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Homo

(Redirected fromEarly humans)
For other uses, seeHomo (disambiguation).
"Genus Homo" redirects here. For the novel, seeGenus Homo(novel).

Homo(fromLatinhomō'human') is agenusofgreat ape(family Hominidae) that emerged from the genusAustralopithecusand encompasses only a single extant species,Homo sapiens(modern humans), along with a number ofextinctspecies (collectively calledarchaic humans) classified as eitherancestralor closely related to modern humans; these includeHomo erectusandHomo neanderthalensis.The oldest member of the genus isHomo habilis,with records of just over 2 million years ago.[a]Homo,together with the genusParanthropus,is probably most closely related to the speciesAustralopithecus africanuswithinAustralopithecus.[4]The closest living relatives ofHomoare of the genusPan(chimpanzeesandbonobos), with the ancestors ofPanandHomoestimated to have diverged around 5.7-11 million years ago during theLate Miocene.[5]

Homo
Temporal range:Piacenzian-present,2.8–0Ma
↓
Notable members ofHomo.
Clockwise from top left: A reconstructedNeanderthal(Homo neanderthalensis) skeleton, amodern human(Homo sapiens) female with a child in India, a reconstructedHomo habilisskull, and a replica skull ofPeking Man(subspecies ofHomo erectus).
Scientific classificationEdit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Primates
Suborder: Haplorhini
Infraorder: Simiiformes
Family: Hominidae
Subfamily: Homininae
Tribe: Hominini
Subtribe: Hominina
Genus: Homo
Linnaeus,1758
Type species
Homo sapiens
Species

For other species or subspecies suggested, seebelow.

Synonyms
Synonyms
  • AfricanthropusDreyer, 1935
  • AtlanthropusArambourg, 1954
  • CyphanthropusPycraft, 1928
  • PalaeanthropusBonarelli, 1909
  • PalaeoanthropusFreudenberg, 1927
  • PithecanthropusDubois, 1894
  • ProtanthropusHaeckel, 1895
  • SinanthropusBlack, 1927
  • TchadanthropusCoppens, 1965
  • TelanthropusBroom & Anderson 1949

H. erectusappeared about 2 million years ago and spread throughoutAfrica(debatably as another species calledHomo ergaster) andEurasiainseveral migrations.The species was adaptive and successful, and persisted for more than a million years before gradually diverging into new species around 500,000 years ago.[b][6]

Anatomically modern humans(H. sapiens) emerged close to 300,000 to 200,000 years ago[7]in Africa, andH. neanderthalensisemerged around the same time inEuropeandWestern Asia.H. sapiensdispersed from Africa inseveral waves,from possibly as early as 250,000 years ago, and certainly by 130,000 years ago, with the so-calledSouthern Dispersal,beginning about 70–50,000 years ago,[8][9][10]leading to thelasting colonisationof Eurasia andOceaniaby 50,000 years ago.H. sapiensmet andinterbred witharchaic humansin Africa and in Eurasia.[11][12]Separate archaic (non-sapiens) human species including Neanderthals are thought to have survived until around 40,000 years ago.

Names and taxonomy

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Main articles:Human taxonomy,Names for the human species,andHomininae
Evolutionary treechart emphasizing the subfamily Homininae and the tribe Hominini. After diverging from the line toPonginae,the early Homininae split into the tribesHomininiandGorillini.The early Hominini split further, separating the line toHomofrom the lineage ofPan.Currently,tribeHominini designates thesubtribesHominina,containing genusHomo;Panina,genusPan;andAustralopithecina,with several extinct genera—the subtribes are not labelled on this chart.

TheLatinnounhomō(genitivehominis) means "human being" or "man"in the generic sense of" human being, mankind ".[c]Thebinomial nameHomo sapienswas coined byCarl Linnaeus(1758).[d][15]Names for other species of the genus were introduced from the second half of the 19th century (H. neanderthalensis1864,H. erectus1892).

