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Homo

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This is an old revision of this page, as edited by Sumanuil (talk | contribs) at 00:05, 14 October 2018 (Reverted to revision 862160828 by AGIOS PAISIOS (talk): It ain't around any more. I'd call that 'extinct'. (TW)). The present address (URL) is a permanent link to this revision, which may differ significantly from the current revision.

"Genus Homo" redirects here. For the novel by L. Sprague de Camp and P. Schuyler Miller, see Genus Homo (novel).
For other uses, see Homo (disambiguation).

Homo
Temporal range: Piacenzian-Present, 2.1–0 Ma
↓
File:Homo erectus adult female - head model - Smithsonian Museum of Natural History - 2012-05-17.jpg
Forensic reconstruction of an adult female Homo erectus[1]
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Primates
Suborder: Haplorhini
Infraorder: Simiiformes
Family: Hominidae
Subfamily: Homininae
Tribe: Hominini
Subtribe: Hominina
Genus: Homo
Linnaeus, 1758
Type species
Homo sapiens
Linnaeus, 1758
Species

For other species or subspecies suggested, see below.

Synonyms
Synonyms
  • Africanthropus Dreyer, 1935
  • Atlanthropus Arambourg, 1954
  • Cyphanthropus Pycraft, 1928
  • Palaeanthropus Bonarelli, 1909
  • Palaeoanthropus Freudenberg, 1927
  • Pithecanthropus Dubois, 1894
  • Protanthropus Haeckel, 1895
  • Sinanthropus Black, 1927
  • Tchadanthropus Coppens, 1965
  • Telanthropus Broom & Anderson 1949

Homo (Latin homō "human being") is the genus that encompasses the extant species Homo sapiens (modern humans), plus several extinct species classified as either ancestral to or closely related to modern humans (depending on a species), most notably Homo erectus and Homo neanderthalensis. The genus is taken to emerge with the appearance of Homo habilis, just more than two million years ago.[2] Homo is derived from the genus Australopithecus, which itself had previously split from the lineage of Pan, the chimpanzees.[3] Taxonomically, Homo is the only genus assigned to the subtribe Hominina which, with the subtribes Australopithecina and Panina, comprise the tribe Hominini.

Homo erectus appeared about two million years ago and, in several early migrations, it spread throughout Africa (where it is dubbed Homo ergaster) and Eurasia. It was likely the first human species to live in a hunter-gatherer society and to control fire. An adaptive and successful species, Homo erectus persisted for more than a million years, and gradually diverged into new species by around 500,000 years ago[4].

Homo sapiens (anatomically modern humans) emerges close to 300,000 to 200,000 years ago,[5] most likely in Africa, and Homo neanderthalensis emerges at 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, the so-called Southern Dispersal beginning about 70,000 years ago leading to the lasting colonisation of Eurasia and Oceania by 50,000 years ago. Both in Africa and Eurasia, H. sapiens met with and interbred with[6][7] archaic humans. Separate archaic (non-sapiens) human species are thought to have survived until around 40,000 years ago (Neanderthal extinction), with possible late survival of hybrid species as late as 12,000 years ago (Red Deer Cave people).

Among extant populations of Homo sapiens, the deepest temporal division is found in the San people of Southern Africa, estimated at close to 130,000 years.[8]

Names and taxonomy

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.
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. Late survival of robust australopithecines (Paranthropus) alongside Homo until 1.2 Mya is indicated in purple.

Main articles: Human taxonomy and Names for the human species
See Homininae for an overview of taxonomy.

The Latin noun homō (genitive hominis) means "human being" or "man" in the generic sense of "human being, mankind".[9] The binomial name Homo sapiens was coined by Carl Linnaeus (1758).[10] 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.[11][12][13] 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 Linnaeus did not even bother to 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, 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 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 and singular species with a large geographic spread of early migrations. Many such names are now dubbed as "synonyms" with Homo, including Pithecanthropus,[14] Protanthropus,[15] Sinanthropus,[16] Cyphanthropus,[17] Africanthropus,[18] Telanthropus,[19] Atlanthropus,[20] and Tchadanthropus.[21]

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.[22] Some recently extinct species in the genus Homo are only recently discovered and do not as yet have consensus binomial names (see Denisova hominin and Red Deer Cave people).[23] Since the beginning of the Holocene, it is likely that Homo sapiens (anatomically modern humans) have been the only extant species of Homo.

