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For other species or subspecies suggested, see below.

Synonyms

Synonyms

Africanthropus Dreyer, 1935 Atlanthropus Arambourg, 1954 Cyphanthropus Pycraft, 1928 Pithecanthropus Dubois, 1894 Protanthropus Haeckel, 1895 Sinanthropus Black, 1927 Tchadanthropus Coppens, 1965 Telanthropus Broom & Anderson 1949

Homo
Homo
is the genus that encompasses the extant species Homo
Homo
sapiens (modern humans), plus several extinct species classified as ancestral to or closely related to modern humans, most notably Homo
Homo
erectus. The genus is between 2 and 3 million years old, taken to emerge with the appearance of Homo
Homo
habilis[1] and possibly that of Homo gautengensis.[2] Homo
Homo
is derived from the genus Australopithecus, which itself had previously split from the lineage of Pan, the chimpanzees.[3] Taxonomically, Homo
Homo
is the only genus assigned to the subtribe Hominina
Hominina
which, with the subtribes Australopithecina
Australopithecina
and Panina, comprise the tribe Hominini
Hominini
(see evolutionary tree below). All species of the genus Homo
Homo
plus those species of the australopithecines that arose after the split from Pan are called hominins. Homo erectus
Homo erectus
appeared about two million years ago in East Africa (where it is dubbed Homo
Homo
ergaster) and, in several early migrations, it spread throughout Africa and Eurasia. It was likely the first hominin to live in a hunter-gatherer society and to control fire. An adaptive and successful species, Homo erectus
Homo erectus
persisted for almost 2 million years before suddenly becoming extinct about 70,000 years ago (0.07 Ma)—perhaps a casualty of the Toba supereruption catastrophe. The subspecies Homo sapiens
Homo sapiens
sapiens, anatomically modern humans, emerged about 200,000 years ago (0.2 Ma) in East Africa (see Omo remains). Modern humans migrated from Africa as recently as 60,000 years ago. During Upper Paleolithic
Upper Paleolithic
times they spread throughout Africa, Eurasia, Oceania, and the Americas, and they encountered archaic humans along the way during these migrations. Homo
Homo
sapiens sapiens is the only surviving species and subspecies of the genus Homo. Archaic humans
Archaic humans
survived until about 40,000 years ago (see H. neanderthalensis),[4] and possibly until as late as the times of the Epipaleolithic
Epipaleolithic
culture (about 12,000 years ago). DNA
DNA
analysis provides some evidence of interbreeding between archaic and modern humans.[5][6]

Contents

1 Names and taxonomy 2 Evolution 3 Migration 4 List of species 5 See also 6 References 7 Further reading 8 External links

Names and taxonomy

Evolutionary tree
Evolutionary tree
chart emphasizing the subfamily Homininae
Homininae
and the tribe Hominini. After diverging from the line to Ponginae
Ponginae
the early Homininae
Homininae
split into the tribes Hominini
Hominini
and Gorillini. The early Hominini
Hominini
split further, separating the line to Homo
Homo
from the lineage of Pan. Currently, tribe Hominini
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 Homininae
Homininae
for an overview of taxonomy.

The Latin
Latin
noun homō (genitive hominis) means "human being" or "man" in the generic sense of "human being, mankind".[7] The binomial name Homo sapiens
Homo sapiens
was coined by Carl Linnaeus
Carl Linnaeus
(1758).[8] Names for other species of the genus were introduced beginning in the second half of the 19th century ( H. neanderthalensis
H. neanderthalensis
1864, H. erectus 1892). Even today, the genus Homo
Homo
has not been properly defined.[9][10][11] Since the early human fossil record began to slowly emerge from the earth, the boundaries and definitions of the genus Homo
Homo
have been poorly defined and constantly in flux. Because there was no reason to think it would ever have any additional members, Carl Linnaeus
Carl Linnaeus
did not even bother to define Homo
Homo
when he first created it for humans in the 18th century. The discovery of Neanderthal
Neanderthal
brought the first addition.

A model of the evolution of the genus Homo
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. Late survival of robust australopithecines (Paranthropus) alongside Homo
Homo
until 1.2 Mya is indicated in purple.

