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Homo

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"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, 2.865–0 Ma
↓
File:Homo georgicus.jpg
Reconstruction of Homo erectus georgicus (Élisabeth Daynès, Musée de Préhistoire, Quinson, France)
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

Homo sapiens
†Homo erectus
†Homo floresiensis
†Homo habilis
†Homo heidelbergensis
†Homo naledi
†Homo neanderthalensis
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 is the genus that comprises the species Homo sapiens, which includes modern humans, as well as several extinct species classified as ancestral to or closely related to modern humans—as for examples Homo habilis and Homo neanderthalensis. The genus is about 2.8 million years old;[1][2][3][4][5] it first appeared as its earliest species Homo habilis, which emerged from the genus Australopithecus, which itself had previously split from the lineage of Pan, the chimpanzees.[6][7] Taxonomically, Homo is the only genus assigned to the subtribe Hominina which, with the subtribes Australopithecina and Panina, comprise the tribe Hominini (see evolutionary tree below). All species of the genus Homo plus those species of the australopithecines that arose after the split from Pan are called hominins.

Homo erectus appeared about two million years ago in East Africa (where it is dubbed 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 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.

Homo sapiens sapiens, or anatomically modern humans, emerged about 200,000 years ago (0.2 Ma) in East Africa (see Omo remains). There is division among scholars as to when H. s. sapiens became behaviorally modern; the debate is: modern behavior developed 1) simultaneously with anatomical development, or 2) separately, and was complete by 50,000 years ago (see Modern human behavior). Homo sapiens sapiens is the only surviving species and subspecies of the genus Homo; all others have become extinct.

Modern humans migrated from Africa as recently as 60,000 years ago, and during Upper Paleolithic times they spread throughout Africa, Eurasia, Oceania, and the Americas; and they encountered archaic humans en route of their migrations. Some archaic humans outside Africa survived alongside modern humans until about 40,000 years ago (see H. neanderthalensis),[8] and possibly until as late as the times of the Epipaleolithic culture (about 12,000 years ago). DNA analysis provides some evidence of interbreeding between archaic and modern humans.[9][10]

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.

Further information: List of alternative names for the human species
See Hominidae for an overview of taxonomy.

The Latin noun homō (genitive hominis) means "human being" or "man" in the generic sense of "human being, mankind".[11] The binomial name Homo sapiens was coined by Carl Linnaeus (1758).[12] 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.[13][14][15] 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.

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.[16]

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 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,[17] Protanthropus,[18] Sinanthropus,[19] Cyphanthropus,[20] Africanthropus,[21] Telanthropus,[22] Atlanthropus,[23] and Tchadanthropus.[24]

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.[25] 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).

John Edward Gray (1825) was an early advocate of classifying taxa by designating tribes and families.[26] 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.[27] Designations alternative to Hominina existed, or were offered: Australopithecinae (Gregory & Hellman 1939) and Preanthropinae (Cela-Conde & Altaba 2002);[28][29][30] and later, Cela-Conde and Ayala (2003) proposed that the four genera Australopithecus, Ardipithecus, Praeanthropus, and Sahelanthropus be grouped with Homo within Hominina.[31]

Evolution

Further information: Timeline of human evolution, Hominina, Archaic humans, and Australopithecus

Several species, including Australopithecus garhi, Australopithecus sediba, Australopithecus africanus, and Australopithecus afarensis, have been proposed as the direct ancestor of the Homo lineage.[32][33] 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.[34] The emergence of Homo also coincides roughly with the onset of Quaternary glaciation, the beginning of the current ice age.

A fossil jawbone dated to 2.8 million years ago which may represent an intermediate stage between Australopithecus and Homo was discovered in 2015 in Afar, Ethiopia.[35] 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.[36]

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.

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.[37]

Migration

See also: Human evolution and Archaic human admixture with modern humans

Some of H. ergaster migrated to Asia, where they are named Homo erectus, and to Europe with Homo georgicus. H. ergaster in Africa and H. erectus in Eurasia evolved separately for almost two million years and presumably separated into two different species.

Homo rhodesiensis, who were descended from H. ergaster, migrated from Africa to Europe and became Homo heidelbergensis and later (about 250,000 years ago) Homo neanderthalensis and the Denisova hominin in Asia. The first 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 and met with resident Neanderthals, with some admixture.[38] Later, about 70,000 years ago, perhaps after the Toba catastrophe, a small group left the Levant to populate Eurasia, Australia and later the Americas. A subgroup among them met the Denisovans[39] 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 also show that 30,000 years ago at least three major subspecies coexisted: Denisovans, Neanderthals and anatomically modern humans.[40] 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 Neanderthal specimen from Vindija Cave has 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 DNA results indicate about 30% of derived alleles in H. sapiens are also in the Neanderthal lineage. This high frequency may suggest some gene flow between ancestral human and Neanderthal populations due to mating between the two.[41]

Homo naledi was discovered near Johannesburg, South Africa in 2013 and announced on 10 September 2015. Fossils indicate the hominid was 1.45-1.5 meters tall and had a small brain.[42] The fossils have yet to be dated.[43]

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[46][c] 2010 2010
H. erectus 1,900–140[47][d][48][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[50] 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[51] Italy 1,000 1 skull cap 1994 2003
H. longi 309–138[52] Northeast China 1,420[53] 1 individual 1933 2021
H. rhodesiensis
early H. sapiens 		c. 300 Zambia 1,300 Single or very few 1921 1921
H. naledi c. 300[54] 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[57][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[60][61] Philippines 3 individuals 2007 2019
Denisova hominin 40 Siberia 2 sites 2000
 2010[l]

See also

References

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    • 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.
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  19. ^ "Sinic man", from Sinanthropus pekinensis (Peking Man), Davidson Black (1927).
  20. ^ "crooked man", from Cyphanthropus rhodesiensis (Rhodesian Man) William Plane Pycraft (1928).
  21. ^ "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.
  22. ^ "remote man"; from Telanthropus capensis (Broom and Robinson 1949), see (1961), p. 487.
  23. ^ 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.
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  25. ^ Alexandra Vivelo (2013), Characterization of Unique Features of the Denisovan Exome
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  34. ^ In 2010, evidence was presented that seems to attribute the use of stone tools to 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 ago [...] Our discovery extends by approximately 800,000 years the antiquity of stone tools and of stone-tool-assisted consumption of ungulates by hominins; furthermore, this behaviour can now be attributed to Australopithecus afarensis."
  35. ^ Erin N. DiMaggio EN; Campisano CJ; Rowan J; Dupont-Nivet G; Deino AL; et al. (2015). "Late Pliocene fossiliferous sedimentary record and the environmental context of early 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. {{cite web}}: Italic or bold markup not allowed in: |publisher= (help), Ghosh, Pallab (4 March 2015). "'First human' discovered in Ethiopia". BBC News. Retrieved 5 March 2015.
  36. ^ 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. doi:10.1073/pnas.0832372100. PMC 164648. PMID 12794185.
  37. ^ "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. doi:10.1038/nature05986. PMID 17687323.
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Further reading

Wikimedia Commons has media related to Homo.
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