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Peopling of China

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In the course of the peopling of the World by Homo sapiens, East Asia was reached about 50,000 years ago (50 kya). The "recent African origin" lineage from 70 kya diverged into identifiable East Eurasian and West Eurasian lineages by about 50 kya.[1][2] The East Eurasian ancestors of East Asians used a southern route to reach South and Southeast Asia, along which they rapidly diverged into the ancestors of Ancient Ancestral South Indians (AASI), Papuans, East Asians and Andamanese peoples (such as the Onge).[3] This early East Asian lineage diverged further during the Last Glacial Maximum, spreading northwards from Mainland Southeast Asia where it significantly contributed to the peopling of the Americas via Beringia about 25 kya.[4] After the last ice age, China was cut off from neighboring island groups. The previous phenotypes of early East Asians were either replaced or prevailed among more geographically distant groups.[5][6]

Genetic history

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Further information: Han Chinese § DNA and genetics analysis, Mongoloid, and Proto-Mongoloid
A population genomic PCA graph, showing the substructure of East Asian populations, including Han Chinese (2019)[7]

Overview

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A "Basal-East Asian population" referred to as the East- and Southeast Asian lineage (ESEA); is ancestral to modern East Asians, Southeast Asians, Polynesians, and Siberians, but also ancestral to the Hoabinhian hunter-gatherers of Southeast Asia and the ~40,000 year old Tianyuan lineage found in Northern China. The ESEA lineage descend from an earlier "eastern non-African" (ENA) or "Ancient East Eurasian" meta-population, which used a single southern route to reach South, Southeast Asia, and Oceania, and along which they rapidly diverged into the ancestors of Ancient South Asians (AASI), East/Southeast Asians (ESEA), as well as Australasians. This ESEA lineage later diverged into the Hoabinhian, the Tianyuan, and Ancient East Asian lineages, and expanded northward. There is "a strong correlation with latitude, with diversity decreasing from south to north".[8][3]

Archaeogenetic studies in the Central Plains

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Neolithic Northern China can be divided into 4 periods: the Pre-Peiligang period (before 7000 BC), the Peiligang period (7000–5000 BC), the Yangshao period (5000–3000 BC), and the Longshan period (3000–2000 BC). The first two correlate with the initial development of Neolithic Chinese culture whilst the latter two correlate with the accelerated development of the Chinese civilization.[9]

The progenitors of the Han Chinese were Neolithic Yellow River farmers,[10][11][9] who were predominantly of Ancient Northern East Asian ancestry, with minor Ancient Southern East Asian ancestry. Or they could be modeled as an earlier admixture of a northern inland lineage that branched early from a common East Asian Tianyuan-like lineage (90%) and an equally deep branching coastal group, related to Önge peoples (10%).[12][13] Located at the estuary of the Yellow River, 9500 to 8000 year old individuals from Shandong province have northern East Asian ancestry that is distinct, yet branching, from that found in inland populations, represented by the 8000 year old Yumin individual. This ancestry is also common in present northern East Asians.[13] Early Neolithic Shandong populations also mixed with peoples from Siberia and Inner Mongolia and to a lesser extent, from the south in Fujian.[13][14]

Yangshao populations clustered with northern East Asian populations,[13][9] and contributed to the genetic makeup of contemporary populations in West Liao River, Inner Mongolia and Tibet. Languages spoken by Early to Middle Neolithic Yellow River populations were believed to be the prototype for Sino-Tibetan languages, with Tibeto-Burman and Sinitic splitting between 8000 and 7200 years ago.[13] Among Longshan populations, influence from southern East Asian populations gradually increased, although there is evidence of continuity with previous Yangshao populations. Between the late Neolithic and late Bronze and Iron Age, there is no evidence of additional foreign influence.[13][14] Both Yangshao and Longshan populations also influenced contemporary Shandong populations.[14][13] According to one study, this 'southern' ancestry is closely related to Kra-Dai populations, which peaks in southern Han due to Han conquests of southern China.[15]

