{{Short description|Genus of hominins}} {{hatnote group|{{other uses}}{{Redirect|Genus Homo|the novel|Genus Homo (novel){{!}}''Genus Homo'' (novel)}}}} {{Use dmy dates|date=November 2025}} {{Automatic taxobox | fossil_range = Piacenzian–present, {{Fossil range|2.8|0}}{{Period fossil range|Neogene-Quaternary|2.8|0}} | image = Members of Genus Homo 2.png | image_caption = Notable members of ''Homo''.<br/>Clockwise from top left: A Neanderthal (''Homo neanderthalensis'') skull found in Iraq, a modern human (''Homo sapiens'') female in Mexico, a reconstructed ''Homo habilis'' skull, and a replica skull of Peking Man (subspecies of ''Homo erectus''). | taxon = Homo | authority = Linnaeus, 1758 | type_species = ''Homo sapiens'' | type_species_authority = Linnaeus, 1758 | subdivision_ranks = Species | subdivision = *''Homo sapiens'' * {{Ext}}''Homo antecessor'' * {{Ext}}''Homo erectus'' * {{Ext}}''Homo ergaster'' * {{Ext}}''Homo floresiensis'' * {{Ext}}''Homo georgicus'' * {{Ext}}''Homo habilis'' * {{Ext}}''Homo heidelbergensis'' * {{Ext}}''Homo juluensis'' * {{Ext}}''Denisovans / Homo longi'' * {{Ext}}''Homo luzonensis'' * {{Ext}}''Homo naledi'' * {{Ext}}''Homo neanderthalensis'' * {{Ext}}''Homo rudolfensis''
''For other species or subspecies suggested, see below.'' | synonyms = {{collapsible list|bullets = true|title=<small>Synonyms</small> |''Africanthropus'' <small>Dreyer, 1935</small> |''Atlanthropus'' <small>Arambourg, 1954</small> |''Cyphanthropus'' <small>Pycraft, 1928</small> |''Palaeanthropus'' <small>Bonarelli, 1909</small> |''Palaeoanthropus'' <small>Freudenberg, 1927</small> |''Pithecanthropus'' <small>Dubois, 1894</small> |''Protanthropus'' <small>Haeckel, 1895</small> |''Sinanthropus'' <small>Black, 1927</small> |''Tchadanthropus'' <small>Coppens, 1965</small> |''Telanthropus'' <small>Broom & Anderson 1949</small> }} }}
'''''Homo''''' ({{ety|la|homō|human}}) is a genus of great ape (family Hominidae) that emerged from the early hominin genus ''Australopithecus'', encompassing a single extant species, ''Homo sapiens'' ('''modern humans'''), along with a number of extinct species (e.g. ''Homo erectus'' and ''Homo neanderthalensis'') classified as either ancestral or closely related to modern humans, collectively called '''archaic humans'''. ''Homo'', together with the genus ''Paranthropus'', is probably most closely related to the species ''Australopithecus africanus'' within ''Australopithecus''.''<ref name="Saylor 2015 483–488">{{cite journal |display-authors=6 |vauthors=Haile-Selassie Y, Gibert L, Melillo SM, Ryan TM, Alene M, Deino A, Levin NE, Scott G, Saylor BZ |date=May 2015 |title=New species from Ethiopia further expands Middle Pliocene hominin diversity |journal=Nature |volume=521 |issue=7553 |pages=483–8 |bibcode=2015Natur.521..483H |doi=10.1038/nature14448 |pmid=26017448 |s2cid=4455029}}</ref>'' The closest living relatives of ''Homo'' are of the hominin genus ''Pan'' (chimpanzees and bonobos), with the ancestors of ''Pan'' and ''Homo'' estimated to have diverged around 5.7–11 million years ago during the Late Miocene.<ref>{{Cite journal |last1=Foley |first1=Robert A. |last2=Mirazón Lahr |first2=Marta |date=January 2024 |title=Ghosts of extinct apes: genomic insights into African hominid evolution |journal=Trends in Ecology & Evolution |volume=39 |issue=5 |pages=456–466 |language=en |doi=10.1016/j.tree.2023.12.009|doi-access=free |pmid=38302324 |bibcode=2024TEcoE..39..456F }}</ref>
The oldest member of the genus is ''Homo habilis'', with fossil records of just over 2 million years ago.{{efn|The conventional estimate on the age of ''H. habilis'' is at roughly 2.1 to 2.3 million years.<ref>{{cite book|title= The Cambridge Encyclopedia of Human Evolution |last=Stringer |first=C.B. |author-link=Chris Stringer |chapter=Evolution of early humans |editor-last1=Jones |editor-first1=S. |editor-last2=Martin |editor-first2=R. |editor-last3=Pilbeam |editor-first3=D. |year=1994 |publisher=Cambridge University Press |location=Cambridge |page=242}}</ref><ref>{{cite book |last1=Schrenk |first1=F. |last2=Kullmer |first2=O. |last3=Bromage |first3=T. |chapter=Chapter 9: The Earliest Putative ''Homo'' Fossils |editor-last1=Henke |editor-first1=W. |editor-last2=Tattersall |editor-first2=I. |title=Handbook of Paleoanthropology |date=2007 |pages=1611–1631 |doi=10.1007/978-3-540-33761-4_52}}</ref> Suggestions for pushing back the age to 2.8 Mya were made in 2015 based on the discovery of a jawbone.<ref>{{cite journal |last1=Spoor |first1=F. |last2=Gunz |first2=P. |last3=Neubauer |first3=S. |last4=Stelzer |first4=S. |last5=Scott |first5=N. |last6=Kwekason |first6=A. |last7=Dean |first7=M.C. |title=Reconstructed Homo habilis type OH 7 suggests deep-rooted species diversity in early Homo |journal=Nature |volume=519 |issue=7541 |pages=83–86 |date=March 2015 |pmid=25739632 |doi=10.1038/nature14224 |s2cid=4470282 |bibcode=2015Natur.519...83S}}</ref>|name=fn1}} ''H. erectus'' appeared about 2 million years ago and spread throughout Africa (debatably as another species called ''Homo ergaster'') and Eurasia in several 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.{{Efn|name=fn3|''Homo erectus'' in the narrow sense (the Asian species) was extinct by 140,000 years ago; ''H. erectus soloensis'', found in Java, is considered the latest known survival of ''H. erectus''. Formerly dated to as late as 50,000 to 40,000 years ago, a 2011 study pushed the ''H. e. soloensis'' extinction date back to 143,000 years ago at the latest, more likely before 550,000 years ago.<ref name="Indriati-2011">{{cite journal |vauthors=Indriati E, Swisher CC, Lepre C, Quinn RL, Suriyanto RA, Hascaryo AT, Grün R, Feibel CS, Pobiner BL, Aubert M, Lees W, Antón SC |display-authors=6 |title=The age of the 20 meter Solo River terrace, Java, Indonesia and the survival of Homo erectus in Asia |journal=PLOS One |volume=6 |issue=6 |article-number=e21562 |year=2011 |pmid=21738710 |pmc=3126814 |doi=10.1371/journal.pone.0021562 |bibcode=2011PLoSO...621562I |doi-access=free}}.</ref>}}<ref name="Indriati-2011" />
Anatomically modern humans (early ''H. sapiens'') emerged close to 300,000 to 200,000 years ago<ref name="NAT-20170607a">{{cite journal |last=Callaway |first=E. |title= Oldest Homo sapiens fossil claim rewrites our species' history |url=http://www.nature.com/news/oldest-homo-sapiens-fossil-claim-rewrites-our-species-history-1.22114 |date=7 June 2017 |journal=Nature |doi=10.1038/nature.2017.22114 |access-date=11 June 2017|url-access=subscription |doi-access=free }}</ref> in Africa, and ''H. neanderthalensis'' emerged around the same time in Europe and Western Asia. ''H. sapiens'' dispersed from Africa in several waves, from possibly as early as 250,000 years ago, and certainly by 130,000 years ago, with the so-called Southern Dispersal, beginning about 70,000–50,000 years ago,<ref name="Posth">{{cite journal |vauthors=Posth C, Renaud G, Mittnik A, Drucker DG, Rougier H, Cupillard C, Valentin F, Thevenet C, Furtwängler A, Wißing C, Francken M, Malina M, Bolus M, Lari M, Gigli E, Capecchi G, Crevecoeur I, Beauval C, Flas D, Germonpré M, van der Plicht J, Cottiaux R, Gély B, Ronchitelli A, Wehrberger K, Grigorescu D, Svoboda J, Semal P, Caramelli D, Bocherens H, Harvati K, Conard NJ, Haak W, Powell A, Krause J |display-authors=6 |title=Pleistocene Mitochondrial Genomes Suggest a Single Major Dispersal of Non-Africans and a Late Glacial Population Turnover in Europe |journal=Current Biology |volume=26 |issue=6 |pages=827–33 |date=March 2016 |pmid=26853362 |doi=10.1016/j.cub.2016.01.037 |bibcode=2016CBio...26..827P |hdl-access=free |s2cid=140098861 |hdl=2440/114930|url=https://pure.rug.nl/ws/files/79384808/Pleistocene_Mitochondrial_Genomes_Suggest_a_Single_Major_Dispersal_of_Non_Africans.pdf }}</ref><ref>See: * {{cite journal |vauthors=Karmin M, Saag L, Vicente M, Wilson Sayres MA, Järve M, Talas UG, Rootsi S, Ilumäe AM, Mägi R, Mitt M, Pagani L, Puurand T, Faltyskova Z, Clemente F, Cardona A, Metspalu E, Sahakyan H, Yunusbayev B, Hudjashov G, DeGiorgio M, Loogväli EL, Eichstaedt C, Eelmets M, Chaubey G, Tambets K, Litvinov S, Mormina M, Xue Y, Ayub Q, Zoraqi G, Korneliussen TS, Akhatova F, Lachance J, Tishkoff S, Momynaliev K, Ricaut FX, Kusuma P, Razafindrazaka H, Pierron D, Cox MP, Sultana GN, Willerslev R, Muller C, Westaway M, Lambert D, Skaro V, Kovačevic L, Turdikulova S, Dalimova D, Khusainova R, Trofimova N, Akhmetova V, Khidiyatova I, Lichman DV, Isakova J, Pocheshkhova E, Sabitov Z, Barashkov NA, Nymadawa P, Mihailov E, Seng JW, Evseeva I, Migliano AB, Abdullah S, Andriadze G, Primorac D, Atramentova L, Utevska O, Yepiskoposyan L, Marjanovic D, Kushniarevich A, Behar DM, Gilissen C, Vissers L, Veltman JA, Balanovska E, Derenko M, Malyarchuk B, Metspalu A, Fedorova S, Eriksson A, Manica A, Mendez FL, Karafet TM, Veeramah KR, Bradman N, Hammer MF, Osipova LP, Balanovsky O, Khusnutdinova EK, Johnsen K, Remm M, Thomas MG, Tyler-Smith C, Underhill PA, Willerslev E, Nielsen R, Metspalu M, Villems R, Kivisild T |display-authors=6 |title=A recent bottleneck of Y chromosome diversity coincides with a global change in culture |journal=Genome Research |volume=25 |issue=4 |pages=459–66 |date=April 2015 |pmid=25770088 |pmc=4381518 |doi=10.1101/gr.186684.114}} * {{cite journal |vauthors=Pagani L, Lawson DJ, Jagoda E, Mörseburg A, Eriksson A, Mitt M, Clemente F, Hudjashov G, DeGiorgio M, Saag L, Wall JD, Cardona A, Mägi R, Wilson Sayres MA, Kaewert S, Inchley C, Scheib CL, Järve M, Karmin M, Jacobs GS, Antao T, Iliescu FM, Kushniarevich A, Ayub Q, Tyler-Smith C, Xue Y, Yunusbayev B, Tambets K, Mallick CB, Saag L, Pocheshkhova E, Andriadze G, Muller C, Westaway MC, Lambert DM, Zoraqi G, Turdikulova S, Dalimova D, Sabitov Z, Sultana GN, Lachance J, Tishkoff S, Momynaliev K, Isakova J, Damba LD, Gubina M, Nymadawa P, Evseeva I, Atramentova L, Utevska O, Ricaut FX, Brucato N, Sudoyo H, Letellier T, Cox MP, Barashkov NA, Skaro V, Mulahasanovic L, Primorac D, Sahakyan H, Mormina M, Eichstaedt CA, Lichman DV, Abdullah S, Chaubey G, Wee JT, Mihailov E, Karunas A, Litvinov S, Khusainova R, Ekomasova N, Akhmetova V, Khidiyatova I, Marjanović D, Yepiskoposyan L, Behar DM, Balanovska E, Metspalu A, Derenko M, Malyarchuk B, Voevoda M, Fedorova SA, Osipova LP, Lahr MM, Gerbault P, Leavesley M, Migliano AB, Petraglia M, Balanovsky O, Khusnutdinova EK, Metspalu E, Thomas MG, Manica A, Nielsen R, Villems R, Willerslev E, Kivisild T, Metspalu M |display-authors=6 |title=Genomic analyses inform on migration events during the peopling of Eurasia |journal=Nature |volume=538 |issue=7624 |pages=238–242 |date=October 2016 |pmid=27654910 |pmc=5164938 |doi=10.1038/nature19792 |bibcode=2016Natur.538..238P}}</ref><ref name="HaberM">{{cite journal |vauthors=Haber M, Jones AL, Connell BA, Arciero E, Yang H, Thomas MG, Xue Y, Tyler-Smith C |display-authors=6 |title=A Rare Deep-Rooting D0 African Y-Chromosomal Haplogroup and Its Implications for the Expansion of Modern Humans Out of Africa |journal=Genetics |volume=212 |issue=4 |pages=1421–1428 |date=August 2019 |pmid=31196864 |pmc=6707464 |doi=10.1534/genetics.119.302368}}</ref> leading to the lasting colonisation of Eurasia and Oceania by 50,000 years ago. ''H. sapiens'' met and interbred with archaic humans in Africa and in Eurasia.<ref name="A draft sequence of the Neandertal">{{cite journal |vauthors=Green RE, Krause J, Briggs AW, Maricic T, Stenzel U, Kircher M, Patterson N, Li H, Zhai W, Fritz MH, Hansen NF, Durand EY, Malaspinas AS, Jensen JD, Marques-Bonet T, Alkan C, Prüfer K, Meyer M, Burbano HA, Good JM, Schultz R, Aximu-Petri A, Butthof A, Höber B, Höffner B, Siegemund M, Weihmann A, Nusbaum C, Lander ES, Russ C, Novod N, Affourtit J, Egholm M, Verna C, Rudan P, Brajkovic D, Kucan Ž, Gušic I, Doronichev VB, Golovanova LV, Lalueza-Fox C, de la Rasilla M, Fortea J, Rosas A, Schmitz RW, Johnson PL, Eichler EE, Falush D, Birney E, Mullikin JC, Slatkin M, Nielsen R, Kelso J, Lachmann M, Reich D, Pääbo S |display-authors=6 |title=A draft sequence of the Neandertal genome |journal=Science |volume=328 |issue=5979 |pages=710–722 |date=May 2010 |pmid=20448178 |pmc=5100745 |doi=10.1126/science.1188021 |bibcode=2010Sci...328..710G}}</ref><ref>{{cite journal |vauthors=Lowery RK, Uribe G, Jimenez EB, Weiss MA, Herrera KJ, Regueiro M, Herrera RJ |title=Neanderthal and Denisova genetic affinities with contemporary humans: introgression versus common ancestral polymorphisms |journal=Gene |volume=530 |issue=1 |pages=83–94 |date=November 2013 |pmid=23872234 |doi=10.1016/j.gene.2013.06.005}} This study raises the possibility of observed genetic affinities between archaic and modern human populations being mostly due to common ancestral polymorphisms.</ref> Separate archaic (non-''sapiens'') human species including Neanderthals are thought to have survived until around 40,000 years ago.
