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=== Demic diffusion and north-south differences === The estimated genetic contribution of northern Han to southern Han is substantial in the ancestral patrilineage in addition to a geographic [[cline (biology)|cline]] that exists for the corresponding matrilineage. These genetic findings align with historical records outlining the continuous and large migratory waves of northern Han Chinese inhabitants settling in southern China, escaping dynastic changes, geopolitical upheavals, instability, warfare and famine.<ref>{{cite book |last=Gernet |first=Jacques |url=https://archive.org/details/historyofchinese00gern_0 |title=A History of Chinese Civilization |publisher=Cambridge University Press |year=1996 |isbn=978-0-521-49781-7 |edition=2nd}}</ref><ref name="Genetic evidence supports demic dif" /><ref>{{Cite book |last1=Crawford |first1=Dorothy H. |title=Cancer Virus: The story of Epstein-Barr Virus |last2=Rickinson |first2=Alan |last3=Johannessen |first3=Ingolfur |publisher=Oxford University Press |year=2014 |publication-date=March 14, 2014 |pages=98}}</ref><ref name="Springer" /><ref name="Six Dynasties" /><ref name="MS" /><ref name="QC" /><ref name="China: A New Cultural History" /><ref name="The History of China" /><ref name="books.google.com" /><ref name="Hunter Publishing, Inc" /><ref name="China: An Environmental History" /><ref name="Lexington Books" /><ref name="Cambridge University Press" /><ref name="Newspaper Enterprise Limited" /><ref name="Rowman Altamira" />{{Excessive citations inline|date=September 2024}} The subsequent intermarriages between northern Han migrants and southern aborigines over the past few thousand years gave rise to modern Chinese demographics - a Han Chinese super-majority and minority non-Han Chinese indigenous peoples.<ref name="Genetic evidence supports demic dif" /> Han Chinese from Fujian and Guangdong show excessive ancestries from Late Neolithic Fujianese sources (35.0–40.3%), which are more significant in modern [[Amis people|Ami]], [[Atayal people|Atayal]] and [[Kankanaey people|Kankanaey]] (66.9–74.3%), and less significant in Han Chinese from Zhejiang (22%), Jiangsu (17%) and Shandong (8%).<ref>{{Cite journal |last1=Huang |first1=Xiufeng |last2=Xia |first2=Zi-Yang |last3=Bin |first3=Xiaoyun |last4=He |first4=Guanglin |display-authors=3 |date=2022 |title=Genomic Insights Into the Demographic History of the Southern Chinese |journal=Frontiers in Ecology and Evolution |volume=10 |doi=10.3389/fevo.2022.853391 |doi-access=free}}</ref> Ancient population admixture with Ami and Atayal also exists for Han Chinese from Guangdong and [[Sichuan]]<ref>{{cite journal |last1=Chiang |first1=Charleston W K |last2=Mangul |first2=Serghei |last3=Robles |first3=Christopher |last4=Sankararaman |first4=Sriram |title=A Comprehensive Map of Genetic Variation in the World's Largest Ethnic Group—Han Chinese |journal=Molecular Biology and Evolution |date=November 2018 |volume=35 |issue=11 |pages=2736–2750 |doi=10.1093/molbev/msy170 |pmid=30169787 |pmc=6693441 }}</ref><ref>{{cite journal |last1=Lo |first1=Yun-Hua |last2=Cheng |first2=Hsueh-Chien |last3=Hsiung |first3=Chia-Ni |last4=Yang |first4=Show-Ling |last5=Wang |first5=Han-Yu |last6=Peng |first6=Chia-Wei |last7=Chen |first7=Chun-Yu |last8=Lin |first8=Kung-Ping |last9=Kang |first9=Mei-Ling |last10=Chen |first10=Chien-Hsiun |last11=Chu |first11=Hou-Wei |last12=Lin |first12=Chiao-Feng |last13=Lee |first13=Mei-Hsuan |last14=Liu |first14=Quintin |last15=Satta |first15=Yoko |last16=Lin |first16=Cheng-Jui |last17=Lin |first17=Marie |last18=Chaw |first18=Shu-Miaw |last19=Loo |first19=Jun-Hun |last20=Shen |first20=Chen-Yang |last21=Ko |first21=Wen-Ya |title=Detecting Genetic Ancestry and Adaptation in the Taiwanese Han People |journal=Molecular Biology and Evolution |date=27 September 2021 |volume=38 |issue=10 |pages=4149–4165 |doi=10.1093/molbev/msaa276 |pmid=33170928 |pmc=8476137 }}</ref> and the ancestors of [[Taiwanese Han]].