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Matrilineal origins and genetic structure of Neolithic Shi'aan populations revealed by 172 ancient genomes in the north

Recently, "Ancient Mitogenomes Reveal the Origins and Genetic Structure of the Neolithic Shimao Population in Northern China", a joint project by Fu Qiaomei's team at the Institute of Paleontology, Chinese Academy of Sciences and Shaanxi Provincial Institute of Archaeology, was published in the journalFrontiers in GeneticsThe magazine. Mitochondrial DNA from 172 ancient humans was captured from 13 archaeological sites in northern Shaanxi and southern Shanxi provinces, and the latest DNA evidence reveals the early origin of the Shiyuan population from the Yangshao population. In addition, the closest matrilineal kinship to the Shiyuan among other populations in the ancient Yellow River basin is with the Longshan Taosi population of the same period. In the modern population, the Shihuang crowd is closer to the present-day northern Han Chinese than to the Chinese minority and southern Han Chinese. The present study provides a new perspective on the genesis, structure and demographic dynamics of the Shiyuan people in the middle reaches of the Yellow River during the Neolithic period.

Located in Shenmu County, Yulin, Shaanxi Province, 4300-3800 years ago, the site of Shi Ahn (also known as "Shi Ahn City") is one of the largest city sites of the Longshan Period of the Late Neolithic in northern China, and is a capital city site with the dual nature of a political and religious center. In the northern Shaanxi region, Shihuangcheng has similar cultural characteristics with other neighboring sites of the same period, such as Zhaishan, Xinhua, Shenzhuzhuliang, and Muzhuzhuliang, and is known as the "Shihuang culture". Due to the importance of Shi Ahn city and Shi Ahn culture, the origin of the inhabitants of Shi Ahn and the genetic relationship with other ancient populations, especially those related to Taosi Temple, show many cultural similarities, but the genetic link between the two cultural populations is still unclear. Therefore, the study of population evolution through DNA has become an effective means to solve these problems.

Matrilineal origins and genetic structure of Neolithic Shi'aan populations revealed by 172 ancient genomes in the north

In order to investigate the origin of the Shihuang population and the potential integration and exchange between them and the surrounding areas, especially the pottery temple sites in the southern Jin region, Fu Qiaomei's team has successfully obtained 172 cases of mitochondrial whole genomes from a total of 13 sites in northern Shaanxi (132 cases from 11 sites) and southern Jin (40 cases from 2 sites) during the late Neolithic period from the Yangshao to the Longshan era (4836-3253 years ago) through collaboration. The whole mitochondrial genomes of ancient humans from the Late Neolithic period to the Longshan period (4836-3253 years ago) were successfully obtained. Six individuals with high contamination rates (> 4%) and four individuals with close relatives (defined as identical mtDNA sequences) were removed, resulting in a final dataset of 162 individuals.

I. The early Shihshan crowd was mostly of local origin

To understand the genetic links between the Late Neolithic Shihuang population and the pre-Middle Neolithic population, the researchers collected 21 individuals from the Middle Neolithic Miaoliang and Wuzhuangguo earworm, and 66 individuals from the Late Neolithic of Shihuangcheng (Fig. 1).

Haplogroup analysis revealed that MT:A, C, D, and G show high proportions in ancient and modern northeastern Asian populations, with a current north-south decreasing trend. MT:B, F, and M are common in ancient and modern Southeast Asians, with a north-south increasing trend. (MT: Contraction of mitochondrial DNA haplogroups)

Early stages of the Middle Neolithic of northern Shaanxi Miaoliang and Wuzhuangguo are known to carry MT: A (9.52%), C (4.76%), D (23.81%), G (4.76%), B (9.52%, B4'5), F (14.29%), M (14.29%), Z (4.76%), R# (14.29%), showing a higher proportion of Asian MT type in Northeast Asia. PCA-based analysis showed that PC1 explained the geographic east-to-west population variation, while PC2 explained the north-to-south variation (Figure 2B).

Generally, populations are genetically divided into three clusters: Northeast Asia (NEA), Southeast Asia (SEA), and Central and Western Asia (CWE). preShimao_MW "Miaoliang-Wuzhuang" skews to the Northeast Asia population and clusters with the Yellow River population (Figure 2B). In addition, the genetic distance ofFThe ST heat map also showed that the correlation between the "Miaoliang-Wuzhuang" population in northern Shaanxi and the NEA population in the Yellow River Basin was higher than that in other regions of East Asia. Although the "Miaoliang-Wuzhuang" population is not obviously related to the SD_MN Yellow River population, the DAPC map shows that there is some connection between the "Miaoliang~Wuzhuang" and SD_MN.

