Homo erectus (meaning "upright man") is an extinct species of archaic
humans that lived throughout most of the
Pleistocene geological epoch.
Its earliest fossil evidence dates to 1.9 million years ago. It likely
East Africa and spread from there, beginning 1.8 million
years ago, migrating throughout Eurasia.
There is an ongoing debate regarding the classification, ancestry, and
Homo erectus, especially in relation to
Homo ergaster, 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
Homo neanderthalensis, and Homo
sapiens; or, 2) it is in fact an Asian species distinct from African
H. ergaster. There is also another view—an alternative to
1): 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.
H. erectus in the narrow sense (the Asian species) was extinct by
140,000 years ago.
A new debate appeared in 2013, with the documentation of the Dmanisi
skulls. Considering the large morphological variation among all
Dmanisi skulls, researchers now suggest that several early human
ancestors variously classified, for example, as
Homo ergaster, or Homo
rudolfensis, and perhaps even
Homo habilis, should instead be
2 Discovery and representative fossils
2.1 African genesis
Homo erectus georgicus
3 Classification and distinctions
3.1 Interpreting evolution: H. erectus /
H. ergaster / H. sapiens
4 Use of tools and fire
4.1 Use of fire
6 Descendants and subspecies
6.2 Related species
6.3 Previously referred taxa
7 Individual fossils
8 Development relative to other Hominins
10 See also
12 Further reading
13 External links
view • discuss • edit
Earliest stone tools
Earliest exit from Africa
Earliest fire use
Earliest in Europe
Axis scale: million years
Also see: Life timeline and Nature timeline
The first hypothesis of origin is that
Homo erectus rose from the
East Africa sometime during—or perhaps even
Early Pleistocene geological epoch, which itself dates to
2.58 million years ago (see below, at African genesis, re earlier date
Ledi-Geraru Research Area). From there it migrated, in part, by 2.0
M.Y.A., probably as a result of broad desertifying conditions
developing then in eastern and northern Africa; it joined the
migrations through the "Saharan pump" and dispersed around much of the
Old World. The fossil record shows that its development from about 1.8
M.Y.A. to one M.Y.A. was widely distributed: in Africa (Lake Turkana
 and Olduvai Gorge), the Transcaucasus (
Dmanisi in Georgia),
Central Java and Trinil, East Java), and in
Zhoukoudian and Shaanxi), and India.
The second hypothesis is that H. erectus evolved in
Eurasia and then
migrated to Africa. They occupied the
Dmanisi site from 1.85 million
to 1.77 million years ago, which was about the same time or slightly
before their earliest evidence in Africa. There are several
proposed explanations of the dispersal of H. erectus
georgicus—including whether or not Africa is the source.
Discovery and representative fossils
The Dutch anatomist
Eugène Dubois was fascinated by Darwin's theory
of evolution especially as it applied to humanity. In 1886, he set out
for Asia—which then was the region accepted as the cradle of human
evolution despite Darwin's theory of African origin; see Haeckel
§ Research—to find a human ancestor. In 1891, his team
discovered 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 named the species Pithecanthropus erectus—from the
Greek πίθηκος, "ape", and ἄνθρωπος,
"man"—based on a skullcap (calotte) and a femur like that of Homo
Dubois' 1891 find was the first fossil of a Homo-species (or any
hominin species) found as result of a directed expedition and
search—and which was inspired by Darwin's radical theory that
humans, like all other species, evolved from ancestral species, see
human evolution. (The first found and recognized human fossil was the
accidental 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
the other apes.
Java Man is now classified as
Most of the spectacular discoveries of H. erectus next took place at
Zhoukoudian Project, now known as the
Peking Man Site, in
Zhoukoudian, China. This site was first discovered by Johan Gunnar
Andersson in 1921 and was first excavated in 1921, and produced
two human teeth. Canadian anatomist Davidson Black's initial
description (1921) of a lower molar as belonging to a previously
unknown species (which he named Sinanthropus pekinensis) prompted
widely publicized interest. Extensive excavations followed, which
altogether uncovered 200 human fossils from more than 40 individuals
including five nearly complete skullcaps. German anatomist Franz
Weidenreich provided much of the detailed description of this material
in several monographs published in the journal Palaeontologica Sinica
Nearly all of the original specimens were lost during World War II;
however, authentic casts were made by Weidenreich which exist at the
American Museum of Natural History
American Museum of Natural History in
New York City
New York City and at the
Institute of Vertebrate Paleontology and Paleoanthropology in Beijing,
and are considered to be reliable evidence.
