Homo

Homo
Temporal range: Piacenzian-Present, 2.865–0 Ma
↓
A basic interpretation of Homo habilis
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Clade: Synapsida
Class: Mammalia
Order: Primates
Suborder: Haplorhini
Infraorder: Simiiformes
Family: Hominidae
Tribe: Hominini
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

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, most notably Homo erectus. The genus is between 2 and 3 million years old, taken to emerge with the appearance of Homo habilis[1] and possibly that of Homo gautengensis.[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 (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.

The subspecies 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 times they spread throughout Africa, Eurasia, Oceania, and the Americas, and they encountered archaic humans along the way during these migrations. Homo sapiens sapiens is the only surviving species and subspecies of the genus Homo. Archaic humans survived until about 40,000 years ago (see H. neanderthalensis),[4] 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.[5][6]

Names and taxonomy

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".[7] The binomial name Homo sapiens was coined by Carl Linnaeus (1758).[8] 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.[9][10][11] 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.[12]

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,[13] Protanthropus,[14] Sinanthropus,[15] Cyphanthropus,[16] Africanthropus,[17] Telanthropus,[18] Atlanthropus,[19] and Tchadanthropus.[20]

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.[21] 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.[22] 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.[23] Designations alternative to Hominina existed, or were offered: Australopithecinae (Gregory & Hellman 1939) and Preanthropinae (Cela-Conde & Altaba 2002);[24][25][26] and later, Cela-Conde and Ayala (2003) proposed that the four genera Australopithecus, Ardipithecus, Praeanthropus, and Sahelanthropus be grouped with Homo within Hominina.[27]

Evolution

Several species, including Australopithecus garhi, Australopithecus sediba, Australopithecus africanus, and Australopithecus afarensis, have been proposed as the direct ancestor of the Homo lineage.[28][29] 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.[30] The emergence of Homo also coincides roughly with the onset of Quaternary glaciation, the beginning of the current ice age.

A fossil mandible fragment dated to 2.8 million years ago which may represent an intermediate stage between Australopithecus and Homo was discovered in 2015 in Afar, Ethiopia (LD 350-1).[31] 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.[32]

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.[33]

Migration

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.[34] 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[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 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

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.[37]

Homo naledi was discovered near Johannesburg, 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.

Comparative table of Homo species
Species Temporal range kya Habitat Adult height Adult mass Cranial capacity (cm³) Fossil record Discovery / publication of name
H. habilis 2,100 – 1,500[39] Africa 110-140 cm (4 ft 11 in) 33–55 kg (73–121 lb) 510–660 Many 1960/1964
H. erectus 1,900 – 70

[40]

 Africa, Eurasia (Java, China, India, Caucasus) 180 cm (5 ft 11 in) 60 kg (130 lb) 850 (early) – 1,100 (late) Many[41] 1891/1892
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[42] 2010/2010
H. ergaster
also classified as H. erectus		 1,800 – 1,300[43] Eastern and Southern Africa 700–850 Many 1975
H. antecessor
also classified as H. heidelbergensis		 1,200 – 800 Spain 175 cm (5 ft 9 in) 90 kg (200 lb) 1,000 2 sites 1997
H. cepranensis
a single fossil, possibly H. erectus		 900 – 350 Italy 1,000 1 skull cap 1994/2003
H. heidelbergensis 600 – 350[44] Europe, Africa, China 180 cm (5 ft 11 in) 90 kg (200 lb) 1,100–1,400 Many 1908
H. neanderthalensis
possibly a subspecies of H. sapiens		 350 – 40[45] 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[46] Taiwan 1 individual pre-2008/2015
H. rhodesiensis
also classified as H. heidelbergensis		 300 – 120 Zambia 1,300 Very few 1921
H. sapiens
(modern humans) 		300[47]

 – 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
Red Deer Cave people
possible H. sapiens subspecies or hybrid		 14.5–11.5 China Very few 2012