Matrilineal origins and genetic structure of Neolithic Shi'aan populations revealed by 172 ancient genomes in the north

(A) Haplogroup frequency. Green haplogroups are common among northeastern Asians, while blue haplogroups are common among Southeast Asians. There are no haplogroups classified as "other" in the Shihuang related groups. Haplogroups R # and N # represent haplotypes shown in East Eurasians (e.g., haplotype R+16189, subhaplogroups R11 and N9, observed in the Shijuan-associated population)

preShimao_MW - Miaoliang, Wuzhuangguo football n=21

Shimao_HCT--Shimao Imperial City Terrace n=10

Shimao_NC--The Inner City of the Stone Mountains n=44

Shimao_DM--The Outer City of the Stone Mountains n=12

MZZSGDL - Wooden pillar column beam, God Ohm beam n=16

XH - Xinhua n=9

ZS - Zhaosan n=10

Matrilineal origins and genetic structure of Neolithic Shi'aan populations revealed by 172 ancient genomes in the north

(B) Haplogroup frequency principal component analysis (PCA). Circles and triangles represent ancient and present-day populations, respectively. The color and shape symbols correspond to Figure 1-A. Gray circles represent ancient populations from the Yellow River Basin (YR), Mongolia or Inner Mongolia (IM), and the Baikal Region (BR) of the NEA. cWE: Central and Western Eurasia.

Matrilineal origins and genetic structure of Neolithic Shi'aan populations revealed by 172 ancient genomes in the north

(C) Heat map of genetic distance ( F ST ) between the population associated with the Stone Mountains and other ancient populations. Different labels and colors correspond to the PCA plots, and different color shades are used to mark different regional populations. values of F ST= 0.00 are white, indicating close kinship. sd_en is excluded from the heat map because the genetic distance is very large (F ST> 0.10) between them and other populations. Gray squares represent ancient populations from the Yellow River Basin (YR), Mongolia and Inner Mongolia (IM), and the Baikal Region (BR) in the Northeast.

Matrilineal origins and genetic structure of Neolithic Shi'aan populations revealed by 172 ancient genomes in the north

Figure 3 (A) Discriminant analysis of principal components (DAPC) of ancient populations in the Yellow River basin. The red dots represent the population related to the Shihuang Mountains. The dark green dots represent TSZJZ (Taosi Zhoujiazhuang) individuals, and the other green dots represent individuals in the "Yellow River".

(B) Haplogroup frequencies in ancient populations. Green haplogroups are common in northeast Asia, while blue-tinged haplogroups are common in Southeast Asia. Haplogroups that are not present in the Shiyuan-related groups are classified as "other". Haplogroups R # and N # represent haplotypes found in East Eurasians (e.g., haplotype R+16189, subhaplotypes R11 and N9, which are also observed in the Shijuan-associated population). Other subhaplogroups R and N found in Western Eurasians are assigned to "other".

(C) Haplogroup sharing analysis. Different colors correspond to F ST heat map and network.

Median linkage network of haplotypes A17 (D) and D4e1a (E) associated with ancient northern Chinese populations. The different population groups are shown in different colors, consistent with those in the F ST heat map.

For the Late Neolithic Longshan population of Shimiao_HCT (Imperial City Terrace), Shimiao_NC (Inner City) and Shimao_DM (Outer City), haplogroup analysis showed similarities.

Matrilineal origins and genetic structure of Neolithic Shi'aan populations revealed by 172 ancient genomes in the north

The Shihuang Mountains population also shows a higher proportion of haplogroups dominated by northeast Asia than by Southeast Asia, and the Outer City (66.67%) shows a higher proportion of haplogroups dominated by northeast Asia than the Royal City Terrace (60.00%) and the Outer City (52.27%) (?). There seems to be an error here in the original article). In addition, the inner city has R# haplogroup (4.55%). The highest percentage of MT-D (27.27-33.33%) is also found in the inner and outer cities, similar to the "Miaoliang-Wuzhuang" and most of the Yellow River populations. PCA shows that the three Shihuang ethnic groups clustered and mapped among the NEA populations in the Yellow River region, which is consistent with the results of haplogroup analysis. Meanwhile, the genetic distance (FST value) between the three Shihuang populations was found to be approximately zero (FST<0.01, P>0.05), indicating that they are closely related (Figure 2C). The same haplotypes (B4a4, C4a2, G2a1, and G1c) were found in these three populations, further suggesting that they are closely related to each other. These results suggest that the populations from different areas of the Late Neolithic Shihuangcheng have close affinities.