Besides the Han Chinese, populations who are closely related to Neolithic Yellow River farmers include Naxi, Yi,[13] Gelao,[16][14] and western Japanese (i.e. Kansai) people.[17] Northern and southern Han also mixed with neighboring minorities in northern and southern China respectively, which is reflected in their matrilineage.[13][18]

Archaeogenetic studies in southern China

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According to Wang et. al (2021), these ancient individuals from southern China play a key role in the ethnogenesis of present-day southern Chinese populations and to an extent, the Austroasiatic and Austronesian diaspora: [19]

  • Archaic individuals from 12,000–10,000 BP:
    • Qihe-3 (奇和) (11,747–11,356 cal BP; Qihe Cave 奇和洞, Zhangping, Fujian) is an Upper Paleolithic individual from the mountainous interior of Fujian, located about 100 km north of present-day Zhangzhou city. Like other late Homo sapiens, Qihe-3 exhibited a long head, large cranial capacity, high narrow face, broad and low/short nasal shape and exhibits features from northern and southern populations in Neolithic China. Another study suggests they have large upper face and nasal height, along with large prominent cheekbones, flat upper face, thin narrow cheeks, a larger head and facial hair. Their features are not necessarily representative of populations living during the Neolithic and Paleolithic transition but rather, reflect the great phenotypic variation that exists.[20][21] Qihe-3 can be modeled as a mixture of coastal Neolithic East Asian ancestry (e.g. Boshan in Neolithic Shandong and Liangdao in Neolithic Fujian) and deeply diverging East Eurasian ancestry (e.g. Longlin, Indus Periphery).
    • Qihe-2, a more recent specimen from a different layer of the same site dating to 8,428-8,359 cal BP, was sequenced and found to be closely related to Iron Age Taiwanese and present-day Austronesians although Qihe-3 displayed similar affinities too.
    • Longlin (隆林) (10,686–10,439 cal BP; Laomocao Cave 老磨槽洞, Longlin Autonomous County, Guangxi, China) is an Upper Paleolithic individual with deeply diverging East Asian ancestry. Despite their uniquely archaic phenotypes, Longlin carried similar levels of archaic human ancestry as Neolithic and present-day East Asians although it is likely that their phenotypes directly derive from hybridization with archaic hominins.[22] Longlin is closely related to the Maludong or Red Deer Cave people and are also closely related to Ikawazu (ca. 2,700 BP; Ikawazu Shell Midden site (伊川津貝塚), Atsumi Peninsula, southern Aichi Prefecture, central Honshu, Japan), a Jōmon individual, than to basal Asian lineages such as Hoabinhian. Longlin, Ikawazu, and coastal Neolithic East Asians from Shandong and Fujian likely all diverged from each other at the same time, with Longlin being located at the basal position on the lineage leading to M71d, sharing a maternal genetic connection with present-day populations from mainland Southeast Asia.
    • Liangdao-2 (~7,600 BP; Liangdao, Fujian),[23] was found to have mostly Qihe-3-related ancestry (82%–90%), as well as a smaller percentage of northern East Asian ancestry (10%–18%) that can be associated with Neolithic Shandong and other northern East Asian sites. As a result, there is more genetic continuity from the early to late Neolithic in Fujian than there is in Guangxi. However, Liangdao-2 lacks the deeply diverging East Eurasian ancestry found in Qihe-3. They also lack Basal Sunda/Australasian ancestry.[19] Cordillerans, who are the least admixed group among the Austronesians of East Asia, are closely related to Liangdao-2.[24] Nonetheless, Late Neolithic Fujian populations, represented by Xitoucun and Tanshishan, are best modeled as a mixture of Dushan-related ancestry (35/54%), northern East Asian ancestry (44/34%), Qihe3-related ancestry (17/8%) and IndusPeriphery-related ancestry (4/3%).[19]
  • Archaic individuals from 9,000–6,000 BP:
    • Dushan (独山) (8,974–8,593 cal BP; Linfeng Town 林逢镇, Tiandong County, Guangxi) is a male individual that can be modeled as a mixture of Longlin-related ancestry (17%) and Qihe-related ancestry (83%). This suggests a mass migration of ancient Fujian populations into Guangxi, where they intermixed with the indigenous inhabitants rather than completely replacing them.[13] Dushan has high affinities with Early Neolithic Shandong and Fujian populations, Mán Bạc populations from Late Neolithic Vietnam and Austroasiatic populations.
    • Baojianshan (宝剑山) (8,335–6,400 cal BP; Baojianshan Cave site (Baojianshan Cave A), Longzhou County, Chongzuo City, Guangxi) can be modeled as a mixture of 72.3% Dushan-related ancestry and 27.7% Hoabinhian-related ancestry.
  • Historical Guangxi populations from 1,500–500 BP:
    • Layi can be described as a mixture of Boshan-related ancestry (22%–27%) and either Longlin-related (78%) or Dushan-related (73%) ancestry.
    • Shenxian can be described a mixture of northern East Asian-related ancestry (9%–22%) and southern East Asian-related ancestry (78%–91%).
    • Yiyang can be modeled as a mixture of northern East Asian (18%–42%), and southern East Asian (Liangdao2, 58%–83%) ancestry. However, Yiyang can also be modeled as a mixture of northern East Asian-related ancestry (27%–42%) and Dushan-related ancestry (58%–73%).
    • LaCen can be described as a mixture of northern East Asian ancestry (22%–30%) and Dushan-related ancestry (70%–78%).
    • BaBanQinCen can be described as a mixture of ancestry related to Dushan (5%–64%), northern East Asians (19%–40%) and southern East Asians (5%–72%). They significantly contributed to the genetic makeup of present Kra-Dai groups in Guangxi. Zhuang and Dong from Congjiang County in Guizhou, China also cluster with BaBanQinCen, along with GaoHuaHua.[25]
    • GaoHuaHua can be described as a mixture of northern East Asian ancestry (Boshan, 34%) and Dushan-related ancestry (66%). They significantly contributed to the genetic makeup of Hmong-Mien groups in Guangxi. Zhuang and Dong from Congjiang County in Guizhou, China also cluster with GaoHuaHua, along with BaBanQinCen.[25]

