Temporal range: Piacenzian-Present, 2.865–0 Ma
Human taxonomy is the classification of the human species (systematic name Homo sapiens) within zoological taxonomy. The systematic genus, Homo, is designed to include both anatomically modern humans and extinct varieties of archaic humans.
Since the introduction of systematic names in the 18th century, knowledge of human evolution has increased drastically, and a number of intermediate taxa have been proposed in the 20th to early 21st century. The most widely accepted taxonomy groups takes the genus Homo as originating between two and three million years ago, divided into at least two species, archaic Homo erectus and modern Homo sapiens, with about a dozen further suggestions for species without universal recognition.
The genus Homo is placed in the tribe Hominini alongside Pan (chimpanzees) . The two genera are estimated to have diverged over an extended time of hybridization spanning roughly 10 to 6 million years ago, with possible admixture as late as 4 million years ago. A subtribe of uncertain validity, grouping archaic "pre-human" or "para-human" species younger than the Homo-Pan split is Australopithecina (proposed in 1939).
A proposal by Wood and Richmond (2000) would introduce Hominina as a subtribe alongside Australopithecina, with Homo the only known genus within Hominina. Alternatively, following Cela-Conde and Ayala (2003), the "pre-human" or "proto-human" genera of Australopithecus, Ardipithecus, Praeanthropus, and possibly Sahelanthropus may be placed on equal footing alongside the genus Homo. An even more radical view rejects the division of Pan and Homo as separate genera, which based on the Principle of Priority would imply the re-classification of chimpanzees as Homo paniscus (or similar).
Human taxonomy on one hand involves the placement of humans within the Taxonomy of the hominids (great apes), and on the other the division of archaic and modern humans into species and, if applicable, subspecies. Modern zoological taxonomy was developed by Carl Linnaeus during the 1730s to 1750s. He named the human species as Homo sapiens in 1758, as the only member species of the genus Homo, divided into several subspecies corresponding to the great races. The Latin noun homō (genitive hominis) means "human being". The systematic name Hominidae for the family of the great apes was introduced by John Edward Gray (1825). Gray also supplied Hominini as the name of the tribe including both chimpanzees (genus Pan) and humans (genus Homo).
The discovery of the first extinct archaic human species from the fossil record dates to the mid 19th century, Homo neanderthalensis, classified in 1864. Since then, a number of other archaic species have been named, but there is no universal consensus as to their exact number. After the discovery of H. neanderthalensis, which even if "archaic" is recognizable as clearly human, late 19th to early 20th century anthropology for a time was occupied with finding the supposedly "missing link" between Homo and Pan. The "Piltdown Man" hoax of 1912 was the (fraudulent) presentation of such a transitional species. Since the mid-20th century, knowledge of the development of Hominini has become much more detailed, and taxonomical terminology has been altered a number of times to reflect this.
The introduction of Australopithecus as a third genus, alongside Homo and Pan, in the Hominini tribe is due to Raymond Dart (1925). Australopithecina as a subtribe containing Australopithecus as well as Paranthropus (Broom 1938) is a proposal by Gregory & Hellman (1939). More recently proposed additions to the Australopithecina subtribe include Ardipithecus (1995) and Kenyanthropus (2001). The position of Sahelanthropus (2002) relative to Australopithecina within Hominini is unclear. Cela-Conde and Ayala (2003) propose the recognition of Australopithecus, Ardipithecus, Praeanthropus, and Sahelanthropus (the latter incertae sedis) as separate genera.
Other proposed genera, now mostly considered part of Homo, include: Pithecanthropus (Dubois, 1894), Protanthropus (Haeckel, 1895), Sinanthropus (Black, 1927), Cyphanthropus (Pycraft, 1928) Africanthropus (Dreyer, 1935), Telanthropus (Broom & Anderson 1949), Atlanthropus (Arambourg, 1954), Tchadanthropus (Coppens, 1965).