The genusHomohas not been strictly defined, even today.[16][17][18]Since the early human fossil record began to slowly emerge from the earth, the boundaries and definitions of the genus have been poorly defined and constantly in flux. Because there was no reason to think it would ever have any additional members,Carl Linnaeusdid not even bother to defineHomowhen he first created it for humans in the 18th century. The discovery of Neanderthal brought the first addition.

The genusHomowas given its taxonomic name to suggest that its member species can be classified as human. And, over the decades of the 20th century, fossil finds of pre-human and early human species from lateMioceneand earlyPliocenetimes produced a rich mix for debating classifications. There is continuing debate on delineatingHomofromAustralopithecus—or, indeed, delineatingHomofromPan.Even so, classifying the fossils ofHomocoincides with evidence of: (1) competenthuman bipedalisminHomo habilisinherited from the earlierAustralopithecusof more than four million years ago, as demonstrated by theLaetoli footprints;and (2)human tool culturehaving begun by 2.5 million years ago to 3 million years ago.[19]

From the late-19th to mid-20th centuries, a number of new taxonomic names, including new generic names, were proposed for early human fossils; most have since been merged withHomoin recognition thatHomo erectuswas a single species with a large geographic spread of early migrations. Many such names are now regarded as "synonyms"withHomo,includingPithecanthropus,[20]Protanthropus,[21]Sinanthropus,[22]Cyphanthropus,[23]Africanthropus,[24]Telanthropus,[25]Atlanthropus,[26]andTchadanthropus.[27][28]

Classifying the genusHomointo species and subspecies is subject to incomplete information and remains poorly done. This has led to using common names ( "Neanderthal" and "Denisovan" ), even in scientific papers, to avoid trinomial names or the ambiguity of classifying groups asincertae sedis(uncertain placement)—for example,H. neanderthalensisvs.H. sapiens neanderthalensis,orH. georgicusvs.H. erectus georgicus.[29]Some recently extinct species in the genus have been discovered only lately and do not as yet have consensus binomial names (seeDenisova hominin).[30]Since the beginning of theHolocene,it is likely thatHomo sapiens(anatomically modern humans) has been the only extant species ofHomo.

John Edward Gray(1825) was an early advocate of classifying taxa by designating tribes and families.[31]Wood and Richmond (2000) proposed thatHominini( "hominins" ) be designated as atribethat comprised all species of early humans and pre-humans ancestral to humans back toafterthechimpanzee–human last common ancestor,and that Homininabe designated asubtribeof Hominini to includeonlythe genusHomo— that is,notincluding the earlier upright walking hominins of thePliocenesuch asAustralopithecus,Orrorin tugenensis,Ardipithecus,orSahelanthropus.[32]Designations alternative to Hominina existed, or were offered:Australopithecinae(Gregory & Hellman 1939) andPreanthropinae(Cela-Conde & Altaba 2002);[33][34][35]and later, Cela-Conde and Ayala (2003) proposed that the four generaAustralopithecus,Ardipithecus,Praeanthropus,andSahelanthropusbe grouped withHomowithinHominini(sansPan).[34]

Evolution

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Main article:Human evolution
Further information:Timeline of human evolution,Hominini,andChimpanzee–human last common ancestor

Australopithecusand the appearance ofHomo

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Further information:Australopithecus

Several species, includingAustralopithecus garhi,Australopithecus sediba,Australopithecus africanus,andAustralopithecus afarensis,have been proposed as the ancestor or sister of theHomolineage.[36][37]These species have morphological features that align them withHomo,but there is no consensus as to which gave rise toHomo.