<ref>{{cite journal |last1=Lo |first1=Yun-Hua |last2=Cheng |first2=Hsueh-Chien |last3=Hsiung |first3=Chia-Ni |last4=Yang |first4=Show-Ling |last5=Wang |first5=Han-Yu |last6=Peng |first6=Chia-Wei |last7=Chen |first7=Chun-Yu |last8=Lin |first8=Kung-Ping |last9=Kang |first9=Mei-Ling |last10=Chen |first10=Chien-Hsiun |last11=Chu |first11=Hou-Wei |last12=Lin |first12=Chiao-Feng |last13=Lee |first13=Mei-Hsuan |last14=Liu |first14=Quintin |last15=Satta |first15=Yoko |last16=Lin |first16=Cheng-Jui |last17=Lin |first17=Marie |last18=Chaw |first18=Shu-Miaw |last19=Loo |first19=Jun-Hun |last20=Shen |first20=Chen-Yang |last21=Ko |first21=Wen-Ya |title=Detecting Genetic Ancestry and Adaptation in the Taiwanese Han People |journal=Molecular Biology and Evolution |date=27 September 2021 |volume=38 |issue=10 |pages=4149–4165 |doi=10.1093/molbev/msaa276 |pmid=33170928 |pmc=8476137 }}</ref> Overall, Cantonese people represent the southernmost Han sub-group, splitting from northern Han between the Qin and Tang dynasties,<ref>{{cite journal |last1=Chen |first1=Jieming |last2=Zheng |first2=Houfeng |last3=Bei |first3=Jin-Xin |last4=Sun |first4=Liangdan |last5=Jia |first5=Wei-hua |last6=Li |first6=Tao |last7=Zhang |first7=Furen |last8=Seielstad |first8=Mark |last9=Zeng |first9=Yi-Xin |last10=Zhang |first10=Xuejun |last11=Liu |first11=Jianjun |title=Genetic Structure of the Han Chinese Population Revealed by Genome-wide SNP Variation |journal=The American Journal of Human Genetics |date=December 2009 |volume=85 |issue=6 |pages=775–785 |doi=10.1016/j.ajhg.2009.10.016 |pmid=19944401 |pmc=2790583 }}</ref><ref>{{cite report |type=Preprint |doi=10.1101/2020.07.03.166413 |title=Fine-scale Population Structure and Demographic History of Han Chinese Inferred from Haplotype Network of 111,000 Genomes |date=2020 |last1=Lan |first1=Ao |last2=Kang |first2=Kang |last3=Tang |first3=Senwei |last4=Wu |first4=Xiaoli |last5=Wang |first5=Lizhong |last6=Li |first6=Teng |last7=Weng |first7=Haoyi |last8=Deng |first8=Junjie |last9=Zheng |first9=Qiang |last10=Yao |first10=Xiaotian |last11=Chen |first11=Gang }}</ref> although there is evidence of additional admixture with Teochew and Hakka people.<ref>{{cite journal |last1=Xiao |first1=Cheng |last2=Yang |first2=Xingyi |last3=Yu |first3=Zhonghao |last4=Wu |first4=Weibin |last5=Wang |first5=Yuan |last6=Xu |first6=Quyi |last7=Chen |first7=Ling |title=Genetic polymorphism and variability in the Guangdong Hakka, Teochew, and Cantonese groups: A comprehensive analysis of 19 X-STRs |journal=Annals of Human Genetics |date=September 2023 |volume=87 |issue=5 |pages=232–240 |doi=10.1111/ahg.12518 |pmid=37337755 }}</ref> Other studies suggest that Han Chinese from [[Fujian]] and Taiwan are equally as southern.