Skip to main content

Thank you for visiting nature.com. You are using a browser version with limited support for CSS. To obtain the best experience, we recommend you use a more up to date browser (or turn off compatibility mode in Internet Explorer). In the meantime, to ensure continued support, we are displaying the site without styles and JavaScript.

Advertisement

Scientific Reports
  • View all journals
  • Search
  • My Account Login
  • Content Explore content
  • About the journal
  • Publish with us
  • Sign up for alerts
  • RSS feed
  1. nature
  2. scientific reports
  3. articles
  4. article
Population history and subsistence of farming communities in an agro-pastoral transition zone of northern China: ancient DNA and isotopic evidence from the Erdaojingzi site
Download PDF
Download PDF
  • Article
  • Open access
  • Published: 17 March 2026

Population history and subsistence of farming communities in an agro-pastoral transition zone of northern China: ancient DNA and isotopic evidence from the Erdaojingzi site

  • Xiaohong Lv1,2 na1,
  • Yao Yu3 na1,
  • Lin Ban1,2 na1,
  • Panxin Du4,
  • Jianxue Xiong4,
  • Xin Chang4,
  • Baoshuai Zhang5,
  • Bangyan Wang6,
  • Yiran Xu7,
  • Chunxue Wang1,2,
  • Jianen Cao8 &
  • …
  • Shaoqing Wen4 

Scientific Reports , Article number:  (2026) Cite this article

  • 1284 Accesses

  • 7 Altmetric

  • Metrics details

We are providing an unedited version of this manuscript to give early access to its findings. Before final publication, the manuscript will undergo further editing. Please note there may be errors present which affect the content, and all legal disclaimers apply.

Subjects

  • Ecology
  • Evolution
  • Genetics

Abstract

This study investigates the population history and subsistence of farming communities at the Erdaojingzi site (ca. 3700–3330 cal. BP), a well-preserved Bronze Age settlement located in the agro-pastoral transition zone of the West Liao River Basin in northern China. We apply a multidisciplinary approach, combining ancient DNA, carbon and nitrogen stable isotope analysis, zooarchaeological data and archaeobotanical evidence, to evaluate population affinities at the site and to characterise subsistence practices in relation to environmental and cultural conditions. Ancient DNA from two individuals shows ancestry profiles closely related to farming populations of the Yellow River Basin, rather than to previously published Neolithic groups from the western Liao region, indicating strong genetic connections with Central Plains related farming communities. Stable isotope results from 43 animal and two human bone samples demonstrate that millet and its by-products formed dietary staples for both humans and domestic animals. Humans, pigs and dogs primarily relied on C4 plants, whereas cattle and sheep consumed mixed diets of C3 and C4 plants. Nitrogen isotope values point to a diet rich in animal protein for humans. Combined age at death and isotopic data from pig remains further indicate that young pigs constituted an important component of meat consumption. Variation in nitrogen isotope values among sheep suggests differences in grazing and foddering regimes, and may also reflect localised soil enrichment by dung and other management practices. Taken together, these results provide new insight into how farming communities organised their subsistence and how population history and economic strategies were intertwined in an agro-pastoral transition zone of prehistoric East Asia.

Similar content being viewed by others

Experimental archaeological study in China: implications for reconstruction of past manuring and dietary practices indicated by δ15N values of Setaria italica and Panicum miliaceum

Article Open access 15 February 2024

Localized management of non-indigenous animal domesticates in Northwestern China during the Bronze Age

Article Open access 03 August 2021

Isotopic insights into han period coastal agriculture on the liaodong peninsula in northeast China

Article Open access 13 February 2026

Data availability

The raw alignment file (in BAM format) can be accessed publicly at the Genome Warehouse in the National Genomics Data Center, Beijing Institute of Genomics (part of the China National Center for Bioinformation), under the Chinese Academy of Sciences. The data set will be catalogued under the Accession Number: HRA010456 and can be retrieved from (https://bigd.big.ac.cn/gsa-human/). The other original contributions presented in the study are included in the article. The other original contributions presented in the study are included in the article.

References

  1. Li, D. Overview of Environmental Behavior (Tsinghua University Press, 1999).

    Google Scholar 

  2. Wang, Y. The late Paleolithic environmental change and human migration in North China. Acta Anthropol. Sin. 37(3), 11 (2018).

    Google Scholar 

  3. Anthony, D. W. The Horse, the Wheel, and Language: How Bronze-Age Riders from the Eurasian Steppes Shaped the Modern World (Princeton University Press, 2007).

    Google Scholar 

  4. Frachetti, M. D. Multiregional emergence of mobile pastoralism and nonuniform institutional complexity across Eurasia. Curr. Anthropol. 53(1), 2–38 (2012).

    Google Scholar 

  5. Zhao, Z. The floatation results at Xinglonggou site and its implications for understanding the dry-land farming in North China. Dongya Guwu 1, 188–189 (2004).

    Google Scholar 

  6. Liu, X. Y., Hunt, H. V. & Jones, M. K. River valleys and foothills: Changing archaeological perceptions of North China’s earliest farms. Antiquity https://doi.org/10.1017/S0003598X00098100 (2009).

    Google Scholar 

  7. Harrison, R. J. & Heyd, V. The transformation of Europe in the third millennium BC: The example of ‘Le Petit Chasseur I + III’ (Sion, Valais, Switzerland). Prähist. Z. 82(2), 129–214 (2007).

    Google Scholar 

  8. Jia, X. et al. The transition of human subsistence strategies in relation to climate change during the Bronze Age in the West Liao River Basin, Northeast China. Holocene 26, 781–789 (2016).

    Google Scholar 

  9. Alexiades, M. N. (Ed.). Mobility and Migration in Indigenous Amazonia: Contemporary Ethnoecological Perspectives (Berghahn Books, 2022).

  10. Ren, L. et al. Foraging and farming: archaeobotanical and zooarchaeological evidence for Neolithic exchange on the Tibetan Plateau. Antiquity 94(375), 637–652 (2020).