The genus Homo has been taken to originate some two millions ago since the discovery of stone tools in Olduvai Gorge, Tanzania, in the 1960s. Homo habilis (Leakey et al., 1964) would be the first "human" species (member of genus Homo) by definition, its type specimen being the OH 7 fossils. However, the discovery of more fossils of this type has opened up the debate on the delineation of H. habilis from Australopithecus. Especially, the LD 350-1 jawbone fossil discovered in 2013, dated to 2.8 Mya, has been argued as being transitional between the two.  It is also disputed whether H. habilis was the first hominin to use stone tools, as Australopithecus garhi, dated to c. 2.5 Mya, has been found along with stone tool implements. Fossil KNM-ER 1470 (discovered in 1972, designated Pithecanthropus rudolfensis by Alekseyev 1978) is now seen as either a third early species of Homo (alongside H. habilis and H. erectus) at about 2 million years ago, or alternatively as transitional between Australopithecus and Homo.
Wood and Richmond (2000) proposed that Gray's tribe Hominini ("hominins") be designated as comprising all species after the chimpanzee-human last common ancestor by definition, to the inclusion of Australopithecines and other possible pre-human or para-human species (such as Ardipithecus and Sahelanthropus) not known in Gray's time. In this suggestion, the new subtribe of Hominina was to be designated as including the genus Homo exclusively, so that Hominini would have two subtribes, Australopithecina and Hominina, with the only known genus in Hominina being Homo. Orrorin (2001) has been proposed as a possible ancestor of Hominina but not Australopithecina.
Designations alternative to Hominina have been proposed: Australopithecinae (Gregory & Hellman 1939) and Preanthropinae (Cela-Conde & Altaba 2002);
At least a dozen species of Homo other than Homo sapiens has been proposed, with varying degrees on consensus. Homo erectus is widely recognized as the species directly ancestral to Homo sapiens. Most other proposed species are proposed as alternatively belonging to either Homo erectus or Homo sapiens as a subspecies. This concerns Homo ergaster in particular: The proposal of a broad division of Homo erectus into an "African" and an "Asian" variety would consider the Asian variety Homo erectus sensu stricto and Homo erectus sensu lato would include both Asian Homo erectus and African Homo ergaster. There appears to be a recent trend, with the availability of ever more difficult-to-classify fossils such as the Dmanisi skulls (2013) or Homo naledi fossils (2015) to subsume all archaic varieties under Homo erectus.
|Species||Temporal range kya||Habitat||Adult height||Adult mass||Cranial capacity (cm³)||Fossil record||Discovery / publication of name|
membership in Homo uncertain
|2,100 – 1,500||East Africa||110-140 cm (4 ft 11 in)||33–55 kg (73–121 lb)||510–660||Many||1960/1964|
membership in Homo uncertain
also classified as H. habilis
|1,900 – 600||South Africa||100 cm (3 ft 3 in)||3 individuals||2010/2010|
|H. erectus||1,900 – 140||Africa, Eurasia||180 cm (5 ft 11 in)||60 kg (130 lb)||850 (early) – 1,100 (late)||Many||1891/1892|
African H. erectus
|1,800 – 1,300||East and Southern Africa||700–850||Many||1949/1975|
also classified as H. heidelbergensis
|1,200 – 800||Western Europe||175 cm (5 ft 9 in)||90 kg (200 lb)||1,000||2 sites||1994/1997|
|H. heidelbergensis||600 – 300||Europe, Africa||180 cm (5 ft 11 in)||90 kg (200 lb)||1,100–1,400||Many||1907/1908|
a single fossil, possibly H. erectus
|c. 450||Italy||1,000||1 skull cap||1994/2003|
also classified as H. heidelbergensis or a subspecies of H. sapiens
|c. 300||Zambia||1,300||single or very few||1921/1921|
||c. 300||South Africa||150 cm (4 ft 11 in)||45 kg (99 lb)||450||15 individuals||2013/2015|
(anatomically modern humans)
|300 – present||Worldwide||150 - 190 cm (4 ft 7 in - 6 ft 3 in)||50–100 kg (110–220 lb)||950–1,800||(extant)||—/1758|
possibly a subspecies of H. sapiens
|240 – 40||Europe, Western Asia||170 cm (5 ft 7 in)||55–70 kg (121–154 lb) (heavily built)||1,200–1,900||Many||1829/1864|
|190 – 50||Indonesia||100 cm (3 ft 3 in)||25 kg (55 lb)||400||7 individuals||2003/2004|
