* _Africanthropus_ Dreyer, 1935 * _Atlanthropus_ Arambourg, 1954 * _Cyphanthropus_ Pycraft, 1928 * _Pithecanthropus_ Dubois, 1894 * _Protanthropus_ Haeckel, 1895 * _Sinanthropus_ Black, 1927 * _Tchadanthropus_ Coppens, 1965 * _Telanthropus_ Broom ">_ Evolutionary treechart emphasizing the subfamily Homininaeand the tribe Hominini. After diverging from the line to Ponginaethe early Homininaesplit into the tribes Hominini and Gorillini. The early Homininisplit 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.
Further information: List of alternative names for the human species _See Homininaefor an overview of taxonomy._
The Latinnoun _homō_ (genitive _hominis_) means "human being" or "man " in the generic sense of "human being, mankind". The binomial name _ Homosapiens_ was coined by Carl Linnaeus(1758). Names for other species of the genus were introduced beginning in the second half of the 19th century (_H. neanderthalensis_ 1864, _H. erectus_ 1892).
Even today, the genus _Homo_ has not been properly defined. Since the early human fossil record began to slowly emerge from the earth, the boundaries and definitions of the genus _Homo_ 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 define _Homo_ when he first created it for humans in the 18th century. The discovery of Neanderthalbrought the first addition. _ A model of the evolution of the genus Homo_ over the last 2 million years (vertical axis). The rapid "Out of Africa " expansion of _H. sapiens_ is indicated at the top of the diagram, with admixture indicated with Neanderthals, Denisovans, and unspecified archaic African hominins.
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 Mioceneand early Pliocenetimes produced a rich mix for debating classifications. There is continuing debate on delineating _Homo_ from _Australopithecus_—or, indeed, delineating _Homo_ from _Pan _, as one body of scientists argue that the two species of chimpanzee should be classed with genus _Homo_ rather than _Pan_. Even so, classifying the fossils of _Homo_ coincides with evidences of: 1) competent human bipedalism in _ Homo habilis_ inherited from the earlier _ Australopithecus_ of more than four million years ago, (see Laetoli); and 2) human tool culture having begun by 2.5 million years ago.
From the late-19th to mid-20th century, 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 and singular species with a large geographic spread of early migrations. Many such names are now dubbed as "synonyms " with _Homo_, including _Pithecanthropus_, _Protanthropus_, _Sinanthropus_, _Cyphanthropus_, _Africanthropus_, _Telanthropus_, _Atlanthropus_, and _Tchadanthropus_.
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") in even 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_. Some recently extinct species in the genus _Homo_ are only recently discovered and do not as yet have consensus binomial names (see Denisova homininand Red Deer Cave people ).
John Edward Gray(1825) was an early advocate of classifying taxa by designating tribes and families. 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 Hominina be designated a subtribe of Homininito include _only_ the genus _Homo_—that is, _not_ including the earlier upright walking hominins of the Pliocenesuch as _ Australopithecus_, _ Orrorin tugenensis_, _ Ardipithecus_, or _ Sahelanthropus_. Designations alternative to Homininaexisted, or were offered: _Australopithecinae_ (Gregory & Hellman 1939) and _Preanthropinae_ (Cela-Conde and later, Cela-Conde and Ayala (2003) proposed that the four genera _Australopithecus_, _Ardipithecus_, _Praeanthropus_, and _Sahelanthropus_ be grouped with _Homo_ within Hominina.
Humantimeline view • discuss • edit -10 — – -9 — – -8 — – -7 — – -6 — – -5 — – -4 — – -3 — – -2 — – -1 — – 0 — Human-like apes _ Nakalipithecus_ _ Ouranopithecus_ _ Sahelanthropus_ _ Orrorin_ _ Ardipithecus_ _ Australopithecus_ _HOMO HABILIS _ _HOMO ERECTUS _ _NEANDERTHAL _ _HOMO SAPIENS _ ← Earlier apes ← Possibly bipedal ← Earliest bipedal ← Earliest stone tools ← Earliest exit from Africa ← Earliest fire use ← Earliest cooking ← Earliest clothes ← Modern humans P l e i s t o c e n e P l i o c e n e M i o c e n e H
Several species, including _ Australopithecusgarhi _, _ Australopithecussediba _, _ Australopithecusafricanus _, and _ Australopithecusafarensis _, have been proposed as the direct ancestor of the _Homo_ lineage. These species have morphological features that align them with _Homo_, but there is no consensus as to which gave rise to _Homo_. The advent of _Homo_ was traditionally taken to coincide with the first use of stone tools (the Oldowan industry), and thus by definition with the beginning of the Lower Palaeolithic . The emergence of _Homo_ also coincides roughly with the onset of Quaternary glaciation, the beginning of the current ice age .