== Names and taxonomy == {{anchor|Name}}{{anchor|Taxonomy}}{{anchor|Hominina}} {{main|Human taxonomy|Names for the human species|Homininae}} thumb|upright=1.5|The hominin fossil record as known in 2024 [[File:Homininae.svg|frame|Evolutionary tree chart emphasizing the subfamily Homininae and the tribe Hominini. After diverging from the line to Ponginae, the early Homininae split into the tribes Hominini and Gorillini. The early Hominini split further, separating the line to ''Homo'' from the lineage of ''Pan''. Currently, ''tribe'' Hominini designates the ''subtribes'' Hominina, containing genus ''Homo''; Panina, genus ''Pan''; and Australopithecina, with several extinct genera—the subtribes are not labelled on this chart.]]
The Latin noun {{lang|la|homō}} (genitive {{lang|la|hominis}}) means "human being" or "man" in the generic sense of "human being, mankind".{{efn|The word "human" itself is from Latin {{lang|la|humanus}}, an adjective formed on the root of ''homo'', thought to derive from a Proto-Indo-European word for "earth" reconstructed as {{lang|ine-x-proto|dʰǵʰem-}}.<ref>{{cite web|url=http://www.bartleby.com/61/roots/IE104.html |title=The American Heritage Dictionary of the English Language |edition=4th |date=2000}}</ref>}} The binomial name ''Homo sapiens'' was coined by Carl Linnaeus (1758).{{efn|In 1959, Carl Linnaeus was designated as the lectotype for ''Homo sapiens'',<ref>{{cite journal |last1=Stearn |first1=W.T. |year=1959 |title=The background of Linnaeus's contributions to the nomenclature and methods of systematic biology |journal=Systematic Zoology |volume=8 |issue=1 |pages=4–22 |doi=10.2307/2411603 |jstor=2411603}}</ref> which means that following the nomenclatural rules, ''Homo sapiens'' was validly defined as the animal species to which Linnaeus belonged.}}<ref>{{cite book|url=https://www.biodiversitylibrary.org/item/80764#page/28/mode/1up |title=Systema naturæ. Regnum animale. |last1=von Linné |first1=C. |publisher=Sumptibus Guilielmi Engelmann |year=1758 |edition=10 |pages=18, 20 |access-date=19 November 2012}}</ref> Names for other species of the genus were introduced from the second half of the 19th century (''H. neanderthalensis'' 1864, ''H. erectus'' 1892).
The genus ''Homo'' has not been strictly defined, even today.<ref name="Schwartz 2015">{{cite journal |last1=Schwartz |first1=J.H. |last2=Tattersall |first2=I. |title=Defining the genus ''Homo'' |journal=Science |volume=349 |issue=6251 |pages=931–932 |date=August 2015 |pmid=26315422 |doi=10.1126/science.aac6182 |s2cid=206639783 |bibcode=2015Sci...349..931S}}</ref><ref name="Homo definition 2015">{{cite news |last1=Lents |first1=N. |url=https://www.thewildernist.org/2015/10/homo-nadeli-problems-with-homo-genus/ |archive-url=https://web.archive.org/web/20151118084131/https://www.thewildernist.org/2015/10/homo-nadeli-problems-with-homo-genus/ |archive-date=18 November 2015 |title=''Homo naledi'' and the problems with the ''Homo'' genus |magazine=The Wildernist |date=4 October 2014 |access-date=2 November 2015 }}</ref><ref>{{cite journal |last1=Wood |first1=B. |last2=Collard |first2=M. |title=The human genus |journal=Science |volume=284 |issue=5411 |pages=65–71 |date=April 1999 |pmid=10102822 |doi=10.1126/science.284.5411.65 |s2cid=7018418 |bibcode=1999Sci...284...65.}}</ref> 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 Linnaeus did not define ''Homo'' when he first created it for humans in the 18th century. The discovery of Neanderthal brought the first addition.
The genus ''Homo'' was 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 late Miocene and early Pliocene times produced a rich mix for debating classifications. There is continuing debate on delineating ''Homo'' from ''Australopithecus''—or, indeed, delineating ''Homo'' from ''Pan''. Even so, classifying the fossils of ''Homo'' coincides with evidence of: (1) competent human bipedalism in ''Homo habilis'' inherited from the earlier ''Australopithecus'' of more than four million years ago, as demonstrated by the Laetoli footprints; and (2) human tool culture having begun by 2.5 million years ago to 3 million years ago.<ref>{{Cite journal |last=Plummer |first=Thomas |date=9 February 2023 |title=Expanded geographic distribution and dietary strategies of the earliest Oldowan hominins and Paranthropus |url=https://www.science.org/doi/10.1126/science.abo7452 |journal=Science|volume=379 |issue=6632 |pages=561–566 |doi=10.1126/science.abo7452 |pmid=36758076 |bibcode=2023Sci...379..561P |s2cid=256697931 }}</ref>
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 with ''Homo'' in recognition that ''Homo erectus'' was a single species with a large geographic spread of early migrations. Many such names are now regarded as "synonyms" with ''Homo'', including ''Pithecanthropus'',<ref>"ape-man", from ''Pithecanthropus erectus'' (Java Man), Eugène Dubois, ''Pithecanthropus erectus: eine menschenähnliche Übergangsform aus Java'' (1894), identified with the ''Pithecanthropus alalus'' (i.e. "non-speaking ape-man") hypothesized earlier by Ernst Haeckel</ref> ''Protanthropus'',<ref>{{cite journal|quote=early man |title=Protanthropus primigenius |author-first=Ernst |author-last=Haeckel |author-link=Ernst Haeckel |journal=Systematische Phylogenie |volume=3 |date=1895 |pages=[https://www.biodiversitylibrary.org/page/40568382#page/655/mode/1up p. 625]}}</ref> ''Sinanthropus'',<ref>"Sinic man", from ''Sinanthropus pekinensis'' (Peking Man), Davidson Black (1927).</ref> ''Cyphanthropus'',<ref>"crooked man", from ''Cyphanthropus rhodesiensis'' (Rhodesian Man) William Plane Pycraft (1928).</ref> ''Africanthropus'',<ref>"African man", used by T.F. Dreyer (1935) for the Florisbad Skull he found in 1932 (also ''Homo florisbadensis'' or ''Homo helmei''). Also the genus suggested for a number of archaic human skulls found at Lake Eyasi by Weinert (1938). Leaky, ''Journal of the East Africa Natural History Society'' (1942), [https://www.biodiversitylibrary.org/page/36831937#page/287/mode/1up p. 43].</ref> ''Telanthropus'',<ref>"remote man"; from ''Telanthropus capensis'' (Broom and Robinson 1949), see (1961), [https://www.biodiversitylibrary.org/page/7952323#page/571/mode/1up p. 487].</ref> ''Atlanthropus'',<ref>from ''Atlanthropus mauritanicus'', name given to the species of fossils (three lower jaw bones and a parietal bone of a skull) discovered in 1954 to 1955 by Camille Arambourg in Tighennif, Algeria. {{cite journal |last1=Arambourg |first1=C. |year=1955 |title=A recent discovery in human paleontology: Atlanthropus of ternifine (Algeria) |journal=American Journal of Physical Anthropology |volume=13 |issue=2 |pages=191–201 |doi=10.1002/ajpa.1330130203 |bibcode=1955AJPA...13..191A }}</ref> and ''Tchadanthropus''.<ref>{{cite journal |last1=Coppens |first1=Y. |title=L'Hominien du Tchad |journal=Actes V Congr. PPEC |volume=I |date=1965 |pages=329f}}</ref><ref>{{cite journal|last1=Coppens |first1=Y. |title=Le Tchadanthropus |journal=Anthropologia |volume=70 |date=1966 |pages=5–16}}</ref>
Classifying the genus ''Homo'' into 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 as ''incertae sedis'' (uncertain placement)—for example, ''H. neanderthalensis'' vs. ''H. sapiens neanderthalensis'', or ''H. georgicus'' vs. ''H. erectus georgicus''.<ref>{{cite thesis|last=Vivelo |first=Alexandra |date=25 August 2013 |url=https://scholarworks.calstate.edu/concern/theses/7w62f909h |title=Characterization of Unique Features of the Denisovan Exome |archive-url=https://web.archive.org/web/20131029211406/http://csueastbay-dspace.calstate.edu:9000/handle/10211.3/47490 |archive-date=29 October 2013 |degree=Masters |hdl=10211.3/47490 |hdl-access=free |url-status=live |publisher=California State University}}</ref> Some recently extinct species in the genus have been discovered only lately and do not as yet have consensus binomial names (see Denisova hominin).<ref name="Barras2012">{{cite web |title=Chinese human fossils unlike any known species |last1=Barras |first1=C. |url=https://www.newscientist.com/article/dn21586-chinese-human-fossils-unlike-any-known-species.html |date=14 March 2012 |access-date=15 March 2012 |magazine=New Scientist}}</ref> Since the beginning of the Holocene, it is likely that ''Homo sapiens'' (anatomically modern humans) has been the only extant species of ''Homo''.