<ref>{{Cite journal |last=He |first=Guanglin |last2=Li |first2=Yinxiang |last3=Zou |first3=Xing |last4=Yeh |first4=Hui-Yuan |display-authors=3 |date=2021 |title=The northern gene flow into southeastern East Asians inferred from genome wide array genotyping |url=https://www.biorxiv.org/content/10.1101/2021.07.25.453681v1.full |journal=bioRxiv |quote=Fujian and Taiwan populations were localized at the southmost end of the Han Chinese cline and showed a close relationship with modern Tai-Kadai-speaking populations and ancient southern East Asians (Liangdao, Hanben, Tanshishan, Xitoucun and so on). |via=bioRxiv}}</ref> Han Chinese from Guangdong also show affinities with local Kra-Dai and Hmong-Mien groups<ref>{{cite journal |last1=Du |first1=Weian |last2=Feng |first2=Chunlei |last3=Yao |first3=Ting |last4=Xiao |first4=Cheng |last5=Huang |first5=Hongyan |last6=Wu |first6=Weibin |last7=Zhu |first7=Linnan |last8=Qiao |first8=Honghua |last9=Liu |first9=Chao |last10=Chen |first10=Ling |title=Genetic variation and forensic efficiency of 30 indels for three ethnic groups in Guangxi: relationships with other populations |journal=PeerJ |date=3 May 2019 |volume=7 |pages=e6861 |doi=10.7717/peerj.6861 |doi-access=free |pmid=31110924 |pmc=6501771 }}</ref><ref>{{cite journal |last1=Wang |first1=Fei |last2=He |first2=Guanglin |last3=Wang |first3=Zheng |last4=Wang |first4=Mengge |last5=Liu |first5=Jing |last6=Zou |first6=Xing |last7=Wang |first7=Shouyu |last8=Song |first8=Mengyuan |last9=Ye |first9=Ziwei |last10=Xie |first10=Mingkun |last11=Hou |first11=Yiping |title=Population Genetics and Forensic Efficiency of 30 InDel Markers in Four Chinese Ethnic Groups Residing in Sichuan |journal=Forensic Sciences Research |date=3 July 2022 |volume=7 |issue=3 |pages=498–502 |doi=10.1080/20961790.2020.1737470 |pmid=36353334 |pmc=9639512 }}</ref><ref>{{cite report |type=Preprint |doi=10.1101/2021.06.19.449013 |title=New insights from the combined discrimination of modern/Ancient genome-wide shared alleles and haplotypes: Differentiated demographic history reconstruction of Tai-Kadai and Sinitic people in South China |date=2021 |last1=Wang |first1=Mengge |last2=He |first2=Guanglin |last3=Zou |first3=Xing |last4=Chen |first4=Pengyu |last5=Wang |first5=Zheng |last6=Tang |first6=Renkuan |last7=Yang |first7=Xiaomin |last8=Chen |first8=Jing |last9=Yang |first9=Meiqing |last10=Li |first10=Yingxiang |last11=Liu |first11=Jing |last12=Wang |first12=Fei |last13=Zhao |first13=Jing |last14=Guo |first14=Jianxin |last15=Hu |first15=Rong |last16=Wei |first16=Lan-Hai |last17=Chen |first17=Gang |last18=Yeh |first18=Hui-Yuan |last19=Wang |first19=Chuan-Chao }}</ref><ref>{{cite journal |last1=Wang |first1=Fei |last2=Song |first2=Feng |last3=Song |first3=Mengyuan |last4=Luo |first4=Haibo |last5=Hou |first5=Yiping |title=Genetic structure and paternal admixture of the modern Chinese Zhuang population based on 37 Y-STRs and 233 Y-SNPs |journal=Forensic Science International: Genetics |date=May 2022 |volume=58 |pages=102681 |doi=10.1016/j.fsigen.2022.102681 |pmid=35263703 }}</ref> and cluster with Han Chinese from Guangxi and Hainan, who themselves mixed with the minority peoples of their respective provinces.<ref>{{cite journal |last1=Cong |first1=Pei-Kuan |last2=Bai |first2=Wei-Yang |last3=Li |first3=Jin-Chen |last4=Yang |first4=Meng-Yuan |last5=Khederzadeh |first5=Saber |last6=Gai |first6=Si-Rui |last7=Li |first7=Nan |last8=Liu |first8=Yu-Heng |last9=Yu |first9=Shi-Hui |last10=Zhao |first10=Wei-Wei |last11=Liu |first11=Jun-Quan |last12=Sun |first12=Yi |last13=Zhu |first13=Xiao-Wei |last14=Zhao |first14=Pian-Pian |last15=Xia |first15=Jiang-Wei |last16=Guan |first16=Peng-Lin |last17=Qian |first17=Yu |last18=Tao |first18=Jian-Guo |last19=Xu |first19=Lin |last20=Tian |first20=Geng |last21=Wang |first21=Ping-Yu |last22=Xie |first22=Shu-Yang |last23=Qiu |first23=Mo-Chang |last24=Liu |first24=Ke-Qi |last25=Tang |first25=Bei-Sha |last26=Zheng |first26=Hou-Feng |title=Genomic analyses of 10,376 individuals in the Westlake BioBank for Chinese (WBBC) pilot project |journal=Nature Communications |date=26 May 2022 |volume=13 |issue=1 |page=2939 |doi=10.1038/s41467-022-30526-x |pmid=35618720 |pmc=9135724 }}</ref><ref>{{cite journal |last1=Zou |first1=Xing |last2=Nie |first2=Qianyun |last3=Li |first3=Wenhui |last4=Chen |first4=Yinyu |last5=Song |first5=Tao |last6=Zhang |first6=Peng |title=Genetic variation and phylogenetic analysis of 23 STR in Chinese Han population from Hainan, Southern China |journal=Medicine |date=31 May 2024 |volume=103 |issue=22 |pages=e38428 |doi=10.1097/MD.0000000000038428 |pmid=39259071 |pmc=11142786 }}</ref><ref name=":1">{{cite journal |last1=Sun |first1=Qiuxia |last2=Wang |first2=Mengge |last3=Lu |first3=Tao |last4=Duan |first4=Shuhan |last5=Liu |first5=Yan |last6=Chen |first6=Jing |last7=Wang |first7=Zhiyong |last8=Sun |first8=Yuntao |last9=Li |first9=Xiangping |last10=Wang |first10=Shaomei |last11=Lu |first11=Liuyi |last12=Hu |first12=Liping |last13=Yun |first13=Libing |last14=Yang |first14=Junbao |last15=Yan |first15=Jiangwei |last16=Nie |first16=Shengjie |last17=Zhu |first17=Yanfeng |last18=Chen |first18=Gang |last19=Wang |first19=Chuan-Chao |last20=Liu |first20=Chao |last21=He |first21=Guanglin |last22=Tang |first22=Renkuan |title=Differentiated adaptative genetic architecture and language-related demographical history in South China inferred from 619 genomes from 56 populations |journal=BMC Biology |date=6 March 2024 |volume=22 |issue=1 |page=55 |doi=10.1186/s12915-024-01854-9 |doi-access=free |pmid=38448908 |pmc=10918984 }}</ref> One study shows that Han Chinese from northern Guangxi have more ancestry related to Austronesian, Kra-Dai and Austroasiatic groups, than those from southern Guangxi.<ref name=":1" /> ==== Patrilineal DNA ==== Typical Y-DNA haplogroups of present-day Han Chinese include Haplogroup O-M95, [[Haplogroup O-M122]], [[Haplogroup O-M175]], C,[[Haplogroup N-M231]] and [[Haplogroup Q-M120]].<ref name="AD">{{cite journal |last1=Zhao |first1=Yong-Bin |last2=Zhang |first2=Ye |last3=Zhang |first3=Quan-Chao |last4=Li |first4=Hong-Jie |last5=Cui |first5=Ying-Qiu |last6=Xu |first6=Zhi |last7=Jin |first7=Li |last8=Zhou |first8=Hui |last9=Zhu |first9=Hong |date=4 May 2015 |title=Ancient DNA Reveals That the Genetic Structure of the Northern Han Chinese Was Shaped Prior to 3,000 Years Ago |journal=PLOS ONE |volume=10 |issue=5 |pages=e0125676 |bibcode=2015PLoSO..1025676Z |doi=10.1371/journal.pone.0125676 |pmc=4418768 |pmid=25938511 |doi-access=free}}</ref> The Y-chromosome haplogroup distribution between Southern Han Chinese and Northern Han Chinese populations and principal core component analysis indicates that almost all modern Han Chinese populations form a tight cluster in their Y chromosome: * Haplogroups prevalent in non-Han southern natives such as O1b-M110, O2a1-M88 and O3d-M7, which are prevalent in non-Han southern natives, were observed in 4% of Southern Han Chinese and not at all in the Northern Han.