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Homo sapiens
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{{short description|Humans as a biological species}}{{about|Homo sapiens as a biological species (its evolution and habitat)|a more general perspective on humanity|Human|other uses|Homo sapiens (disambiguation)}}{{pp-vandalism|small=yes}}{{speciesbox
0.35Middle Pleistocene–Holocene>Present| image = Akha cropped hires.JPG| image_upright = 0.9H s. sapiens(Akha people>Akha in northern Thailand, 2007 photograph)| image_alt = Akha man and woman in northern Thailand – husband carries stem of banana-plant, which will be fed to their pigs| status = LC| status_system = IUCN3.1JOURNAL=THE IUCN RED LIST OF THREATENED SPECIES PAGE=E.T136584A4313662 DOI=10.2305/IUCN.UK.2008.RLTS.T136584A4313662.EN, | taxon = Homo sapiensCarl Linnaeus>Linnaeus, 1758| subdivision_ranks = SubspeciesHomo sapiens sapiens>H. s. sapiens†''Homo sapiens idaltu†Homo sapiens neanderthalensis>H. s. neanderthalensis(?) †Homo sapiens rhodesiensis''(?)(human subspecies>others proposed) }}Homo sapiens is the only extant human species. The name is Latin for "(:wikt:sapiens|wise) (:wikt:homo#Latin|man)" and was introduced in 1758 by Carl Linnaeus (who is himself the lectotype for the species).Extinct species of the genus Homo include Homo erectus, extant during roughly 1.9 to 0.4 million years ago, and a number of other species (by some authors considered subspecies of either H. sapiens or H. erectus). The age of speciation of H. sapiens out of ancestral H. erectus (or an intermediate species such as Homo antecessor) is estimated to have been roughly 350,000 years ago.{{refn|group=note|name=350kiloyearsAgo|Based on Schlebusch et al., "Southern African ancient genomes estimate modern human divergence to 350,000 to 260,000 years ago",JOURNAL, Schlebusch, etal, Southern African ancient genomes estimate modern human divergence to 350,000 to 260,000 years ago, Science, 358, 6363, 3 November 2017, 652–655, 10.1126/science.aao6266, 28971970, 2017Sci...358..652S, Fig. 3 (H. sapiens divergence times) and Stringer (2012),JOURNAL, Stringer, C, What makes a modern human, Nature, 2012, 485, 7396, 33–35, 10.1038/485033a, 22552077, 2012Natur.485...33S, (archaic admixture).}} Sustained archaic admixture is known to have taken place both in Africa and (following the recent Out-Of-Africa expansion) in Eurasia, between about 100,000 and 30,000 years ago.DOCUMENT,weblink Harrod (2014) Suppl File Table 1 mtDNA language myth Database rev May 17 2019.doc, Harrod, James, The term anatomically modern humansBOOK,weblink Nitecki, Matthew H, Nitecki, Doris V, 1994, Origins of Anatomically Modern Humans, Springer, 1489915079, (AMH) is used to distinguish H. sapiens having an anatomy consistent with the range of phenotypes seen in contemporary humans from varieties of extinct archaic humans. This is useful especially for times and regions where anatomically modern and archaic humans co-existed, for example, in Paleolithic Europe.