John Edward Gray (1825) was an early advocate of classifying taxa by designating tribes and families.[24] 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 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.[25] Designations alternative to Hominina existed, or were offered: Australopithecinae (Gregory & Hellman 1939) and Preanthropinae (Cela-Conde & Altaba 2002);[26][27][28] and later, Cela-Conde and Ayala (2003) proposed that the four genera Australopithecus, Ardipithecus, Praeanthropus, and Sahelanthropus be grouped with Homo within Hominina.[failed verification][29]

Evolution

Further information: Timeline of human evolution
See Hominini and Chimpanzee–human last common ancestor for the separation of Australopithecina and Panina.

Australopithecus

Further information: Australopithecus
File:Australopithecus afarensis adult male - head model - Smithsonian Museum of Natural History - 2012-05-17.jpg
Forensic reconstruction of A. afarensis[30]

Several species, including Australopithecus garhi, Australopithecus sediba, Australopithecus africanus, and Australopithecus afarensis, have been proposed as the direct ancestor of the Homo lineage.[31][32] 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 from 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.[33] LD 350-1, a fossil mandible fragment dated to 2.8 Mya, discovered in 2015 in Afar, Ethiopia, was described as combining "primitive traits seen in early Australopithecus with derived morphology observed in later Homo.[34] Some authors would push the development of Homo close to or even past 3 Mya.[35] 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. [36]

The most salient physiological development between the earlier australopithecine species and Homo is the increase in endocranial volume (ECV), from about 460 cm3 (28 cu in) in A. garhi to 660 cm3 (40 cu in) in H. habilis and further to 760 cm3 (46 cu in) in H. erectus, 1,250 cm3 (76 cu in) in H. heidelbergensis and up to 1,760 cm3 (107 cu in) in H. neanderthalensis. However, a steady rise in cranial capacity is observed already in Autralopithecina 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.[37]

Homo habilis

File:Homo habilis.JPG
Forensic reconstruction of Homo habilis, exhibit in LWL-Museum für Archäologie, Herne, Germany (2007 photograph).[38]

Homo habilis emerged about 2.1 Mya. Already before 2010, there were suggestions that H. habilis should not be placed in genus Homo but rather in Australopithecus.[39][40] 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.[33] 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.[41] 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[42] and widely dispersed throughout Eurasia (including Europe, Indonesia, China) by 0.5 Mya.[43]

Homo erectus

Main article: Homo erectus

Homo erectus has often been assumed to have developed anagenetically from Homo 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. 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, 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.[44]

A separate South African species Homo gautengensis has been postulated as contemporary with Homo erectus in 2010.[45]

Cladogram

The cladogram below depicts most species of Homo, with Australopithecus as an outgroup, according to Strait, Fleagle and Grine (2015).[46]

Hominina (4.5 Ma)

†Australopithecus

Homo (2.5 Ma)
2.1 Ma
1.9 Ma

†Homo ergaster

†Homo erectus

1.2 Ma

†Homo antecessor ssp.

0.8 Ma
0.3 Ma

Homo sapiens

Dispersal

See also: Early human expansions out of Africa, Archaic human admixture with modern humans, and Early human migrations

By about 1.8 million years ago, Homo erectus is present in both East Africa (Homo ergaster) and in Western Asia (Homo georgicus). The ancestors of Indonesian Homo floresiensis may have left Africa even earlier.[47]

Homo erectus and related or derived archaic human species over the next 1.5 million years spread throughout Africa and Eurasia.[48] Europe is reached by about 0.5 Mya by Homo heidelbergensis.

