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''Homo erectus'' (meaning "
upright Body relative directions (also known as egocentric coordinates) are geometrical orientations relative to a body such as a human Humans (''Homo sapiens'') are the most abundant and widespread species In biology Biology is t ...
man") is an extinct
species In biology, a species is the basic unit of biological classification, classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individu ...

species
of
archaic human A number of varieties of ''Homo ''Homo'' () is the that emerged in the (otherwise extinct) genus ' that encompasses the extant species ' (), plus several extinct species classified as either to or closely related to modern humans (dependin ...
from the
Pleistocene The Pleistocene ( , often referred to as the ''Ice Age'') is the geological Epoch (geology), epoch that lasted from about 2,580,000 to 11,700 years ago, spanning the earth’s most recent period of repeated glaciations. Before a change finally ...
, with its earliest occurrence about 2 million years ago, and its specimens are among the first recognizable members of the genus ''
Homo ''Homo'' () is the that emerged in the (otherwise extinct) genus ' that encompasses the extant species ' (), plus several extinct species classified as either to or closely related to modern humans (depending on the species), most notably ' ...

Homo
''. ''H. erectus'' was the first human ancestor to spread throughout Eurasia, with a continental range extending from the
Iberian Peninsula The Iberian Peninsula , ** * Aragonese language, Aragonese and Occitan language, Occitan: ''Peninsula Iberica'' ** ** * french: Péninsule Ibérique * mwl, Península Eibérica * eu, Iberiar penintsula also known as Iberia, is a peni ...

Iberian Peninsula
to
Java Java ( id, Jawa, ; jv, ꦗꦮ; su, ) is one of the Greater Sunda Islands in Indonesia. It is bordered by the Indian Ocean to the south and the Java Sea to the north. With a population of 147.7 million people, Java is the world's List of ...

Java
. African populations of ''H. erectus'' are likely to be the ancestors to several human species, such as '' H. heidelbergensis'' and '' H. antecessor'', with the former generally considered to have been the ancestor to
Neanderthal Neanderthals (, also Neandertals, ''Homo neanderthalensis'' or ''Homo sapiens neanderthalensis'') are an extinct species In biology Biology is the natural science that studies life and living organisms, including their anatomy, phys ...
s,
Denisovan The Denisovans or Denisova hominins ) are an extinct species or subspecies of archaic human that ranged across Asia during the Lower Paleolithic, Lower and Middle Paleolithic. Denisovans are known from few remains, and, consequently, most of wha ...
s, and
modern humans Early modern human (EMH) or anatomically modern human (AMH) are terms used to distinguish ''Homo sapiens Humans (''Homo sapiens'') are the most abundant and widespread of , characterized by and large, complex brains. This has enabled ...
. Asian populations of ''H. erectus'' may be ancestral to '' H. floresiensis'' and possibly to '' H. luzonensis''. As a
chronospecies A chronospecies is a species In biology, a species is the basic unit of biological classification, classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organ ...

chronospecies
, the time of the disappearance of ''H. erectus'' is a matter of contention. There are also several proposed subspecies with varying levels of recognition. The last known population of ''H. erectus'' is '' H. e. soloensis'' from Java, around 117,000–108,000 years ago. ''H. erectus'' had a humanlike gait and body proportions, and was the first human species to have exhibited a flat face, prominent nose, and possibly sparse body hair coverage. Though brain size certainly exceeds that of ancestor species, capacity varied widely depending on the population. In older populations, brain development seemed to cease early in childhood, suggesting that offspring were largely self-sufficient at birth, thus limiting cognitive development through life. Nonetheless, sites generally show consumption of medium to large animals, such as
bovine The biological subfamily In biological classification In biology, taxonomy () is the scientific study of naming, defining (Circumscription (taxonomy), circumscribing) and classifying groups of biological organisms based on shared characte ...

bovine
s or
elephant Elephants are the largest existing land animals. Three living species In biology, a species is the basic unit of biological classification, classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A specie ...

elephant
s, and suggest the development of predatory behaviour and coordinated hunting. ''H. erectus'' is associated with the
Acheulean . The types shown are (clockwise from top) cordate, ficron, and ovate. in Nice, France as postulated by Henry de Lumley dated to 400 thousand years ago. Acheulean (; also Acheulian and Mode II), from the French ''acheuléen'' after the ty ...

Acheulean
stone tool
industry Industry may refer to: Economics * Industry (economics) In macroeconomics, an industry is a branch of an economy that produces a closely related set of raw materials, goods, or services. For example, one might refer to the wood industry ...
, and is postulated to have been the earliest human ancestor capable of using fire, hunting and gathering in coordinated groups, caring for injured or sick group members, and possibly seafaring and art (though examples of art are controversial, and are otherwise rudimentary and few and far between). ''H. erectus'' men and women may have been roughly the same size as each other (i.e. exhibited reduced
sexual dimorphism Sexual dimorphism is the condition where the sex Sex is either of two divisions, typically male Male (♂) is the sex of an organism that produces the gamete known as sperm. A male gamete can fuse with a larger female gamete, or ovum, in ...
) like modern humans, which could indicate
monogamy Monogamy ( ) is a form of dyadic relationship Relationship most often refers to: * Interpersonal relationship The concept of interpersonal relationship involves social associations, connections, or affiliations between two or more peo ...
in line with general trends exhibited in primates. Size, nonetheless, ranged widely from in height and in weight. It is unclear if ''H. erectus'' was anatomically capable of speech, though it is postulated they communicated using some
proto-language In the tree model In historical linguistics Historical linguistics, also termed diachronic linguistics, is the scientific study of language change over time. Principal concerns of historical linguistics include: # to describe and accoun ...
.


Taxonomy


Naming

Despite what English naturalist
Charles Darwin Charles Robert Darwin (; ; 12 February 1809 – 19 April 1882) was an English natural history#Before 1900, naturalist, geologist and biologist, best known for his contributions to the science of evolution. His proposition that all sp ...

Charles Darwin
had hypothesised in his 1871 book '' Descent of Man'', many late-19th century evolutionary naturalists postulated that Asia, not Africa, was the birthplace of humankind as it is midway between Europe and America, providing optimal dispersal routes throughout the world (the Out of Asia theory). Among these was German naturalist
Ernst Haeckel Ernst Heinrich Philipp August Haeckel (; 16 February 1834 – 9 August 1919) was a German zoologist Zoology ()The pronunciation of zoology as is typically regarded as nonstandard, though it is not uncommon. is the branch of biology that stu ...

Ernst Haeckel
who argued that the first human species evolved on the now-disproven hypothetical continent "
Lemuria
Lemuria
" in what is now Southeast Asia, from a species he termed "'' Pithecanthropus alalus''" ("speechless apeman"). "Lemuria" had supposedly sunk below the
Indian Ocean The Indian Ocean is the third-largest of the world's five ocean The ocean (also the or the world ocean) is the body of that covers approximately 70.8% of the surface of and contains 97% of . Another definition is "any of the large ...

Indian Ocean
, so no fossils could be found to prove this. Nevertheless, Haeckel's model inspired Dutch scientist
Eugène Dubois Marie Eugène François Thomas Dubois (; 28 January 1858 – 16 December 1940) was a Dutch Dutch commonly refers to: * Something of, from, or related to the Netherlands * Dutch people () * Dutch language () *Dutch language , spoken in Belgium (al ...
to journey to the
Dutch East Indies The Dutch East Indies (or Netherlands East-Indies; nl, Nederlands(ch)-Indië; ) was a Dutch colony The Dutch colonial empire ( nl, Nederlandse koloniale rijk) comprised the overseas territories and trading posts controlled and administer ...
. Because no directed expedition had ever discovered human fossils (the few known had all been discovered by accident), and the economy was strained by the
Long Depression The Long Depression was a worldwide price and economic recession In economics, a recession is a business cycle contraction when there is a general decline in economic activity. Recessions generally occur when there is a widespread drop in spend ...
, the Dutch government refused to fund Dubois. In 1887, he enlisted in the Dutch East India Army as a medical officer, and was able to secure a post in 1887 in the Indies to search for his " missing link" in his spare time. On
Java Java ( id, Jawa, ; jv, ꦗꦮ; su, ) is one of the Greater Sunda Islands in Indonesia. It is bordered by the Indian Ocean to the south and the Java Sea to the north. With a population of 147.7 million people, Java is the world's List of ...

Java
, he found a skullcap in 1891 and a
femur The femur (; ), or thigh bone, is the proximal Standard anatomical terms of location deal unambiguously with the anatomy of animals, including humans. Terms used generally derive from Latin or Greek language, Greek roots and used to describe s ...

femur
in 1892 (
Java Man Java Man (''Homo erectus erectus'', formerly also ''Anthropopithecus erectus'', ''Pithecanthropus erectus'') is an early human fossil discovered in 1891 and 1892 on the island of Java (Dutch East Indies, now part of Indonesia). Estimated to be b ...