possibly H. erectus
|c. 100||Taiwan||1 individual||2008(?)/2015|
possible H. sapiens subspecies or hybrid
|Red Deer Cave people
possible H. sapiens subspecies or hybrid
|15–12||Southwest China||Very few||2012/—|
H. sapiens ssp.
The recognition or non-recognition of subspecies of Homo sapiens has a complicated history. The rank of subspecies in zoology is introduced for convenience, and not by objective criteria, based on pragmatic consideration of factors such as geographic isolation and sexual selection. The informal taxonomic rank of race is variously considered equivalent or subordinate to the rank of subspecies, and the division of anatomically modern humans (H. sapiens) into subspecies is closely tied to the recognition of major racial groupings based on human genetic variation.
A subspecies cannot be recognized independently: a species will either be recognized as having no subspecies at all or at least two (including any that are extinct). Therefore, the designation of an extant subspecies Homo sapiens sapiens (for which Carl Linnaeus would be the type specimen) only makes sense if at least one other subspecies is recognized.
During the 19th to mid-20th century, it was common practice to classify the major divisions of extant H. sapiens as subspecies, following Linnaeus (1758), who had recognized H. s. americanus, H. s. europaeus, H. s. asiaticus and H. s. afer as grouping the native populations of the Americas, West Eurasia, East Asia and Sub-Saharan Africa, respectively, besides H. s. ferus (for the "wild" form which he identified with feral children) and two further "wild" forms for reported specimens now considered part of cryptozoology, H. s. monstrosus and H. s. troglodytes. There were variations and additions to the categories of Linnaeus, such as H. s. tasmanianus for the native population of Australia. Bory de St. Vincent in his Essai sur l'Homme (1825) extended Linné's "racial" categories to as many as fifteen: Leiotrichi ("smooth-haired"): japeticus (with subraces), arabicus, indicus, scythicus, sinicus, hyperboreus, neptunianus, australasicus, columbicus, americanus, patagonicus; Oulotrichi ("crisp-haired"): aethiopicus, cafer, hottentotus, melaninus. Similarly, Georges Vacher de Lapouge (1899) also had categories based on race, such as priscus, spelaeus (etc.);
Homo sapiens neanderthalensis was proposed by King (1864) as an alternative to Homo neanderthalensis. There have been "taxonomic wars" over whether Neanderthals were a separate species since their discovery in the 1860s. Pääbo (2014) frames this as a debate that is unresolvable in principle, "since there is no definition of species perfectly describing the case." Louis Lartet (1869) proposed Homo sapiens fossilis based on the Cro-Magnon fossils.
There are a number of proposals of extinct varieties of Homo sapiens made in the 20th century. Many of the original proposals were not using explicit trinomial nomenclature, even though they are still cited as valid synonyms of H. sapiens by Wilson & Reeder (2005). These include: Homo grimaldii (Lapouge, 1906), Homo aurignacensis hauseri (Klaatsch & Hauser, 1910), Notanthropus eurafricanus (Sergi, 1911), Homo fossilis infrasp. proto-aethiopicus (Giuffrida-Ruggeri, 1915), Telanthropus capensis (Broom, 1917), Homo wadjakensis (Dubois, 1921), Homo sapiens cro-magnonensis, Homo sapiens grimaldiensis (Gregory, 1921), Homo drennani (Kleinschmidt, 1931), Homo galilensis (Joleaud, 1931) = Paleanthropus palestinus (McCown & Keith, 1932). Rightmire (1983) proposed Homo sapiens rhodesiensis.