See also

References

  1. ↑ 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 Evolution. Cambridge: Cambridge University Press. p. 242. 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. 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 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 type OH 7 suggests deep-rooted species diversity in early Homo". Nature. 519 (7541): 83–86. ISSN 0028-0836. PMID 25739632. doi:10.1038/nature14224.. Villmoare, Brian; Kimbel, William H.; Seyoum, Chalachew; Campisano, Christopher J.; DiMaggio, Erin N.; Rowan, John; Braun, David R.; Arrowsmith, J. Ramón; Reed, Kaye E. (2015-03-20). "Early Homo at 2.8 Ma from Ledi-Geraru, Afar, Ethiopia". Science. 347 (6228): 1352–1355. ISSN 0036-8075. PMID 25739410. doi:10.1126/science.aaa1343.
  2. ↑ Curnoe, D (2010). "A review of early Homo in southern Africa focusing on cranial, mandibular and dental remains, with the description of a new species (Homo gautengensis sp. nov.)". HOMO – Journal of Comparative Human Biology. 61: 151–177.
  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 made final separation from the lineage of Pan by late Miocene or early 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. PMID 9847414. doi:10.1007/PL00006431.
    • Patterson, N; Richter, DJ; Gnerre, S; Lander, ES; Reich, D (2006). "Genetic evidence for complex speciation of humans and chimpanzees". Nature. 441 (7097): 1103–8. PMID 16710306. doi:10.1038/nature04789.
    • Wakeley, J (2008). "Complex speciation of humans and chimpanzees". Nature. 452 (7184): E3–4. PMID 18337768. doi:10.1038/nature06805. "Patterson et al. suggest that the apparently short divergence time between humans and chimpanzees on the X chromosome is explained by a massive interspecific hybridization event in the ancestry of these two species. However, Patterson et al. do not statistically test their own null model of simple speciation before concluding that speciation was complex, and—even if the null model could be rejected—they do not consider other explanations of a short divergence time on the X chromosome. These include natural selection on the X chromosome in the common ancestor of humans and chimpanzees, changes in the ratio of male-to-female mutation rates over time, and less extreme versions of divergence with gene flow. I therefore believe that their claim of hybridization is unwarranted." see current estimates regarding complex speciation.
  4. ↑ Ghosh, Pallab (20 August 2014). "New dates rewrite Neanderthal story". BBC News.
  5. ↑ Green, R.E.; Krause, J.; Briggs, A.W.; Maricic, T.; Stenzel, U.; Kircher, M.; Patterson, N.; Li, H.; Zhai, W.; Fritz, M.H.Y.; Hansen, N.F. (2010). "A draft sequence of the Neandertal genome". Science. 328 (5979): 710–722. PMID 20448178. doi:10.1126/science.1188021.
  6. ↑ 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. PMID 23872234. doi:10.1016/j.gene.2013.06.005. This study raises the possibility of observed genetic affinities between archaic and modern human populations being mostly due to common ancestral polymorphisms.
  7. ↑ 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.
  8. ↑ 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.
  9. ↑ 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.
  10. ↑ 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.
  11. ↑ Wood, B.; Collard, M. (2 April 1999). "The human genus". Science. 284 (5411): 65–71. PMID 10102822. doi:10.1126/science.284.5411.65.
  12. ↑ Stringer, C. (2012). "What makes a modern human". Nature. 485 (7396): 33–35. PMID 22552077. doi:10.1038/485033a.
  13. ↑ "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
  14. ↑ "early man", Protanthropus primigenius Ernst Haeckel, Systematische Phylogenie vol. 3 (1895), p. 625
  15. ↑ "Sinic man", from Sinanthropus pekinensis (Peking Man), Davidson Black (1927).
  16. ↑ "crooked man", from Cyphanthropus rhodesiensis (Rhodesian Man) William Plane Pycraft (1928).
  17. ↑ "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.
  18. ↑ "remote man"; from Telanthropus capensis (Broom and Robinson 1949), see (1961), p. 487.
  19. ↑ 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.
  20. ↑ Y. Coppens, "L'Hominien du Tchad", Actes V Congr. PPEC I (1965), 329f.; "Le Tchadanthropus", Anthropologia 70 (1966), 5–16.
  21. ↑ Alexandra Vivelo (2013), Characterization of Unique Features of the Denisovan Exome
  22. ↑ 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.
  23. ↑ Wood and Richmond; Richmond, BG (2000). "Human evolution: taxonomy and paleobiology". Journal of Anatomy. 197 (Pt 1): 19–60. PMC 1468107 Freely accessible. PMID 10999270. doi:10.1046/j.1469-7580.2000.19710019.x.
  24. ↑ Brunet, M.; et al. (2002). "A new hominid from the upper Miocene of Chad, central Africa". Nature. 418: 145–151. PMID 12110880. doi:10.1038/nature00879.
  25. ↑ Cela-Conde, C.J.; Ayala, F.J. (2003). "Genera of the human lineage". PNAS. 100 (13): 7684–7689. PMC 164648 Freely accessible. PMID 12794185. doi:10.1073/pnas.0832372100.
  26. ↑ Wood, B.; Lonergan, N. (2008). "The hominin fossil record: taxa, grades and clades" (PDF). J. Anat. 212: 354–376. PMC 2409102 Freely accessible. PMID 18380861. doi:10.1111/j.1469-7580.2008.00871.x.
  27. ↑ Cela-Conde, C. J.; Ayala, F. J. (2003). "Genera of the human lineage". Proceedings of the National Academy of Sciences. 100 (13): 7684–7689. PMC 164648 Freely accessible. PMID 12794185. doi:10.1073/pnas.0832372100.
  28. ↑ 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. PMID 21903808. doi:10.1126/science.1203697.
