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=== Encephalization === [[File:Skull evolution.png|center|thumb|920px|Skulls of successive (or near-successive, depending on the source) human evolutionary ancestors,{{efn|There is no general agreement on the line of [[species|special]] descent of ''H. sapiens'' from ''H. erectus''. Some of the species depicted in the image may not actually represent a direct evolutionary ancestor to ''H. sapiens'', and may not directly derive from one another, namely: * ''H. heidelbergensis'' likely did not descend from ''H. antecessor''.<ref name="Welker2020">{{Cite journal |doi=10.1038/s41586-020-2153-8 |issn=1476-4687 |volume=580 |issue=7802 |pages=235–238 |last1=Welker |first1=F. |last2=Ramos-Madrigal |first2=J. |last3=Gutenbrunner |first3=P. |display-authors=etal |title=The dental proteome of ''Homo antecessor'' |journal=[[Nature (journal)|Nature]] |date=April 1, 2020 |pmid=32269345 |pmc=7582224 |bibcode=2020Natur.580..235W |s2cid=214736611 |url= http://eprints.whiterose.ac.uk/159068/1/Welker_etal_Hominin1_AAM.docx |access-date=November 10, 2021 |archive-date=November 10, 2021 |archive-url= https://web.archive.org/web/20211110200345/https://eprints.whiterose.ac.uk/159068/1/Welker_etal_Hominin1_AAM.docx |url-status=live}}{{collapsible list |title=Full list of authors |bullets=true |Frido Welker |Jazmín Ramos-Madrigal |Petra Gutenbrunner |Meaghan Mackie |Shivani Tiwary |Rosa Rakownikow Jersie-Christensen |Cristina Chiva |Marc R. Dickinson |Martin Kuhlwilm |Marc de Manuel |Pere Gelabert |María Martinón-Torres |Ann Margvelashvili |Juan Luis Arsuaga |Eudald Carbonell |Tomas Marques-Bonet |Kirsty Penkman |Eduard Sabidó |Jürgen Cox |Jesper V. Olsen |David Lordkipanidze |Fernando Racimo |Carles Lalueza-Fox |José María Bermúdez de Castro |Eske Willerslev |Enrico Cappellini}}</ref> * ''H. heidelbergensis'' is likely not an ancestor to ''H. sapiens'', nor is ''H. antecessor''.<ref name="Welker2020" /> * ''H. ergaster'' is often considered the next evolutionary ancestor to ''H. sapiens'' following ''H. erectus'', however, there is considerable uncertainty as to the accuracy of classifying it as a separate species from ''H. erectus'' at all.<ref>{{Cite journal |last1=Dennell |first1=Robin |last2=Roebroeks |first2=Wil |date=2005 |title=An Asian perspective on early human dispersal from Africa |journal=[[Nature (journal)|Nature]] |volume=438 |issue=7071 |pages=1099–1104 |doi=10.1038/nature04259 |pmid=16371999 |bibcode=2005Natur.438.1099D |s2cid=4405913 |url= https://www.nature.com/articles/nature04259 |access-date=November 10, 2021 |archive-date=October 31, 2021 |archive-url= https://web.archive.org/web/20211031154723/https://www.nature.com/articles/nature04259 |url-status=live}}</ref>}} up until 'modern' ''Homo sapiens''<br /> * [[Mya (unit)|Mya]] – million years ago, [[kya (unit)|kya]] – thousand years ago]] [[File:Brain size and tooth size in hominins.jpg|thumb|upright=1.5|Brain size and tooth size in hominins]] The human species eventually developed a much larger brain than that of other primates—typically {{convert|1330|cm3|abbr=on}} in modern humans, nearly three times the size of a chimpanzee or gorilla brain.