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, it was widely accepted that
modern humans first evolved in Asia. 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
From the 1950s forward, numerous finds in
East Africa confirmed the
hypothesis of an African genesis, providing fossil evidence that the
earliest hominins originated there. It is now generally accepted that
H. erectus descended from either: 1) the earliest hominin genera (such
as Australopithecus, and possibly Ardipithecus—of which is still
debated whether it is hominin or hominid); or 2) the earliest
Homo-species (such as
Homo habilis or
Homo ergaster). East Africa
provided sympatric coexistence for H. erectus and
H. habilis for
several hundred-thousand years, 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
habilis—or of a common ancestor of habilis and erectus—became
reproductively isolated from the main-group population, eventually
evolving into the new species
Homo erectus, Indian Museum
In the 1950s, archaeologists
John T. Robinson
John T. Robinson and
Robert Broom named
Telanthropus capensis; 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.
Yves Coppens discovered a skull of Tchadanthropus uxoris,
then the earliest fossil human discovered in north Africa. 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". Although at first considered to be a specimen of H.
habilis, T. uxoris is no longer considered a valid taxon, and has
been subsumed into H. erectus.
In 2013, a fragment of fossilized jawbone, dated to around 2.8 million
years ago, was discovered in the
Ledi-Geraru Research Area in the Afar
depression, Ethiopia. The fossil is considered the earliest
evidence of the
Homo genus known to date, and seems to be intermediate
Australopithecus and H. habilis. The individual lived just
after a major climate shift in the region, when forests and waterways
were rapidly replaced by arid savannah, which was a domain favored by
the early hominins.
Homo erectus georgicus
Dmanisi skull 3, Fossils skull
D2700 and D2735 jaw, two of several
Dmanisi in the Georgian Caucasus.
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.
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. The fossils are about 1.8 million years old.
After their initial assessment, some scientists were persuaded to name
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
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. 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 M.Y.A. The assemblage includes one of the largest
Homo mandibles (D2600), one of the smallest Lower
Pleistocene mandibles (D211), a nearly complete sub-adult (D2735), and
a toothless specimen D3444/D3900.
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
Hominina skulls from the
Pleistocene period. The
variation in these skulls were compared to variations in modern humans
and within a sample group of chimpanzees. The researchers found that,
despite appearances, the variations in the
Dmanisi skulls were no
greater than those seen among modern people and among chimpanzees.
These findings suggest that previous fossil finds that were classified
as different species on the basis of the large morphological variation
Homo gautengensis, H.
ergaster, and potentially even H. habilis—should perhaps be
re-classified to the same lineage as
Classification and distinctions
Dmanisi discovery, Georgia
Paleoanthropologists continue to debate the classification of Homo
Homo ergaster as separate species. One school of thought
suggests dropping the taxon
Homo erectus and equating H. erectus with
the archaic H. sapiens. Another 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. 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".
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
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.
Finding an extant (i.e., living) model suitable for field study,
analysis, and comparison is very important; and selecting a living
sample population of an appropriate species can be difficult. (For
example, the morphological variation among the global population of H.
sapiens is small, so our own species diversity may not be a
trustworthy comparison. Fossils found in Dmanisi, Georgia were
originally designated as a separate (but closely related) species; but
subsequent specimens showed their variation to be within the range of
Homo erectus. and they are now classified as
Homo erectus georgicus.)
New foot tracks found in 2009 in
Kenya and reported in Science by
Matthew Bennett of
Bournemouth University in Britain and his
colleagues, confirmed that the gait of
Homo erectus was heel-to-toe,
walking as a modern human does, rather than with the
australopithecine-like method of its own ancestors.
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, 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)—only 17
percent of modern male humans are taller—and were
extraordinarily slender, with long arms and legs.
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
Kamoya Kimeu in 1984—one of the most complete hominin
skeletons ever discovered—has contributed greatly to the
Interpreting evolution: H. erectus /
H. ergaster / H. sapiens
Model of the evolution of several species of genus
Homo over the last
2 million years (vertical axis) based on Stringer (2012).
Stringer (2003, 2012) and Reed, et al. (2004) and others have produced
schematic graph-models for interpreting the evolution of
from earlier species of Homo, including
Homo erectus and/or Homo
ergaster, see graphs at right. Blue areas denote the existence of one
or more hominin species at a given time and place (that is, region).
These and other interpretations differ mainly in the taxonomy and
geographical distribution of species.
Stringer (see upper graph-model) depicts the presence of H. erectus as
dominating the temporal and geographic development of human evolution;
and as persisting broadly throughout Africa and
Eurasia for nearly 2
million years, eventually evolving into
H. heidelbergensis / H.
rhodesiensis, which in turn evolved into H. sapiens. Reed, et al.