In exploring the relationship between the Shihuang Mountains (4148-3253 cal BP period) and the early populations (before 4500 BP) from northern Shaanxi and beyond, it was found that the Shihuang Mountains from the pre-Miaoliang-Wuzhuang period to the late period carried similar MTs: A, C, D, G, Z, B, F, and M. The highest proportion of Ds was found in both the Huangchengtai and the outer city, which was the same as that of the preexisting outer city (Figure 2A).

Some of these types are not found in early populations outside of Shaanxi. Like QT_MN (∼5,500-5,000 BP), SD_EN (∼9,600-7,700 BP) and SD_MN (∼5,500-4,600 BP) lacked C, while SD_EN and SD_MN lacked F (Figure 3B).

The DAPC also shows that the Shihuang people clustered with the "Miaoliang-Wuzhuang" in northern Shaanxi (Figure 3A). In addition, the FST heat map shows that the three Shiyuan populations clustered with "Miaoliang-Wuzhuang" and showed minimal genetic differentiation among them (FST<0.01, P>0.05), while QT_MN, SD_EN and SD_MN showed larger FST values (FST>0.05) with earlier populations outside of northern Shaanxi. 0.05). These results suggest that the Late Neolithic population of Shihuangcheng has the closest genetic relationship with the Middle Neolithic Yangshao population ("Miaoliang-Wuzhuang" 4836-4530 cal BP) from northern Shaanxi. This close relationship also shows the haplotypes shared by both: D4j3, D4b2b, A+152+16362, F1a1c, R11, and are mapped on the same branches of the median linkage network. However some connections were also found between the Shishuang population and the early and middle QT_MN. This is supported by the presence of haplotypes M9a1a1 and M10a1b in the network analysis, which differ by only one and four mutations in these two populations, respectively.

Therefore, the populations in different areas of the Long Shan period in the Stone Mountains (4148-3253 cal BP) are closely related to the Yangshao in northern Shaanxi. The findings show that the Yangshao people of northern Shaanxi were largely not displaced with the establishment of the Shihuang Mountains, which supports the hypothesis that the Shihuang people were mainly of local genetic origin. However, considering the shared haplotypes with other Yellow River populations (i.e., the Middle Neolithic Qingtai), additional genetic contributions from populations outside northern Shaanxi cannot be excluded.

Matrilineal origins and genetic structure of Neolithic Shi'aan populations revealed by 172 ancient genomes in the northMatrilineal origins and genetic structure of Neolithic Shi'aan populations revealed by 172 ancient genomes in the north

Second, the kinship between the stone mount and the ancient surrounding people

Three populations around Shihuangcheng: MZZSGDL (MuZhuZhuLiang, ShenYaoLiang), XH (Xinhua), and ZS (ZhaiShan), mainly carry MT: D (22.22-31.25%), B (6.25-22.22%, B4′5), F (6.25-11.11%), and M (10.00-25.00%). In addition, MZZSGDL also carried haplogroups C (6.25%) and G (18.75%), XH carried haplogroups G (11.11%), Z (11.11%) and R# (11.11%), and ZS had haplogroup A (10.00%) and R# (10.00%). All three populations showed the highest percentage of haplogroup D (31.25% for MZZSGDL, 22.22% for XH and 30.00% for ZS) (Figure 2A; S4).

In the PCA map, the populations adjacent to the Shihuangcheng are represented in the northeastern Asian populations in the Yellow River region and are relatively close to each other (Fig. 2B). The small genetic differences between them also showed this close relationship (FST < 0.01) (Figure 2C; Supplementary Table S6). In the median linkage network, these populations share the same branches in haplotypes Z3, F2g, and M10a1a1b (Supplementary Fig. S2). Thus, these three populations in the neighboring Shihuangcheng are closely related.