Homo neanderthalensis and Homo 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. 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.[49] H. sapiens interbred with archaic humans both in Africa and in Eurasia, in Eurasia notably with Neanderthals and Denisovans. [50]

Among extant populations of Homo sapiens, the deepest temporal division is found in the San people of Southern Africa, estimated at close to 130,000 years,[8] or possibly more than 300,000 years ago.[51] 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 (Red Deer Cave people), although they were mostly extinct or absorbed by the expanding H. sapiens populations by 40 kya (Neanderthal extinction).

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.

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, Homo habilis (2.1–1.5 Mya, membership in Homo questionable), Homo erectus (1.8–0.1 Mya, including the majority of the age of the genus, and the majority of archaic varieties as subspecies,[52] including H. heidelbergensis as a late or transitional variety[53]) and Homo sapiens (300 kya to present, including H. neanderthalensis and other varieties as subspecies).

Comparative table of Homo lineages
Lineages Temporal range
(kya) 		Habitat Adult height Adult mass Cranial capacity
(cm3) 		Fossil record Discovery Publication
of name
H. habilis
membership in Homo uncertain 		2,100–1,500[a][b] Tanzania 110–140 cm (3 ft 7 in – 4 ft 7 in) 33–55 kg (73–121 lb) 510–660 Many 1960 1964
H. rudolfensis
membership in 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[56][c] 2010 2010
H. erectus 1,900–140[57][d][58][e] Africa, Eurasia 180 cm (5 ft 11 in) 60 kg (130 lb) 850 (early) – 1,100 (late) Many[f][g] 1891 1892
H. ergaster
African H. erectus 		1,800–1,300[60] East and Southern Africa 700–850 Many 1949 1975
H. antecessor 1,200–800 Western Europe 175 cm (5 ft 9 in) 90 kg (200 lb) 1,000 2 sites 1994 1997
H. heidelbergensis
early H. neanderthalensis 		600–300[h] Europe, Africa 180 cm (5 ft 11 in) 90 kg (200 lb) 1,100–1,400 Many 1907 1908
H. cepranensis
a single fossil, possibly H. heidelbergensis 		c. 450[61] Italy 1,000 1 skull cap 1994 2003
H. longi 309–138[62] Northeast China 1,420[63] 1 individual 1933 2021
H. rhodesiensis
early H. sapiens 		c. 300 Zambia 1,300 Single or very few 1921 1921
H. naledi c. 300[64] South Africa 150 cm (4 ft 11 in) 45 kg (99 lb) 450 15 individuals 2013 2015
H. sapiens
(anatomically modern humans) 		c. 300–present[i] Worldwide 150–190 cm (4 ft 11 in – 6 ft 3 in) 50–100 kg (110–220 lb) 950–1,800 (extant) —— 1758
H. neanderthalensis
 240–40[67][j] 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. floresiensis
classification uncertain 		190–50 Indonesia 100 cm (3 ft 3 in) 25 kg (55 lb) 400 7 individuals 2003 2004
Nesher Ramla Homo
classification uncertain 		140–120 Israel several individuals 2021
H. tsaichangensis
possibly H. erectus or Denisova 		c. 100[k] Taiwan 1 individual 2008(?) 2015
H. luzonensis
 c. 67[70][71] Philippines 3 individuals 2007 2019
Denisova hominin 40 Siberia 2 sites 2000
 2010[l]