Java Man
) dating to the
late Pliocene Late may refer to: * LATE, an acronym which could stand for: ** Limbic-predominant age-related TDP-43 encephalopathy Limbic-predominant age-related TDP-43 encephalopathy (LATE) is a term that describes a prevalent condition with impaired memory ...
or
early Pleistocene The Early Pleistocene is an unofficial sub-epoch in the international geologic timescale The geologic time scale (GTS) is a system of chronological datingChronological dating, or simply dating, is the process of attributing to an object or ev ...
at the
Trinil upright=1.45, The '' Homo Erectus'' "Java Man" in the Naturalis Biodiversity Center">Java_Man.html" ;"title="Homo Erectus'' "Java Man">Homo Erectus'' "Java Man" in the Naturalis Biodiversity Center, Netherlands. Trinil is a paleoanthropology, pal ...
site along the
Solo River The Solo River (known in Indonesian as Bengawan Solo, with ''Bengawan'' being an Old Javanese word for ''river A river is a natural flowing watercourse, usually freshwater, flowing towards an ocean, sea, lake or another river. In some cases ...
, which he named "''P.''" ''erectus'' ("upright apeman") in 1893. He attempted unsuccessfully to convince the European scientific community that he had found an upright-walking ape-man. Given few fossils of ancient humans had even been discovered at the time, they largely dismissed his findings as a malformed non-human ape. The significance of these fossils would not be realized until the 1927 discovery of what Canadian palaeoanthropologist Davidson Black called "''Sinanthropus pekinensis''" (Peking Man) at the
Zhoukoudian Zhoukoudian Peking Man Site (), also romanized as Choukoutien, is a cave system in suburban Fangshan District, Beijing Beijing ( ), Chinese postal romanization, alternatively romanized as Peking ( ), is the Capital city, capital of the ...
cave near
Beijing Beijing ( ), as Peking ( ), is the of the . It is the world's , with over 21 million residents within an of 16,410.5 km2 (6336 sq. mi.). It is located in , and is governed as a under the direct administration of the with .Figures ...

Beijing
, China. Black lobbied across North America and Europe for funding to continue excavating the site, which has since become the most productive ''H. erectus'' site in the world. Continued interest in Java led to further ''H. erectus'' fossil discoveries at Ngandong (
Solo Man Solo Man (''Homo erectus soloensis'') is a subspecies In Taxonomy (biology), biological classification, the term subspecies refers to one of two or more populations of a species living in different subdivisions of the species' range and vary ...
) in 1931,
Mojokerto Mojokerto ( jv, ꦩꦗꦏꦼꦂꦠ (''Majakerta'')) is a city in East Java East Java ( id, Jawa Timur) is a Provinces of Indonesia, province of Indonesia. It has a land border only with the province of Central Java to the west; the Java Sea an ...
(Java Man) in 1936, and
Sangiran Sangiran is an archaeological excavation site in Java in Indonesia Indonesia ( ), officially the Republic of Indonesia ( id, Republik Indonesia, links=yes ), is a country in Southeast Asia and Oceania between the Indian Ocean, Indian and ...
(Java Man) in 1937. The Sangiran site yielded the best preserved Java Man skull. German paleoanthropologist Franz Weidenreich provided much of the detailed description of this material in several monographs. The original specimens were lost during the
Second Sino-Japanese War The Second Sino-Japanese War (1937–1945) was a military conflict that was primarily waged between the Republic of China (1912–1949), Republic of China and the Empire of Japan. The war made up the Chinese theater of the wider Pacific War, Pac ...
after an attempt to smuggle them out of China for safekeeping. Only casts remain. Similarities between Java Man and Peking Man led
Ernst Mayr Ernst Walter Mayr (; 5 July 1904 – 3 February 2005) was one of the 20th century's leading evolutionary biologists File:Francesco Redi.jpg, Francesco Redi, the founder of biology, is recognized to be one of the greatest biologists of all ti ...
to rename both as ''Homo erectus'' in 1950. Throughout much of the 20th century, anthropologists debated the role of ''H. erectus'' in
human evolution Human evolution is the evolution Evolution is change in the heritable Heredity, also called inheritance or biological inheritance, is the passing on of Phenotypic trait, traits from parents to their offspring; either through asexual ...

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 Charles Robert Darwin (; ; 12 February 1809 – 19 April 1882) was an English natural history#Before 1900, naturalist, geologist and biologist, best known for his contributions to the science of evolution. His proposition that all sp ...

Charles Darwin
the 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. In 1949, the species was reported in Swartkraans Cave, South Africa, by South African paleoanthropologists
Robert Broom Robert Broom FRS FRSE (30 November 1866 6 April 1951) was a South Africa South Africa, officially the Republic of South Africa (RSA), is the southernmost country in Africa. With over Demographics of South Africa, 59 million people, ...
and
John Talbot Robinson John Talbot Robinson FRSSAf The Royal Society of South Africa is a learned society composed of eminent South African scientists and academics. The Society was granted its royal charter by King Edward VII in 1908, nearly a century after Capeton ...
, who described it as "''Telanthropus capensis''". ''Homo'' fossils have also been reported from nearby caves, but their species designation has been a tumultuous discussion. A few North African sites have additionally yielded ''H. erectus'' remains, which at first were classified as "''Atlantanthropus mauritanicus''" in 1951. Beginning in the 1970s, propelled most notably by
Richard Leakey Richard Erskine Frere Leakey (19 December 1944 – 2 January 2022) was a Kenyan paleoanthropologist Paleoanthropology or paleo-anthropology is a branch of paleontology Paleontology, also spelled palaeontology or palæontology (), is the sc ...

Richard Leakey
, more were being unearthed in East Africa predominantly at the
Koobi Fora Koobi Fora refers primarily to a region around Koobi Fora Ridge, located on the eastern shore of Lake Turkana Lake Turkana (), formerly known as Lake Rudolf, is a lake in the Kenyan Rift Valley, in northern Kenya, with its far northern en ...
site, Kenya, and
Olduvai Gorge The Olduvai Gorge or Oldupai Gorge in Tanzania is one of the most important paleoanthropology, paleoanthropological sites in the world; it has proven invaluable in furthering understanding of early human evolution. A steep-sided ravine in the Greg ...

Olduvai Gorge
, Tanzania. Archaic human fossils unearthed across Europe used to be assigned to ''H. erectus'', but have since been separated as ''
H. heidelbergensis
H. heidelbergensis
'' as a result of namely British physical anthropologist
Chris Stringer Christopher Brian Stringer (born 1947) is a British physical anthropologist noted for his work on human evolution. Biography Growing up in a working-class family in the East End of London The East End of London, often referred to withi ...

Chris Stringer
's work.


Evolution

It has been proposed that ''H. erectus'' evolved from ''
H. habilis
H. habilis
'' about 2 Mya, though this has been called into question because they coexisted for at least a half a million years. Alternatively, a group of ''H. habilis'' may have been
reproductively isolated The mechanisms of reproductive isolation are a collection of evolutionary mechanisms, ethology, behaviors and physiology, physiological processes critical for speciation. They prevent members of different species from producing offspring, or ensur ...
, and only this group developed into ''H. erectus'' (
cladogenesis Cladogenesis is an evolutionary splitting of a parent species In biology, a species is the basic unit of biological classification, classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defi ...
). Because the earliest remains of ''H. erectus'' are found in both Africa and East Asia (in China as early as 2.1 Mya, in South Africa 2.04 Mya), it is debated where ''H. erectus'' evolved. A 2011 study suggested that it was ''H. habilis'' who reached West Asia from Africa, that early ''H. erectus'' developed there, and that early ''H. erectus'' would then have dispersed from West Asia to East Asia (
Peking Man Peking Man (''Homo erectus pekinensis'') is a subspecies In biological classification In biology Biology is the natural science that studies life and living organisms, including their anatomy, physical structure, Biochemis ...
), Southeast Asia (
Java Man Java Man (''Homo erectus erectus'', formerly also ''Anthropopithecus erectus'', ''Pithecanthropus erectus'') is an early human fossil discovered in 1891 and 1892 on the island of Java (Dutch East Indies, now part of Indonesia). Estimated to be b ...

Java Man
), back to Africa (''
Homo ergaster ''Homo ergaster'' is an extinct species In biology Biology is the natural science that studies life and living organisms, including their anatomy, physical structure, Biochemistry, chemical processes, Molecular biology, molecular int ...

Homo ergaster
''), and to Europe ( Tautavel Man), eventually evolving into modern humans in Africa. Others have suggested that ''H. erectus''/''H. ergaster'' developed in Africa, where it eventually evolved into modern humans. ''H. erectus'' had reached
Sangiran Sangiran is an archaeological excavation site in Java in Indonesia Indonesia ( ), officially the Republic of Indonesia ( id, Republik Indonesia, links=yes ), is a country in Southeast Asia and Oceania between the Indian Ocean, Indian and ...
, Java, by 1.6 Mya, and a second and distinct wave of ''H. erectus'' had colonized
Zhoukoudian Zhoukoudian Peking Man Site (), also romanized as Choukoutien, is a cave system in suburban Fangshan District, Beijing Beijing ( ), Chinese postal romanization, alternatively romanized as Peking ( ), is the Capital city, capital of the ...
, China, about 780 kya. Early teeth from Sangiran are bigger and more similar to those of basal (ancestral) Western ''H. erectus'' and ''H. habilis'' than to those of the derived Zhoukoudian ''H. erectus''. However, later Sangiran teeth seem to reduce in size, which could indicate a secondary colonization event of Java by the Zhoukoudian or some closely related population.


Subspecies

* '' Homo erectus erectus'' (
Java Man Java Man (''Homo erectus erectus'', formerly also ''Anthropopithecus erectus'', ''Pithecanthropus erectus'') is an early human fossil discovered in 1891 and 1892 on the island of Java (Dutch East Indies, now part of Indonesia). Estimated to be b ...