Name and taxonomy

{{further|Homo|Names for the human species}}The binomial name Homo sapiens was coined by Linnaeus, 1758.BOOK, Linné, Carl von, Systema naturæ. Regnum animale, 1758, 18, 20,weblink 10th, 2019-05-06, Sumptibus Guilielmi Engelmann, The Latin noun (wikt:homo#Latin|homō) (genitive hominis) means "human being", while the participle (wikt:sapiens#Latin|sapiēns) means "discerning, wise, sensible".The species was initially thought to have emerged from a predecessor within the genus Homo around 300,000 to 200,000 years ago.{{refn|group=note|This is a matter of convention (rather than a factual dispute), and there is no universal consensus on terminology. Some scholars include humans of up to 600,000 years ago under the same species. See Bryant (2003), p. 811.BOOK,weblink Handbook of Death and Dying, Bryant, Clifton D, 2003, SAGE, 0761925147, See also Tattersall (2012), Page 82 (cf. Unfortunately this consensus in principle hardly clarifies matters much in practice. For there is no agreement on what the 'qualities of a man' actually are," [...]).BOOK,weblink Masters of the Planet: The Search for Our Human Origins, Tattersall, Ian, 2012, St Martin's Press, 978-1137000385, }} A problem with the morphological classification of "anatomically modern" was that it would not have included certain extant populations. For this reason, a lineage-based (cladistic) definition of H. sapiens has been suggested, in which H. sapiens would by definition refer to the modern human lineage following the split from the Neanderthal lineage. Such a cladistic definition would extend the age of H. sapiens to over 500,000 years.{{refn|group=note|WerdelinBOOK, Lars, Werdelin, William Joseph, Sanders, Cenozoic Mammals of Africa, 2010,weblink 517, Univ of California Press, 9780520257214, citing Lieberman et al.JOURNAL, DE, Lieberman, BM, McBratney, G, Krovitz, The evolution and development of cranial form in Homo sapiens, PNAS, 2002, 99, 3, 1134–39, 10.1073/pnas.022440799, 11805284, 122156, 2002PNAS...99.1134L, }}Extant human populations have historically been divided into subspecies, but since around the 1980s all extant groups have tended to be subsumed into a single species, H. sapiens, avoiding division into subspecies altogether.{{refn|group=note|The history of claimed or proposed subspecies of H. sapiens is complicated and fraught with controversy. The only widely recognized archaic subspecies{{citation needed|date=February 2019}} is H. sapiens idaltu (2003). The name H. s. sapiens is due to Linnaeus (1758), and refers by definition the subspecies of which Linnaeus himself is the type specimen. However, Linnaeus postulated four other extant subspecies, viz. H. s. afer, H. s. americanus, H. s. asiaticus and H. s. ferus for Africans, Americans, Asians and Malay. This classification remained in common usage until the mid 20th century, sometimes alongside H. s. tasmanianus for Australians. See, for example, Bailey, 1946;BOOK,weblink John Wendell, Bailey, The Mammals of Virginia, 1946, 356, Hall, 1946.JOURNAL, Hall, E, 1946, Zoological Subspecies of Man at the Peace Table, Journal of Mammalogy, 27, 4, 358–364, 10.2307/1375342, 1375342, The division of extant human populations into taxonomic subspecies was gradually given up in the 1970s (for example, Grzimek's Animal Life EncyclopediaENCYCLOPEDIA,weblink Grzimek's Animal Life Encyclopedia, 11, 55, 1970, ).}}
Some sources show Neanderthals (H. neanderthalensis) as a subspecies (H. sapiens neanderthalensis).JOURNAL, 10.1073/pnas.0904119106, The origin of Neandertals, 2009, Hublin, J. J., Proceedings of the National Academy of Sciences, 106, 38, 16022–27, 19805257, 40485013, 2009PNAS..10616022H, 2752594, JOURNAL, Neanderthal taxonomy reconsidered: implications of 3D primate models of intra- and interspecific differences, 14745010, Harvati, K., Frost, S.R., McNulty, K.P., 2004, 10.1073/pnas.0308085100, 337021, 101, 5, Proc. Natl. Acad. Sci. U.S.A., 1147–52, 2004PNAS..101.1147H, Similarly, the discovered specimens of the H. rhodesiensis species have been classified by some as a subspecies (H. sapiens rhodesiensis), although it remains more common to treat these last two as separate species within the genus Homo rather than as subspecies within H. sapiens.ENCYCLOPEDIA, Homo neanderthalensis King, 1864, Wiley-Blackwell Encyclopedia of Human Evolution, 2013, Wiley-Blackwell, Chichester, West Sussex, 328–31, The subspecies name H. sapiens sapiens is sometimes used informally instead of "modern humans" or "anatomically modern humans". It has no formal authority associated with it.{{refn|group=note|Homo sapiens sapiens is first used in the 1940s as a synonym of Linnaeus' H. s. europaeus, i. e. Caucasoids. This usage is abandoned by the 1970s, and H. s. sapiens was now used for Cro-Magnon by authors who wished to classify Neanderthals as subspecies of H. sapiens taken in a wider sense, for example Seely.JOURNAL,weblink The Journal of the American Scientific Affiliation, Not a Viable Theory, Seely, Paul H, 1971, 23, 4, 134, }} By the early 2000s, it had become common to use H. s. sapiens for the ancestral population of all contemporary humans, and as such it is equivalent to the binomial H. sapiens in the more restrictive sense (considering H. neanderthalensis a separate species).{{refn|group=note|For example, "DMA studies have revealed that the first anatomically modern humans (H. s. sapiens) arose in Africa between 200,000 and 140,000 years ago".BOOK,weblink Geoffrey, Parker, Compact history of the world, 2001, 14, 9780760725757, This usage persists alongside H. s. sapiens designating Upper Paleolithic Cro Magnon, for example. "About 200,000 years ago our own species, Homo sapiens (the thinking human), evolved [...] About 60,000 years ago we became elaborate artisans, building boats and intricate shelters; at this stage, scientists refer to us as Homo sapiens sapiens."BOOK,weblink Nina, Rosenstand, The Human Condition: An Introduction to Philosophy of Human Nature, 2002, 42, 1559347643, McGraw-Hill, }}

Age and speciation process

{{further|Human evolution|Homo|Timeline of human evolution|Early human migrations}}File:Homo lineage 2017update.svg|thumb|Schematic representation of the emergence of H. sapiens from earlier species of Homo. The horizontal axis represents geographic location; the vertical axis represents time in millions of years ago (blue areas denote the presence of a certain species of Homo at a given time and place; late survival of robust australopithecinesrobust australopithecines