A fossil mandible fragment dated to 2.8 million years ago which may represent an intermediate stage between _Australopithecus_ and _Homo_ was discovered in 2015 in Afar, Ethiopia ( LD 350-1). Some authors would push the development of _Homo_ past 3 Mya, by including _ Kenyanthropus_ (a fossil dated 3.2 to 3.5 Mya, usually classified as an australopithecine species) into the genus _Homo_.
The most salient physiological development between the earlier australopithecine species and _Homo_ is the increase in cranial capacity , from about 450 cm3 (27 cu in) in _A. garhi_ to 600 cm3 (37 cu in) in _H. habilis_. Within the genus _Homo_, cranial capacity again doubled from _H. habilis_ through _ Homo ergaster_ or _H. erectus _ to _ Homo heidelbergensis_ by 0.6 million years ago. The cranial capacity of _H. heidelbergensis_ overlaps with the range found in modern humans.
_ Homoerectus_ has often been assumed to have developed anagenetically from _ Homohabilis_ from about 2 million years ago. This scenario was strengthened with the discovery of _ Homoerectus georgicus _, early specimens of _H. erectus_ found in the Caucasus, which seemed to exhibit transitional traits with _H. habilis_. As the earliest evidence for _H. erectus_ was found outside of Africa, it was considered plausible that _H. erectus_ developed in Eurasiaand then migrated back to Africa. Based on fossils from the Koobi Fora Formation, east of Lake Turkana in Kenya, Spoor et al. (2007) argued that _H. habilis_ may have survived beyond the emergence of _H. erectus_, so that the evolution of _H. erectus_ would not have been anagenetically, and _H. erectus_ would have existed alongside _H. habilis_ for about half a million years (1.9 to 1.4 million years ago ), during the early Calabrian .
See also: Human evolutionand Archaic human admixture with modern humans
Some of _H. ergaster_ migrated to Asia, where they are named _Homo erectus_, and to Europe with _ Homogeorgicus_. _H. ergaster_ in Africa and _H. erectus_ in Eurasiaevolved separately for almost two million years and presumably separated into two different species.
_ Homorhodesiensis_, who were descended from _H. ergaster_, migrated from Africa to Europe and became _ Homoheidelbergensis_ and later (about 250,000 years ago) _ Homoneanderthalensis_ and the Denisova hominin in Asia. The first _ Homosapiens_, descendants of _H. rhodesiensis_, appeared in Africa about 250,000 years ago. About 100,000 years ago, some _H. sapiens sapiens_ migrated from Africa to the Levantand met with resident Neanderthals, with some admixture . Later, about 70,000 years ago, perhaps after the Toba catastrophe, a small group left the Levantto populate Eurasia, Australiaand later the Americas . A subgroup among them met the Denisovans and, after further admixture, migrated to populate Melanesia. In this scenario, non-African people living today are mostly of African origin ("Out of Africa model "). However, there was also some admixture with Neanderthals and Denisovans, who had evolved locally (the "multiregional hypothesis "). Recent genomic results from the group of Svante Pääboalso show that 30,000 years ago at least three major subspecies coexisted: Denisovans, Neanderthals and anatomically modern humans . Today, only _H. sapiens_ remains, with no other extant species.
LIST OF SPECIES
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, Red Deer Cave people, and _ H. floresiensis_ remains under debate. _ H. heidelbergensis_ and _H. neanderthalensis_ are closely related to each other and have been considered to be subspecies of _H. sapiens_. Recently, nuclear DNA from a Neanderthalspecimen from Vindija Cavehas been sequenced using two different methods that yield similar results regarding Neanderthal and _H. sapiens_ lineages, with both analyses suggesting a date for the split between 460,000 and 700,000 years ago, though a population split of around 370,000 years is inferred. The nuclear DNAresults indicate about 30% of derived alleles in _H. sapiens_ are also in the Neanderthallineage. This high frequency may suggest some gene flow between ancestral human and Neanderthalpopulations due to mating between the two.