Java Man
, 1.6–0.5 Ma) * '''' (1.9–1.4 Ma) * '' Homo erectus georgicus'' (1.8–1.6 Ma) * '' Homo erectus lantianensis'' (
Lantian Man Lantian Man (), ''Homo erectus lantianensis'') is a subspecies of ''Homo erectus ''Homo erectus'' (meaning " upright man") is an extinct species In biology, a species is the basic unit of biological classification, classification and ...
, 1.6 Ma) * '' Homo erectus nankinensis'' ( Nanjing Man, 0.6 Ma) * ''
Homo erectus pekinensis Peking Man (''Homo erectus pekinensis'', formerly known by the junior synonym ''Sinanthropus pekinensis'') is a group of fossil specimens of ''Homo erectus'', dated from roughly 750,000 years ago, discovered in 1929–37 during excavations at Z ...

Homo erectus pekinensis
'' (
Peking Man Peking Man (''Homo erectus pekinensis'') is a subspecies In biological classification In biology Biology is the natural science that studies life and living organisms, including their anatomy, physical structure, Biochemis ...
, 0.7 Ma) * '' Homo erectus soloensis'' (
Solo Man Solo Man (''Homo erectus soloensis'') is a subspecies In Taxonomy (biology), biological classification, the term subspecies refers to one of two or more populations of a species living in different subdivisions of the species' range and vary ...
, 0.546–0.143 Ma) * '''' ( Tautavel Man, 0.45 Ma) * '' Homo erectus yuanmouensis'' ( Yuanmou Man) " Wushan Man" was proposed as ''Homo erectus wushanensis'', but is now thought to be based upon fossilized fragments of an extinct non-hominin ape. Since its discovery in 1893 (
Java man Java Man (''Homo erectus erectus'', formerly also ''Anthropopithecus erectus'', ''Pithecanthropus erectus'') is an early human fossil discovered in 1891 and 1892 on the island of Java (Dutch East Indies, now part of Indonesia). Estimated to be b ...

Java man
), 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 The Lower Paleolithic (or Lower Palaeolithic) is the earliest subdivision of the Paleolithic The Paleolithic or Palaeolithic or Palæolithic (), also called the Old Stone Age (from Greek wikt:παλαιός, palaios - old, lithos - stone) ...
) forms of ''Homo'' sufficiently derived from '''' and distinct from early '''' (in Africa also known as ''''). It is sometimes considered as a wide-ranging, polymorphous species. Due to such a wide range of variation, it has been suggested that the ancient '''' and '''' should be considered early varieties of ''H. erectus''. The primitive ''H. e. georgicus'' from
Dmanisi Dmanisi ( ka, დმანისი, tr, ) is a town and archaeological site in the Kvemo Kartli region of Georgia (country), Georgia approximately 93 km southwest of the nation’s capital Tbilisi in the river valley of Mashavera. The hominin ...
, Georgia has the smallest brain capacity of any known Pleistocene hominin (about 600 cc), and its inclusion in the species would greatly expand the range of variation of ''H. erectus'' to perhaps include species as ''H. rudolfensis'', '' H. gautengensis'', '''', and perhaps ''H. habilis''. However, a 2015 study suggested that ''H. georgicus'' represents an earlier, more primitive species of ''Homo'' derived from an older dispersal of hominins from Africa, with ''H. ergaster/erectus'' possibly deriving from a later dispersal. ''H. georgicus'' is sometimes not even regarded as ''H. erectus''. It is debated whether the African ''H. e. ergaster'' is a separate species (and that ''H. erectus'' evolved in Asia, then migrated to Africa), or is the African form (''
sensu lato ''Sensu'' is a Latin Latin (, or , ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoken in the area around Rome, known as Latium. Through the power of the Ro ...
'') of ''H. erectus (
sensu stricto ''Sensu'' is a Latin Latin (, or , ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoken in the area around Rome, known as Latium. Through the power of the Ro ...
)''. In the latter, ''H. ergaster'' has also been suggested to represent the immediate ancestor of ''H. erectus''. It has also been suggested that ''H. ergaster'' instead of ''H. erectus'', or some hybrid between the two, was the immediate ancestor of other archaic humans and modern humans. It has been proposed that Asian ''H. erectus'' have several unique characteristics from non-Asian populations (
autapomorphies In phylogenetics In biology Biology is the natural science that studies life and living organisms, including their anatomy, physical structure, Biochemistry, chemical processes, Molecular biology, molecular interactions, Physiology, physiol ...
), but there is no clear consensus on what these characteristics are or if they are indeed limited to only Asia. Based on supposed derived characteristics, the 120 ka Javan ''H. e. soloensis'' has been proposed to have speciated from ''H. erectus'', as ''H. soloensis'', but this has been challenged because most of the basic cranial features are maintained. In a wider sense, ''H. erectus'' had mostly been replaced by ''H. heidelbergensis'' by about 300 kya years ago, with possible late survival of '' H. erectus soloensis'' in Java an estimated 117-108kya.


Descendants and synonyms

''Homo erectus'' is the most long-lived species of ''Homo'', having survived for almost two million years. By contrast, ''
Homo sapiens Humans (''Homo sapiens'') are the most abundant and widespread species of primate, characterized by bipedality, bipedalism and large, complex brains. This has enabled the development of advanced tools, culture, and language. Humans are highl ...

Homo sapiens
'' emerged about a third of a million years ago. Regarding many
archaic humans A number of varieties of ''Homo ''Homo'' () is the genus Genus /ˈdʒiː.nəs/ (plural genera /ˈdʒen.ər.ə/) is a taxonomic rank In biological classification In biology, taxonomy () is the scientific study of naming, definin ...
, there is no definite consensus as to whether they should be classified as
subspecies In biological classification In biology Biology is the natural science that studies life and living organisms, including their anatomy, physical structure, Biochemistry, chemical processes, Molecular biology, molecular interacti ...
of ''H. erectus'' or ''H. sapiens'' or as separate species. * African ''H. erectus'' candidates ** ''
Homo ergaster ''Homo ergaster'' is an extinct species In biology Biology is the natural science that studies life and living organisms, including their anatomy, physical structure, Biochemistry, chemical processes, Molecular biology, molecular int ...

Homo ergaster
'' ("African ''H. erectus''") ** ''
Homo naledi '' Homo naledi'' is a species of Archaic humans, archaic human discovered in 2013 in the Rising Star Cave, Cradle of Humankind, South Africa dating to the Middle Pleistocene 335,000–236,000 years ago. The initial discovery comprises 1,550 spe ...

Homo naledi
'' (or ''H. e. naledi'') * Eurasian ''H. erectus'' candidates: ** ''
Homo antecessor ''Homo antecessor'' (Latin Latin (, or , ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoken in the area around Rome, known as Latium. Through the power of the ...

Homo antecessor
'' (or ''H. e. antecessor'') ** ''
Homo heidelbergensis ''Homo heidelbergensis'' (also ''H. sapiens heidelbergensis'') is an extinct species In biology, a species is the basic unit of biological classification, classification and a taxonomic rank of an organism, as well as a unit of biodiversit ...

Homo heidelbergensis
'' (or ''H. e. heidelbergensis'') ** '''' (or ''H. e. cepranensis'') * ''
Homo floresiensis ''Homo floresiensis'' ("Flores Man"; nicknamed "Hobbit") is a species of small archaic human A number of varieties of ''Homo ''Homo'' () is the that emerged in the (otherwise extinct) genus ' that encompasses the extant species ' (), p ...

Homo floresiensis
'' * ''
Homo sapiens Humans (''Homo sapiens'') are the most abundant and widespread species of primate, characterized by bipedality, bipedalism and large, complex brains. This has enabled the development of advanced tools, culture, and language. Humans are highl ...

Homo sapiens
'' candidates ** ''
Homo neanderthalensis Neanderthals (, also Neandertals, ''Homo neanderthalensis'' or ''Homo sapiens neanderthalensis'') are an extinct species In biology, a species is the basic unit of biological classification, classification and a taxonomic rank of an org ...
'' (or ''H. s. neanderthalensis'') **
Denisovan The Denisovans or Denisova hominins ) are an extinct species or subspecies of archaic human that ranged across Asia during the Lower Paleolithic, Lower and Middle Paleolithic. Denisovans are known from few remains, and, consequently, most of wha ...
s (or ''Homo'' sp. 'Denisova' ''H. sapiens'' ssp. 'Denisova' or ''H.'' sp. 'Altai') ** ''
Homo rhodesiensis ''Homo rhodesiensis'' is the species name proposed by Arthur Smith Woodward (1921) to classify Kabwe 1 (the "Kabwe skull" or "Broken Hill skull", also "Rhodesian Man"), a Middle Stone Age fossil recovered from a cave at Broken Hill, or Kabwe, No ...

Homo rhodesiensis
'' (or ''H. s. rhodesiensis'') ** ''
Homo heidelbergensis ''Homo heidelbergensis'' (also ''H. sapiens heidelbergensis'') is an extinct species In biology, a species is the basic unit of biological classification, classification and a taxonomic rank of an organism, as well as a unit of biodiversit ...

Homo heidelbergensis
'' (or ''H. s. heidelbergensis'') ** ''
Homo sapiens idaltu Herto Man refers to the 160,000- to 154,000-year-old human remains (''Homo sapiens Humans (''Homo sapiens'') are the most abundant and widespread species of primate, characterized by bipedality, bipedalism and large, complex brains. This ...

Homo sapiens idaltu
'' ** the Narmada fossil, discovered in 1982 in
Madhya Pradesh Madhya Pradesh (, ; meaning ''Central Province'') is a state in central India. Its capital city, capital is Bhopal, and the largest city is Indore, with Jabalpur, Ujjain, Gwalior, Satna being the other major cities. Madhya Pradesh is the List o ...