John Edward Gray (1825) was an early advocate of classifying taxa by designating tribes and families.<ref>{{cite journal |last=Gray |first=J.E. |title=An outline of an attempt at the disposition of Mammalia into Tribes and Families, with a list of genera apparently appertaining to each Tribe |journal=Annals of Philosophy |series=new series |year=1825 |pages=337–344}}</ref> Wood and Richmond (2000) proposed that Hominini ("hominins") be designated as a tribe that comprised all species of early humans and pre-humans ancestral to humans back to ''after'' the chimpanzee–human last common ancestor, and that Hominin<u>a</u> be designated a subtribe of Hominini to include ''only'' the genus ''Homo'' — that is, ''not'' including the earlier upright walking hominins of the Pliocene such as ''Australopithecus'', ''Orrorin tugenensis'', ''Ardipithecus'', or ''Sahelanthropus''.<ref>{{Harvp|Wood|Richmond|2000|pages=19–60}}</ref> Designations alternative to Hominina existed, or were offered: ''Australopithecinae'' (Gregory & Hellman 1939) and ''Preanthropinae'' (Cela-Conde & Altaba 2002);<ref>{{cite journal |last1=Brunet |first1=M. |last2=Guy |first2=F. |last3=Pilbeam |first3=D. |last4=Mackaye |first4=H.T. |last5=Likius |first5=A. |last6=Ahounta |first6=D. |last7=Beauvilain |first7=A. |last8=Blondel |first8=C. |last9=Bocherens |first9=H. |last10=Boisserie |first10=J.R. |last11=De Bonis |first11=L. |last12=Coppens |first12=Y. |last13=Dejax |first13=J. |last14=Denys |first14=C. |last15=Duringer |first15=P. |last16=Eisenmann |first16=V. |last17=Fanone |first17=G. |last18=Fronty |first18=P. |last19=Geraads |first19=D. |last20=Lehmann |first20=T. |last21=Lihoreau |first21=F. |last22=Louchart |first22=A. |last23=Mahamat |first23=A. |last24=Merceron |first24=G. |last25=Mouchelin |first25=G. |last26=Otero |first26=O. |last27=Campomanes |first27=P. |last28=Ponce De Leon |first28=M. |last29=Rage |first29=J.C. |last30=Sapanet |first30=M. |last31=Schuster |first31=M. |last32=Sudre |first32=J. |last33=Tassy |first33=P. |last34=Valentin |first34=X. |last35=Vignaud |first35=P. |last36=Viriot |first36=L. |last37=Zazzo |first37=A. |last38=Zollikofer |first38=C. |display-authors=6 |title=A new hominid from the Upper Miocene of Chad, Central Africa |journal=Nature |volume=418 |issue=6894 |pages=145–51 |date=July 2002 |pmid=12110880 |doi=10.1038/nature00879 |s2cid=1316969 |bibcode=2002Natur.418..145B |url=http://doc.rero.ch/record/13388/files/PAL_E190.pdf}}</ref><ref name="Cela-Conde-2003">{{cite journal |last1=Cela-Conde |first1=C.J. |last2=Ayala |first2=F.J. |title=Genera of the human lineage |journal=Proceedings of the National Academy of Sciences of the United States of America |volume=100 |issue=13 |pages=7684–7689 |date=June 2003 |pmid=12794185 |pmc=164648 |doi=10.1073/pnas.0832372100 |bibcode=2003PNAS..100.7684C |doi-access=free}}</ref><ref>{{cite journal |last1=Wood |first1=B. |last2=Lonergan |first2=N. |title=The hominin fossil record: taxa, grades and clades |journal=Journal of Anatomy |volume=212 |issue=4 |pages=354–76 |date=April 2008 |pmid=18380861 |pmc=2409102 |doi=10.1111/j.1469-7580.2008.00871.x |url=http://www.gwu.edu/~hogwash/BW_PDFs/RP156.pdf}}</ref> and later, Cela-Conde and Ayala (2003) proposed that the four genera ''Australopithecus'', ''Ardipithecus'', ''Praeanthropus'', and ''Sahelanthropus'' be grouped with ''Homo'' within Hominini (sans ''Pan'').<ref name="Cela-Conde-2003" />
==Evolution== {{main|Human evolution}} {{further|Timeline of human evolution|Hominini|Chimpanzee–human last common ancestor}}
===''Australopithecus'' and the appearance of ''Homo''=== {{further|Australopithecus}} [[File:Australopithecus afarensis Naturhistorisches Museum Wenen 7-06-2023 11-13-12.jpg|thumb|200px|Reconstruction of ''Australopithecus afarensis'', Natural History Museum, Vienna]] Several species, including ''Australopithecus garhi'', ''Australopithecus sediba'', ''Australopithecus africanus'', and ''Australopithecus afarensis'', have been proposed as the ancestor or sister of the ''Homo'' lineage.<ref>{{cite journal |last1=Pickering |first1=R. |last2=Dirks |first2=P.H. |last3=Jinnah |first3=Z. |last4=de Ruiter |first4=D.J. |last5=Churchill |first5=S.E. |last6=Herries |first6=A.I. |last7=Woodhead |first7=J.D. |last8=Hellstrom |first8=J.C. |last9=Berger |first9=L.R. |display-authors=6 |title=Australopithecus sediba at 1.977 Ma and implications for the origins of the genus Homo |journal=Science |volume=333 |issue=6048 |pages=1421–1423 |date=September 2011 |pmid=21903808 |doi=10.1126/science.1203697 |s2cid=22633702 | bibcode =2011Sci...333.1421P|doi-access=free }}</ref><ref>{{cite journal |last1=Asfaw |first1=B. |last2=White |first2=T. |last3=Lovejoy |first3=O. |last4=Latimer |first4=B. |last5=Simpson |first5=S. |last6=Suwa |first6=G. |title=Australopithecus garhi: a new species of early hominid from Ethiopia |journal=Science |volume=284 |issue=5414 |pages=629–635 |date=April 1999 |pmid=10213683 |doi=10.1126/science.284.5414.629 |bibcode=1999Sci...284..629A}}</ref> These species have morphological features that align them with ''Homo'', but there is no consensus as to which gave rise to ''Homo''.
Especially since the 2010s, the delineation of ''Homo'' in ''Australopithecus'' has become more contentious. Traditionally, the advent of ''Homo'' has been taken to coincide with the first use of stone tools (the Oldowan industry), and thus by definition with the beginning of the Lower Palaeolithic. But in 2010, evidence was presented that seems to attribute the use of stone tools to ''Australopithecus afarensis'' around 3.3 million years ago, close to a million years before the first appearance of ''Homo''.<ref name=Dikika>{{cite journal |last1=McPherron |first1=S.P. |last2=Alemseged |first2=Z. |last3=Marean |first3=C.W. |last4=Wynn |first4=J.G. |last5=Reed |first5=D. |last6=Geraads |first6=D. |last7=Bobe |first7=R. |last8=Béarat |first8=H.A. |display-authors=6 |title=Evidence for stone-tool-assisted consumption of animal tissues before 3.39 million years ago at Dikika, Ethiopia |journal=Nature |volume=466 |issue=7308 |pages=857–860 |date=August 2010 |pmid=20703305 |doi=10.1038/nature09248 |s2cid=4356816 |bibcode=2010Natur.466..857M |quote=The oldest direct evidence of stone tool manufacture comes from Gona (Ethiopia) and dates to between 2.6 and 2.5 million years (Myr) ago. [...] Here we report stone-tool-inflicted marks on bones found during recent survey work in Dikika, Ethiopia [... showing] unambiguous stone-tool cut marks for flesh removal [..., dated] to between 3.42 and 3.24 Myr ago [...] Our discovery extends by approximately 800,000 years the antiquity of stone tools and of stone-tool-assisted consumption of ungulates by hominins; furthermore, this behaviour can now be attributed to Australopithecus afarensis.}}</ref> LD 350-1, a fossil mandible fragment dated to 2.8 Mya, discovered in 2013 in Afar, Ethiopia, was described as combining "primitive traits seen in early ''Australopithecus'' with derived morphology observed in later ''Homo''.<ref>See: * {{cite journal |last1=Villmoare |first1=B. |last2=Kimbel |first2=W.H. |last3=Seyoum |first3=C. |last4=Campisano |first4=C.J. |last5=DiMaggio |first5=E.N. |last6=Rowan |first6=J. |last7=Braun |first7=D.R. |last8=Arrowsmith |first8=J.R. |last9=Reed |first9=K.E. |display-authors=6 |title=Paleoanthropology. Early Homo at 2.8 Ma from Ledi-Geraru, Afar, Ethiopia |journal=Science |volume=347 |issue=6228 |pages=1352–1355 |date=March 2015 |pmid=25739410 |doi=10.1126/science.aaa1343 |doi-access=free |bibcode=2015Sci...347.1352V}}.
See also: * {{cite journal |vauthors=DiMaggio EN, Campisano CJ, Rowan J, Dupont-Nivet G, Deino AL, Bibi F, Lewis ME, Souron A, Garello D, Werdelin L, Reed KE, Arrowsmith JR |display-authors=6 |title=Paleoanthropology. Late Pliocene fossiliferous sedimentary record and the environmental context of early Homo from Afar, Ethiopia |journal=Science |volume=347 |issue=6228 |pages=1355–1359 |date=March 2015 |pmid=25739409 |doi=10.1126/science.aaa1415 |doi-access=free |bibcode=2015Sci...347.1355D}}</ref> Some authors would push the development of ''Homo'' close to or even past 3 Mya.{{efn|{{harvp|Cela-Conde|Ayala|2003}} recognize five genera within Hominina: ''Ardipithecus'', ''Australopithecus'' (including ''Paranthropus''), ''Homo'' (including ''Kenyanthropus''), ''Praeanthropus'' (including ''Orrorin''), and ''Sahelanthropus''.<ref name="Cela-Conde-2003" />}} This finds support in a recent phylogenetic study in hominins that by using morphological, molecular and radiometric information, dates the emergence of ''Homo'' at 3.3 Mya (4.30 – 2.56 Mya).<ref name="research.ed.ac.uk">{{cite journal |last1=Püschel |first1=Hans P. |last2=Bertrand |first2=Ornella C. |last3=O'Reilly |first3=Joseph E. |last4=Bobe |first4=René |last5=Püschel |first5=Thomas A. |title=Divergence-time estimates for hominins provide insight into encephalization and body mass trends in human evolution |journal=Nature Ecology & Evolution |date=June 2021 |volume=5 |issue=6 |pages=808–819 |doi=10.1038/s41559-021-01431-1 |pmid=33795855 |bibcode=2021NatEE...5..808P |s2cid=232764044 |url=https://www.research.ed.ac.uk/en/publications/35151870-c7b5-477e-aca8-2c75c8382002|hdl=20.500.11820/35151870-c7b5-477e-aca8-2c75c8382002 |hdl-access=free }}</ref> Others have voiced doubt as to whether ''Homo habilis'' should be included in ''Homo'', proposing an origin of ''Homo'' with ''Homo erectus'' at roughly 1.9 Mya instead.<ref>{{cite journal |last=Wood |first=Bernard |date=28 June 2011 |title=Did early Homo migrate "out of" or "in to" Africa? |journal=Proceedings of the National Academy of Sciences |volume=108 |issue=26 |pages=10375–10376 |doi=10.1073/pnas.1107724108 |issn=0027-8424 |pmid=21677194 |pmc=3127876 |bibcode=2011PNAS..10810375W |quote=the adaptive coherence of ''Homo'' would be compromised if ''H. habilis'' is included in ''Homo''. Thus, if these arguments are accepted the origins of the genus ''Homo'' are coincident in time and place with the emergence of ''H. erectus'', not ''H. habilis''. |doi-access=free}}</ref>
The most salient physiological development between the earlier australopithecine species and ''Homo'' is the increase in endocranial volume (ECV), from about {{Convert|460|cm3|cuin|0|abbr=on}} in ''A. garhi'' to {{Convert|660|cm3|cuin|0|abbr=on}} in ''H. habilis'' and further to {{Convert|760|cm3|cuin|0|abbr=on}} in ''H. erectus'', {{Convert|1250|cm3|cuin|0|abbr=on}} in ''H. heidelbergensis'' and up to {{Convert|1760|cm3|cuin|0|abbr=on}} in ''H. neanderthalensis''. However, a steady rise in cranial capacity is observed already in ''Australopithecina'' and does not terminate after the emergence of ''Homo'', so that it does not serve as an objective criterion to define the emergence of the genus.<ref>{{cite journal|last1=Kimbel |first1=W.H. |last2=Villmoare |first2=B. |date=July 2016 |title=From Australopithecus to Homo: the transition that wasn't |journal=Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences |volume=371 |issue=1698 |article-number=20150248 |doi=10.1098/rstb.2015.0248 |pmc=4920303 |pmid=27298460 |quote=A fresh look at brain size, hand morphology and earliest technology suggests that a number of key ''Homo'' attributes may already be present in generalized species of Australopithecus, and that adaptive distinctions in ''Homo'' are simply amplifications or extensions of ancient hominin trends. [...] the adaptive shift represented by the ECV of ''Australopithecus'' is at least as significant as the one represented by the ECV of early ''Homo'', and that a major 'grade-level' leap in brain size with the advent of ''H. erectus'' is probably illusory.}}</ref>