    Google Scholar 

  11. Liu, M. & Sun, K. A survey study of migration and life adaptation of Tajiks on the Pamir Plateau. Soc. Sci. Xinjiang. 03, 56–62 (2012).

    Google Scholar 

  12. Wang, C. C. et al. Genomic insights into the formation of human populations in East Asia. Nature 591(7850), 413–419. https://doi.org/10.1038/s41586-021-03336-2 (2021).

    Google Scholar 

  13. Holt, R. D. & Gomulkiewicz, R. How does immigration influence local adaptation? A reexamination of a familiar paradigm. Am. Nat. 149(3), 563–572 (1997).

    Google Scholar 

  14. Durkee, L. F., Olazcuaga, L., Melbourne, B. A., & Hufbauer, R. A. Immigration delays but does not prevent adaptation following environmental change: experimental evidence. J. Evolut. Biol. voae031 (2024).

  15. Qu, Y. Understanding mammal resource choices and subsistence strategies during the Holocene Climate Optimum: Integration of evidence from palaeodistribution modelling, animal bones and archaeological remains in the farming-pastoral ecotone, Northern China. J. Archaeol. Sci. 171, 106071 (2024).

    Google Scholar 

  16. Mazzucco, N. et al. Migration, adaptation, innovation: The spread of Neolithic harvesting technologies in the Mediterranean. PLoS ONE 15(4), e0232455. https://doi.org/10.1371/journal.pone.0232455 (2020).

    Google Scholar 

  17. Gao, J. Survival Environments and Adaptation Strategies of Prehistoric Humans in the Yellow River Basin of the Qinghai-Tibetan Plateau (Doctoral dissertation, Qinghai Normal University, 2023). https://doi.org/10.27778/d.cnki.gqhzy.2023.000006

  18. Yang, J. et al. Sustainable intensification of millet–pig agriculture in Neolithic North China. Nat. Sustain. 5(9), 780–786 (2022).

    Google Scholar 

  19. Lazaridis, I. et al. Ancient human genomes suggest three ancestral populations for present-day Europeans. Nature 513(7518), 409–413 (2014).

    Google Scholar 

  20. Stevens, C. J. & Fuller, D. Q. Did Neolithic farming fail? The case for a Bronze Age agricultural revolution in the British Isles. Antiquity 86(333), 707–722 (2012).

    Google Scholar 

  21. Olalde, I. et al. The Beaker phenomenon and the genomic transformation of northwest Europe. Nature 555(7695), 190–196 (2018).

    Google Scholar 

  22. Nørgaard, H. W. Review of: Parker Pearson, M., Sheridan, A., Jay, M., Chamberlain, A., Richards, MP & Evans, J. (2019). The Beaker People. Isotopes, mobility and diet in prehistoric Britain 43, (Prehistoric Society Research Paper, 7) (Oxford: Oxbow. Archaeologische Informationen, 2020).

  23. Jeong, C. et al. A Dynamic 6000-year genetic history of Eurasia’s Eastern Steppe. Cell 183(4), 890-904.e29. https://doi.org/10.1016/j.cell.2020.10.015 (2020).

    Google Scholar 

  24. Dong, Y. et al. The potential of stable carbon and nitrogen isotope analysis of foxtail and broomcorn millets for investigating ancient farming systems. Front. Plant Sci. 13, 1018312 (2022).

    Google Scholar 

  25. Ran, M. & Chen, L. The 4.2 ka BP climatic event and its cultural responses. Quat. Int. 521, 158–167 (2019).

    Google Scholar 

  26. Allen, E. et al. Multidisciplinary lines of evidence reveal East/Northeast Asian origins of agriculturalist/pastoralist residents at a Han dynasty military outpost in ancient Xinjiang. Front. Ecol. Evolut. 10, 932004 (2022).

    Google Scholar 

  27. Chen, G. et al. Multipurpose animal utilization underpinned early Bronze Age subsistence of an oasis in an arid area of northwestern China. Archaeol. Anthropol. Sci. 16(5), 66 (2024).

    Google Scholar 

  28. Wang, Z. The historical “Go West”. Grand Garden of Sci. 24, 26 (2009).

  29. Ning, C. et al. Ancient genomes from northern China suggest links between subsistence changes and human migration. Nat. Commun. 11, 2700 (2020).

    Google Scholar 

  30. Huang, Y. Analysis of the foreign communication of Xiajiadian Lower Culture. J. Chifeng Univ. (Philosophy and Social Science Chinese Edition) 09, 1–5. https://doi.org/10.13398/j.cnki.issn1673-2596.2024.09.002 (2024).

    Google Scholar 

  31. Guo, D. & Zhang, X. Early Chinese Civilization-Northeastern Culture and Youyan Civilization (Hardcover) (Jiangsu Education Press, 2005).

    Google Scholar 

  32. Ni, Y. Exploring the origins of animal mask motifs on Shang and Zhou Dynasty bronze vessels. Study of Art 00, 637–652 (2014).

    Google Scholar 

  33. Wang, P. Preliminary analysis of painted pottery motifs (geometric and zoomorphic designs) unearthed from the Dadianzi Cemetery, Inner Mongolia. Sichuan Cultural Relics 06, 44–48 (2019).

    Google Scholar 

  34. Qin, X. The production and circulation of turquoise ornaments in the Erlitou culture period. Cultural Relics of Central China 02, 64–74 (2022).

    Google Scholar 

  35. Deng, C. et al. A comparative analysis of ancient turquoise tesserae technology in China and Mesoamerica. Jianghan Archaeol. 04, 114–122 (2022).

    Google Scholar 

  36. Dou, Y. Study of Jade Artifacts in Xiajiadian Lower Culture. (Master’s thesis, Chifeng University, Chifeng, 2024).

  37. Xu, Z. Preliminary study on the bone of divination in Xiajiadian lower culture. Wenwuchunqiu 04, 14–18+27. https://doi.org/10.13635/j.cnki.wwcq.2010.04.002 (2010).