Madhya Pradesh
, India, was at first suggested as ''H. erectus'' (''Homo erectus narmadensis'') but later recognized as ''H. sapiens''. ''
Meganthropus ''Meganthropus'' is an extinct genus of non-hominin hominid ape, known from the Pleistocene of Indonesia. It is known from a series of large jaw and Hominid skull, skull fragments found at the Sangiran site near Surakarta in Central Java (island), ...
'', based on fossils found in Java, dated to between 1.4 and 0.9 Mya, was tentatively grouped with ''H. erectus'' in contrast to earlier interpretations of it as a giant species of early human although older literature has placed the fossils outside of ''Homo'' altogether. However, Zanolli et al. (2019) judged ''Meganthropus'' to be a distinct genus of extinct ape.


Anatomy


Head

''Homo erectus'' featured a flat face compared to earlier hominins; pronounced brow ridge; and a low, flat skull. The presence of
sagittal In anatomy, the sagittal plane (), or longitudinal plane, is an anatomical plane which divides the body into right and left parts. The plane may be in the center of the body and split it into two halves (mid-sagittal plane, mid-sagittal) or away ...

sagittal
,
frontal Front may refer to: Arts, entertainment, and media Films * The Front (1943 film), ''The Front'' (1943 film), a 1943 Soviet drama film * ''The Front'', 1976 film Music *The Front (band), an American rock band signed to Columbia Records and activ ...

frontal
, and
coronal Coronal may refer to: * a nuptial crown * anything relating to a Corona (disambiguation), corona * Coronal plane, an anatomical term of location * The Commonly used terms of relationship and comparison in dentistry, coronal direction on a tooth * Co ...

coronal
keels, which are small crests that run along these suture lines, has been proposed to be evidence of significant thickening of the skull, specifically the
cranial vault The cranial vault is the space in the skull The skull is a bone A bone is a Stiffness, rigid tissue (anatomy), tissue that constitutes part of the vertebrate skeleton in animals. Bones protect the various organs of the body, produce red blo ...
.
CT scan A CT scan or computed tomography scan (formerly known as computed axial tomography or CAT scan) is a medical image, imaging Scientific technique, technique used in radiology to obtain detailed internal images of the body noninvasively for Diagno ...

CT scan
analyses reveal this to not be the case. However, the
squamous part of occipital bone The squamous part of occipital bone, is situated above and behind the foramen magnum The foramen magnum ( la, great hole) is a large, oval-shaped opening in the occipital bone The occipital bone () is a neurocranium, cranial dermal bone and t ...
, particularly the , at the rear of the skull is notably thicker than that of modern humans, likely a
basal Basal or basilar is a term meaning ''base'', ''bottom'', or ''minimum''. Science * Basal (anatomy), an anatomical term of location for features associated with the base of an organism or structure * Basal (medicine), a minimal level that is neces ...
(ancestral) trait. The fossil record indicates that ''H. erectus'' was the first human species to have featured a projecting nose, which is generally thought to have evolved in response to breathing dry air in order to retain moisture. American psychologist Lucia Jacobs hypothesized that the projecting nose instead allowed for distinguishing the direction different smells come from (stereo olfaction) to facilitate navigation and long-distance migration. The average brain size of Asian ''H. erectus'' is about . However, markedly smaller specimens have been found in Dmanisi, Georgia (''H. e. georgicus'');
Koobi Fora Koobi Fora refers primarily to a region around Koobi Fora Ridge, located on the eastern shore of Lake Turkana Lake Turkana (), formerly known as Lake Rudolf, is a lake in the Kenyan Rift Valley, in northern Kenya, with its far northern en ...
and
Olorgesailie Olorgesailie is a geological formation in East Africa containing a group of Lower Paleolithic Archeology, archaeological sites. It is on the floor of the Eastern Rift Valley in southern Kenya, southwest of Nairobi along the road to Lake Magadi ...
, Kenya; and possibly
Gona Gona is a coastal village in Oro Province Oro Province, formerly (and officially still) Northern Province, is a coast The coast, also known as the coastline or seashore, is defined as the area where land meets the sea or ocean, or as a line th ...
, Ethiopia. Overall, ''H. erectus'' brain size varies from , which is greater than the range of variation seen in modern humans and chimps, though less than that of gorillas. Dentally, ''H. erectus'' have the thinnest enamel of any Plio–Pleistocene hominin. Enamel prevents the tooth from breaking from hard foods, but impedes shearing through tough foods. The of ''H. erectus'', and all early ''Homo'', are thicker than those of modern humans and all living apes. The mandibular body resists torsion from the
bite force Bite force quotient (BFQ) is the regression analysis, regression of the quotient of an animal bite, animal's bite force in Newton (unit), newtons divided by its body mass in kilograms. It does not take into account sharpness of teeth or other dif ...
or chewing, meaning their jaws could produce unusually powerful stresses while eating, but the practical application of this is unclear. Nonetheless, the mandibular bodies of ''H. erectus'' are somewhat thinner than those of early ''Homo''. The premolars and molars also have a higher frequency of pits than ''H. habilis'', suggesting ''H. erectus'' ate more brittle foods (which cause pitting). These all indicate that the ''H. erectus'' mouth was less capable of processing hard foods and more at shearing through tougher foods, thus reducing the variety of foods it could process, likely as a response to tool use.


Body

Like modern humans, ''H. erectus'' varied widely in size, ranging from in height and in weight, thought to be due to regional differences in climate, mortality rates, and nutrition. Like modern humans and unlike other great apes, there does not seem to have been a great size disparity between ''H. erectus'' men and women (size-specific
sexual dimorphism Sexual dimorphism is the condition where the sex Sex is either of two divisions, typically male Male (♂) is the sex of an organism that produces the gamete known as sperm. A male gamete can fuse with a larger female gamete, or ovum, in ...
), though there is not much fossil data regarding this. Brain size in two adults from Koobi Fora measured , and another significantly smaller adult measured , which could possibly indicate sexual dimorphism, though sex was undetermined. If ''H. erectus'' did not exhibit sexual dimorphism, then it is possible that they were the first in the human line to do so, though the fragmentary fossil record for earlier species makes this unclear. If yes, then there was a substantial and sudden increase in female height. ''H. erectus'' had about the same limb configurations and proportions as modern humans, implying humanlike locomotion. ''H. erectus'' tracks near
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 ...
, Kenya, also indicate a
human gait A gait is a pattern of Limb (anatomy), limb movements made during Animal locomotion, locomotion. Human gaits are the various ways in which a human can move, either naturally or as a result of specialized training. Human gait is defined as Bipeda ...
. A humanlike shoulder suggests an ability for high speed throwing.Roach, & Richmond. (2015). "Clavicle length, throwing performance and the reconstruction of the Homo erectus shoulder". ''Journal of Human Evolution'', 80(C), 107–113. It was once thought that Turkana boy had 6
lumbar vertebra The lumbar vertebrae are, in human anatomy, the five vertebrae In the vertebrate spinal column The vertebral column, also known as the backbone or spine, is part of the axial skeleton. The vertebral column is the defining characteristic ...
instead of the 5 seen in modern humans and 11 instead of 12
thoracic vertebra In vertebrates, thoracic vertebrae compose the middle segment of the vertebral column, between the cervical vertebrae and the lumbar vertebrae. In humans, there are twelve thoracic vertebra (anatomy), vertebrae and they are intermediate in size be ...

thoracic vertebra
e, but this has since been revised, and the specimen is now considered to have exhibited a humanlike curvature of the spine (
lordosis Lordosis is historically defined as an ''abnormal'' inward curvature of the lumbar spine. However, the terms ''lordosis'' and ''lordotic'' are also used to refer to the normal inward curvature of the lumbar and cervical vertebrae, cervical region ...

lordosis
) and the same number of respective vertebrae. It is largely unclear when human ancestors lost most of their body hair. Genetic analysis suggests that high activity in the
melanocortin 1 receptor The melanocortin 1 receptor (MC1R), also known as melanocyte-stimulating hormone receptor (MSHR), melanin-activating peptide receptor, or melanotropin receptor, is a G protein–coupled receptor that binds to a class of pituitary peptide hormones ...
, which would produce dark skin, dates back to 1.2 Mya. This could indicate the evolution of hairlessness around this time, as a lack of body hair would have left the skin exposed to harmful
UV radiation Ultraviolet (UV) is a form of electromagnetic radiation In physics Physics (from grc, φυσική (ἐπιστήμη), physikḗ (epistḗmē), knowledge of nature, from ''phýsis'' 'nature'), , is the natural science that stud ...
. It is possible that exposed skin only became maladaptive in the Pleistocene, because the increasing
tilt Tilt may refer to: Music * Tilt (American band), a punk rock group, formed in 1992 * Tilt (British band), an electronic music group, formed in 1993 * Tilt (Polish band), a rock band, formed in 1979 Albums * Tilt (Cozy Powell album), ''Tilt'' (Coz ...
of the Earth (which also caused the
ice ages#REDIRECT Ice age {{Redirect category shell, 1= {{R from move {{R from ambiguous term {{R from other capitalisation {{R unprintworthy ...
) would have increased solar radiation bombardment- which would suggest that hairlessness first emerged in the australopithecines. However, australopithecines seem to have lived at much higher, much colder elevations—typically where the nighttime temperature can drop to —so they may have required hair to stay warm, unlike early ''Homo'' which inhabited lower, hotter elevations. Populations in higher latitudes potentially developed lighter skin to prevent
vitamin D deficiency Vitamin D deficiency, or hypovitaminosis D is defined as a vitamin D level that is below normal. It most commonly occurs in people when they have inadequate sunlight exposure (in particular sunlight with adequate ultraviolet B rays (UVB)). Vita ...
. A 500–300 ka ''H. erectus'' specimen from Turkey was diagnosed with the earliest known case of
tuberculous meningitis Tuberculous meningitis, also known as TB meningitis or tubercular meningitis, is a specific type of bacterial meningitis Meningitis is an Acute (medical), acute inflammation of the protective membranes covering the brain and spinal cord, known c ...
, which is typically exacerbated in dark-skinned people living in higher latitudes due to vitamin D deficiency. Hairlessness is generally thought to have facilitated sweating, but reduction of parasite load and
sexual selection Sexual selection is a mode of natural selection Natural selection is the differential survival and reproduction of individuals due to differences in phenotype right , Here the relation between genotype and phenotype is illustrat ...
have also been proposed.


