Stable Carbon and Nitrogen Isotope Analysis for subsistence Shift and Dietary Differentiation from the Late Neolithic to the early Bronze Ages in the Central Plains, China

Stable Carbon and Nitrogen Isotope Analysis for subsistence Shift and Dietary Differentiation from the Late Neolithic to the early Bronze Ages in the Central Plains, China | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Article Stable Carbon and Nitrogen Isotope Analysis for subsistence Shift and Dietary Differentiation from the Late Neolithic to the early Bronze Ages in the Central Plains, China Xianglong Chen, Yue You, Guangkuo Yuan, Yaowu Hu This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-8751401/v1 This work is licensed under a CC BY 4.0 License Status: Under Revision Version 1 posted 11 You are reading this latest preprint version Abstract The Central Plains, a core region for early Chinese civilization, underwent profound sociocultural transformations during the third and second millennia BC, as well as the intensification and diversification of the agricultural economy. This study explores the subsistence changes through isotopic analysis of human and animal remains from the Pan Mount Song Area (PMSA), the hinterland of the Central Plains. We present new stable carbon and nitrogen isotope data from the MeiShan site in the PMSA, complemented by a comprehensive analysis of published datasets. Findings reveal distinct subsistence patterns across different periods. During the Longshan period (ca. 4400 − 3800 BP), millet remained the staple crop for both humans and domesticated animals, while some individuals consumed significant amounts of rice, indicating a notable shift in subsistence compared to the Yangshao culture period (ca. 6300 − 4500 BP), when millet farming was highly emphasized. From the Erlitou culture (ca. 3800 − 3500 BP) and Erligang culture (ca. 3500 − 3350 BP) to the late Shang periods (ca. 3210 − 3000 BP), millet-derived nutrients became predominant. When integrated with the published isotopic data focusing on PMSA, it is found that the dietary patterns of inhabitants from the late Neolithic to early Bronze Ages were predominantly millet-based during most periods. However, a relatively high proportion of C 3 or rice-derived foods could be found in the diets of some residents at several late Longshan sites, as well as the Early Bronze Age capitals and regional centers. The disparities in human dietary patterns at these sites are presumably ascribable to the increasing complexity of the agricultural economy. Furthermore, they may also be associated with human mobility and the disparity in resource allocation during the urbanization process, which are more prone to have exerted substantial impacts at the central sites of the Early Bronze Age. Late Neolithic Bronze Age MeiShan Subsistence Dietary Differentiation Figures Figure 1 Figure 2 Figure 3 Introduction The agricultural landscape of ancient China was marked by a distinct dichotomy. The millet-based farming system dominated the middle and lower reaches of the Yellow River basin and its northern regions, whereas the rice-based farming system prevailed in the middle and lower reaches of the Yangtze River basin and its southern areas 1 – 4 . The origin and early development of Chinese civilization were also spurred by the interaction between these two types of agricultural economies, a phenomenon unique in world prehistory. It is well-established that broomcorn millet ( Panicum miliaceum ) and foxtail millet ( Setaria italica ) functioned as the staple crops in the Central Plains since the middle Neolithic Age 5 – 7 . Commencing around 8000 BP, the northward expansion of rice cultivation led to the emergence of a millet-rice mixed farming zone in central China 8 , 9 , with the Central Plains located at its northern periphery 10 , 11 . During the Longshan culture period (approximately 4400 BP to 3800 BP), East Asia became integrated into the broader Bronze Age exchange networks 12 , especially through its involvement in prehistoric Food Globalization events 13 . This process was characterized by the eastward dissemination of multiple domesticated species, such as wheat ( Triticum aestivum ), sheep ( Ovis aries ), and cattle ( Bos taurus) , from western Eurasia, which were subsequently incorporated into the subsistence strategies of various regions, particularly the Central Plains. The archaeological evidence indicates a significant increase in both the frequencies and absolute quantities of cereal grains and domesticated animal remains during the Longshan culture period, reflecting a substantial shift towards a more diversified and intensified agricultural economy 14 . The transformation of subsistence regimes was paralleled by substantial changes in material culture and sociopolitical systems, particularly in the Pan Mount Song Area (PMSA), the core region of the Central Plains. Comprehensive archaeological surveys in the Luoyang Basin, situated in the northern part of PMSA, have revealed a remarkable transition. This transition was from a decentralized and competitive settlement pattern, characterized by several subgroups with a three-tiered hierarchy during the Longshan culture period, to a more complex four-tiered settlement hierarchy. This new structure reached its peak in a large-scale state-level society centered on the ErLiTou site during the Erlitou culture period (ca. 3800 BP to 3500 BP), which is argued to be the onset of the Bronze Age in Chinese Archaeology 15 , 16 . This socioeconomic transformation has attracted considerable attention from archaeologists and historians, and is often associated with the origin of the legendary Xia dynasty in Chinese history. Although the diversified and intensified agricultural economy undoubtedly increased food provisions to support population growth, the degree of its association with material culture and social dynamics remains a, matter of contention. A comprehensive exploration from the paleodietary perspective, integrating archaeobotany and zooarchaeology, and concentrating on the diachronic variations of subsistence practices in prehistory of PMSA, especially from the Longshan culture period to the early Bronze Age, is crucial for comprehending the aforementioned issues. This study presents novel stable carbon and nitrogen isotope data from human (n = 17) and animal (n = 47) remains dating from the late 3rd to the middle 2nd millennium BC at the MeiShan site, located in the southern region of the PMSA. Furthermore, these findings are combined with previously published isotopic data, as well as archaeobotanical and zooarchaeological research, from the late Neolithic