    Google Scholar 

  38. Song, R. & Chen, Q. A study of pre-Han dynasty faunal remains in Chifeng. Steppe Cultural Relics 02, 85–101 (2004).

    Google Scholar 

  39. Chen, Q. Animal remains excavated from the Xiliang site, Jinggouzi, Linxi County, Inner Mongolia. Steppe Cultural Relics 02, 105–110 (2006).

    Google Scholar 

  40. Chen, J., Ta, L., Cai, J. & Xiong, Z. Findings and recognition of the Neolithic site of Weijiawopu, Chifeng, Inner Mongolia. Cultural Relics 11, 47–52. https://doi.org/10.13619/j.cnki.cn11-1532/k.2014.11.005 (2014).

    Google Scholar 

  41. Chen, Q. & Zhang, Z. Research on the faunal remains of the Weijiawopu site, 2015~2016. Steppe Cultural Relics 01, 104–114. https://doi.org/10.16327/j.cnki.cn15-1361/k.2017.01.016 (2017).

    Google Scholar 

  42. Wang, Y., Chen, Q., Bao, S. & Wang, L. Study on the faunal remains of the Nailingao site, Inner Mongolia,2015. Steppe Cultural Relics 01, 115–124. https://doi.org/10.16327/j.cnki.cn15-1361/k.2017.01.017 (2017).

    Google Scholar 

  43. Yang, M. A. et al. Ancient DNA indicates human population shifts and admixture in northern and southern China. Science 369(6501), 282–288 (2020).

    Google Scholar 

  44. Cao, J., Dang, Y. & Sun, J. A perfect re-enactment of the Bronze Age “Pompeii in the East” in Inner Mongolia Erdaojingzi site excavation record. China Cultural Herit. 03, 78–87 (2010).

    Google Scholar 

  45. Cao, J., Sun, J. & Dang, Y. The excavations at the Erdaojingzi Site in Chifeng City, Inner Mongolia. Archaeology 8, 13–26 (2010).

    Google Scholar 

  46. Cao, J. et al. Excavation report on workshop area of the Lower Xiajiadian Culture settlement of Erdaojingzi Site,Chifeng City. Steppe Cult. Relics. 02, 12–26. https://doi.org/10.16327/j.cnki.cn15-1361/k.2022.02.003 (2022).

    Google Scholar 

  47. Zhu, K. et al. The genetic diversity in the ancient human population of Upper Xiajiadian culture. J. Syst. Evol. https://doi.org/10.1111/jse.13029 (2023).

    Google Scholar 

  48. Peterson, C. E., Lu, X., Drennan, R. D. & Zhu, D. Hongshan chiefly communities in Neolithic northeastern China. Proc. Natl. Acad. Sci. U. S. A. 107(13), 5756–5761. https://doi.org/10.1073/pnas.1000949107 (2010).

    Google Scholar 

  49. Sun, Y., Zhao, Z., Cao, J., Sun, J. & Dang, Y. Report on the analysis of the 2009 flotation results from the Erdaojingzi site in Inner Mongolia. Agric. Archaeol. 06, 1–9 (2014).

    Google Scholar 

  50. Pääbo, S. Ancient DNA: Extraction, characterization, molecular cloning, and enzymatic amplification. Proc. Natl. Acad. Sci. U. S. A. 86(6), 1939–1943 (1989).

    Google Scholar 

  51. Knapp, M. & Hofreiter, M. Next generation sequencing of ancient DNA: Requirements, strategies and perspectives. Genes 1(2), 227–243 (2010).

    Google Scholar 

  52. Orlando, L. et al. Ancient DNA analysis. Nat. Rev. Methods Primers 1(1), 14 (2021).

    Google Scholar 

  53. Reich, D. Who We are and How We Got Here: Ancient DNA and the New Science of the Human Past (Oxford University Press, 2018).

    Google Scholar 

  54. DeNiro, M. J. & Epstein, S. Influence of diet on the distribution of carbon isotopes in animals. Geochim. Cosmochim. Acta 42, 495–506. https://doi.org/10.1016/0016-7037(78)90199-0 (1978).

    Google Scholar 

  55. Katzenberg, M. A., & Waters‐Rist, A. L. Stable isotope analysis: a tool for studying past diet, demography, and life history. In Biological Anthropology of the Human Skeleton 467–504 (2018).

  56. Reitsema, L. J., Kozłowski, T. & Makowiecki, D. Human–environment interactions in medieval Poland: A perspective from the analysis of faunal stable isotope ratios. J. Archaeol. Sci. 40(10), 3636–3646 (2013).

    Google Scholar 

  57. Britton, K. A stable relationship: Isotopes and bioarchaeology are in it for the long haul. Antiquity 91(358), 853–864 (2017).

    Google Scholar 

  58. Heaton, T. H. E., Vogel, J. C., von la Chevallerie, G. & Collett, G. Climatic influence on the isotopic composition of bone nitrogen. Nature 322, 822–823. https://doi.org/10.1038/322822a0 (1986).

    Google Scholar 

  59. Schwarcz, H. P., Dupras, T. L. & Fairgrieve, S. I. 15N enrichment in the Sahara: In search of a global relationship. J. Archaeol. Sci. 26, 629–636. https://doi.org/10.1006/jasc.1998.0380 (1999).

    Google Scholar 

  60. Fuller, B. T., Fuller, J. L., Harris, D. A. & Hedges, R. E. M. Detection of breastfeeding and weaning in modern human infants with carbon and nitrogen stable isotope ratios. Am. J. Phys. Anthropol. 129, 279–293. https://doi.org/10.1002/ajpa.20249 (2006).

    Google Scholar 

  61. Zhu, K. et al. Cultural and demic co-diffusion of Tubo Empire on Tibetan Plateau. iScience 25, 105636. https://doi.org/10.1016/j.isci.2022.105636 (2022).

    Google Scholar 

  62. Knapp, M., Clarke, A. C., Horsburgh, K. A. & Matisoo-Smith, E. A. Setting the stage—building and working in an ancient DNA laboratory. Ann. Anat. 194, 3–6. https://doi.org/10.1016/j.aanat.2011.03.008 (2012).