Metabolism

The 1.8 Ma Mojokerto child specimen from Java, who died at about 1 year of age, presented 72–84% of the average adult brain size, which is more similar to the faster brain growth trajectory of great apes than modern humans. This indicates that ''H. erectus'' was probably not cognitively comparable to modern humans, and that secondary altriciality—an extended childhood and long period of dependency due to the great amount of time required for brain maturation—evolved much later in human evolution, perhaps in the modern human/Neanderthal last common ancestor. It was previously believed that, based on the narrow pelvis of Turkana boy, ''H. erectus'' could only safely deliver a baby with a brain volume of about , equating to a similar brain growth rate as modern humans to achieve the average adult brain size of . However, a 1.8 Ma female pelvis from Gona, Ethiopia, shows that ''H. erectus'' babies with a brain volume of could have been safely delivered, which is 34–36% the mean adult size, compared to 40% in chimps and 28% in modern humans. This more aligns with the conclusions drawn from the Mojokerto child. A faster development rate could indicate a lower expected lifespan. Based on an average mass of for males and for females, the total energy expenditure (TEE)—the amount of calories consumed in one day—was estimated to be about 2271.8 and 1909.5
kcal The calorie is a unit of energy Unit may refer to: Arts and entertainment * UNIT Unit may refer to: Arts and entertainment * UNIT, a fictional military organization in the science fiction television series ''Doctor Who'' * Unit of action, a di ...
, respectively. This is similar to that of earlier ''Homo'', despite a marked increase in activity and migratory capacity, likely because the longer legs of ''H. erectus'' were more energy-efficient in long-distance movement. Nonetheless, the estimate for ''H. erectus'' females is 84% higher than that for ''Australopithecus'' females, possibly due to an increased body size and a decreased growth rate. A 2011 study, assuming high energy or dietary fat requirements based on the abundance of large game animals at ''H. erectus'' sites, calculated a TEE of 2,700–3,400 kcal of which 27–44% derived from fat, and 44–62% of the fat from animal sources. In comparison, modern humans with a similar activity level have a DEE of 2,450 calories, of which 33% derives from fat, and 49% of the fat from animals.


Bone thickness

The
cortical bone A bone is a Stiffness, rigid tissue (anatomy), tissue that constitutes part of the vertebrate skeleton in animals. Bones protect the various organs of the body, produce red blood cell, red and white blood cells, store minerals, provide structu ...
(the outer layer of the bone) is extraordinarily thickened, particularly in East Asian populations. The skullcaps have oftentimes been confused with fossil turtle
carapace A carapace is a Dorsum (biology), dorsal (upper) section of the exoskeleton or shell in a number of animal groups, including arthropods, such as crustaceans and arachnids, as well as vertebrates, such as turtles and tortoises. In turtles and tor ...
s, and the
medullary canal The medullary cavity (''medulla'', innermost part) is the central cavity of bone shafts where red bone marrow and/or yellow bone marrow (adipose tissue) is stored; hence, the medullary cavity is also known as the marrow cavity. Located in the main ...
in the
long bone The long bones are those that are longer than they are wide. They are one of five types of bone A bone is a rigid tissue Tissue may refer to: Biology * Tissue (biology), an ensemble of similar cells that together carry out a specific funct ...

long bone
s (where the
bone marrow Bone marrow is a semi-solid tissue Tissue may refer to: Biology * Tissue (biology), an ensemble of similar cells that together carry out a specific function * ''Triphosa haesitata'', a species of geometer moth found in North America * ''Triphos ...
is stored, in the limbs) is extremely narrowed (medullary
stenosis A stenosis (from Ancient Greek στενός, "narrow") is an abnormal narrowing in a blood vessel or other tubular Organ (anatomy), organ or structure such as foramina and canals. It is also sometimes called a stricture (as in urethral stricture) ...
). This degree of thickening is usually exhibited in semi-aquatic animals which used their heavy ( pachyosteosclerotic) bones as ballasts to help them sink, induced by
hypothyroidism Hypothyroidism (also called ''underactive thyroid'', ''low thyroid'' or ''hypothyreosis'') is a disorder of the endocrine system in which the thyroid gland The thyroid, or thyroid gland, is an endocrine gland in the neck consisting of two ...

hypothyroidism
. Male specimens have thicker cortical bone than females. It is largely unclear what function this could have served. All pathological inducers would leave scarring or some other indicator not normally exhibited in ''H. erectus''. Before more complete skeletons were discovered, Weidenreich suggested ''H. erectus'' was a gigantic species, thickened bone required to support the massive weight. It was hypothesised that intense physical activity could have induced bone thickening, but in 1970, human biologist
Stanley Marion GarnStanley Marion Garn Ph.D. (October 27, 1922 – August 31, 2007) was a human biologist and educator. He was Professor of Anthropology at the College for Literature, Science and Arts and Professor of Nutrition at the School of Public Health at the Uni ...
demonstrated there is a low correlation between the two at least in modern humans. Garn instead noted different races have different average cortical bone thicknesses, and concluded it is genetic rather than environmental. It is unclear if the condition is caused by increased bone apposition (bone formation) or decreased
bone resorption Bone resorption is resorption Resorption is the absorption into the circulatory system of cells or tissue, usually by osteoclast, osteoclasts. Types of resorption include: * Bone resorption * Herniated Disc Resorption * Tooth resorption * Fetal r ...
, but Garn noted the stenosis is quite similar to the
congenital A birth defect, also known as a congenital disorder, is a condition present at birth Birth is the act or process of bearing or bringing forth offspring, also referred to in technical contexts as parturition. In mammals, the process is initiat ...
condition in modern humans induced by hyper-apposition. In 1985, biological anthropologist Gail Kennedy argued for resorption as a result of
hyperparathyroidism Hyperparathyroidism is an increase in parathyroid hormone (PTH) levels in the blood. This occurs from a disorder either within the parathyroid glands (primary hyperparathyroidism) or outside the parathyroid glands (secondary hyperparathyroidism). ...
caused by
hypocalcemia Hypocalcemia is low calcium Calcium is a chemical element In chemistry, an element is a pure Chemical substance, substance consisting only of atoms that all have the same numbers of protons in their atomic nucleus, nuclei. Unlike ch ...
(
calcium Calcium is a chemical element In chemistry, an element is a pure Chemical substance, substance consisting only of atoms that all have the same numbers of protons in their atomic nucleus, nuclei. Unlike chemical compounds, chemical elem ...
deficiency), a consequence of a dietary shift to low-calcium meat. Kennedy could not explain why the
calcium metabolism Calcium metabolism Metabolism (, from el, μεταβολή ''metabolē'', "change") is the set of life Life is a characteristic that distinguishes physical entities that have biological processes, such as signaling and self-sus ...
of ''H. erectus'' never adjusted. In 1985, American palaeoanthropologist Mary Doria Russell and colleagues argued the supraorbital torus is a response to withstanding major
bending stress In applied mechanics, bending (also known as flexure) characterizes the behavior of a slender structural element subjected to an external Structural load, load applied perpendicularly to a longitudinal axis of the element. The structural elemen ...
which localises in that region when significant force is applied through the front teeth, such as while using the mouth as a third hand to carry objects. In 2004, Noel Boaz and Russel Ciochon suggested it was a result of a cultural practice, wherein ''H. erectus'' would fight each other with fists, stones, or clubs to settle disputes or battle for mates, since the skull is reinforced in key areas. The mandible is quite robust, capable of absorbing heavy blows (no "glass jaw"); the heavy brow ridge protects the eyes, and transitions into a bar covering the ears, connecting all the way in the back of the skull, meaning blows to any of these regions can be effectively dissipated across the skull; and the sagittal keel protects the top of the braincase. Many skullcaps bear usually debilitating fractures, such as the Peking Man skull X, yet they can show signs of surviving and healing. Anthropologist Peter Brown suggested a similar reason for the unusual thickening of the modern
Australian Aboriginal Aboriginal Australians are the various Indigenous peoples of the Mainland Australia, Australian mainland and many of its islands, such as Tasmania, Fraser Island, Hinchinbrook Island, the Tiwi Islands, and Groote Eylandt, but excluding the Tor ...
skull, a result of a ritual popular in central and southeast Australian tribes where adversaries would wack each other with waddies (sticks) until
knockout A knockout (abbreviated to KO or K.O.) is a fight-ending, winning criterion in several full-contact Contact sports are sports that emphasize or require physical contact between players. Some sports, such as mixed martial arts, are scored on im ...

knockout
.


