Temporal range: Gelasian - Recent 2.5–0Ma
|Reconstruction of Homo erectus georgicus (Élisabeth Daynès, Musée de Préhistoire, Quinson, France)|
Homo is the genus of hominids that includes modern humans and species closely related to them. The genus is estimated to be about 2.5 million years old, emerging with the appearance of Homo habilis from australopithecine ancestors. It is the only genus in the subtribe Hominina, which together with the subtribe Australopithecina forms the tribe Hominini, estimated to have diverged from the genus Pan in the late Miocene, by about 7 million years ago.
Homo erectus appears about 2 to 4 million years ago and spreads throughout Eurasia. It was likely the first hominin to live in a hunter-gatherer society and to control fire. Homo sapiens (anatomically modern humans) emerges about 0.2 million years ago, and is the only surviving species in the genus, all others having become extinct. Archaic humans did survive alongside H. sapiens until at least some 40,000 years ago (H. neanderthalensis, and possibly until as late as the Epipaleolithic (about 12,000 years ago), and there is evidence of limited interbreeding of archaic and modern humans. Modern humans migrated from Africa some 60,000 years ago, and in the course of the Upper Paleolithic spread throughout Eurasia, Oceania and the Americas.
Name and taxonomy
- See Hominidae for an overview of taxonomy.
The binomial name Homo sapiens was coined by Carl Linnaeus (1758). Names for other species were introduced beginning in the second half of the 19th century (H. neanderthalensis 1864, H. erectus 1892).
The genus Homo was given this taxonomic name to suggest that its members can be classified as human, and while there is substantial debate on the delineation of the genus from Australopithecus or indeed from Pan, classification of fossils within Homo coincides with evidence of early tool use beginning some 2.5 million years ago. Some recently discovered, recently extinct, species in the genus Homo do not have accepted binomial names yet see (Denisova hominin and Red Deer Cave people).
In the history of paleoanthropological research during the late 19th to mid 20th century, a number of taxonomic names proposed for individual early human fossils included suggestions for new generic names which have since been merged with Homo (mostly due to the recognition of Homo erectus as a single species with a large geographic spread) and are now considered "synonyms" with Homo for systematic reasons. These generic names include Pithecanthropus, Protanthropus, Sinanthropus, Cyphanthropus, Africanthropus, Telanthropus, Atlanthropus, and Tchadanthropus.
Classification of the genus Homo into species and subspecies remains poorly defined and subject to incomplete information, leading to difficulties in binomial naming, and the use of common names, such as "Neanderthal" and "Denisovan", even in scientific papers, to avoid the ambiguity in the classification of species or subspecies incertae sedis within Homo (e.g. H. neanderthalensis vs. H. sapiens neanderthalensis, or H. georgicus vs. H. erectus georgicus).
Going back to a suggestion by John Edward Gray (1825) Hominini ("hominins") is defined as a tribe including all early human and pre-human species ancestral to modern humans, back to the chimpanzee–human last common ancestor. Hominina ("hominans") is a subtribe of Hominini, including only the genus Homo but not earlier upright walking hominins of the Pliocene such as Australopithecus, Orrorin tugenensis, Ardipithecus, or Sahelanthropus. Alternative names for Hominina are Australopithecinae (Gregory & Hellman 1939) and Preanthropinae (Cela-Conde & Altaba 2002). Cela-Conde and Ayala (2003) recognize five genera within Hominina: Ardipithecus, Australopithecus (including Paranthropus), Homo (including Kenyanthropus), Praeanthropus (including Orrorin), and Sahelanthropus.
Several species, including Australopithecus garhi, Australopithecus sediba, Australopithecus africanus, and Australopithecus afarensis, 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 co-incide 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 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. 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 autralopithecine species) into the Homo genus.
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 Homo genus, 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 evindence 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 with H. habilis for about half a million years ( ), during the early Calabrian.
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. 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 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. Today, only H. sapiens remains, with no other extant species.