Derivation from H. erectus

{{see|Homo antecessor|Homo heidelbergensis|Homo rhodesiensis|Acheulean}}File:Homo sapiens lineage.svg|thumb|upright=1.35|A model of the phylogeny of H. sapiens during the Middle Paleolithic. The horizontal axis represents geographic location; the vertical axis represents time in thousands of years ago.{{refn|group=note|name=350kiloyearsAgo}} Neanderthals, Denisovans and unspecified archaic African hominins are shown as admixed into the H. sapiens lineage. In addition, prehistoric Archaic Human and Eurasian admixture events in modern African populations are indicated.]]The speciation of H. sapiens out of archaic human varieties derived from H. erectus is estimated as having taken place over 350,000 years ago, as the Khoisan split from other populations is dated between 260,000 and 350,000 years ago.JOURNAL,weblink Ancient genomes from southern Africa pushes modern human divergence beyond 260,000 years ago, Carina M., Schlebusch, Helena, Malmström, Torsten, Günther, Per, Sjödin, Alexandra, Coutinho, Hanna, Edlund, Arielle R., Munters, Maryna, Steyn, Himla, Soodyall, Marlize, Lombard, Mattias, Jakobsson, 5 June 2017, bioRxiv, 145409, 10.1101/145409, An alternative suggestion defines H. sapiens cladistically as including the lineage of modern humans since the split from the lineage of Neanderthals, roughly 500,000 to 800,000 years ago.The time of divergence between archaic H. sapiens and ancestors of Neanderthals and Denisovans caused by a genetic bottleneck of the latter was dated at 744,000 years ago, combined with repeated early admixture events and Denisovans diverging from Neanderthals 300 generations after their split from H. sapiens, as calculated by Rogers et al. (2017).JOURNAL, Rogers, Alan R., Bohlender, Ryan J., Huff, Chad D., Early history of Neanderthals and Denisovans, Proceedings of the National Academy of Sciences, 12 September 2017, 114, 37, 9859–9863, 10.1073/pnas.1706426114, 28784789, 5604018, The derivation of a comparatively homogeneous single species of H. sapiens from more diverse varieties of archaic humans (all of which were descended from the early dispersal of H. erectus some 1.8 million years ago) was debated in terms of two competing models during the 1980s: "recent African origin" postulated the emergence of H. sapiens from a single source population in Africa, which expanded and led to the extinction of all other human varieties, while the "multiregional evolution" model postulated the survival of regional forms of archaic humans, gradually converging into the modern human varieties by the mechanism of clinal variation, via genetic drift, gene flow and selection throughout the Pleistocene.JOURNAL, Wolpoff, M. H., Spuhler, J. N., Smith, F. H., Radovcic, J., Pope, G., Frayer, D. W., Eckhardt, R., Clark, G., 1988, Modern Human Origins, Science, 241, 4867, 772–74, 10.1126/science.3136545, 3136545, 1988Sci...241..772W, Since the 2000s, the availability of data from archaeogenetics and population genetics has led to the emergence of a much more detailed picture, intermediate between the two competing scenarios outlined above: The recent Out-of-Africa expansion accounts for the predominant part of modern human ancestry, while there were also significant admixture events with regional archaic humans.JOURNAL, Green, RE, Krause, J, Briggs, Adrian W., Maricic, Tomislav, Stenzel, Udo, Kircher, Martin, Patterson, Nick, Li, Heng, Zhai, Weiwei, Fritz, Markus Hsi-Yang, Hansen, Nancy F., Durand, Eric Y., Malaspinas, Anna-Sapfo, Jensen, Jeffrey D., Marques-Bonet, Tomas, Alkan, Can, Prüfer, Kay, Meyer, Matthias, Burbano, Hernán A., Good, Jeffrey M., Schultz, Rigo, Aximu-Petri, Ayinuer, Butthof, Anne, Höber, Barbara, Höffner, Barbara, Siegemund, Madlen, Weihmann, Antje, Nusbaum, Chad, Lander, Eric S., Russ, Carsten, vanc, May 2010, A draft sequence of the Neandertal genome, Science, 328, 5979, 710–22, 10.1126/science.1188021, 20448178, 2010Sci...328..710G, 29, 5100745, JOURNAL, Reich, D, Patterson, Nick, Kircher, Martin, Delfin, Frederick, Nandineni, Madhusudan R., Pugach, Irina, Ko, Albert Min-Shan, Ko, Ying-Chin, Jinam, Timothy A., Phipps, Maude E., Saitou, Naruya, Wollstein, Andreas, Kayser, Manfred, Pääbo, Svante, Stoneking, Mark, vanc, 2011, Denisova admixture and the first modern human dispersals into southeast Asia and oceania, Am J Hum Genet, 89, 4, 516–28, 10.1016/j.ajhg.2011.09.005, 21944045, 3188841, Since the 1970s, the Omo remains, dated to some 195,000 years ago, have often been taken as the conventional cut-off point for the emergence of "anatomically modern humans". Since the 2000s, the discovery of older remains with comparable characteristics, and the discovery of ongoing hybridization between "modern" and "archaic" populations after the time of the Omo remains, have opened up a renewed debate on the age of H. sapiens in journalistic publications.WEB,weblink New Clues Add 40,000 Years to Age of Human Species, NSF – National Science Foundation, www.nsf.gov, NEWS,weblink BBC News, Age of ancient humans reassessed, February 16, 2005, April 10, 2010, WEB,weblink The Oldest Homo Sapiens: Fossils Push Human Emergence Back To 195,000 Years Ago, February 28, 2005, 2019-05-06, ScienceDaily, JOURNAL, Alemseged, Z., Coppens, Y., Geraads, D., Hominid cranium from Homo: Description and taxonomy of Homo-323-1976-896, Am J Phys Anthropol, 117, 2, 103–12, 2002, 11815945, 10.1002/ajpa.10032, JOURNAL, Stoneking, Mark, Soodyall, Himla, Human evolution and the mitochondrial genome, Current Opinion in Genetics & Development, 6, 6, 731–36, 1996, 10.1016/S0959-437X(96)80028-1, H. s. idaltu, dated to 160,000 years ago, has been postulated as an extinct subspecies of H. sapiens in 2003.Human evolution: the fossil evidence in 3D, by Philip L. Walker and Edward H. Hagen, Dept. of Anthropology, University of California, Santa Barbara. Retrieved April 5, 2005.{{better source|date=May 2019|reason=Shockwave sites are not legitimate sources anymore}} H. neanderthalensis, which became extinct about 40,000 years ago, has also been classified as a subspecies, H. s. neanderthalensis.H. heidelbergensis, dated 600,000 to 300,000 years ago, has long been thought to be a likely candidate for the last common ancestor of the Neanderthal and modern human lineages. However, genetic evidence from the Sima de los Huesos fossils published in 2016 seems to suggest that H. heidelbergensis in its entirety should be included in the Neanderthal lineage, as "pre-Neanderthal" or "early Neanderthal", while the divergence time between the Neanderthal and modern lineages has been pushed back to before the emergence of H. heidelbergensis, to close to 800,000 years ago, the approximate time of disappearance of H. antecessor.JOURNAL, Meyer, Matthias, Arsuaga, Juan-Luis, de Filippo, Cesare, Nagel, Sarah, Aximu-Petri, Ayinuer, Nickel, Birgit, Martínez, Ignacio, Gracia, Ana, de Castro, José María Bermúdez, Carbonell, Eudald, Viola, Bence, Kelso, Janet, Prüfer, Kay, Pääbo, Svante, Nuclear DNA sequences from the Middle Pleistocene Sima de los Huesos hominins, Nature, 14 March 2016, 531, 7595, 504–507, 10.1038/nature17405, 26976447, 2016Natur.531..504M, JOURNAL, Callaway, Ewen, Oldest ancient-human DNA details dawn of Neanderthals, Nature, 14 March 2016, 531, 7594, 296–286, 10.1038/531286a, 26983523, 2016Natur.531..296C,