to early Bronze Ages, enabling a more robust and comprehensive analysis of subsistence transitions as well as dietary differentiation across the PMSA region. This integrated approach aims to offer an alternative interpretation for interpreting the complex interconnections between subsistence economy transformations, cultural evolution, and social structural changes, thereby enhancing our understanding of these critical processes in prehistoric societies. Subsistence in the PMSA Previous archaeobotanical research has indicated that the assemblages of charred seeds at late Longshan sites in the PMSA were primarily composed of foxtail and broomcorn millets, with rice and soybean functioning as supplementary crops. Only a limited number of charred wheat remains have been identified at several Longshan sites, including WangGeDang 17 , WaDian 18 , 19 , WangChengGang 20 . It is crucial to emphasize that all currently available absolute dates for charred wheat remains in the PMSA postdate the Longshan culture period 21 . Whether wheat was actually utilized by the inhabitants during the Longshan culture period in this region remains undetermined. During the Erlitou culture period and the subsequent Shang dynasty, which includes the Early Shang (Erligang culture period, ca. ~3500 − 3300 BP) and Late Shang (Yinxu period, ca. 3300 − 3000 BP), archaeological evidence reveals a gradual yet consistent increase in both the quantity and ubiquity of wheat 22 – 28 . The increasing significance of soybean and rice, along with the adoption of wheat, has led to hypothesis that a multiple cropping regime was established during the Longshan culture period and continued to be strengthened throughout the early Bronze Age 20 . Zooarchaeological evidence indicates that sheep and cattle husbandry was integrated into the agricultural systems alongside local domesticated pigs and dogs during the Longshan culture period in the Central Plains 14 . The diversification of domesticated animal species provided communities with a broader range of animal products and facilitated the secondary exploitation of animals for wool, traction, and other purposes during the late Neolithic and Bronze Ages 29 – 31 . Archaeobotanical and zooarchaeological investigations provide essential insights into human subsistence strategies in PMSA, while stable isotope analysis offer a method to assess the relative contributions of different food production techniques to human diet. Recently published δ 13 C and δ 15 N values have revealed that the inhabitants of Longshan settlements, such as WaDian and HaoJiaTai 32 , 33 , can be classified into at least two distinct dietary groups based on the proportion of millet- versus rice-derived nutrients in their diets. During the subsequent Erlitou culture period and the Shang dynasty, isotopic evidence indicates a predominant focus on C 4 -based foods at sites including XinZhai (ca. 3800 − 3700 BP) 34 , 35 , NanWa (ca. 3800-3000BP) 36 , and Xiaoshuangqiao (ca. 3350 − 3210 BP) 37 , with the exception of some outliers showing relatively negative δ 13 C values at ErLiTou 38 – 40 and XiaoHuCun location at the ZhengZhou ShangCheng site (ca. 3300 − 3000 BP) 41 . Regarding animal husbandry practices, multiple studies have demonstrated that millet farming byproducts served as the primary fodder for both omnivorous (pigs and dogs) and herbivorous animals (cattle and some sheep) across nearly all PMSA sites from the Longshan culture to the Shang dynasty 32 – 35 , 38 , 40 – 43 . This consistent pattern of animal feeding practices illustrates the integral role of millet agriculture in supporting diverse animal husbandry strategies throughout the region's subsistence development in the Neolithic and Bronze Ages. Substantially more research efforts are still needed to comprehensively document the transition process of dietary complexity in prehistory of the PMSA, particularly during the critical period of early state formation from Neolithic to Bronze Ages. To address these research questions, a dual approach integrating both detailed case studies and broader regional comparisons is indispensable. Archaeological background The study site, MeiShan, is situated in RuZhou city, Henan province, on the left terrace of the Xi'Er River, a minor tributary of the BeiRu River within the upper Ying River Valley of the PMSA (Fig. 1 ). Archaeological surveys have delineated an area of approximately 20 hectares, which was predominantly occupied from the Longshan culture through the late Shang or Yinxu periods. In 1995, an excavation spanning ca. 1200 m 2 unearthed dozens of trash pits and several house foundations dating back to the Longshan culture, Erlitou culture, and late Shang periods. Additionally, six earth-pit tombs from the Longshan culture were discovered as well. Typological examination of pottery indicates that the Longshan Culture period remains predominantly mirror the Wangwan Phase III cultural tradition, while also integrating non-local elements similar to the Qujialing-Shijiahe cultures of the middle Yangtze River catchment area in the south and the Dawenkou-Longshan cultures of Shandong Province in the east 44 . During the 1995 field season, numerous fragmented human skeleton remains were identified within diverse contexts, such as strata and trash pits, etc., frequently intermingled with animal bones and artifacts—a common phenomenon during the Longshan and Erlitou culture periods in the Central Plains. Zooarchaeological investigations at the MeiShan site revealed a diverse faunal assemblage comprising both domesticated and wild species. The domesticates included pigs, dogs, sheep, and cattle, while the wild fauna consisted of deer, porcupine, and various bird species. Quantitative analysis of the faunal remains showed that pig bones constituted the majority of the assemblage, accounting for approximately 53%, followed by deer (31%), dogs (6%), sheep (5%), and cattle (2.5%). Stable carbon and nitrogen isotope data of 4 individuals from late Longshan tombs have been published, showing a dichotomy in human diet at MeiShan. While three individuals exhibited diets dominated by 15 N-enriched C 3 food resources, the remaining individuals primarily relied on C 4 resources 45 . In this study, seventeen human bones from strata, trash pits of different periods were sampled for δ 13 C and δ 15 N measurement, allowing a paleodietary comparison of different contexts, prior to comprehensive analysis of subsistence transitions across the PMSA region. To establish a reliable isotopic baseline and investigate domesticated animal husbandry practices from the late Neolithic to the early Bronze Ages, both