See also

References

  1. ^ Reconstruction by John Gurche (2010), Smithsonian Museum of Natural History, based on KNM ER 3733 and 992. Abigail Tucker, "A Closer Look at Evolutionary Faces", Smithsonian.com, 25 February 2010. H. erectus has the most extensive range of all species of Homo, from 1.8 to 0.14 Mya, or some 80% of the entire lifetime of the genus.
  2. ^ 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". The Cambridge Encyclopedia of Human Evolution. Cambridge: Cambridge University Press. p. 242. {{cite book}}: Unknown parameter |editors= ignored (|editor= suggested) (help) Friedemann Schrenk, Ottmar Kullmer, Timothy Bromage, "The Earliest Putative Homo Fossils", chapter 9 in: Winfried Henke, Ian Tattersall (eds.), Handbook of Paleoanthropology, 2007, pp 1611–1631, doi:10.1007/978-3-540-33761-4_52. Suggestions for pushing back the age to 2.8 Mya were made in 2015 based on the discovery of a jawbone: Spoor, Fred; Gunz, Philipp; Neubauer, Simon; Stelzer, Stefanie; Scott, Nadia; Kwekason, Amandus; Dean, M. Christopher (March 5, 2015). "Reconstructed Homo habilis type OH 7 suggests deep-rooted species diversity in early Homo". Nature. 519 (7541): 83–86. Bibcode:2015Natur.519...83S. doi:10.1038/nature14224. ISSN 0028-0836. PMID 25739632..
  3. ^ 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 were derived from Australopithecus, a genus which had separated from the Chimpanzee–human last common ancestor by late Miocene or early Pliocene times.
  4. ^ H. erectus in the narrow sense (the Asian species) was extinct by 140,000 years ago, 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. Indriati E, Swisher CC III, Lepre C, Quinn RL, Suriyanto RA, et al. 2011 The Age of the 20 Meter Solo River Terrace, Java, Indonesia and the Survival of Homo erectus in Asia.PLoS ONE 6(6): e21562. doi:10.1371/journal.pone.0021562.
  5. ^ Callaway, Ewan (7 June 2017). "Oldest Homo sapiens fossil claim rewrites our species' history". Nature. doi:10.1038/nature.2017.22114. Retrieved 11 June 2017.
  6. ^ 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. Bibcode:2010Sci...328..710G. doi:10.1126/science.1188021. PMC 5100745. PMID 20448178.
  7. ^ Lowery, R.K.; Uribe, G.; Jimenez, E.B.; Weiss, M.A.; Herrera, K.J.; Regueiro, M.; Herrera, R.J. (2013). "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.
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  9. ^ 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.
  10. ^ 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, Homo sapiens was validly defined as the animal species to which Linnaeus belonged.
  11. ^ Schwartz, Jeffrey H.; Tattersall, Ian (28 August 2015). "Defining the genus Homo". Science. 349 (6251): 931–932. Bibcode:2015Sci...349..931S. doi:10.1126/science.aac6182. Retrieved 2015-11-02.
  12. ^ Lents, Nathan (4 October 2014). "Homo naledi and the Problems with the Homo Genus". The Wildernist. Archived from the original on 18 November 2015. Retrieved 2015-11-02. {{cite news}}: Unknown parameter |dead-url= ignored (|url-status= suggested) (help)
  13. ^ Wood, B.; Collard, M. (2 April 1999). "The human genus". Science. 284 (5411): 65–71. doi:10.1126/science.284.5411.65. PMID 10102822.
  14. ^ "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
  15. ^ "early man", Protanthropus primigenius Ernst Haeckel, Systematische Phylogenie vol. 3 (1895), p. 625
  16. ^ "Sinic man", from Sinanthropus pekinensis (Peking Man), Davidson Black (1927).
  17. ^ "crooked man", from Cyphanthropus rhodesiensis (Rhodesian Man) William Plane Pycraft (1928).
  18. ^ "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), p. 43.
  19. ^ "remote man"; from Telanthropus capensis (Broom and Robinson 1949), see (1961), p. 487.
  20. ^ 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.
  21. ^ Y. Coppens, "L'Hominien du Tchad", Actes V Congr. PPEC I (1965), 329f.; "Le Tchadanthropus", Anthropologia 70 (1966), 5–16.
  22. ^ Alexandra Vivelo (2013), Characterization of Unique Features of the Denisovan Exome Archived 2013-10-29 at the Wayback Machine
  23. ^ Barras, Colin (2012-03-14). "Chinese human fossils unlike any known species". New Scientist. Retrieved 2012-03-15.
  24. ^ 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.
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  30. ^ Reconstruction by John Gurche (2010),Smithsonian Museum of Natural History. Abigail Tucker, "A Closer Look at Evolutionary Faces", Smithsonian.com, February 25, 2010.
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