The species was firstdescribed byEugène Dubois in 1893 as "Pithecanthropus erectus" using a skullcap,molar, andfemur fromJava, Indonesia. Further discoveries around East Asia were used to contend that humanity evolved out of Asia. Based onhistorical race concepts, it was argued that localH. erectus populations evolved directly into local modern human populations (polycentrism) rather than all humanity sharing a singleanatomically 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). WesternH. erectus brain volume could be as low as 546 cc (33.3 cu in) inH. e. georgicus.H. erectus probably had a faster apelikegrowth trajectory, lacking theextended childhood required forlanguage 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 theAcheuleantool industry, a major innovation of large, heavy-dutystone tools. These may have been used in butchery, vegetable processing, and woodworking ofspears anddigging sticks.H. erectus was a major predator of large herbivores on the expanding savannas during theQuaternary glaciation. The species is usually characterized as the firsthunter-gatherer and the first to practicesexual 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 ofH. 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.
In 1868,Ernst Haeckel suggested early humans dispersed from the now-disproven hypothetical continent "Lemuria".[3][4]
WhileCharles Darwin had hypothesized in his 1871Descent of Man that humans most likely evolved in Africa,[note 2] 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 wasErnst Haeckel, who argued that the first human species (which he speculatively namedHomo primigenius) evolved on a now-disproven hypothetical continent "Lemuria" from a species he termedPithecanthropus alalus (speechless ape-man). "Lemuria" had supposedly sunk below theIndian Ocean, accounting for the lack of fossil evidence.[3]
Dubois argued that "P. erectus" was agibbon-like ape which was the precursor to a more familiar human body plan, but in the 1930s, German-American anatomistFranz Weidenreich noticed a striking similarity with ancient human remains recently being unearthed in China (Peking Man, "Sinanthropus pekinensis").[8][9] This characterization became better supported as German-Dutch palaeontologistGustav Heinrich Ralph von Koenigswald discovered more Indonesian ancient human remains over the decade atMojokerto,Sangiran, andNgandong.[note 4][11][12] Weidenreich believed that they were the direct ancestors of the local modern humanHomo sapiens subspecies, in accord withhistorical race concepts (polycentricism) — that is, Peking Man was the direct ancestor of specificallyChinese people, and Java Man ofAboriginal Australians.[13][14] As the significance of racial distinction diminished with the development ofmodern evolutionary synthesis, many fossil human species and genera around Asia, Africa, and Europe (including "Pithecanthropus" and "Sinanthropus") were reclassified as subspecies ofHomo erectus.[15][16]
During the late 20th century, some of the oldestH. erectus fossils were discovered across Africa, the first being Kenyan archeologistLouis Leakey'sOlduvai Hominin 9 in 1960.[17] As the human fossil record expanded, the "Out of Africa" theory andmonogenism became the consensus: that all modern humans share a fullyanatomically 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.[18]
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.[19] In 1940, Weidenreich was the first to suggest reclassifying "Sinanthropus pekinensis" and "Pithecanthropus erectus" as subspecies ofH. erectus.[13] In 1950, German-American evolutionary biologistErnst Mayr entered this field. Surveying a "bewildering diversity of names" and many proposals for consolidation, he decided to reclassify human fossils into three species ofHomo: "H. transvaalensis" (theaustralopithecines),H. erectus (including "Sinanthropus", "Pithecanthropus", and various other Asian, African, and European taxa), andH. sapiens (including anything younger thanH. erectus, such as modern humans and Neanderthals). Mayr defined these species as a sequential lineage, each evolving into the next (chronospecies).[14] Though later Mayr changed his opinion on the australopithecines (recognizingAustralopithecus), his moreconservative view ofarchaic human diversity became widely adopted in the subsequent decades.[15]
...never more than one species of man existed on the earth at any one time... If fossils ofCongo pygmies and ofWatusi 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.
In the 1970s, aspopulation genetics was being formulated, the anatomical variation ofH. erectus across its wide geographic and temporal range (the basis for the subspecies distinctions) became better understood asclines — different populations which attained some anatomical regionality but were not reproductively isolated.[18] In general, subspecies names forH. erectus are now used for convenience to indicate time and region rather than specific anatomical trends.[20]
...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.