The genus Homo
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
Miocene
and early Pliocene
Pliocene
times produced a rich mix for debating classifications. There is continuing debate on delineating Homo
Homo
from Australopithecus—or, indeed, delineating Homo
Homo
from Pan, as one body of scientists argue that the two species of chimpanzee should be classed with genus Homo
Homo
rather than Pan. Even so, classifying the fossils of Homo
Homo
coincides with evidences of: 1) competent human bipedalism in Homo habilis
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 centuries, a number of new taxonomic names including new generic names were proposed for early human fossils; most have since been merged with Homo
Homo
in recognition that Homo erectus
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,[12] Protanthropus,[13] Sinanthropus,[14] Cyphanthropus,[15] Africanthropus,[16] Telanthropus,[17] Atlanthropus,[18] and Tchadanthropus.[19] Classifying the genus Homo
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
H. georgicus
vs. H. erectus georgicus.[20] Some recently extinct species in the genus Homo
Homo
are only recently discovered and do not as yet have consensus binomial names (see Denisova hominin
Denisova hominin
and Red Deer Cave
Cave
people). John Edward Gray
John Edward Gray
(1825) was an early advocate of classifying taxa by designating tribes and families.[21] Wood and Richmond (2000) proposed that Hominini
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
Hominina
be designated a subtribe of Hominini
Hominini
to include only the genus Homo—that is, not including the earlier upright walking hominins of the Pliocene
Pliocene
such as Australopithecus, Orrorin
Orrorin
tugenensis, Ardipithecus, or Sahelanthropus.[22] Designations alternative to Hominina
Hominina
existed, or were offered: Australopithecinae (Gregory & Hellman 1939) and Preanthropinae (Cela-Conde & Altaba 2002);[23][24][25] and later, Cela-Conde and Ayala (2003) proposed that the four genera Australopithecus, Ardipithecus, Praeanthropus, and Sahelanthropus
Sahelanthropus
be grouped with Homo
Homo
within Hominina.[not in citation given][26] Evolution

Human
Human
timeline

view • discuss • edit

-10 — – -9 — – -8 — – -7 — – -6 — – -5 — – -4 — – -3 — – -2 — – -1 — – 0 —

Human-like apes

Nakalipithecus

Ouranopithecus

Sahelanthropus

Orrorin

Ardipithecus

Australopithecus

Homo
Homo
habilis

Homo
Homo
erectus

Neanderthal

Homo
Homo
sapiens

Earlier apes

LCA-Gorilla separation

Possibly bipedal

LCA-Chimpanzee separation

Earliest bipedal

Earliest stone tools

Earliest exit from Africa

Earliest fire use

Earliest in Europe

Earliest cooking

Earliest clothes

Modern speech

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

o

m

i

n

i

d

s

Axis scale: million years Also see: Life timeline and Nature timeline

Further information: Timeline of human evolution, Archaic humans, and Australopithecus Several species, including Australopithecus
Australopithecus
garhi, Australopithecus sediba, Australopithecus
Australopithecus
africanus, and Australopithecus
Australopithecus
afarensis, have been proposed as the direct ancestor of the Homo
Homo
lineage.[27][28] 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
Homo
was traditionally taken to coincide with the first use of stone tools (the Oldowan
Oldowan
industry), and thus by definition with the beginning of the Lower Palaeolithic.[29] The emergence of Homo
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
Australopithecus
and Homo
Homo
was discovered in 2015 in Afar, Ethiopia (LD 350-1).[30] Some authors would push the development of Homo
Homo
past 3 Mya, by including Kenyanthropus
Kenyanthropus
(a fossil dated 3.2 to 3.5 Mya, usually classified as an australopithecine species) into the genus Homo.[31] The most salient physiological development between the earlier australopithecine species and Homo
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
Homo heidelbergensis
by 0.6 million years ago. The cranial capacity of H. heidelbergensis
H. heidelbergensis
overlaps with the range found in modern humans. Homo erectus
Homo erectus
has often been assumed to have developed anagenetically from Homo habilis
Homo habilis
from about 2 million years ago. This scenario was strengthened with the discovery of Homo erectus
Homo erectus
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 Eurasia
Eurasia
and 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.[32] The cladogram below depicts most species of Homo, with Australopithecus
Australopithecus
as an outgroup, according to Strait, Fleagle and Grine (2015).[33]