=== ''Homo habilis'' === [[File:Homo ergaster reconstruction, American Museum of Natural History.jpg|thumb|Reconstruction of ''H. ergaster'', American Museum of Natural History]] ''Homo habilis'' emerged about 2.1 Mya. Already before 2010, there were suggestions that ''H. habilis'' should not be placed in the genus ''Homo'' but rather in ''Australopithecus''.<ref>{{Harvp|Wood|Richmond|2000|page=41|ps=: "A recent reassessment of cladistic and functional evidence concluded that there are few, if any, grounds for retaining ''H. habilis'' in ''Homo'', and recommended that the material be transferred (or, for some, returned) to Australopithecus (Wood & Collard, 1999)."}}</ref><ref name="Miller_2000">{{cite journal |last1=Miller |first1=J.M. |title=Craniofacial variation in Homo habilis: an analysis of the evidence for multiple species |journal=American Journal of Physical Anthropology |volume=112 |issue=1 |pages=103–128 |date=May 2000 |pmid=10766947 |doi=10.1002/(SICI)1096-8644(200005)112:1<103::AID-AJPA10>3.0.CO;2-6}}</ref> The main reason to include ''H. habilis'' in ''Homo'', its undisputed tool use, has become obsolete with the discovery of ''Australopithecus'' tool use at least a million years before ''H. habilis''.<ref name=Dikika/> Furthermore, ''H. habilis'' was long thought to be the ancestor of the more gracile ''Homo ergaster'' (''Homo erectus''). In 2007, it was discovered that ''H. habilis'' and ''H. erectus'' coexisted for a considerable time, suggesting that ''H. erectus'' is not immediately derived from ''H. habilis'' but instead from a common ancestor.<ref name="Spoor.et.al.2007" /> With the publication of Dmanisi skull 5 in 2013, it has become less certain that Asian ''H. erectus'' is a descendant of African ''H. ergaster'' which was in turn derived from ''H. habilis''. Instead, ''H. ergaster'' and ''H. erectus'' appear to be variants of the same species, which may have originated in either Africa or Asia<ref>{{cite journal |vauthors=Agustí J, Lordkipanidze D |date=June 2011 |title=How "African" was the early human dispersal out of Africa? |volume=30 |issue=11–12 |pages=1338–1342 |journal=Quaternary Science Reviews |doi=10.1016/j.quascirev.2010.04.012 |bibcode=2011QSRv...30.1338A}}</ref> and widely dispersed throughout Eurasia (including Europe, Indonesia, China) by 0.5 Mya.<ref>{{cite book |last1=Prins |first1=H.E. |last2=Walrath |first2=D. |last3=McBride |first3=B. |title=Evolution and prehistory: the human challenge |publisher=Wadsworth Publishing |year=2007 |page=162 |url=https://books.google.com/books?id=LfYirloa_rUC&pg=PA162 |isbn=978-0-495-38190-7}}.</ref>
=== ''Homo erectus'' === {{main|Homo erectus}} [[File:Le musée de préhistoire (Tautavel) (14498190867).jpg|thumb|''Homo heidelbergensis'' may have evolved from ''H. ergaster'', possibly following an intense population bottleneck 800,000 to 900,000 years ago.]] ''Homo erectus'' has often been assumed to have developed anagenetically from ''H. habilis'' from about 2 million years ago. This scenario was strengthened with the discovery of ''Homo erectus georgicus'', early specimens of ''H. erectus'' found in the Caucasus, which seemed to exhibit transitional traits with ''H. habilis.'' For example they showed increased cranial capacity from around 575 cm<sup>3</sup> in ''H. habilis'' to around 850 cm<sup>3</sup> in ''H. erectus'' {{Citation needed|date=July 2025|reason=Examples of these}}. As the earliest evidence for ''H. erectus'' was found outside of Africa, it was considered plausible that ''H. erectus'' developed in Eurasia and then migrated back to Africa. Based on fossils from the Koobi Fora Formation, east of Lake Turkana in Kenya, a 2007 study argued that ''H. habilis'' may have survived beyond the emergence of ''H. erectus'', so that the evolution of ''H. erectus'' would not have been anagenetical, and ''H. erectus'' would have existed alongside ''H. habilis'' for about half a million years ({{Mya|1.9|1.4}}), during the early Calabrian.<ref name="Spoor.et.al.2007"> {{cite journal|display-authors=6 |last1=Spoor |first1=F. |last2=Leakey |first2=M.G. |author-link2=Meave Leakey |last3=Gathogo |first3=P.N. |last4=Brown |first4=F.H. |last5=Antón |first5=S.C. |last6=McDougall |first6=I. |last7=Kiarie |first7=C. |last8=Manthi |first8=F.K. |last9=Leakey |first9=L.N. |date=August 2007 |title=Implications of new early Homo fossils from Ileret, east of Lake Turkana, Kenya |journal=Nature |volume=448 |issue=7154 |pages=688–691 |bibcode=2007Natur.448..688S |doi=10.1038/nature05986 |pmid=17687323 |quote=A partial maxilla assigned to H. habilis reliably demonstrates that this species survived until later than previously recognized, making an anagenetic relationship with H. erectus unlikely. The discovery of a particularly small calvaria of H. erectus indicates that this taxon overlapped in size with H. habilis, and may have shown marked sexual dimorphism. The new fossils confirm the distinctiveness of H. habilis and H. erectus, independently of overall cranial size, and suggest that these two early taxa were living broadly sympatrically in the same lake basin for almost half a million years. |s2cid=35845}}</ref> On 31 August 2023, researchers reported, based on genetic studies, that a human ancestor population bottleneck (from a possible 100,000 to 1,000 individuals) occurred "around 930,000 and 813,000 years ago ... lasted for about 117,000 years and brought human ancestors close to extinction."<ref name="NYT-20230831">{{cite news |last=Zimmer |first=Carl |author-link=Carl Zimmer |title=Humanity's Ancestors Nearly Died Out, Genetic Study Suggests - The population crashed following climate change about 930,000 years ago, scientists concluded. Other experts aren't convinced by the analysis. |url=https://www.nytimes.com/2023/08/31/science/human-survival-bottleneck.html |date=31 August 2023 |work=the New York Times |url-status=live |archive-url= https://archive.today/20230831182259/https://www.nytimes.com/2023/08/31/science/human-survival-bottleneck.html |archive-date=31 August 2023 |access-date=2 September 2023 }}</ref><ref name="SCI-20230831">{{cite journal |author=Hu, Wangjie |display-authors=et al. |title=Genomic inference of a severe human bottleneck during the Early to Middle Pleistocene transition |url=http://www.science.org/doi/10.1126/science.abq7487 |date=31 August 2023 |journal=Science |volume=381 |issue=6661 |pages=979–984 |doi=10.1126/science.abq7487 |pmid=37651513 |bibcode=2023Sci...381..979H |s2cid=261396309 |url-status=live |archive-url=https://archive.today/20230901024052/https://www.science.org/doi/10.1126/science.abq7487 |archive-date=1 September 2023 |access-date=2 September 2023 }}</ref>
Kenneth M. Weiss estimated that there have been about 44 billion individuals of the genus ''Homo'' from its origins to the evolution of ''H. erectus'', about 56 billion individuals from ''H. erectus'' to the Neolithic, and another 51 billion individuals since the Neolithic.<ref>{{cite journal |author=Weiss, Kenneth M. |title=On the Number of Members of the Genus Homo Who Have Ever Lived, and Some Evolutionary Implications |date=December 1984 |journal=Human Biology |volume=56 |issue=4 |pages=637–649 |jstor=41463610 |pmid=6442261 }}</ref>
A separate South African species ''Homo gautengensis'' was postulated as contemporary with ''H. erectus'' in 2010.<ref>{{cite journal |last1=Curnoe |first1=D. |title=A review of early Homo in southern Africa focusing on cranial, mandibular and dental remains, with the description of a new species (Homo gautengensis sp. nov.) |journal=Homo |volume=61 |issue=3 |pages=151–77 |date=June 2010 |pmid=20466364 |doi=10.1016/j.jchb.2010.04.002}}</ref>
== Phylogeny == {{Human timeline}} A taxonomy of ''Homo'' within the great apes is assessed as follows, with ''Paranthropus'' and ''Homo'' emerging within ''Australopithecus'' (shown here cladistically granting ''Paranthropus'', ''Kenyanthropus'', and ''Homo'').{{efn|name=fn1}}{{Efn|The line to the earliest members of ''Homo'' were derived from ''Australopithecus'', a genus that had separated from the chimpanzee–human last common ancestor by late Miocene or early Pliocene times.<ref name="Schuster" /> <!--why is the entire Chimpanzee–human last common ancestor article reproduced in this footnote? with date estimates by several specialists ranging from 13 million years ago to more recently than six million years ago. * {{cite journal |last1=Arnason |first1=U. |last2=Gullberg |first2=A. |last3=Janke |first3=A. |title=Molecular timing of primate divergences as estimated by two nonprimate calibration points |journal=Journal of Molecular Evolution |volume=47 |issue=6 |pages=718–727 |date=December 1998 |pmid=9847414 |doi=10.1007/PL00006431 |bibcode=1998JMolE..47..718A}} * {{cite journal | vauthors = Patterson N, Richter DJ, Gnerre S, Lander ES, Reich D |title=Genetic evidence for complex speciation of humans and chimpanzees |journal=Nature | volume = 441 | issue = 7097 | pages = 1103–8 | date = June 2006 | pmid = 16710306 |doi=10.1038/nature04789 |bibcode=2006Natur.441.1103P}} * {{cite journal | vauthors = Wakeley J | title = Complex speciation of humans and chimpanzees |journal=Nature | volume = 452 | issue = 7184 |pages=E3-4; discussion E4 |date=March 2008 | pmid = 18337768 | doi = 10.1038/nature06805 |bibcode=2008Natur.452....3W |quote=Patterson et al. suggest that the apparently short divergence time between humans and chimpanzees on the X chromosome is explained by a massive interspecific hybridization event in the ancestry of these two species. However, Patterson et al. do not statistically test their own null model of simple speciation before concluding that speciation was complex, and—even if the null model could be rejected—they do not consider other explanations of a short divergence time on the X chromosome. These include natural selection on the X chromosome in the common ancestor of humans and chimpanzees, changes in the ratio of male-to-female mutation rates over time, and less extreme versions of divergence with gene flow. I therefore believe that their claim of hybridization is unwarranted.}} see current estimates regarding complex speciation.-->|name=fn2}}<ref name="Indriati-2011" /><ref name="Berger-2017" /><ref name="Schuster">{{cite journal |vauthors=Schuster AM |year=1997 |title=Earliest Remains of Genus ''Homo'' |url=https://archive.archaeology.org/9701/newsbriefs/homo.html |journal=Archaeology |volume=50 |access-date=5 March 2015 |number=1}}</ref><ref name="Saylor 2015 483–488"/><ref name="Mondal-2019">{{cite journal |last1=Mondal |first1=M. |last2=Bertranpetit |first2=J. |last3=Lao |first3=O. |title=Approximate Bayesian computation with deep learning supports a third archaic introgression in Asia and Oceania |journal=Nature Communications |volume=10 |issue=1 |page=246 |date=January 2019 |pmid=30651539 |pmc=6335398 |doi=10.1038/s41467-018-08089-7 |bibcode=2019NatCo..10..246M}}</ref><ref name="Zeitoun 2003 148–156">{{cite journal |last=Zeitoun |first=V. |title=High occurrence of a basicranial feature in Homo erectus: anatomical description of the preglenoid tubercle |journal=The Anatomical Record Part B: The New Anatomist |volume=274 |issue=1 |pages=148–156 |date=September 2003 |pmid=12964205 |doi=10.1002/ar.b.10028 |doi-access=free}}</ref><ref name="Proceedings 2015">{{cite journal |vauthors = Dembo M, Matzke NJ, Mooers AØ, Collard M |title=Bayesian analysis of a morphological supermatrix sheds light on controversial fossil hominin relationships |journal=Proceedings. Biological Sciences |volume=282 |issue=1812 |article-number=20150943 |date=August 2015 |pmid=26202999 |pmc=4528516 |doi=10.1098/rspb.2015.0943 |bibcode=2015PBioS.28250943D }}</ref><ref name="Dembo-2016">{{cite journal | vauthors = Dembo M, Radovčić D, Garvin HM, Laird MF, Schroeder L, Scott JE, Brophy J, Ackermann RR, Musiba CM, de Ruiter DJ, Mooers AØ, Collard M | display-authors = 6 | title = The evolutionary relationships and age of Homo naledi: An assessment using dated Bayesian phylogenetic methods |journal=Journal of Human Evolution |volume=97 |pages=17–26 |date=August 2016 |pmid=27457542 | doi = 10.1016/j.jhevol.2016.04.008 | bibcode = 2016JHumE..97...17D |hdl-access=free |hdl=2164/8796}}</ref><ref name="Ko-2016"/><ref>{{cite book|url=https://books.google.com/books?id=bL2XDwAAQBAJ&q=Sahelanthropus+tchadensis&pg=PA7 |title=The Origins of Europeans and Their Pre-Historic Innovations from 6 Million to 10,000 BCE: From 6 Million to 10,000 BCE |last1=Harrison |first1=N. |date=1 May 2019 |publisher=Algora Publishing |isbn=978-1-62894-379-5}}</ref><ref>See: * {{cite book|url=https://www.researchgate.net/publication/278682043 |title=Analyzing Hominin Hominin Phylogeny: Cladistic Approach |last1=Strait |first1=David |last2=Grine |first2=Frederick |last3=Fleagle |first3=John |name-list-style=vanc |date=2015 |isbn=978-3-642-39978-7 |pages=1989–2014 (cladogram p. 2006)|publisher=Springer }}. * {{cite journal|last1=Mounier |first1=A. |last2=Caparros |first2=M. |date=2015 |title=The phylogenetic status of Homo heidelbergensis – a cladistic study of Middle Pleistocene hominins |journal=BMSAP |language=fr |volume=27 |issue=3–4 |pages=110–134 |doi=10.1007/s13219-015-0127-4 |s2cid=17449909 |issn=0037-8984}} * {{cite journal | vauthors = Rogers AR, Harris NS, Achenbach AA | title = Neanderthal-Denisovan ancestors interbred with a distantly related hominin | journal = Science Advances | volume = 6 |issue=8 |article-number=eaay5483 |date=February 2020 |pmid=32128408 |pmc=7032934 | doi = 10.1126/sciadv.aay5483 | doi-access = free |bibcode=2020SciA....6.5483R}}</ref>{{Excessive citations inline|reason=Over four citations inline, unbundled|date=December 2021}} The exact phylogeny within ''Australopithecus'' is still highly controversial. Approximate radiation dates of daughter clades are shown in millions of years ago (Mya).<ref name="research.ed.ac.uk"/><ref name="Dembo-2016"/> ''Sahelanthropus'' and ''Orrorin'', possibly sisters to ''Australopithecus'', are not shown here. The naming of groupings is sometimes muddled as often certain groupings are presumed before any cladistic analysis is performed.<ref name="Zeitoun 2003 148–156"/>