<ref name="Schoeneman">{{cite journal |last=Schoenemann |first=P. Thomas |date=October 2006 |title=Evolution of the Size and Functional Areas of the Human Brain |journal=[[Annual Review of Anthropology]] |volume=35 |pages=379–406 |doi=10.1146/annurev.anthro.35.081705.123210 |s2cid=7611321 |issn=0084-6570}}</ref> After a period of stasis with ''Australopithecus anamensis'' and ''Ardipithecus'', species which had smaller brains as a result of their bipedal locomotion,<ref>{{cite web |last1=Brown |first1=Graham |last2=Fairfax |first2=Stephanie |last3=Sarao |first3=Nidhi |title=Tree of Life Web Project: Human Evolution |url= http://tolweb.org/treehouses/?treehouse_id=3710 |website=www.tolweb.org |access-date=August 24, 2015 |archive-date=June 6, 2020 |archive-url= https://web.archive.org/web/20200606091921/http://tolweb.org/treehouses/?treehouse_id=3710 |url-status=live}}</ref> the pattern of [[encephalization]] started with ''Homo habilis'', whose {{convert|600|cm3|abbr=on}} brain was slightly larger than that of chimpanzees. This evolution continued in ''Homo erectus'' with {{convert|800–1,100|cm3|abbr=on}}, and reached a maximum in Neanderthals with {{convert|1200–1,900|cm3|abbr=on}}, larger even than modern ''Homo sapiens''. This brain increase manifested during postnatal [[neural development|brain growth]], far exceeding that of other apes ([[heterochrony]]). It also allowed for extended periods of [[Observational learning|social learning]] and [[language acquisition]] in juvenile humans, beginning as much as 2 million years ago. Encephalization may be due to a dependency on calorie-dense, difficult-to-acquire food.<ref>{{Cite journal |last1=Kaplan |first1=Hillard |last2=Hill |first2=Kim |last3=Lancaster |first3=Jane |last4=Hurtado |first4=Magdelena |date=August 16, 2000 |title=A Theory of Human Life History Evolution: Diet, Intelligence, and Longevity |url= https://www.unm.edu/~hkaplan/KaplanHillLancasterHurtado_2000_LHEvolution.pdf |journal=Evolutionary Anthropology |volume=9 |issue=4 |pages=156–185 |doi=10.1002/1520-6505(2000)9:4<156::AID-EVAN5>3.0.CO;2-7 |s2cid=2363289 |access-date=September 20, 2019 |archive-date=February 15, 2020 |archive-url= https://web.archive.org/web/20200215120245/http://www.unm.edu/~hkaplan/KaplanHillLancasterHurtado_2000_LHEvolution.pdf |url-status=live}}</ref> Furthermore, the changes in the structure of [[human brain]]s may be even more significant than the increase in size.<ref name="Park2007">{{cite journal |last1=Park |first1=M. S. |last2=Nguyen |first2=A. D. |last3=Aryan |first3=H. E. |last4=U |first4=H. S. |last5=Levy |first5=M. L. |last6=Semendeferi |first6=K. |date=March 2007 |title=Evolution of the human brain: Changing brain size and the fossil record |journal=[[Neurosurgery (journal)|Neurosurgery]] |volume=60 |issue=3 |pages=555–562 |issn=0148-396X |pmid=17327801 |s2cid=19610624 |doi=10.1227/01.NEU.0000249284.54137.32}}</ref><ref name="Bruner2007">{{cite journal |last=Bruner |first=Emiliano |date=December 2007 |title=Cranial shape and size variation in human evolution: Structural and functional perspectives |citeseerx=10.1.1.391.288 |journal=Child's Nervous System |volume=23 |issue=12 |pages=1357–1365 |doi=10.1007/s00381-007-0434-2 |issn=0256-7040 |pmid=17680251 |s2cid=16163137}}</ref><ref>{{cite journal |last=Potts |first=Richard |author-link=Rick Potts |date=October 2012 |title=Evolution and Environmental Change in Early Human Prehistory |journal=Annual Review of Anthropology |volume=41 |pages=151–167 |doi=10.1146/annurev-anthro-092611-145754 |issn=0084-6570}}</ref><ref