Hominini

Australopithecus
Australopithecus

Homo

Homo habilis
Homo habilis

Homo rudolfensis
Homo rudolfensis

Homo ergaster
Homo ergaster

Homo erectus
Homo erectus

Homo antecessor
Homo antecessor

Homo
Homo
heidelbergensis

Homo neanderthalensis
Homo neanderthalensis

Homo sapiens
Homo sapiens

Migration See also: Human evolution
Human evolution
and Archaic human admixture with modern humans Some of H. ergaster
H. ergaster
migrated to Asia, where they are named Homo erectus, and to Europe with Homo
Homo
georgicus. H. ergaster
H. ergaster
in Africa and H. erectus in Eurasia
Eurasia
evolved separately for almost two million years and presumably separated into two different species. Homo
Homo
rhodesiensis, who were descended from H. ergaster, migrated from Africa to Europe and became Homo heidelbergensis
Homo heidelbergensis
and later (about 250,000 years ago) Homo neanderthalensis
Homo neanderthalensis
and the Denisova hominin
Denisova hominin
in Asia. The first Homo
Homo
sapiens, 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 Levant
Levant
and met with resident Neanderthals, with some admixture.[34] Later, about 70,000 years ago, perhaps after the Toba catastrophe, a small group left the Levant
Levant
to populate Eurasia, Australia
Australia
and later the Americas. A subgroup among them met the Denisovans[35] 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 Pääbo
Svante Pääbo
also show that 30,000 years ago at least three major subspecies coexisted: Denisovans, Neanderthals and anatomically modern humans.[36] 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
Cave
people, and H. floresiensis
H. floresiensis
remains under debate. H. heidelbergensis
H. heidelbergensis
and H. neanderthalensis
H. neanderthalensis
are closely related to each other and have been considered to be subspecies of H. sapiens. Recently, nuclear DNA
DNA
from a Neanderthal
Neanderthal
specimen from Vindija Cave
Vindija Cave
has been sequenced using two different methods that yield similar results regarding Neanderthal
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 DNA
DNA
results indicate about 30% of derived alleles in H. sapiens are also in the Neanderthal
Neanderthal
lineage. This high frequency may suggest some gene flow between ancestral human and Neanderthal
Neanderthal
populations due to mating between the two.[37] Homo naledi
Homo naledi
was discovered near Johannesburg, South Africa
South Africa
in 2013 and announced on 10 September 2015. Fossils indicate the hominin was 1.45–1.5 meters tall and had a small brain.[38] The fossils have been dated to be between 335,000 and 236,000 years old, long after much larger-brained and more modern-looking hominins had appeared.[39][40] It is thought that H. naledi is probably an offshoot within the genus Homo.[41]

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Comparative table of Homo
Homo
species

Species Temporal range kya Habitat Adult height Adult mass Cranial capacity
Cranial capacity
(cm³) Fossil record Discovery / publication of name

H. habilis 2,100 – 1,500[42] Africa 110-140 cm (4 ft 11 in) 33–55 kg (73–121 lb) 510–660 Many 1960/1964

H. erectus 1,900 – 70[43][44][45][46] Africa, Eurasia
Eurasia
(Java, China, India, Caucasus) 180 cm (5 ft 11 in) 60 kg (130 lb) 850 (early) – 1,100 (late) Many[47] 1891/1892

H. rudolfensis membership in Homo
Homo
uncertain 1,900 Kenya

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[48] 2010/2010

H. ergaster also classified as H. erectus 1,800 – 1,300[49] Eastern and Southern Africa

700–850 Many 1949/1975

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 1994/1997

H. cepranensis a single fossil, possibly H. erectus 900 – 350 Italy

1,000 1 skull cap 1994/2003

H. heidelbergensis 600 – 300[50] Europe, Africa, China 180 cm (5 ft 11 in) 90 kg (200 lb) 1,100–1,400 Many 1907/1908

H. neanderthalensis possibly a subspecies of H. sapiens 350 – 40[51] 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[52] Taiwan

1 individual 2008(?)/2015

H. rhodesiensis also classified as H. heidelbergensis
H. heidelbergensis
or a subspecies of H. sapiens 300 – 120 Zambia

1,300 Very few 1921/1921

H. sapiens (anatomically modern humans) 200[53] – present Worldwide 150 - 190 cm (4 ft 7 in - 6 ft 3 in) 50–100 kg (110–220 lb) 950–1,800 (extant) —/1758

H. floresiensis classification uncertain 190 – 50 Indonesia 100 cm (3 ft 3 in) 25 kg (55 lb) 400 7 individuals 2003/2004

Denisova hominin possible H. sapiens subspecies or hybrid 40 Russia

1 site 2000/2010[54]

Red Deer Cave
Cave
people possible H. sapiens subspecies or hybrid 14.5–11.5 China

Very few 2012/—

See also

List of human evolution fossils
List of human evolution fossils
(with images) Nature timeline