    Google Scholar 

  63. Rohland, N., Glocke, I., Aximu-Petri, A. & Meyer, M. Extraction of highly degraded DNA from ancient bones, teeth and sediments for high-throughput sequencing. Nat. Protoc. 13, 2447–2461. https://doi.org/10.1038/s41596-018-0050-5 (2018).

    Google Scholar 

  64. Meyer, M. & Kircher, M. Illumina sequencing library preparation for highly multiplexed target capture and sequencing. Cold Spring Harb. Protoc. https://doi.org/10.1101/pdb.prot5448 (2010).

    Google Scholar 

  65. Gamba, C. et al. Genome flux and stasis in a five millennium transect of European prehistory. Nat. Commun. 5, 5257. https://doi.org/10.1038/ncomms6257 (2014).

    Google Scholar 

  66. Allentoft, M. E. et al. Population genomics of Bronze Age Eurasia. Nature 522, 167–172. https://doi.org/10.1038/nature14507 (2015).

    Google Scholar 

  67. Du, P. et al. Ancient genome of the Chinese Emperor Wu of Northern Zhou. Curr. Biol. 34(7), 1587–1595 (2024).

    Google Scholar 

  68. Du, P. et al. Genomic dynamics of the Lower Yellow River Valley since the Early Neolithic. Curr. Biol. 34(17), 3996–4006 (2024).

    Google Scholar 

  69. Rohland, N. et al. Three assays for in-solution enrichment of ancient human DNA at more than a million SNPs. Genome Res. 32, 2068–2078. https://doi.org/10.1101/gr.276728.122 (2022).

    Google Scholar 

  70. Schubert, M., Lindgreen, S. & Orlando, L. AdapterRemoval v2: Rapid adapter trimming, identification, and read merging. BMC Res. Notes 9, 88. https://doi.org/10.1186/s13104-016-1900-2 (2016).

    Google Scholar 

  71. Li, H. & Durbin, R. Fast and accurate short read alignment with Burrows-Wheeler transform. Bioinformatics (Oxford, England) 25(14), 1754–1760. https://doi.org/10.1093/bioinformatics/btp324 (2009).

    Google Scholar 

  72. Peltzer, A. et al. EAGER: Efficient ancient genome reconstruction. Genome Biol. 17, 60. https://doi.org/10.1186/s13059-016-0918-z (2016).

    Google Scholar 

  73. Renaud, G., Slon, V., Duggan, A. T. & Kelso, J. Schmutzi: Estimation of contamination and endogenous mitochondrial consensus calling for ancient DNA. Genome Biol. 16, 224. https://doi.org/10.1186/s13059-015-0776-0 (2015).

    Google Scholar 

  74. Fu, Q. et al. An early modern human from Romania with a recent Neanderthal ancestor. Nature 524(7564), 216–219. https://doi.org/10.1038/nature14558 (2015).

    Google Scholar 

  75. Weissensteiner, H. et al. HaploGrep 2: Mitochondrial haplogroup classification in the era of high-throughput sequencing. Nucleic Acids Res. 44(W1), W58–W63. https://doi.org/10.1093/nar/gkw233 (2016).

    Google Scholar 

  76. Patterson, N., Price, A. L. & Reich, D. Population structure and eigenanalysis. PLoS Genet. 2(12), e190. https://doi.org/10.1371/journal.pgen.0020190 (2006).

    Google Scholar 

  77. Damgaard, P. B. et al. 137 ancient human genomes from across the Eurasian steppes. Nature 557(7705), 369–374. https://doi.org/10.1038/s41586-018-0094-2 (2018).

    Google Scholar 

  78. Sikora, M. et al. The population history of northeastern Siberia since the Pleistocene. Nature 570(7760), 182–188. https://doi.org/10.1038/s41586-019-1279-z (2019).

    Google Scholar 

  79. Mao, X. et al. The deep population history of northern East Asia from the Late Pleistocene to the Holocene. Cell 184(12), 3256-3266.e13. https://doi.org/10.1016/j.cell.2021.04.040 (2021).

    Google Scholar 

  80. Peter, B. M. Admixture, population structure, and F-statistics. Genetics 202(4), 1485–1501. https://doi.org/10.1534/genetics.115.183913 (2016).

    Google Scholar 

  81. Lawson, D. J., van Dorp, L. & Falush, D. A tutorial on how not to over-interpret STRUCTURE and ADMIXTURE bar plots. Nat. Commun. 9(1), 3258. https://doi.org/10.1038/s41467-018-05257-7 (2018).

    Google Scholar 

  82. Chang, C. C. et al. Second-generation PLINK: Rising to the challenge of larger and richer datasets. Gigascience 4(1), s13742–s14015 (2015).

    Google Scholar 

  83. Alexander, D. H., Novembre, J. & Lange, K. Fast model-based estimation of ancestry in unrelated individuals. Genome Res. 19(9), 1655–1664. https://doi.org/10.1101/gr.094052.109 (2009).

    Google Scholar 

  84. Patterson, N. et al. Ancient admixture in human history. Genetics 192(3), 1065–1093. https://doi.org/10.1534/genetics.112.145037 (2012).

    Google Scholar 

  85. Jay, M. & Richards, M. P. Diet in the iron age cemetery population at Wetwang slack, east Yorkshire, UK: carbon and nitrogen stable isotope evidence. J. Archaeol. Sci. 33, 653–662. https://doi.org/10.1016/j.jas.2005.09.020 (2006).

    Google Scholar 

  86. Haak, W. et al. Massive migration from the steppe was a source for Indo-European languages in Europe. Nature 522(7555), 207–211. https://doi.org/10.1038/nature14317 (2015).

    Google Scholar 

  87. Mathieson, I. et al. Genome-wide patterns of selection in 230 ancient Eurasians. Nature 528(7583), 499–503. https://doi.org/10.1038/nature16152 (2015).

