a height of 163 cm (5 ft 4 in) if he had reached adulthood.[65]

Growth and development[edit | edit source]

The dimensions of a 1.8 million years old adult female H. e. ergaster pelvis from Gona, Ethiopia, suggests that she would have been capable of birthing children with a maximum prenatal brain size of 315 cc (19.2 cu in), about 30–50% of adult brain size, falling between chimpanzees (~40%) and modern humans (28%).[66] Similarly, a 1.5 million year old infant skull from Mojokerto had a brain volume of about 72–84% the size of an adult, which suggests a brain growth trajectory more similar to that of non-human apes.[67] This suggests that the childhood growth and development of H. erectus was intermediate between that of chimpanzees and modern humans,[66] and the faster development rate suggests that altriciality (an extended childhood) evolved at a later stage in human evolution.[67] The faster development rate might also indicate a shorter expected lifespan compared to later Homo.[68]

Bone thickness[edit | edit source]

Cross sections of Peking Man Skulls III (A) and XII (B), and Java Man Skull II (C)

The bones are extraordinarily thickened, particularly in Homo erectus sensu stricto, so much so that skull fragments have sometimes been confused for fossil turtle carapaces.[69] The medullary canal in the long bones (where the bone marrow is stored, in the limbs) is extremely narrowed (medullary stenosis). This degree of thickening is usually exhibited in semi-aquatic animals which use their heavy (pachyosteosclerotic) bones as ballasts to help them sink, induced by hypothyroidism.[70]

It is unclear what function intense bone thickening could have served. Before more complete skeletons were discovered, Weidenreich suggested H. erectus was a gigantic species.[71] Other explanations include a far more violent and impact-prone lifestyle than other Homo, or pathological nutrient deficiencies.[72] The supraorbital torus thickens with age, and may be a response to bending stresses from habitual loading of the front teeth.[73]

Culture[edit | edit source]

Subsistence[edit | edit source]

H. erectus overhunting may have led to the extinction of Megalochelys (above).[74]

H. erectus was early-on portrayed as the earliest hunter-gatherer and a skilled predator of big game, relying on running. The few identified specimens of the H. e. ergaster torso and pelvis may indicate a body plan more conducive for power running, unlike modern humans better adapted for endurance running.[57] The gradual shift to "top predator" may have led to its dispersal throughout Afro-Eurasia.[41] Though scavenging may have instead played a bigger role at least in some populations, H. erectus fossils are often associated with the butchered remains of large herbivores,[75] especially elephants, rhinos, hippos, bovines, and boars. Tracking complex prey behaviors as well as the nutritional value of meat have been connected to brain volume growth.[76]

H. erectus is usually assumed to have practiced sexual division of labor much like recent hunter-gatherer societies, with men hunting and women gathering. This model is supported by a fossil trackway from Ileret, Kenya, made by a probably all-male band of over 20 H. erectus individuals, possibly a hunting party or (similar to chimpanzees) a border patrol group.[77]

Since common modern human tapeworms began to diverge from those of other predators roughly 1.7 million years ago (specifically the pork tapeworm, beef tapeworm, and Asian tapeworm), not only was H. erectus consuming meat regularly enough for speciation to occur in these parasites, but meat was probably consumed raw more often than not.[78] Some populations were collecting aquatic resources like fish, shellfish, and turtles at waterside sites, such as Lake Turkana[79] and Trinil.[80] Underground storage organs (roots, tubers, etc.) were likely also major dietary components, and traces of the edible plant Celtis have been documented at several H. erectus sites.[81]

Possibly due to overhunting of the biggest game available, the dispersal of H. erectus and descendant species may be implicated in the extinctions of large herbivores and the gradual reduction of average herbivore size over the Pleistocene.[82] H. erectus overhunting has been blamed by some authors for the decline of proboscidean species as well as competing carnivores,[76][83][84] but their decline may be better attributed to the spread of grasslands.[84][85] The giant tortoise Megalochelys may have been driven to extinction by H. erectus in Sundaland (what is now Island Southeast Asia), since species went extinct shortly after the arrival of H. erectus.[74]

Technology[edit | edit source]

Stone tools[edit | edit source]

A handaxe from the Saint-Acheul site at the Musée d'Archéologie nationale, France

H. erectus manufactured Lower Paleolithic technologies, and is credited with the invention of the Acheulean stone tool industry at latest 1.95 million years ago.[86] This was a major technological breakthrough featuring large, heavy-duty tools; most iconically, the handaxe. Over hundreds of thousands of years, the Acheulean eventually replaced its predecessor — the Oldowan (a chopper and flake industry) — in Africa, and spread out across Western Eurasia.[87] This sudden innovation was typically explained as a response to environmental instability in order to process more types of food and broaden the diet combined with increasing brain size, which also allowed H. erectus to colonize Eurasia. Despite this characterization of the Acheulean, the small-brained H. e. georgicus was able to leave Africa despite only manufacturing Oldowan-style tools,[41][87] the 1.6 million year old DAN5/P1 specimen from Gona, Ethiopia is associated with Acheulean style tools despite its low brain volume of 598 cc (36.5 cu in),[29] and the handaxe does not seem to have been manufactured commonly in East Asia.[88] The lack of East Asian handaxes was first noted by American archaeologist Hallam L. Movius in 1948, who drew the "Movius Line", dividing the East into a "chopping-tool culture" and the West into a "hand axe culture".[89] Movius took this as evidence of inferiority of Far Eastern populations:

...as early as Lower Palaeolithic times Southern and Eastern Asia as a whole was a region of cultural retardation...very primitive forms of Early Man apparently persisted there long after types at a comparable stage of physical evolution had become extinct elsewhere.

