Homo erectus

Homo erectus (meaning 'upright man') is a species of archaic humans that lived throughout most of the Pleistocene geological epoch. Its earliest fossil evidence dates to 1.8 million years ago (discovered 1991 in Dmanisi, Georgia).[5]

A debate regarding the classification, ancestry, and progeny of H. erectus, especially in relation to Homo ergaster, is ongoing, with two major positions:

1) H. erectus is the same species as H. ergaster, and thereby H. erectus is a direct ancestor of the later hominins including Homo heidelbergensis, Homo antecessor, Homo neanderthalensis, Homo denisova, and Homo sapiens; or,
2) it is in fact an Asian species or subspecies distinct from African H. ergaster.[6]

Some paleoanthropologists consider H. ergaster to be a variety, that is, the "African" variety, of H. erectus; the labels "Homo erectus sensu stricto" (strict sense) for the Asian species and "Homo erectus sensu lato" (broad sense) have been offered for the greater species comprising both Asian and African populations.[7][8]

H. erectus eventually became extinct throughout its range in Africa, Europe and Asia, but developed into derived species, notably Homo heidelbergensis. As a chronospecies, the time of its disappearance is thus a matter of contention. The species name proposed in 1950 defines Java Man as the type specimen (now H. e. erectus). Since then, there has been a trend in palaeoanthropology of reducing the number of proposed species of Homo, to the point where H. erectus includes all early (Lower Paleolithic) forms of Homo sufficiently derived from H. habilis and distinct from early H. heidelbergensis (in Africa also known as H. rhodesiensis).[9] In this wider sense, H. erectus had mostly been replaced by H. heidelbergensis by about 300,000 years ago, with possible late survival in Java as late as 70,000 years ago.[1] The discovery of the morphologically divergent Dmanisi skull 5 in 2013 has reinforced the trend of subsuming fossils formerly given separate species names under H. erectus considered as a wide-ranging, polymorphous species.[10] Thus, H. ergaster is now well within the accepted morphological range of H. erectus, and it has been suggested that even H. rudolfensis and H. habilis (alternatively suggested as late forms of Australopithecus rather than early Homo) should be considered early varieties of H. erectus.[11][12]

Homo erectus
Temporal range: 2–0.07 Ma
Early PleistoceneLate Pleistocene[1]
Homme de Tautavel 01-08
Reconstructed skeleton of
Tautavel Man[2]
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Primates
Suborder: Haplorhini
Infraorder: Simiiformes
Family: Hominidae
Subfamily: Homininae
Tribe: Hominini
Genus: Homo
Species:
H. erectus
Binomial name
Homo erectus
(Dubois, 1893)
Synonyms
Homo erectus adult female - head model - Smithsonian Museum of Natural History - 2012-05-17
Forensic reconstruction of an adult female Homo erectus.[3]
Homo erectus new
Forensic reconstruction of an adult male Homo erectus.[4]

Discovery and type specimen

The Dutch anatomist Eugène Dubois, inspired by Darwin's theory of evolution as it applied to humanity, set out in 1886 for Asia (despite Darwin's theory of African origin) to find a human ancestor. In 1891–92, his team discovered first a tooth, then a skullcap, and finally a femur of a human fossil on the island of Java, Dutch East Indies (now Indonesia). Excavated from the bank of the Solo River at Trinil, in East Java, he first (1893) allocated the material to a genus of fossil chimpanzees as Anthropopithecus erectus, then the following year assigned his species to a new genus as Pithecanthropus erectus (the genus name had been coined by Ernst Haeckel in 1868 for the hypothetical link between humans and fossil Apes)—from the Greek πίθηκος (píthēkos, "ape") and ἄνθρωπος (ánthrōpos, "human")—based on the proposal that the femur suggested that the creature had been bipedal, like Homo sapiens.

Dubois' 1891 find was the first fossil of a Homo-species (or any hominin species) found as result of a directed expedition and search (the first recognized human fossil had been the circumstantial discovery of Homo neanderthalensis in 1856; see List of human evolution fossils). The Java fossil from Indonesia aroused much public interest. It was dubbed by the popular press as Java Man; but few scientists accepted Dubois' argument that his fossil was the transitional form—the so-called "missing link"—between humans and nonhuman apes.[13]

Ficha del homo georgicus. Museo Arqueológico Nacional de España
Poster of homo georgicus. National Archaeological Museum of Spain.

Most of the spectacular discoveries of H. erectus next took place at the Zhoukoudian Project, now known as the Peking Man site, in Zhoukoudian, China. This site was first discovered by Johan Gunnar Andersson in 1921[14] and was first excavated in 1921, and produced two human teeth.[15] Davidson Black's initial description (1921) of a lower molar as belonging to a previously unknown species (which he named Sinanthropus pekinensis)[16] prompted widely publicized interest. Extensive excavations followed, which altogether uncovered 200 human fossils from more than 40 individuals including five nearly complete skullcaps.[17] Franz Weidenreich provided much of the detailed description of this material in several monographs published in the journal Palaeontologica Sinica (Series D).

Nearly all of the original specimens were lost during World War II during an attempt to smuggle them out of China for safekeeping; however, authentic casts were made by Weidenreich, which exist at the American Museum of Natural History in New York City and at the Institute of Vertebrate Paleontology and Paleoanthropology in Beijing, and are considered to be reliable evidence.

Similarities between Java Man and Peking Man led Ernst Mayr to rename both Homo erectus in 1950.