Especially since the 2010s, the delineation ofHomoinAustralopithecushas become more contentious. Traditionally, the advent ofHomohas been taken to coincide with the first use ofstone tools(theOldowanindustry), and thus by definition with the beginning of theLower Palaeolithic.But in 2010, evidence was presented that seems to attribute the use ofstone toolstoAustralopithecus afarensisaround 3.3 million years ago, close to a million years before the first appearance ofHomo.[38]LD 350-1,a fossil mandible fragment dated to 2.8 Mya, discovered in 2013 inAfar, Ethiopia,was described as combining "primitive traits seen in earlyAustralopithecuswith derived morphology observed in laterHomo.[39]Some authors would push the development ofHomoclose to or even past 3 Mya.[e]This finds support in a recent phylogenetic study in hominins that by using morphological, molecular and radiometric information, dates the emergence ofHomoat 3.3 Ma (4.30 – 2.56 Ma).[40]Others have voiced doubt as to whetherHomo habilisshould be included inHomo,proposing an origin ofHomowithHomo erectusat roughly 1.9 Mya instead.[41]

The most salient physiological development between the earlier australopithecine species andHomois the increase inendocranial volume(ECV), from about 460 cm3(28 cu in) inA. garhito 660 cm3(40 cu in) inH. habilisand further to 760 cm3(46 cu in) inH. erectus,1,250 cm3(76 cu in) inH. heidelbergensisand up to 1,760 cm3(107 cu in) inH. neanderthalensis.However, a steady rise in cranial capacity is observed already inAutralopithecinaand does not terminate after the emergence ofHomo,so that it does not serve as an objective criterion to define the emergence of the genus.[42]

Homo habilis

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Homo habilisemerged about 2.1 Mya. Already before 2010, there were suggestions thatH. habilisshould not be placed in the genusHomobut rather inAustralopithecus.[43][44]The main reason to includeH. habilisinHomo,its undisputed tool use, has become obsolete with the discovery ofAustralopithecustool use at least a million years beforeH. habilis.[38]Furthermore,H. habiliswas long thought to be the ancestor of the more gracileHomo ergaster(Homo erectus). In 2007, it was discovered thatH. habilisandH. erectuscoexisted for a considerable time, suggesting thatH. erectusis not immediately derived fromH. habilisbut instead from a common ancestor.[45]With the publication ofDmanisi skull 5in 2013, it has become less certain that AsianH. erectusis a descendant of AfricanH. ergasterwhich was in turn derived fromH. habilis.Instead,H. ergasterandH. erectusappear to be variants of the same species, which may have originated in either Africa or Asia[46]and widely dispersed throughout Eurasia (includingEurope,Indonesia,China) by 0.5 Mya.[47]

Homo erectus

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Main article:Homo erectus

Homo erectushas often been assumed to have developedanageneticallyfromH. habilisfrom about 2 million years ago. This scenario was strengthened with the discovery ofHomo erectus georgicus,early specimens ofH. erectusfound in theCaucasus,which seemed to exhibit transitional traits withH. habilis.As the earliest evidence forH. erectuswas found outside of Africa, it was considered plausible thatH. erectusdeveloped in Eurasia and then migrated back to Africa. Based on fossils from theKoobi ForaFormation, east of Lake Turkana in Kenya, Spoor et al. (2007) argued thatH. habilismay have survived beyond the emergence ofH. erectus,so that the evolution ofH. erectuswould not have been anagenetically, andH. erectuswould have existed alongsideH. habilisfor about half a million years (1.9 to 1.4million years ago), during the earlyCalabrian.[45]On 31 August 2023, researchers reported, based on genetic studies, that ahuman ancestorpopulation bottleneck(from a possible 100,000 to 1000 individuals) occurred "around 930,000 and 813,000 years ago... lasted for about 117,000 years and brought human ancestors close to extinction."[48][49]

Weiss(1984) estimated that there have been about 44billion (short scale)members of the genusHomofrom its origins to the evolution ofH. erectus,about 56 billion individuals fromH. erectusto theNeolithic,and another 51 billion individuals since the Neolithic. This provides the opportunity for an immense amount of new mutational variation to have arisen during human evolution.[50]

A separate South African speciesHomo gautengensishas been postulated as contemporary withH. erectusin 2010.[51]

Phylogeny

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A taxonomy ofHomowithin thegreat apesis assessed as follows, withParanthropusandHomoemerging withinAustralopithecus(shown herecladisticallygrantingParanthropus,Kenyanthropus,andHomo).[a][f][6][53][52][4][54][55][56][57][58][59][60][excessive citations]The exact phylogeny withinAustralopithecusis still highly controversial. Approximate radiation dates of daughter clades are shown in millions of years ago (Mya).[61][57]SahelanthropusandOrrorin,possibly sisters toAustralopithecus,are not shown here. The naming of groupings is sometimes muddled as often certain groupings are presumed before any cladistic analysis is performed.[55]