Early Homo sapiens

{{see|Human subspecies|Middle Paleolithic|Mousterian|Archaic human admixture with modern humans|Homo sapiens idaltu|Skhul and Qafzeh hominins}}File:Skhul.JPG|thumb|right|upright=0.9|Skhul VSkhul VThe term Middle Paleolithic is intended to cover the time between the first emergence of H. sapiens (roughly 300,000 years ago) and the emergence of full behavioral modernity (roughly 50,000 years ago, corresponding to the start of the Upper Paleolithic).Many of the early modern human finds, like those of Omo, Herto, Skhul, Jebel Irhoud and PeÈ™tera cu Oase exhibit a mix of archaic and modern traits.BOOK, Oppenheimer, S., Out of Eden: The Peopling of the World, 2003, 978-1-84119-697-8, JOURNAL, 10.1073/pnas.2035108100, Trinkaus, E., Moldovan, O., Milota, Ș., Bîlgăr, A., Sarcina, L., Athreya, S., Bailey, S. E., Rodrigo, R., Gherase, M., Higham, T., Ramsey, C. B., Van Der Plicht, J., 2003, An early modern human from PeÈ™tera cu Oase, Romania, PNAS, 100, 20, 11231–36, 14504393, 208740, 2003PNAS..10011231T, 8, JOURNAL, Callaway, Ewan, Oldest Homo sapiens fossil claim rewrites our species' history,weblink 7 June 2017, Nature (journal), Nature, 10.1038/nature.2017.22114, 11 June 2017, Skhul V, for example, has prominent brow ridges and a projecting face. However, the brain case is quite rounded and distinct from that of the Neanderthals and is similar to the brain case of modern humans. It is uncertain whether the robust traits of some of the early modern humans like Skhul V reflects mixed ancestry or retention of older traits.JOURNAL, David, Reich, Richard E., Green, Martin, Kircher, Johannes, Krause, Nick, Patterson, Eric Y., Durand, Bence, Viola, Adrian W., Briggs, Udo, Stenzel, Philip L. F., Johnson, Tomislav, Maricic, Jeffrey M., Good, Tomas, marques-Bonet, Can, Alkan, Qiaomei, Fu, Swapan, Mallick, Heng, Li, Matthias, Meyer, Evan E., Eichler, Mark, Stoneking, Michael, Richards, Sahra, Talamo, Michael V., Shunkov, Anatoli P., Derevianko, Jean-Jacques, Hublin, Janet, Kelso, Montgomery, Slatkin, yes, Svante, Pääbo, 8, 2010, Genetic history of an archaic hominin group from Denisova Cave in Siberia, Nature (journal), Nature, 468, 7327, 1053–60, 10.1038/nature09710, 21179161, 2010Natur.468.1053R, 4306417, 10230/25596, JOURNAL, Trinkaus, Erik, Early modern humans, Annual Review of Anthropology, October 2005, 34, 1, 207–30, 10.1146/annurev.anthro.34.030905.154913, The "gracile" or lightly built skeleton of anatomically modern humans has been connected to a change in behavior, including increased cooperation and "resource transport".BOOK, Meldrum, Jeff, Hilton, Charles E., From Biped to Strider: The Emergence of Modern Human Walking, Running, and Resource Transport,weblink 31 March 2004, Springer Science & Business Media, 978-0-306-48000-3, BOOK, Vonk, Jennifer, Shackelford, Todd K., The Oxford Handbook of Comparative Evolutionary Psychology,weblink 13 February 2012, Oxford University Press, USA, 978-0-19-973818-2, 429–, There is evidence that the characteristic human brain development, especially the prefrontal cortex, was due to "an exceptional acceleration of metabolome evolution ... paralleled by a drastic reduction in muscle strength. The observed rapid metabolic changes in brain and muscle, together with the unique human cognitive skills and low muscle performance, might reflect parallel mechanisms in human evolution."JOURNAL, 10.1371/journal.pbio.1001871, 24866127, Exceptional Evolutionary Divergence of Human Muscle and Brain Metabolomes Parallels Human Cognitive and Physical Uniqueness, PLoS Biology, 12, 5, e1001871, 2014, Bozek, Katarzyna, Wei, Yuning, Yan, Zheng, Liu, Xiling, Xiong, Jieyi, Sugimoto, Masahiro, Tomita, Masaru, Pääbo, Svante, Pieszek, Raik, Sherwood, Chet C., Hof, Patrick R., Ely, John J., Steinhauser, Dirk, Willmitzer, Lothar, Bangsbo, Jens, Hansson, Ola, Call, Josep, Giavalisco, Patrick, Khaitovich, Philipp, 4035273, The Schöningen spears and their correlation of finds are evidence that complex technological skills already existed 300,000 years ago, and are the first obvious proof of an active (big game) hunt. H. heidelbergensis already had intellectual and cognitive skills like anticipatory planning, thinking and acting that so far have only been attributed to modern man.ENCYCLOPEDIA, Thieme, H, 2007, Der große Wurf von Schöningen: Das neue Bild zur Kultur des frühen Menschen, 224–28, Die Schöninger Speere – Mensch und Jagd vor 400 000 Jahren, Konrad Theiss Verlag, 978-3-89646-040-0, ENCYCLOPEDIA, Haidle, M.N., 2006, Menschenaffen? Affenmenschen? Mensch! Kognition und Sprache im Altpaläolithikum, 69–97, Conard, N.J., Woher kommt der Mensch, Attempto Verlag, 3-89308-381-2, The ongoing admixture events within anatomically modern human populations make it difficult to estimate the age of the matrilinear and patrilinear most recent common ancestors of modern populations (Mitochondrial Eve and Y-chromosomal Adam). Estimates of the age of Y-chromosomal Adam have been pushed back significantly with the discovery of an ancient Y-chromosomal lineage in 2013, to likely beyond 300,000 years ago.{{refn|group=note|(95% confidence interval 237–581 kya)JOURNAL, Mendez, Fernando, Krahn, Thomas, Schrack, Bonnie, Krahn, Astrid-Maria, Veeramah, Krishna, Woerner, August, Fomine, Forka Leypey Mathew, Bradman, Neil, Thomas, Mark, An African American paternal lineage adds an extremely ancient root to the human Y chromosome phylogenetic tree, American Journal of Human Genetics, 7 March 2013, 10.1016/j.ajhg.2013.02.002,weblink 92, 3, 454–59, 23453668, 3591855, }} There have, however, been no reports of the survival of Y-chromosomal or mitochondrial DNA clearly deriving from archaic humans (which would push back the age of the most recent patrilinear or matrilinear ancestor beyond 500,000 years).JOURNAL, Krings M, Stone A, Schmitz RW, Krainitzki H, Stoneking M, Pääbo S, Neandertal DNA sequences and the origin of modern humans, Cell, 90, 1, 19–30, July 1997, 9230299, 10.1016/S0092-8674(00)80310-4, WEB,weblink Hill, Deborah, 16 March 2004, No Neandertals in the Gene Pool, Science, 2019-05-06, JOURNAL, Serre, 2004, No evidence of Neandertal mtDNA contribution to early modern humans, PLoS Biology, 2, 3, 313–17, 15024415, 10.1371/journal.pbio.0020057, Langaney, A, Chech, M, Teschler-Nicola, M, Paunovic, M, Mennecier, P, Hofreiter, M, Possnert, G, Pääbo, S, 368159, D, Maria Teschler-Nicola, Fossil teeth found at Qesem Cave (Israel) and dated to between 400,000 and 200,000 years ago have been compared to the dental material from the younger (120,000–80,000 years ago) Skhul and Qafzeh hominins.{{refn|group=note|"Although none of the Qesem teeth shows a suite of Neanderthal characters, a few traits may suggest some affinities with members of the Neanderthal evolutionary lineage. However, the balance of the evidence suggests a closer similarity with the Skhul/Qafzeh dental material, although many of these resemblances likely represent plesiomorphous features."JOURNAL, Hershkovitz, I, Smith, P, Sarig, R, Quam, R, Rodríguez, L, García, R, Arsuaga, JL, Barkai, R, Gopher, A, Middle pleistocene dental remains from Qesem Cave (Israel), American Journal of Physical Anthropology, 2011, 144, 4, 575–592, 10.1002/ajpa.21446, 21404234, }}