Culture


Social structure

The only fossil evidence regarding ''H. erectus'' group composition comes from 4 sites outside of
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 ...
, Kenya, where 97 footprints made 1.5 Mya were likely left by a group of at least 20 individuals. One of these trackways, based on the size of the footprints, may have been an entirely male group, which could indicate they were some specialised task group, such as a hunting or foraging party, or a border patrol. If correct, this would also indicate sexual division of labour, which distinguishes human societies from those of other great apes and social mammalian carnivores. In modern hunter gatherer societies who target large prey items, typically male parties are dispatched to bring down these high-risk animals, and, due to the low success rate, female parties focus on more predictable foods. Based on modern day savanna chimp and
baboon Baboons are primate A primate ( ) (from Latin , from 'prime, first rank') is a eutherian mammal constituting the Taxonomy (biology), taxonomic order (biology), order Primates (). Primates arose 85–55 million years ago first fro ...

baboon
group composition and behaviour, ''H. erectus ergaster'' may have lived in large, multi-male groups in order to defend against large savanna predators in the open and exposed environment. However, dispersal patterns indicate that ''H. erectus'' generally avoided areas with high carnivore density. It is possible that male–male bonding and male–female friendships were important societal aspects. Because ''H. erectus'' children had faster brain growth rates, ''H. erectus'' likely did not exhibit the same degree of maternal investment or child-rearing behaviours as modern humans. Because ''H. erectus'' men and women are thought to have been about the same size compared to other great apes (exhibit less size-specific sexual dimorphism), it is generally hypothesised that they lived in a monogamous society, as reduced sexual dimorphism in primates is typically correlated with this mating system. However, it is unclear if ''H. erectus'' did in fact exhibit humanlike rates of sexual dimorphism. If they did, then it would mean only female height increased from the ancestor species, which could have been caused by a shift in female fertility or diet, and/or reduced pressure on males for large size. This in turn could imply a shift in female behaviour which made it difficult for males to maintain a harem.


Food

Increasing brain size is often directly associated with a meatier diet and resultant higher caloric intake. However, it is also possible that the energy-expensive guts decreased in size in ''H. erectus'', because the large ape gut is used to synthesize fat by fermenting plant matter which was replaced by dietary animal fat, allowing more energy to be diverted to brain growth. This would have increased brain size indirectly while maintaining the same caloric requirements of ancestor species. ''H. erectus'' may have also been the first to use a
hunting and gathering A hunter-gatherer is a human living in a society in which most or all food is obtained by foraging (collecting wild plants and pursuing Wildlife, wild animals). Hunter-gatherer societies stand in contrast to agriculture, agricultural societies, wh ...
food collecting strategy as a response to the increasing dependence on meat. With an emphasis on teamwork, division of labor, and food sharing, hunting and gathering was a dramatically different subsistence strategy from previous modes. ''H. erectus'' sites frequently are associated with assemblages of medium- to large-sized game, namely
elephant Elephants are the largest existing land animals. Three living species In biology, a species is the basic unit of biological classification, classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A specie ...

elephant
s,
rhino A rhinoceros (, , ), commonly abbreviated to rhino, is a member of any of the five extant Extant is the opposite of the word extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. T ...

rhino
s,
hippo The hippopotamus ( ; ''Hippopotamus amphibius''), also called the hippo, common hippopotamus or river hippopotamus, is a large, mostly herbivorous File:Land_Snail_radula_tracks.jpg#, 250px, Tracks made by terrestrial gastropods with their ...

hippo
s,
bovine The biological subfamily In biological classification In biology, taxonomy () is the scientific study of naming, defining (Circumscription (taxonomy), circumscribing) and classifying groups of biological organisms based on shared characte ...

bovine
, and
boar The wild boar (''Sus scrofa''), also known as the wild swine, common wild pig, Eurasian wild pig, or simply wild pig, is a suid native to much of Eurasia Eurasia () is the largest continent A continent is any of several large ...

boar
. ''H. erectus'' would have had considerable leftovers, potentially pointing to food sharing or long-term
food preservation Food preservation includes food processing practices which prevent the growth of microorganisms, such as yeasts (although some methods work by introducing benign bacteria or fungi to the food), and slow the redox, oxidation of fats that cause Ran ...

food preservation
(such as by drying) if most of the kill was indeed utilized. It is possible that ''H. erectus'' grew to become quite dependent on large-animal meat, and the disappearance of ''H. erectus'' from the
Levant The Levant () is an term referring to a large area in the region of . In its narrowest sense, it is equivalent to the , which included present-day , , , , and most of southwest of the middle . In its widest historical sense, the Levant ...

Levant
is correlated with the local extinction of the
straight-tusked elephant The straight-tusked elephant (''Palaeoloxodon antiquus'') is an extinct species of elephant that inhabited Europe and Western Asia during the Middle and Late Pleistocene (781,000–30,000 years before present). Recovered individuals have reac ...
. Nonetheless, ''H. erectus'' diet likely varied widely depending upon location. For example, at the 780 ka Gesher Benot Ya‘aqov site, Israel, the inhabitants gathered and ate 55 different types of fruits, vegetables, seeds, nuts, and tubers, and it appears that they used fire to roast certain plant materials that otherwise would have been inedible; they also consumed amphibians, reptiles, birds, aquatic and terrestrial invertebrates, in addition to the usual large creatures such as elephant and
fallow deer ''Dama'' is a genus Genus /ˈdʒiː.nəs/ (plural genera /ˈdʒen.ər.ə/) is a taxonomic rank In biological classification In biology, taxonomy () is the scientific study of naming, defining (Circumscription (taxonomy), circumscrib ...

fallow deer
. At the 1.95 Ma FwJJ20 lakeside site in the East Turkana Basin, Kenya, the inhabitants ate (alongside the usual bovids, hippos, and rhinos) aquatic creatures such as turtles, crocodiles, and catfish. The large animals were likely scavenged at this site, but the turtles and fish were possibly collected live. At the 1.5 Ma Trinil H. K. Fauna, Trinil H. K. site, Java, ''H. erectus'' likely gathered fish and shellfish. Dentally, ''H. erectus'' mouths were not as versatile as those of ancestor species, capable of processing a narrower range of foods. However, tools were likely used to process hard foods, thus affecting the chewing apparatus, and this combination may have instead increased dietary flexibility (though this does not equate to a highly varied diet). Such versatility may have permitted ''H. erectus'' to inhabit a range of different environments, and migrate beyond Africa. In 1999, British anthropologist Richard Wrangham proposed the "cooking hypothesis" which states that ''H. erectus'' speciated from the ancestral ''H. habilis'' because of fire usage and cooking 2 Million years ago to explain the rapid doubling of brain size between these two species in only a 500,000 year timespan, and the sudden appearance of the typical human body plan. Cooking makes protein more easily digestible, speeds up nutrient absorption, and destroys food-borne pathogens, which would have increased the environment's natural carrying capacity, allowing group size to expand, causing selective pressure for sociality, requiring greater brain function. However, the fossil record does not associate the emergence of ''H. erectus'' with fire usage nor with any technological breakthrough for that matter, and cooking likely did not become a common practice until after 400 kya. Java Man's dispersal through Southeast Asia coincides with the extirpation of the giant turtle ''Megalochelys'', possibly due to overhunting as the turtle would have been an easy, slow-moving target which could have been stored for quite some time.


Technology


Tool production

''H. erectus'' is credited with inventing the
Acheulean . The types shown are (clockwise from top) cordate, ficron, and ovate. in Nice, France as postulated by Henry de Lumley dated to 400 thousand years ago. Acheulean (; also Acheulian and Mode II), from the French ''acheuléen'' after the ty ...

Acheulean
stone tool industry, succeeding the Oldowan industry, and were the first to make lithic flakes bigger than , and hand axes (which includes bifacial tools with only 2 sides, such as picks, knives, and cleaver (tool), cleavers). Though larger and heavier, these hand axes had sharper, chiseled edges. They were likely multi-purpose tools, used in variety of activities such as cutting meat, wood, or edible plants. In 1979, American paleontologist Thomas Wynn stated that Acheulean technology required operational intelligence (foresight and planning), being markedly more complex than Oldowan technology which included lithics of unstandardized shape, cross-sections, and symmetry. Based on this, he concluded that there is not a significant disparity in intelligence between ''H. erectus'' and modern humans and that, for the last 300,000 years, increasing intelligence has not been a major influencer of cultural evolution. However, a 1 year old ''H. erectus'' specimen shows that this species lacked an extended childhood required for greater brain development, indicating lower cognitive capabilities. A few sites, likely due to occupation over several generations, features hand axes en masse, such as at Melka Kunture, Ethiopia;
Olorgesailie Olorgesailie is a geological formation in East Africa containing a group of Lower Paleolithic Archeology, archaeological sites. It is on the floor of the Eastern Rift Valley in southern Kenya, southwest of Nairobi along the road to Lake Magadi ...
, Kenya; Isimila, Tanzania; and Kalambo Falls, Zambia. The earliest record of Acheulean technology comes from West Turkana, Kenya 1.76 Mya. Oldowan lithics are also known from the site, and the two seemed to coexist for some time. The earliest records of Acheulean technology outside of Africa date to no older than 1 Mya, indicating it only became widespread after some secondary ''H. erectus'' dispersal from Africa. On Java, ''H. erectus'' produced tools from shells at
Sangiran Sangiran is an archaeological excavation site in Java in Indonesia Indonesia ( ), officially the Republic of Indonesia ( id, Republik Indonesia, links=yes ), is a country in Southeast Asia and Oceania between the Indian Ocean, Indian and ...
and Trinil. Spherical stones, measuring in diameter, are frequently found in African and Chinese Lower Paleolithic sites, and were potentially used as bolas; if correct, this would indicate string and cordage technology.


