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{{Short description|Hominin fossil found in Siberia}}
'''{{coor|57.744|N|71.200|E|display=title}}'''
{{Infobox artefact
| name = Ust'-Ishim man
| image = Femur ust-ishim 2.jpg
| image_caption = Femur from the Ust'-Ishim man
| image_size = 300
| material =
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| writing =
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| discovered =
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|lat_deg = 57
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'''Ust'-Ishim man''' is the term given to the 45,000-year-old remains of one of the [[anatomically modern humans|early modern humans]] to inhabit western [[Siberia]].<ref name="SciAm">{{cite web| url=http://www.scientificamerican.com/article/45-000-year-old-mans-genome-sequenced/ |title= 45,000-Year-Old Man's Genome Sequenced| author= Callaway, Ewen & Nature magazine |website=Scientific American |date= 23 October 2014|access-date= 24 October 2014}}</ref> The fossil is notable in that it had intact DNA which permitted the complete [[genome sequence|sequencing]] of its [[genome]], one of the oldest modern [[human genome]]s to be so decoded.<ref name="SciAm"/><ref>{{Cite journal |last1=Prüfer |first1=Kay |last2=Posth |first2=Cosimo |last3=Yu |first3=He |last4=Stoessel |first4=Alexander |last5=Spyrou |first5=Maria A. |last6=Deviese |first6=Thibaut |last7=Mattonai |first7=Marco |last8=Ribechini |first8=Erika |last9=Higham |first9=Thomas |last10=Velemínský |first10=Petr |last11=Brůžek |first11=Jaroslav |last12=Krause |first12=Johannes |date=2021 |title=A genome sequence from a modern human skull over 45,000 years old from Zlatý kůň in Czechia |journal=Nature Ecology & Evolution |language=en |volume=5 |issue=6 |pages=820–825 |doi=10.1038/s41559-021-01443-x |issn=2397-334X |pmc=8175239 |pmid=33828249}}</ref>▼
 
▲{{Infobox fossil|catalog number=|image=Femur ust-ishim 2.jpg|species=[[Human]]|age=45,000 years|place discovered=[[Omsk Oblast|Omsk]], [[Russia]]|date discovered=2008|common name=Ust'-Ishim man|discovered by=Nikolai Peristov|caption=Femur from the Ust'-Ishim man}}'''Ust'-Ishim man''' is the term given to the 45,000-year-old remains of one of the [[anatomically modern humans|early modern humans]] to inhabit western [[Siberia]].<ref name="SciAm">{{cite web| url=http://www.scientificamerican.com/article/45-000-year-old-mans-genome-sequenced/ |title= 45,000-Year-Old Man's Genome Sequenced| author= Callaway, Ewen & Nature magazine |website=Scientific American |date= 23 October 2014|access-date= 24 October 2014}}</ref> The fossil is notable in that it had intact DNA which permitted the complete [[genome sequence|sequencing]] of its [[genome]], one of the oldest modern [[human genome]]s to be so decoded.<ref name="SciAm"/><ref>{{Cite journal |last1=Prüfer |first1=Kay |last2=Posth |first2=Cosimo |last3=Yu |first3=He |last4=Stoessel |first4=Alexander |last5=Spyrou |first5=Maria A. |last6=Deviese |first6=Thibaut |last7=Mattonai |first7=Marco |last8=Ribechini |first8=Erika |last9=Higham |first9=Thomas |last10=Velemínský |first10=Petr |last11=Brůžek |first11=Jaroslav |last12=Krause |first12=Johannes |date=2021 |title=A genome sequence from a modern human skull over 45,000 years old from Zlatý kůň in Czechia |journal=Nature Ecology & Evolution |language=en |volume=5 |issue=6 |pages=820–825 |doi=10.1038/s41559-021-01443-x |issn=2397-334X |pmc=8175239 |pmid=33828249|bibcode=2021NatEE...5..820P }}</ref>