    Google Scholar 

  88. Ambrose, S. H. Preparation and characterization of bone and tooth collagen for isotopic analysis. J. Archaeol. Sci. 17(4), 431–451 (1990).

    Google Scholar 

  89. Ambrose, S. H., Butler, B. M., Hanson, D. B., Hunter-Anderson, R. L. & Krueger, H. W. Stable isotopic analysis of human diet in the Marianas Archipelago, Western Pacific. Am. J. Phys. Anthropol.: Off. Publ. Am. Assoc. Phys. Anthropol. 104(3), 343–361 (1997).

    Google Scholar 

  90. DeNiro, M. J. Postmortem preservation and alteration of in vivo bone collagen isotope ratios in relation to palaeodietary reconstruction. Nature 317, 806–809. https://doi.org/10.1038/317806a0 (1985).

    Google Scholar 

  91. Hu, Y., Ambrose, S. H. & Wang, C. Stable isotopic analysis of human bones from Jiahu site, Henan, China: Implications for the transition to agriculture. J. Archaeol. Sci. 33, 1319–1330. https://doi.org/10.1016/j.jas.2006.01.007 (2006).

    Google Scholar 

  92. Hu, Y., Luan, F., Wang, S., Wang, C. & Richards, M. P. Preliminary attempt to distinguish the domesticated pigs from wild boars by the methods of carbon and nitrogen stable isotope analysis. Sci. China D Earth Sci. 52, 85–92. https://doi.org/10.1007/s11430-008-0151-z (2009).

    Google Scholar 

  93. Keith, D. et al. A new perspective for the study of the origin of the domestic pig. Archaeology 11, 74–80 (2006).

    Google Scholar 

  94. Liu, X., Jones, M. K., Zhao, Z., Liu, G. & O’Connell, T. C. The earliest evidence of millet as a staple crop: New light on neolithic foodways in North China. Am. J. Phys. Anthropol. 149, 283–290. https://doi.org/10.1002/ajpa.22127 (2012).

    Google Scholar 

  95. Lv, X. et al. Pig management strategies in the east Liao River basin from the Bronze Age (c. 2000-256 BC) to the Liaojin dynasties (907-1234 AD): Stable isotope analysis of animals at the Changshan site, Jilin Province, China. Front. Earth Sci. 10, 883481. https://doi.org/10.3389/feart.2022.883481 (2022).

    Google Scholar 

  96. Makarewicz, C. A. Winter is coming: Seasonality of ancient pastoral nomadic practices revealed in the carbon (δ13C) and nitrogen (δ15N) isotopic record of Xiongnu caprines. Archaeol. Anthropol. Sci. 9, 405–418 (2017).

    Google Scholar 

  97. Si, Y., Li, Z., Hu, Y., Yuan, J. & Wang, C. Hydrogen and oxygen stable isotopic analysis of animal bone collagen from Erlitou site, Yanshi Henan Province. Quat. Sci. 34(1), 196–203 (2014).

    Google Scholar 

  98. Hou, L. et al. Stable isotopic analysis of faunal remains from the Cishan site in Wuan Hebei. Cultural Relics South. China 02, 142–149 (2023).

    Google Scholar 

  99. Albarella, U., Dobney, K. & Rowley-Conwy, P. Documenting Domestication: New Genetic and Archaeological Paradigms 209–227 (University of California Press, 2006).

    Google Scholar 

  100. Wang, G., Han, J. & Liu, D. The carbon isotope composition of C 3 herbaceous plants in loess area of northern China. Sci. China D Earth Sci. 46, 1069–1076 (2003).

    Google Scholar 

  101. Liu, X., Martin J. Under one roof: people, crops and animals in Neolithic North China. In Living in the landscape: essays in honour of Graeme Barker 227–234 (McDonald Institute for Archaeological Research, Cambridge, 2014).

  102. Hou, L. A proxy for reconstruction of subsistence economy of the past populations on stable isotopic perspective. Cultural Relics South. China 2, 165–183. https://doi.org/10.3969/j.issn.1004-6275.2019.02.018 (2019).

    Google Scholar 

  103. Ambrose, S. H. & Norr, L. Experimental evidence for the relationship of the carbon isotope ratios of whole diet and dietary protein to those of bone collagen and carbonate. In Molecular Archaeology of Prehistoric Human Bone (Prehistoric Human Bone-Archaeology at the Molecular Level) (eds Lambert, J. B. & Grupe, G.) 1–37 (Springer, Berlin, 1993).

    Google Scholar 

  104. Guan, L. et al. Carbon and nitrogen stable isotope analysis of pig bones from Wanfa Bozi site in Tonghua. Chin. Sci. Bull. 52(14), 1678–1680. https://doi.org/10.3321/j.issn:0023-074x.2007.14.013 (2007) (in Chinese).

    Google Scholar 

  105. O’ Leary, M. H. Carbon isotope fractionation in plants. Phytochemistry 20(4), 553–567. https://doi.org/10.1016/0031-9422(81)85134-5 (1981).

    Google Scholar 

  106. Chen, G. Huainanzi淮南子 (China Publishing House, 2012).

    Google Scholar 

  107. Lv, P. Pigs with White Trotter: Archaeological Studies of Ancient Chinese Domestic Pigs有豕白蹢: 中国古代家猪的考古研究 (Elephant Press: Zhenzhou, 2024).

  108. Wang, X., Shang, X., Bian, H. & Hu, Y. Evaluating the influence of manuring on stable isotope values of foxtail millet: An experimental planting study. Quat. Sci. 06, 1806–1814 (2022).

    Google Scholar 

  109. Hedges, R. E. M. & Reynard, L. M. Nitrogen isotopes and the trophic level of humans in archaeology. J. Archaeol. Sci. 34, 1240–1251. https://doi.org/10.1016/j.jas.2006.10.015 (2007).