domesticated and wild animal bones were selected for stable carbon and nitrogen isotope analysis. The animal samples included 20 pigs, 5 dogs, 5 cattle, 8 sheep, and 8 deer. The majority of these samples were dated to the Longshan culture period, with the remaining specimens originating from contexts associated with the Erlitou culture and Shang periods, as detailed in Table 1. Results The collagen yields, as well as carbon and nitrogen contents of the sampled bones from MeiShan, are summarized in Table S1 . The majority of the samples were well-preserved, with collagen yields ranging from 1.1% to 12.3%. Moreover, these samples yielded high quality collagen suitable for radiocarbon dating and dietary reconstruction 46 , with carbon contents ranging from 24.1% to 45.2% and nitrogen contents from 8.8% to 16.4%, and the atomic C:N ratios fell within the range of 3.1 to 3.3. The radiocarbon dating results are presented in Table S2 . The five samples ascribed to the Longshan culture period are concentrated within the range of ca. 4240 to 3720 BP, which is consistent with the chronological framework of the Wangwan Phase III Longshan culture. In contrast, the two samples from the Erlitou culture period date to ca. 3560 to 3390 BP, corresponding to the final stage of the Erlitou culture or slightly later. The isotopic results of human and animal bone collagen are summarized in Table S1 and plotted in Fig. 2 . The isotopic values of animal bones exhibited a broad range, reflecting diverse dietary patterns. Deer displayed mean δ 13 C (-18.4 ± 1.7‰) and δ 15 N (3.8 ± 0.8‰) values, indicative of a C 3 plant-based diet. In contrast, domesticated animals exhibited higher δ 13 C values (ranging from − 18.9‰ to -6.0‰), with mean δ 13 C values for pigs, dogs, cattle, and sheep being − 11.7 ± 3.6‰, -9.3 ± 0.8‰, -10.2 ± 2.5‰, and − 16.6 ± 2.6‰, respectively, suggesting varying degrees of C 4 fodder consumption. The mean δ 15 N values for dogs (7.4 ± 0.7‰), pigs (7.5 ± 1.0‰), cattle (6.8 ± 2.4‰), and sheep (3.9 ± 1.1‰) were all elevated in comparison to those of deer. The MeiShan inhabitants exhibited considerable variation in δ 13 C (-16.9‰ to -6.8‰) and δ 15 N (7.5‰ to 12.3‰) values, indicating dietary diversity. Their consumption of C 4 versus C 3 foods likely derived primarily from millet foods (i.e. millet grains and domesticated animals raised with millet byproducts) or rice and related food products. Among the 13 Longshan individuals, three displayed δ 13 C values no more positive than − 14.1‰ and δ 15 N values no less than 10.8‰, while the remaining individuals exhibited less negative δ 13 C values (-13.4‰ to -6.8‰) and lower δ 15 N values (7.5‰ to 9.5‰). This pattern suggests differential access to food resources within the MeiShan community during the Longshan culture period. In contrast, the isotopic values (δ 13 C range = -10.9‰ to -7.1‰, δ 15 N range = 8.5‰ to 9.5‰) indicate a primary reliance on millet-based nutrients with only minor supplementation from C 3 foods such as rice. The individual labelled MS01, lacking chronological context, displayed δ 13 C and δ 15 N values of -9.5‰ and 8.1‰, respectively, suggesting a diet predominantly based on millet farming. Discussion Animal husbandry practices In prehistoric farming communities of the Central Plains and adjacent regions, pigs and dogs, which are typical indigenous domesticates in East Asia, often exhibited comparable isotopic patterns. This suggests that their diets were similar, predominantly characterized by C 4 resources, such as millet byproducts, human leftovers, or/and even millet grains 35 , 47 . The isotopic data (δ 13 C ranging from − 10.3‰ to -8.4‰, δ 15 N values ranging from 6.3‰ to 8.3‰, n = 5) indicate that this dietary pattern likely applied to dogs at MeiShan from the Longshan culture to the YinXu periods. In contrast, pigs of the Longshan culture period at MeiShan displayed a wider range of δ 13 C values (-18.2‰ to -6.0‰), with a relatively negative mean value (-12.0 ± 3.8‰, n = 14). This implies a mixed C 3 /C 4 fodder and more flexible feeding practices. Similar isotopic data have also been identified at contemporaneous sites like WaDian 32 and XinZhai in the PMSA 34 , 35 , where considerable amounts of rice remains were discovered alongside charred millets 18 , 19 , 22 . This has given rise to the hypothesis that C 3 fodder, presumably derived from rice, was employed for pig rearing at these sites, a practice which might have also occurred at MeiShan. During the Erlitou culture period, the mean δ 13 C value of pigs (-10.8 ± 3.1‰, n = 5) increased slightly, ranging from − 14.0‰ to -6.8‰, while δ 15 N values remained relatively stable (7.5 ± 0.5‰, ranging: 6.6‰ to 7.9‰), indicating a decreased reliance on rice in pig diets. This trend appears to have been prevalent in the PMSA between approximately 4000 and 3500 BP, as pigs at WangGeDang, ErLiTou 40 and XinZhai 34 , 35 exhibiting similar millet-dominated diet patterns. The introduction of domesticated sheep and cattle to the Central Plains can be traced back to no later than the Longshan period, as evidenced by the extensive distribution of their remains 14 . Sheep exhibited δ 15 N patterns (3.9 ± 1.1‰) similar to those of deer, while their δ 13 C values were more positive (-16.6 ± 2.6‰, range: -18.9‰ to -11.6‰). The isotopic pattern has been observed at numerous Longshan culture period and Bronze Age sites in the Central Plains, and is attributed to the consumption of millet byproducts, particularly during winter 32 , 47 , 48 . At MeiShan, the five sheep from the Longshan culture period displayed δ 13 C values ranging from − 18.6‰ to -14.9‰, while some sheep from the Erlitou culture (#MS48, -11.6‰) and late Shang (#MS49, -15.0‰) periods exhibited less negative δ 13 C values, indicating an increased intake of millet straws. All MeiShan cattle, including one from the Longshan and three from the Erlitou culture periods, exhibited elevated δ 13 C values (-11.7‰ to -7.0‰), reflecting a focus on millet-related fodder. This pattern is consistent with cattle husbandry practices in other PMSA agricultural communities during the Longshan and Erlitou culture periods, indicating that cattle husbandry became increasingly dependent on millet farming 32 , 47 , 48 . The utilization of sheep and cattle offered novel opportunities for diversifying livestock husbandry practices. This was not only manifested in the provision of protein, wool, hides, bone materials, and traction power, but also in a more efficient utilization of agricultural byproducts, and grasses and stubble from fallow and marginal lands. As a result, the rearing of these ruminants contributed to a more