Throughout much of the 20th century, anthropologists debated the role of H. erectus in human evolution. Early in the century, due in part to the discoveries at Java and Zhoukoudian, the belief that modern humans first evolved in Asia was widely accepted. A few naturalists—Charles Darwin most prominent among them—theorized that humans' earliest ancestors were African: Darwin pointed out that chimpanzees and gorillas, humans' closest relatives, evolved and exist only in Africa.[18]

Origin and dispersal

Carte hachereaux
Map of the distribution of Middle Pleistocene (Acheulean) cleaver finds

The derivation of the genus Homo from Australopithecina took place in East Africa after 3 million years ago. The inclusion of species dated to just before 2 million years ago, Homo habilis and Homo rudolfensis, into Homo is somewhat contentious.[19] Especially as H. habilis appears to have coexisted with H. ergaster/erectus for a substantial period after 2 Mya, it has been proposed that ergaster may not be directly derived from habilis.[20]

Homo erectus emerged about 2 million years ago. Fossils dated close to 1.8 million years ago have been found both in Africa and in West Asia, so it is unclear whether H. erectus emerged in Africa or in Asia. Ferring et al. (2011) suggest that it was still H. habilis which reached West Asia, and that early H. erectus developed there. Early H. erectus would then have dispersed from West Asia, to East Asia (Peking Man) Southeast Asia (Java Man), back to Africa (Homo ergaster), and to Europe (Tautavel Man).[21][22]

Africa

Homo ergaster
KNM-ER 3733 (1.6 Mya, discovered 1975 at Koobi Fora, Kenya)

In the 1950s, archaeologists John T. Robinson and Robert Broom named Telanthropus capensis;[23] Robinson had discovered a jaw fragment in 1949 in Swartkrans, South Africa. Later, Simonetta proposed to re-designate it to Homo erectus, and Robinson agreed.[24]

From the 1950s forward, numerous finds in East Africa suggested sympatric coexistence for H. ergaster and H. habilis for several hundred millennia, which tends to confirm the hypothesis that they represent separate lineages from a common ancestor; that is, the ancestral relationship between them was not anagenetic, but was cladogenetic, which here suggests that a subgroup population of H. habilis—or of a common ancestor of H. habilis and H. ergaster/erectus—became reproductively isolated from the main-group population, eventually evolving into the new species Homo ergaster (Homo erectus sensu lato).[25]

In 1961, Yves Coppens discovered a skull in northern Chad. He coined the name Tchadanthropus uxoris for what he considered the earliest fossil human discovered in north Africa.[26] Although once considered to be a specimen of H. habilis,[27] T. uxoris has been subsumed into H. erectus but it is no longer considered a valid taxon.[26][28] It was reported that the fossil "had been so eroded by wind-blown sand that it mimicked the appearance of an australopith, a primitive type of hominid".[29] It is probably only 10,000 years old according to stratigraphy, paleontology and C14 dating presented in Michel Servant's PhD as early as 1973.[30]

Eurasia

Caucasus

Homo Georgicus IMG 2921
Dmanisi skull 3 (fossils skull D2700 and jaw D2735, two of several found in Dmanisi in the Georgian Transcaucasus)
MEH Homo georgicus 29-04-2012 11-35-22 2372x3863
Reconstruction of Homo georgicus based on D2700, by Élisabeth Daynès, Museo de la Evolución Humana, Burgos, Spain.

Homo erectus georgicus is the subspecies name assigned to fossil skulls and jaws found in Dmanisi, Georgia. First proposed as a separate species, it is now classified within H. erectus.[31][32][33] The site was discovered in 1991 by Georgian scientist David Lordkipanidze. Five skulls were excavated from 1991 forward, including a "very complete" skull in 2005. Excavations at Dmanisi have yielded 73 stone tools for cutting and chopping and 34 bone fragments from unidentified fauna.[34]

After their initial assessment, some scientists were persuaded to name the Dmanisi find as a new species, Homo georgicus, which they posited as a descendant of African Homo habilis and an ancestor to Asian Homo erectus. This classification, however, was not supported, and the fossil was instead designated a divergent subgroup of Homo erectus.[35][36][37][38]

The fossil skeletons present a species primitive in its skull and upper body but with relatively advanced spine and lower limbs, implying greater mobility than the previous morphology.[39] It is now thought not to be a separate species, but to represent a stage soon after the transition between H. habilis to H. erectus; it has been dated at 1.8 Mya.[32][40] The assemblage includes one of the largest Pleistocene Homo mandibles (D2600), one of the smallest Lower Pleistocene mandibles (D211), a nearly complete sub-adult (D2735), and a toothless specimen D3444/D3900.[41]

Two of the skulls—D2700, with a brain volume of 600 cubic centimetres (37 cu in), and D4500 or Dmanisi Skull 5, with a brain volume of about 546 centimetres—present the two smallest and most primitive Hominina skulls from the Pleistocene period.[11] The variation in these skulls were compared to variations in modern humans and within a sample group of chimpanzees. The researchers found that, despite appearances, the variations in the Dmanisi skulls were no greater than those seen among modern people and among chimpanzees. These findings suggest that previous fossil finds that were classified as different species on the basis of the large morphological variation among them—including Homo rudolfensis, Homo gautengensis, H. ergaster, and potentially even H. habilis—should perhaps be re-classified to the same lineage as Homo erectus.[42]

East and Southeast Asia

H. erectus is attested with certainty in East and Southeast Asia from about 0.7 Ma, with possible early presence before 1 Ma; stone tools from Shangchen discovered in 2018 were even claimed to be older than 2 Mya.[43][44]

Meganthropus refers to a group of fossils found in Java, dated to between 1.4 and 0.9 Mya, which are tentatively grouped with H. erectus, at least in the wider sense of the term in which "all earlier Homo populations that are sufficiently derived from African early Homo belong to H. erectus",[9] although older literature has placed the fossils outside of Homo altogether.[45]

Java Man (H. e. erectus, the type specimen for H. erectus), discovered on the island of Java in 1891/2, is dated to 1.0–0.7 Mya. Lantian Man (H. e. lantianensis), discovered in 1963 in Lantian County, Shaanxi province, China, is roughly contemporary with Java Man.