The ancient Georgia fossils have variably been classified asH. e. ergaster (or quadrinomialH. e. ergaster georgicus),[25] as their own subspecies asH. e. georgicus, or as their own speciesH. georgicus.[23] Some authors may also elevateH. ergaster,[26]H. soloensis,[27] andH. pekinensis to species level.[28] Fossils relegated toH. e. tautavelensis are traditionally assigned toH. heidelbergensis.[24]
H. e. georgicus (above) represents one of the earliest dispersals out of Africa about 1.8 million years ago.[29]
H. erectus evolved in Africa from a population ofH. habilis[30][31] and they coexisted for about half a million years.[32] The oldest identifiedH. erectus specimen is a 2.04 million year old skull, DNH 134, fromDrimolen, South Africa, coexisting with the australopithecineParanthropus robustus.[33]H. erectus dispersed out of Africa soon after evolution, the earliest recorded instances beingH. e. georgicus 1.78 to 1.85 million years ago in Georgia[29] and the Indonesian Mojokerto and Sangiran sites 1.6 to 1.8 million years ago.[34][35] Populations may have pushed into northwestern Europe at around the same time.[36] WhileH. erectus is usually considered the firsthominin to leave Africa, old stone tools that possibly date to as far back as 2.48 and 2.1 million year from respectivelyZarqa Valley, Jordan, andShangchen, China, could indicate thatH. erectus or earlier hominin species left Africa earlier than the current estimated range of dates.[37] SinceH. erectus was first defined in East Asia, those populations are sometimes distinguished asH. erectus sensu stricto ("in the strict sense"), and African and West Eurasian populations asH. erectus sensu lato ("in the broad sense"), but this may not reflect how these populations are actually related to each other.[note 5][20][31]
Once established around theOld World,H. erectus evolved into other later species in the genusHomo, including:H. heidelbergensis,H. antecessor,[38]H. floresiensis,[39] andH. luzonensis.[40]H. heidelbergensis, in turn, is usually placed as thelast common ancestor ofNeanderthals (H. neanderthalensis),Denisovans,[38] and, formerly,[41][42][43] modern humans (H. sapiens).H. erectus is thus a non-natural,paraphyletic grouping of fossils and does not include all the descendants of a last common ancestor.[44] Despite being designated as a different species,H. erectus may haveinterbred with some of its descendant species, namely the common ancestor of Neanderthals and Denisovans ("Neandersovans").[45]
The dispersal ofH. erectus is generally ascribed to the evolution of obligatebipedalism, better technology, and adoption of a carnivorous diet.[46] However, the sudden adoption of carnivory could besampling bias, with earlier species consuming the same amount of meat.[47] Populations spread out via open grassland and woodlandsavannas, which were expanding due to a globalaridification trend at the onset of theQuaternary glaciation.[46]H. erectus is usually thought to have occupied theSahara and West Asia duringhumid periods, but populations may have persisted into desert periods.[48]
MostH. 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 ofMiddle PleistoceneHomo has been a controversial topic, termed "the muddle in the middle".[24][49]H. erectus sensu stricto persisted much longer thansensu lato, with the youngest population (H. e. soloensis) dating to 108,000 to 117,000 years ago inLate Pleistocene Java.[1] This population appears to have died out when the savannah corridors closed and tropical jungle took over.[50]
A 2021 phylogeny of someH. erectus fossils usingtip dating:[44]
As such a widely distributed species both across regions and through time, the anatomy ofH erectus can vary considerably. Among livingprimates, the degree of regionality achieved byH. erectus (phenotypic plasticity) is only observed in modern humans.[51]
Dubois originally described the species using a skullcap, noting the traits of a low and thickenedcranial vault and a continuous bar of bone forming the brow ridge (supraorbital torus).[52]H. erectus fossils typically share these traits, but the KenyanKoobi Fora skulls notably have thinner skulls and weaker supraorbital tori.[53] He also used several other traits now considered more typical ofH. erectus sensu stricto, such as asagittal 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 themastoid part of the temporal bone.[52] These traits can be still be found, nonetheless, in a fewH. erectus sensu lato specimens, namely the 1.47 million year old Olduvai Hominin 9.[54]
Compared toH. erectus sensu lato, the skullcap ofsensu stricto narrows considerably at the front, the face is bigger and presumably moreprognathic (it juts out more, but the face is poorly documented), and the molars are larger particularly in Indonesian fossils.[55]H. erectus was the first human species with a fleshynose, which is generally thought to have evolved in response to breathing dry air in order to retain moisture.[56] Compared to earlierHomo,H. erectus has smaller teeth, thinnerenamel, and weakermandibles (jawbone), likely due to a greater reliance on tool use and food processing.[57]
The brain size ofH. erectus varies considerably, but is generally smaller inH. erectus sensu lato, as low as 546 cc (33.3 cu in) inDmanisi skull 5.[58] East AsianH. erectus overall are rather big-brained, averaging roughly 1,000 cc,[51] staying within the range of variation for modern humans.[59] The late-survivingH. e. soloensis has the biggest brain volume with one specimen measuring 1,251 cc (76.3 cu in).[54]
The rest of the body is primarily understood by three partial skeletons from the KenyanLake Turkana site, notablyTurkana Boy. Other postcranial fossils (all bones aside from the skull) attributed toH. erectus are not associated with a skull, making attribution unverifiable. Though the body plan of earlierHomo is poorly understood,H. erectus is usually characterized as the firstHomo species with a human body plan, distinct from non-human apes.[60][51][61] The chest may have been short andbarrel-shaped, like other archaic humans.[62]Fossil tracks nearIleret, Kenya, suggest ahuman gait. This adaptation is implicated in the dispersal ofH. erectus across the Old World.[63]