The remains consist of a single bone—left femur—of a male [[hunter-gatherer]], which was discovered in 2008<ref name="MaxPlanck" /> protruding from the bank of the [[Irtysh River]] by Nikolai Peristov, a Russian sculptor who specialises in carving [[mammoth ivory]].<ref name="SciAm"/> Peristov showed the fossil to a forensic investigator who suggested that it might be of human origin.<ref name="SciAm"/> The fossil was named after the [[Ust'-Ishim District]] of Siberia where it had been discovered.<ref name="SciAm"/>▼
 
▲The remains consist of a single bone—left femur—of a male [[hunter-gatherer]], which was discovered in 2008<ref name="MaxPlanck" /> protruding from the bank of the [[Irtysh River]] by Nikolai Peristov, a Russian sculptor who specialises in carving [[mammoth ivory]].<ref name="SciAm"/> Peristov showed the fossil to a forensic investigator who suggested that it might be of human origin.<ref name="SciAm"/> The fossil was named after the [[Ust'-IshimIshimsky District]] of Siberia where it had been discovered.<ref name="SciAm"/>
 
==Genome sequencing==
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===Y-DNA and mtDNA===
Ust'-Ishim man belongs to [[Y-DNA haplogroup]] [[Haplogroup K2|K2]]. The two subclades of K2 are K2a and K2b, and he has been found to be positive for some but not all SNPs of the K2a (or NO*) subclade, such as M2308.<ref>{{cite web | url=http://www.yfull.com/tree/K-M2308 | title=K-M2308 YTree }}</ref><ref>{{Cite web|url=https://yfull.com/tree/K-Y28299/|title = K-Y28299 YTree}}</ref><ref name="poznik">{{cite journal|pmc=4884158|year=2016|last1=Poznik|first1=G. D|title=Punctuated bursts in human male demography inferred from 1,244 worldwide Y-chromosome sequences|journal=Nature Genetics|volume=48|issue=6|pages=593–599|last2=Xue|first2=Y|last3=Mendez|first3=F. L|last4=Willems|first4=T. F|last5=Massaia|first5=A|last6=Wilson Sayres|first6=M. A|last7=Ayub|first7=Q|last8=McCarthy|first8=S. A|last9=Narechania|first9=A|last10=Kashin|first10=S|last11=Chen|first11=Y|last12=Banerjee|first12=R|last13=Rodriguez-Flores|first13=J. L|last14=Cerezo|first14=M|last15=Shao|first15=H|last16=Gymrek|first16=M|last17=Malhotra|first17=A|last18=Louzada|first18=S|last19=Desalle|first19=R|last20=Ritchie|first20=G. R|last21=Cerveira|first21=E|last22=Fitzgerald|first22=T. W|last23=Garrison|first23=E|last24=Marcketta|first24=A|last25=Mittelman|first25=D|last26=Romanovitch|first26=M|last27=Zhang|first27=C|last28=Zheng-Bradley|first28=X|last29=Abecasis|first29=G. R|last30=McCarroll|first30=S. A|display-authors=29|doi=10.1038/ng.3559|pmid=27111036}}</ref> In the original paper, he was classified only as [[Haplogroup K-M9]] (KxLT).<ref name = "poznik" /><ref name = "YFull">[https://www.yfull.com/tree/NO/ YFull] Haplogroup YTree v5.06 at 25 September 2017</ref><ref name = "Karmin2015">{{cite journal | last1 = Karmin | first1 = Monika | last2 = Saag | first2 = Lauri | last3 = Vicente | first3 = Mário |display-authors=etal | year = 2015 | title = ", "A recent bottleneck of Y chromosome diversity coincides with a global change in culture | journal = Genome Research | volume = 25 | issue = 4| pages = 459–466 | doi = 10.1101/gr.186684.114 | pmid = 25770088 | pmc = 4381518 }}</ref>
 
He belonged to [[Human mitochondrial DNA haplogroup|mitochondrial DNA haplogroup]] [[haplogroup R (mtDNA)|R*]], differing from the root sequence of R by a single mutation.
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Both of these haplogroups and descendant [[subclades]] are now found among populations throughout [[Eurasia]], [[Oceania]] and [[The Americas]], although no direct descendants of Ust Ishim man's specific lineages are known from modern populations.
