The interaction between prehistoric humans and plant resources is key to understanding the origin and spread of agriculture. As a pivotal region for the transmission of East Asian continental cultures to the islands of Southeast Asia, the trajectory of rice (Oryza sativa) cultivation along the South China coast continues to be debated, largely due to the lack of direct evidence. This study employs micro-CT scanning to analyze Neolithic pottery sherds from the Hengyu site (6194–5582 cal a BP) in the lower Min River region of Fujian province. The findings reveal plant impressions composed primarily of rice chaff, accompanied by small quantities of minute weed seeds. The proportions of non-shattering rice spikelet bases (38%) and pedicels (33%) are significantly lower than those recorded in the contemporaneous lower Yangtze region. This indicates a lower degree of domestication and a delayed development of rice cultivation in this area. The findings suggest that within a mixed subsistence economy combining a broad-spectrum subsistence strategy with agriculture, the lag in rice domestication may have stemmed from the combined effects of low population pressure, a mobile lifestyle, and extensive rice management strategies. This pattern reflects a distinct prehistoric trajectory of rice cultivation in the coastal regions of South China. The study offers new evidence of localized agricultural adaptations in these areas and underscores the considerable potential of micro-CT scanning for advancing archaeobotanical research.
AbboSLev-YadunSGopherA (2010) Agricultural origins: Centers and noncenters; a near eastern reappraisal. Critical Reviews in Plant Sciences29(5): 317–328.
2.
AnJKirleisWZhaoC, et al. (2022) Understanding crop processing and its social meaning in the Xinzhai period (1850–1750 cal BCE): A case study on the Xinzhai site, China. Vegetation History and Archaeobotany31(3): 261–277.
3.
AnTZhangZZhengY, et al. (2024) MicroCT analysis reveals insights into the beginning of rice domestication in the lower Yangtze during the 10th millennium BP. Holocene34(11): 1733–1740.
4.
AnTZhangZZhengY, et al. (2025) Cultivating diversity at the onset of agriculture: Insights from the lower Yangtze in the 10th millennium BP. Vegetation History and Archaeobotany34(4): 489–500.
5.
Arranz-OtaeguiAColledgeSZapataL, et al. (2016) Regional diversity on the timing for the initial appearance of cereal cultivation and domestication in southwest Asia. Proceedings of the National Academy of Sciences113(49): 14001–14006.
6.
BarronADatanIBellwoodP, et al. (2020a) Sherds as archaeobotanical assemblages: Gua Sireh reconsidered. Antiquity94(377): 1325–1336.
7.
BarronAFullerDQStevensC, et al. (2020b) Snapshots in time: MicroCT scanning of pottery sherds determines early domestication of Sorghum (Sorghum bicolor) in East Africa. Journal of Archaeological Science123: 105259.
8.
BarronATurnerMBeechingL, et al. (2017) MicroCT reveals domesticated rice (Oryza sativa) within pottery sherds from early Neolithic sites (4150–3265 cal BP) in southeast Asia. Scientific Reports7(1): 7410.
9.
Bronk RamseyC (2009) Bayesian analysis of radiocarbon dates. Radiocarbon51(1): 337–360.
10.
DaiJCaiXJinJ, et al. (2021) Earliest arrival of millet in the south China coast dating back to 5,500 years ago. Journal of Archaeological Science129: 105356.
11.
Dal MartelloRWangYVMir MakhamadB, et al. (2025) Contrasting diachronic regional trends in cereal grain evolution across Eurasia: A metadata analysis of linear morphometrics from the ninth millennium BCE to today. Philosophical Transactions of the Royal Society B: Biological Sciences380(1926): 20240193.
12.
DengZHungH-CFanX, et al. (2018) The ancient dispersal of millets in southern China: New archaeological evidence. Holocene28(1): 34–43.
13.
DzhanfezovaT (2021) Exploring the broad spectrum: Vegetal inclusions in early Neolithic eastern Balkan pottery. Open Archaeology7(1): 1138–1159.