  29. ↑ 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. PMID 10213683. doi:10.1126/science.284.5414.629.
  30. ↑ 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. PMID 20703305. doi:10.1038/nature09248. "The oldest direct evidence of stone tool manufacture comes from Gona (Ethiopia) and dates to between 2.6 and 2.5 million years (Myr) ago. [...] Here we report stone-tool-inflicted marks on bones found during recent survey work in Dikika, Ethiopia [... 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."
  31. ↑ 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. PMID 25739409. doi:10.1126/science.aaa1415. See also: "Oldest known member of human family found in Ethiopia". New Scientist. 4 March 2015. Retrieved 7 March 2015., Ghosh, Pallab (4 March 2015). "'First human' discovered in Ethiopia". BBC News. Retrieved 5 March 2015.
  32. ↑ Cela-Conde and Ayala (2003) recognize five genera within Hominina: Ardipithecus, Australopithecus (including Paranthropus), Homo (including Kenyanthropus), Praeanthropus (including Orrorin), and Sahelanthropus. Cela-Conde, C. J.; Ayala, F. J. (2003). "Genera of the human lineage". Proceedings of the National Academy of Sciences. 100 (13): 7684–7689. PMC 164648 Freely accessible. PMID 12794185. doi:10.1073/pnas.0832372100.
  33. ↑ "A partial maxilla assigned to H. habilis reliably demonstrates that this species survived until later than previously recognized, making an anagenetic relationship with H. erectus unlikely. The discovery of a particularly small calvaria of H. erectus indicates that this taxon overlapped in size with H. habilis, and may have shown marked sexual dimorphism. The new fossils confirm the distinctiveness of H. habilis and H. erectus, independently of overall cranial size, and suggest that these two early taxa were living broadly sympatrically in the same lake basin for almost half a million years." Spoor, F; Leakey, M.G; Gathogo, P.N; Brown, F.H; Antón, S.C; McDougall, I; Kiarie, C; Manthi, F.K.; Leakey, L.N. (2007). "Implications of new early Homo fossils from Ileret, east of Lake Turkana, Kenya". Nature. 448 (7154): 688–691. PMID 17687323. doi:10.1038/nature05986.
  34. ↑ Green, RE; Krause, J; et al. (2010). "A draft sequence of the Neanderthal genome". Science. 328 (5979): 710–22. PMID 20448178. doi:10.1126/science.1188021.
  35. ↑ 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. PMID 21179161. doi:10.1038/nature09710.
  36. ↑ 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. PMC 3188841 Freely accessible. PMID 21944045. doi:10.1016/j.ajhg.2011.09.005.
  37. ↑ Biological Anthropology: 2nd Edition. 2009. Craig Stanford et al.
  38. ↑ Shaun Smillie, "Homo naledi—New human ancestor buried its dead," Times Live, 10 Sept 2015.
  39. ↑ Schrenk, Friedemann; Kullmer, Ottmar; Bromage, Timothy (2007). "The Earliest Putative Homo Fossils". In Henke, Winfried; Tattersall, Ian. Handbook of Paleoanthropology. 1. In collaboration with Thorolf Hardt. Berlin, Heidelberg: Springer. pp. 1611–1631. ISBN 978-3-540-32474-4. doi:10.1007/978-3-540-33761-4_52. Confirmed H. habilis fossils are dated to between 2.1 and 1.5 million years ago. This date range overlaps with the emergence of 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 fossiliferous sedimentary record and the environmental context of early Homo from Afar, Ethiopia". Science. Washington, D.C.: American Association for the Advancement of Science. 347 (6228): 1355–1359. ISSN 0036-8075. PMID 25739409. doi:10.1126/science.aaa1415. 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.
  40. ↑ Haviland, William A.; Walrath, Dana; Prins, Harald E. L.; McBride, Bunny (2007). Evolution and Prehistory: The 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 and migrated back to Africa.
  41. ↑ 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.
  42. ↑ Curnoe, Darren (June 2010). "A review of early Homo in southern Africa focusing on cranial, mandibular and dental remains, with the description of a new species (Homo gautengensis sp. nov.)". HOMO - Journal of Comparative Human Biology. Amsterdam, the Netherlands: Elsevier. 61 (3): 151–177. ISSN 0018-442X. PMID 20466364. doi:10.1016/j.jchb.2010.04.002. A species proposed in 2010 based on the fossil remains of three individuals dated between 1.9 and 0.6 million years ago. The same fossils were also classified as H. habilis, H. ergaster or Australopithecus by other anthropologists.
  43. ↑ Hazarika, Manjil (2007). "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"
  44. ↑ 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).
  45. ↑ Bischoff, James L.; Shamp, Donald D.; Aramburu, Arantza; et al. (March 2003). "The Sima de los Huesos Hominids Date to Beyond U/Th Equilibrium (>350 kyr) and Perhaps to 400–500 kyr: New Radiometric Dates". Journal of Archaeological Science. Amsterdam, the Netherlands: Elsevier. 30 (3): 275–280. ISSN 0305-4403. doi:10.1006/jasc.2002.0834. The first humans with "proto-Neanderthal traits" lived in 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.
  46. ↑ Chang, Chun-Hsiang; Kaifu, Yousuke; Takai, Masanaru; Kono, Reiko T.; Grün, Rainer; Matsu’ura, Shuji; Kinsley, Les; Lin, Liang-Kong (2015). "The first archaic Homo from Taiwan". Nature Communications. 6: 6037. PMC 4316746 Freely accessible. PMID 25625212. doi:10.1038/ncomms7037.
  47. ↑ Callaway, Ewan (7 June 2017). "Oldest Homo sapiens fossil claim rewrites our species' history". Nature (journal). doi:10.1038/nature.2017.22114. Retrieved 7 June 2017.

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