Fire

''H. erectus'' is credited as the first human ancestor to have used fire, though the timing of this invention is debated mainly because campfires very rarely and very poorly preserve over long periods of time, let alone thousands or millions of years. The earliest claimed fire sites are in Kenya, FxJj20 at
Koobi Fora Koobi Fora refers primarily to a region around Koobi Fora Ridge, located on the eastern shore of Lake Turkana Lake Turkana (), formerly known as Lake Rudolf, is a lake in the Kenyan Rift Valley, in northern Kenya, with its far northern en ...
and GnJi 1/6E in the Chemoigut Formation, as far back as 1.5 Mya, and in South Africa, Wonderwerk Cave, 1.7 Mya. The first firekeepers are thought to have simply transported to caves and maintained naturally occurring fires for extended periods of time or only sporadically when the opportunity arose. Maintaining fires would require firekeepers to have knowledge on slow-burning materials such as dung. Fire becomes markedly more abundant in the wider archaeological record after 400,000–300,000 years ago, which can be explained as some advancement in fire management techniques took place at this time or human ancestors only opportunistically used fire until this time. It is possible that firestarting was invented and lost and reinvented multiple times and independently by different communities rather than being invented in one place and spreading throughout the world. The earliest evidence of hearths comes from Gesher Benot Ya’aqov, Israel, over 700,000 years ago, where fire is recorded in multiple layers in an area close to water, both uncharacteristic of natural fires. Artificial lighting may have led to increased waking hours—modern humans have about a 16-hour waking period, whereas other apes are generally awake from only sunup to sundown—and these additional hours were probably used for socializing. Because of this, fire usage is probably also linked to the origin of language. Artificial lighting may have also made sleeping on the ground instead of the trees possible by keeping terrestrial predators at bay. Migration into the frigid climate of Ice Age Europe may have only been possible because of fire, but evidence of fire usage in Europe until about 400–300,000 years ago is notably absent. If these early European ''H. erectus'' did not have fire, it is largely unclear how they stayed warm, avoided predators, and prepared animal fat and meat for consumption; and lightning is less common farther north equating to a reduced availability of naturally occurring fires. It is possible that they only knew how to maintain fires in certain settings in the landscapes and prepared food some distance away from home, meaning evidence of fire and evidence of hominin activity are spaced far apart. Alternatively, ''H. erectus'' may have only pushed farther north during warmer interglacial periods—thus not requiring fire, food storage, or clothing technology— and their dispersal patterns indicate they generally stayed in warmer lower-to-middle latitudes. It is debated if the ''H. e. pekinensis'' inhabitants of
Zhoukoudian Zhoukoudian Peking Man Site (), also romanized as Choukoutien, is a cave system in suburban Fangshan District, Beijing Beijing ( ), Chinese postal romanization, alternatively romanized as Peking ( ), is the Capital city, capital of the ...
, Northern China, were capable of controlling fires as early as 770 kya to stay warm in what may have been a relatively cold climate.


Construction

In 1962, a circle made with volcanic rocks was discovered in
Olduvai Gorge The Olduvai Gorge or Oldupai Gorge in Tanzania is one of the most important paleoanthropology, paleoanthropological sites in the world; it has proven invaluable in furthering understanding of early human evolution. A steep-sided ravine in the Greg ...

Olduvai Gorge
. At intervals, rocks were piled up to high. British palaeoanthropologist Mary Leakey suggested the rock piles were used to support poles stuck into the ground, possibly to support a windbreak or a rough hut. Some modern day nomadic tribes build similar low-lying rock walls to build temporary shelters upon, bending upright branches as poles and using grasses or animal hide as a screen. Dating to 1.75 Mya, it is the oldest claimed evidence of architecture. In Europe, evidence of constructed dwelling structures dating to or following the Holstein Interglacial (which began 424 kya) has been claimed in Bilzingsleben, Germany; Terra Amata (archaeological site), Terra Amata, France; and Fermanville and Saint-Germain-des-Vaux in Normandy. The oldest evidence of a dwelling (and a campfire) in Europe comes from Přezletice, Czech Republic, 700 kya during the Cromerian Interglacial. This dwelling's base measured about on the exterior and on the interior, and is considered to have been a firm surface hut, probably with a vaulted roof made of thick branches or thin poles, supported by a foundation of big rocks and earth, and likely functioned as a winter base camp. The earliest evidence of cave habitation is Wonderwerk Cave, South Africa, about 1.6 Mya, but evidence of cave use globally is sporadic until about 600 kya.


Clothing

It is largely unclear when clothing was invented, with the earliest estimate stretching as far back as 3 Mya to compensate for a lack of insulating body hair. It is known that head lice and body lice (the latter can only inhabit clothed individuals) for modern humans diverged about 170 kya, well before modern humans left Africa, meaning clothes were already well in use before encountering cold climates. One of the first uses of animal hide is thought to have been for clothing, and the oldest hide scrapers date to about 780 kya, though this is not indicative of clothing.


Seafaring

Acheulean artifacts discovered on isolated islands that were never connected to land in the Pleistocene may show seafaring by ''H. erectus'' as early as 1 Mya in Indonesia. They had arrived on the islands of Flores, Timor, and Rote Island, Roti, which would have necessitated crossing the Lombok Strait (the Wallace Line), at least before 800 kya. It is also possible they were the first European mariners as well and crossed the Strait of Gibraltar between North Africa and Spain. A 2021 genetic analysis of these island populations of ''H. erectus'' found no evidence of interbreeding with modern humans. Seafaring capability would show ''H. erectus'' had a great capacity for planning, likely months in advance of the trip. Similarly, ''Homo luzonensis'' is dated between 771,000 to 631,000 years ago. Because Luzon has always been an island in the Quaternary, the ancestors of ''H. luzonensis'' would have had to have made a substantial sea crossing and crossed the Huxley Line.


Healthcare

The earliest probable example of infirming sick group members is a 1.77 Ma ''H. e. georgicus'' specimen who had lost all but one tooth due to age or gum disease, the earliest example of severe chewing impairment, yet still survived for several years afterwards. However, it is possible australopithecines were capable of caring for debilitated group members. Unable to chew, this ''H. e. georgicus'' individual probably ate soft plant or animal foods possibly with assistance from other group members. High-latitude groups are thought to have been predominantly carnivorous, eating soft tissue such as
bone marrow Bone marrow is a semi-solid tissue Tissue may refer to: Biology * Tissue (biology), an ensemble of similar cells that together carry out a specific function * ''Triphosa haesitata'', a species of geometer moth found in North America * ''Triphos ...
or brains, which may have increased survival rates for toothless individuals. The 1.5 Ma Turkana boy was diagnosed with juvenile spinal disc herniation, and, because this specimen was still growing, this caused some scoliosis (abnormal curving of the spine). These usually cause recurrent lower back pain and sciatica (pain running down the leg), and likely restricted Turkana boy in walking, bending, and other daily activities. The specimen appears to have survived into adolescence, which evidences advanced group care. The 1,000–700 ka Java man specimen presents a noticeable osteocyte on the femur, likely Paget's disease of bone, and osteopetrosis, thickening of the bone, likely resulting from skeletal fluorosis caused by ingestion of food contaminated by fluorine-filled volcanic ash (as the specimen was found in ash-filled stratum (geology), strata). Livestock that grazes on volcanic ash ridden fields typically die of acute intoxication within a few days or weeks.


Art and rituals

An engraved Pseudodon shell DUB1006-fL with geometric markings could possibly be evidence of the earliest art-making, dating back to 546–436 kya. Art-making capabilities could be considered evidence of symbolic thinking, which is associated with modern cognition and behavior. In 1976, American archeologist Alexander Marshack asserted that engraved lines on an ox rib, associated with Acheulean lithics, from Pech de l'Azé, France, are similar to a meander (art), meander design found in modern human Upper Paleolithic cave art. Three ostrich eggshell beads associated with Achuelian lithics were found in northwestern Africa, the earliest disc beads ever found, and Acheulian disc beads have also been found in France and Israel. The Middle Pleistocene "Venus of Tan-Tan" and "Venus of Berekhat Ram" are postulated to been crafted by ''H. erectus'' to resemble a human form. They were mostly formed by natural weathering, but slightly modified to emphasize certain grooves to suggest hairline, limbs, and eyes. The former has traces of pigments on the front side, possibly indicating it was colored. ''H. erectus'' was also the earliest human to have intentionally collected red-colored pigments, namely ochre, recorded as early as the Middle Pleistocene. Ochre lumps at
Olduvai Gorge The Olduvai Gorge or Oldupai Gorge in Tanzania is one of the most important paleoanthropology, paleoanthropological sites in the world; it has proven invaluable in furthering understanding of early human evolution. A steep-sided ravine in the Greg ...