{{clade|{{clade |1=Hylobatidae (gibbons) |label2=Hominidae |sublabel2=(15.7) |2={{clade |1=Ponginae (orangutans) |label2=Homininae |sublabel2=(8.8) |2={{clade |1=Gorillini (gorillas) |label2=Hominini |sublabel2=(7.5) |2={{clade |1=Panina (chimpanzees) |2=Australopithecines (incl. ''Australopithecus'', ''Kenyanthropus'', ''Paranthropus'', ''Homo'') }} }} }} }} |label1=Hominoidea |sublabel1=(20.4 Mya) }} {{clear}} Cladogram based on Dembo et al. (2016) with ages of last occurrence based on Wood & Boyle (2016):<ref name="Dembo-2016" /><ref>{{Cite journal |last1=Wood |first1=Bernard |last2=K. Boyle |first2=Eve |date=2016 |title=Hominin taxic diversity: Fact or fantasy? |url=https://onlinelibrary.wiley.com/doi/abs/10.1002/ajpa.22902 |journal=American Journal of Physical Anthropology |language=en |volume=159 |issue=S61 |pages=37–78 |doi=10.1002/ajpa.22902 |bibcode=2016AJPA..159S..37W |issn=1096-8644}}</ref>{{clade|{{clade |1=''Ardipithecus ramidus'' (†4.3) |sublabel2=(5.5) |label2=''Australopithecus s.l.'' |2={{clade |1=''Australopithecus anamensis s.s.'' (†3.9) |2={{clade |1={{clade |1=''Australopithecus afarensis'' (†3.0) |2=''Australopithecus garhi'' (†2.4) }} |2={{clade |1=''Kenyanthropus platyops'' (†3.3) |2={{clade |1={{clade |1=''Australopithecus africanus'' (†2.4) |2=''Paranthropus'' (†1.0) }} |label2=''Homo'' |sublabel2=(3.3) |2={{clade |1=''Homo floresiensis'' (†0.01) |2={{clade |1={{clade |1=''Homo habilis'' (†1.6) 45 px |2=''Australopithecus sediba'' (†2.0) }} |2={{clade |1=''Homo rudolfensis'' (†2.0) 41 px |2={{clade |1=Georgian ''H. erectus'' (†1.8) |2={{clade |1=African ''Homo erectus'' (†1.4) 45 px |2={{clade |1=Asian H. erectus (†0.03) 45 px |2={{clade |1=''Homo naledi'' (†0.2) |2={{clade |1=''Homo antecessor'' (†0.9) 45px |2={{clade |1='''''Homo sapiens'' 39 px''' |2={{clade |1=''Homo heidelbergensis'' (†0.1) |2=''H. neanderthalensis'' (†0.04) 39 px }} }} }} }} }} }} }} }} }} }} }} }} }} }} }} |label1=Australopithecines|sublabel1=(7.3 Mya) }}
Cladogram based on Feng et al. (2025):<ref name=":0">{{Cite journal |last1=Feng |first1=Xiaobo |last2=Yin |first2=Qiyu |last3=Gao |first3=Feng |last4=Lu |first4=Dan |last5=Fang |first5=Qin |last6=Feng |first6=Yilu |last7=Huang |first7=Xuchu |last8=Tan |first8=Chen |last9=Zhou |first9=Hanwen |last10=Li |first10=Qiang |last11=Zhang |first11=Chi |last12=Stringer |first12=Chris |last13=Ni |first13=Xijun |date=2025-09-25 |title=The phylogenetic position of the Yunxian cranium elucidates the origin of Homo longi and the Denisovans |url=https://www.science.org/doi/10.1126/science.ado9202 |journal=Science |volume=389 |issue=6767 |pages=1320–1324 |doi=10.1126/science.ado9202 |pmid=40997177 |bibcode=2025Sci...389.1320F }}</ref>{{Efn|Ages as listed in the supplementary materials for Feng et al. (2025)}} {{Clade |label1='''''Homo''''' (2.84) |1={{Clade |1=''Homo habilis'' (†1.78 Mya) 45 px |label2=''Homo erectus'' s.l. (2.6) |2={{clade |1=Stw53 (†1.9) |label2=(2.3) |2={{clade |1={{clade |1=Turkana (†1.53) 45 px |2={{clade |1=Dmanisi (†1.77) |2=Olduvai Hominids (†1.47) }} }} |label2=(2.0) |2={{clade |1=''Homo naledi'' (†0.33) |label2= (1.8) |2={{clade |1=''Homo floresiensis'' (†0.05) |2={{clade |1={{clade |1={{clade |1=Sambungmacan (†0.2) |2=Ngandong (†0.1) }} |2={{clade |1={{clade |1=Peking Man (†0.28) 45 px |2=Nanjing Man (†0.58) }} |2={{clade |1=Hexian (†0.39) |2=Sangiran (†1.3) }} }} }} |label2=(1.6) |2={{clade |1=''Homo heidelbergensis'' (†0.15) |label2=(1.4) |2={{clade |1=''H. neanderthalensis'' (†0.04) 39 px |label2=(1.3) |2={{clade |1='''''Homo sapiens''''' 39 px |label2=(1.3) |2={{clade |1=''H. antecessor'' (†0.77) 45px |2=''H. longi'' (†0.05) 60 px }} }} }} }} }} }} }} }} }} }} }}
Several of the ''Homo'' lineages 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, perhaps ''H. erectus'', may have interbred into the Denisovans about 55,000 years ago.<ref>See: * {{cite journal|last1=Dediu |first1=D. |last2=Levinson |first2=S.C. |date=1 June 2018 |title=Neanderthal language revisited: not only us |journal=Current Opinion in Behavioral Sciences |series=The Evolution of Language |volume=21 |pages=49–55 |doi=10.1016/j.cobeha.2018.01.001 |issn=2352-1546 |hdl=21.11116/0000-0000-1667-4 |s2cid=54391128 |hdl-access=free}} * {{cite journal |last1=Hubisz |first1=M.J. |last2=Williams |first2=A.L. |last3=Siepel |first3=A. |title=Mapping gene flow between ancient hominins through demography-aware inference of the ancestral recombination graph |journal=PLOS Genetics |volume=16 |issue=8 |article-number=e1008895 |date = August 2020 |pmid=32760067 |pmc=7410169 |doi=10.1371/journal.pgen.1008895 |doi-access=free }} * {{cite journal |vauthors=Kuhlwilm M, Gronau I, Hubisz MJ, de Filippo C, Prado-Martinez J, Kircher M, Fu Q, Burbano HA, Lalueza-Fox C, de la Rasilla M, Rosas A, Rudan P, Brajkovic D, Kucan Ž, Gušic I, Marques-Bonet T, Andrés AM, Viola B, Pääbo S, Meyer M, Siepel A, Castellano S |display-authors=6 |title=Ancient gene flow from early modern humans into Eastern Neanderthals |journal=Nature |volume=530 |issue=7591 |pages=429–33 |date=February 2016 |pmid=26886800 |pmc=4933530 |doi=10.1038/nature16544 |bibcode=2016Natur.530..429K}} * {{cite journal |vauthors=Prüfer K, Racimo F, Patterson N, Jay F, Sankararaman S, Sawyer S, Heinze A, Renaud G, Sudmant PH, de Filippo C, Li H, Mallick S, Dannemann M, Fu Q, Kircher M, Kuhlwilm M, Lachmann M, Meyer M, Ongyerth M, Siebauer M, Theunert C, Tandon A, Moorjani P, Pickrell J, Mullikin JC, Vohr SH, Green RE, Hellmann I, Johnson PL, Blanche H, Cann H, Kitzman JO, Shendure J, Eichler EE, Lein ES, Bakken TE, Golovanova LV, Doronichev VB, Shunkov MV, Derevianko AP, Viola B, Slatkin M, Reich D, Kelso J, Pääbo S |display-authors=6 |title=The complete genome sequence of a Neanderthal from the Altai Mountains |journal=Nature |volume=505 |issue=7481 |pages=43–9 |date=January 2014 |pmid=24352235 |pmc=4031459 |doi=10.1038/nature12886 |bibcode=2014Natur.505...43P}}</ref><ref name="Mondal-2019" /><ref name="Callaway 2016">{{cite journal |last=Callaway |first=E. |title=Evidence mounts for interbreeding bonanza in ancient human species |url=http://www.nature.com/news/evidence-mounts-for-interbreeding-bonanza-in-ancient-human-species-1.19394 |journal=Nature News |doi=10.1038/nature.2016.19394 |year=2016 |s2cid=87029139|url-access=subscription }}</ref> Fossil evidence shows ''H. erectus'' s.s. survived at least until 117,000 yrs ago, and the even more basal ''H. floresiensis'' survived until 50,000 years ago. A 1.5-million-year ''H. erectus''-like lineage appears to have made its way into modern humans through the Denisovans and specifically into the Papuans and aboriginal Australians.<ref name="Mondal-2019" /> 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.<ref>{{cite journal |last1=Varki |first1=A. |title=Why are there no persisting hybrids of humans with Denisovans, Neanderthals, or anyone else? |journal=Proceedings of the National Academy of Sciences of the United States of America |volume=113 |issue=17 |pages=E2354 |date=April 2016 |pmid=27044111 |pmc=4855598 |doi=10.1073/pnas.1602270113 |bibcode=2016PNAS..113E2354V |doi-access=free}}</ref><ref name="Callaway 2016" /> 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.<ref name="Ko-2016">{{cite journal |last1=Ko |first1=K.H. |title=Hominin interbreeding and the evolution of human variation |journal=Journal of Biological Research |volume=23 |issue=1 |article-number=17 |date=December 2016 |pmid=27429943 |pmc=4947341 |doi=10.1186/s40709-016-0054-7 |doi-access=free }}</ref><ref>{{cite journal |vauthors=Pickrell JK, Patterson N, Loh PR, Lipson M, Berger B, Stoneking M, Pakendorf B, Reich D |display-authors=6 |title=Ancient west Eurasian ancestry in southern and eastern Africa |journal=Proceedings of the National Academy of Sciences of the United States of America |volume=111 |issue=7 |pages=2632–2637 |date=February 2014 |pmid=24550290 |pmc=3932865 |doi=10.1073/pnas.1313787111 |arxiv=1307.8014 |bibcode=2014PNAS..111.2632P |doi-access=free}}</ref>
Some evidence suggests that ''Australopithecus sediba'' could be moved to the genus ''Homo'', or placed in its own genus, due to its position with respect to e.g. ''H. habilis'' and ''H. floresiensis''.<ref name="Proceedings 2015" /><ref>{{cite journal |vauthors=Groves C |date=2017 |title=Progress in human systematics. A review |url=http://www.francoangeli.it/riviste/Scheda_Riviste.asp?IDArticolo=59711 |journal=Paradigmi |issue=2 |pages=59–74 |article-number=5 |doi=10.3280/PARA2017-002005 |issn=1120-3404|url-access=subscription }}</ref>
=== Dispersal === {{see also|Early expansions of hominins out of Africa|Interbreeding between archaic and modern humans|Early human migrations}}
By about 1.8 million years ago, ''H. erectus'' is present in both East Africa (''H. ergaster'') and in Western Asia (''H. georgicus''). The ancestors of Indonesian ''H. floresiensis'' may have left Africa even earlier.{{Efn|name=efn4|In a 2015 phylogenetic study, ''H. floresiensis'' was placed with ''Australopithecus sediba'', ''H. habilis'' and Dmanisi Man, raising the possibility that the ancestors of ''H. floresiensis'' left Africa before the appearance of ''H. erectus'', possibly even becoming the first hominins to do so and evolved further in Asia.<ref name="Proceedings 2015" />}}<ref name="Proceedings 2015" />
[[File:Spreading homo sapiens la.svg|thumb|upright=1.5|Successive dispersals of {{color box|yellow||}} ''Homo erectus'' (yellow), {{color box|#e4ca30}} ''H. neanderthalensis'' (ochre) and {{color box|#e9252c}} ''H. sapiens'' (red)]]
''Homo erectus'' and related or derived archaic human species over the next 1.5 million years spread throughout Africa and Eurasia<ref>{{cite journal |last=Beyin |first=A. |title=Upper Pleistocene Human Dispersals out of Africa: A Review of the Current State of the Debate |journal=International Journal of Evolutionary Biology |volume=2011 |article-number=615094 |year=2011 |pmid=21716744 |pmc=3119552 |doi=10.4061/2011/615094 |doi-access=free }}</ref><ref>{{cite journal |last=Callaway |first=E. |title=Oldest ancient-human DNA details dawn of Neanderthals |journal=Nature |volume=531 |issue=7594 |page=286 |date=March 2016 |pmid=26983523 |doi=10.1038/531286a |s2cid=4459329 |bibcode=2016Natur.531..296C |doi-access=free}}</ref> (see: Recent African origin of modern humans). Europe is reached by about 0.5 Mya by ''Homo heidelbergensis''.