<ref name="table">{{cite journal |last1=Wen |first1=Bo |last2=Li |first2=Hui |last3=Lu |first3=Daru |last4=Song |first4=Xiufeng |last5=Zhang |first5=Feng |last6=He |first6=Yungang |last7=Li |first7=Feng |last8=Gao |first8=Yang |last9=Mao |first9=Xianyun |last10=Zhang |first10=Liang |last11=Qian |first11=Ji |last12=Tan |first12=Jingze |last13=Jin |first13=Jianzhong |last14=Huang |first14=Wei |last15=Deka |first15=Ranjan |last16=Su |first16=Bing |last17=Chakraborty |first17=Ranajit |last18=Jin |first18=Li |title=Genetic evidence supports demic diffusion of Han culture |journal=Nature |date=September 2004 |volume=431 |issue=7006 |pages=302–305 |bibcode=2004Natur.431..302W |doi=10.1038/nature02878 |pmid=15372031 }}</ref><ref name="EJH">{{cite journal |last1=Xue |first1=Fuzhong |last2=Wang |first2=Yi |last3=Xu |first3=Shuhua |last4=Zhang |first4=Feng |last5=Wen |first5=Bo |last6=Wu |first6=Xuesen |last7=Lu |first7=Ming |last8=Deka |first8=Ranjan |last9=Qian |first9=Ji |last10=Jin |first10=L |display-authors=9 |year=2008 |title=A spatial analysis of genetic structure of human populations in China reveals distinct difference between maternal and paternal lineages |journal=European Journal of Human Genetics |volume=16 |issue=6 |pages=705–17 |doi=10.1038/sj.ejhg.5201998 |pmid=18212820 |doi-access=free}}</ref> * Biological research findings have also demonstrated that the paternal lineages Y-DNA O-M119,<ref name="Li et al (2008)">{{cite journal |last1=Li |first1=Hui |year=2008 |title=Paternal genetic affinity between western Austronesians and Daic populations |journal=BMC Evolutionary Biology |volume=8 |issue=1 |page=146 |bibcode=2008BMCEE...8..146L |doi=10.1186/1471-2148-8-146 |pmc=2408594 |pmid=18482451 |doi-access=free}}</ref> O-P201,<ref name="Karafet et al (2010)">{{cite journal |last1=Karafet |first1=Tatiana |last2=Hallmark |first2=B |last3=Cox |first3=M.P. |last4=Sudoyo |first4=H |last5=Downey |first5=S |last6=Lansing |first6=J.S. |last7=Hammer |first7=M.F. |date=August 2010 |title=Major East–West Division Underlies Y Chromosome Stratification across Indonesia |journal=Molecular Biology and Evolution |volume=27 |issue=8 |pages=1833–44 |doi=10.1093/molbev/msq063 |pmid=20207712 |doi-access=free}}</ref> O-P203<ref name="Karafet et al (2010)" /> and O-M95<ref name="Karafet et al 2005">{{cite book |last1=Karafet |first1=Tatiana |last2=Hagberg |first2=L |last3=Hanson |first3=L. A. |last4=Korhonen |first4=T |last5=Leffler |first5=H |last6=Olling |first6=S |year=1981 |chapter=Balinese Y-chromosome perspective on the peopling of Indonesia: genetic contributions from pre-Neolithic hunter-gatherers, Austronesian farmers, and Indian traders |title=Ciba Foundation Symposium 80 - Adhesion and Microorganism Pathogenicity |series=Novartis Foundation Symposia |volume=80 |pages=161–87 |doi=10.1002/9780470720639.ch11 |pmid=6114819|isbn=978-0-470-72063-9 }}</ref> are found in commonly Southern non-Han minorities, less commonly in Southern Han, and even less frequently in Northern Han Chinese.<ref name="Yan et al (2011)">{{cite journal |last1=Yan |first1=Shi |last2=Wang |first2=C.C. |last3=Li |first3=H |last4=Li |first4=S.L. |last5=Jin |first5=L |year=2011 |title=An updated tree of Y-chromosome Haplogroup O and revised phylogenetic positions of mutations P164 and PK4 |journal=European Journal of Human Genetics |volume=19 |issue=9 |pages=1013–15 |doi=10.1038/ejhg.2011.64 |pmc=3179364 |pmid=21505448}}</ref> * Haplogroups O1 and O2 significantly peak in the southeastern coastlines and eastern regions of China respectively, according to one study.