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
Use of tools and fire
An alternate graph-model of the temporal and geographical distribution
Homo species, evolving over the last two million
years ; proposed by Reed, et al., redrawn from Stringer. Note
the depiction of
Homo ergaster as an ancestor of
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; thus it closely coincides with the
Pleistocene epoch of
geologic time, which is 2.58 million to 11,700 years ago. 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 made hand axes out of
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. Because the use of
began ca. 1.8 million years ago, 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. And the oldest stone tool found in
Turkey reveals that
hominins passed through the Anatolian gateway from western
Europe approximately 1.2 million years ago—much earlier than
Use of fire
East African sites, such as Chesowanja near Lake Baringo, Koobi Fora,
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.. 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). At a
"hearth-like depression" at a site in Olorgesailie, Kenya, some
microscopic charcoal was found—but that could have resulted from
natural brush fires.
In Gadeb, Ethiopia, fragments of welded tuff that appeared to have
been burned, or scorched, were found alongside H. erectus–created
Acheulean artifacts; but such re-firing of the rocks may have been
caused by local volcanic activity. In the
Middle Awash River
Valley, cone-shaped depressions of reddish clay were found that could
have been created only by temperatures of 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. 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 B.P.; 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. 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
Cooking § History
There is archaeological evidence that
Homo erectus cooked their
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,
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. 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. Both brain size and the presence of the
Broca's area also
support the use of articulate language.
H. erectus was probably the first hominin to live in small, familiar
band-societies similar to modern hunter-gatherer band-societies,
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
Daniel Everett has argued that H. erectus may have been the
first hominin to evolve the capability of language. He argues that
their level of social organization and technical sophistication must
have required a complex communication system. 
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
Homo sapiens have existed for about 200,000 years. If
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 hominins (see above,
"Interpreting evolution: ...").
A model of the face of an adult male
A model of the face of an adult female
Homo erectus erectus (
Homo erectus yuanmouensis (Yuanmou Man)
Homo erectus lantianensis (Lantian Man)
Homo erectus nankinensis (Nanjing Man)
Homo erectus pekinensis (Peking Man)
Homo erectus palaeojavanicus (Meganthropus)
Homo erectus soloensis (Solo Man)
Homo erectus tautavelensis (
Homo erectus georgicus
Homo erectus bilzingslebenensis
Human taxonomy § Species
On many archaic humans there is no definite consensus as to whether
they should be classified as subspecies of either H. erectus or H.
sapiens, or as separate species
African H. erectus candidates
Homo ergaster ("African H. erectus)
Homo naledi (or H. e. naledi)
Eurasian H. erectus candidates:
Homo antecessor (or H. e. antecessor)
Homo heidelbergensis (or H. e. heidelbergensis)
Homo cepranensis (or H. e. cepranensis)
Homo floresiensis (or H. e. florsiensis)
Homo sapiens candidates
Homo neanderthalensis (or H. s. neanderthalensis)
Homo rhodesiensis (or H. s. rhodensis)
Homo heidelbergensis (or H. s. heidelbergensis)
Homo sapiens idaltu
anatomically modern humans (
Homo sapiens or H. s. sapiens)
Previously referred taxa
Homo erectus wushanensis (actually a stem-orangutan)
Homo erectus soloensis, who was long assumed to have lived on
least as late as about 50,000 years ago but was re-dated in 2011 to a
much older age,. New 2017 research suggests that H. floresiensis
was descended from the same (presumably Australopithecine) ancestor as
Homo habilis, making it 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.
Some of the major
Homo erectus fossils:
Indonesia (island of Java):
Trinil 2 (holotype),
Sambungmachan collection, Ngandong collection
China ("Peking Man"): Lantian (Gongwangling and Chenjiawo), Yunxian,
Zhoukoudian, Nanjing, Hexian
Kenya: KNM ER 3883, KNM ER 3733
Vértesszőlős, Hungary "Samu"
Vietnam: Northern, Tham Khuyen, Hoa Binh
Republic of Georgia:
Dmanisi collection ("
Homo erectus georgicus")
Ethiopia: Daka calvaria
Eritrea: Buia cranium (possibly H. ergaster)
Denizli Province, Turkey: Kocabas fossil
Development relative to other Hominins
Hominin species during Pleistocene
Homo erectus tautavelensis skull.
Replica of lower jaws of
Homo erectus from Tautavel, France.
Sangiran II" original, collection Koenigswald, Senckenberg
A reconstruction based on evidence from the Daka Member, Ethiopia
Original fossils of Pithecanthropus erectus (now
Homo erectus) found
Java in 1891.