Reconstruction of Turkana Boy at the Neanderthal Museum

H. erectus seems to have been using stone tools in butchery, vegetable processing, and woodworking (maybe manufacturing spears and digging sticks).[81][90] In Africa, Oldowan sites are typically found alongside major fossil assemblages, but Acheulean sites normally feature more stone tools than fossils, so H. erectus could have been using choppers and handaxes for different activities.[90] These stone tools probably were not hafted onto spears; this innovation is associated with the transition to the Middle Paleolithic and the emergence of Neanderthals and modern humans.[91]

Materials for stone tools were normally sourced locally, and it seems blanks were usually chosen based on size rather than material quality.[87] H. erectus also produced tools from shells at Sangiran[92] and Trinil.[93]

Fire[edit | edit source]

H. erectus is credited as the first human species to wield fire. The earliest claimed fire site is Wonderwerk Cave, South Africa, at 1.7 million years old.[94] While the species' dispersal far out of Africa has often been attributed to fire and cave dwelling, fire does not become common in the archaeological record until 300,000 to 400,000 years ago,[95] and cave-dwelling about 600,000 years ago.[96] Therefore, H. erectus may have only been scavenging fire opportunistically. Similarly, H. erectus sites usually stay within warmer tropical or subtropical latitudes.[41]

The dating of northerly populations (namely Peking Man) could suggest that they were retreating to warmer refugia during glacial periods, but the precise age of the Peking Man fossils is poorly resolved.[47][97] There have been claims of manmade hearths and "clear-cut evidence for intentional fire use",[98] ostensibly as far back as 770,000 years ago in the supposed cave home of Peking Man.[97] At the French Caune de L'Arago, Tautavel Man does not seem to have been using fire at all, even though occupation sequences span two cold periods.[20]

Healthcare[edit | edit source]

The single-toothed H. e. georgicus specimen (above) is the earliest probable example of human group care.[99]

Like other primates, H. erectus probably used medicinal plants[81] and infirmed sick group members. The earliest probable example of human group care is a 1.77 million year old H. e. georgicus specimen who had lost all but one tooth due to age or gum disease (the earliest example of severe chewing impairment) yet still survived for several years afterwards.[99]

Seafaring[edit | edit source]

H. erectus made long sea crossings to arrive on the islands of Flores, Luzon,[100] and some Mediterranean islands. Some authors have asserted that H. erectus intentionally made these crossings by inventing watercrafts and seafaring so early in time, speaking to advanced cognition and language skills. These populations could have also been founded by natural rafting events instead.[101]

Art and rituals[edit | edit source]

In East Asia, H. erectus is usually represented only by skullcaps, which used to be interpreted as widespread cannibalism and ritual headhunting. This had been reinforced by the historic practice of headhunting and cannibalism in some recent Indonesian, Australian, and Polynesian cultures, which were formerly believed to have directly descended from these H. erectus populations. The lack of the rest of the skeleton is now normally explained by natural phenomena.[102]

Engraved Pseudodon shell DUB1006-fL from Trinil, Java

Art-making could be evidence of symbolic thinking. An engraved Pseudodon shell DUB1006-fL from Trinil, Java, with geometric markings could possibly be the earliest example of art-making, dating to 436,000 to 546,000 years ago.[93][103][104] H. erectus was also the earliest human to collect red-colored pigments, namely ochre. Ochre lumps at Olduvai Gorge, Tanzania, associated with the 1.4 million year old Olduvai Hominid 9 may have been purposefully shaped and trimmed by a hammerstone. Red ochre is normally recognized as bearing symbolic value when associated with modern humans.[104]

Language[edit | edit source]

The spinal column of the 1.6 million year old Turkana boy would not have supported properly developed respiratory muscles required to produce speech;[105][106] and a 1.5 million year old infant H. erectus skull from Mojokerto, Java, shows that this population did not have an extended childhood, which is a prerequisite for language acquisition.[67] On the other hand, despite the cochlear (ear) anatomy of Sangiran 2 and 4 retaining several traits reminiscent of australopithecines, the hearing range may have included the higher frequencies used to discern speech.[107]

Given expanding brain size and technological innovation, H. erectus may have been using some basic proto-language in combination with gesturing, and built the basic framework around which fully-fledged languages would eventually be formed.[108]

See also[edit | edit source]

Notes[edit | edit source]

Template:Notelist-num

References[edit | edit source]

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