name="Leonard_2007">{{cite journal |last1=Leonard |first1=William R. |last2=Snodgrass |first2=J. Josh |last3=Robertson |first3=Marcia L. |date=August 2007 |title=Effects of brain evolution on human nutrition and metabolism |journal=[[Annual Review of Nutrition]] |volume=27 |pages=311–327 |doi=10.1146/annurev.nutr.27.061406.093659 |issn=0199-9885 |pmid=17439362 |s2cid=18869516}}</ref> Fossilized skulls shows the brain size in early humans fell within the range of modern humans 300,000 years ago, but only got its present-day brain shape between 100,000 and 35,000 years ago.<ref>{{Cite journal |title=The evolution of modern human brain shape - Science |journal=Science Advances |date=January 5, 2018 |volume=4 |issue=1 |doi=10.1126/sciadv.aao5961 |last1=Neubauer |first1=Simon |last2=Hublin |first2=Jean-Jacques |last3=Gunz |first3=Philipp |pages=eaao5961 |pmid=29376123 |pmc=5783678 |bibcode=2018SciA....4.5961N}}</ref> [[File:Students explore hominid evolution.jpg|alt=Three students hold three different skulls in front of their faces, to show the difference in size and shape compared to the modern head|left|thumb|The size and shape of the skull changed over time. The leftmost, and largest, is a replica of a modern human skull.]] The [[temporal lobe]]s, which contain centers for language processing, have increased disproportionately, as has the [[prefrontal cortex]], which has been related to complex decision-making and moderating social behavior.<ref name="Schoeneman" /> Encephalization has been tied to increased starches<ref name="NYT-20150813">{{cite news |last=Zimmer |first=Carl |author-link=Carl Zimmer |date=August 13, 2015 |title=For Evolving Brains, a 'Paleo' Diet Full of Carbs |work=The New York Times |url= https://www.nytimes.com/2015/08/13/science/for-evolving-brains-a-paleo-diet-full-of-carbs.html |url-access=limited |access-date=August 14, 2015 |archive-url= https://ghostarchive.org/archive/20220101/https://www.nytimes.com/2015/08/13/science/for-evolving-brains-a-paleo-diet-full-of-carbs.html |archive-date=January 1, 2022}}{{cbignore}}</ref> and meat<ref>{{cite journal |last=Mann |first=Neil |date=September 2007 |title=Meat in the human diet: An anthropological perspective |journal=Nutrition & Dietetics |volume=64 |issue=Supplement 4 |pages=S102–S107 |doi=10.1111/j.1747-0080.2007.00194.x |issn=1747-0080 |doi-access=free}}</ref><ref>{{cite press release |last=McBroom |first=Patricia |date=June 14, 1999 |title=Meat-eating was essential for human evolution, says UC Berkeley anthropologist specializing in diet |location=Berkeley |publisher=[[University of California Press]] |url= http://www.berkeley.edu/news/media/releases/99legacy/6-14-1999a.html |access-date=April 25, 2015 |archive-date=April 20, 2015 |archive-url= https://web.archive.org/web/20150420060844/http://www.berkeley.edu/news/media/releases/99legacy/6-14-1999a.html |url-status=live}}</ref> in the diet, however a 2022 meta study called into question the role of meat.