14.
EndoEShodaSFrachettiM, et al. (2023) Pottery impressions reveal earlier westward dispersal of foxtail millet in inner Asian mountain corridor. Agronomy13(7): 1706.
15.
EnyuanWYingfeiXJieC, et al. (2024) Transformation of plant-based tempers (Grog): Evolution and techniques in Songze, Liangzhu, and Guangfulin pottery. Journal of the Australian Ceramic Society60(1): 305–321.
16.
FuLGuanMYangJ, et al. (2021) The 2018 survey of Niubishan site and Shangdongping site in Pucheng, Fujian province. Southern Cultures37(01): 55–62.
17.
FullerDQAllabyRGStevensC (2010) Domestication as innovation: The entanglement of techniques, technology and chance in the domestication of cereal crops. World Archaeology42(1): 13–28.
18.
FullerDQBarronAChampionL, et al. (2021) Transition from wild to domesticated pearl millet (Pennisetum glaucum) revealed in ceramic temper at three Middle Holocene sites in northern Mali. African Archaeological Review38(2): 211–230.
19.
FullerDQDenhamTAllabyR (2023) Plant domestication and agricultural ecologies. Current Biology33(11): R636–R649.
20.
FullerDQQinLZhengY, et al. (2009) The domestication process and domestication rate in rice: Spikelet bases from the lower Yangtze. Science323(5921): 1607–1610.
21.
GaoY (2012) A study of plant remains and the Neolithic Subsistence in Taihu Lake Region. MSc Thesis, Peking University, China.
22.
GaoY (2024a) Hunting and fishing-led economy and early civilization: The case of the Hongshan culture in western Liaoning. Asian Archaeology8(1): 113–122.
23.
GaoY (2024b) A research on pottery impressions from Xiying site and Rongshan site of Fujian province, China. Kumamoto University, Studies in Social and Cultural Sciences22: 57–70.
24.
GaucherelCEvinA (2025) Trends, stasis and trajectories for plant and animal domestications: Possibilistic models alert on resource overexploitation. Philosophical Transactions of the Royal Society B: Biological Sciences380(1926): 20240189.
25.
GeYLuHWangC, et al. (2022) Phytoliths in spikelets of selected Oryzoideae species: New findings from in situ observation. Archaeological and Anthropological Sciences14(4): 73.
26.
GeYWangCHuanX, et al. (2023) Micro-CT-assisted determination of the shattering and non-shattering phenotypes of charred rice spikelet bases from archaeological sites: A new methodological approach. Archaeometry65(6): 1313–1321.
27.
HammerK (1984) Das Domestikationssyndrom. Die Kulturpflanze32(1): 11–34.
28.
HarlanJRDe WetJMJPriceEG (1973) Comparative evolution of cereals. Evolution27(2): 311–325.
29.
HarveyELFullerDQ (2005) Investigating crop processing using phytolith analysis: The example of rice and millets. Journal of Archaeological Science32(5): 739–752.
30.
HeKWangYZhengY, et al. (2024) Geographic mosaics of rice domestication in the lower Yangtze river indicated by morphological characteristics of rice bulliform phytoliths. Archaeological and Anthropological Sciences16(2): 28.
31.
HuanXLuHJiangL, et al. (2021) Spatial and temporal pattern of rice domestication during the Early Holocene in the lower Yangtze region, China. Holocene31(9): 1366–1375.
32.
IshikawaRCastilloCCHtunTM, et al. (2022) A stepwise route to domesticate rice by controlling seed shattering and panicle shape. Proceedings of the National Academy of Sciences119(26): e2121692119.
33.
JinJHuangYZhangY, et al. (2019) Optically stimulated luminescence ages of the Hengyu site and their implications for anthropogenic activities in the Fuzhou basin, south China. Journal of Quaternary Science34(7): 548–557.
34.