_ Homo naledi_ was discovered near Johannesburg, South Africain 2013 and announced on 10 September 2015. Fossils indicate the hominin was 1.45–1.5 meters tall and had a small brain. The fossils have been dated to be between 335,000 and 236,000 years old.
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Comparative table of _Homo_ species SPECIES TEMPORAL RANGE KYA HABITAT ADULT HEIGHT ADULT MASS CRANIAL CAPACITY (CM³) FOSSIL RECORD DISCOVERY / PUBLICATION OF NAME
_H. habilis _ 2,100 – 1,500 Africa 110-140 cm (4 ft 11 in) 33–55 kg (73–121 lb) 510–660 Many 1960/1964
_H. erectus _ 1,900 – 70
700 2 sites 1972/1986
_H. gautengensis _ also classified as _H. habilis_ 1,900 – 600 South Africa 100 cm (3 ft 3 in)
3 individuals 2010/2010
_ H. ergaster_ also classified as _H. erectus_ 1,800 – 1,300 Eastern and Southern Africa
700–850 Many 1975
_ H. antecessor_ also classified as _H. heidelbergensis_ 1,200 – 800 Spain 175 cm (5 ft 9 in) 90 kg (200 lb) 1,000 2 sites 1997
_ H. cepranensis_ a single fossil, possibly _H. erectus_ 900 – 350 Italy
1,000 1 skull cap 1994/2003
_ H. heidelbergensis_ 600 – 350 Europe, Africa, China 180 cm (5 ft 11 in) 90 kg (200 lb) 1,100–1,400 Many 1908
_ H. neanderthalensis_ possibly a subspecies of _H. sapiens_ 350 – 40 Europe, Western Asia 170 cm (5 ft 7 in) 55–70 kg (121–154 lb) (heavily built) 1,200–1,900 Many (1829)/1864
_H. naledi _ 335–236 South Africa 150 centimetres (4 ft 11 in) tall 45 kilograms (99 lb) 450 15 individuals 2013/2015
_H. tsaichangensis _ possibly _H. erectus_ 190 – 10 Taiwan
1 individual pre-2008/2015
1,300 Very few 1921
_H. sapiens _ (modern humans ) 300
– present Worldwide 150 - 190 cm (4 ft 7 in - 6 ft 3 in) 50–100 kg (110–220 lb) 950–1,800 (extant) —/1758
Denisova hominin possible _H. sapiens_ subspecies or hybrid 40 Russia
1 site 2000/2010
Red Deer Cave people possible _H. sapiens_ subspecies or hybrid 14.5–11.5 China
Very few 2012
* List of human evolution fossils_(with images)_ * Nature timeline
* ^ The conventional estimate on the age of _H. habilis_ is at roughly 2.1 to 2.3 million years. Stringer, C.B. (1994). "Evolution of early humans". In Steve Jones, Robert Martin & David Pilbeam (eds.). _The Cambridge Encyclopedia of HumanEvolution_. Cambridge: Cambridge University Press. p. 242. CS1 maint: Uses editors parameter (link ) McHenry, H.M (2009). " HumanEvolution". In Michael Ruse & Joseph Travis. _Evolution: The First Four Billion Years_. Cambridge, Massachusetts: The Belknap Press of Harvard University Press. p. 265. ISBN 978-0-674-03175-3 . CS1 maint: Uses editors parameter (link ) Suggestions for pushing back the age to 2.8 Mya were made in 2015 based on the discovery of a jawbone : Wilford, John Noble (2015-03-04). "Jawbone Fossil Fills a Gap in Early HumanEvolution". _The New York Times_. ISSN 0362-4331 . Retrieved 2015-05-30. Spoor, Fred; Gunz, Philipp; Neubauer, Simon; Stelzer, Stefanie; Scott, Nadia; Kwekason, Amandus; Dean, M. Christopher (March 5, 2015). "Reconstructed _ Homohabilis_ type OH 7 suggests deep-rooted species diversity in early _Homo_". _Nature_. 519 (7541): 83–86. ISSN 0028-0836 . PMID 25739632 . doi :10.1038/nature14224 . . Villmoare, Brian; Kimbel, William H.; Seyoum, Chalachew; Campisano, Christopher J.; DiMaggio, Erin N.; Rowan, John; Braun, David R.; Arrowsmith, J. Ramón; Reed, Kaye E. (2015-03-20). "Early _Homo_ at 2.8 Ma from Ledi-Geraru, Afar, Ethiopia". _Science_. 347 (6228): 1352–1355. ISSN 0036-8075 . PMID 25739410 . doi :10.1126/science.aaa1343 . * ^ Curnoe, D (2010). "A review of early _Homo_ in southern Africa focusing on cranial, mandibular and dental remains, with the description of a new species (_ Homogautengensis_ sp. nov.)". _HOMO – Journal of Comparative HumanBiology_. 61: 151–177.