productive land use model and agricultural system. From this viewpoint, it is contended that ruminant husbandry, especially cattle raising, had a positive influence on and became an essential part of the agricultural system as a supplementary subsistence practice since the end of the 3rd millennium BC. Human diet complexity at MeiShan and contemporary sites in the PSMA Although there is a dearth of archaeobotanical data from MeiShan, flotation investigations at adjacent sites within the PMSA, dating from roughly 4400 BP to 3400 BP or later, have indicated that foxtail millet was the predominant constituent of the charred grain assemblages, with rice occupying the second or third position 49 . Consequently, it is hypothesized that rice cultivation significantly contributed to the subsistence economy and transformed the local cropping system during this period, which had previously been an agricultural regime primarily centered around millet 27 , 50 , 51 . Nevertheless, the role of rice farming in human subsistence and whether disparities in rice consumption existed among sites and individuals remain unresolved queries. Our study offers novel perspectives on these issues. The isotopic results of 13 Longshan human samples from our study, combined with previously published data from 4 individuals, are discussed together to understand human diet and subsistence at MeiShan. Overall, the 17 individuals exhibited a wide spectrum of values, ranging from − 16.9‰ to -6.8‰ for δ 13 C and from 7.5‰ to 12.3‰ for δ 15 N, indicating a diverse intake among the subjects. While the majority of individuals followed a mixed C 3 /C 4 diet, some depended solely on either C 3 (rice-derived) or C 4 (millet-derived) nutrients, as shown by their δ 13 C values. The mean δ 15 N (9.6 ± 1.4‰, n = 17) of these individuals was significantly higher than that of contemporary animals, including omnivorous pigs (7.5 ± 1.2‰, n = 14) and dogs (7.5 ± 1.0‰, n = 3), domesticated herbivorous cattle (5.5‰, n = 1) and sheep (3.7 ± 0.8‰, n = 5), as well as deer (3.4 ± 0.3‰, n = 4). Given that the δ 15 N range is 7.5‰ to 12.3‰, some individuals with δ 15 N values comparable to those of domesticated omnivores might have had limited access to food resources with higher δ 15 N values, while others probably consumed more δ 15 N-enriched nutrients. The dietary diversity, as reflected by δ 15 N values, is primarily attributed to disparities in human trophic levels 52 , which might have resulted from differential access to animal products among individuals. A previous study at WaDian identified two distinct dietary groups based on stable carbon and nitrogen isotope analysis. One group primarily consumed millet with some amount of domesticated animal protein, while the other exhibited a mixed C 3 /C 4 diet enriched in 15 N 32 . This finding has been corroborated by recent publications on WaDian, HaoJiaTai, MeiShan, Jiazhuang, and other Longshan culture sites 33 , 45 . Zhou observed that elites at MeiShan and JiaZhuang, whose grave goods exhibited southern stylistic influences, primarily consumed rice-derived foods. These findings provide evidence for diffusion and popularization in PMSA of southern-oriented rice farming and materials. The high proportion of rice-derived foods in the diets of elites during the Longshan culture period raises questions about whether rice was a highly valued staple crop and whether access to rice was linked to social status. While attributing δ 15 N enrichment to social disparities between rice-eaters and millet-eaters offers a straightforward explanation, it may not fully account for other potential factors. A pilot study on plant isotopes at XinZhai, WaDian, ChengYao, WangChengGang and other contemporary sites in PMSA indicates that rice exhibits higher δ 15 N values than millets, possibly due to nitrogen metabolism in paddy fields 53 . Fox example, at XinZhai, the mean δ 15 N values of C 4 millet and C 3 rice grains are 4.4 ± 1.0‰ (n = 15) and 8.0 ± 1.4‰ (n = 10), respectively, highlighting the disparity in δ 15 N between the two staple crops. Consequently, it remains unclear whether the lower δ 15 N values of millet-eaters reflect a more vegetarian diet. A linear correlation test between human δ 13 C and δ 15 N values at MeiShan provides further insights into human dietary preferences and potential causal factors. The correlation of δ 13 C and δ 15 N for all 17 Longshan culture residents at MeiShan (n = 17, r 2 = 0.69) suggests that individuals with a higher intake of 13 C-depleted or rice-derived nutrients exhibited elevated bone collagen δ 15 N values. Furthermore, the δ 13 C and δ 15 N values of MeiShan pigs showed a significant correlation as well (r 2 = 0.62, n = 20), indicating that the dietary intake of rice products by pigs does not necessarily imply a higher consumption of animal-derived materials. Instead, the correlation is probably related to the difference of δ 15 N values between rice and millet-related fodders. We argue consequently that the correlation among δ 13 C and δ 15 N values of MeiShan residents likely reflects differences in the consumption of millet versus rice. A comprehensive investigation integrating isotopic data and social status at WaDian indicated no distinct hierarchical disparities between millet-eaters and mixed C 3 /C 4 diet consumers. Instead, it was hypothesized that differences in social status existed within each group 33 . As previously noted, individuals with burial customs associated with the Shijiahe culture in the southern region typically presented more negative δ 13 C values at MeiShan and contemporary sites. This suggests that rice farming might have played a more important role in their subsistence economy, analogous to the communities of the Shijiahe culture. Nevertheless, more evidence is required to conclude that rice and rice-derived foods were highly valued during the Longshan culture period in the PMSA in the future. Comparatively speaking, the three MeiShan residents in the Erlitou culture period exhibited more clustered δ 13 C values (ranging from − 10.9‰ to -7.1‰), indicating a focus on millet-derived nutrients supplemented with only a minor supplements of C 3 foods. Their δ 15 N values (ranging from 8.5‰ to 9.5‰) exceeded those of pigs (7.5 ± 0.5‰, n = 5), dogs (7.1‰, 7.6‰, n = 2), cattle (7.2 ± 2.7‰, n = 4) and sheep (2.6‰, 5.9‰ n = 2). This suggests that MeiShan residents during the Longshan culture period experienced diverse diets and lifestyles, while their successors in the Erlitou culture period relied more heavily on millets and millet feeding domesticates. Subsistence shift from the Neolithic to the early Bronze Age in PMSA