Hominoidea

Hylobatidae(gibbons)

Hominidae

Ponginae(orangutans)

Homininae
(8.8)
(15.7)
(20.4 Mya)

Cladogram based on Dembo et al. (2016):[57]

Australopithecines
(7.3 Mya)

Several of theHomolineages appear to have surviving progeny through introgression into other lines. Genetic evidence indicates an archaic lineage separating from the other human lineages 1.5 million years ago, perhapsH. erectus,may have interbred into the Denisovans about 55,000 years ago.[62][54][63]Fossil evidence showsH. erectuss.s. survived at least until 117,000 yrs ago, and the even more basalH. floresiensissurvived until 50,000 years ago. A 1.5-million-yearH. erectus-like lineage appears to have made its way into modern humans through the Denisovans and specifically into the Papuans and aboriginal Australians.[54]The genomes of non-sub-Saharan African humans show what appear to be numerous independent introgression events involving Neanderthal and in some cases also Denisovans around 45,000 years ago.[64][63]The genetic structure of some sub-Saharan African groups seems to be indicative of introgression from a west Eurasian population some 3,000 years ago.[58][65]

Some evidence suggests thatAustralopithecus sedibacould be moved to the genusHomo,or placed in its own genus, due to its position with respect to e.g.H. habilisandH. floresiensis.[56][66]

Dispersal

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See also:Early expansions of hominins out of Africa,Interbreeding between archaic and modern humans,andEarly human migrations

By about 1.8 million years ago,H. erectusis present in both East Africa (H. ergaster) and in Western Asia (H. georgicus). The ancestors of IndonesianH. floresiensismay have left Africa even earlier.[g][56]

Successive dispersals ofHomo erectus(yellow),H. neanderthalensis(ochre) andH. sapiens(red)

Homo erectusand related or derivedarchaic humanspecies over the next 1.5 million years spread throughout Africa and Eurasia[67][68](see:Recent African origin of modern humans). Europe is reached by about 0.5 Mya byHomo heidelbergensis.

Homo neanderthalensisandH. sapiensdevelop after about 300 kya.Homo nalediis present in Southern Africa by 300 kya.

H. sapienssoon after its first emergence spread throughout Africa, and to Western Asia inseveral waves,possibly as early as 250 kya, and certainly by 130 kya. In July 2019, anthropologists reported the discovery of 210,000 year old remains of aH. sapiensand 170,000 year old remains of aH. neanderthalensisinApidima Cave,Peloponnese,Greece,more than 150,000 years older than previousH. sapiensfinds in Europe.[69][70][71]

Most notable is theSouthern DispersalofH. sapiensaround 60 kya, which led to the lasting peopling of Oceania and Eurasia byanatomically modern humans.[11]H. sapiensinterbredwith archaic humans both in Africa and in Eurasia, in Eurasia notably with Neanderthals andDenisovans.[72][73]

Among extant populations ofH. sapiens,the deepest temporal division is found in theSan peopleof Southern Africa, estimated at close to 130,000 years,[74]or possibly more than 300,000 years ago.[75]Temporal division amongnon-Africansis of the order of 60,000 years in the case ofAustralo-Melanesians.Division ofEuropeansandEast Asiansis of the order of 50,000 years, with repeated and significant admixture events throughout Eurasia during theHolocene.

Archaic human species may have survived until the beginning of theHolocene,although they were mostly extinct or absorbed by the expandingH. sapienspopulations by 40 kya (Neanderthal extinction).

List of lineages

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See also:List of human evolution fossils

The species status ofH. rudolfensis,H. ergaster,H. georgicus,H. antecessor,H. cepranensis,H. rhodesiensis,H. neanderthalensis,Denisova hominin,andH. floresiensisremain under debate.H. heidelbergensisandH. neanderthalensisare closely related to each other and have been considered to besubspeciesofH. sapiens.