    Google Scholar 

  110. Zhang, Q., Sun, Y., Hou, L., Ji, P. & Zhu, Y. Carbon and nitrogen stable isotope analysis of the human and animalbones from the Haminmangha site. Acta Anthropol. Sin. 02, 261–273. https://doi.org/10.16359/j.1000-3193/aas.2021.0006 (2022).

    Google Scholar 

  111. Zhang, X., Liu, G., Wang, M. & Lv, P. Carbon and nitrogen stable isotope analysis of human bones unearthed from Xinglonggou site. Cultural Relics South. China 04, 185–195 (2017).

    Google Scholar 

  112. Hu, Y., Wang, S., Luan, F., Wang, C. & Richards, M. P. Stable isotope analysis of humans from Xiaojingshan site: Implications for understanding the origin of millet agriculture in China. J. Archaeol. Sci. 35(11), 2960–2965 (2008).

    Google Scholar 

  113. Barton, L. et al. Agricultural origins and the isotopic identity of domestication in northern China. Proc. Natl. Acad. Sci. U. S. A. 106(14), 5523–5528. https://doi.org/10.1073/pnas.0809960106 (2009).

    Google Scholar 

  114. Dai, L. et al. An isotopic perspective on animal husbandry at the Xinzhai site during the initial stage of the legendary Xia Dynasty (2070–1600 BC). Int. J. Osteoarchaeol. 26(5), 885–896. https://doi.org/10.1002/oa.2503 (2016).

    Google Scholar 

  115. Maes, D. G., Dewulf, J., Piñeiro, C., Edwards, S. & Kyriazakis, I. A critical reflection on intensive pork production with an emphasis on animal health and welfare. J. Anim. Sci. 98(Supplement_1), S15–S26 (2020).

    Google Scholar 

  116. Blavi, L. et al. Management and feeding strategies in early life to increase piglet performance and welfare around weaning: A review. Animals 11(2), 302 (2021).

    Google Scholar 

  117. Tao, L. et al. Ancient genomes reveal millet farming-related demic diffusion from the Yellow River into southwest China. Curr. Biol. 33(22), 4995-5002.e7. https://doi.org/10.1016/j.cub.2023.09.055 (2023).

    Google Scholar 

  118. Wang, H. et al. Human genetic history on the Tibetan Plateau in the past 5100 years. Sci. Adv. 9(11), eadd5582. https://doi.org/10.1126/sciadv.add5582 (2023).

    Google Scholar 

  119. Xiong, J. et al. Inferring the demographic history of Hexi Corridor over the past two millennia from ancient genomes. Sci. Bull. 69(5), 606–611. https://doi.org/10.1016/j.scib.2023.12.031 (2024).

    Google Scholar 

  120. Ma, M. et al. Forager-farmer transition at the crossroads of East and Southeast Asia 4900 years ago. Science bulletin 69(1), 103–113. https://doi.org/10.1016/j.scib.2023.10.015 (2024).

    Google Scholar 

  121. Ping, W., Xiong, J. & Fu, Q. Ancient DNA elucidates the migration and evolutionary history of northern and southern populations in East Asia. Hereditas (Beijing) https://doi.org/10.16288/j.yczz.24-224 (2024).

    Google Scholar 

  122. Zhang, F. et al. Ancient genome analyses shed light on the origin and kinship among humans of a cliff tomb from southwestern China. J. Archaeol. Sci. Rep. https://doi.org/10.1016/j.jasrep.2023.104333 (2024).

    Google Scholar 

  123. Yang, J. Zooarchaeological Research on the Erlitou Site in Yanshi, Henan Province. (Master’s thesis, Graduate School of the Chinese Academy of Social Sciences). Beijing. (2006).

  124. Wang, Q. et al. A preliminary study of prehistoric subsistence economy in the southern part of Northern Shaanxi: Zooarchaeological evidence from field surveys in the upper Beiluohe River valley. Quat. Sci. 42(6), 1709–1722 (2022).

    Google Scholar 

  125. Hu, S., Yang, M., Sun, Z. & Shao, J. A study of faunal remains unearthed at Shenmu Shiqian Site, Shaanxi, China, 2012–2013. Archaeol. Cultural Relics 04, 109–121 (2016).

    Google Scholar 

  126. Sheng, P. et al. Feeding shimao: archaeobotanical and isotopic investigation into early urbanism (4200–3000 BP) on the northern Loess Plateau China. Environ. Archaeol. 29(5), 425–439 (2024).

    Google Scholar 

  127. Wang, Q. Q. Probing into palaeoenvironment and means of livelihood reflected by remains of animals and plants in Jinchankou site. J. Qinghai Normal Uni 6, 75–78 (2014).

    Google Scholar 

  128. Wang, L. The economic pattern and environmental context of Lower Xiajiadian Culture reflected by the excavation data of Dashanqian Site. Res. China’s Front. Archaeol. 00, 350–357 (2007).

    Google Scholar 

  129. Liu, C. & Fang, Y. M. Analysis of plant remains unearthed at Wadian Site, Yuzhou, Henan, China. Cultural Relics South. China 04, 55–64+47 (2010).

    Google Scholar 

  130. Guo, X. Late Longshan period livelihoods in northern Shaanxi—the botanical and faunal remains of the Mujuzhuliang and Shenyangyang sites as examples. Agric. Archaeol. 03, 19–23 (2017).

    Google Scholar 

  131. Wang, S., Wang, Z. & Zhu, Y. The Ecological and Climatic Environments of Lower Xiajiadian Culture Reflectedfrom the First Locality of Dashanqian in Chifeng City, Inner Mongolia. Huaxia Archaeol. 03, 44–51 (2004).

    Google Scholar 

  132. Chen, T., Jia, X., Li, H. & Dong, G. The analysis of spatiotemporal transformations of agricultural and its influence factors during Qijia culture period in GansuQinghai region. Quat. Sci. 39(1), 132–143 (2019).

    Google Scholar 

  133. Yang Y. The Analysis of Charred Plant Seeds at Jinchankou Site and Lijiaping Site During Qijia Culture Period in the Hehuang. Master’s thesis. (Lanzhou Universty. Gansu, 2014).