To gain a more comprehensive understanding of the development of subsistence economies in the PMSA from the Neolithic to the Bronze Age, we have systematically compiled and chronologically plotted the published δ 13 C and δ 15 N values of humans (Table S3 ) in Fig. 3 . Isotopic analysis results indicate that millet-based foods constituted the primary component of the food resources for ancient peoples in the PMSA region from the Yangshao culture to the late Shang periods. Meanwhile, during certain periods, the dietary contribution of C 3 foods increased markedly among a small number of inhabitants. The dietary patterns largely reflect the diachronic developmental trends of the subsistence economy in the PMSA region. Archaeobotanical evidence from Peiligang culture sites such as Peiligang 54 and Zhuzhai 8 indicates that millets and rice were cultivated in the PMSA as early as 8000 BP to 7500 BP. During the subsequent Yangshao culture period, the ubiquity and quantities of foxtail and broomcorn millets increased significantly as the primary crops, while rice cultivation served as the supplement farming practice 44 , 55 . By the Longshan culture period, the agricultural economy shifted from a focus on millet farming to a more complex system incorporating the cultivation of both millet and rice. The shift is evident at WaDian, HaoJiaTai, and MeiShan 32 , 33 , where two distinct dietary groups were isotopically identified; one was reliant on millet, and the other consumed more rice-derived nutrients. The successful integration of diverse dietary and subsistence groups was crucial for the stability and subsequent development of early societies, which was a key factor in the formation of a unified early Chinese civilization. Compared to the Longshan culture period, the δ 13 C and δ 15 N isotope ratios of the Erlitou culture population in the PMSA were more concentrated, reflecting a greater reliance on millet-based agricultural products and a reduced contribution of C 3 foods, namely rice. These contracts with the proto-Shang, and late Shang populations at ZhengZhou ShangCheng 41 and the Erligang culture population at WangJingLou 37 , where some individuals exhibited more diverse diets with greater contribution from C 3 foods, despite millet farming remaining the dominant agricultural economy. From the Yangshao culture to the late Shang period, the transitions in human diet, subsistence, and social complexity process appear to be chronologically aligned. The role of the agricultural economy in supporting or influencing the social complexity process in the PMSA requires further investigation. During the Yangshao culture period, the rapid development of millet farming in the Central Plains coincided with significant cultural changes, including population growth and the expansion of cultural influence to the surrounding areas. Millet-oriented agricultural economy probably provided the food security necessary for these developments. During the Longshan culture period, deteriorating climatic conditions and frequent floods may have prompted the adoption of rice farming in the Central Plains and surrounding areas, potentially influenced by interactions with archaeological cultures in southern lowlands, such as the Qujialing and Shijiahe cultures, as suggested by ancient DNA analysis 56 and artifacts with southern stylistic influences 9 . Our study provides insights into human adaptation, cultural integration, and population interaction in the PSMA during the Longshan culture period, topics that merit further research. At the beginning of the Bronze Age, the rise of Erlitou culture (ca. 3800 BP to 3500 BP) unified the Central Plains hinterland and surrounding regions, with its center in the Luoyang Basin of the northern PMSA 57 . While the contribution of rice to human diet declined, millets remained highly valued 27 , 40 . Millet farming not only provided a staple food for ErLiTou communities but also served as feed for pigs, dogs, as well as cattle. The subsequent Erligang culture (ca. 3500 BP to 3350 BP), corresponding to the early period of the Shang Dynasty, centered on ZhengZhou ShangCheng, where some individuals exhibited more diverse diets with greater contributions from C 3 foods, possibly reflecting migration and cultural interactions during the transition from Erlitou to Erligang cultures. When it comes to the middle (ca. 3350 BP to 3210 BP) and late Shang periods (ca. 3210 BP to 3000 BP), dietary variations among individuals still persisted. Given the profound changes in social structure, namely the rise of large-scale state-level society, the implications reflected by dietary variations among different individuals can no longer be adequately explained solely through the lens of subsistence economy. During the Neolithic Age, social division of labor had not yet manifested distinct specialized characteristics, and the majority of inhabitants at Neolithic sites still relied mainly on agricultural production for their livelihoods. Consequently, the proportion of C 4 foods (e.g., foxtail millet, broomcorn millet) to C 3 foods (e.g., rice) in the diets of inhabitants was directly linked to the cultivation scale and yield of different crops. This phenomenon may have been prevalent at the MeiShan site investigated in this study, as well as at other sites dating from the Yangshao to the Longshan culture periods, such as the WangGou, WaDian and HaoJiaTai sites. In contrast, for urban residents, whose livelihoods focused on non-agricultural activities at the capital and regional centers, the ratio of C 3 to C 4 foods in their diets was largely determined by food acquisition channels and distribution mechanisms developed by the early states. At the ZhengZhou ShangCheng, Erlitou and Wangjinglou sites, the capitals or regional centers of the Bronze Age, the dietary contribution of C 3 foods to urban inhabitants increased markedly, and the combination of food types became more diverse. This presumably suggests that they may have originated from subsistence economic zones distinct from those of the main population, possessed more extensive food acquisition networks, or occupied an advantageous position in the distribution of social resources—allowing them to obtain non-local agricultural products from various regions via trade, tribute and other such means. Such dietary differentiation stemming from disparities in social roles and economic activities provides an important material cultural basis for our understanding of the process of social complexity, patterns of resource distribution, and urban-rural relations in ancient societies. Conclusions The PSMA of the core area of the Central