<ref>{{Cite journal |last1=Barr |first1=W. Andrew |last2=Pobiner |first2=Briana |last3=Rowan |first3=John |last4=Du |first4=Andrew |last5=Faith |first5=J. Tyler |date=February 1, 2022 |title=No sustained increase in zooarchaeological evidence for carnivory after the appearance of ''Homo erectus'' |journal=[[Proceedings of the National Academy of Sciences of the United States of America]] |volume=119 |issue=5 |doi=10.1073/pnas.2115540119 |doi-access=free |issn=0027-8424 |pmid=35074877 |pmc=8812535 |bibcode=2022PNAS..11915540B}}</ref> Other factors are the development of cooking,<ref name="Organ_2011">{{cite journal |last1=Organ |first1=Chris |last2=Nunn |first2=Charles L. |last3=Machanda |first3=Zarin |last4=Wrangham |first4=Richard W. |author4-link=Richard Wrangham |date=August 30, 2011 |title=Phylogenetic rate shifts in feeding time during the evolution of ''Homo'' |journal=[[Proceedings of the National Academy of Sciences of the United States of America]] |volume=108 |issue=35 |pages=14555–14559 |doi=10.1073/pnas.1107806108 |issn=0027-8424 |pmc=3167533 |pmid=21873223 |bibcode=2011PNAS..10814555O |doi-access=free}}</ref> and it has been proposed that intelligence increased as a response to an increased necessity for [[Dunbar's number|solving social problems]] as human society became more complex.<ref name="David-Barrett">{{cite journal |last1=David-Barrett |first1=T. |last2=Dunbar |first2=R. I. M. |date=2013 |title=Processing Power Limits Social Group Size: Computational Evidence for the Cognitive Costs of Sociality |journal=Proceedings of the Royal Society of London B: Biological Sciences |volume=280 |issue=1765 |page=20131151 |doi=10.1098/rspb.2013.1151 |pmid=23804623 |pmc=3712454}}</ref> Changes in skull morphology, such as smaller mandibles and mandible muscle attachments, allowed more room for the brain to grow.{{sfn|Bown|Rose|1987}} The increase in volume of the [[neocortex]] also included a rapid increase in size of the [[cerebellum]]. Its function has traditionally been associated with balance and fine motor control, but more recently with [[speech]] and [[cognition]]. The great apes, including hominids, had a more pronounced cerebellum relative to the neocortex than other primates. It has been suggested that because of its function of sensory-motor control and learning complex muscular actions, the cerebellum may have underpinned human technological adaptations, including the preconditions of speech.<ref>{{cite journal |last1=Barton |first1=Robert A. |last2=Venditti |first2=Chris |date=October 20, 2014 |title=Rapid Evolution of the Cerebellum in Humans and Other Great Apes |journal=[[Current Biology]] |volume=24 |issue=20 |pages=2440–2444 |issn=0960-9822 |doi=10.1016/j.cub.2014.08.056 |pmid=25283776 |s2cid=5041106 |doi-access=free |bibcode=2014CBio...24.2440B}}</ref><ref>{{cite journal |last1=Starowicz-Filip |first1=Anna |last2=Milczarek |first2=Olga |last3=Kwiatkowski |first3=Stanisław |last4=Bętkowska-Korpała |first4=Barbara |last5=Prochwicz |first5=Katarzyna |date=2013 |title=Cerebellar cognitive affective syndrome CCAS – a case report |journal=Archives of Psychiatry and Psychotherapy |volume=15 |issue=3 |pages=57–64 |doi=10.12740/APP/18666 |doi-access=free}}</ref><ref>{{cite journal |last1=Yu |first1=Feng |last2=Jiang |first2=Qing-jun |last3=Sun |first3=Xi-yan |last4=Zhang |first4=Rong-wei |date=August 22, 2014 |title=A new case of complete primary cerebellar agenesis: Clinical and imaging findings in a living patient |journal=[[Brain (journal)|Brain]] |doi=10.1093/brain/awu239 |issn=1460-2156 |pmid=25149410 |volume=138 |issue=Pt 6 |pages=e353 |pmc=4614135}}</ref><ref>{{cite journal |last=Weaver |first=Anne H. |date=March 8, 2005 |title=Reciprocal evolution of the cerebellum and neocortex in fossil