Dispersal and archaic admixture

{{see|Recent African origin of modern humans|Southern Dispersal|Early human migrations|List of first human settlements}}{{see|Interbreeding between archaic and modern humans}}File:Homo sapiens dispersal routes.jpg|thumb|450x450px|Overview map of the peopling of the worldpeopling of the worldDispersal of early H. sapiens begins soon after its emergence, as evidenced by the North African Jebel Irhoud finds (dated to between 280,000 and 350,000 years ago). There is indirect evidence for modern human presence in West Asia around 270,000 years ago and Dali Man from China is dated at 260,000 years ago.JOURNAL, Posth, Cosimo, etal, Deeply divergent archaic mitochondrial genome provides lower time boundary for African gene flow into Neanderthals, 4 July 2017, Nature Communications, 8, 16046, 10.1038/ncomms16046, 28675384, 5500885, 2017NatCo...816046P, Among extant populations, the Khoi-San (or "Capoid") hunters-gatherers of Southern Africa may represent the human population with the earliest possible divergence within the group Homo sapiens sapiens. Their separation time has been estimated in a 2017 study to be as long as between 260,000 and 350,000 years ago, compatible with the estimated age of H. sapiens. H. s. idaltu, found at Middle Awash in Ethiopia, lived about 160,000 years ago,JOURNAL, White, Tim D., Asfaw, Berhane, Degusta, David, Gilbert, Henry, Richards, Gary D., Suwa, Gen, Howell, Clark F., June 2003, Pleistocene Homo sapiens from Middle Awash, Ethiopia, Nature, 423, 6941, 742–7, 12802332, 10.1038/nature01669, 2003Natur.423..742W, and H. sapiens lived at Omo Kibish in Ethiopia about 195,000 years ago.MAGAZINE, Fossil Reanalysis Pushes Back Origin of Homo sapiens, Scientific American, 2005-02-17, 2019-05-06,weblink Fossil evidence for modern human presence in West Asia is ascertained for 177,000 years ago,WEB,weblink A 177,000-year-old jawbone fossil discovered in Israel is oldest human remains found outside Africa, Ankita, Mehta, 26 January 2018, International Business Times, 2019-05-06, and disputed fossil evidence suggests expansion as far as East Asia by 120,000 years ago.JOURNAL, Bae, Christopher J., Douka, Katerina, Petraglia, Michael D., On the origin of modern humans: Asian perspectives, Science (journal), Science, 8 December 2017, 358, 6368, eaai9067, 10.1126/science.aai9067, 29217544, WEB, Kuo, Lily, Early humans migrated out of Africa much earlier than we thought,weblink 10 December 2017, Quartz (publication), Quartz, 2019-05-06, In July 2019, anthropologists reported the discovery of 210,000 year old remains of a H. sapiens and 170,000 year old remains of a H. neanderthalensis in Apidima Cave, Peloponnese, Greece, more than 150,000 years older than previous H. sapiens finds in Europe.NEWS, Zimmer, Carl, Carl Zimmer, A Skull Bone Discovered in Greece May Alter the Story of Human Prehistory - The bone, found in a cave, is the oldest modern human fossil ever discovered in Europe. It hints that humans began leaving Africa far earlier than once thought.,weblink 10 July 2019, The New York Times, 11 July 2019, NEWS, Staff, 'Oldest remains' outside Africa reset human migration clock,weblink 10 July 2019, Phys.org, 10 July 2019, JOURNAL, Harvati, Katerina, et al., Apidima Cave fossils provide earliest evidence of Homo sapiens in Eurasia,weblink 10 July 2019, Nature (journal), Nature, 10.1038/s41586-019-1376-z, 31292546, 10 July 2019, A significant dispersal event, within Africa and to West Asia, is associated with the African megadroughts during MIS 5, beginning 130,000 years ago.JOURNAL, Rito, T, Richards, MB, Fernandes, V, Alshamali, F, Cerny, V, Pereira, L, Soares, P, The first modern human dispersals across Africa, PLOS ONE, 2013, 8, 11, e80031, 10.1371/journal.pone.0080031, 24236171, 3827445, 2013PLoSO...880031R, A 2011 study located the origin of basal population of contemporary human populations at 130,000 years ago, with the Khoi-San representing an "ancestral population cluster" located in southwestern Africa (near the coastal border of Namibia and Angola).JOURNAL, Henn, Brenna, Gignoux, Christopher R., Jobin, Matthew, 2011, Hunter-gatherer genomic diversity suggests a southern African origin for modern humans, Proceedings of the National Academy of Sciences of the United States of America, 108, 13, 5154–62, 10.1073/pnas.1017511108, 2011PNAS..108.5154H, 21383195, 3069156, File:Ksar Akil Fossils.jpg|thumb|Layer sequence at Ksar Akil in the Levantine corridorLevantine corridorWhile early modern human expansion in Sub-Saharan Africa before 130 kya persisted, early expansion to North Africa and Asia appears to have mostly disappeared by the end of MIS5 (75,000 years ago), and is known only from fossil evidence and from archaic admixture. Asia was re-populated by early modern humans in the so-called "recent out-of-Africa migration" post-dating MIS5, beginning around 70,000 years