* ^ Schuster, Angela M. H. (1997). "Earliest Remains of Genus _Homo_". _Archaeology_. 50 (1). Retrieved 5 March 2015. The line to the earliest members of _Homo_ made final separation from the lineage of _Pan_ by late Mioceneor early Pliocenetimes—with date estimates by several specialists ranging from 13 million years ago to more recently than six million years ago.
* Arnason, U; Gullberg, A; Janke, A (1998). "Molecular timing of primate divergences as estimated by two nonprimate calibration points". _J. Mol. Evol_. 47 (6): 718–27. PMID 9847414 . doi :10.1007/PL00006431 . * Patterson, N; Richter, DJ; Gnerre, S; Lander, ES; Reich, D (2006). "Genetic evidence for complex speciation of humans and chimpanzees". _Nature_. 441 (7097): 1103–8. PMID 16710306 . doi :10.1038/nature04789 . * Wakeley, J (2008). "Complex speciation of humans and chimpanzees". _Nature_. 452 (7184): E3–4. PMID 18337768 . doi :10.1038/nature06805 . "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 .
* ^ Ghosh, Pallab (20 August 2014). "New dates rewrite Neanderthal story". BBC News. * ^ Green, R.E.; Krause, J.; Briggs, A.W.; Maricic, T.; Stenzel, U.; Kircher, M.; Patterson, N.; Li, H.; Zhai, W.; Fritz, M.H.Y.; Hansen, N.F. (2010). "A draft sequence of the Neandertal genome". _Science_. 328 (5979): 710–722. PMID 20448178 . doi :10.1126/science.1188021 . * ^ Lowery, R.K.; Uribe, G.; Jimenez, E.B.; Weiss, M.A.; Herrera, K.J.; Regueiro, M.; Herrera, R.J. (2013). " Neanderthaland Denisova genetic affinities with contemporary humans: Introgression versus common ancestral polymorphisms". _Gene_. 530 (1): 83–94. 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. * ^ The word "human" itself is from Latin_humanus_, an adjective formed on the root of _homo_, thought to derive from a Proto-Indo-European word for "earth" reconstructed as _*dhǵhem-_. dhghem The American Heritage Dictionary of the English Language: Fourth Edition. 2000. * ^ Linné, Carl von (1758). _Systema naturæ. Regnum animale._ (10 ed.). pp. 18, 20. Retrieved 19 November 2012. . Note: In 1959, Linnaeus was designated as the lectotype for _ Homo sapiens_ (Stearn, W. T. 1959. "The background of Linnaeus's contributions to the nomenclature and methods of systematic biology", _Systematic Zoology_ 8 (1): 4-22, p. 4) which means that following the nomenclatural rules , _ Homosapiens_ was validly defined as the animal species to which Linnaeus belonged. * ^ Schwartz, Jeffrey H.; Tattersall, Ian (28 August 2015). "Defining the genus _Homo_". _Science_. 349 (6251): 931–932. doi :10.1126/science.aac6182 . Retrieved 2015-11-02. * ^ Lents, Nathan (4 October 2014). "_ Homonaledi_ and the Problems with the _Homo_ Genus". _The Wildernist_. Archived from the original on 18 November 2015. Retrieved 2015-11-02. * ^ Wood, B.; Collard, M. (2 April 1999). "The human genus". _Science_. 284 (5411): 65–71. PMID 10102822 . doi :10.1126/science.284.5411.65 . * ^ Stringer, C. (2012). "What makes a modern human". _Nature_. 485 (7396): 33–35. PMID 22552077 . doi :10.1038/485033a . * ^ "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 * ^ "early man", _Protanthropus primigenius_ Ernst Haeckel, _Systematische Phylogenie_ vol. 3 (1895), p. 625 * ^ "Sinic man", from _Sinanthropus pekinensis_ ( Peking Man), Davidson Black(1927). * ^ "crooked man", from _Cyphanthropus rhodesiensis_ (Rhodesian Man ) William Plane Pycraft(1928). * ^ "African man", used by T. F. Dreyer (1935) for the Florisbad Skull he found in 1932 (also _ Homoflorisbadensis_ or _ Homohelmei_). Also the genus suggested for a number of archaic human skulls found at Lake Eyasiby Weinert (1938). Leaky, _Journal of the East Africa Natural History Society' (1942), p. 43._ * ^ "remote man"; from _Telanthropus capensis_ (Broom and Robinson 1949), see (1961), p. 487. * ^ 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 Arambourgin Tighennif, Algeria. Arambourg, C. (1955). "A recent discovery in human paleontology: Atlanthropus of ternifine (Algeria)". _American Journal of Physical Anthropology_. 