''Homo neanderthalensis'' and ''H. sapiens'' develop after about 300 kya. ''Homo naledi'' is present in Southern Africa by 300 kya.
''H. sapiens'' soon after its first emergence spread throughout Africa, and to Western Asia in several 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 a ''H. sapiens'' and 170,000 year old remains of a ''H. neanderthalensis'' in Apidima Cave, Peloponnese, Greece, more than 150,000 years older than previous ''H. sapiens'' finds in Europe.<ref name="NYT-20190710">{{cite news |last=Zimmer |first=C. |author-link=Carl Zimmer |title=A Skull Bone Discovered in Greece May Alter the Story of Human Prehistory - The bone, found in a cave, is the oldest modern human fossil ever discovered in Europe. It hints that humans began leaving Africa far earlier than once thought. |url=https://www.nytimes.com/2019/07/10/science/skull-neanderthal-human-europe-greece.html |date=10 July 2019 |work=The New York Times |access-date=11 July 2019}}</ref><ref name="PHYS-20190710">{{cite news |author=Staff |title='Oldest remains' outside Africa reset human migration clock |url=https://phys.org/news/2019-07-oldest-africa-reset-human-migration.html |date=10 July 2019 |work=Phys.org |access-date=10 July 2019}}</ref><ref name="NAT-20190710">{{cite journal |last1=Harvati |first1=K. |author-link1=Katerina Harvati |last2=Röding |first2=C. |last3=Bosman |first3=A.M. |last4=Karakostis |first4=F.A. |last5=Grün |first5=R. |last6=Stringer |first6=C. |author-link6=Chris Stringer |last7=Karkanas |first7=P. |last8=Thompson |first8=N.C. |last9=Koutoulidis |first9=V. |last10=Moulopoulos |first10=L.A. |last11=Gorgoulis |first11=V.G. |last12=Kouloukoussa |first12=M. |display-authors=6 |title=Apidima Cave fossils provide earliest evidence of Homo sapiens in Eurasia |journal=Nature |volume=571 |issue=7766 |pages=500–504 |date=July 2019 |pmid=31292546 |doi=10.1038/s41586-019-1376-z |bibcode=2019Natur.571..500H |s2cid=195873640|url=https://zenodo.org/record/6646855|hdl=10072/397334 |hdl-access=free }}</ref>
Most notable is the Southern Dispersal of ''H. sapiens'' around 60 kya, which led to the lasting peopling of Oceania and Eurasia by anatomically modern humans.<ref name="A draft sequence of the Neandertal"/> ''H. sapiens'' interbred with archaic humans both in Africa and in Eurasia, in Eurasia notably with Neanderthals and Denisovans.<ref>{{cite journal |vauthors=Reich D, Green RE, Kircher M, Krause J, Patterson N, Durand EY, Viola B, Briggs AW, Stenzel U, Johnson PL, Maricic T, Good JM, Marques-Bonet T, Alkan C, Fu Q, Mallick S, Li H, Meyer M, Eichler EE, Stoneking M, Richards M, Talamo S, Shunkov MV, Derevianko AP, Hublin JJ, Kelso J, Slatkin M, Pääbo S |author-link28=Svante Pääbo |display-authors=6 |title=Genetic history of an archaic hominin group from Denisova Cave in Siberia |journal=Nature |volume=468 |issue=7327 |pages=1053–1060 |date=December 2010 |pmid=21179161 |pmc=4306417 |doi=10.1038/nature09710 |hdl=10230/25596 |url=http://pubman.mpdl.mpg.de/pubman/item/escidoc:1552828/component/escidoc:2261391/Reich_Genetic_Nature_2010.pdf |bibcode=2010Natur.468.1053R}}</ref><ref>{{cite journal |vauthors=Reich D, Patterson N, Kircher M, Delfin F, Nandineni MR, Pugach I, Ko AM, Ko YC, Jinam TA, Phipps ME, Saitou N, Wollstein A, Kayser M, Pääbo S, Stoneking M |author-link14=Svante Pääbo |display-authors=6 |title=Denisova admixture and the first modern human dispersals into Southeast Asia and Oceania |journal=American Journal of Human Genetics |volume=89 |issue=4 |pages=516–528 |date=October 2011 |pmid=21944045 |pmc=3188841 |doi=10.1016/j.ajhg.2011.09.005}}</ref>
Among extant populations of ''H. sapiens'', the deepest temporal division is found in the San people of Southern Africa, estimated at close to 130,000 years,<ref name="National Academy of Sciences">{{cite journal |vauthors=Henn BM, Gignoux CR, Jobin M, Granka JM, Macpherson JM, Kidd JM, Rodríguez-Botigué L, Ramachandran S, Hon L, Brisbin A, Lin AA, Underhill PA, Comas D, Kidd KK, Norman PJ, Parham P, Bustamante CD, Mountain JL, Feldman MW |display-authors=6 |title=Hunter-gatherer genomic diversity suggests a southern African origin for modern humans |journal=Proceedings of the National Academy of Sciences of the United States of America |volume=108 |issue=13 |pages=5154–5162 |date=March 2011 |pmid=21383195 |pmc=3069156 |doi=10.1073/pnas.1017511108 |bibcode=2011PNAS..108.5154H |doi-access=free}}</ref> or possibly more than 300,000 years ago.<ref>{{cite journal |vauthors=Schlebusch CM, Malmström H, Günther T, Sjödin P, Coutinho A, Edlund H, Munters AR, Vicente M, Steyn M, Soodyall H, Lombard M, Jakobsson M |display-authors=6 |title=Southern African ancient genomes estimate modern human divergence to 350,000 to 260,000 years ago |journal=Science |volume=358 |issue=6363 |pages=652–655 |date=November 2017 |pmid=28971970 |doi=10.1126/science.aao6266 |doi-access=free |bibcode=2017Sci...358..652S}}</ref> Temporal division among non-Africans is of the order of 60,000 years in the case of Australo-Melanesians. Division of Europeans and East Asians is of the order of 50,000 years, with repeated and significant admixture events throughout Eurasia during the Holocene.
Archaic human species may have survived until the beginning of the Holocene, although they were mostly extinct or absorbed by the expanding ''H. sapiens'' populations by 40 kya (Neanderthal extinction).