In addition, these populations adjacent to the Shihuangcheng were found to show the same haplogroups (D, B, F, and M) as the Shihuangcheng population, with some sharing branching types (A+152+16362, B4a4, D4j3, F2g, and Z3) (Figure 2A, D; S2). This close affinity was also found from the DAPC and FST heat maps, as well as the smaller genetic distance between them (FST=0.00, P>0.05 for most of them, and FST=0.04, P<0.05 between Shimao_NC and MZZSGDL) (Figure 2C, 3A; S6).

Among the older populations, individuals near Shihuangcheng showed the closest relationship with the "Miaoliang-Wuzhuang" in northern Shaanxi, which was similar to the Shihuangcheng population, as evidenced by their common haplogroup, close distribution in DAPC, and small FST values between them (FST=0. 00, P>0.05; FST=0.04, P=0.03) (Fig. 2A, C, 3A; S6). 0.04, P=0.03) (Fig. 2A, C, 3A; S6). Similar to the Shiyuan population, the population surrounding the Shiyuan city (e.g., MZZSGDL) indicated some slight association with the Middle Neolithic Qingtai, supported by the same haplotype M9a1b, and only four mutations were shown to differ between them in the network analysis (Fig. S2). Thus, these results suggest a close genetic affinity between the populations of the Shihuangcheng and its surrounding area.

Most importantly, the populations associated with the Shihuang culture during the Longshan period (including those in and around the Shihuang city, called "Shihuang-related populations") show close affinities with each other, revealing extensive connections between the populations in and around the Shihuang city in northern Shaanxi. All the Shihuang-related populations were shown to have local genetic origins from the Yangshao population of the Middle Neolithic period in northern Shaanxi.

Third, the population of the Shihuang Mountains has a close matrilineal genetic link with the population of the Taosi culture of the same period

Given the genetic relationship between the Yangshao and Longshan periods in northern Shaanxi, the research team then focused on the interaction between populations associated with the Shihuang Mountains and populations outside of northern Shaanxi. Previous archaeological studies have shown that the stone carvings excavated in the Shihuang Mountains share cultural characteristics with the Shang Dynasty of the Central Plains. At the same time, the study of pottery from Shihuangcheng found that the Shihuang culture was closely related to the Taosi culture of the same period. To explore the genetic relationship between the population related to the Stone Mountains and the population from different regions, the research team sequenced 40 new individuals from the proximal part of the Taosi culture, including the Zhoujiazhuang site, and collected 198 previously published ancient individuals from different regions of the Yellow River Basin.

It was found that the population associated with the Shihuang Mountains had more affinities with the population of the Yellow River Basin. Among these Yellow River Basin populations, GQMajiaY_EBA from the Early Bronze Age in the upper Yellow River region, TSZJZ from the middle Yellow River, and SD_LN from the lower Yellow River, i.e., the LN Longshan period (after 4500 BP), carry a higher proportion of haplogroups common to northeastern Asia, such as MT: A (6. 90-10.00%), C (6.00-7.50%), D (22.50-44.83%) and G (4.00-24.14%). Among these haplogroups, D showed the highest percentage among these three Yellow River populations (22.50% for TSZJZ, 26.00% for SD_LN and 44.83% for GQMajiaY_EBA). In addition, they all have clusters B (2.50-22.00%, B4′5), F (6.00-15.00%) and M (3.45-12.50%). All of these haplogroups were found in the population associated with the Stone Mountains.

As seen in the DAPC map, in the Middle Yellow River region, the Shihuang related population is closer to the contemporaneous Longshan population (after 4,500 BP) associated with the Taosi culture than to the Middle Neolithic Yangshao population (5,500-5,000 BP) from the Qingtai Village site in Xingyang (Fig. 3A). In the lower Yellow River region, the Shihuang cultural population is closer to the Late Neolithic Longshan population (SD_LN, after 4,500 BP) than to the EN (SD_EN, ∼9,600-7,700 BP) and MN (SD_MN, ∼5,500-4,600 BP) individuals (Fig. 3A).DAPC indicates that the Shihuang cultural population is closer to the contemporaneous Neolithic The DAPC suggests that the Shihuang Mountains culture population is closer to the contemporaneous Late Neolithic population than to the Early and Middle Neolithic populations outside the northern part of Shaanxi in the Yellow River Basin.