  134. Zhang, Q. et al. Stable isotope analysis of the human remains from the ‘Sacrificial Pit’ at the Dashangian Site of the Upper Xiajiadian Culture in Chifeng, Inner Mongolia. Archaeol. Cultural Relics 04, 107–110 (2015).

    Google Scholar 

  135. Chen, X. et al. The subsistence patterns of the Shengedaliang site(~4,000yrBP)revealed by stable carbon and nitrogen isotopes in Northern Shaanxi, China. Sci. China Earth Sci. 60, 268–276. https://doi.org/10.1007/s11430-016-5123-8 (2017).

    Google Scholar 

  136. Chen, X., Guo, X., Hu, Y., Wang, W. & Wang, C. Analysis of the Subsistence Practice at the Muzhuzhuliang Site in Shenmu Shaanxi. Archaeol. Cultural Relics 5, 6 (2015).

    Google Scholar 

  137. Zhang, X., Ye, M. & Qiu, S. A preliminary discussion on the food of the ancestors at the Lajia site—carbon and nitrogen stable isotope analysis of human bones excavated from the Lajia site disaster site. Cultural Relics in South. China 04, 197–202 (2016).

    Google Scholar 

  138. Ma, M. et al. Early emergence and development of pastoralism in Gan-Qing region from the perspective of isotopes. Archaeol. Anthropol. Sci. 13(6), 93 (2021).

    Google Scholar 

  139. Zhou, L. A study of the tombs and society of the Henan Longshan culture by means of stable carbon and nitrogen isotope analysis. Huaxia Archaeol. 03, 145–152. https://doi.org/10.16143/j.cnki.1001-9928.2017.03.012 (2017).

    Google Scholar 

  140. Chen, X. L. et al. Isotopic reconstruction of the Late Longshan period (ca. 4200–3900 BP) dietary complexity before the onset of state‐level societies at the Wadian Site in the Ying River Valley, Central Plains, China. Int. J. Osteoarchaeol. 26(5), 808–817 (2016).

    Google Scholar 

  141. Chen, X. et al. A preliminary exploration to the domestic animal raising strategy: The evidences from carbon and nitrogen isotope analyses. Archaeology 9, 75–82 (2012).

    Google Scholar 

  142. Zhang, X. & Zhao, C. Carbon and nitrogen stable isotope analysis of some animal bones unearthed from the Xinzhai site. Cult. Relics South. China. 232–240 (2015).

  143. Wu, X., Xiao, H., Wei, C., Pan, Y., Huang, Y., Zhao, Q., Xu, X. & Nives, O. Food structure and agricultural pattern of humans and pigs at the Xinzhai site and stable isotopes of domestic pig domestication. Archaeomerty. 49–57 (2007).

  144. Zhang, Q., Eng, J. T., Wang, L. & Ta, L. Paleodiet studies using stable carbon isotopes from human bone sample: Example from Jinggouzi cemetery, Inner Mongolia. Res. China’s Front. Archaeol. 07, 322–327 (2008).

    Google Scholar 

  145. Deng, Z., Zhang, H., Li, W., Liang, F. & Cao, Y. A preliminary study of early agriculture practices at the Haojiatai site in Luohe city, Henan Province. Sci. China. Earth Sci. 64, 307–317 (2021).

    Google Scholar 

  146. Li, W. et al. Interdisciplinary study on dietary complexity in Central China during the Longshan Period (4.5–3.8 kaBP): New isotopic evidence from Wadian and Haojiatai Henan Province. The Holocene 31(2), 258–270 (2021).

    Google Scholar 

  147. Luo, Y. Pig’s raising and using ritually in Dadianzi site. Res. China’s Front. Archaeol. 00, 288–300 (2009).

    Google Scholar 

Download references

Funding

This work was funded by Archaeological Talent Promotion Program of China (No.2025-197 & No.2024-280), Major Project of the Key Research Base for Philosophy and Social Sciences of the Ministry of Education (No.22JJD780009), The Fundamental Research Funds for the Central Universities (No.2022CXTD17), Jilin University “Chinese-style Modernization and New Forms of Human Civilization” Philosophy and Social Sciences Innovation Team Project (No.2023QNTD12), The National Social Science Fund Special Project (No.23VMZ003), Key Projects of the National Social Science Foundation of China (No.23AKG009), Lantai Young Scholars Program of Chinese History Institute (2022LTQN602), theNational Social Science Fund of China (19VJX074), the National Key Research and Development Program (2020YFE0201600), Major Special Project of Philosophy and Social Sciences Research of the Ministry of Education (2022JZDZ023), the Key Research and Development Program Project of Karamay (2025BA0092), The Doctoral Students’ Scientific Research Innovation Ability Enhancement Project in Jilin University (No.2024KC012; No.2024KC011) and Program for Enhancing Doctoral Research Innovation Capacity in Jilin Province (JJKH20250043BS; JJKH20250044BS).

Author information

Author notes
  1. Xiaohong Lv, Yao Yu and Lin Ban are co-first authors.