Peking Man (H. e. pekinensis), discovered in 1923–27 at Zhoukoudian (Chou K'ou-tien) near Beijing, China, dates to about 0.75 Mya.[46] and a new 26Al/10Be dating suggests they are in the range of 680,000–780,000 years old.[47][48] Yuanmou Man (H. e. yuanmouensis), discovered in Yuanmou County in Yunnan, China, in 1965, is likely of similar age as Peking Man (but with dates proposed as early as 1.7 Mya).[49]

Nanjing Man (H. e. nankinensis), discovered in 1993 in the Hulu cave on the Tangshan hills near Nanjing, dates to about 0.6 Mya.[50][51]

Solo Man (H. e. soloensis), discovered between 1931/1933 along the Solo River, Java, is of uncertain age, dated between 0.25 and 0.075 Mya (the younger date would qualify Solo Man as the latest fossil still classified as H. erectus, even though it shows some derived features, notably larger cranial capacity).[52]

Europe

It is conventional to label European archaic humans contemporary with late H. erectus under the separate species name of Homo heidelbergensis, the immediate predecessor of Homo neanderthalensis. H. heidelbergensis fossils are recorded from 600 ka (Mauer 1 mandible), the oldest complete skulls are "Tautavel Man" (Homo erectus tautavelensis), c. 450 ka, and the Atapuerca skull ("Miguelón"), c. 430 ka. The oldest known human fossils found in Europe is a molar from the Sima del Elefante site, Atapuerca Mountains, Spain, dated c. 1.2 Mya. This is associated with the skull fragments of the "Boy of Gran Dolina" found nearby, dated 0.9 Mya and classified as Homo antecessor by the discoverers. The relationship of these fossils to H. erectus is open to debate, as no complete skull has been found. H. e. bilzingslebenensis (Vlcek 1978) refers to skull fragments found at the Bilzingsleben site, Thuringia, Germany.

There is, however, indirect evidence of human presence in Europe as early as 1.6 Mya, in the form of stone tools discovered at Lézignan-la-Cèbe, France, in 2008.[53] Other stone tools found in France and thought to predate 1 Mya are from Chilhac, Haute-Loire) and from the Grotte du Vallonnet, near Menton. Human presence in Great Britain close to 1 Mya is established by stone tools and fossilized footprints found near Happisburgh, Norfolk.[54][55]

Taxonomy

Homo-Stammbaum, Version Stringer-en
One proposed model of the evolution of several species of genus Homo over the last 2 million years (vertical axis) based on Stringer (2012).[56]
Human evolution chart-en
An alternate model proposed by Reed, et al., redrawn from Stringer.[57] Note the depiction of Homo ergaster as an ancestor of Homo erectus.

Paleoanthropologists continue to debate the classification of Homo erectus and Homo ergaster as separate species. One school of thought calls H. ergaster the direct African ancestor of H. erectus, proposing that erectus emigrated out of Africa to Asia while branching into a distinct species.[58] Some scholars dispense with the species name ergaster, making no distinction between such fossils as the Turkana Boy and Peking Man. Still, "Homo ergaster" has gained some acceptance as a valid taxon, and the two species are still usually defined as distinct African and Asian populations of the greater species H. erectus, that is, "Homo erectus sensu lato".

One of the features distinguishing H. erectus from H. Sapiens is the size difference in teeth. H. erectus has large teeth while H. sapiens have smaller teeth.[59] One theory for the reason of H. erectus having larger teeth is because of the requirements for eating raw meat instead of cooked meat like H. sapiens.

Some have insisted that Ernst Mayr's biological species definition cannot be used to test the above hypotheses—that is, that the two species might be considered the same. Alternatively, the amount of variation of cranial morphology between known specimens of H. erectus and H. ergaster can be compared to the same variation within an appropriate population of living primates (that is, one of similar geographical distribution or close evolutionary relationship), such that: if the amount of variation between H. erectus and H. ergaster is greater than that within an appropriately selected population, for example, say, macaques, then H. erectus and H. ergaster may be considered as two different species.

Homo erectus skull
The skull of Homo Erectus; note the large teeth.
Homo sapiens-MGL on loan from MAHL-P5030047-white
The skull of Homo Sapiens, with its smaller teeth.

Finding an extant (i.e., living) model suitable for field study, analysis, and comparison is very important; and selecting a living sample population of an appropriate species can be difficult. (For example, the morphological variation among the global population of H. sapiens is small,[60] so our own species diversity may not be a trustworthy comparison. Fossils found in Dmanisi, Georgia were originally designated as a separate (but closely related) species; but subsequent specimens showed their variation to be within the range of Homo erectus. and they are now classified as Homo erectus georgicus.) New foot tracks found in 2009 in Kenya and reported in Science by Matthew Bennett of Bournemouth University in Britain and his colleagues, confirmed that the gait of Homo erectus was heel-to-toe, walking as a modern human does, rather than with the australopithecine-like method of its own ancestors.[61]

H. erectus fossils show a cranial capacity greater than that of Homo habilis (although the Dmanisi specimens have distinctively small crania): the earliest fossils show a cranial capacity of 850 cm³, while later Javan specimens measure up to 1100 cm³,[60] overlapping that of H. sapiens.; the frontal bone is less sloped and the dental arcade smaller than that of the australopithecines; the face is more orthognatic (less protrusive) than either the australopithecines or H. habilis, with large brow-ridges and less prominent zygomata (cheekbones). The early hominins stood about 1.79 m (5 ft 10 in)[62]—only 17 percent of modern male humans are taller[63]—and were extraordinarily slender, with long arms and legs.[64]

Human arm bones diagram
Diagram of modern human shoulder bones.

Throwing performance may have been an important mode for early hunting and defense in the genus Homo. Throwing performance in the genus has previously been linked to several anatomical shifts in the upper body during the evolution of Homo. Different fossils and skeletal measures used in reconstructing the Homo erectus shoulder make it possible for either an anteriorly facing shoulder configuration or a lateral orientation that is similar to modern humans.[65] These two different orient the throwing ability and hunting behavior of early Homo species. However, it has been found that a commonly used metric for clavicle length (the claviculohumeral ratio) does not predict shoulder position on the torso accurately. Also, no connection between clavicle length and throwing performance was found. This new evidence supports the conclusion that Homo erectus fossil clavicles are similar to modern human variations, [65] with Homo erectus having a shoulder construction that was lateral facing. This suggests that the ability for high speed throwing can be dated back to nearly two million years ago.[65]

Sexual dimorphism in H. erectus—males are about 25% larger than females—is slightly greater than seen in the later H. sapiens, but less than that of the earlier genus Australopithecus. Regarding evolution of human physiology, the discovery of the skeleton of "Turkana boy" (Homo ergaster) near Lake Turkana, Kenya, by Richard Leakey and Kamoya Kimeu in 1984—one of the most complete hominin skeletons ever discovered—has contributed greatly to the interpretation.