Minor West Eurasian ancestry in Shaanxi and Liaoning Han Chinese

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Several studies reveal minor West Eurasian-derived admixture among Shaanxi Han Chinese, especially those living in Guanzhong and Shaanbei (2–5%),[26][27] and Liaoning Chinese (~2%).[28] Ancient North Eurasian admixture is more dominant among Shaanxi Han Chinese compared to other Han subgroups.[26] Some Shaanxi Han Chinese carry maternal haplogroup U[26] although there's evidence that some Han Chinese from other provinces carry haplogroups W6, H and R1a1.[26] Northwestern Han also have very minor West Eurasian ancestral components, dating 4,500–1,200/1,300 years ago.[29]

Paternal lineages

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A PCA graph illustrating the genetic makeup of Han Chinese and other Asian populations.[30]
Hypothetical migration patterns of paternal human lineages
Haplogroup O, also known as O-M175, is primarily found among populations in Southeast Asia and China.

Looking at Y-DNA studies, it would seem that East Asian paternal lineages expanded in Asia approximately 50,000 years ago. People bearing genetic markers ancestral such as C, D, N, and O, as well as P (specifically Q), came through the Himalayan mountain range and proceeded to Southeast Asia.[31] Haplogroup C moved to East Asia and Australia, with at least two subclades of the major East Asian branch migrating into the Americas, and with members of Haplogroup C-M38 spreading throughout Wallacea, New Guinea, Melanesia, and Polynesia. Another group of peoples, bearing the Y-DNA Haplogroup D, has left descendants mostly in the Andaman Islands, Tibet, and Japan.[32][33] Haplogroup Q, believed to have arisen in Central Asia or Southern Siberia approximately 17,000 to 22,000 years ago, went north to populate Northern Siberia and the Americas. Some northern Chinese have this genetic marker. Haplogroups N and O, originated in Southern China and by 10,000 years ago went on to populate first Southeast Asia and then from Southeast Asia, left for East Asia. Roughly 12,000 years ago, during the Neolithic period, farmers settled along the Yellow River. Alongside various other lineages including O2‐M122, they initiated the development of agriculture. About 6000 years ago, ancestors of the Tibetans split off from this parent group. About 5,000 years ago, Neolithic Yellow River farmers experienced rapid expansion, with notable gene flow into surrounding populations.[34] This corresponds to the late period (2600-2000 BC) of the Longshan culture in the middle Yellow River area. As the Neolithic population in China reached its peak, the number of settlements increased. In some locations, such as the basin of the Fen River in southern Shanxi, the Yellow River in western Henan (confined by the Zhongtiao Mountains and Xiao Mountains), and the coastal Rizhao plain of southeast Shandong, a few very large (over 200 ha) centers developed. In more open areas, such as the rest of Shandong, the Central Plain (in Henan) and the Wei River basin in Shaanxi, local centers were more numerous, smaller (generally 20 to 60 ha) and fairly evenly spaced. Walls of rammed earth have been found in 20 towns in Shandong, nine in the Central Plain, and one (Taosi) in southern Shanxi, suggesting conflict between polities in these areas.