References

^ 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 Human
Human
Evolution. Cambridge: Cambridge University Press. p. 242. CS1 maint: Uses editors parameter (link) McHenry, H.M (2009). " Human
Human
Evolution". 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 Human
Human
Evolution". 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 Homo habilis
Homo habilis
type OH 7 suggests deep-rooted species diversity in early Homo". Nature. 519 (7541): 83–86. doi:10.1038/nature14224. ISSN 0028-0836. PMID 25739632. . 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
Homo
at 2.8 Ma from Ledi-Geraru, Afar, Ethiopia". Science. 347 (6228): 1352–1355. doi:10.1126/science.aaa1343. ISSN 0036-8075. PMID 25739410.  ^ Curnoe, D (2010). "A review of early Homo
Homo
in southern Africa focusing on cranial, mandibular and dental remains, with the description of a new species ( Homo gautengensis
Homo gautengensis
sp. nov.)". HOMO – Journal of Comparative Human
Human
Biology. 61: 151–177. doi:10.1016/j.jchb.2010.04.002.  ^ Schuster, Angela M. H. (1997). "Earliest Remains of Genus
Genus
Homo". Archaeology. 50 (1). Retrieved 5 March 2015.  The line to the earliest members of Homo
Homo
made final separation from the lineage of Pan by late Miocene
Miocene
or early Pliocene
Pliocene
times—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. doi:10.1007/PL00006431. PMID 9847414.  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. doi:10.1038/nature04789. PMID 16710306.  Wakeley, J (2008). "Complex speciation of humans and chimpanzees". Nature. 452 (7184): E3–4. doi:10.1038/nature06805. PMID 18337768.  "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
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. doi:10.1126/science.1188021. PMC 5100745 . PMID 20448178.  ^ Lowery, R.K.; Uribe, G.; Jimenez, E.B.; Weiss, M.A.; Herrera, K.J.; Regueiro, M.; Herrera, R.J. (2013). " Neanderthal
Neanderthal
and Denisova genetic affinities with contemporary humans: Introgression versus common ancestral polymorphisms". Gene. 530 (1): 83–94. doi:10.1016/j.gene.2013.06.005. PMID 23872234.  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
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
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, Homo sapiens
Homo sapiens
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). " Homo naledi
Homo naledi
and the Problems with the Homo
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. doi:10.1126/science.284.5411.65. PMID 10102822.  ^ "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
William Plane Pycraft
(1928). ^ "African man", used by T. F. Dreyer (1935) for the Florisbad Skull he found in 1932 (also Homo
Homo
florisbadensis or Homo
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), 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 Arambourg in 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 Denisovan
Denisovan
Exome Archived 2013-10-29 at the Wayback Machine. ^ 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). " Human
Human
evolution: taxonomy and paleobiology". Journal of Anatomy. 197 (Pt 1): 19–60. doi:10.1046/j.1469-7580.2000.19710019.x. PMC 1468107 . PMID 10999270.  ^ Brunet, M.; et al. (2002). "A new hominid from the upper Miocene
Miocene
of Chad, central Africa". Nature. 418: 145–151. doi:10.1038/nature00879. PMID 12110880.  ^ Cela-Conde, C.J.; Ayala, F.J. (2003). "Genera of the human lineage". PNAS. 100 (13): 7684–7689. doi:10.1073/pnas.0832372100. PMC 164648 . PMID 12794185.  ^ Wood, B.; Lonergan, N. (2008). "The hominin fossil record: taxa, grades and clades" (PDF). J. Anat. 212: 354–376. doi:10.1111/j.1469-7580.2008.00871.x. PMC 2409102 . PMID 18380861.  ^ Cela-Conde, C. J.; Ayala, F. J. (2003). "Genera of the human lineage". Proceedings of the National Academy of Sciences. 100 (13): 7684–7689. doi:10.1073/pnas.0832372100. PMC 164648 . PMID 12794185.  ^ Pickering, R.; Dirks, P. H.; Jinnah, Z.; De Ruiter, D. J.; Churchill, S. E.; Herries, A. I.; Berger, L. R. (2011). " Australopithecus
Australopithecus
sediba at 1.977 Ma and implications for the origins of the genus Homo". Science. 333 (6048): 1421–1423. doi:10.1126/science.1203697. PMID 21903808.  ^ Asfaw, B.; White, T.; Lovejoy, O.; Latimer, B.; Simpson, S.; Suwa, G. (1999). " Australopithecus
Australopithecus
garhi: a new species of early hominid from Ethiopia". Science. 284 (5414): 629–635. doi:10.1126/science.284.5414.629. PMID 10213683.  ^ In 2010, evidence was presented that seems to attribute the use of stone tools to Australopithecus
Australopithecus
afarensis, 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. doi:10.1038/nature09248. PMID 20703305.  "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
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
Australopithecus
afarensis." ^ Erin N. DiMaggio EN; Campisano CJ; Rowan J; Dupont-Nivet G; Deino AL; et al. (2015). "Late Pliocene
Pliocene
fossiliferous sedimentary record and the environmental context of early Homo
Homo
from Afar, Ethiopia". Science. 347: 1355–1359. doi:10.1126/science.aaa1415. PMID 25739409.  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
BBC
News. Retrieved 5 March 2015.  ^ Cela-Conde and Ayala (2003) recognize five genera within Hominina: Ardipithecus, Australopithecus
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. doi:10.1073/pnas.0832372100. PMC 164648 . PMID 12794185.  ^ "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
Homo
fossils from Ileret, east of Lake Turkana, Kenya". Nature. 448 (7154): 688–691. doi:10.1038/nature05986. PMID 17687323.  ^ Strait, David; Grine, Frederick; Fleagle, John (2015-01-01). Analyzing Hominin Hominin Phylogeny: Cladistic Approach. pp. 1989–2014. ISBN 9783642399787.  ^ Green, RE; Krause, J; et al. (2010). "A draft sequence of the Neanderthal
Neanderthal
genome". Science. 328 (5979): 710–22. doi:10.1126/science.1188021. PMC 5100745 . PMID 20448178.  ^ Reich, D; Green, RE; Kircher, M; et al. (December 2010). "(December 2010). "Genetic history of an archaic hominin group from Denisova Cave in Siberia"". Nature. 468 (7327): 1053–60. doi:10.1038/nature09710. PMC 4306417 . PMID 21179161.  ^ Reich; et al. (October 2011). "Denisova admixture and the first modern human dispersals into southeast Asia and Oceania". Am J Hum Genet. 89 (4): 516–28. doi:10.1016/j.ajhg.2011.09.005. PMC 3188841 . PMID 21944045.  ^ Biological Anthropology: 2nd Edition. 2009. Craig Stanford et al. ^ Shaun Smillie, " Homo
Homo
naledi—New human ancestor buried its dead," Times Live, 10 Sept 2015. ^ Dirks, Paul H.G.M.; Roberts, Eric M.; et al. (2017). "The age of Homo naledi
Homo naledi
and associated sediments in the Rising Star Cave, South Africa". eLife (published 9 May 2017). 6: e24231. doi:10.7554/eLife.24231 . 