ago. In this expansion, bearers of mt-DNA haplogroup L3 left East Africa, likely reaching Arabia via the Bab-el-Mandeb, and in the Great Coastal Migration spread to South Asia, Maritime South Asia and Oceania by 65,000 years ago,JOURNAL, Chris, Clarkson, Zenobia, Jacobs, Colin, Pardoe, 2017, Human occupation of northern Australia by 65,000 years ago, Nature, 10.1038/nature22968, 28726833, 547, 7663, 306–310, 2017Natur.547..306C, 2440/107043,weblink NEWS, St. Fleu, Nicholas, Humans First Arrived in Australia 65,000 Years Ago, Study Suggests,weblink New York Times, July 19, 2017, while Europe, East and North Asia, and possibly the Americas, were reached by 50,000 years ago.Evidence for the overwhelming contribution of this "recent" (L3-derived) expansion to all non-African populations was established based on mitochondrial DNA, combined with evidence based on physical anthropology of archaic specimens, during the 1990s and 2000s.{{refn|group=note|"Currently available genetic and archaeological evidence is generally interpreted as supportive of a recent single origin of modern humans in East Africa."JOURNAL, Liu, Hua, etal, 2006, A Geographically Explicit Genetic Model of Worldwide Human-Settlement History, 10.1086/505436, The American Journal of Human Genetics, 79, 2, 230–237, 16826514, 1559480, }}JOURNAL, Out of Africa Revisited, 10.1126/science.308.5724.921g, 2005-05-13, 308, 5724, Science, 921g, The assumption of complete replacement has been revised in the 2010s with the discovery of admixture events (introgression) of populations of H. sapiens with populations of archaic humans over the period of between roughly 100,000 and 30,000 years ago, both in Eurasia and in Sub-Saharan Africa. Neanderthal admixture, in the range of 1-4%, is found in all modern populations outside of Africa, including in Europeans, Asians, Papuan New Guineans, Australian Aboriginals, and Native Americans.JOURNAL, Sankararaman, Sriram, Mallick, Swapan, Patterson, Nick, Reich, David, David Reich (geneticist), The Combined Landscape of Denisovan and Neanderthal Ancestry in Present-Day Humans, Current Biology, 26, 9, 2016, 1241–47, 0960-9822, 10.1016/j.cub.2016.03.037, 27032491, 4864120, This suggests that interbreeding between Neanderthals and anatomically modern humans took place after the recent "out of Africa" migration, likely between 60,000 and 40,000 years ago.JOURNAL, October 17, 2012, North African Populations Carry the Signature of Admixture with Neandertals, PLOS ONE, 10.1371/journal.pone.0047765, 7, 10, e47765, 23082212, 3474783, Sánchez-Quinto, F, Botigué, LR, Civit, S, Arenas, C, Avila-Arcos, MC, Bustamante, CD, Comas, D, Lalueza-Fox, C, 2012PLoSO...747765S, JOURNAL, October 23, 2014, Genome sequence of a 45,000-year-old modern human from western Siberia, Nature, 514, 7523, 445–49, 10.1038/nature13810, 25341783, 4753769, Fu, Q, Li, H, Moorjani, P, Jay, F, Slepchenko, SM, Bondarev, AA, Johnson, PL, Aximu-Petri, A, Prüfer, K, de Filippo, C, Meyer, M, Zwyns, N, Salazar-García, DC, Kuzmin, YV, Keates, SG, Kosintsev, PA, Razhev, DI, Richards, MP, Peristov, NV, Lachmann, M, Douka, K, Higham, TF, Slatkin, M, Hublin, JJ, Reich, D, Kelso, J, Viola, TB, Pääbo, S, 2014Natur.514..445F, MAGAZINE, Brahic, Catherine,weblink Humanity's forgotten return to Africa revealed in DNA, The New Scientist, February 3, 2014, 2019-05-06, Recent admixture analyses have added to the complexity, finding that Eastern Neanderthals derive up to 2% of their ancestry from anatomically modern humans who left Africa some 100 (wikt:kya|kya).JOURNAL, Kuhlwilm, Martin, Ancient gene flow from early modern humans into Eastern Neanderthals, Nature, 17 February 2016, 530, 7591, 429–433, 10.1038/nature16544, 26886800, 4933530, 2016Natur.530..429K, The extent of Neanderthal admixture (and introgression of genes acquired by admixture) varies significantly between contemporary racial groups, being absent in Africans, intermediate in Europeans and highest in East Asians. Certain genes related to UV-light adaptation introgressed from Neanderthals have been found to have been selected for in East Asians specifically from 45,000 years ago until around 5,000 years ago.JOURNAL, Ding, Q., Hu, Y., Xu, S., Wang, J., Jin, L., Neanderthal Introgression at Chromosome 3p21.31 was Under Positive Natural Selection in East Asians, Molecular Biology and Evolution, 2014, Online 2013, 31, 3, 683–695, 10.1093/molbev/mst260, 24336922, The extent of archaic admixture is of the order of about 1% to 4% in Europeans and East Asians, and highest among Melanesians (Denisova hominin admixture), at 4% to 6%. Cumulatively, about 20% of the Neanderthal genome is estimated to remain present spread in contemporary populations.JOURNAL, Vernot, B., Akey, J. M., Resurrecting Surviving Neandertal Lineages from Modern Human Genomes, Science, 2014, 343, 6174, 1017–1021, 10.1126/science.1245938, 24476670, 2014Sci...343.1017V,