Plains of China featured an economic model dominated by millet farming alongside rice cultivation of a certain scale from the Yangshao period to the Early Bronze Age. Roughly from the 3rd millennium to the 2nd millennium BC, the Central Plains underwent a transformation of social structure, evolving from the polycentric settlement pattern of the late Longshan period into early large-scale state-level society with overarching authority. Meanwhile, domesticated cattle and sheep as well as wheat originating from West Asia were also introduced into this region. Although dietary patterns of humans at different sites and in different periods can reflect, to a certain extent, the developmental trends of the subsistence economy in this region, specifically millet farming versus rice farming, the subsistence economic conditions during the formation of early states, particularly the impact of early state formation on the dietary patterns of different populations, still warrant further research. To gain a deeper understanding of these issues, we conducted stable carbon and nitrogen isotope analyses on human and animal bones from the Longshan culture period, the Erlitou culture period, and the late Shang period at the MeiShan site in Ruzhou, Henan province. To facilitate a more rigorous and comprehensive examination of subsistence transitions and dietary differentiation across the PMSA region, the data from the MeiShan site are integrated with previous publications spanning from the late Neolithic to the early Bronze Ages. It is revealed that during the Longshan culture period, the dietary pattern of the inhabitants at the MeiShan site showed a strong signal of rice consumption, although millet-related foods were still widely consumed. However, during the Erlitou culture period, the contribution of rice to the human diet notably declined, whereas millet farming continued to be highly emphasized. This shift in agricultural practices was also reflected in the feeding practices of domesticated animals. From the Longshan culture to the Erlitou culture periods, the proportion of rice-related foods in the fodder of domesticated pigs decreased significantly, while millet-related feed increased. By integrating previous research findings, we propose that the subsistence economy in the PSMA, represented by the MeiShan site during the Longshan culture period, was likely influenced by the Shijiahe culture from the middle Yangtze River catchment. Under this influence, the inhabitants of MeiShan and nearby Longshan culture sites such as HaoJiaTai and WaDian consumed a notable amount of rice products. However, the trend toward agricultural diversification did not rapidly progress during the Erlitou culture period or the subsequent Erligang culture and late Shang periods. Instead, millet farming once again became the dominant form of local agricultural economy, with rice and wheat cultivation serving only as supplementary practices. By comprehensively examining the isotopic data of the inhabitants at the MeiShan site in conjunction with those from sites in this region spanning the Neolithic to the Early Bronze Ages, and contextualizing these findings within the macro backdrop of social transformation, we argue that the dietary patterns of Neolithic residents from the Yangshao to the Longshan periods likely primarily reflected the subsistence economic characteristics of the respective sites. The changes in the subsistence economy of the PSMA from the late Neolithic to the early Bronze Age were the result of the development of the traditional millet farming oriented agricultural economy in the area. At the same time, these changes were also related to the frequent cultural interactions between the Central Plains and the middle reaches of the Yangtze River, which began during the Longshan culture period. Besides, the dietary variations within the resident groups at early Bronze Age capital sites and regional centers may also have been associated with the social complexity phenomena driven by factors such as individual migration and food resource distribution during the urbanization process. Declarations Competing interests The authors declare no competing interests. Author Contribution X.C. collected the samples, analyzed the data, prepared all figures and tables, and wrote the manuscript; Y.Y. contributed to the interpretation of the data and revised the manuscript; G.Y. and Y.H. offered valuable suggestions and contributed to the revision of the manuscript. 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Supplementary Files TableS1Sampleinformationandisotopedataofth.docx TableS2SamplesinformationandresultsforAMS1.docx TableS3Stablecarbonandnitrogenisotopevalues.docx Cite Share Download PDF Status: Under Revision Version 1 posted Editorial decision: Revision requested 31 Mar, 2026 Reviews received at journal 31 Mar, 2026 Reviews received at journal 10 Mar, 2026 Reviews received at journal 02 Mar, 2026 Reviewers agreed at journal 26 Feb, 2026 Reviewers agreed at journal 22 Feb, 2026 Reviewers agreed at journal 19 Feb, 2026 Reviewers invited by journal 18 Feb, 2026 Editor assigned by journal 05 Feb, 2026 Submission checks completed at journal 05 Feb, 2026 First submitted to journal 31 Jan, 2026 You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. 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study\u003c/strong\u003e\u003c/p\u003e","description":"","filename":"1.png","url":"https://assets-eu.researchsquare.com/files/rs-8751401/v1/486cb9340ec0c42eb1407771.png"},{"id":103505868,"identity":"c80d1d96-ae0d-4893-9ad2-ed78cfaf0dfe","added_by":"auto","created_at":"2026-02-26 13:33:17","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":83523,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eδ\u003c/strong\u003e\u003csup\u003e\u003cstrong\u003e13\u003c/strong\u003e\u003c/sup\u003e\u003cstrong\u003eC and δ\u003c/strong\u003e\u003csup\u003e\u003cstrong\u003e15\u003c/strong\u003e\u003c/sup\u003e\u003cstrong\u003eN values for humans and animals (mean ± SD) from the MeiShan site in the PSMA, Henan province, China.\u003c/strong\u003e\u003c/p\u003e","description":"","filename":"2.png","url":"https://assets-eu.researchsquare.com/files/rs-8751401/v1/3dd208df371d8e6424d22cc5.png"},{"id":103252955,"identity":"d881dd31-7be2-4fac-bbe1-2e10195c115f","added_by":"auto","created_at":"2026-02-23 16:16:46","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":502513,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eThe comparison of human δ\u003c/strong\u003e\u003csup\u003e\u003cstrong\u003e13\u003c/strong\u003e\u003c/sup\u003e\u003cstrong\u003eC and δ\u003c/strong\u003e\u003csup\u003e\u003cstrong\u003e15\u003c/strong\u003e\u003c/sup\u003e\u003cstrong\u003eN values of archaeological sites in the PSMA from the Neolithic Yangshao culture periods to the Early Bronze Age Shang dynasty, Henan province, China.