It is unclear when human ancestorslost most of their body hair. Genetic analysis suggests that high activity in themelanocortin 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 harmfulUV radiation.[64] It is possible that populations in higher latitudes developed lighter skin to preventvitamin D deficiency,[65] though a 300,000 to 500,000 year old TurkishH. erectus specimen presents the earliest case oftuberculous meningitis, which is typically exacerbated by vitamin D deficiency in dark-skinned people living in higher latitudes.[66] Hairlessness is generally thought to have facilitated sweating,[67] but it may also have helped to reduce parasite load, and was possibly reinforced bysexual selection.[68][69]
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-specificsexual dimorphism, but the variability of postcranial material makes this unclear.[51] A 2010 study estimates that theTurkana Boy would have reached a height of 163 cm (5 ft 4 in) if he had reached adulthood.[70]
The dimensions of a 1.8 million years old adult femaleH. e. ergaster pelvis fromGona, 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 betweenchimpanzees (~40%) and modern humans (28%).[71] 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.[72] This suggests that the childhood growth and development ofH. erectus was intermediate between that of chimpanzees and modern humans,[71] and the faster development rate suggests thataltriciality (an extended childhood) evolved at a later stage in human evolution.[72] The faster development rate might also indicate a shorter expected lifespan compared to laterHomo.[73]
Cross sections ofPeking Man Skulls III (A) and XII (B), andJava Man Skull II (C)
The bones are extraordinarily thickened, particularly inHomo erectus sensu stricto, so much so that skull fragments have sometimes been confused for fossil turtlecarapaces.[74] Themedullary canal in thelong bones (where thebone marrow is stored, in the limbs) is extremely narrowed (medullarystenosis). This degree of thickening is usually exhibited in semi-aquatic animals which use their heavy (pachyosteosclerotic) bones as ballasts to help them sink, induced byhypothyroidism.[75]
It is unclear what function intense bone thickening could have served. Before more complete skeletons were discovered, Weidenreich suggestedH. erectus was a gigantic species.[76] Other explanations include a far more violent and impact-prone lifestyle than otherHomo, or pathological nutrient deficiencies.[77] The supraorbital torus thickens with age, and may be a response tobending stresses from habitual loading of the front teeth.[78]
H. erectus overhunting may have led to the extinction ofMegalochelys (above).[79]
H. erectus was early-on portrayed as the earliesthunter-gatherer and a skilled predator of big game, relying on running. The few identified specimens of theH. e. ergaster torso and pelvis may indicate a body plan more conducive for power running, unlike modern humans betteradapted for endurance running.[62] The gradual shift to "top predator" may have led to its dispersal throughout Afro-Eurasia.[46] 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,[80] especiallyelephants,rhinos,hippos,bovines, andboars. The complexities of prey behaviors and the nutritional value of meat have been connected to brain volume growth.[81]
H. erectus is usually assumed to have practicedsexual division of labor much like recent hunter-gatherer societies, with men hunting and women gathering. This ideation is supported by a fossil trackway from Ileret, Kenya, made by a probably all-male band of over 20H. erectus individuals, possibly a hunting party or (similar to chimpanzees) a border patrol group.[82]
Since common modern human tapeworms began to diverge from those of other predators roughly 1.7 million years ago (specifically thepork tapeworm,beef tapeworm, andAsian tapeworm), not only wasH. erectus consuming meat regularly enough for speciation to occur in these parasites, but meat was probably consumed raw more often than not.[83] Some populations were collecting aquatic resources, include fish, shellfish, and turtles such as at Lake Turkana[84] and Trinil.[85] Undergroundstorage organs (roots, tubers, etc.) were likely also major dietary components, and traces of the edible plantCeltis have been documented at severalH. erectus sites.[86]
Possibly due to overhunting of the biggest game available, the dispersal ofH. erectus and descendant species may be implicated in the extinctions of large herbivores and the gradual reduction of average herbivore size over the Pleistocene.[87]H. erectus overhunting has been blamed by some authors for the decline ofproboscidean species as well as competing carnivores,[81][88][89] but their decline may be better attributed to the spread of grasslands.[89][90] The giant tortoiseMegalochelys may have been driven to extinction byH. erectus inSundaland (what is nowIsland Southeast Asia), since species went extinct shortly after the arrival ofH. erectus.[79]
H. erectus manufacturedLower Paleolithic technologies, and is credited with the invention of theAcheulean stone toolindustry at latest 1.95 million years ago.[91] This was a major technological breakthrough featuring large, heavy-duty tools; most iconically, thehandaxe. Over hundreds of thousands of years, the Achuelean eventually replaced its predecessor — theOldowan (achopper andflake industry) — in Africa, and spread out across Western Eurasia.[92] This sudden innovation was typically explained as a response to environmental instability in order to process more types of food and broaden the diet, which allowedH. erectus to colonize Eurasia. Despite this characterization of the Acheulean,H. e. georgicus was able to leave Africa despite only manufacturing Oldowan-style tools,[46][92] and the handaxe does not seem to have been manufactured commonly in East Asia.[93] This conspicuous pattern was first noted by American archaeologistHallam 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".[94] Movius took this as evidence of inferiority of Far Eastern populations:
...as early asLower 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.