Anatomy

{{see also|Human anatomy|Human physical appearance|Anthropometry}}(File:Anatomically Modern Humans archaeological remains, Europe and Africa, directly dated, calibrated carbon dates as of 2013.jpg|thumb|Known archaeological remains of Anatomically Modern Humans in Europe and Africa, directly dated, calibrated carbon dates as of 2013.JOURNAL, Higham, Thomas F. G., Wesselingh, Frank P., Hedges, Robert E. M., Bergman, Christopher A., Douka, Katerina, 2013-09-11, Chronology of Ksar Akil (Lebanon) and Implications for the Colonization of Europe by Anatomically Modern Humans, PLOS ONE, en, 8, 9, e72931, 10.1371/journal.pone.0072931, 1932-6203, 3770606, 24039825, 2013PLoSO...872931D, )Generally, modern humans are more lightly built (or more "gracile") than the more "robust" archaic humans. Nevertheless, contemporary humans exhibit high variability in many physiological traits, and may exhibit remarkable "robustness". There are still a number of physiological details which can be taken as reliably differentiating the physiology of Neanderthals vs. anatomically modern humans.

Anatomical modernity

{{see also|Behavioral modernity}}The term "anatomically modern humans" (AMH) is used with varying scope depending on context, to distinguish "anatomically modern" Homo sapiens from archaic humans such as Neanderthals and Middle and Lower Paleolithic hominins with transitional features intermediate between H. erectus, Neanderthals and early AMH called archaic Homo sapiens.BOOK,weblink Processes in Human Evolution: The Journey from Early Hominins to Neanderthals and Modern Humans, 9780198739906, Ayala, Francisco José, Conde, Camilo José Cela, 2017, BOOK,weblink Human Adaptation: The Biosocial Background, 9781351514712, Cohen, Yehudi A, 2017-07-12, In a convention popular in the 1990s, Neanderthals were classified as a subspecies of H. sapiens, as H. s. neanderthalensis, while AMH (or European early modern humans, EEMH) was taken to refer to "Cro-Magnon" or H. s. sapiens. Under this nomenclature (Neanderthals considered H. sapiens), the term "anatomically modern Homo sapiens" (AMHS) has also been used to refer to EEMH ("Cro-Magnons").BOOK, Schopf, J. William, Major Events in the History of Life,weblink 1992, Jones & Bartlett Learning, 978-0-86720-268-7, 168–, It has since become more common to designate Neanderthals as a separate species, H. neanderthalensis, so that AMH in the European context refers to H. sapiens (but the question is by no means resolved{{refn|group=note|It is important to note that this is a question of conventional terminology, not one of a factual disagreement. Pääbo (2014) frames this as a debate that is unresolvable in principle, "since there is no definition of species perfectly describing the case."BOOK, Pääbo, Svante, Neanderthal Man: In Search of Lost Genomes, Basic Books, New York, 2014, 237, }}).In this more narrow definition of H. sapiens, the subspecies H. s. idaltu, discovered in 2003, also falls under the umbrella of "anatomically modern".WEB,weblink Robert, Sanders, UC Berkeley News, 160,000-year-old fossilized skulls uncovered in Ethiopia are oldest anatomically modern humans, 11 June 2003, 2019-05-07, The recognition of H. s. idaltu as a valid subspecies of the anatomically modern human lineage would justify the description of contemporary humans with the subspecies name H. s. sapiens.A further division of AMH into "early" or "robust" vs. "post-glacial" or "gracile" subtypes has since been used for convenience. The emergence of "gracile AMH" is taken to reflect a process towards a smaller and more fine-boned skeleton beginning around 50,000–30,000 years ago.JOURNAL, 10.1073/pnas.0707650104, Recent acceleration of human adaptive evolution, Proceedings of the National Academy of Sciences, 104, 52, 20753–8, 2007, Hawks, J., Wang, E. T., Cochran, G. M., Harpending, H. C., Moyzis, R. K., 2007PNAS..10420753H, 18087044, 2410101,

Braincase anatomy

{{see|Brain size}}File:Sapiens neanderthal comparison en.png|thumb|upright=1.35|Anatomical comparison of skulls of H. sapiens (left) and H. neanderthalensis (right)(in Cleveland Museum of Natural History)Features compared are the braincase shape, forehead, browridge, nasal bone, projection, cheek bone angulation, chin and occipital contour ]]The cranium lacks a pronounced occipital bun in the neck, a bulge that anchored considerable neck muscles in Neanderthals. Modern humans, even the earlier ones, generally have a larger fore-brain than the archaic people, so that the brain sits above rather than behind the eyes. This will usually (though not always) give a higher forehead, and reduced brow ridge. Early modern people and some living people do however have quite pronounced brow ridges, but they differ from those of archaic forms by having both a supraorbital foramen or notch, forming a groove through the ridge above each eye.JOURNAL, Bhupendra, P., Forehead Anatomy,weblink Medscape references, 2019-05-06, April 2019, This splits the ridge into a central part and two distal parts. In current humans, often only the central section of the ridge is preserved (if it is preserved at all). This contrasts with archaic humans, where the brow ridge is pronounced and unbroken.WEB, How to ID a modern human?,weblink News, 2012, Natural History Museum, London, 11 December 2013, Modern humans commonly have a steep, even vertical forehead whereas their predecessors had foreheads that sloped strongly backwards.ENCYCLOPEDIA, Encarta, Human Evolution,weblinkweblink" title="web.archive.org/web/20091029044339weblink">weblink 29 October 2009, yes, dmy, According to Desmond Morris, the vertical forehead in humans plays an important role in human communication through eyebrow movements and forehead skin wrinkling.BOOK, The Naked Woman: A Study of the Female Body, The Brow,weblink Desmond, Morris, Desmond Morris, 2007, 978-0-312-33853-4, Brain size in both Neanderthals and AMH is significantly larger on average (but overlapping in range) than brain size in H. erectus. Neanderthal and AMH brain sizes are in the same range, but there are differences in the relative sizes of individual brain areas, with significantly larger visual systems in Neanderthals than in AMH.JOURNAL, Pearce, Eiluned, Stringer, Chris, Dunbar, R. I. M., 2013-05-07, New insights into differences in brain organization between Neanderthals and anatomically modern humans, Proceedings of the Royal Society of London B: Biological Sciences, 280, 1758, 20130168, 10.1098/rspb.2013.0168, 0962-8452, 3619466, 23486442, {{refn|group=note|Contemporary human endocranial volume averages at {{cvt|1350|cm3}}, with significant differences between populations, global group means range {{cvt|1085|-|1580|cm3}}.JOURNAL, Smith, C. L., Beals, K. L., Cultural correlates with cranial capacity, American Anthropologist, 1990, 92, 193–200, 10.1525/aa.1990.92.1.02a00150, Neanderthal average is close to {{cvt|1450|cm3}} (male average {{cvt|1600|cm3}}, female average {{cvt|1300|cm3}}), with a range extending up to {{cvt|1736|cm3}} (Amud 1).ENCYCLOPEDIA, Stringer, C, Foley, R, Hominid evolution and community ecology, Human evolution and biological adaptation in the Pleistocene, 1984, Academic Press, New York, 978-0122619205, }}