Stringer (2003, 2012) and Reed, et al. (2004) and others have produced schematic graph-models for interpreting the evolution of Homo sapiens from earlier species of Homo, including Homo erectus and/or Homo ergaster, see graphs at right. Blue areas denote the existence of one or more hominin species at a given time and place (that is, region). These and other interpretations differ mainly in the taxonomy and geographical distribution of species.[56][57]

Stringer (see upper graph-model) depicts the presence of H. erectus as dominating the temporal and geographic development of human evolution; and as persisting broadly throughout Africa and Eurasia for nearly 2 million years, eventually evolving into H. heidelbergensis / H. rhodesiensis, which in turn evolved into H. sapiens. Reed, et al. shows Homo ergaster as the ancestor of Homo erectus; then it is ergaster, or a variety of ergaster, or perhaps a hybrid of ergaster and erectus, which develops into species that evolve into archaic and then modern humans and then out of Africa.

Both models show the Asian variety of Homo erectus going extinct recently. And both models indicate species admixture: early modern humans spread from Africa across different regions of the globe and interbred with earlier descendants of H. heidelbergensis / H. rhodesiensis, namely the Neanderthals, Denisovans, as well as unknown archaic African hominins. See admixture; and see Neanderthal admixture theory.[66]

Habitat

There has been evidence of H. erectus inhabiting a cave in Zhoukoudian, China.[59] This evidence consisted of remains, stones, charred animal bones, collections of seeds, and possibly ancient hearths and charcoal.[59] Although this does not prove that H. erectus lived in caves, it does show that H. erectus spent periods of time in caves of Zhoukoudian. Remains of H. erectus have more consistently been found in lake beds and in stream sediments.[59] This suggest that H. erectus also lived in open encampments near streams and lakes.

Behaviour

Tool use

Canto tallado 2-Guelmim-Es Semara
Early Homo erectus appears to have inherited Oldowan technology and refined it into the Acheulean industry beginning 1.7 million years ago.[67]
Bifaz cordiforme
A cordiform biface as commonly found in the Acheulean, associated with Homo erectus and derived species such as Homo heidelbergensis.

The Paleolithic Age (Old Stone Age) of prehistoric human history and industry is dated from 2.6 million years ago to about 10,000 years ago;[68] thus it closely coincides with the Pleistocene epoch of geologic time, which is 2.58 million to 11,700 years ago.[69] The beginning of early human evolution reaches back to the earliest innovations of primitive technology and tool culture. H. erectus were the first to use fire to cook and to make hand axes out of stone.

Homo ergaster used more diverse and sophisticated stone tools than its predecessors, where early Homo erectus used comparatively primitive tools. This is probably because H. ergaster inherited, used, and created tools first of Oldowan technology and later advanced the technology to the Acheulean.[70] Because the use of Acheulean tools began ca. 1.8 million years ago,[71] and the line of H. erectus diverged some 200,000 years before the general innovation of Acheulean industry in Africa, then it is plausible that the Asian migratory descendants of H. erectus made no use of Acheulean technology. It has been suggested that the Asian H. erectus may have been the first humans to use rafts to travel over bodies of water, including oceans.[72] And the oldest stone tool found in Turkey reveals that hominins passed through the Anatolian gateway from western Asia to Europe approximately 1.2 million years ago—much earlier than previously thought.[73]

Use of fire

East African sites, such as Chesowanja near Lake Baringo, Koobi Fora, and Olorgesailie in Kenya, show potential evidence that fire was utilized by early humans. At Chesowanja, archaeologists found fire-hardened clay fragments, dated to 1.42 M.Y.A.[74] Analysis showed that, in order to harden it, the clay must have been heated to about 400 °C (752 °F). At Koobi Fora, two sites show evidence of control of fire by Homo erectus at about 1.5 M.Y.A., with reddening of sediment associated with heating the material to 200–400 degrees Celsius (392–752 degrees Fahrenheit).[74] At a "hearth-like depression" at a site in Olorgesailie, Kenya, some microscopic charcoal was found—but that could have resulted from natural brush fires.[74]

In Gadeb, Ethiopia, fragments of welded tuff that appeared to have been burned, or scorched, were found alongside H. erectus–created Acheulean artifacts; but such re-firing of the rocks may have been caused by local volcanic activity.[74] In the Middle Awash River Valley, cone-shaped depressions of reddish clay were found that could have been created only by temperatures of 200 °C (392 °F) or greater. These features are thought to be burnt tree stumps such that the fire was likely away from a habitation site.[74] Burnt stones are found in the Awash Valley, but naturally burnt (volcanic) welded tuff is also found in the area.

A site at Bnot Ya'akov Bridge, Israel is reported to show evidence that H. erectus or H. ergaster controlled fire there between 790,000 and 690,000 years ago;[75] to date this claim has been widely accepted. Some evidence is found that H. erectus was controlling fire less than 250,000 years ago. Evidence also exists that H. erectus were cooking their food as early as 500,000 years ago.[76] Re-analysis of burnt bone fragments and plant ashes from the Wonderwerk Cave, South Africa, has been dubbed evidence supporting human control of fire there by 1 M.Y.A.[77]

There is archaeological evidence that Homo erectus cooked their food.[76]

Engravings and religion

Homo Erectus shell with geometric incisions circa 500,000 BP, Naturalis Biodiversity Center, Netherlands (with detail)
Homo Erectus shell with geometric incisions, circa 500,000 BP, has been claimed as the first known work of art. From Trinil, Java. Now in the Naturalis Biodiversity Center, Netherlands.[78][79]

It has previously been thought that engraving is indicative of modern cognition and behavior.[80] In the Dubois collection discovered by Eugène Dubois in 1891, a shell with a geometric engraving was found.[81][80][82] The engraving of the shell was dated back to a maximum age of 0.5460 ± .10 million years and a minimum age of 0.4360 ± .05 million years. It was made around the time Homo erectus was present and is the oldest geometric engraving found.[80][83][84] This shows that engraved patterns were a part of the Asian Homo erectus' cognition and neuromotor control.[80]

There is very little evidence for religion among Homo erectus, however there is a possibility [85]. Then again due to the span of time most religious artifacts would have been gone anyway, and seeing how their descendants, Homo naledi, and Homo neanderthalensis/sapiens have religious or pre-religious behavior, it is probable that this species had it.