H. erectus seems to have been using stone tools in butchery, vegetable processing, and woodworking (maybe manufacturingspears anddigging sticks).[86][95] In Africa, Oldowan sites are typically found alongside major fossil assemblages, but Acheulean sites normally feature more stone tools than fossils, soH. erectus could have been using choppers and handaxes for different activities.[95]
Materials for stone tools were normally sourced locally, and it seemsblanks were usually chosen based on size rather than material quality.[92]H. erectus also produced tools from shells at Sangiran[96] and Trinil.[97]
H. erectus is credited as the first human species to wield fire. The earliest claimed fire site isWonderwerk Cave, South Africa, at 1.7 million years old.[98] 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,[99] and cave-dwelling about 600,000 years ago.[100] Therefore,H. erectus may have only been scavenging fire opportunistically. Similarly,H. erectus sites usually stay within warmer tropical or subtropical latitudes.[46]
The dating of northerly populations (namely Peking Man) could suggest that they were retreating to warmer refugia duringglacial periods, but the precise age of the Peking Man fossils is poorly resolved.[52][101] There have been claims of manmade hearths and "clear-cut evidence for intentional fire use",[102] ostensibly as far back as 770,000 years ago in the supposed cave home of Peking Man.[101] 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.[24]
The single-toothedH. e. georgicus specimen (above) is the earliest probable example of group care.[103]
Like other primates,H. erectus probably usedmedicinal plants[86] and infirmed sick group members. The earliest probable example of this is a 1.77 million year oldH. e. georgicus specimen who had lost all but one tooth due to age orgum disease (the earliest example of severe chewing impairment) yet still survived for several years afterwards.[103]
H. erectus made long sea crossings to arrive on the islands ofFlores,Luzon,[104] and someMediterranean islands. Some authors have asserted thatH. 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 bynatural rafting events instead.[105]
In East Asia,H. erectus is usually represented only by skullcaps, which used to be interpreted as widespreadcannibalism and ritualheadhunting. 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 theseH. erectus populations. The lack of the rest of the skeleton is now normally explained by natural phenomena.[106]
Art-making could be evidence of symbolic thinking. An engravedPseudodon shellDUB1006-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.[97][107][108]H. erectus was also the earliest human to collect red-colored pigments, namelyochre. 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 ahammerstone. Red ochre is normally recognized as bearing symbolic value when associated with modern humans.[108]
Thespinal column of the 1.6 million year old Turkana boy would not have supported properly developed respiratory muscles required to produce speech;[109][110] and a 1.5 million year old infantH. erectus skull from Mojokerto, Java, shows that this population did not have an extended childhood, which is a prerequisite forlanguage acquisition.[72] On the other hand, despite thecochlear (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.[111]
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.[112]
^Darwin's work mainly aimed to demonstrate that histheory ofcommon descent with modification bynatural andsexual 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 thedifferences between the so-calledraces of man."[5]
^Dubois was studying the anatomy and evolution of thelarynx invertebrates inAmsterdam with German anatomistMax Fürbringer, but fed up with both the research and Fürbringer, he decided to drop everything and board theSS Prinses Amalia to the Dutch East Indies on 29 October 1887. Dubois said his interest in human evolution began in secondary school (the StateHogere Burgerschool), where he heard of lectures given byCarl Vogt in 1868.[6]
^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. SirArthur 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."[10]
^There may have been two differentH. erectus dispersals into East Asia: an early one that led to the SangiranJava Man (specifically the older material), and a later one which led to the northern ChinesePeking Man. That is, the older Sangiran material may be more closely related toH. erectus sensu lato than to Peking Man. The younger Sangiran material, though, possibly descended from or interbred with ChineseH. erectus.[31]
^H. heidelbergensis is also aparaphyletic assemblage of fossils.[44]
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