humans |journal=[[Proceedings of the National Academy of Sciences of the United States of America]] |volume=102 |issue=10 |pages=3576–3580 |doi=10.1073/pnas.0500692102 |issn=0027-8424 |pmc=553338 |pmid=15731345 |bibcode=2005PNAS..102.3576W |doi-access=free}}</ref> The immediate survival advantage of encephalization is difficult to discern, as the major brain changes from ''Homo erectus'' to ''Homo heidelbergensis'' were not accompanied by major changes in technology. It has been suggested that the changes were mainly social and behavioural, including increased empathic abilities,<ref>{{cite book |last=Klein |first=Stefan |date=2014 |title=Survival of the Nicest |isbn=978-1-61519-090-4 |publisher=The Experiment}}</ref><ref>{{Cite journal |title=Social Network Size Affects Neural Circuits in Macaques |journal=[[Science (journal)|Science]] |date=November 4, 2011 |issn=0036-8075 |pmid=22053054 |pages=697–700 |volume=334 |issue=6056 |doi=10.1126/science.1210027 |last1=Sallet |first1=J. |last2=Mars |first2=R. B. |last3=Noonan |first3=M. P. |last4=Andersson |first4=J. L. |last5=O'Reilly |first5=J. K. |last6=Jbabdi |first6=S. |last7=Croxson |first7=P. L. |last8=Jenkinson |first8=M. |last9=Miller |first9=K. L. |bibcode=2011Sci...334..697S |s2cid=206536017}}</ref> increases in size of social groups,<ref name="David-Barrett" /><ref>{{cite journal |last1=Dunbar |first1=R. I. M. |date=1992 |title=Neocortex size as a constraint on group size in primates |journal=Journal of Human Evolution |volume=22 |issue=6 |pages=469–493 |doi=10.1016/0047-2484(92)90081-j |doi-access=free |bibcode=1992JHumE..22..469D}}</ref><ref>{{Cite journal |title=Encephalization is not a universal macroevolutionary phenomenon in mammals but is associated with sociality |journal=[[Proceedings of the National Academy of Sciences of the United States of America]] |date=December 14, 2010 |issn=0027-8424 |pmc=3003036 |pmid=21098277 |pages=21582–21586 |volume=107 |issue=50 |doi=10.1073/pnas.1005246107 |first1=Susanne |last1=Shultz |first2=Robin |last2=Dunbar |bibcode=2010PNAS..10721582S |doi-access=free}}</ref> and increased behavioral plasticity.<ref>{{Cite journal |last=Richard |first=Potts |date=1998 |title=Environmental Hypotheses of Hominin Evolution |journal=American Journal of Physical Anthropology |volume=107 |issue=S27 |pages=93–136 |doi=10.1002/(sici)1096-8644(1998)107:27+<93::aid-ajpa5>3.0.co;2-x |pmid=9881524}}</ref> Humans are unique in the ability to acquire information through social transmission and adapt that information.<ref>{{Cite book |last=Richerson |first=Peter J. |title=Not by Genes Alone: How Culture Transformed Human Evolution |date=2006 |publisher=[[University of Chicago Press]] |isbn=0-226-71212-5 |oclc=642503808}}</ref> The emerging field of [[cultural evolution]] studies human sociocultural change from an evolutionary perspective.<ref>{{Cite journal |last1=Mesoudi |first1=Alex |last2=Whiten |first2=Andrew |last3=Laland |first3=Kevin N. |date=August 2006 |title=Towards a unified science of cultural evolution |journal=Behavioral and Brain Sciences |volume=29 |issue=4 |pages=329–347 |doi=10.1017/s0140525x06009083 |pmid=17094820 |issn=0140-525X}}</ref> [[File:Homo skull changes.png|center|thumb|750px|Evolution of the shape, size, and contours of the human (''Homo'') skull<ref>{{cite journal |last1=Berger |first1=L. R. |last2=Hawks |first2=J. |last3=Dirks |first3=P. H. |last4=Elliott |first4=M. |last5=Roberts |first5=E. M. |title=''Homo naledi'' and Pleistocene hominin evolution in subequatorial Africa |journal=eLife |volume=6 |date=May 9, 2017 |pmid=28483041 |pmc=5423770 |doi=10.7554/elife.24234 |doi-access=free}}</ref><ref>{{cite book |url= https://www.researchgate.net/publication/278682043 |title=Analyzing Hominin Hominin Phylogeny: Cladistic Approach |last1=Strait |first1=David |last2=Grine |first2=Frederick |last3=Fleagle |first3=John |name-list-style=vanc |date=2015 |isbn=978-3-642-39978-7 |pages=1989–2014 (cladogram p. 2006) |publisher=Springer}}.</ref><!--ref name="Saylor 2015 483–488" /-->{{efn|name=habilis|The conventional estimate on the age of ''H. habilis'' is at roughly 2.1 to 2.3 million years.{{sfn|Stringer|1994|p=242}}<ref>{{cite book |last1=Schrenk |first1=F. |last2=Kullmer |first2=O. |last3=Bromage |first3=T. |chapter=Chapter 9: The Earliest Putative ''Homo'' Fossils |editor1-last=Henke |editor1-first=W. |editor2-last=Tattersall |editor2-first=I. |title=Handbook of Paleoanthropology |date=2007 |pages=1611–1631 |doi=10.1007/978-3-540-33761-4_52}}</ref> Suggestions for pushing back the age to 2.8 Mya were made in 2015 based on the discovery of [[LD 350-1|a jawbone]].<ref>{{cite journal |last1=Spoor |first1=F. |last2=Gunz |first2=P. |last3=Neubauer |first3=S. |last4=Stelzer |first4=S. |last5=Scott |first5=N. |last6=Kwekason |first6=A. |last7=Dean |first7=M. C. |title=Reconstructed ''Homo habilis'' type OH 7 suggests deep-rooted species diversity in early ''Homo'' |journal=[[Nature (journal)|Nature]] |volume=519 |issue=7541 |pages=83–6 |date=March 2015 |pmid=25739632 |doi=10.1038/nature14224 |bibcode=2015Natur.519...83S |s2cid=4470282}}</ref>}}<ref>{{cite journal |last=Schuster |first=A. M. |date=1997 |title=Earliest Remains of Genus ''Homo'' |url= http://archive.archaeology.org/9701/newsbriefs/homo.html |journal=Archaeology |volume=50 |access-date=March 5, 2015 |number=1 |archive-date=March 17, 2015 |archive-url= https://web.archive.org/web/20150317002614/http://archive.archaeology.org/9701/newsbriefs/homo.html |url-status=live}} The line to the earliest members of ''Homo'' was derived from ''[[Australopithecus]]'', a genus which had separated from the Chimpanzee–human last common ancestor by late [[Miocene]] or early [[Pliocene]] times. <!--Why is so much of the [[Chimpanzee–human last common ancestor]] article reproduced in this footnote? with date estimates by several specialists ranging from 13 million years ago to more recently than six million years ago. * {{cite journal |last1=Arnason |first1=U. |last2=Gullberg |first2=A. |last3=Janke |first3=A. |title=Molecular timing of primate divergences as estimated by two nonprimate calibration points |journal=Journal of Molecular Evolution |volume=47 |issue=6 |pages=718–27 |date=December 1998 |pmid=9847414 |doi=10.1007/PL00006431 |bibcode=1998JMolE..47..718A}} * {{cite journal |last1=Patterson |first1=N. |last2=Richter |first2=D. J. |last3=Gnerre |first3=S. |last4=Lander |first4=E. S. |last5=Reich |first5=D. |title=Genetic evidence for complex speciation of humans and chimpanzees |journal=[[Nature (journal)|Nature]] |volume=441 |issue=7097 |pages=1103–1108 |date=June 2006 |pmid=16710306 |doi=10.1038/nature04789 |bibcode=2006Natur.441.1103P}} * {{cite journal |last=Wakeley |first=J. |title=Complex speciation of