Full list of authors

Paul H.G.M. Dirks Eric M. Roberts Hannah Hilbert-Wolf Jan D. Kramers John Hawks Anthony Dosseto Mathieu Duval Marina Elliott Mary Evans Rainer Grün John Hellstrom Andy I.R. Herries Renaud Joannes-Boyau Tebogo V. Makhubela Christa J. Placzek Jessie Robbins Carl Spandler Jelle Wiersma Jon Woodhead Lee R. Berger

^ Rincon, Paul (9 May 2017). "Amazing haul of ancient human finds unveiled". BBC. Retrieved 9 May 2017.  ^ Berger, L. R.; Hawks, J.; Dirks, P. HGM; Elliott, M.; Roberts, E. M. (9 May 2017). " Homo naledi
Homo naledi
and Pleistocene
Pleistocene
hominin evolution in subequatorial Africa". eLife. 6. doi:10.7554/eLife.24234.  ^ Schrenk, Friedemann; Kullmer, Ottmar; Bromage, Timothy (2007). "The Earliest Putative Homo
Homo
Fossils". In Henke, Winfried; Tattersall, Ian. Handbook of Paleoanthropology. 1. In collaboration with Thorolf Hardt. Berlin, Heidelberg: Springer. pp. 1611–1631. doi:10.1007/978-3-540-33761-4_52. ISBN 978-3-540-32474-4.  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. Wilford, John Noble (August 9, 2007). "Fossils in Kenya Challenge Linear Evolution". The New York Times. Retrieved 2015-05-04. 