Jaw anatomy

Compared to archaic people, anatomically modern humans have smaller, differently shaped teeth.JOURNAL, Townsend G, Richards L, Hughes T, Molar intercuspal dimensions: genetic input to phenotypic variation, Journal of Dental Research, 82, 5, 350–5, May 2003, 12709500, 10.1177/154405910308200505, JOURNAL, Keith A, Problems relating to the Teeth of the Earlier Forms of Prehistoric Man, Proceedings of the Royal Society of Medicine, 6, Odontol Sect, 103–124, 1913, 19977113, 2005996, This results in a smaller, more receded dentary, making the rest of the jaw-line stand out, giving an often quite prominent chin. The central part of the mandible forming the chin carries a triangularly shaped area forming the apex of the chin called the mental trigon, not found in archaic humans.BOOK, Tattersall, Jeffrey H, Schwartz, Ian, The human fossil record Craniodental Morphology of Genus Homo (Africa and Asia) (vol 2), 2003, Wiley-Liss, 978-0471319283, 327–328, Particularly in living populations, the use of fire and tools requires fewer jaw muscles, giving slender, more gracile jaws. Compared to archaic people, modern humans have smaller, lower faces.

Body skeleton structure

The body skeletons of even the earliest and most robustly built modern humans were less robust than those of Neanderthals (and from what little we know from Denisovans), having essentially modern proportions. Particularly regarding the long bones of the limbs, the distal bones (the radius/ulna and tibia/fibula) are nearly the same size or slightly shorter than the proximal bones (the humerus and femur). In ancient people, particularly Neanderthals, the distal bones were shorter, usually thought to be an adaptation to cold climate.JOURNAL, Steegmann, A. Theodore, Cerny, Frank J., Holliday, Trenton W., Neandertal cold adaptation: Physiological and energetic factors, American Journal of Human Biology, 2002, 14, 5, 566–583, 10.1002/ajhb.10070, 12203812, The same adaptation can be found in some modern people living in the polar regions.JOURNAL, Stock, J.T., Hunter-gatherer postcranial robusticity relative to patterns of mobility, climatic adaptation, and selection for tissue economy, American Journal of Physical Anthropology, October 2006, 131, 2, 194–204, 10.1002/ajpa.20398, 16596600, Height ranges overlap between Neanderthals and AMH, with Neanderthal averages cited as {{cvt|164|to|168|cm|0}} and {{cvt|152|to|156|cm|0}} for males and females, respectively.{{refn|group=note|"Based on 45 long bones from maximally 14 males and 7 females, Neanderthals' height averages between 164 and 168 (males) resp. 152 to 156 cm (females). This height is indeed 12-14 cm lower than the height of post-WWII Europeans, but compared to Europeans some 20,000 or 100 years ago, it is practically identical or even slightly higher."JOURNAL, Helmuth H, Body height, body mass and surface area of the Neanderthals, Zeitschrift für Morphologie und Anthropologie, 82, 1, 1–12, 1998, 9850627, }} By comparison, contemporary national averages range between {{cvt|158|to|184|cm|0}} in males and {{cvt|147|to|172|cm|0}} in females. Neanderthal ranges approximate the height distribution measured among Malay people, for one.{{refn|group=note|Malay, 20–24 (N{{=}} m:749 f:893, Median{{=}} m:{{Height|cm=166}} f:{{Height|cm=155}}, SD{{=}} m:{{Height|cm=6.46}} f:{{Height|cm=6.04}})JOURNAL,weblink Distribution of Body Weight, Height and Body Mass Index in a National Sample of Malaysian Adults, 11072496, 55, 1, March 2000, Med. J. Malaysia, 108–28, Lim TO, Ding LM, Zaki M, etal, }}

Recent evolution

{{see|Human genetic variability|Race and genetics|Sexual selection in humans}}Following the peopling of Africa some 130,000 years ago, and the recent Out-of-Africa expansion some 70,000 to 50,000 years ago, some sub-populations of H. sapiens have been essentially isolated for tens of thousands of years prior to the early modern Age of Discovery. Combined with archaic admixture this has resulted in significant genetic variation, which in some instances has been shown to be the result of directional selection taking place over the past 15,000 years, i.e. significantly later than possible archaic admixture events.NEWS,weblink Still Evolving, Human Genes Tell New Story, Wade, N, 2006-03-07, 2008-07-10, The New York Times, Some climatic adaptations, such as high-altitude adaptation in humans, are thought to have been acquired by archaic admixture. Introgression of genetic variants acquired by Neanderthal admixture have different distributions in European and East Asians, reflecting differences in recent selective pressures. A 2014 study reported that Neanderthal-derived variants found in East Asian populations showed clustering in functional groups related to immune and haematopoietic pathways, while European populations showed clustering in functional groups related to the lipid catabolic process.{{refn|group=note|"Specifically, genes in the LCP [lipid catabolic process] term had the greatest excess of NLS in populations of European descent, with an average NLS frequency of 20.8±2.6% versus 5.9±0.08% genome wide (two-sided t-test, P


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