 
Examination of the sequenced genome indicates that Ust'-Ishim man lived at a point in time (270,000 to 45,000 years ago) between the [[Recent African origin of modern humans|first wave]] of [[anatomically modern humans]] (270,000 years ago) that migrated out of Africa and the divergence of that population into distinct populations (45,000 years ago), in terms of [[autosomal DNA]] in different parts of Eurasia.<ref name="MaxPlanck">{{cite web|url=http://www.mpg.de/8710423/genome-earliest-modern-human|title=Earliest modern human sequenced |date=22 October 2014 |website=Max-Planck-Gesellschaft |access-date=30 October 2014}}</ref> Consequently, Ust'-Ishim man is not more closely related to the first two major migrations of ''Homo Sapiens'' eastward from Africa into Asia: a group that migrated along the coast of [[South Asia]], or a group that moved north-east through [[Central Asia]].<ref name=Nature>{{cite journal|author= Qiaomei Fu, Heng Li, Priya Moorjani, Flora Jay, Sergey M. Slepchenko, Aleksei A. Bondarev, Philip L. F. Johnson, Ayinuer Aximu-Petri, Kay Prüfer, Cesare de Filippo, Matthias Meyer, Nicolas Zwyns, Domingo C. Salazar-García, Yaroslav V. Kuzmin, Susan G. Keates, Pavel A. Kosintsev, Dmitry I. Razhev, Michael P. Richards, Nikolai V. Peristov, Michael Lachmann, Katerina Douka, Thomas F. G. Higham, Montgomery Slatkin, Jean-Jacques Hublin, David Reich, Janet Kelso, T. Bence Viola & Svante Pääbo|title=Genome sequence of a 45,000-year-old modern human from western Siberia |journal=Nature | issue= 7523| pages=445–449|date=23 October 2014|doi=10.1038/nature13810 | pmid=25341783 | volume=514 | pmc=4753769|bibcode=2014Natur.514..445F |hdl= 10550/42071}}</ref> When compared to other ancient remains, Ust'-Ishim man is more closely related, in terms of autosomal DNA to [[Tianyuan man]], found near Beijing and dating from 42,000 to 39,000 years ago; [[Mal'ta boy]] (or MA-1), a child who lived 24,000 years ago along the [[Bolshaya Belaya River]] near today's [[Irkutsk]] in Siberia, or; [[La Braña man]] – a hunter-gatherer who lived in [[La Braña]] (modern [[Spain]]) about 8,000 years ago.<ref name="ScienceMag">{{cite web| url=https://www.science.org/content/article/oldest-human-genome-reveals-when-our-ancestors-had-sex-neandertals |title=Oldest human genome reveals when our ancestors had sex with Neandertals | author= Wade, Lizzie | date= 22 October 2014| website=Science |publisher=American Association for the Advancement of Science | access-date= 24 October 2014}}</ref><ref name="ScienceMag25Oct13">{{cite journal |title=Ancient DNA Links Native Americans With Europe |author=Balter, Michael |journal=Science |volume=342 |issue=6157 |pages=409–410 | date= 25 October 2013|doi=10.1126/science.342.6157.409 |pmid=24159019 |bibcode=2013Sci...342..409B |doi-access=free }}</ref><ref name="ScienceMag26Jan14">{{cite web |url=https://www.science.org/content/article/how-farming-reshaped-our-genomes |title=How Farming Reshaped Our Genomes |author=Balter, Michael |website=Science |publisher=American Association for the Advancement of Science | date= 26 January 2014|access-date=25 October 2014}}</ref>
 
===Relationship with Neanderthals===
Analysis of modern human genomes reveals that humans interbred with [[Neanderthal]]s between 3786,000 and 8637,000 years ago,<ref name="LiveScience04Oct12">{{cite web|url=http://www.livescience.com/23730-neanderthals-modern-humans-interbreeding.html|title=Humans Broke Off Neanderthal Sex After Discovering Eurasia|author=Choi, Charles Q.|website=LiveScience |date=4 October 2012|access-date= 25 October 2014}}</ref> resulting in the DNA of modern humans outside Africa containing between 1.5 and 2.1 percent DNA of Neanderthal origin.<ref name="LiveScience18Dec13">{{cite web|url=http://www.livescience.com/42056-neanderthal-woman-genome-sequenced.html|title=Neanderthal Woman's Genome Reveals Unknown Human Lineage|author=Choi, Charles Q. |website=LiveScience |date=18 December 2013|access-date= 25 October 2014}}</ref> Neanderthal DNA in modern humans occurs in broken fragments; however, the Neanderthal DNA in Ust'-Ishim man occurs in clusters, indicating that Ust'-Ishim man lived in the immediate aftermath of the genetic interchange.<ref name="ScienceMag"/> The genomic sequencing of Ust'-Ishim man has led to refinement of the estimated date of mating between the two hominin species to between 52,000 and 58,000 years ago.<ref name="ScienceMag"/>
 
No relationship between [[Denisovans]] and the Ust'-Ishim man has been checked, although Denisovans have some descendants in Oceania and Asia.