By the 1980s, the practice of dividing extant populations of homo sapiens into subspecies declined. An early authority explicitly avoiding the division of H. sapiens into subspecies was Grzimeks Tierleben, published 1967–1972. A late example of an academic authority proposing that the human racial groups should be considered taxonomical subspecies is John Baker (1974). The trinomial nomenclature Homo sapiens sapiens became popular for "modern humans" in the context of Neanderthals being considered a subspecies of H. sapiens in the second half of the 20th century, without having any formal authority introducing it. Derived from the convention, widespread in the 1980s, of considering two subspecies, H. s. neanderthalensis and H. s. sapiens, the explicit claim that "H. s. sapiens is the only extant human subspecies" appears in the early 1990s.
Since the 2000s, the extinct Homo sapiens idaltu (White et al., 2003) has gained wide recognition as a subspecies of Homo sapiens, but even in this case there is a dissenting view arguing that "the skulls may not be distinctive enough to warrant a new subspecies name". H. s. neanderthalensis and H. s. rhodesiensis continue to be considered separate species by some authorities, but the genetic evidence of archaic human admixture with modern humans discovered in the 2010s has re-opened the details of taxonomy of archaic humans.
Homo erectus ssp.
Homo erectus since its introduction in 1892 has been divided into numerous subspecies, many of them formerly considered individual species of Homo. None of these subspecies have universal consensus among paleontologists.
- Homo erectus erectus (Java Man) (1970s)
- Homo erectus yuanmouensis (Yuanmou Man) (Li et al., 1977)
- Homo erectus lantianensis (Lantian Man) (Woo Ju-Kang, 1964)
- Homo erectus nankinensis (Nanjing Man) (1993)
- Homo erectus pekinensis (Peking Man) (1970s)
- Homo erectus palaeojavanicus (Meganthropus) (Tyler, 2001)
- Homo erectus soloensis (Solo Man) (Oppenoorth, 1932)
- Homo erectus tautavelensis (Tautavel Man) (de Lumley and de Lumley, 1971)
- Homo erectus georgicus (1991)
- Homo erectus bilzingslebenensis (Vlček, 2002)
- Jared Diamond in The Third Chimpanzee (1991), and Morris Goodman (2003) Hecht, Jeff (19 May 2003). "Chimps are human, gene study implies". New Scientist. Retrieved 2011-12-08.
- 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.
- 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 . PMID 12794185.
- Introduced for the Florisbad Skull (discovered in 1932, 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.
- Villmoare B, Kimbel H, Seyoum C, Campisano C, DiMaggio E, Rowan J, Braun D, Arrowsmith J, Reed K. (2015). Early Homo at 2.8 Ma from Ledi-Geraru, Afar, Ethiopia. Science. doi:10.1126/science.aaa1343. Some paleoanthropologists regard the H. habilis taxon as invalid, made up of fossil specimens of Australopithecus and Homo. Tattersall, I. & Schwartz, J.H., Extinct Humans, Westview Press, New York, 2001, p. 111.
- De Heinzelin, J; Clark, JD; White, T; Hart, W; Renne, P; Woldegabriel, G; Beyene, Y; Vrba, E (1999). "Environment and behavior of 2.5-million-year-old Bouri hominids". Science. 284 (5414): 625–9. doi:10.1126/science.284.5414.625. PMID 10213682.
- Kaplan, Matt (8 August 2012). "Fossils point to a big family for human ancestors". Nature. Retrieved 8 August 2012.
- 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 . PMID 10999270.
- Reynolds, Sally C; Gallagher, Andrew (2012-03-29). African Genesis: Perspectives on Hominin Evolution. ISBN 9781107019959.