Authors and Affiliations

  1. Research Center for Chinese Frontier Archaeology, Jilin University, Changchun, China

    Xiaohong Lv, Lin Ban & Chunxue Wang

  2. Bioarchaeology Laboratory, School of Archaeology, Jilin University, Changchun, China

    Xiaohong Lv, Lin Ban & Chunxue Wang

  3. Department of History, Fudan University, Shanghai, China

    Yao Yu

  4. Institute of Archaeological Science, Fudan University, Shanghai, China

    Panxin Du, Jianxue Xiong, Xin Chang & Shaoqing Wen

  5. USTC Archaeometry Laboratory, University of Science and Technology of China, Hefei, China

    Baoshuai Zhang

  6. Human Phenome Institute, Fudan University, Shanghai, China

    Bangyan Wang

  7. School of Life Sciences, Fudan University, Shanghai, 200433, China

    Yiran Xu

  8. Inner Mongolia Normal University, Hohhot, China

    Jianen Cao

Authors
  1. Xiaohong Lv
    View author publications

    Search author on:PubMed Google Scholar

  2. Yao Yu
    View author publications

    Search author on:PubMed Google Scholar

  3. Lin Ban
    View author publications

    Search author on:PubMed Google Scholar

  4. Panxin Du
    View author publications

    Search author on:PubMed Google Scholar

  5. Jianxue Xiong
    View author publications

    Search author on:PubMed Google Scholar

  6. Xin Chang
    View author publications

    Search author on:PubMed Google Scholar

  7. Baoshuai Zhang
    View author publications

    Search author on:PubMed Google Scholar

  8. Bangyan Wang
    View author publications

    Search author on:PubMed Google Scholar

  9. Yiran Xu
    View author publications

    Search author on:PubMed Google Scholar

  10. Chunxue Wang
    View author publications

    Search author on:PubMed Google Scholar

  11. Jianen Cao
    View author publications

    Search author on:PubMed Google Scholar

  12. Shaoqing Wen
    View author publications

    Search author on:PubMed Google Scholar

Contributions

Xiaohong Lv: Formal analysis, methodology, data curation, visualization, writing—original draft, writing—review & editing. Yao Yu: Formal analysis, methodology, data curation, visualization, writing—original draft, writing—review & editing. Lin Ban: Formal analysis, methodology, data curation, visualization, writing—original draft, writing—review & editing. Panxin Du: Formal analysis, data curation, visualization, supervision, writing—review & editing. Jianxue Xiong Validation, Xin Chang: Validation. Baoshuai Zhang: Validation. Bangyan Wang: Validation. Yiran Xu: Validation. Chunxue Wang: Conceptualization, Data curation, Formal analysis, Funding acquisition, Investigation, Methodology, Project administration, Resources, Supervision, Validation, Writing—original draft, Writing—review & editing. Jianen Cao: Investigation, Methodology, Resources, Writing—review & editing. Shaoqing Wen: Conceptualization, Formal analysis, Methodology, Visualization, Validation, Writing—review & editing.

Corresponding authors

Correspondence to Chunxue Wang, Jianen Cao or Shaoqing Wen.

Ethics declarations

Competing interests

The authors declare no competing interests.

Additional information

Publisher’s note

Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.

Supplementary Information

Supplementary Information 1. (download PDF )

Supplementary Information 2. (download PDF )

Supplementary Information 3. (download PDF )

Supplementary Information 4. (download XLSX )

Supplementary Information 5. (download XLSX )

Supplementary Information 6. (download XLSX )

Supplementary Information 7. (download XLSX )

Rights and permissions

Open Access This article is licensed under a Creative Commons Attribution-NonCommercial-NoDerivatives 4.0 International License, which permits any non-commercial use, sharing, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons licence, and indicate if you modified the licensed material. You do not have permission under this licence to share adapted material derived from this article or parts of it. The images or other third party material in this article are included in the article’s Creative Commons licence, unless indicated otherwise in a credit line to the material. If material is not included in the article’s Creative Commons licence and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of this licence, visit http://creativecommons.org/licenses/by-nc-nd/4.0/.

Reprints and permissions

About this article

Check for updates. Verify currency and authenticity via CrossMark

Cite this article

Lv, X., Yu, Y., Ban, L. et al. Population history and subsistence of farming communities in an agro-pastoral transition zone of northern China: ancient DNA and isotopic evidence from the Erdaojingzi site. Sci Rep (2026). https://doi.org/10.1038/s41598-026-42242-3

Download citation

  • Received: 18 November 2025

  • Accepted: 25 February 2026

  • Published: 17 March 2026

  • DOI: https://doi.org/10.1038/s41598-026-42242-3

Share this article

Anyone you share the following link with will be able to read this content:

Sorry, a shareable link is not currently available for this article.

Provided by the Springer Nature SharedIt content-sharing initiative

Keywords

  • Farming communities
  • Subsistence
  • Agro-pastoral transition zone
  • Erdaojingzi site
  • Ancient DNA
  • Stable isotopes
Download PDF

Advertisement

Explore content

  • Research articles
  • News & Comment
  • Collections
  • Subjects
  • Follow us on Facebook
  • Follow us on X
  • Sign up for alerts
  • RSS feed

About the journal

  • About Scientific Reports
  • Contact
  • Journal policies
  • Guide to referees
  • Calls for Papers
  • Editor's Choice
  • Journal highlights
  • Open Access Fees and Funding

Publish with us

  • For authors
  • Language editing services
  • Open access funding
  • Submit manuscript

Search

Advanced search

Quick links

  • Explore articles by subject
  • Find a job
  • Guide to authors
  • Editorial policies

Scientific Reports (Sci Rep)

ISSN 2045-2322 (online)

nature.com footer links

About Nature Portfolio

  • About us
  • Press releases
  • Press office
  • Contact us

Discover content

  • Journals A-Z
  • Articles by subject
  • protocols.io
  • Nature Index

Publishing policies

  • Nature portfolio policies
  • Open access

Author & Researcher services

  • Reprints & permissions
  • Research data
  • Language editing
  • Scientific editing
  • Nature Masterclasses
  • Research Solutions

Libraries & institutions

  • Librarian service & tools
  • Librarian portal
  • Open research
  • Recommend to library

Advertising & partnerships

  • Advertising
  • Partnerships & Services
  • Media kits
  • Branded content

Professional development

  • Nature Awards
  • Nature Careers
  • Nature Conferences

Regional websites

  • Nature Africa
  • Nature China
  • Nature India
  • Nature Japan
  • Nature Middle East
  • Privacy Policy
  • Use of cookies
  • Legal notice
  • Accessibility statement
  • Terms & Conditions
  • Your US state privacy rights
Springer Nature

© 2026 Springer Nature Limited

Nature Briefing Anthropocene

Sign up for the Nature Briefing: Anthropocene newsletter — what matters in anthropocene research, free to your inbox weekly.

Get the most important science stories of the day, free in your inbox. Sign up for Nature Briefing: Anthropocene