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| image = Principal component analysis of ancient and present-day individuals from worldwide populations (clear).png
| image-width = <!-- Do not change image width (300) as this will affect the layout --> 301
| caption = Genetic proximity of Ust'-Ishim to [[Ancient North Eurasian]] populations ([[Yana RHS|Yana]], [[Mal'ta]] and [[Afontova Gora]]), within a principal component analysis of ancient and present-day individuals from worldwide populations.<ref name="TG">{{cite journal |last1=Gakuhari |first1=Takashi |last2=Nakagome |first2=Shigeki |last3=Rasmussen |first3=Simon |last4=Allentoft |first4=Morten E. |title=Ancient Jomon genome sequence analysis sheds light on migration patterns of early East Asian populations |journal=Communications Biology |date=25 August 2020 |volume=3 |issue=1 |page=Fig.1 A, B |doi=10.1038/s42003-020-01162-2 |url=https://www.nature.com/articles/s42003-020-01162-2 |language=en |issn=2399-3642|hdl=20.500.12000/50006 |hdl-access=free }}</ref>
| annotations = {{Annotation|125|104|[[File:Red circle 50%.svg|30px]]|text-align=center|font-weight=bold|font-style=normal|font-size=7|color=#000000}}
}}
Ust'-Ishim was equally related to modern [[East Asians]], [[Oceanians]] and certain ancient [[West Eurasians|West Eurasian]] populations, such as the ancient[[Goyet EuropeansCaves|Goyet specimen]].<ref name="fu">{{cite journal | last1=Fu | first1=Qiaomei | last2=Li | first2=Heng | last3=Moorjani | first3=Priya | last4=Jay | first4=Flora | last5=Slepchenko | first5=Sergey M. | last6=Bondarev | first6=Aleksei A. | last7=Johnson | first7=Philip L. F. | last8=Aximu-Petri | first8=Ayinuer | last9=Prüfer | first9=Kay | last10=de Filippo | first10=Cesare | last11=Meyer | first11=Matthias | last12=Zwyns | first12=Nicolas | last13=Salazar-García | first13=Domingo C. | last14=Kuzmin | first14=Yaroslav V. | last15=Keates | first15=Susan G. | last16=Kosintsev | first16=Pavel A. | last17=Razhev | first17=Dmitry I. | last18=Richards | first18=Michael P. | last19=Peristov | first19=Nikolai V. | last20=Lachmann | first20=Michael | last21=Douka | first21=Katerina | last22=Higham | first22=Thomas F. G. | last23=Slatkin | first23=Montgomery | last24=Hublin | first24=Jean-Jacques | last25=Reich | first25=David | last26=Kelso | first26=Janet | last27=Viola | first27=T. Bence | last28=Pääbo | first28=Svante | title=Genome sequence of a 45,000-year-old modern human from western Siberia | journal=Nature | volume=514 | issue=7523 | date=2014-10-23 | issn=0028-0836 | pmid=25341783 | pmc=4753769 | doi=10.1038/nature13810 | pages=445–449| bibcode=2014Natur.514..445F }}</ref><ref name="ScienceMag"/> Modern Europeans are more closely related to other ancient remains.<ref name="ScienceMag4Sep14">{{cite web |url=http://news.sciencemag.org/biology/2014/09/three-part-ancestry-europeans |title=Three-part ancestry for Europeans |author=Gibbons, Ann |website=Science |publisher=American Association for the Advancement of Science |date=4 September 2014 |access-date=30 October 2014 |archive-url=https://web.archive.org/web/20141011034223/http://news.sciencemag.org/biology/2014/09/three-part-ancestry-europeans |archive-date=11 October 2014 |url-status=dead |df=dmy-all }}</ref> "The finding that the Ust’-Ishim individual is equally closely related to present-day Asians and to 8,000- to 24,000-year-old individuals from western Eurasia, but not to present-day Europeans, is compatible with the hypothesis that present-day Europeans derive some of their ancestry from a population that did not participate in the initial dispersals of modern humans into Europe and Asia."