- Brunet, M.; et al. (2002). "A new hominid from the upper Miocene of Chad, central Africa". Nature. 418: 145–151. doi:10.1038/nature00879. PMID 12110880. Cela-Conde, C.J.; Ayala, F.J. (2003). "Genera of the human lineage". PNAS. 100 (13): 7684–7689. doi:10.1073/pnas.0832372100. PMC . PMID 12794185. Wood, B.; Lonergan, N. (2008). "The hominin fossil record: taxa, grades and clades" (PDF). J. Anat. 212: 354–376. doi:10.1111/j.1469-7580.2008.00871.x. PMC . PMID 18380861.
- Hazarika, Manji (16–30 June 2007). "Homo erectus/ergaster and Out of Africa: Recent Developments in Paleoanthropology and Prehistoric Archaeology" (PDF).; Klein, R. (1999). The Human Career: Human Biological and Cultural Origins. Chicago: University of Chicago Press, ISBN 0226439631.
- Antón, S. C. (2003). "Natural history of Homo erectus". Am. J. Phys. Anthropol. 122: 126–170. doi:10.1002/ajpa.10399.
By the 1980s, the growing numbers of H. erectus specimens, particularly in Africa, led to the realization that Asian H. erectus (H. erectus sensu stricto), once thought so primitive, was in fact more derived than its African counterparts. These morphological differences were interpreted by some as evidence that more than one species might be included in H. erectus sensu lato (e.g., Stringer, 1984; Andrews, 1984; Tattersall, 1986; Wood, 1984, 1991a, b; Schwartz and Tattersall, 2000) ... Unlike the European lineage, in my opinion, the taxonomic issues surrounding Asian vs. African H. erectus are more intractable. The issue was most pointedly addressed with the naming of H. ergaster on the basis of the type mandible KNM-ER 992, but also including the partial skeleton and isolated teeth of KNM-ER 803 among other Koobi Fora remains (Groves and Mazak, 1975). Recently, this specific name was applied to most early African and Georgian H. erectus in recognition of the less-derived nature of these remains vis à vis conditions in Asian H. erectus (see Wood, 1991a, p. 268; Gabunia et al., 2000a). It should be noted, however, that at least portions of the paratype of H. ergaster (e.g., KNM-ER 1805) are not included in most current conceptions of that taxon. The H. ergaster question remains famously unresolved (e.g., Stringer, 1984; Tattersall, 1986; Wood, 1991a, 1994; Rightmire, 1998b; Gabunia et al., 2000a; Schwartz and Tattersall, 2000), in no small part because the original diagnosis provided no comparison with the Asian fossil record
- Skull suggests three early human species were one : Nature News & Comment David Lordkipanidze, Marcia S. Ponce de Leòn, Ann Margvelashvili, Yoel Rak, G. Philip Rightmire, Abesalom Vekua, Christoph P. E. Zollikofer (18 October 2013). "A Complete Skull from Dmanisi, Georgia, and the Evolutionary Biology of Early Homo". Science. 342 (6156): 326–331. doi:10.1126/science.1238484. Switek, Brian (17 October 2013). "Beautiful Skull Spurs Debate on Human History". National Geographic. Retrieved 22 September 2014.
- 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. 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. doi:10.1007/978-3-540-33761-4_52. ISBN 978-3-540-32474-4. 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. doi:10.1126/science.aaa1415. ISSN 0036-8075. PMID 25739409. 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.
- 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. doi:10.1016/j.jchb.2010.04.002. ISSN 0018-442X. PMID 20466364. A species proposed in 2010 based on the fossil remains of three individuals dated between 1.9 and 0.6 million years ago. The same fossils were also classified as H. habilis, H. ergaster or Australopithecus by other anthropologists.
- 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.
- 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. PMC . PMID 21646521. 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.
- 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.
- 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"
- The type fossil is Mauer 1, dated to ca. 0.6 million years ago. The transition from H. heidelbergensis to H. neanderthalensis between 300 and 243 thousand years ago is conventional, and makes use of the fact that there is no known fossil in this period. Examples of H. heidelbergensis are fossils found at Bilzingsleben (also classified as Homo erectus bilzingslebensis).