13 (2): 191–201. doi :10.1002/ajpa.1330130203 . * ^ Y. Coppens, "L'Hominien du Tchad", _Actes V Congr. PPEC_ I (1965), 329f.; "Le Tchadanthropus", _Anthropologia_ 70 (1966), 5–16. * ^ Alexandra Vivelo (2013), Characterization of Unique Features of the DenisovanExome * ^ J. E. Gray, "An outline of an attempt at the disposition of Mammalia into Tribes and Families, with a list of genera apparently appertaining to each Tribe", _Annals of Philosophy', new series (1825), pp. 337–344._ * ^ Wood and Richmond; Richmond, BG (2000). " Humanevolution: taxonomy and paleobiology" . _Journal of Anatomy_. 197 (Pt 1): 19–60. PMC 1468107 _. PMID 10999270 . doi :10.1046/j.1469-7580.2000.19710019.x . * ^ Brunet, M.; et al. (2002). "A new hominid from the upper Mioceneof Chad, central Africa". Nature_. 418: 145–151. PMID 12110880 . doi :10.1038/nature00879 . * ^ Cela-Conde, C.J.; Ayala, F.J. (2003). "Genera of the human lineage" . _PNAS_. 100 (13): 7684–7689. PMC 164648 _. PMID 12794185 . doi :10.1073/pnas.0832372100 . * ^ Wood, B.; Lonergan, N. (2008). "The hominin fossil record: taxa, grades and clades" (PDF). J. Anat_. 212: 354–376. PMC 2409102 _. PMID 18380861 . doi :10.1111/j.1469-7580.2008.00871.x . * ^ Cela-Conde, C. J.; Ayala, F. J. (2003). "Genera of the human lineage" . Proceedings of the National Academy of Sciences_. 100 (13): 7684–7689. PMC 164648 _. PMID 12794185 . doi :10.1073/pnas.0832372100 . * ^ Pickering, R.; Dirks, P. H.; Jinnah, Z.; De Ruiter, D. J.; Churchill, S. E.; Herries, A. I.; Berger, L. R. (2011). " Australopithecussediba_ at 1.977 Ma and implications for the origins of the genus _Homo_". _Science_. 333 (6048): 1421–1423. PMID 21903808 . doi :10.1126/science.1203697 . * ^ Asfaw, B.; White, T.; Lovejoy, O.; Latimer, B.; Simpson, S.; Suwa, G. (1999). " Australopithecusgarhi: a new species of early hominid from Ethiopia". _Science_. 284 (5414): 629–635. PMID 10213683 . doi :10.1126/science.284.5414.629 . * ^ In 2010, evidence was presented that seems to attribute the use of stone tools to _ Australopithecusafarensis_, close to a million years before the first appearance of _Homo_. McPherron, S. P.; Alemseged, Z.; Marean, C. W.; Wynn, J. G.; Reed, D.; Geraads, D.; Bobe, R.; Bearat, H. A. (2010). "Evidence for stone-tool-assisted consumption of animal tissues before 3.39 million years ago at Dikika, Ethiopia". _Nature_. 466: 857–860. PMID 20703305 . doi :10.1038/nature09248 . "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 unambiguous stone-tool cut marks for flesh removal to between 3.42 and 3.24 Myrago 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 Australopithecusafarensis." * ^ Erin N. DiMaggio EN; Campisano CJ; Rowan J; Dupont-Nivet G; Deino AL; et al. (2015). "Late Pliocenefossiliferous sedimentary record and the environmental context of early _Homo_ from Afar, Ethiopia". _Science _. 347: 1355–1359. PMID 25739409 . doi :10.1126/science.aaa1415 . See also: "Oldest known member of human family found in Ethiopia". _ New Scientist_. 