Sociality

Homo erectus was probably the first hominin to live in a hunter-gatherer society, and anthropologists such as Richard Leakey believe that erectus was socially more like modern humans than the more Australopithecus-like species before it. Likewise, increased cranial capacity generally coincides with the more sophisticated tools occasionally found with fossils.

The discovery of Turkana boy (H. ergaster) in 1984 evidenced that, despite its Homo sapiens-like anatomy, ergaster may not have been capable of producing sounds comparable to modern human speech. It likely communicated in a proto-language lacking the fully developed structure of modern human language but more developed than the non-verbal communication used by chimpanzees.[86] This inference is challenged by the find in Dmanisi, Georgia, of an H. ergaster / erectus vertebrae (at least 150,000 years earlier than the Turkana Boy) that reflects vocal capabilities within the range of H. sapiens.[39] Both brain size and the presence of the Broca's area also support the use of articulate language.[87]

H. erectus was probably the first hominin to live in small, familiar band-societies similar to modern hunter-gatherer band-societies,[88] and is thought to be the first hominin species to hunt in coordinated groups, to use complex tools, and to care for infirm or weak companions. It is also unknown if they wore clothes and had tools such as bowls and utensils, however it is likely they did as they migrated to the colder regions of Asia and Europe and would have needed to carry tools, food, and water.

Descendants and subspecies

Homo erectus is the most, or one of the most, long-lived species of Homo, having existed well over one million years and perhaps over two million years; by contrast, Homo sapiens emerged about a quarter million years ago. If considering Homo erectus in its strict sense (that is, as referring to only the Asian variety) no consensus has been reached as to whether it is ancestral to H. sapiens or any later human species.

Homo erectus

"Wushan Man" was proposed as Homo erectus wushanensis, but is now thought to be based upon fossilized fragments of an extinct non-hominin ape.[91]

Related species

Regarding many archaic humans, there is no definite consensus as to whether they should be classified as subspecies of H. erectus or H. sapiens or as separate species.

Fossils

The lower cave of the Zhoukoudian cave, China, is one of the most important archaeological sites worldwide.[94] There have been remains of 45 homo erectus individuals found and thousands of tools recovered.[94] 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'.[94]

New evidence has shown that homo erectus does not have uniquely thick vault bones, like what was previously thought.[95] Testing showed that neither Asian or African homo erectus had uniquely large vault bones.[95]

Individual fossils

Some of the major Homo erectus fossils:

Homo erectus KNM ER 3733
Homo erectus KNM ER 3733 actual skull

Gallery

Tautavel UK 2

Replica of lower jaws of Homo erectus from Tautavel, France.

Calvaria Sangiran II (A)

Calvaria "Sangiran II" original, collection Koenigswald, Senckenberg Museum.

Homo erectus hand axe Daka Ethiopia

A reconstruction based on evidence from the Daka Member, Ethiopia

Pithecanthropus-erectus

Original fossils of Pithecanthropus erectus (now Homo erectus) found in Java in 1891.

See also

General:

References

  1. ^ a b Homo erectus soloensis, found in Java, is considered the latest known survival of H. erectus. Formerly dated to as late as 50,000 to 40,000 years ago, a 2011 study pushed back the date of its extinction of H. e. soloensis to 75,000 years ago at the latest. Indriati E, Swisher CC III, Lepre C, Quinn RL, Suriyanto RA, et al. 2011 The Age of the 20 Meter Solo River Terrace, Java, Indonesia and the Survival of Homo erectus in Asia.PLoS ONE 6(6): e21562. doi:10.1371/journal.pone.0021562.
  2. ^ based on numerous fossil remains of H. erectus. Museum of Prehistory Tautavel, France (2008 photograph)
  3. ^ Reconstruction by John Gurche (2010), Smithsonian Museum of Natural History, based on KNM ER 3733 and 992. Abigail Tucker, "A Closer Look at Evolutionary Faces", Smithsonian.com, 25 February 2010.
  4. ^ Reconstruction by W. Schnaubelt & N. Kieser (Atelier Wild Life Art), 2006, Westfälisches Museum für Archäologie, Herne, Germany.
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Further reading

External links

Early expansions of hominins out of Africa

Several expansions of populations of archaic humans (genus Homo) out of Africa and throughout Eurasia took place in the course of the Lower Paleolithic, and into the beginning Middle Paleolithic, between about 2.1 million and 0.2 million years ago (Ma).

These expansions are collectively known as Out of Africa I, in contrast to the expansion of Homo sapiens

(anatomically modern humans) into Eurasia, which may have begun shortly after 0.2 million years ago (known in this context as "Out of Africa II").The earliest presence of Homo (or indeed any hominin) outside of Africa dates to close to 2 million years ago.

A 2018 study claims human presence at Shangchen, central China, as early as 2.12 Ma based on

magnetostratigraphic dating of the lowest layer containing stone artefacts.

The oldest known human skeletal remains outside of Africa are from Dmanisi, Georgia (Dmanisi skull 4), and are dated to 1.8 Ma. These remains are classified as Homo erectus georgicus.

Later waves of expansion are proposed around 1.4 Ma (early Acheulean industries), associated with Homo antecessor and 0.8 Ma (cleaver-producing Acheulean groups, associated with Homo heidelbergensis).Until the early 1980s, early humans were thought to have been restricted to the African continent in the Early Pleistocene, or until about 0.8 Ma; Hominin migrations outside East Africa were apparently rare in the Early Pleistocene, leaving a fragmentary record of events.

Homo

Homo (Latin: homō, "human being") is the genus which emerged in the otherwise extinct genus Australopithecus that encompasses the extant species Homo sapiens (modern humans), plus several extinct species classified as either ancestral to or closely related to modern humans (depending on a species), most notably Homo erectus and Homo neanderthalensis.