DiMaggio, Erin N.; Campisano, Christopher J.; Rowan, John; et al. (March 20, 2015). "Late Pliocene
Pliocene
fossiliferous sedimentary record and the environmental context of early Homo
Homo
from Afar, Ethiopia". Science. Washington, D.C.: American Association for the Advancement of Science. 347 (6228): 1355–1359. doi:10.1126/science.aaa1415. ISSN 0036-8075. PMID 25739409.  Hominins
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
Australopithecus
and Homo
Homo
discovered in 2015.

^ Haviland, William A.; Walrath, Dana; Prins, Harald E. L.; McBride, Bunny (2007). Evolution and Prehistory: The Human
Human
Challenge (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 Eurasia
Eurasia
and 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
Dmanisi
(Georgian Caucasus) dated to 1.85-1.78 Ma". Proceedings of the National Academy of Sciences. 108 (26): 10432. doi:10.1073/pnas.1106638108. PMC 3127884 . PMID 21646521.  ^ "New discovery suggests Homo erectus
Homo erectus
originated 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 Kenya
Kenya
Museum's Human
Human
Fossils". 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
Peking Man
(formerly Sinanthropus pekinensis) and Java Man
Java Man
(formerly Pithecanthropus erectus). H. erectus is now grouped into various subspecies, including Homo erectus
Homo erectus
erectus, Homo erectus
Homo erectus
yuanmouensis, Homo
Homo
erectus lantianensis, Homo erectus
Homo erectus
nankinensis, Homo erectus
Homo erectus
pekinensis, Homo erectus palaeojavanicus, Homo erectus
Homo erectus
soloensis, Homo
Homo
erectus tautavelensis, Homo erectus
Homo erectus
georgicus. The distinction from descendant species such as Homo
Homo
ergaster, Homo
Homo
floresiensis, Homo
Homo
antecessor, Homo heidelbergensis
Homo heidelbergensis
and indeed Homo sapiens
Homo sapiens
is not entirely clear. ^ Curnoe, Darren (June 2010). "A review of early Homo
Homo
in southern Africa focusing on cranial, mandibular and dental remains, with the description of a new species ( Homo gautengensis
Homo gautengensis
sp. nov.)". HOMO - Journal of Comparative Human
Human
Biology. Amsterdam, the Netherlands: Elsevier. 61 (3): 151–177. doi:10.1016/j.jchb.2010.04.002. ISSN 0018-442X. PMID 20466364.  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
H. ergaster
or Australopithecus
Australopithecus
by other anthropologists. ^ Hazarika, Manjil (2007). " Homo
Homo
erectus/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 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
H. heidelbergensis
are fossils found at Bilzingsleben (also classified as Homo erectus
Homo erectus
bilzingslebensis). ^ 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. doi:10.1006/jasc.2002.0834. ISSN 0305-4403.  The first humans with "proto-Neanderthal traits" lived in Eurasia
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), 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
Homo
from Taiwan". Nature Communications. 6: 6037. doi:10.1038/ncomms7037. PMC 4316746 . PMID 25625212.  ^ 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 has high as 300,000 years. In 2017, fossils found in Jebel Irhoud
Jebel Irhoud
(Morocco) suggest that Homo sapiens
Homo sapiens
may have speciated by as early as 315,000 years ago. Callaway, Ewan (7 June 2017). "Oldest Homo sapiens
Homo sapiens
fossil claim rewrites our species' history". Nature. doi:10.1038/nature.2017.22114. Retrieved 11 June 2017.  Genetic evidence has been adduced for an age of roughly 270,000 years. Posth, Cosimo; et al. (4 July 2017). "Deeply divergent archaic mitochondrial genome provides lower time boundary for African gene flow into Neanderthals". Nature Communications. doi:10.1038/ncomms16046. Retrieved 4 July 2017. CS1 maint: Explicit use of et al. (link) ^ provisional names Homo
Homo
sp. Altai or Homo sapiens
Homo sapiens
ssp. Denisova.

Further reading

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 DNA
DNA
contribution to early modern humans". PLoS Biology. 2 (3): 313–7. doi:10.1371/journal.pbio.0020057. PMC 368159 . PMID 15024415. 

External links

Wikimedia Commons has media related to Homo.

Wikispecies
Wikispecies
has information related to Homo

Wikibooks has a book on the topic of: Introduction to Paleoanthropology

Exploring the Hominid Fossil Record (Center for the Advanced Study of Hominid Paleobiology at George Washington University) Hominid species Prominent Hominid Fossils Mikko's Phylogeny archive Homo
Homo
at the Encyclopedia of Life
Encyclopedia of Life
Human
Human
Timeline (Interactive) – Smithsonian, National Museum of Natural History (August 2016).

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