JonesG (1987) A statistical approach to the archaeological identification of crop processing. Journal of Archaeological Science14(3): 311–323.
35.
KislevME (1989) Pre-domesticated cereal in the pre-pottery Neolithic A. In: H I (ed.) People and Culture in Change. British Archaeological Reports International Series 508. Oxford, UK: British Archaeological Reports, pp.147–151.
36.
KistlerLMaezumiSYGregoriodeSouzaJ, et al. (2018) Multiproxy evidence highlights a complex evolutionary legacy of maize in south America. Science362(6420): 1309–1313.
37.
Lev-YadunSGopherAAbboS (2000) Archaeology. The cradle of agriculture. Science288(5471): 1602–1603.
38.
LiHWangCZhangX, et al. (2024) Mosaic pattern of sustained rice domestication and its environmental and cultural implications in Neolithic east China. Journal of Archaeological Science171: 106084.
39.
LiHWangZQianY, et al. (2025) Subsistence economic changes and their influencing factors in the lower Yangtze river region during 11,000–2,300 BP: Archaeobotanical and zooarchaeological evidence. Quaternary International740: 109891.
40.
LiHZhengDLiuZ, et al. (2022) Formation of agriculture and its impact factors in the Songze culture period in the lower reaches of the Yangtze river. Scientia Geographica Sinica42(10): 1807–1817.
41.
LimayeA (2012) Drishti: A volume exploration and presentation tool. Proc.SPIE. 85060X.
42.
LinYZuoXFuL, et al. (2025) Phytolith evidence of agricultural emergence and its dispersal routes in the southeastern mountain area of China. Journal of Archaeological Science Reports61: 104975.
43.
LordKALarsonGAllabyRG, et al. (2025) A universally applicable definition for domestication. Proceedings of the National Academy of Sciences122(22): e2413207122.
44.
MureauCD’OliveiraLPeyronO, et al. (2025) 8,000 years of wild and domestic animal body size data reveal long-term synchrony and recent divergence due to intensified human impact. Proceedings of the National Academy of Sciences122(36): e2503428122.
45.
MurphyCFullerDQ (2017) Seed coat thinning during horsegram (Macrotyloma uniflorum) domestication documented through synchrotron tomography of archaeological seeds. Scientific Reports7(1): 5369.
46.
ObataH (2015) Assessment of foxtail millet impression assemblages based on threshing and winnowing experiments and grain morphology. Japanese Association of Historical Botany23(2): 43–54.
47.
PanY (2023) Revisiting the archaeological investigations of rice domestication in China during 10,000–7,000 BP in a human behavioral context. Frontiers in Earth Science11: 1180376.
48.
PanYYuanJ (2018) A study on subsistence patterns from the Neolithic to the pre-Qin period in the lower Yangtze river valley (part 1). Cultural Relics in Southern China30(04): 111–125.
49.
PuruggananMDFullerDQ (2009) The nature of selection during plant domestication. Nature457(7231): 843–848.
50.
QinLFullerDQZhuT (2024) Why rice farmers don’t sail far: Coastal subsistence economic traditions and maritime trends in early China. Cultural Relics in Southern China 36(6): 76–96. DOI: 10.3969/j.issn.1004-6275.2024.06.008.(6)
51.
ReimerPJAustinWENBardE, et al. (2020) The IntCal20 northern hemisphere radiocarbon age calibration curve (0–55 cal kBP). Radiocarbon62(4): 725–757.
52.
RicePM (1987) Pottery Analysis: A Sourcebook. Chicago: The University of Chicago Press.
53.
RolettBVZhengZYueY (2011) Holocene sea-level change and the emergence of Neolithic seafaring in the Fuzhou basin (Fujian, China). Quaternary Science Reviews30(7-8): 788–797.
54.
SongJZhaoZFullerDQ (2013) The archaeobotanical significance of immature millet grains: An experimental case study of Chinese millet crop processing. Vegetation History and Archaeobotany22(2): 141–152.
55.