\u003c/strong\u003e\u003c/p\u003e","description":"","filename":"3.png","url":"https://assets-eu.researchsquare.com/files/rs-8751401/v1/4d99c80fc24ef6cb42f57445.png"},{"id":104410180,"identity":"b1a29466-09a4-4993-894f-a3af42b59222","added_by":"auto","created_at":"2026-03-11 12:49:53","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":2627493,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-8751401/v1/82b5aa8b-00ad-45d5-b4e7-83e6a9abc51d.pdf"},{"id":103252923,"identity":"8c9c3159-fca4-4ab5-beaa-ef19f831a250","added_by":"auto","created_at":"2026-02-23 16:16:39","extension":"docx","order_by":1,"title":"","display":"","copyAsset":false,"role":"supplement","size":41782,"visible":true,"origin":"","legend":"","description":"","filename":"TableS1Sampleinformationandisotopedataofth.docx","url":"https://assets-eu.researchsquare.com/files/rs-8751401/v1/ecf77988c1c8ae9cc336cc2e.docx"},{"id":103252992,"identity":"00d78999-079f-4d43-9fdb-a29e1f2ab76d","added_by":"auto","created_at":"2026-02-23 16:16:56","extension":"docx","order_by":2,"title":"","display":"","copyAsset":false,"role":"supplement","size":13440,"visible":true,"origin":"","legend":"","description":"","filename":"TableS2SamplesinformationandresultsforAMS1.docx","url":"https://assets-eu.researchsquare.com/files/rs-8751401/v1/b78eecb00381dd418e9c5465.docx"},{"id":103252962,"identity":"f38a74a3-79ae-476d-a28e-ccfc50e7c9b8","added_by":"auto","created_at":"2026-02-23 16:16:48","extension":"docx","order_by":3,"title":"","display":"","copyAsset":false,"role":"supplement","size":370898,"visible":true,"origin":"","legend":"","description":"","filename":"TableS3Stablecarbonandnitrogenisotopevalues.docx","url":"https://assets-eu.researchsquare.com/files/rs-8751401/v1/c878cdfbea1556716a5ba7a0.docx"}],"financialInterests":"No competing interests reported.","formattedTitle":"Stable Carbon and Nitrogen Isotope Analysis for subsistence Shift and Dietary Differentiation from the Late Neolithic to the early Bronze Ages in the Central Plains, China","fulltext":[{"header":"Introduction","content":"\u003cp\u003eThe agricultural landscape of ancient China was marked by a distinct dichotomy. The millet-based farming system dominated the middle and lower reaches of the Yellow River basin and its northern regions, whereas the rice-based farming system prevailed in the middle and lower reaches of the Yangtze River basin and its southern areas\u003csup\u003e\u003cspan additionalcitationids=\"CR2 CR3\" citationid=\"CR1\" class=\"CitationRef\"\u003e1\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e4\u003c/span\u003e\u003c/sup\u003e. The origin and early development of Chinese civilization were also spurred by the interaction between these two types of agricultural economies, a phenomenon unique in world prehistory. It is well-established that broomcorn millet (\u003cem\u003ePanicum miliaceum\u003c/em\u003e) and foxtail millet (\u003cem\u003eSetaria italica\u003c/em\u003e) functioned as the staple crops in the Central Plains since the middle Neolithic Age\u003csup\u003e\u003cspan additionalcitationids=\"CR6\" citationid=\"CR5\" class=\"CitationRef\"\u003e5\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e7\u003c/span\u003e\u003c/sup\u003e. Commencing around 8000 BP, the northward expansion of rice cultivation led to the emergence of a millet-rice mixed farming zone in central China\u003csup\u003e\u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e8\u003c/span\u003e, \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e9\u003c/span\u003e\u003c/sup\u003e, with the Central Plains located at its northern periphery\u003csup\u003e\u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e10\u003c/span\u003e, \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e11\u003c/span\u003e\u003c/sup\u003e.\u003c/p\u003e \u003cp\u003eDuring the Longshan culture period (approximately 4400 BP to 3800 BP), East Asia became integrated into the broader Bronze Age exchange networks\u003csup\u003e\u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e12\u003c/span\u003e\u003c/sup\u003e, especially through its involvement in prehistoric Food Globalization events\u003csup\u003e\u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e13\u003c/span\u003e\u003c/sup\u003e. This process was characterized by the eastward dissemination of multiple domesticated species, such as wheat (\u003cem\u003eTriticum aestivum\u003c/em\u003e), sheep (\u003cem\u003eOvis aries\u003c/em\u003e), and cattle (\u003cem\u003eBos taurus)\u003c/em\u003e, from western Eurasia, which were subsequently incorporated into the subsistence strategies of various regions, particularly the Central Plains. The archaeological evidence indicates a significant increase in both the frequencies and absolute quantities of cereal grains and domesticated animal remains during the Longshan culture period, reflecting a substantial shift towards a more diversified and intensified agricultural economy\u003csup\u003e\u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e14\u003c/span\u003e\u003c/sup\u003e.\u003c/p\u003e \u003cp\u003eThe transformation of subsistence regimes was paralleled by substantial changes in material culture and sociopolitical systems, particularly in the Pan Mount Song Area (PMSA), the core region of the Central Plains. Comprehensive archaeological surveys in the Luoyang Basin, situated in the northern part of PMSA, have revealed a remarkable transition. This transition was from a decentralized and competitive settlement pattern, characterized by several subgroups with a three-tiered hierarchy during the Longshan culture period, to a more complex four-tiered settlement hierarchy. This new structure reached its peak in a large-scale state-level society centered on the ErLiTou site during the Erlitou culture period (ca. 3800 BP to 3500 BP), which is argued to be the onset of the Bronze Age in Chinese Archaeology\u003csup\u003e\u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e15\u003c/span\u003e, \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e16\u003c/span\u003e\u003c/sup\u003e. This socioeconomic transformation has attracted considerable attention from archaeologists and historians, and is often associated with the origin of the legendary Xia dynasty in Chinese history. Although the diversified and intensified agricultural economy undoubtedly increased food provisions to support population growth, the degree of its association with material culture and social dynamics remains a, matter of contention.