There has historically been a trend to postulate new human species based on as little as an individual fossil. A "minimalist" approach to human taxonomy recognizes at most three species,H. habilis(2.1–1.5 Mya, membership inHomoquestionable),H. erectus(1.8–0.1 Mya, including the majority of the age of the genus, and the majority of archaic varieties assubspecies,[76][77][78]includingH. heidelbergensisas a late or transitional variety[79][80][81]) andHomo sapiens(300 kya to present, includingH. neanderthalensisand other varieties assubspecies). Consistent definitions and methodology of species delineation are not generally agreed upon in anthropology or paleontology. Indeed, speciating populations of mammals can typically interbreed for several million years after they begin to genetically diverge,[82][83]so all contemporary "species" in the genusHomowould potentially have been able to interbreed at the time, and introgression from beyond the genusHomocan nota prioribe ruled out.[84]It has been suggested thatH. naledimay have been a hybrid with a late survivingAustralipith(taken to mean beyondHomo,ed.),[53]despite the fact that these lineages generally are regarded as long extinct. As discussed above, many introgressions have occurred between lineages, with evidence of introgression after separation of 1.5 million years.

Comparative table ofHomolineages
Lineages Temporal range
(kya) 		Habitat Adult height Adult mass Cranial capacity
(cm3) 		Fossil record Discovery/
publication
of name
H. habilis
membership inHomouncertain 		2,100–1,500[h][i] Tanzania 110–140 cm (3 ft 7 in – 4 ft 7 in) 33–55 kg (73–121 lb) 510–660 Many 1960
1964
H. rudolfensis
membership inHomouncertain 		1,900 Kenya 700 2 sites 1972
1986
H. gautengensis
also classified asH. habilis 		1,900–600 South Africa 100 cm (3 ft 3 in) 3 individuals[87][j] 2010
2010
H. erectus 1,900–140[88][k][89][l] Africa,Eurasia 180 cm (5 ft 11 in) 60 kg (130 lb) 850 (early) – 1,100 (late) Many[m][n] 1891
1892
H. ergaster
AfricanH. erectus 		1,800–1,300[91] East and Southern Africa 700–850 Many 1949
1975
H. antecessor 1,200–800 Western Europe 175 cm (5 ft 9 in) 90 kg (200 lb) 1,000 2 sites 1994
1997
H. floresiensis
classification uncertain 		1,000–50 Indonesia 100 cm (3 ft 3 in) 25 kg (55 lb) 400 7 individuals 2003
2004
H. heidelbergensis
earlyH. neanderthalensis 		600–300[o] Europe, Africa 180 cm (5 ft 11 in) 90 kg (200 lb) 1,100–1,400 Many 1907
1908
H. cepranensis
a single fossil, possiblyH. heidelbergensis 		c. 450[92] Italy 1,000 1 skull cap 1994
2003
H. naledi 335—236[93] South Africa 150 cm (4 ft 11 in) 45 kg (99 lb) 450 15 individuals 2013
2015
H. longi 309–138[94] Northeast China 1,420[95] 1 individual 1933
2021
H. rhodesiensis
earlyH. sapiens 		c. 300 Zambia 1,300 Single or very few 1921
1921
H. sapiens
(anatomically modern humans) 		c. 300–present[p] Worldwide 150–190 cm (4 ft 11 in – 6 ft 3 in) 50–100 kg (110–220 lb) 950–1,800 (extant) ——
1758
Denisova hominin c. 285 - c. 51 Siberia 2 sites 2000
2010[q]
H. neanderthalensis
 240–40[98][r] Europe, Western Asia 170 cm (5 ft 7 in) 55–70 kg (121–154 lb)
(heavily built) 		1,200–1,900 Many 1829
1864
Nesher RamlaHomo
classification uncertain 		140–120 Israel several individuals 2021
H. tsaichangensis
possiblyH. erectusor Denisova 		c. 100[s] Taiwan 1 individual 2008(?)
2015
H. luzonensis
 c. 67[101][102] Philippines 3 individuals 2007
2019