SpenglerRN (2022) Insularity and early domestication: Anthropogenic ecosystems as habitat islands. Oikos2022(12): e09549.
56.
SpenglerRNTangLDal CorsoM, et al. (2025) Seeking consensus on the domestication concept. Philosophical Transactions of the Royal Society B: Biological Sciences380(1926): 20240188.
57.
ThurberCSReagonMGrossBL, et al. (2010) Molecular evolution of shattering loci in U.S. weedy rice. Molecular Ecology19(16): 3271–3284.
58.
WangBLiuNLiuX, et al. (2025) Phytolith analysis reveals plant tempering in bronze-casting moulds from bronze age China (1250-221 BC) as a technological choice. Journal of Archaeological Science182: 106358.
59.
WangCLuHGuW, et al. (2018) Temporal changes of mixed millet and rice agriculture in Neolithic-bronze age central plain, China: Archaeobotanical evidence from the Zhuzhai site. Holocene28(5): 738–754.
60.
WangMLimayeALuJ (2024) Drishti paint 3.2: A new open-source tool for both 2D and 3D segmentation. Vertebrata Palasiatica62(4): 313–320.
61.
WuGZuoXWuW, et al. (2024) Late Neolithic to bronze age water management and upland rice cultivation in the mountainous areas of southeastern China coast. Quaternary International680: 55–63.
62.
WuJAoXLiuF, et al. (2025) Chemical insights into pottery production and use at Neolithic Zoumaling earthen-walled town in China. NPJ Heritage Science13(1): 115.
63.
WuWZuoXLiF, et al. (2022a) Development and influence of planting agriculture in Tanshishan culture: Based on the phytolith evidence from the Tanshishan site. Agricultural Archaeology42(04): 7–12.
64.
WuYWangCZhangZ, et al. (2022b) Subsistence, environment, and society in the Taihu lake area during the Neolithic era from a dietary perspective. Land11(8): 1229.
65.
YangXChenQMaY, et al. (2018) New radiocarbon and archaeobotanical evidence reveal the timing and route of southward dispersal of rice farming in south China. Science Bulletin63(22): 1495–1501.
66.
ZhangJJiangLYuL, et al. (2024a) Rice’s trajectory from wild to domesticated in east Asia. Science384(6698): 901–906.
67.
ZhangYBarronAPritchardJ, et al. (2024b) A microCT assessment of archaeological parenchyma from three riverine shell middens dating to the Early Holocene, Yongjiang River Basin, Guangxi, southern China. Journal of Archaeological Science Reports56: 104553.
68.
ZhaoZ (2010) New data and new issues for the study of origin of rice agriculture in China. Archaeological and Anthropological Sciences2(2): 99–105.
69.
ZhengXShaoDLiuB, et al. (2021) Subsistence of the Majiabang culture: New archaeobotanical evidence from the Yangjia and Majiabang sites. Jianghan Archaeology 42(5): 119–127.
70.
ZhouGZuoXZhouZ, et al. (2025a) Early island rice farmers on the south China coast during the 7th millennium BP. Journal of Archaeological Science175: 106158.
71.
ZhouXSpenglerRNSayfullaevB, et al. (2025b) 9,000-year-old barley consumption in the foothills of central Asia. Proceedings of the National Academy of Sciences122(36): e2424093122.
72.
ZhouZHuangYFanX (2018) The Nanshan site in Mingxi county, Fujian. Archaeology19(1): 15–12.
73.
ZoharyDHopfMWeissE (2012) Domestication of Plants in the Old World. Oxford, UK: Oxford Univ Press.
74.
ZongYYaoSCrawfordGW, et al. (2017) Selection for oil content during soybean domestication revealed by X-ray tomography of ancient beans. Scientific Reports7(1): 43595.
75.
ZuoXDaiJWuW, et al. (2022) Microfossil evidence of rice cultivation on the southeast China coast 7500 years ago. Science China Earth Sciences65(11): 2115–2126.