A Different Flesh, alternate history novel by
Harry Turtledove where
erectus survived to modern times
Dawn of Humanity (2015 PBS documentary)
List of fossil sites
List of fossil sites (with link directory)
List of human evolution fossils
List of human evolution fossils (with images)
Origins of Us (2011 BBC documentary)
Prehistoric Autopsy (2012 BBC documentary)
^ 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 143,000 years ago at the latest, more likely
before 550,000 years ago. Indriati E, Swisher CC III, Lepre C, Quinn
RL, Suriyanto RA, et al. 2011 The Age of the 20 Meter Solo River
Indonesia and the Survival of
Homo erectus in Asia.
PLoS ONE 6(6): e21562. doi:10.1371/journal.pone.0021562.
^ a b Hazarika, Manji (16–30 June 2007). "
Homo erectus/ergaster and
Out of Africa: Recent Developments in Paleoanthropology and
Prehistoric Archaeology" (PDF).
^ Chauhan, Parth R. (2003) "Distribution of Acheulian sites in the
Siwalik region" Archived 4 January 2012 at the Wayback Machine. in An
Overview of the Siwalik Acheulian & Reconsidering Its
Chronological Relationship with the Soanian – A Theoretical
^ See overview of theories on human evolution.
^ Klein, R. (1999). The
Human Biological and Cultural
Origins. Chicago: University of Chicago Press, ISBN 0226439631.
^ Antón, S. C. (2003). "Natural history of
Homo erectus". Am. J.
Phys. Anthropol. 122: 126–170. doi:10.1002/ajpa.10399. By the 1980s,
the growing numbers of H. erectus specimens, particularly in Africa,
led to the realization that Asian H. erectus (H. erectus sensu
stricto), once thought so primitive, was in fact more derived than its
African counterparts. These morphological differences were interpreted
by some as evidence that more than one species might be included in H.
erectus sensu lato (e.g., Stringer, 1984; Andrews, 1984; Tattersall,
1986; Wood, 1984, 1991a, b; Schwartz and Tattersall, 2000) ... Unlike
the European lineage, in my opinion, the taxonomic issues surrounding
Asian vs. African H. erectus are more intractable. The issue was most
pointedly addressed with the naming of
H. ergaster on the basis of the
type mandible KNM-ER 992, but also including the partial skeleton and
isolated teeth of KNM-ER 803 among other
Koobi Fora remains (Groves
and Mazak, 1975). Recently, this specific name was applied to most
early African and Georgian H. erectus in recognition of the
less-derived nature of these remains vis à vis conditions in Asian H.
erectus (see Wood, 1991a, p. 268; Gabunia et al., 2000a). At least
portions of the paratype of
H. ergaster (e.g., KNM-ER 1805) are not
included in most current conceptions of that taxon. The H. ergaster
question remains famously unresolved (e.g., Stringer, 1984;
Tattersall, 1986; Wood, 1991a, 1994; Rightmire, 1998b; Gabunia et al.,
2000a; Schwartz and Tattersall, 2000), in no small part because the
original diagnosis provided no comparison with the Asian fossil
^ Suwa G, Asfaw B, Haile-Selassie Y, White T, Katoh S, WoldeGabriel G,
Hart W, Nakaya H, Beyene Y (2007). "
fossils from Konso, southern Ethiopia". Anthropological Science. 115
(2): 133–151. doi:10.1537/ase.061203.
^ Skull suggests three early human species were one : Nature News
^ a b David Lordkipanidze, Marcia S. Ponce de Leòn, Ann
Margvelashvili, Yoel Rak, G. Philip Rightmire, Abesalom Vekua,
Christoph P. E. Zollikofer (18 October 2013). "A Complete Skull from
Dmanisi, Georgia, and the Evolutionary Biology of Early Homo".
Science. 342 (6156): 326–331. doi:10.1126/science.1238484. CS1
maint: Multiple names: authors list (link)
^ Switek, Brian (17 October 2013). "Beautiful Skull Spurs Debate on
Human History". National Geographic. Retrieved 22 September
^ Frazier, Kendrick (Nov–Dec 2006). "Leakey Fights Church Campaign
Human Fossils". Skeptical Inquirer. 30
(6). Archived from the original on 2009-01-10.
^ Prins, Harald E. L.; Walrath, Dana; McBride, Bunny (2007). Evolution
and prehistory: the human challenge. Wadsworth Publishing.
p. 162. ISBN 978-0-495-38190-7.
^ a b Ferring, R.; Oms, O.; Agusti, J.; Berna, F.; Nioradze, M.;
Shelia, T.; Tappen, M.; Vekua, A.; Zhvania, D.; Lordkipanidze, D.
(2011). "Earliest human occupations at
Dmanisi (Georgian Caucasus)
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