See also

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Footnotes

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  1. ^abThe conventional estimate on the age ofH. habilisis at roughly 2.1 to 2.3 million years.[1][2]Suggestions for pushing back the age to 2.8 Mya were made in 2015 based on the discovery ofa jawbone.[3]
  2. ^Homo erectusin the narrow sense (the Asian species) was extinct by 140,000 years ago;H. erectus soloensis,found inJava,is considered the latest known survival ofH. erectus.Formerly dated to as late as 50,000 to 40,000 years ago, a 2011 study pushed theH. e. soloensisextinction date back to 143,000 years ago at the latest, more likely before 550,000 years ago.[6]
  3. ^The word "human" itself is from Latinhumanus,an adjective formed on the root ofhomo,thought to derive from aProto-Indo-Europeanword for "earth" reconstructed as*dhǵhem-.[13]
  4. ^In 1959,Carl Linnaeuswas designated as thelectotypeforHomo sapiens,[14]which means that following thenomenclatural rules,Homo sapienswas validly defined as the animal species to which Linnaeus belonged.
  5. ^Cela-Conde & Ayala (2003)recognize five genera within Hominina:Ardipithecus,Australopithecus(includingParanthropus),Homo(includingKenyanthropus),Praeanthropus(includingOrrorin), andSahelanthropus.[34]
  6. ^The line to the earliest members ofHomowere derived fromAustralopithecus,a genus that had separated from thechimpanzee–human last common ancestorby lateMioceneor earlyPliocenetimes.[52]
  7. ^In a 2015 phylogenetic study,H. floresiensiswas placed withAustralopithecus sediba,H. habilisandDmanisi Man,raising the possibility that the ancestors ofH. floresiensisleft Africa before the appearance ofH. erectus,possibly even becoming the first hominins to do so and evolved further in Asia.[56]
  8. ^ConfirmedH. habilisfossils are dated to between 2.1 and 1.5 million years ago. This date range overlaps with the emergence ofHomo erectus.[85][86]
  9. ^Hominins with "proto-Homo" traits may have lived as early as 2.8 million years ago, as suggested by a fossil jawbone classified as transitional betweenAustralopithecusandHomodiscovered in 2015.
  10. ^A species proposed in 2010 based on the fossil remains of three individuals dated between 1.9 and 0.6 million years ago. The same fossils were also classified asH. habilis,H. ergasterorAustralopithecusby other anthropologists.
  11. ^H. erectusmay have appeared some 2 million years ago. Fossils dated to as much as 1.8 million years ago have been found both in Africa and in Southeast Asia, and the oldest fossils by a narrow margin (1.85 to 1.77 million years ago) were found in the Caucasus, so that it is unclear whetherH. erectusemerged in Africa and migrated to Eurasia, or if, conversely, it evolved in Eurasia and migrated back to Africa.
  12. ^Homo erectus soloensis,found inJava,is considered the latest known survival ofH. erectus.Formerly dated to as late as 50,000 to 40,000 years ago, a 2011 study pushed back the date of its extinction ofH. e. soloensisto 143,000 years ago at the latest, more likely before 550,000 years ago. [90]
  13. ^Now also included inH. erectusarePeking Man(formerlySinanthropus pekinensis) andJava Man(formerlyPithecanthropus erectus).
  14. ^H. erectusis now grouped into various subspecies, includingHomo erectus erectus,Homo erectus yuanmouensis,Homo erectus lantianensis,Homo erectus nankinensis,Homo erectus pekinensis,Homo erectus palaeojavanicus,Homo erectus soloensis,Homo erectus tautavelensis,Homo erectus georgicus.The distinction from descendant species such asHomo ergaster,Homo floresiensis,Homo antecessor,Homo heidelbergensis and indeedHomo sapiensis not entirely clear.
  15. ^The type fossil isMauer 1,dated to ca. 0.6 million years ago. The transition fromH. heidelbergensistoH. neanderthalensisbetween 300 and 243 thousand years ago is conventional, and makes use of the fact that there is no known fossil in this period. Examples ofH. heidelbergensisare fossils found atBilzingsleben(also classified asHomo erectus bilzingslebensis).
  16. ^The age ofH. sapienshas long been assumed to be close to 200,000 years, but since 2017 there have been a number of suggestions extending this time to as high as 300,000 years. In 2017, fossils found inJebel Irhoud(Morocco) suggest thatHomo sapiensmay have speciated by as early as 315,000 years ago.[96] Genetic evidence has been adduced for an age of roughly 270,000 years.[97]
  17. ^provisional namesHomo sp. AltaiorHomo sapiens ssp. Denisova.
  18. ^The first humans with "proto-Neanderthal traits" lived in Eurasia as early as 0.6 to 0.35 million years ago (classified asH. heidelbergensis,also called achronospeciesbecause it represents a chronological grouping rather than being based on clear morphological distinctions from eitherH. erectusorH. neanderthalensis). There is a fossil gap in Europe between 300 and 243 kya, and by convention, fossils younger than 243 kya are called "Neanderthal".[99]
  19. ^younger than 450 kya, either between 190–130 or between 70–10 kya[100]