4 March 2015. Retrieved 7 March 2015. , Ghosh, Pallab (4 March 2015). "\'First human\' discovered in Ethiopia". _BBC News_. Retrieved 5 March 2015. * ^ Cela-Conde and Ayala (2003) recognize five genera within Hominina: _Ardipithecus_, _Australopithecus_ (including _Paranthropus_), _Homo_ (including _Kenyanthropus_), _Praeanthropus_ (including _Orrorin_), and _Sahelanthropus_. Cela-Conde, C. J.; Ayala, F. J. (2003). "Genera of the human lineage" . _Proceedings of the National Academy of Sciences_. 100 (13): 7684–7689. PMC 164648 _. PMID 12794185 . doi :10.1073/pnas.0832372100 . * ^ "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." Spoor, F; Leakey, M.G; Gathogo, P.N; Brown, F.H; Antón, S.C; McDougall, I; Kiarie, C; Manthi, F.K.; Leakey, L.N. (2007). "Implications of new early _Homo_ fossils from Ileret, east of Lake Turkana, Kenya". _Nature_. 448 (7154): 688–691. PMID 17687323 . doi :10.1038/nature05986 . * ^ Green, RE; Krause, J; et al. (2010). "A draft sequence of the Neanderthalgenome". _Science_. 328 (5979): 710–22. PMID 20448178 . doi :10.1126/science.1188021 . * ^ Reich, D; Green, RE; Kircher, M; et al. (December 2010). "(December 2010). 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* ^ Schrenk, Friedemann; Kullmer, Ottmar; Bromage, Timothy (2007). "The Earliest Putative _Homo_ Fossils". In Henke, Winfried; Tattersall, Ian . _Handbook of Paleoanthropology_. 1. In collaboration with Thorolf Hardt. Berlin, Heidelberg: Springer . pp. 1611–1631. ISBN 978-3-540-32474-4 . doi :10.1007/978-3-540-33761-4_52 . Confirmed _H. habilis_ fossils are dated to between 2.1 and 1.5 million years ago. This date range overlaps with the emergence of _ Homoerectus_. Wilford, John Noble (August 9, 2007). "Fossils in KenyaChallenge Linear Evolution". _ The New York Times_. Retrieved 2015-05-04.
* DiMaggio, Erin N.; Campisano, Christopher J.; Rowan, John; et al. (March 20, 2015). "Late Pliocenefossiliferous sedimentary record and the environmental context of early _Homo_ from Afar, Ethiopia". _Science _. Washington, D.C.: American Association for the Advancement of Science . 347 (6228): 1355–1359. ISSN 0036-8075 . PMID 25739409 . doi :10.1126/science.aaa1415 . Homininswith "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.
* ^ Haviland, William A.; Walrath, Dana; Prins, Harald E. L. ; McBride, Bunny (2007). _Evolution and Prehistory: The HumanChallenge_ (8th ed.). Belmont, CA: Thomson Wadsworth . p. 162. ISBN 978-0-495-38190-7 . _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 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 whether _H. erectus_ emerged in Africa and migrated to Eurasia, or if, conversely, it evolved in Eurasiaand migrated back to Africa.
* Ferring, R.; Oms, O.; Agusti, J.; Berna, F.; Nioradze, M.; Shelia, T.; Tappen, M.; Vekua, A.; Zhvania, D.; Lordkipanidze, D. (2011). "Earliest human occupations at Dmanisi(Georgian Caucasus) dated to 1.85-1.78 Ma" . _Proceedings of the National Academy of Sciences_. 108 (26): 10432. PMC 3127884 _. PMID 21646521 . doi :10.1073/pnas.1106638108 . * "New discovery suggests Homo erectusoriginated from Asia". Daily News and Analysis _. Mumbai, India: Diligent Media Corporation Ltd. Asian News International. June 8, 2011. Retrieved 2015-05-04. * Frazier, Kendrick (November–December 2006). "Leakey Fights Church Campaign to Downgrade KenyaMuseum\'s HumanFossils". _ Skeptical Inquirer_. Amherst, NY: Committee for Skeptical Inquiry. 30 (6). ISSN 0194-6730 . Retrieved 2015-05-04.