- Muttoni, Giovanni; Scardia, Giancarlo; Kent, Dennis V.; Swisher, Carl C.; Manzi, Giorgio (2009). "Pleistocene magnetochronology of early hominin sites at Ceprano and Fontana Ranuccio, Italy". Earth and Planetary Science Letters. ScienceDirect. 286: 255–268. Bibcode:2009E&PSL.286..255M. doi:10.1016/j.epsl.2009.06.032. Retrieved December 10, 2015.
- Dirks, P.; et al. (9 May 2017). "The age of Homo naledi and associated sediments in the Rising Star Cave, South Africa". eLife. 6: e24231. doi: .
- The age of H. sapiens has long been assumed to be close to 200,000 years, but since 2017 there have been a number of suggestions extending this time to has high as 300,000 years. In 2017, fossils found in Jebel Irhoud (Morocco) suggest that Homo sapiens may have speciated by as early as 315,000 years ago. 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. Genetic evidence has been adduced for an age of roughly 270,000 years. Posth, Cosimo; et al. (4 July 2017). "Deeply divergent archaic mitochondrial genome provides lower time boundary for African gene flow into Neanderthals". Nature Communications. doi:10.1038/ncomms16046. Retrieved 4 July 2017.
- Bischoff, James L.; Shamp, Donald D.; Aramburu, Arantza; et al. (March 2003). "The Sima de los Huesos Hominids Date to Beyond U/Th Equilibrium (>350 kyr) and Perhaps to 400–500 kyr: New Radiometric Dates". Journal of Archaeological Science. Amsterdam, the Netherlands: Elsevier. 30 (3): 275–280. doi:10.1006/jasc.2002.0834. ISSN 0305-4403. The first humans with "proto-Neanderthal traits" lived in Eurasia 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). There is a fossil gap in Europe between 300 and 243 kya, and by convention, fossils younger than 243 kya are called "Neanderthal". D. Dean; J.-J. Hublin; R. Holloway; R. Ziegler (1998). "On the phylogenetic position of the pre-Neandertal specimen from Reilingen, Germany". Journal of Human Evolution. 34 (5). pp. 485–508. doi:10.1006/jhev.1998.0214.
- younger than 450 kya, either between 190-130 or between 70-10 kya. 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. doi:10.1038/ncomms7037. PMC . PMID 25625212.
- provisional names Homo sp. Altai or Homo sapiens ssp. Denisova.
- Bølling-Allerød warming period; Curnoe, D.; et al. (2012). Caramelli, David, ed. "Human remains from the Pleistocene-Holocene transition of southwest China Suggest a complex evolutionary history for East Asians". PLoS ONE. 7 (3): e31918. doi:10.1371/journal.pone.0031918. PMC . PMID 22431968.
- Linné, Carl von (1758). Systema naturæ. Regnum animale (10 ed.). pp. 18ff.
- See e.g. John Wendell Bailey, The Mammals of Virginia (1946), p. 356.; Journal of Mammalogy 26-27 (1945), p. 359.; The Mankind Quarterly 1-2 (1960), 113ff ("Zoological Subspecies of Man"), J. Desmond Clark (ed.), The Cambridge History of Africa, Cambridge University Press (1982), p. 141 (with references).
- Annals of Philosophy 11, London (1826), p. 71
- Frederick S. Szalay, Eric Delson, Evolutionary History of the Primates (2013), 508
- Pääbo, Svante (2014). Neanderthal Man: In Search of Lost Genomes. New York: Basic Books. p. 237.
- Groves, C.P. (2005). Wilson, D.E.; Reeder, D.M., eds. Mammal Species of the World: A Taxonomic and Geographic Reference (3rd ed.). Baltimore: Johns Hopkins University Press. ISBN 0-801-88221-4. OCLC 62265494.