References

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  1. ^Stringer, C.B.(1994). "Evolution of early humans". In Jones, S.; Martin, R.; Pilbeam, D. (eds.).The Cambridge Encyclopedia of Human Evolution.Cambridge:Cambridge University Press.p. 242.
  2. ^Schrenk, F.; Kullmer, O.; Bromage, T. (2007). "Chapter 9: The Earliest PutativeHomoFossils ". In Henke, W.; Tattersall, I. (eds.).Handbook of Paleoanthropology.pp. 1611–1631.doi:10.1007/978-3-540-33761-4_52.
  3. ^Spoor, F.; Gunz, P.; Neubauer, S.; Stelzer, S.; Scott, N.; Kwekason, A.; Dean, M.C. (March 2015). "Reconstructed Homo habilis type OH 7 suggests deep-rooted species diversity in early Homo".Nature.519(7541): 83–86.Bibcode:2015Natur.519...83S.doi:10.1038/nature14224.PMID25739632.S2CID4470282.
  4. ^abHaile-Selassie Y, Gibert L, Melillo SM, Ryan TM, Alene M, Deino A, et al. (May 2015). "New species from Ethiopia further expands Middle Pliocene hominin diversity".Nature.521(7553): 483–8.Bibcode:2015Natur.521..483H.doi:10.1038/nature14448.PMID26017448.S2CID4455029.
  5. ^Foley, Robert A.; Mirazón Lahr, Marta (January 2024)."Ghosts of extinct apes: genomic insights into African hominid evolution".Trends in Ecology & Evolution.39(5): 456–466.Bibcode:2024TEcoE..39..456F.doi:10.1016/j.tree.2023.12.009.PMID38302324.
  6. ^abcIndriati E, Swisher CC, Lepre C, Quinn RL, Suriyanto RA, Hascaryo AT, et al. (2011)."The age of the 20 meter Solo River terrace, Java, Indonesia and the survival of Homo erectus in Asia".PLOS One.6(6): e21562.Bibcode:2011PLoSO...621562I.doi:10.1371/journal.pone.0021562.PMC3126814.PMID21738710..
  7. ^Callaway, E. (7 June 2017)."Oldest Homo sapiens fossil claim rewrites our species' history".Nature.doi:10.1038/nature.2017.22114.Retrieved11 June2017.
  8. ^Posth C, Renaud G, Mittnik A, Drucker DG, Rougier H, Cupillard C, et al. (March 2016). "Pleistocene Mitochondrial Genomes Suggest a Single Major Dispersal of Non-Africans and a Late Glacial Population Turnover in Europe".Current Biology.26(6): 827–33.Bibcode:2016CBio...26..827P.doi:10.1016/j.cub.2016.01.037.hdl:2440/114930.PMID26853362.S2CID140098861.
  9. ^See:
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Bibliography

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