<ref>{{cite journal |last1=Li |first1=Jienan |last2=Song |first2=Feng |last3=Lang |first3=Min |last4=Xie |first4=Mingkun |title=Comprehensive insights into the genetic background of Chinese populations using Y chromosome markers |journal=Royal Society Open Science |date=September 2023 |volume=10 |issue=9 |doi=10.1098/rsos.230814 |pmid=37736526 |pmc=10509572 |bibcode=2023RSOS...1030814L }}</ref> Patrilineal DNA indicates the northern Han Chinese were the primary contributors to the paternal gene pool of modern southern Han Chinese.<ref name="AD" /><ref name="table" /><ref name="EJH" /><ref name="Yan et al (2011)" /> The data also indicates that the contribution of southern non-Han aboriginals to the southern Han Chinese genetics is limited. In short, male Han Chinese were the primary drivers of Han Chinese expansion in successive migratory waves from the north into what is now modern southern China as is shown by a greater contribution to the Y-chromosome than the mtDNA from northern to Southern Han.<ref name="Genetic evidence supports demic dif" /> During the Zhou dynasty, or earlier, peoples with haplogroup Q-M120 also contributed to the ethnogenesis of Han Chinese people. This haplogroup is implied to be spread across in the Eurasian steppe and north Asia since it is found among [[Cimmerians]] in [[Moldova]] and Bronze Age natives of [[Khövsgöl Province|Khövsgöl]]. But it is currently near-absent in these regions except for East Asia. In modern China, haplogroup Q-M120 can be found in the northern and eastern regions.<ref>{{cite journal |last1=Sun |first1=Na |last2=Ma |first2=Peng-Cheng |last3=Yan |first3=Shi |last4=Wen |first4=Shao-Qing |last5=Sun |first5=Chang |last6=Du |first6=Pan-Xin |last7=Cheng |first7=Hui-Zhen |last8=Deng |first8=Xiao-Hua |last9=Wang |first9=Chuan-Chao |last10=Wei |first10=Lan-Hai |title=Phylogeography of Y-chromosome haplogroup Q1a1a-M120, a paternal lineage connecting populations in Siberia and East Asia |journal=Annals of Human Biology |date=3 April 2019 |volume=46 |issue=3 |pages=261–266 |doi=10.1080/03014460.2019.1632930 |pmid=31208219 }}</ref> '''Matrilineal DNA''' MtDNA of Han Chinese increases in diversity as one looks from northern to Southern China, which suggests that the influx of male Han Chinese migrants intermarried with the local female non-Han aborigines after arriving in what is now modern-day Fujian, Guangdong, Guangxi, Hainan and other regions of Southern China.<ref name="table" /><ref name="EJH" /> In these populations, the contribution to mtDNA from Han Chinese and indigenous tribes is evenly matched, representing a substantial mtDNA contribution from non-Han groups, collectively known as the Bai Yue or Hundred Yue.<ref name="table" /><ref name="EJH" /> A study by the Chinese Academy of Sciences into the gene frequency data of Han sub-populations and ethnic minorities in China, showed that Han sub-populations in different regions are also genetically quite close to the local ethnic non-Han minorities, meaning that in many cases, the blood of ethnic minorities had mixed into Han genetic substrate through varying degrees of intermarriage, while at the same time, the blood of the Han had also mixed into the genetic substrates of the local ethnic non-Han minorities.<ref>{{cite journal |last1=Du |first1=R |last2=Xiao |first2=C |last3=Cavalli-Sforza |first3=LL |year=1997 |title=Genetic distances between Chinese populations calculated on gene frequencies of 38 loci |journal=Science China Life Sciences |volume=40 |issue=6 |pages=613–21 |doi=10.1007/BF02882691 |pmid=18726285 }}</ref>
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