== List of lineages == {{see also|List of human evolution fossils}} The species status of ''H. rudolfensis'', ''H. ergaster'', ''H. georgicus'', ''H. antecessor'', ''H. cepranensis'', ''H. rhodesiensis'', ''H. neanderthalensis'', Denisova hominin, and ''H. floresiensis'' remain under debate. ''H. heidelbergensis'' and ''H. neanderthalensis'' are closely related to each other and have been considered to be subspecies of ''H. 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 in ''Homo'' questionable), ''H. erectus'' (1.8–0.1 Mya, including the majority of the age of the genus, and the majority of archaic varieties as subspecies,<ref>{{cite news |first=Sid |last=Perkins |url=http://www.nature.com/news/skull-suggests-three-early-human-species-were-one-1.13972 |title=Skull suggests three early human species were one |date=17 October 2013 |work=Nature News & Comment}}</ref><ref> {{cite news |last=Switek |first=B. |date=17 October 2013 |title=Beautiful Skull Spurs Debate on Human History |url=http://news.nationalgeographic.com/news/2013/10/131017-skull-human-origins-dmanisi-georgia-erectus/ |archive-url=https://web.archive.org/web/20131017184738/http://news.nationalgeographic.com/news/2013/10/131017-skull-human-origins-dmanisi-georgia-erectus/ |archive-date=17 October 2013 |newspaper=National Geographic |access-date=22 September 2014}}</ref><ref> {{cite journal |vauthors=Lordkipanidze D, Ponce de León MS, Margvelashvili A, Rak Y, Rightmire GP, Vekua A, Zollikofer CP |title=A complete skull from Dmanisi, Georgia, and the evolutionary biology of early Homo |journal=Science |volume=342 |issue=6156 |pages=326–31 |date=October 2013 |pmid=24136960 |doi=10.1126/science.1238484 |s2cid=20435482 |bibcode=2013Sci...342..326L}} </ref> including ''H. heidelbergensis'' as a late or transitional variety<ref>{{cite web |title=''Homo heidelbergensis'' - The evolutionary dividing line between ''Homo erectus'' and modern humans was not sharp. |publisher=Dennis O'Neil |url=http://anthro.palomar.edu/homo2/mod_homo_1.htm |access-date=29 November 2015 |archive-date=9 March 2016 |archive-url=https://web.archive.org/web/20160309161650/http://humanorigins.si.edu/resources/whats-hot/mystery-pit-bones-atapuerca-spain }}</ref><ref> {{cite journal |last1=Mounier |first1=A. |last2=Marchal |first2=F. |last3=Condemi |first3=S. |title=Is Homo heidelbergensis a distinct species? New insight on the Mauer mandible |journal=Journal of Human Evolution |volume=56 |issue=3 |pages=219–246 |date=March 2009 |pmid=19249816 |doi=10.1016/j.jhevol.2008.12.006 |bibcode=2009JHumE..56..219M |url=https://www.researchgate.net/publication/24144485}}</ref><ref> {{cite journal |last1=Lieberman |first1=D.E. |last2=McBratney |first2=B.M. |last3=Krovitz |first3=G. |title=The evolution and development of cranial form in Homosapiens |journal=Proceedings of the National Academy of Sciences of the United States of America |volume=99 |issue=3 |pages=1134–1139 |date=February 2002 |pmid=11805284 |pmc=122156 |doi=10.1073/pnas.022440799 |bibcode=2002PNAS...99.1134L |doi-access=free}}</ref>) and ''Homo sapiens'' (300 kya to present, including ''H. neanderthalensis'' and other varieties as subspecies). 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,<ref>{{cite journal |last1=Wilson |first1=A.C. |last2=Maxson |first2=L.R. |last3=Sarich |first3=V.M. |title=Two types of molecular evolution. Evidence from studies of interspecific hybridization |journal=Proceedings of the National Academy of Sciences of the United States of America |volume=71 |issue=7 |pages=2843–2847 |date=July 1974 |pmid=4212492 |pmc=388568 |doi=10.1073/pnas.71.7.2843 |bibcode=1974PNAS...71.2843W |doi-access=free}}</ref><ref>{{cite journal|last1=Popadin |first1=K. |last2=Gunbin |first2=K. |last3=Peshkin |first3=L. |last4=Annis |first4=S. |last5=Kraytsberg |first5=Y. |last6=Markuzon |first6=N. |last7=Ackermann |first7=R.R. |last8=Khrapko |first8=K. |display-authors=6 |title=Mitochondrial Pseudogenes Suggest Repeated Inter-Species Hybridization among Direct Human Ancestors |journal=Genes |date=2022 |volume=13 |issue=5 |at=134502 |doi=10.3390/genes13050810 |doi-access=free |pmid=35627195 |pmc=9140377 |biorxiv=10.1101/134502 }}</ref> so all contemporary "species" in the genus ''Homo'' would potentially have been able to interbreed at the time, and introgression from beyond the genus ''Homo'' can not ''a priori'' be ruled out.<ref>{{cite journal |last1=Ackermann |first1=R.R. |last2=Arnold |first2=M.L. |last3=Baiz |first3=M.D. |last4=Cahill |first4=J.A. |last5=Cortés-Ortiz |first5=L. |last6=Evans |first6=B.J. |last7=Grant |first7=B.R. |last8=Grant |first8=P.R. |last9=Hallgrimsson |first9=B. |last10=Humphreys |first10=R.A. |last11=Jolly |first11=C.J. |last12=Malukiewicz |first12=J. |last13=Percival |first13=C.J. |last14=Ritzman |first14=T.B. |last15=Roos |first15=C. |last16=Roseman |first16=C.C. |last17=Schroeder |first17=L. |last18=Smith |first18=F.H. |last19=Warren |first19=K.A. |last20=Wayne |first20=R.K. |last21=Zinner |first21=D. |display-authors=6 |title=Hybridization in human evolution: Insights from other organisms |journal=Evolutionary Anthropology |volume=28 |issue=4 |pages=189–209 |date=July 2019 |pmid=31222847 |pmc=6980311 |doi=10.1002/evan.21787 |url=http://osf.io/y3bp7/ |hdl=2027.42/151330}}</ref> It has been suggested that ''H. naledi'' may have been a hybrid with a late surviving ''Australipith'' (taken to mean beyond ''Homo'', ed.),<ref name="Berger-2017">{{cite journal |last1=Berger |first1=L.R. |last2=Hawks |first2=J. |last3=Dirks |first3=P.H. |last4=Elliott |first4=M. |last5=Roberts |first5=E.M. |title=Homo naledi and Pleistocene hominin evolution in subequatorial Africa |journal=eLife |volume=6 |article-number=e24234 |date=May 2017 |pmid=28483041 |pmc=5423770 |doi=10.7554/eLife.24234 |doi-access=free |editor-last=Perry |editor-first=G.H.}}</ref> 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.
{| class="wikitable sortable" |+ Comparative table of ''Homo'' lineages |- ! Lineages !data-sort-type=number| Temporal range<br />(kya) ! Habitat !data-sort-type=number| Adult height !data-sort-type=number| Adult mass !data-sort-type=number| Cranial capacity<br />(cm<sup>3</sup>) !data-sort-type=number| Fossil record !data-sort-type=number| Discovery/<br />publication<br />of name |- | ''H. habilis''<br /><small>membership in ''Homo'' uncertain</small> | 2,100–1,500{{efn|Confirmed ''H. habilis'' fossils are dated to between 2.1 and 1.5 million years ago. This date range overlaps with the emergence of ''Homo erectus''.<ref>{{cite book |last1=Schrenk |first1=Friedemann |last2=Kullmer |first2=Ottmar |last3=Bromage |first3=Timothy |name-list-style=vanc |year=2007 |editor-last1=Henke |editor-first1=Winfried |editor-last2=Tattersall |editor-first2=Ian |editor-link2=Ian Tattersall |title=Handbook of Paleoanthropology |volume=1 |others=In collaboration with Thorolf Hardt |chapter=9 the Earliest Putative Homo Fossils |location=Berlin, Heidelberg |publisher=Springer |pages=1611–1631 |doi=10.1007/978-3-540-33761-4_52 |isbn=978-3-540-32474-4}}</ref><ref>{{cite journal |last1=DiMaggio |first1=E.N. |last2=Campisano |first2=C.J. |last3=Rowan |first3=J. |last4=Dupont-Nivet |first4=G. |last5=Deino |first5=A.L. |last6=Bibi |first6=F. |last7=Lewis |first7=M.E. |last8=Souron |first8=A. |last9=Garello |first9=D. |last10=Werdelin |first10=L. |last11=Reed |first11=K.E. |last12=Arrowsmith |first12=J.R. |display-authors=6 |title=Paleoanthropology. Late Pliocene fossiliferous sedimentary record and the environmental context of early Homo from Afar, Ethiopia |journal=Science |volume=347 |issue=6228 |pages=1355–9 |date=March 2015 |pmid=25739409 |doi=10.1126/science.aaa1415 |bibcode=2015Sci...347.1355D |s2cid=43455561|doi-access=free }}</ref>}}{{efn|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 between ''Australopithecus'' and ''Homo'' discovered in 2015.}} | Tanzania | 110–140 cm (3 ft 7 in – 4 ft 7 in) | {{convert|33|–|55|kg|lb|abbr=on}} | 510–660 |data-sort-value=50| Many | 1960<br/>1964 |- | ''H. rudolfensis''<br /><small>membership in ''Homo'' uncertain</small> | 1,900 | Kenya | | | 700 |data-sort-value=2| 2 sites | 1972<br/>1986 |- | ''H. gautengensis''<br /><small>also classified as ''H. habilis''</small> | 1,900–600 | South Africa | 100 cm (3 ft 3 in) | | |data-sort-value=3| 3 individuals<ref>{{cite journal |last1=Curnoe |first1=D. |title=A review of early Homo in southern Africa focusing on cranial, mandibular and dental remains, with the description of a new species (Homo gautengensis sp. nov.) |journal=Homo |volume=61 |issue=3 |pages=151–177 |date=June 2010 |pmid=20466364 |doi=10.1016/j.jchb.2010.04.002}}</ref>{{efn|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 as ''H. habilis'', ''H. ergaster'' or ''Australopithecus'' by other anthropologists.}} | 2010<br/>2010 |- | ''H. erectus'' | 2,000–140<ref>{{cite book |last1=Haviland |first1=William A. |last2=Walrath |first2=Dana |last3=Prins |first3=Harald E.L. |author-link3=Harald E. L. Prins |last4=McBride |first4=Bunny |name-list-style=vanc |year=2007 |title=Evolution and Prehistory: The Human Challenge |url=https://books.google.com/books?id=LfYirloa_rUC&pg=PA162 |edition=8th |location=Belmont, CA |publisher=Thomson Wadsworth |page=162 |isbn=978-0-495-38190-7}}</ref>{{efn|''H. erectus'' may 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 some of the oldest fossils (1.85 to 1.77 million years ago) were found in the Caucasus. The oldest identified ''H. erectus'' specimen is a 2.04 million year old skull, DNH 134, from Drimolen, South Africa.<ref>{{Cite journal |last1=Herries |first1=Andy I. R. |last2=Martin |first2=Jesse M. |year=2020 |title=Contemporaneity of ''Australopithecus'', ''Paranthropus'', and early ''Homo erectus'' in South Africa |journal=Science |volume=368 |issue=6486 |article-number=eaaw7293 |doi=10.1126/science.aaw7293 |pmid=32241925 |bibcode=2020Sci...368w7293H |ref=CITEREFHerriesMartinLeeceAdams2020 |hdl=11568/1040368 |s2cid=214763272 |hdl-access=free}}</ref>}}<ref>{{cite journal |vauthors=Ferring R, Oms O, Agustí J, Berna F, Nioradze M, Shelia T, Tappen M, Vekua A, Zhvania D, Lordkipanidze D |display-authors=6 |title=Earliest human occupations at Dmanisi (Georgian Caucasus) dated to 1.85-1.78 Ma |journal=Proceedings of the National Academy of Sciences of the United States of America |volume=108 |issue=26 |pages=10432–6 |date=June 2011 |pmid=21646521 |pmc=3127884 |doi=10.1073/pnas.1106638108 |bibcode=2011PNAS..10810432F |doi-access=free}}</ref>{{efn|''Homo erectus soloensis'', found in Java, is considered the latest known survival of ''H. erectus''. Formerly dated to as late as 50,000 to 40,000 years ago, a 2011 study pushed back the date of its extinction of ''H. e. soloensis'' to 143,000 years ago at the latest, more likely before 550,000 years ago. <ref>{{cite journal |last1=Indriati |first1=E. |last2=Swisher |first2=C.C. |last3=Lepre |first3=C. |last4=Quinn |first4=R.L. |last5=Suriyanto |first5=R.A. |last6=Hascaryo |first6=A.T. |last7=Grün |first7=R. |last8=Feibel |first8=C.S. |last9=Pobiner |first9=B.L. |last10=Aubert |first10=M. |last11=Lees |first11=W. |last12=Antón |first12=S.C. |display-authors=6 |title=The age of the 20 meter Solo River terrace, Java, Indonesia and the survival of Homo erectus in Asia |journal=PLOS One |volume=6 |issue=6 |article-number=e21562 |year=2011 |pmid=21738710 |pmc=3126814 |doi=10.1371/journal.pone.0021562 |bibcode=2011PLoSO...621562I |doi-access=free}}</ref>}} | Africa, Eurasia | 180 cm (5 ft 11 in) | {{convert|60|kg|lb|abbr=on}} | 850 (early) – 1,100 (late) |data-sort-value=1000| Many{{efn|Now also included in ''H. erectus'' are Peking Man (formerly ''Sinanthropus pekinensis'') and Java Man (formerly ''Pithecanthropus erectus'').}}{{efn|''H. erectus'' is now grouped into various subspecies, including ''Homo 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 as ''Homo ergaster'', ''Homo floresiensis'', ''Homo antecessor'', ''Homo heidelbergensis'' and indeed ''Homo sapiens'' is not entirely clear.