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
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
Li, D. Overview of Environmental Behavior (Tsinghua University Press, 1999).
Wang, Y. The late Paleolithic environmental change and human migration in North China. Acta Anthropol. Sin. 37(3), 11 (2018).
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).
Frachetti, M. D. Multiregional emergence of mobile pastoralism and nonuniform institutional complexity across Eurasia. Curr. Anthropol. 53(1), 2–38 (2012).
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).
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).
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).
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).
Alexiades, M. N. (Ed.). Mobility and Migration in Indigenous Amazonia: Contemporary Ethnoecological Perspectives (Berghahn Books, 2022).
Ren, L. et al. Foraging and farming: archaeobotanical and zooarchaeological evidence for Neolithic exchange on the Tibetan Plateau. Antiquity 94(375), 637–652 (2020).
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).
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).
Holt, R. D. & Gomulkiewicz, R. How does immigration influence local adaptation? A reexamination of a familiar paradigm. Am. Nat. 149(3), 563–572 (1997).
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).
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).
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).
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
Yang, J. et al. Sustainable intensification of millet–pig agriculture in Neolithic North China. Nat. Sustain. 5(9), 780–786 (2022).
Lazaridis, I. et al. Ancient human genomes suggest three ancestral populations for present-day Europeans. Nature 513(7518), 409–413 (2014).
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).
Olalde, I. et al. The Beaker phenomenon and the genomic transformation of northwest Europe. Nature 555(7695), 190–196 (2018).
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).
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).
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).
Ran, M. & Chen, L. The 4.2 ka BP climatic event and its cultural responses. Quat. Int. 521, 158–167 (2019).
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).
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).
Wang, Z. The historical “Go West”. Grand Garden of Sci. 24, 26 (2009).
Ning, C. et al. Ancient genomes from northern China suggest links between subsistence changes and human migration. Nat. Commun. 11, 2700 (2020).
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).
Guo, D. & Zhang, X. Early Chinese Civilization-Northeastern Culture and Youyan Civilization (Hardcover) (Jiangsu Education Press, 2005).
Ni, Y. Exploring the origins of animal mask motifs on Shang and Zhou Dynasty bronze vessels. Study of Art 00, 637–652 (2014).
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).
Qin, X. The production and circulation of turquoise ornaments in the Erlitou culture period. Cultural Relics of Central China 02, 64–74 (2022).
Deng, C. et al. A comparative analysis of ancient turquoise tesserae technology in China and Mesoamerica. Jianghan Archaeol. 04, 114–122 (2022).
Dou, Y. Study of Jade Artifacts in Xiajiadian Lower Culture. (Master’s thesis, Chifeng University, Chifeng, 2024).
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).
Song, R. & Chen, Q. A study of pre-Han dynasty faunal remains in Chifeng. Steppe Cultural Relics 02, 85–101 (2004).
Chen, Q. Animal remains excavated from the Xiliang site, Jinggouzi, Linxi County, Inner Mongolia. Steppe Cultural Relics 02, 105–110 (2006).
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).
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).
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).
Yang, M. A. et al. Ancient DNA indicates human population shifts and admixture in northern and southern China. Science 369(6501), 282–288 (2020).
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).
Cao, J., Sun, J. & Dang, Y. The excavations at the Erdaojingzi Site in Chifeng City, Inner Mongolia. Archaeology 8, 13–26 (2010).
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).
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).
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).
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).
Pääbo, S. Ancient DNA: Extraction, characterization, molecular cloning, and enzymatic amplification. Proc. Natl. Acad. Sci. U. S. A. 86(6), 1939–1943 (1989).
Knapp, M. & Hofreiter, M. Next generation sequencing of ancient DNA: Requirements, strategies and perspectives. Genes 1(2), 227–243 (2010).
Orlando, L. et al. Ancient DNA analysis. Nat. Rev. Methods Primers 1(1), 14 (2021).
Reich, D. Who We are and How We Got Here: Ancient DNA and the New Science of the Human Past (Oxford University Press, 2018).
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).
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).
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).
Britton, K. A stable relationship: Isotopes and bioarchaeology are in it for the long haul. Antiquity 91(358), 853–864 (2017).
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).
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).
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).
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).
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).
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).
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).
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).
Allentoft, M. E. et al. Population genomics of Bronze Age Eurasia. Nature 522, 167–172. https://doi.org/10.1038/nature14507 (2015).
Du, P. et al. Ancient genome of the Chinese Emperor Wu of Northern Zhou. Curr. Biol. 34(7), 1587–1595 (2024).
Du, P. et al. Genomic dynamics of the Lower Yellow River Valley since the Early Neolithic. Curr. Biol. 34(17), 3996–4006 (2024).
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).
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).
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).
Peltzer, A. et al. EAGER: Efficient ancient genome reconstruction. Genome Biol. 17, 60. https://doi.org/10.1186/s13059-016-0918-z (2016).
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).
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).
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).
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).
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).
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).
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).
Peter, B. M. Admixture, population structure, and F-statistics. Genetics 202(4), 1485–1501. https://doi.org/10.1534/genetics.115.183913 (2016).
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).
Chang, C. C. et al. Second-generation PLINK: Rising to the challenge of larger and richer datasets. Gigascience 4(1), s13742–s14015 (2015).
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).
Patterson, N. et al. Ancient admixture in human history. Genetics 192(3), 1065–1093. https://doi.org/10.1534/genetics.112.145037 (2012).
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).
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).
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).
Ambrose, S. H. Preparation and characterization of bone and tooth collagen for isotopic analysis. J. Archaeol. Sci. 17(4), 431–451 (1990).
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).
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).
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).
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).
Keith, D. et al. A new perspective for the study of the origin of the domestic pig. Archaeology 11, 74–80 (2006).
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).
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).
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).
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).
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).
Albarella, U., Dobney, K. & Rowley-Conwy, P. Documenting Domestication: New Genetic and Archaeological Paradigms 209–227 (University of California Press, 2006).
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).
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).
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).
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).
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).
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).
Chen, G. Huainanzi淮南子 (China Publishing House, 2012).
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).
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).
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).
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).
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).
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).
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).
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).
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).
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).
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).
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).
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).
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).
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).
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).
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).
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).
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).
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).
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).
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).
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).
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).
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).
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).
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).
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).
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).
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).
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).
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).
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).
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).
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).
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).
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).
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).
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).
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).
Luo, Y. Pig’s raising and using ritually in Dadianzi site. Res. China’s Front. Archaeol. 00, 288–300 (2009).
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
Authors and Affiliations
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
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
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/.
About this article
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
Received:
Accepted:
Published:
DOI: https://doi.org/10.1038/s41598-026-42242-3