Olduvai Gorge
, Tanzania—associated with the 1.4 Ma Olduvai Hominid 9—and Ambrona, Spain—which dates to 424–374 kya—were suggested to have been struck by a hammerstone and purposefully shaped and trimmed. At Terra Amata, France—which dates to 425–400 or 355–325 kya—red, yellow, and brown ochres were recovered in association with pole structures; ochre was probably heated to achieve such a wide color range. As it is unclear if ''H. erectus'' could have used ochre for any practical application, ochre collection might indicate that ''H. erectus'' was the earliest human to have exhibited a sense of aesthetics and to think beyond simply survival. Later human species are postulated to have used ochre as body paint, but in the case of ''H. erectus'', it is contested if body paint was used so early in time. Further, it is unclear if these few examples are not simply isolated incidents of ochre use, as ochre is much more prevalent in Middle and Upper Paleolithic sites attributed to Neanderthals and ''H. sapiens''. In 1935, Jewish German anthropologist Franz Weidenreich speculated that the inhabitants of the Chinese Zhoukoudian, Zhoukoudian Peking Man site were members of some Lower Paleolithic Skull Cult because the skulls all showed fatal blows to the head, breaking in of the foramen magnum at the base of the skull, by-and-large lack of preserved facial aspects, an apparently consistent pattern of breaking on the mandible, and a lack of post-cranial remains (elements that are not the skull). He believed that the inhabitants were headhunting, headhunters, and smashed open the skulls and ate the brains of their victims. However, scavenging animals and natural forces such as flooding can also inflict the same kind of damage to skulls, and there is not enough evidence to suggest manhunting or cannibalism. In 1999, British science writers Marek Kohn and Steven Mithen said that many hand axes exhibit no wear and were produced en masse, and concluded that these symmetrical, tear-drop shaped lithics functioned primarily as display (zoology), display tools so males could prove their fitness to females in some courting ritual, and were discarded afterwards. However, an apparent lack of reported wearing is likely due to a lack of use-wear studies, and only a few sites yield an exorbitant sum of hand axes likely due to gradual accumulation over generations instead of mass production.


Language

In 1984, the vertebral column of the 1.6 Ma adolescent Turkana boy indicated that this individual did not have properly developed respiratory muscles in order to produce speech. In 2001, American anthropologists Bruce Latimer and James Ohman concluded that Turkana boy was afflicted by skeletal dysplasia and scoliosis. In 2006, American anthropologist Marc Meyer and colleagues described a 1.8 Ma ''H. e. georgicus'' specimen as having a spine within the range of variation of modern human spines, contending that Turkana boy had spinal stenosis and was thus not representative of the species. Also, because he considered ''H. e. georgicus'' ancestral to all non-African ''H. erectus'', Meyer concluded that the respiratory muscles of all ''H. erectus'' (at least non-''ergaster'') would not have impeded vocalisation or speech production. However, in 2013 and 2014, anthropologist Regula Schiess and colleagues concluded that there is no evidence of any congenital defects in Turkana boy, and considered the specimen representative of the species. Neurologically, all ''Homo'' have similarly configured brains, and, likewise, the Broca's area, Broca's and Wernicke's area, Wernicke's areas (in charge of sentence formulation and speech production in modern humans) of ''H. erectus'' were comparable to those of modern humans. However, this is not indicative of anything in terms of speech capability as even large chimpanzees can have similarly expanded Broca's area, and it is unclear if these areas served as language centers in archaic humans. A 1-year-old ''H. erectus'' specimen shows that an extended childhood to allow for brain growth, which is a prerequisite in language acquisition, was not exhibited in this species. The hyoid bone supports the tongue and makes possible modulation of the vocal tract to control pitch and volume. A 400 ka ''H. erectus'' hyoid bone from Castel di Guido, Italy, is bar-shaped—more similar to that of other ''Homo'' than to that of non-human apes and ''Australopithecus''—but is devoid of muscle impressions, has a shield-shaped body, and is implied to have had reduced greater horns, meaning ''H. erectus'' lacked a humanlike vocal apparatus and thus anatomical prerequisites for a modern human level of speech. Increasing brain size and cultural complexity in tandem with technological refinement, and the hypothesis that articulate Neanderthals and modern humans may have inherited speech capabilities from the last common ancestor, could possibly indicate that ''H. erectus'' used some
proto-language In the tree model In historical linguistics Historical linguistics, also termed diachronic linguistics, is the scientific study of language change over time. Principal concerns of historical linguistics include: # to describe and accoun ...
and built the basic framework which fully fledged languages would eventually be built around. However, this ancestor may have instead been ''H. heidelbergensis'', as a hyoid bone of a 530 ka ''H. heidelbergensis'' specimen from the Spanish Sima de los Huesos Cave is like that of modern humans, and another specimen from the same area shows an auditory capacity sensitive enough to pick up human speech.


Extinction

The last known occurrence of ''Homo erectus'' is 117,000–108,000 years ago in Ngandong, Java (island), Java according to a study published in 2019. In 2020 researchers reported that ''Homo erectus'' and ''
Homo heidelbergensis ''Homo heidelbergensis'' (also ''H. sapiens heidelbergensis'') is an extinct species In biology, a species is the basic unit of biological classification, classification and a taxonomic rank of an organism, as well as a unit of biodiversit ...

Homo heidelbergensis
'' lost more than half of their climate Ecological niche, niche – climate they were adapted to – space, with no corresponding reduction in physical range, just before extinction and that climate change played a substantial role in extinctions of past ''Homo'' species.


Fossils

The lower cave of the Zhoukoudian cave, China, is one of the most important archaeological sites worldwide.Zanolli, Clément, et al. “Inner Tooth Morphology of Homo Erectus from Zhoukoudian. New Evidence from an Old Collection Housed at Uppsala University, Sweden.” ''Journal of Human Evolution'', vol. 116, Mar. 2018, pp. 1–13. There have been remains of 45 homo erectus individuals found and thousands of tools recovered. 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'. New evidence has shown that ''Homo erectus'' does not have uniquely thick vault bones, as was previously thought.Copes, Lynn E., and William H. Kimbel. “Cranial Vault Thickness in Primates: Homo Erectus Does Not Have Uniquely Thick Vault Bones.” ''Journal of Human Evolution'', vol. 90, Jan. 2016, pp. 120–134. Testing showed that neither Asian or African ''Homo erectus'' had uniquely large vault bones.


Individual fossils

Some of the major ''Homo erectus'' fossils: * Indonesia (island of Java): Trinil 2 (holotype),
Sangiran Sangiran is an archaeological excavation site in Java in Indonesia Indonesia ( ), officially the Republic of Indonesia ( id, Republik Indonesia, links=yes ), is a country in Southeast Asia and Oceania between the Indian Ocean, Indian and ...
collection, Sambungmachan collection, Solo Man, Ngandong collection * China ("
Peking Man Peking Man (''Homo erectus pekinensis'') is a subspecies In biological classification In biology Biology is the natural science that studies life and living organisms, including their anatomy, physical structure, Biochemis ...
"): Lantian (Gongwangling and Chenjiawo), Yunxian,
Zhoukoudian Zhoukoudian Peking Man Site (), also romanized as Choukoutien, is a cave system in suburban Fangshan District, Beijing Beijing ( ), Chinese postal romanization, alternatively romanized as Peking ( ), is the Capital city, capital of the ...
, Nanjing, Hexian * Kenya: KNM ER 3883, KNM ER 3733 * Vietnam: Northern, Tham Khuyen, Hoa Binh * Republic of Georgia: Dmanisi collection (" Homo erectus georgicus") * Ethiopia: Daka skull, Daka calvaria * Eritrea: Buia cranium (possibly H. ergaster) * Denizli Province, Turkey: Kocabas fossil * Drimolen, South Africa: DNH 134


Gallery

File:Homo erectus tautavelensis.jpg, '''' skull. File:Tautavel UK 2.JPG, Replica of lower jaws of ''Homo erectus'' from Tautavel, France. File:Calvaria Sangiran II (A).jpg, Calvaria (skull), Calvaria "
Sangiran Sangiran is an archaeological excavation site in Java in Indonesia Indonesia ( ), officially the Republic of Indonesia ( id, Republik Indonesia, links=yes ), is a country in Southeast Asia and Oceania between the Indian Ocean, Indian and ...
II" original, collection Gustav Heinrich Ralph von Koenigswald, Koenigswald, Senckenberg Museum. File:Homo erectus hand axe Daka Ethiopia.jpg, A reconstruction based on evidence from the Daka skull, Daka Member, Ethiopia File:Pithecanthropus-erectus.jpg, Original fossils of ''Pithecanthropus erectus'' (now ''Homo erectus'') found in
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Java
in 1891.


See also

* ''Anthropopithecus'' * Kozarnika * Multiregional origin of modern humans General: * List of fossil sites ''(with link directory)'' * List of human evolution fossils ''(with images)''


References


Further reading

* * *


External links


Homo erectus
Origins – Exploring the Fossil Record – Bradshaw Foundation


Homo erectus
– The Smithsonian Institution's Human Origins Program
Possible co-existence with Homo Habilis
– BBC News * John D. Hawks, John Hawks'
discussion of the Kocabas fossil



The Age of Homo erectus
– Interactive Map of the Journey of Homo erectus out of Africa
Human Timeline (Interactive)
– Smithsonian Institution, Smithsonian, National Museum of Natural History (August 2016). {{DEFAULTSORT:Homo Erectus Homo erectus, Fossil taxa described in 1892 Mammals described in 1892 Pliocene primates Pleistocene primates Pleistocene mammals of Africa Prehistoric Indonesia Prehistoric China Prehistoric India Prehistoric Kenya Prehistoric Tanzania Prehistoric Hungary Prehistoric Vietnam Prehistoric Georgia (country) Prehistoric Ethiopia Prehistoric Eritrea Prehistoric Anatolia Prehistoric Spain Pleistocene mammals of Asia Pleistocene mammals of Europe Taxa named by Eugène Dubois Tool-using mammals Early species of Homo