The genus is taken to emerge with the appearance of Homo habilis, just over two million years ago. Genus Homo, together with the genus Paranthropus is probably sister to A. africanus in the genus Australopithecus, which itself had previously split from the lineage of Pan, the chimpanzees.Homo erectus appeared about two million years ago and, in several early migrations, it spread throughout Africa (where it is dubbed Homo ergaster) and Eurasia.

It was likely the first human species to live in a hunter-gatherer society and to control fire.

An adaptive and successful species, Homo erectus persisted for more than a million years, and gradually diverged into new species by around 500,000 years ago.Homo sapiens (anatomically modern humans) emerged close to 300,000 to 200,000 years ago, most likely in Africa, and Homo neanderthalensis emerged at around the same time in Europe and Western Asia.

H. sapiens dispersed from Africa in several waves, from possibly as early as 250,000 years ago, and certainly by 130,000 years ago, the so-called Southern Dispersal beginning about 70,000 years ago leading to the lasting colonisation of Eurasia and Oceania by 50,000 years ago.

Both in Africa and Eurasia, H. sapiens met with and interbred with archaic humans. Separate archaic (non-sapiens) human species are thought to have survived until around 40,000 years ago (Neanderthal extinction), with possible late survival of hybrid species as late as 12,000 years ago (Red Deer Cave people).

Homo Erectus (film)

Homo Erectus (released on DVD in the United States as National Lampoon's Stoned Age) is a 2007 comedy film written and directed by Adam Rifkin, and starring Giuseppe Andrews, Gary Busey, David Carradine, Ron Jeremy, Ali Larter, Hayes MacArthur, Adam Rifkin, and Talia Shire. The film premiered at the 2007 Slamdance Film Festival in January 2007.

Homo ergaster

Homo ergaster, also Homo erectus ergaster or African Homo erectus is an extinct chronospecies of the genus Homo that lived in eastern and southern Africa during the early Pleistocene, between about 1.9 million and 1.4 million years ago.

Originally proposed as a separate species, H. ergaster is now mostly considered either an early form, or an African variety, of H. erectus.The binomial name was published in 1975 by Groves and Mazák. The specific epithet, "ergaster", is derived from the Ancient Greek ἐργαστήρ ergastḗr - "workman", in reference to the advanced lithic technology developed by the species, thereby introducing the Acheulean industry.

KNM-ER 2598, a "H. erectus-like" occipital bone stands as the earliest evidence for H. erectus in Africa at approximately 1.9 million years ago (contemporary with Homo rudolfensis). There is a fossil gap between 1.9 and 1.6 million years ago, KNM-ER 3733 is the oldest known H. ergaster skull dated to about 1.6 million years ago. Its survival past 1.4 million years ago is uncertain, again due to a fossil gap, the next available African fossils allowing reliable morphological analysis are those of Homo rhodesiensis (African H. heidelbergensis), at 0.6 million years ago.

Homo heidelbergensis

Homo heidelbergensis is an extinct species or subspecies of archaic humans in the genus Homo, which radiated in the Middle Pleistocene from about 700,000 to 300,000 years ago, known from fossils found in Southern Africa, East Africa and Europe. African H. heidelbergensis has several subspecies. The subspecies are Homo heidelbergensis heidelbergensis, Homo heidelbergensis daliensis, Homo rhodesiensis, and Homo heidelbergensis steinheimensi. The derivation of Homo sapiens from Homo rhodesiensis has often been proposed, but is obscured by a fossil gap from 400–260 kya. The species was originally named Homo heidelbergensis due to the skeleton's first discovery near Heidelberg, Germany.The first discovery—a mandible—was made in 1907 by Otto Schoetensack. The skulls of this species share features with both Homo erectus and the anatomically modern Homo sapiens; its brain was nearly as large as that of Homo sapiens. The Sima de los Huesos cave at Atapuerca in northern Spain holds particularly rich layers of deposits where excavations were still in progress as of 2018.H. heidelbergensis was dispersed throughout Eastern and Southern Africa (Ethiopia, Namibia, Southern Africa) as well as Europe (England, France, Germany, Greece, Hungary, Italy, Portugal, Spain). Its exact relation both to the earlier Homo antecessor and Homo ergaster, and to the later lineages of Neanderthals, Denisovans, and modern humans is unclear.Homo sapiens has been proposed as derived from H. heidelbergensis via Homo rhodesiensis, present in East and North Africa from around 400,000 years ago. The correct assignment of many fossils to a particular chronospecies is difficult and often differences in opinion ensue among paleoanthropologists due to the absence of universally accepted dividing lines (autapomorphies) between Homo erectus, Homo heidelbergensis, Homo rhodesiensis and Neanderthals.

It is uncertain whether H. heidelbergensis is ancestral to Homo sapiens, as a fossil gap in Africa between 400,000 and 260,000 years ago obscures the presumed derivation of H. sapiens from H. rhodesiensis. Genetic analysis of the Sima de los Huesos fossils (Meyer et al. 2016) seems to suggest that H. heidelbergensis in its entirety should be included in the Neanderthal lineage, as "pre-Neanderthal" or "archaic Neanderthal" or "early Neanderthal", while the divergence time between the Neanderthal and modern lineages has been pushed back to before the emergence of H. heidelbergensis, to about 600,000 to 800,000 years ago, the approximate time of disappearance of Homo antecessor.The delineation between early H. heidelbergensis and H. erectus is also unclear.

Human taxonomy

Human taxonomy is the classification of the human species (systematic name Homo sapiens, Latin: "knowing man") within zoological taxonomy. The systematic genus, Homo, is designed to include both anatomically modern humans and extinct varieties of archaic humans. Current humans have been designated as subspecies Homo sapiens sapiens, differentiated from the direct ancestor, Homo sapiens idaltu.

Since the introduction of systematic names in the 18th century, knowledge of human evolution has increased drastically, and a number of intermediate taxa have been proposed in the 20th to early 21st century. The most widely accepted taxonomy groups takes the genus Homo as originating between two and three million years ago, divided into at least two species, archaic Homo erectus and modern Homo sapiens, with about a dozen further suggestions for species without universal recognition.

The genus Homo is placed in the tribe Hominini alongside Pan (chimpanzees). The two genera are estimated to have diverged over an extended time of hybridization spanning roughly 10 to 6 million years ago, with possible admixture as late as 4 million years ago. A subtribe of uncertain validity, grouping archaic "pre-human" or "para-human" species younger than the Homo-Pan split is Australopithecina (proposed in 1939).