- T. Harrison in: William H. Kimbel, Lawrence B. Martin (eds.), Species, Species Concepts and Primate Evolution (2013), 361.
- M. R. Drennan, "An Australoid Skull from the Cape Flats", The Journal of the Royal Anthropological Institute of Great Britain and Ireland Vol. 59 (Jul. - Dec., 1929), 417-427.
- among other names suggested for fossils later subsumed under neanderthalensis, see: Eric Delson, Ian Tattersall, John Van Couvering, Alison S. Brooks, Encyclopedia of Human Evolution and Prehistory: Second Edition, Routledge (2004).
- Rightmire GP (June 3, 1983). "The Lake Ndutu cranium and early Homo sapiens in Africa". Am. J. Phys. Anthropol. 61 (2): 245–54. doi:10.1002/ajpa.1330610214. PMID 6410925.
- English translation (1972–1975): Grzimek's Animal Life Encyclopedia, Volume 11, p. 55.
- John R. Baker, Race, Oxford University Press (1974).
- Homo sapiens sapiens is rarely used before the 1940s. In 1946, John Wendell Bailey attributes the name to Linnaeus (1758) explicitly: "Linnaeus. Syst. Nat. ed. 10, Vol. 1. pp. 20, 21, 22, lists five races of man, viz: Homo sapiens sapiens (white — Caucasian) [...]", This is a misattribution, but H. s. sapiens has since often been attributed to Linnaeus. In actual fact, Linnaeus, Syst. Nat. ed. 10 Vol. 1. p. 21 does not have Homo sapiens sapiens, the "white" or "Caucasian" race being instead called Homo sapiens Europaeus. This is explicitly pointed out in Bulletin der Schweizerische Gesellschaft für Anthropologie und Ethnologie Volume 21 (1944), p. 18 (arguing not against H. s. sapiens but against "H. s. albus L." proposed by von Eickstedt and Peters): "die europide Rassengruppe, als Subspecies aufgefasst, [würde] Homo sapiens eurpoaeus L. heissen" ("the Europid racial group, considered as a subspecies, would be named H. s. europeaeus L."). See also: John R. Baker, Race, Oxford University Press (1974), 205.
- "We are the only surviving subspecies of Homo sapiens." Michio Kitahara, The tragedy of evolution: the human animal confronts modern society (1991), p. xi.
- Chris Stringer (June 12, 2003). "Human evolution: Out of Ethiopia". Nature.
- Hublin, J. J. (2009). "The origin of Neandertals". Proceedings of the National Academy of Sciences. 106 (38): 16022–7. Bibcode:2009PNAS..10616022H. doi:10.1073/pnas.0904119106. JSTOR 40485013. PMC . PMID 19805257. Harvati, K.; Frost, S.R.; McNulty, K.P. (2004). "Neanderthal taxonomy reconsidered: implications of 3D primate models of intra- and interspecific differences". Proc. Natl. Acad. Sci. U.S.A. 101 (5): 1147–52. Bibcode:2004PNAS..101.1147H. doi:10.1073/pnas.0308085100. PMC . PMID 14745010. "Homo neanderthalensis King, 1864". Wiley-Blackwell Encyclopedia of Human Evolution. Chichester, West Sussex: Wiley-Blackwell. 2013. pp. 328–331.
- In the 1970s a tendency developed to regard the Javanese variety of H. erectus as a subspecies, Homo erectus erectus, with the Chinese variety being referred to as Homo erectus pekinensis. See: Sartono, S. Implications arising from Pithecanthropus VIII In: Paleoanthropology: Morphology and Paleoecology. Russell H. Tuttle (Ed.), p. 328.
- Emanuel Vlček: Der fossile Mensch von Bilzingsleben (= Bilzingsleben. Bd. 6 = Beiträge zur Ur- und Frühgeschichte Mitteleuropas 35). Beier & Beran, Langenweißbach 2002.