}} | 1891<br/>1892 |- | ''H. ergaster''<br/><small>African ''H. erectus''</small> | 1,800–1,300<ref name="Hazarika2007">{{cite book |last=Hazarika |first=Manjil |name-list-style=vanc |year=2007 |chapter=''Homo erectus/ergaster'' and Out of Africa: Recent Developments in Paleoanthropology and Prehistoric Archaeology |chapter-url=http://www.himalayanlanguages.org/files/hazarika/Manjil%20Hazarika%20EAA.pdf |title=EAA Summer School eBook |volume=1 |publisher=European Anthropological Association |pages=35–41 |quote=Intensive Course in Biological Anthrpology, 1st Summer School of the European Anthropological Association, 16–30 June 2007, Prague, Czech Republic}}</ref> | East and Southern Africa | | | 700–850 |data-sort-value=100| Many | 1949<br/>1975 |- | ''H. antecessor'' | 1,200–800 | Western Europe | 175 cm (5 ft 9 in) | {{convert|90|kg|lb|abbr=on}} | 1,000 |data-sort-value=6| 2 sites | 1994<br/>1997 |- | ''H. floresiensis''<br /><small>classification uncertain</small> | 1,000–50 | Indonesia | 100 cm (3 ft 3 in) | {{convert|25|kg|lb|abbr=on}} | 400 | data-sort-value="7" | 7 individuals | 2003<br />2004 |- | ''H. heidelbergensis''<br/><small>early ''H. neanderthalensis''?</small> | 600–300{{efn|The type fossil is Mauer 1, dated to ca. 0.6 million years ago. The transition from ''H. heidelbergensis'' to ''H. neanderthalensis'' between 300 and 243 thousand years ago is conventional, and makes use of the fact that there is no known fossil in this period. Examples of ''H. heidelbergensis'' are fossils found at Bilzingsleben (also classified as ''Homo erectus bilzingslebensis'').}} | Europe, Africa | 180 cm (5 ft 11 in) | {{convert|90|kg|lb|abbr=on}} | 1,100–1,400 |data-sort-value=100| Many | 1907<br/>1908 |- | ''H. cepranensis''<br/><small>a single fossil, possibly ''H. heidelbergensis''</small> |data-sort-value=450| c. 450<ref>{{cite journal |title=Pleistocene magnetochronology of early hominin sites at Ceprano and Fontana Ranuccio, Italy |doi=10.1016/j.epsl.2009.06.032 |bibcode=2009E&PSL.286..255M |volume=286 |issue= 1–2 |journal=Earth and Planetary Science Letters |pages=255–268 |year=2009 |last1=Muttoni |first1=Giovanni |last2=Scardia |first2=Giancarlo |last3=Kent |first3=Dennis V. |last4=Swisher |first4=Carl C. |last5=Manzi |first5=Giorgio |name-list-style=vanc|hdl=2434/164132 |hdl-access=free }}</ref> | Italy | | | 1,000 |data-sort-value=1.0| 1 skull cap | 1994<br/>2003 |- | ''H. naledi'' | data-sort-value="300" | 335—236<ref name="eLIFE-2017a">{{cite journal |last1=Dirks |first1=P.H. |last2=Roberts |first2=E.M. |last3=Hilbert-Wolf |first3=H. |last4=Kramers |first4=J.D. |last5=Hawks |first5=J. |last6=Dosseto A. |last7=Duval |first7=M. |last8=Elliott |first8=M. |last9=Evans |first9=M. |last10=Grün |first10=R. |last11=Hellstrom |first11=J. |last12=Herries |first12=A.I. |last13=Joannes-Boyau |first13=R. |last14=Makhubela |first14=T.V. |last15=Placzek |first15=C.J. |last16=Robbins |first16=J. |last17=Spandler |first17=C. |last18=Wiersma |first18=J. |last19=Woodhead |first19=J. |last20=Berger |first20=L.R. |display-authors=6 |title=Homo naledi and associated sediments in the Rising Star Cave, South Africa |journal=eLife |volume=6 |article-number=e24231 |date=May 2017 |pmid=28483040 |pmc=5423772 |doi=10.7554/eLife.24231 |doi-access=free}}</ref> | South Africa | {{height|cm=150}} | {{convert|45|kg|abbr=on}} | 450 | data-sort-value="15" |15 individuals | 2013<br />2015 |- | ''H. rhodesiensis''<br /><small>possibly ''H. heidelbergensis''</small> |data-sort-value=300| c. 300 | Zambia | | | 1,300 |data-sort-value=1.5| Single or very few | 1921<br/>1921 |- | ''H. sapiens''<br />{{small|(anatomically modern humans)}}<!-- To avoid confusion with "behaviourally modern humans", this phrase is almost never shortened to "modern humans". --> | c. 300–present{{efn|The age of ''H. sapiens'' has 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 in Jebel Irhoud (Morocco) suggest that ''Homo sapiens'' may have speciated by as early as 315,000 years ago.<ref>{{cite journal |last=Callaway |first=Ewan |title=Oldest Homo sapiens fossil claim rewrites our species' history |url=http://www.nature.com/news/oldest-homo-sapiens-fossil-claim-rewrites-our-species-history-1.22114 |date=7 June 2017 |journal=Nature |doi=10.1038/nature.2017.22114 |access-date=11 June 2017|url-access=subscription |doi-access=free }}</ref> Genetic evidence has been adduced for an age of roughly 270,000 years.<ref>{{cite journal |last1=Posth |first1=C. |last2=Wißing |first2=C. |last3=Kitagawa |first3=K. |last4=Pagani |first4=L. |last5=van Holstein |first5=L. |last6=Racimo |first6=F. |last7=Wehrberger |first7=K. |last8=Conard |first8=N.J. |last9=Kind |first9=C.J. |last10=Bocherens |first10=H. |last11=Krause |first11=J. |display-authors=6 |title=Deeply divergent archaic mitochondrial genome provides lower time boundary for African gene flow into Neanderthals |journal=Nature Communications |volume=8 |article-number=16046 |date=July 2017 |pmid=28675384 |pmc=5500885 |doi=10.1038/ncomms16046 |bibcode=2017NatCo...816046P}}</ref>}} | Worldwide | 150–190 cm (4 ft 11 in – 6 ft 3 in) | {{convert|50|–|100|kg|lb|abbr=on}} | 950–1,800 |data-sort-value=7500000000| (extant) |data-sort-value=1758| ——<br/>1758 |- | ''H. longi'' | c. 1,000<ref name=":0" />–c. 51 | Siberia | | |1,420<ref>{{cite journal |vauthors=Ni X, Ji Q, Wu W, Shao Q, Ji Y, Zhang C, Liang L, Ge J, Guo Z, Li J, Li Q, Grün R, Stringer C |date=25 June 2021 |title=Massive cranium from Harbin in northeastern China establishes a new Middle Pleistocene human lineage |journal=The Innovation |volume=2 |issue=3 |bibcode=2021Innov...200130N |doi=10.1016/j.xinn.2021.100130 |pmc=8454562 |pmid=34557770 |article-number=100130}}</ref> | data-sort-value="4.5" | Many<ref name=":0" /> | 2000<br />2021 |- | ''H. neanderthalensis''<br /> | 240–40<ref>{{cite journal |last1=Bischoff |first1=James L. |last2=Shamp |first2=Donald D. |last3=Aramburu |first3=Arantza |last4=Arsuaga |first4=Juan Luis |last5=Carbonell |first5=Eudald |last6=Bermúdez de Castro |first6=José María |name-list-style=vanc |date=March 2003 |title=The Sima de los Huesos Hominids Date to Beyond U/Th Equilibrium (>350 kyr) and Perhaps to 400–500 kyr: New Radiometric Dates |journal=Journal of Archaeological Science |volume=30 |issue=3 |pages=275–280 |doi=10.1006/jasc.2002.0834 |bibcode=2003JArSc..30..275B |issn=0305-4403 |display-authors=3}}</ref>{{efn|The first humans with "proto-Neanderthal traits" lived in Eurasia as early as 0.6 to 0.35 million years ago (classified as ''H. heidelbergensis'', also called a chronospecies because it represents a chronological grouping rather than being based on clear morphological distinctions from either ''H. erectus'' or ''H. neanderthalensis''). There is a fossil gap in Europe between 300 and 243 kya, and by convention, fossils younger than 243 kya are called "Neanderthal".<ref>{{cite journal |last1=Dean |first1=D. |last2=Hublin |first2=J.J. |last3=Holloway |first3=R. |last4=Ziegler |first4=R. |title=On the phylogenetic position of the pre-Neandertal specimen from Reilingen, Germany |journal=Journal of Human Evolution |volume=34 |issue=5 |pages=485–508 |date=May 1998 |pmid=9614635 |doi=10.1006/jhev.1998.0214 |doi-access=free|bibcode=1998JHumE..34..485D }}</ref>}} | Europe, Western Asia | 170 cm (5 ft 7 in) | {{convert|55|–|70|kg|lb|abbr=on}}<br />(heavily built) | 1,200–1,900 |data-sort-value=400| Many | 1829<br/>1864 |- | Nesher Ramla ''Homo''<br /><small>classification uncertain</small> | 140–120 | Israel | | | |data-sort-value=7| several individuals | 2021<br/> |- | Penghu 1<br/><small>possibly ''H. erectus'' or Denisova</small> |data-sort-value=100| c. 100{{efn|younger than 450 kya, either between 190–130 or between 70–10 kya<ref name="Changetal2015">{{cite journal |last1=Chang |first1=C.H. |last2=Kaifu |first2=Y. |last3=Takai |first3=M. |last4=Kono |first4=R.T. |last5=Grün |first5=R. |last6=Matsu'ura |first6=S. |last7=Kinsley |first7=L. |last8=Lin |first8=L.K. |display-authors=6 |title=The first archaic Homo from Taiwan |journal=Nature Communications |volume=6 |article-number=6037 |date=January 2015 |pmid=25625212 |pmc=4316746 |doi=10.1038/ncomms7037 |bibcode=2015NatCo...6.6037C}}</ref>}} | Taiwan | | | |data-sort-value=1.1| 1 individual | 2008(?)<br/>2015 |- | ''H. luzonensis''<ref name="NYT-20190410">{{cite news |last=Zimmer |first=Carl |name-list-style=vanc |author-link=Carl Zimmer |title=A new human species once lived in this Philippine cave – Archaeologists in Luzon Island have turned up the bones of a distantly related species, Homo luzonensis, further expanding the human family tree |url=https://www.nytimes.com/2019/04/10/science/homo-luzonensis-philippines-evolution.html |date=10 April 2019 |work=The New York Times |access-date=10 April 2019 }}</ref><br/> | 134–49<ref name="Détroit2019">{{cite journal |last1=Détroit |first1=F. |last2=Mijares |first2=A.S. |last3=Corny |first3=J. |last4=Daver |first4=G. |last5=Zanolli |first5=C. |last6=Dizon |first6=E. |last7=Robles |first7=E. |last8=Grün |first8=R. |last9=Piper |first9=P.J. |display-authors=6 |title=A new species of Homo from the Late Pleistocene of the Philippines |journal=Nature |volume=568 |issue=7751 |pages=181–186 |date=April 2019 |pmid=30971845 |doi=10.1038/s41586-019-1067-9 |bibcode=2019Natur.568..181D |hdl=10072/386785 |s2cid=106411053 |url=https://hal.archives-ouvertes.fr/hal-02296712/file/Detroit_%26_al_2019_Nature_postprint.pdf}}</ref><ref name="Grün_2023">{{cite journal| author1= Rainer Grün| author2= Chris Stringer | url= https://www.sciencedirect.com/science/article/pii/S0277379123004274/pdfft?md5=100788ed2ba65a9d9ec703067d360f27&pid=1-s2.0-S0277379123004274-main.pdf | title= Direct dating of human fossils and the ever-changing story of human evolution | doi= 10.1016/j.quascirev.2023.108379 | journal= Quaternary Science Reviews | volume= 322 | year= 2023 | issue= 108379 | article-number= 108379 | bibcode= 2023QSRv..32208379G | issn= 0277-3791 }}</ref> | Philippines | | | |data-sort-value=3| 3 individuals | 2007<br/>2019 |}
== See also == {{Portal|Evolutionary biology}} * List of human evolution fossils ''(with images)'' * Multiregional origin of modern humans
== Footnotes == {{notelist}}
== References == {{reflist}}
== Bibliography == {{Refbegin}} * {{cite journal |last1=Wood |first1=Bernard |last2=Richmond |first2=Brian G. |title=Human evolution: taxonomy and paleobiology |journal=Journal of Anatomy |volume=197 |issue=1 |pages=19–60 |date=July 2000 |pmid=10999270 |pmc=1468107 |doi=10.1046/j.1469-7580.2000.19710019.x}} {{Refend}}
== External links == {{Commons category|Homo}} {{Wikispecies|Homo}} {{Wikibooks|Introduction to Paleoanthropology}} * [http://www.bradshawfoundation.com/origins/index.php Exploring the Hominid Fossil Record] (Center for the Advanced Study of Hominid Paleobiology at George Washington University) * [http://www.talkorigins.org/faqs/homs/species.html Hominid species] * [http://www.talkorigins.org/faqs/homs/specimen.html Prominent Hominid Fossils] * [http://www.helsinki.fi/~mhaaramo/metazoa/deuterostoma/chordata/synapsida/eutheria/primates/hominoidea/hominidae_1.html Mikko's Phylogeny archive] * {{EOL}} * [http://humanorigins.si.edu/evidence/human-evolution-timeline-interactive Human Timeline (Interactive)] – Smithsonian, National Museum of Natural History (August 2016). {{Navboxes |title = Articles related to genus ''Homo'' |list = {{Human_Evolution}} {{Hominidae nav}} {{Haplorhini|Ho.}} }} {{Taxonbar|from=Q171283}} {{Authority control}}
Category:Homo * * Category:Australopithecines Category:Primate genera Category:Human evolution Category:Animal taxa named by Carl Linnaeus Category:Mammal genera with one living species Category:Extant Pliocene first appearances Category:Cradle of Humankind fauna