* ^ Now also included in _H. erectus_ are Peking Man(formerly _Sinanthropus pekinensis_) and Java Man(formerly _Pithecanthropus erectus_). _H. erectus_ is now grouped into various subspecies, including _ Homo erectuserectus _, _ Homo erectusyuanmouensis _, _Homo erectus lantianensis _, _ Homo erectusnankinensis _, _ Homoerectus pekinensis _, _ Homo erectuspalaeojavanicus _, _ Homo erectussoloensis _, _ Homo erectustautavelensis _, _ Homo erectusgeorgicus _. The distinction from descendant species such as _ Homo ergaster_, _Homo floresiensis _, _ Homo antecessor_, _ Homo heidelbergensis_ and indeed _ Homo sapiens_ is not entirely clear. * ^ Curnoe, Darren (June 2010). "A review of early _Homo_ in southern Africa focusing on cranial, mandibular and dental remains, with the description of a new species (_ Homogautengensis_ sp. nov.)". _HOMO - Journal of Comparative HumanBiology_. Amsterdam, the Netherlands: Elsevier. 61 (3): 151–177. ISSN 0018-442X . PMID 20466364 . doi :10.1016/j.jchb.2010.04.002 . 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. * ^ Hazarika, Manjil (2007). "_ Homoerectus/ergaster_ and Out of Africa: Recent Developments in Paleoanthropology and Prehistoric Archaeology" (PDF). _EAA Summer School eBook_. 1. European Anthropological Association. pp. 35–41. Retrieved 2015-05-04. "Intensive Course in Biological Anthrpology, 1st Summer School of the European Anthropological Association, 16–30 June, 2007, Prague, Czech Republic" * ^ The type fossil is Mauer 1, dated to ca. 0.6 million years ago. The transition from _H. heidelbergensis_ to _H. neanderthalensis_ at about 0.35 to 0.25 million years ago is largely conventional. Relevant examples are fossils found at Bilzingsleben (also classified as _ Homo erectusbilzingslebensis_).
* ^ Bischoff, James L.; Shamp, Donald D.; Aramburu, Arantza; et al. (March 2003). "The Sima de los Huesos Hominids Date to Beyond U/Th Equilibrium (>350 kyr) and Perhaps to 400–500 kyr: New Radiometric Dates". _ Journal of Archaeological Science_. Amsterdam, the Netherlands: Elsevier. 30 (3): 275–280. ISSN 0305-4403 . doi :10.1006/jasc.2002.0834 . The first humans with "proto-Neanderthal traits" lived in Eurasiaas 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_), with the first "true Neanderthals" appearing between 0.25 and 0.2 million years ago.
* Papagianni, Dmitra; Morse, Michael A. (2013). _The Neanderthals Rediscovered: How Modern Science is Rewriting Their Story_. New York: Thames & Hudson. ISBN 978-0-500-05177-1 .
* ^ Chang, Chun-Hsiang; Kaifu, Yousuke; Takai, Masanaru; Kono, Reiko T.; Grün, Rainer; Matsu’ura, Shuji; Kinsley, Les; Lin, Liang-Kong (2015). "The first archaic _Homo_ from Taiwan". _Nature Communications _. 6: 6037. PMC 4316746 _. PMID 25625212 . doi :10.1038/ncomms7037 . * ^ Zimmer, Carl (7 June 2017). "Oldest Fossils of HomoSapiens Found in Morocco, Altering History of Our Species". New York Times_. Retrieved 7 June 2017. * ^ Callaway, Ewan (7 June 2017). "Oldest Homo sapiensfossil claim rewrites our species\' history". _ Nature (journal)_. doi :10.1038/nature.2017.22114 . Retrieved 7 June 2017.
* Serre; Langaney, André; Chech, Mario; Teschler-Nicola, Maria; Paunovic, Maja; Mennecier, Philippe; Hofreiter, Michael; Possnert, Göran; Pääbo, Svante; et al. (2004). "No evidence of Neandertal mt DNAcontribution to early modern humans" . _PLoS Biology_. 2 (3): 313–7. PMC 368159 . PMID 15024415 . doi :10.1371/journal.pbio.0020057 .
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