<ref>{{Cite journal |last1=Fu |first1=Qiaomei |last2=Li |first2=Heng |last3=Moorjani |first3=Priya |last4=Jay |first4=Flora |last5=Slepchenko |first5=Sergey M. |last6=Bondarev |first6=Aleksei A. |last7=Johnson |first7=Philip L.F. |last8=Petri |first8=Ayinuer A. |last9=Prüfer |first9=Kay |last10=de Filippo |first10=Cesare |last11=Meyer |first11=Matthias |last12=Zwyns |first12=Nicolas |last13=Salazar-Garcia |first13=Domingo C. |last14=Kuzmin |first14=Yaroslav V. |last15=Keates |first15=Susan G. |date=2014-10-23 |title=The genome sequence of a 45,000-year-old modern human from western Siberia |journal=Nature |volume=514 |issue=7523 |pages=445–449 |doi=10.1038/nature13810 |issn=0028-0836 |pmc=4753769 |pmid=25341783|bibcode=2014Natur.514..445F }}</ref>{{Phylogenetic tree for ancient Eurasians}}
 
In a 2016 study, modern [[Tibetan people|Tibetans]] were identified as the modern population that has the most alleles in common with Ust'-Ishim man.<ref name="Lu 2016">{{cite journal|last1=Lu|first1=Dongsheng|display-authors=etal|title=Ancestral Origins and Genetic History of Tibetan Highlanders|journal=[[The American Journal of Human Genetics]]|date=September 1, 2016|volume=99|issue=3|pages=580–594|doi=10.1016/j.ajhg.2016.07.002|pmid=27569548|pmc=5011065}}</ref> According to a 2017 study, "Siberian and East Asian populations shared 38% of their ancestry" with Ust’-Ishim man.<ref>{{Cite journal|last1=Wong|first1=Emily H. M.|last2=Khrunin|first2=Andrey|last3=Nichols|first3=Larissa|last4=Pushkarev|first4=Dmitry|last5=Khokhrin|first5=Denis|last6=Verbenko|first6=Dmitry|last7=Evgrafov|first7=Oleg|last8=Knowles|first8=James|last9=Novembre|first9=John|date=2017-01-01|title=Reconstructing genetic history of Siberian and Northeastern European populations|journal=Genome Research|language=en|volume=27|issue=1|pages=1–14|doi=10.1101/gr.202945.115|issn=1088-9051|pmid=27965293|pmc=5204334}}</ref> A 2021 study foundargues that "the Ust’Ishim and Oase1[[Oase 1]] individuals showed no more affinity to westernany thanmodern towestern or eastern Eurasian populations, suggesting that they did not contribute ancestry to later Eurasian populations, as previously shown.<ref>{{Cite journal |last1=Hajdinjak |first1=Mateja |last2=Mafessoni |first2=Fabrizio |last3=Skov |first3=Laurits |last4=Vernot |first4=Benjamin |last5=Hübner |first5=Alexander |last6=Fu |first6=Qiaomei |last7=Essel |first7=Elena |last8=Nagel |first8=Sarah |last9=Nickel |first9=Birgit |last10=Richter |first10=Julia |last11=Moldovan |first11=Oana Teodora |last12=Constantin |first12=Silviu |last13=Endarova |first13=Elena |last14=Zahariev |first14=Nikolay |last15=Spasov |first15=Rosen |date=April 2021 |title=Initial Upper Palaeolithic humans in Europe had recent Neanderthal ancestry |journal=Nature |language=en |volume=592 |issue=7853 |pages=253–257 |doi=10.1038/s41586-021-03335-3 |issn=1476-4687 |pmc=8026394 |pmid=33828320|bibcode=2021Natur.592..253H }}</ref>▼
 