List of species
The species status of Homo rudolfensis, Homo ergaster, H. georgicus, H. antecessor, H. cepranensis, H. rhodesiensis, Homo neanderthalensis, Denisova hominin, Red Deer Cave people and Homo 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.
|Species||Temporal range Mya||Habitat||Adult height||Adult mass||Cranial capacity (cm³)||Fossil record||Discovery / publication of name|
|H. habilis||2.1 – 1.4||Africa||150 cm (4 ft 11 in)||33–55 kg (73–121 lb)||510–660||Many||1960/1964|
|H. erectus||1.9 – 0.07||Africa, Eurasia (Java, China, India, Caucasus)||180 cm (5 ft 11 in)||60 kg (130 lb)||850 (early) – 1,100 (late)||Many||1891/1892|
membership in Homo uncertain
also classified as H. habilis
|1.9 – 0.6||South Africa||100 cm (3 ft 3 in)||3 individuals||2010/2010|
also classified as H. erectus
|1.8 – 1.3||Eastern and Southern Africa||700–850||Many||1975|
also classified as H. heidelbergensis
|1.2 – 0.8||Spain||175 cm (5 ft 9 in)||90 kg (200 lb)||1,000||2 sites||1997|
a single fossil, possibly H. erectus
|0.9 – 0.35||Italy||1,000||1 skull cap||1994/2003|
|H. heidelbergensis||0.6 – 0.35||Europe, Africa, China||180 cm (5 ft 11 in)||90 kg (200 lb)||1,100–1,400||Many||1908|
possibly a subspecies of H. sapiens
|0.35 – 0.04||Europe, Western Asia||170 cm (5 ft 7 in)||55–70 kg (121–154 lb) (heavily built)||1,200–1,900||Many||(1829)/1864|
also classified as H. heidelbergensis
|0.3 – 0.12||Zambia||1,300||Very few||1921|
possibly H. erectus
|0.25 – 0.2||Taiwan||1 individual||pre-2008/2015|
|Worldwide||150 - 190 cm (4 ft 7 in - 6 ft 3 in)||50–100 kg (110–220 lb)||950–1,800||(extant)||—/1758|
|0.10 – 0.012||Indonesia||100 cm (3 ft 3 in)||25 kg (55 lb)||400||7 individuals||2003/2004|
possible H. sapiens subspecies or hybrid
|Red Deer Cave people
possible H. sapiens subspecies or hybrid
- List of human evolution fossils (with images)
- Stringer, C.B. (1994). "Evolution of early humans". In Steve Jones, Robert Martin & David Pilbeam (eds.). The Cambridge Encyclopedia of Human Evolution. Cambridge: Cambridge University Press. p. 242. ISBN 0-521-32370-3. Also ISBN 0-521-46786-1 (paperback)
- McHenry, H.M (2009). "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.
- Schuster, Angela M. H. (1997). "Earliest Remains of Genus Homo". Archaeology 50 (1). Retrieved 5 March 2015.
- , BBC
- 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, Homo sapiens was validly defined as the animal species to which Linnaeus belonged.
- "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 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.
- "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. C. Arambourg, "A recent discovery in human paleontology: Atlanthropus of ternifine (Algeria)", American Journal of Physical Anthropology 13.2 (June 1955), 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 Exome
- 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 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 of Chad, central Africa. Nature (London), 418: 145-151. Cela-Conde, C.J. and Ayala, F.J., 2003: Genera of the human lineage. PNAS, 100(13): 7684-7689. Wood, B.; Lonergan, N., 2008: The hominin fossil record: taxa, grades and clades. J. Anat., 212: 354–376. PDF
- C. J. Cela-Conde and F. J. Ayala. 2003. "Genera of the human lineage". Proceedings of the National Academy of Sciences 100(13):7684-7689.
- Stringer, C. (2012). "What makes a modern human". Nature 485 (7396): 33–35. doi:10.1038/485033a. PMID 22552077.
- Pickering, R.; Dirks, P. H.; Jinnah, Z.; De Ruiter, D. J.; Churchill, S. E.; Herries, A. I.; Berger, L. R. (2011). "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.
- Asfaw, B.; White, T.; Lovejoy, O.; Latimer, B.; Simpson, S.; Suwa, G. (1999). "Australopithecus garhi: a new species of early hominid from Ethiopia". Science 284 (5414): 629–635. doi:10.1126/science.284.5414.629.
- 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. "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."
- Erin N. DiMaggio EN, Campisano CJ, Rowan J, Dupont-Nivet G, Deino AL et al. "Late Pliocene fossiliferous sedimentary record and the environmental context of early Homo from Afar, Ethiopia". Science. 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. C. J. Cela-Conde and F. J. Ayala. 2003. "Genera of the human lineage". Proceedings of the National Academy of Sciences 100(13):7684-7689.
- "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.
- Green, RE; Krause, J et al. (2010). "A draft sequence of the Neanderthal genome". Science 328 (5979): 710–22. doi:10.1126/science.1188021. 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. PMID 21179161.