A proposal by Wood and Richmond (2000) would introduce Hominina as a subtribe alongside Australopithecina, with Homo the only known genus within Hominina. Alternatively, following Cela-Conde and Ayala (2003), the "pre-human" or "proto-human" genera of Australopithecus, Ardipithecus, Praeanthropus, and possibly Sahelanthropus may be placed on equal footing alongside the genus Homo. An even more radical view rejects the division of Pan and Homo as separate genera, which based on the Principle of Priority would imply the re-classification of chimpanzees as Homo paniscus (or similar).Prior to the current scientific classification of humans, philosophers and scientists have made various attempts to classify humans. They offered definitions of the human being and schemes for classifying types of humans. Biologists once classified races as subspecies, but today anthropologists reject the concept of race and view humanity as an interrelated genetic continuum. Taxonomy of the hominins continues to evolve.

Ileret

Ileret (also spelled Illeret) is a village in Marsabit County, Kenya. It is located in Northern Kenya, on the eastern shore of Lake Turkana, north of Sibiloi National Park and near the Ethiopian border.

Numerous hominin fossils have been found near Ileret, including Homo erectus footprints dating back to about 1.5 million years ago, making them the second oldest hominin footprints ever found after those at Laetoli, Tanzania.

Java Man

Java Man (Homo erectus erectus; Javanese: Manungsa Jawa; Indonesian: Manusia Jawa) is an early human fossil discovered on the island of Java (Indonesia) in 1891 and 1892. Led by Eugène Dubois, the excavation team uncovered a tooth, a skullcap, and a thighbone at Trinil on the banks of the Solo River in East Java. Arguing that the fossils represented the "missing link" between apes and humans, Dubois gave the species the scientific name Anthropopithecus erectus, then later renamed it Pithecanthropus erectus.

The fossil aroused much controversy. Less than ten years after 1891, almost eighty books or articles had been published on Dubois's finds. Despite Dubois's argument, few accepted that Java Man was a transitional form between apes and humans. Some dismissed the fossils as apes and others as modern humans, whereas many scientists considered Java Man as a primitive side branch of evolution not related to modern humans at all. In the 1930s Dubois made the claim that Pithecanthropus was built like a "giant gibbon", a much misinterpreted attempt by Dubois to prove that it was the "missing link".

Eventually, similarities between Pithecanthropus erectus (Java Man) and Sinanthropus pekinensis (Peking Man) led Ernst Mayr to rename both Homo erectus in 1950, placing them directly in the human evolutionary tree. To distinguish Java Man from other Homo erectus populations, some scientists began to regard it as a subspecies, Homo erectus erectus, in the 1970s. Other fossils found in the first half of the twentieth century in Java at Sangiran and Mojokerto, all older than those found by Dubois, are also considered part of the species Homo erectus. Estimated to be between 700,000 and 1,000,000 years old, at the time of their discovery the fossils of Java Man were the oldest hominin fossils ever found. The fossils of Java Man have been housed at the Naturalis in the Netherlands since 1900.

Lantian Man

Lantian Man (simplified Chinese: 蓝田人; traditional Chinese: 藍田人; pinyin: Lántián rén), formerly Sinanthropus lantianensis (currently Homo erectus lantianensis) is a subspecies of Homo erectus. Its discovery in 1963 was first described by J. K. Woo the following year. The cranial capacity is estimated to be 780 cubic centimetres (48 cu in), somewhat similar to that of its contemporary, Java Man.

Remnants of Lantian Man were found in Lantian County, in China's Shaanxi province, approximately 50 km southeast of Xi'an. Shortly after the discovery of the mandible (jaw bone) of the first Lantian Man at Chenjiawo (陈家窝), also in Lantian, a cranium (skull) with nasal bones, right maxilla, and three teeth of another specimen of Lantian Man were found at Gongwangling (公王岭), another site in Lantian.

List of human evolution fossils

The following tables give an overview of notable finds of hominin fossils and remains relating to human evolution, beginning with the formation of the tribe Hominini (the divergence of the human and chimpanzee lineages) in the late Miocene, roughly 7 to 8 million years ago.

As there are thousands of fossils, mostly fragmentary, often consisting of single bones or isolated teeth with complete skulls and skeletons rare, this overview is not complete, but does show some of the most important finds. The fossils are arranged by approximate age as determined by radiometric dating and/or incremental dating and the species name represents current consensus; if there is no clear scientific consensus the other possible classifications are indicated.

Most of the early fossils shown are not considered direct ancestors to Homo sapiens but are closely related to direct ancestors and are therefore important to the study of the lineage. After 1.5 million years ago (extinction of Paranthropus), all fossils shown are human (genus Homo). After 11,500 years ago (11.5 ka, beginning of the Holocene), all fossils shown are Homo sapiens (anatomically modern humans), illustrating recent divergence in the formation of modern human sub-populations.

Meganthropus

Meganthropus is a series of large jaw and skull fragments found at the Sangiran site near Surakarta in Central Java, Indonesia. The original fossils were ascribed to a new species, Meganthropus palaeojavanicus, and while it is commonly considered invalid today, the genus name has survived as an informal name for the fossils.

As of 2005, the taxonomy and phylogeny for the specimens are still uncertain, although most paleoanthropologists consider them related to Homo erectus in some way. However, the names Homo palaeojavanicus and even Australopithecus palaeojavanicus are sometimes used as well, indicating the classification uncertainty.

After the discovery of a robust skull in Swartkrans in 1948 (SK48), the name Meganthropus africanus was briefly applied. However, that specimen is now formally known as Paranthropus robustus and the earlier name is a junior synonym.

Some of these finds were accompanied by evidence of tool use similar to that of Homo erectus. This is the reason Meganthropus is often linked with that species as H. e. palaeojavanicus.