The expansion and rise of these various settlements could be due to the impetus of the collective benefit of the construction of irrigation works in the late Neolithic:

"Most of the labor to dike and drain an area is associated with digging a ditch and sidecasting the soil to make an earthen dike. To make the culvert and tide gate you can use an old worn-out canoe for the pipe… and use just about any good-sized flat stones you can get your hands on. If you have twice as many people making your dike, you can make twice as many linear feet of dike. Doubling the perimeter of a square dike results in quadrupling the area within the dike."[This quote needs a citation]

The agricultural surplus would have allowed for a rapidly expanding population, which would provide more labor for irrigation. By the time of the establishment of the Xia and Shang dynasties, population estimates were at approximately 13 million people.[35]

Studies of DNA remnants from the Central Plains area of China 3000 years ago show close affinity between that population and those of Northern Han today in both the Y-DNA and mtDNA. Both northern and southern Han show similar Y-DNA genetic structure.[36]

Y-chromosome haplogroup O2-M122 is a common DNA marker in Han Chinese, as it appeared in China in prehistoric times. It is found in more than 50% of Chinese males, with frequencies tending to be high toward the east of the country (30/101 = 29.7% Guangxi Pinghua Han,[37] 13/40 = 32.5% Guangdong Han,[38] 11/30 = 36.7% Lanzhou Han,[39] 26/60 = 43.3% Yunnan Han,[40] 251/565 = 44.4% Zhaotong Han,[41] 15/32 = 46.9% Yili Han,[39] 23/49 = 46.9% Lanzhou Han,[42][43] 32/65 = 49.2% South China Han,[44] 18/35 = 51.4% Meixian Han,[39] 22/42 = 52.4% Northern Han,[45] 43/82 = 52.4% Northern Han,[46] 18/34 = 52.9% Chengdu Han,[39] 154/280 = 55.0% Southern Han,[46] 27/49 = 55.1% Northern Han,[47] 73/129 = 56.6% North China Han,[44] 49/84 = 58.3% Taiwan Han,[38] 35/60 = 58.3% Taiwan Minnan,[48] 99/167 = 59.3% East China Han,[44] 33/55 = 60.0% Fujian Han,[48] 157/258 = 60.9% Taiwan Han,[48] 13/21 = 61.9% Taiwan Han,[47] 189/305 = 62.0% Zibo Han,[41] 23/35 = 65.7% Harbin Han,[39] 29/44 = 65.9% Northern Han,[38] 23/34 = 67.6% Taiwan Hakka,[48] 35/51 = 68.6% Beijing Han[40]).[49][50]

During the Zhou dynasty, or earlier, peoples with haplogroup Q-M120 likewise also contributed to the ethnogenesis of Han Chinese people. This haplogroup is implied to be widespread in the Eurasian steppe and north Asia since it is found among Cimmerians in Moldova and Bronze Age natives of 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.[51] Ancient individuals from the Yangshao culture at the Xiaowu site dating back 7,000 years ago carried the Q-M120 lineage.[52]