- Reich et al. (Oct 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.
- Confirmed H. habilis fossils are dated to between 2.1 and 1.44 million years ago. 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 This date range overlaps with the emergence of Homo erectus. New York Times article Fossils in Kenya Challenge Linear Evolution published August 9, 2007. Hominins with "proto-Homo" traits may have lived as early as 2.8 million years ago, as suggested by a fossil jawbone classified as transitional between Australopithecus and Homo discovered in 2015. Erin N. DiMaggio EN, Campisano CJ, Rowan J, Dupont-Nivet G, Deino AL et al. "Late Pliocene fossiliferous sedimentary record and the environmental context of early Homo from Afar, Ethiopia". Science. doi:10.1126/science.aaa1415.
- 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 and migrated back to Africa. Frazier, Kendrick (Nov–Dec 2006). "Leakey Fights Church Campaign to Downgrade Kenya Museum’s Human Fossils". Skeptical Inquirer magazine 30 (6). Archived from the original on 2009-01-10. Prins, Harald E. L.; Walrath, Dana and McBride, Bunny (2007). Evolution and prehistory: the human challenge. Wadsworth Publishing. p. 162. ISBN 978-0-495-38190-7. 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. doi:10.1073/pnas.1106638108. New discovery suggests Homo erectus originated from Asia. Dnaindia.com. 8 June 2011.
- 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 erectus erectus, Homo erectus yuanmouensis, Homo erectus lantianensis, Homo erectus nankinensis, Homo erectus pekinensis, Homo erectus palaeojavanicus, Homo erectus soloensis, Homo erectus tautavelensis, Homo erectus georgicus. The distinction from descendant species such as Homo ergaster, Homo floresiensis, Homo antecessor, Homo heidelbergensis and indeed Homo sapiens is not entirely clear.
- 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. D. Curnoe, "A review of early Homo in southern Africa focusing on cranial, mandibular and dental remains, with the description of a new species (Homo gautengensis sp. nov.)." HOMO - Journal of Comparative Human Biology, vol.61 (2010), pp. 151–177.
- Hazarika, Manji (16–30 June 2007). "Homo erectus/ergaster and Out of Africa: Recent Developments in Paleoanthropology and Prehistoric Archaeology".
- 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 erectus bilzingslebensis).
- The first humans with "proto-Neanderthal traits" lived in Eurasia as early as 0.6 to 0.35 million years ago (classified as H. heidelbergensis, also called a chronospecies because it represents a chronological grouping rather than being based on clear morphological distinctions from either H. erectus or H. neanderthalensis), with the first "true Neanderthals" appearing between 0.25 and 0.2 million years ago. Bischoff, James L.; Shamp, Donald D.; Aramburu, Arantza; Arsuaga, Juan Luis; Carbonell, Eudald; Bermudez de Castro, J.M. (2003). "The Sima de los Huesos Hominids Date to Beyond U/Th Equilibrium (>350kyr) and Perhaps to 400–500kyr: New Radiometric Dates". Journal of Archaeological Science 30 (3): 275–80. doi:10.1006/jasc.2002.0834. Papagianni, Dmitra; Morse, Michael (2013). The Neandethals Rediscovered. Thames & Hudson. ISBN 978-0-500-05177-1.
- The oldest fossil remains of anatomically modern humans are the Omo remains, which date to 195,000 (±5,000) years ago and include two partial skulls as well as arm, leg, foot and pelvis bones. "Fossil Reanalysis Pushes Back Origin of Homo sapiens". Scientific American. February 17, 2005. McDougall, Ian; Brown, Francis H.; Fleagle, John G. (17 February 2005). "Stratigraphic placement and age of modern humans from Kibish, Ethiopia". Nature 433 (7027): 733–736. Bibcode:2005Natur.433..733M. doi:10.1038/nature03258. PMID 15716951. H. sapiens idaltu is a confirmed subspecies, based on 3 craniums dated 0.16 – 0.15 Mya found in Ethiopia (1997/2003).
- 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 mtDNA contribution to early modern humans". PLoS Biology 2 (3): 313–7. doi:10.1371/journal.pbio.0020057. PMC 368159. PMID 15024415.
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- Bradshaw Foundation Origins – Exploring the Fossil Record (with Center for the Advanced Study of Hominid Paleobiology at George Washington University)
- Hominid species
- Prominent Hominid Fossils
- Mikko's Phylogeny archive
- Homo at the Encyclopedia of Life