Nanjing Man

Nanjing Man (Homo erectus nankinensis) is a subspecies of Homo erectus found in China. Large fragments of one male and one female skull and a molar tooth of H. e. nankinensis were discovered in 1993 in the Hulu cave on the Tangshan hills near Nanjing, the former capital city of China. The term Nanjing man is used to describe the subspecies of Homo erectus but is also used when referring to the three fossils. The specimens were found in the Hulu limestone cave at a depth of 60–97 cm by Liu Luhong, a local worker. Dating the fossils yielded an estimated age of 580,000 to 620,000 years old.

Peking Man

Peking Man (Chinese: 北京猿人; pinyin: Běijīng Yuánrén), Homo erectus pekinensis (formerly known by the junior synonym Sinanthropus pekinensis), is an example of Homo erectus. Discovered in 1923–27 during excavations at Zhoukoudian (Chou K'ou-tien) near Beijing (written "Peking" before the adoption of the Pinyin romanization system), China, in 2009 this group of fossil specimens dated from roughly 750,000 years ago, and a new 26Al/10Be dating suggests they are in the range of 680,000–780,000 years old.Between 1929 and 1937, 15 partial crania, 11 mandibles, many teeth, some skeletal bones and large numbers of stone tools were discovered in the Lower Cave at Locality 1 of the Peking Man site at Zhoukoudian. Their age is estimated to be between 500,000 and 300,000 years old. (A number of fossils of modern humans were also discovered in the Upper Cave at the same site in 1933.) The most complete fossils, all of which were portions of the skullcap (calvariae), are:

Skull II, discovered at Locus D in 1929 but only recognized in 1930, is an adult or adolescent with a brain size of 1030 cc.

Skull III, discovered at Locus E in 1929 is an adolescent or juvenile with a brain size of 915 cc.

Skulls X, XI and XII (sometimes called LI, LII and LIII) were discovered at Locus L in 1936. They are thought to belong to an adult man, an adult woman and a young adult, with brain sizes of 1225 cc, 1015 cc and 1030 cc respectively.

Skull V: two cranial fragments were discovered in 1966 which fit with (casts of) two other fragments found in 1934 and 1936 to form much of a skullcap with a brain size of 1140 cc. These pieces were found at a higher level, and appear to be more modern than the other skullcaps.Most of the study on these fossils was done by Davidson Black until his death in 1934. Pierre Teilhard de Chardin took over until Franz Weidenreich replaced him and studied the fossils until he left China in 1941. The original fossils disappeared in 1941, but excellent casts and descriptions remain.

Samu (fossil)

Samu is the nickname given to a fragmentary human occipital bone (also known as "Vertesszolos man", or "Vertesszolos occipital") found in Vértesszőlős, Central Transdanubia, Hungary.

The discovery was made on 21 August 1965 during a dig led by László Vértes, and the fossil was named Sámuel, 21 August being the name day of biblical judge Samuel in Hungarian tradition.

It has since become widely known as Samu, a Hungarian short form of the name.

Hungarian anthropologist Andor Thoma (1928–2003) initially described it as Homo erectus seu sapiens paleohungaricus. The fossil at the time was believed to be about 500,000 years old, and some literature of the 1970s classifies it as Homo erectus.

Analyses performed in the 1990s have revealed a significantly younger age, at between 250,000 and 300,000 years old (Mindel glaciation), and the fossil is now classified as Homo heidelbergensis.The Vértesszőlős archaeological site itself had been discovered by Márton Pécsi in 1962. Also found at the site were two child teeth, Abbevillian stone tools and a fireplace. The site is now open to the public.

A replica of the Samu occipital bone is on exhibit in the local museum (the original is kept in the Hungarian National Museum), as well as associated tools and fossilized animal footprints."Samu" has become a common name for plastic skeletons shown in biology classes in Hungarian student slang.

Solo Man

Solo Man (Homo erectus soloensis) is a subspecies of Homo erectus, identified based on fossil evidence discovered between 1931 and 1933 by Gustav Heinrich Ralph von Koenigswald from sites along the Solo River, on the Indonesian island of Java, dated to between 550,000 and 143,000 years old. The remains are also commonly referred to as Ngandong (now at Kradenan district, Blora Regency), after the village near where they were first recovered.

It is a late variant of H. erectus, dated to after 550,000 years ago, overlapping with Homo heidelbergensis and possibly with early Homo sapiens. Though its morphology was, for the most part, typical of Homo erectus, its cranial capacity of 1,013–1,251 cm³ places it amongst the larger-brained representatives of its species (compared to 900 cm³ for the older Java Man), and its culture was also unusually advanced.Due to the tools found with the fossils and many of their more gracile anatomical features, Solo Man was first classified as a subspecies of Homo sapiens (dubbed Homo sapiens soloensis) and long thought to be the ancestor of modern Australo-Melanesians. More rigorous studies in the 1990s have concluded that this is not the case. Analysis of 18 crania from Sangiran, Trinil, Sambungmacan, and Ngandong show chronological development from the Bapang-AG to Ngandong periods.

Tautavel

Tautavel (Catalan: Talteüll) is a commune in the Pyrénées-Orientales department in southern France.

It is home to The European Centre for Prehistoric Research (CERP). Tautavel Man, an early hominid and some of the oldest human remains in Europe, was found in Caune de l'Arago, a cave in the commune.

Tautavel Man

Tautavel Man (Homo erectus tautavelensis) is a proposed subspecies of Homo erectus, the type specimen being 450,000-year-old fossil remains discovered in the Arago Cave in Tautavel, France.

Excavations began in 1964, with the first notable discovery occurring in 1969.

Trinil

Trinil is a palaeoanthropological site on the banks of the Bengawan Solo River in Ngawi Regency, East Java Province, Indonesia. It was at this site in 1891 that the Dutch anatomist Eugène Dubois discovered the first early hominin remains to be found outside of Europe: the famous "Java Man" (Homo erectus erectus) specimen.

Yuanmou Man

Yuanmou Man (simplified Chinese: 元谋人; traditional Chinese: 元謀人; pinyin: Yuánmóu Rén), Homo erectus yuanmouensis, refers to a member of the genus Homo whose remnants, two incisors, were discovered near Danawu Village in Yuanmou County in southwestern province of Yunnan, China. Later, stone artifacts, pieces of animal bone showing signs of human work and ash from campfires were also dug up from the site. The fossils are on display at the National Museum of China, Beijing.

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