humans and chimpanzees |journal=[[Nature (journal)|Nature]] |volume=452 |issue=7184 |pages=E3-4; discussion E4 |date=March 2008 |pmid=18337768 |doi=10.1038/nature06805 |bibcode=2008Natur.452....3W}} "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 [[CHLCA#Current estimates|current estimates regarding complex speciation]]. --></ref>{{efn|''H. erectus'' in the narrow sense (the Asian species) was extinct by 140,000 years ago, ''[[Homo erectus soloensis]]'', found in [[Java (island)|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.<ref>{{cite journal |last1=Indriati |first1=E. |last2=Swisher |first2=C. C. |last3=Lepre |first3=C. |last4=Quinn |first4=R. L. |last5=Suriyanto |first5=R. A. |last6=Hascaryo |first6=A. T. |last7=Grün |first7=R. |last8=Feibel |first8=C. S. |last9=Pobiner |first9=B. L. |last10=Aubert |first10=M. |last11=Lees |first11=W. |last12=Antón |first12=S. C. |display-authors=5 |title=The age of the 20 meter Solo River terrace, Java, Indonesia and the survival of ''Homo erectus'' in Asia |journal=[[PLoS One]] |volume=6 |issue=6 |pages=e21562 |date=2011 |pmid=21738710 |pmc=3126814 |doi=10.1371/journal.pone.0021562 |bibcode=2011PLoSO...621562I |doi-access=free}}{{collapsible list |title=Full list of authors |bullets=true |Etty Indriati |Carl C. Swisher III |Christopher Lepre |Rhonda L. Quinn |Rusyad A. Suriyanto |Agus T. Hascaryo |Rainer Grün |Craig S. Feibel |Briana L. Pobiner |Maxime Aubert |Wendy Lees |Susan C. Antón}}</ref>}}<ref name=":3">{{cite journal |last1=Mondal |first1=M. |last2=Bertranpetit |first2=J. |last3=Lao |first3=O. |title=Approximate Bayesian computation with deep learning supports a third archaic introgression in Asia and Oceania |journal=Nature Communications |volume=10 |issue=1 |page=246 |date=January 2019 |pmid=30651539 |pmc=6335398 |doi=10.1038/s41467-018-08089-7 |bibcode=2019NatCo..10..246M}}</ref><ref name="Zeitoun 2003 148–156">{{cite journal |last=Zeitoun |first=V. |title=High occurrence of a basicranial feature in ''Homo erectus'': Anatomical description of the preglenoid tubercle |journal=The Anatomical Record Part B: The New Anatomist |volume=274 |issue=1 |pages=148–56 |date=September 2003 |pmid=12964205 |doi=10.1002/ar.b.10028 |doi-access=free}}</ref><ref name="Proceedings 2015">{{cite journal |last1=Dembo |first1=M. |last2=Matzke |first2=N. J. |last3=Mooers |first3=A. Ø. |last4=Collard |first4=M. |title=Bayesian analysis of a morphological supermatrix sheds light on controversial fossil hominin relationships |journal=Proceedings. Biological Sciences |volume=282 |issue=1812 |page=20150943 |date=August 2015 |pmid=26202999 |pmc=4528516 |doi=10.1098/rspb.2015.0943}}</ref><ref name=":2">{{cite journal |last1=Dembo |first1=M. |last2=Radovčić |first2=D. |last3=Garvin |first3=H. M. |last4=Laird |first4=M. F. |last5=Schroeder |first5=L. |last6=Scott |first6=J. E. |last7=Brophy |first7=J. |last8=Ackermann |first8=R. R. |last9=Musiba |first9=C. M. |last10=de Ruiter |first10=D. J. |last11=Mooers |first11=A. 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A. |title=Neanderthal-Denisovan ancestors interbred with a distantly related hominin |journal=Science Advances |volume=6 |issue=8 |pages=eaay5483 |date=February 2020 |pmid=32128408 |pmc=7032934 |doi=10.1126/sciadv.aay5483 |doi-access=free |bibcode=2020SciA....6.5483R}}</ref>]]
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