The FST results also show that genetic differentiation between individuals associated with the Shihuang Mountains and contemporaneous Longshan populations of the Late Neolithic period is smaller (FST=0.00, p>0.05 with TSZJZ; FST<0.02, p>0.02 between most Shihuang Mountains culture populations and SD_LN; FST=0. 05, p=0.00 between MZZSGDL and SD_LN) than in the Yellow River region for those early populations (FST=0.04-0.07, p<0.05 with QT_MN; FST=0.04-0.80, p<0.05 with SD_MN; and FST=0.12-0.49, p>0.05 with SD_EN). The haplogroup sharing analysis also showed that the proportion of haplogroups shared by the late Neolithic population of the River Yellow region and the population group of the Shihuang culture was higher than that of the early Neolithic and middle Neolithic (9.80% for TSZJZ compared to 9.27% for QT_MN in the middle YR region; 9.05% for SD_LN compared to 0.98% for SD_EN and 2.79% for SD_MN in the lower Yellow River region). These results also suggest that the Shiyuan cultural population is closer to the late Neolithic population (before 4500 BP) than the early population (after 4500 BP) found elsewhere in the Yellow River.

Among the Late Neolithic populations throughout the Yellow River, the population associated with the Stone Mountains is closest to that associated with the Taosi culture (FST = 0.01 in TSZJZ(p > 0.05). Similarly, DAPC results showed that the Shihuang-related population was clearly clustered with TSZJZ (Taosi Zhoujiazhuang), excluding other LN and BA populations. In addition, haplogroup sharing analysis also showed that the haplotype proportion of Taoji-related individuals was slightly higher (9.80% for TSZJZ; 9.05% for SD_LN) than that of the Shihuang-related population. The network results further indicated that haplotypes A17, C4a1a2, C4a2a1, D4b2b, D4e1a, F1a1c and F2g carried by TSZJZ were also present in the Shihuang-related population, and they shared the same branches. These results suggest that among the populations in other regions of the Yangtze River basin, the Shiyuan-related population is the closest in genetic affinity to the Taosi culture-related population in southern Shanxi Province.

In conclusion, ancient individuals associated with the Shihuang Mountains during the Longshan period in northern Shaanxi have more matrilineal relationships with contemporaneous (but not earlier) populations in the Yellow River region outside of northern Shaanxi. Among these Late Neolithic Longshan populations, the Shishuang-associated populations are most closely related to those associated with the Middle Taosi culture. These results suggest that the Late Neolithic Longshan period populations interacted strongly and extensively not only in northern Shaanxi, but also between northern Shaanxi and southern Jin.

Fourth, the relationship between the people related to the Stone Mountains and modern people

To explore the genetic relationship between the populations associated with the Shihuang Mountains and present-day people, their genetic affinities were compared, including ethnic minorities (such as the Daur, Mongolian, Dai, and Miao), Han populations from the north and south, and populations from Tibet and Taiwan in China.

Among these present-day Chinese populations, the Han population carries NEA-dominated haplogroups A (5.95-6.46%), C (1.79-5.94%), D (19.05-25.84%), and G (1.19-5.94%) and SEA-dominated haplogroups B (11. 37-14.88%, B4′5), F (13.69-13.95% ) and M (18.60-24.40%), and showed the highest proportion of haplogroup D (19.05%-25.84%), consistent with the Shishuang-associated population (Fig. 4A; S5).

Genetic distance analysis also showed that the Shimao-related populations (e.g., Shimao_HCT, XH, ZS) were closer to the Han populations (FST < 0.03, P > 0.06 for NChina_Han; FST < 0.04, P > 0.06 for SChina_Han) than to other present-day minority populations, including those in Tibet and Taiwan (Figure 4B. S6). Haplogroup sharing analysis showed that the haplogroup sharing ratio of the population associated with the Shihuang Mountains was higher than that of other present-day populations (0.00-3.65%) with Han (NChina_Han, 6.04%; SChina_Han, 4.70%) (Fig. 4C; S8). In addition, the populations associated with the Shihuang Mountains (e.g. Shimao_HCT, XH, ZS) were genetically closer to the northern Han population (SChina_Han, FST<0.03, P>0.06) than the southern Han population (SChina_Han, FST>0.03, P>0.03) (Figure 4B; S6).

In addition, haplogroup sharing analysis also showed that the proportion of haplogroups shared between the northern Han population and the Shihuang-related population (NChina_Han, 6.04%) was higher than that of the southern Han population (SChina_Han, 4.70%) (Figure 4C; Table S8). Network analysis also showed that multiple haplotypes (D4g2a1, G1c, and F1a1) were shared between the Shishuang-related and northern Han populations (Figure 4D-F). Therefore, it is concluded that the Shiyuan-related populations are closer to the Han populations in northern China compared to the Chinese minority and southern Han populations.