In 2022, a study determined that the Ust'Ishim man was part of an [[Initial Upper Paleolithic]] wave (>45kya) "ascribed to a population movement with uniform genetic features and material culture" ([[Ancient East Eurasians]]), and sharing deep ancestry with [[Bacho Kiro cave|Bacho Kiro]] and the [[Tianyuan man]], as well as ancestors of modern-day [[Papuans]] (Australasians). The Ust’Ishim man is best described as basal to all modern East Eurasian populations, and diverged from their ancestor shortly after the divergence from Ancient Western Eurasians (represented by the [[Kostenki-14]] specimen).<ref>{{Cite web |title=Genetics and Material Culture Support Repeated Expansions into Paleolithic Eurasia from a Population Hub Out of Africa |url=https://academic.oup.com/gbe/article/14/4/evac045/6563828 |access-date=2023-11-10 |website=academic.oup.com}}</ref>
▲In a 2016 study, modern [[Tibetan people|Tibetans]] were identified as the modern population that has the most alleles in common with Ust'-Ishim man.<ref name="Lu 2016">{{cite journal|last1=Lu|first1=Dongsheng|display-authors=etal|title=Ancestral Origins and Genetic History of Tibetan Highlanders|journal=[[The American Journal of Human Genetics]]|date=September 1, 2016|volume=99|issue=3|pages=580–594|doi=10.1016/j.ajhg.2016.07.002|pmid=27569548|pmc=5011065}}</ref> According to a 2017 study, "Siberian and East Asian populations shared 38% of their ancestry" with Ust’-Ishim man.<ref>{{Cite journal|last1=Wong|first1=Emily H. M.|last2=Khrunin|first2=Andrey|last3=Nichols|first3=Larissa|last4=Pushkarev|first4=Dmitry|last5=Khokhrin|first5=Denis|last6=Verbenko|first6=Dmitry|last7=Evgrafov|first7=Oleg|last8=Knowles|first8=James|last9=Novembre|first9=John|date=2017-01-01|title=Reconstructing genetic history of Siberian and Northeastern European populations|journal=Genome Research|language=en|volume=27|issue=1|pages=1–14|doi=10.1101/gr.202945.115|issn=1088-9051|pmid=27965293|pmc=5204334}}</ref> A 2021 study found that "the Ust’Ishim and Oase1 individuals showed no more affinity to western than to eastern Eurasian populations, suggesting that they did not contribute ancestry to later Eurasian populations, as previously shown.<ref>{{Cite journal |last1=Hajdinjak |first1=Mateja |last2=Mafessoni |first2=Fabrizio |last3=Skov |first3=Laurits |last4=Vernot |first4=Benjamin |last5=Hübner |first5=Alexander |last6=Fu |first6=Qiaomei |last7=Essel |first7=Elena |last8=Nagel |first8=Sarah |last9=Nickel |first9=Birgit |last10=Richter |first10=Julia |last11=Moldovan |first11=Oana Teodora |last12=Constantin |first12=Silviu |last13=Endarova |first13=Elena |last14=Zahariev |first14=Nikolay |last15=Spasov |first15=Rosen |date=April 2021 |title=Initial Upper Palaeolithic humans in Europe had recent Neanderthal ancestry |journal=Nature |language=en |volume=592 |issue=7853 |pages=253–257 |doi=10.1038/s41586-021-03335-3 |issn=1476-4687 |pmc=8026394 |pmid=33828320|bibcode=2021Natur.592..253H }}</ref>
 
==References==
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[[Category:Upper Paleolithic Homo sapiens fossils]]
[[Category:People from Siberia]]
[[Category:History of Omsk Oblast]]
[[Category:2008 in Russia]]
[[Category:2008 archaeological discoveries]]
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