Han Chinese are genetically distinguishable from Yamato Japanese and Koreans, and internally the different Han Chinese subgroups are genetically closer to each other than any of them are to Koreans and Japanese. However, some Southern Han Chinese, such as Guangxi Han, are genetically closer to Vietnamese and Dai people than Northern Han. But meanwhile, when compared to Europeans genetics, the Han Chinese, Southeast Asian, Japanese and Koreans are closer to each other than Europeans and South Asians. Genealogical research has indicated extremely similar genetic profiles of a less than 1% total variation in spectrum between these three groups.[53] Some Southern Han Chinese and Northern Han Chinese are closest to each other and show the smallest differences when they are compared to other Asians. Vietnamese Ho Chi Minh City Kinh are close to Xishuangbanna's Dai ethnic minority and Guangdong Han Chinese.[54] Koreans are also relatively close to northern Han Chinese. Japanese are more genetically distant from Koreans than Koreans are from northern Han Chinese. However Buryat Mongols and Qinghai Mongols are further from each other than Japanese and Korean despite both being Mongols.[55] Comparisons between the Y chromosome SNP and MtDNA of modern Northern Han Chinese and 3,000 year old Hengbei ancient samples from China's Central Plains show they are extremely similar to each other and show continuity between ancient Chinese of Hengbei and current Northern Han Chinese while Southern Han Chinese were different from the people of Hengbei. This showed that already 3,000 years ago the current northern Han Chinese genetic structure was already formed.[36]

Maternal lineages

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The mitochondrial-DNA haplogroups of the Han Chinese can be classified into the northern East Asian-dominating haplogroups, including A, C, D, G, M8, M9, and Z, and the southern East Asian-dominating haplogroups, including B, F, M7, N*, and R.[56]

These haplogroups account for 52.7% and 33.85% of those in the Northern Han, respectively.

Haplogroup mtDNA D descend from Dravidian Haplogroup M mtdna is the modal mtDNA haplogroup among northern East Asians. Among these haplogroups, D, B, F, and A were predominant in the Northern Han, with frequencies of 25.77%, 11.54%, 11.54%, and 8.08%, respectively.

However, in the Southern Han, the northern and southern East Asian-dominating mtDNA haplogroups accounted for 35.62% and 51.91%, respectively. The frequencies of haplogroups D, B, F, and A reached 15.68%, 20.85%, 16.29%, and 5.63%, respectively.[57][58][59][60][61]

Climate history

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During the Last Glacial Maximum, 29,000 to 18,000 years ago, northern China was a treeless steppe with areas of permafrost and southern China lost much of its forest cover. The sea level was much lower. Borneo, Indonesia, the Philippines, and the Japanese archipelago may have been accessible by land. With the end of the last ice age, a period of warming occurred lasting from 18,000 to 10,000 years ago. The oceans rose and inundated vast regions leaving little trace of coastal settlements used by these people. We know little about their languages. Their cultures are likely to have been diverse. There are many limestone cave sites in southern China which show human settlements. There is evidence of pottery making. The inhabitants had bone tools, fished, and hunted pigs and deer.[62]

It is believed that the climate in southern China was warmer and wetter south of the Qinling mountains; elephants are known to have inhabited the Yangtze river region. The climate in Northeast China north of present-day Beijing was characterized as a cold steppe environment during this period. The presence of woolly mammoth is well documented.[63]

The climate was also much warmer between 8,000 and 3,500 years ago. In the Shandong region, excavations have found the bones of alligators and elephants.

The development of agriculture about 10,000 years ago, with the domestication of millet in the Yellow River valley region and rice in the Yangtze River valley, may have been associated with accelerated growth in the number and size of settlements and the intensified development of local cultures and languages.[64]

Settlement patterns

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Early settlements in the Chinese Upper Paleolithic were either hunter-gatherer societies, or marine environment-based societies characterized by shell middens.[65] Relatively speaking the land was sparsely populated, as the peoples followed the coastal regions and the river valleys.

Neolithic settlements have been found from Liaoning province in the northeast to the Chengdu region in the southwest; from Gansu province in the northwest to sites in Fujian in the southeast. The settlement pattern in the Tibetan region is still unclear as there is debate as to whether there was a pre-Neolithic population movement into the region.[66]

See also

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References

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