Matrilineal origins and genetic structure of Neolithic Shi'aan populations revealed by 172 ancient genomes in the north

Figure 4.(A) Haplogroup frequencies. Green haplogroups are common among northeastern Asians, while blue-tinged haplogroups are common among Southeast Asians. Haplogroups that are not present in the Ishuan-related groups are classified as "other". Haplogroups R # and N # represent haplotypes found in East Eurasians (e.g., haplotype R+16189, subhaplogroups R11 and N9, which are also observed in the Shijuan-associated population). Other subhaplogroups R and N found in Western Eurasians are assigned to "other". (B) Genetic distance ( F ST) heat map of Shimao-related and present-day populations. nChina_Minor: minority in northern China; SChina_Minor: minority in southern China, SChina_Tibet: Tibetan population; SChina_Taiwan: Taiwanese population. (C) Haplogroup sharing analysis. Green and blue are used to represent the populations in northern and southern China. Purple represents the ancient population of the Yellow River. The median linkage network of haplotypes D4g2a1 (D), G1c (E) and F1a1c (F) shared by the Shiyuan-related and northern Han populations. The different colored groups correspond to the heat map in Figure 4B.

To explore which ancient populations have the closest genetic proximity to the Han people of northern China, the researchers compared the kinship of ancient individuals from the new Shiyuan-associated populations and other regions of China to the Han people of the north. These include EBA (GQMajiaY_EBA), BA (GQQijia_BA), late BA (GQKayue_LBA), and IA (LTP_IA) individuals from Gansu-Qinghai Province; MN individuals from Henan Province (QT_MN); LN (TSZJZ) individuals from southern Shanxi Province; and EN, MN, and LN individuals from Shandong Province (SD_ EN, SD_MN, SD_LN) in Shandong Province.

Haplogroups A, C, D, G, Z, B (B4′5), F, M, and R# were found to be observed in the Shiyuan-related population, Shandong LN individuals, and NChina_Han, while some of these haplogroups did not occur in other populations (Figs. 2A, 3B, 4A). For example, TSZJZ lacked haplogroup R#, SD_MN lacked haplogroups C, F and R#, and QT_MN lacked haplogroups C, R# and Z (Figure 3B; Table S4).

In addition, most of the populations associated with the Stone Mountains (22.20-33.30%), SD_LN (26.00%) and NChina_Han (25.84%) carried the highest proportion of haplogroup D (Supplementary Tables S4, S5). Haplogroup sharing analysis showed that NChina_Han shared the highest proportion of haplogroups (6.04%) with the Shishuang-related population compared to QT_MN (4.09%), TSZJZ (4.18%) and SD_LN (5.75%) (Figure 4C; Supplementary Table S8). Genetic distance analysis also revealed that NChina_Han had the closest genetic affinity to the Shimao-related population (FST = 0.02, p = 0.10 in XH and ZS; FST = 0. 03, p = 0.06 in Shimao_HCT), while QT_MN (FST = 0.03, p = 0.00); TSZJZ (FST = 0.05, p = 0.00); SD_LN (FST = 0.06, p = 0.00) (Table S6). Thus, the Han population in northern China is more closely related to the population associated with the Shihuang Mountains than to other published ancient Chinese individuals.

The study found that the population in and around the Shihuang Mountains was more closely related to the northern Han population than to the southern Han and minority populations. It was also found that Han groups in northern China were closer to the groups associated with the Shihuang Mountains than to other ancient individuals in China.

Matrilineal origins and genetic structure of Neolithic Shi'aan populations revealed by 172 ancient genomes in the northMatrilineal origins and genetic structure of Neolithic Shi'aan populations revealed by 172 ancient genomes in the northMatrilineal origins and genetic structure of Neolithic Shi'aan populations revealed by 172 ancient genomes in the northMatrilineal origins and genetic structure of Neolithic Shi'aan populations revealed by 172 ancient genomes in the northMatrilineal origins and genetic structure of Neolithic Shi'aan populations revealed by 172 ancient genomes in the northMatrilineal origins and genetic structure of Neolithic Shi'aan populations revealed by 172 ancient genomes in the northMatrilineal origins and genetic structure of Neolithic Shi'aan populations revealed by 172 ancient genomes in the north