\u003c/p\u003e \u003cp\u003eA comprehensive exploration from the paleodietary perspective, integrating archaeobotany and zooarchaeology, and concentrating on the diachronic variations of subsistence practices in prehistory of PMSA, especially from the Longshan culture period to the early Bronze Age, is crucial for comprehending the aforementioned issues. This study presents novel stable carbon and nitrogen isotope data from human (n\u0026thinsp;=\u0026thinsp;17) and animal (n\u0026thinsp;=\u0026thinsp;47) remains dating from the late 3rd to the middle 2nd millennium BC at the MeiShan site, located in the southern region of the PMSA. Furthermore, these findings are combined with previously published isotopic data, as well as archaeobotanical and zooarchaeological research, from the late Neolithic to early Bronze Ages, enabling a more robust and comprehensive analysis of subsistence transitions as well as dietary differentiation across the PMSA region. This integrated approach aims to offer an alternative interpretation for interpreting the complex interconnections between subsistence economy transformations, cultural evolution, and social structural changes, thereby enhancing our understanding of these critical processes in prehistoric societies.\u003c/p\u003e\n\u003ch3\u003eSubsistence in the PMSA\u003c/h3\u003e\n\u003cp\u003ePrevious archaeobotanical research has indicated that the assemblages of charred seeds at late Longshan sites in the PMSA were primarily composed of foxtail and broomcorn millets, with rice and soybean functioning as supplementary crops. Only a limited number of charred wheat remains have been identified at several Longshan sites, including WangGeDang\u003csup\u003e\u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e17\u003c/span\u003e\u003c/sup\u003e, WaDian\u003csup\u003e\u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e18\u003c/span\u003e, \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e19\u003c/span\u003e\u003c/sup\u003e, WangChengGang\u003csup\u003e\u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e20\u003c/span\u003e\u003c/sup\u003e. It is crucial to emphasize that all currently available absolute dates for charred wheat remains in the PMSA postdate the Longshan culture period\u003csup\u003e\u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e21\u003c/span\u003e\u003c/sup\u003e. Whether wheat was actually utilized by the inhabitants during the Longshan culture period in this region remains undetermined. During the Erlitou culture period and the subsequent Shang dynasty, which includes the Early Shang (Erligang culture period, ca. ~3500\u0026thinsp;\u0026minus;\u0026thinsp;3300 BP) and Late Shang (Yinxu period, ca. 3300\u0026thinsp;\u0026minus;\u0026thinsp;3000 BP), archaeological evidence reveals a gradual yet consistent increase in both the quantity and ubiquity of wheat\u003csup\u003e\u003cspan additionalcitationids=\"CR23 CR24 CR25 CR26 CR27\" citationid=\"CR22\" class=\"CitationRef\"\u003e22\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e28\u003c/span\u003e\u003c/sup\u003e. The increasing significance of soybean and rice, along with the adoption of wheat, has led to hypothesis that a multiple cropping regime was established during the Longshan culture period and continued to be strengthened throughout the early Bronze Age\u003csup\u003e\u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e20\u003c/span\u003e\u003c/sup\u003e. Zooarchaeological evidence indicates that sheep and cattle husbandry was integrated into the agricultural systems alongside local domesticated pigs and dogs during the Longshan culture period in the Central Plains \u003csup\u003e\u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e14\u003c/span\u003e\u003c/sup\u003e. The diversification of domesticated animal species provided communities with a broader range of animal products and facilitated the secondary exploitation of animals for wool, traction, and other purposes during the late Neolithic and Bronze Ages\u003csup\u003e\u003cspan additionalcitationids=\"CR30\" citationid=\"CR29\" class=\"CitationRef\"\u003e29\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e31\u003c/span\u003e\u003c/sup\u003e.\u003c/p\u003e \u003cp\u003eArchaeobotanical and zooarchaeological investigations provide essential insights into human subsistence strategies in PMSA, while stable isotope analysis offer a method to assess the relative contributions of different food production techniques to human diet. Recently published δ\u003csup\u003e13\u003c/sup\u003eC and δ\u003csup\u003e15\u003c/sup\u003eN values have revealed that the inhabitants of Longshan settlements, such as WaDian and HaoJiaTai\u003csup\u003e\u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e32\u003c/span\u003e, \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e33\u003c/span\u003e\u003c/sup\u003e, can be classified into at least two distinct dietary groups based on the proportion of millet- versus rice-derived nutrients in their diets. During the subsequent Erlitou culture period and the Shang dynasty, isotopic evidence indicates a predominant focus on C\u003csub\u003e4\u003c/sub\u003e-based foods at sites including XinZhai (ca. 3800\u0026thinsp;\u0026minus;\u0026thinsp;3700 BP)\u003csup\u003e\u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e34\u003c/span\u003e, \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e35\u003c/span\u003e\u003c/sup\u003e, NanWa (ca. 3800-3000BP)\u003csup\u003e\u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e36\u003c/span\u003e\u003c/sup\u003e, and Xiaoshuangqiao (ca. 3350\u0026thinsp;\u0026minus;\u0026thinsp;3210 BP)\u003csup\u003e\u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e37\u003c/span\u003e\u003c/sup\u003e, with the exception of some outliers showing relatively negative δ\u003csup\u003e13\u003c/sup\u003eC values at ErLiTou\u003csup\u003e\u003cspan additionalcitationids=\"CR39\" citationid=\"CR38\" class=\"CitationRef\"\u003e38\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e40\u003c/span\u003e\u003c/sup\u003e and XiaoHuCun location at the ZhengZhou ShangCheng site (ca. 3300\u0026thinsp;\u0026minus;\u0026thinsp;3000 BP)\u003csup\u003e\u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e41\u003c/span\u003e\u003c/sup\u003e. Regarding animal husbandry practices, multiple studies have demonstrated that millet farming byproducts served as the primary fodder for both omnivorous (pigs and dogs) and herbivorous animals (cattle and some sheep) across nearly all PMSA sites from the Longshan culture to the Shang dynasty\u003csup\u003e\u003cspan additionalcitationids=\"CR33 CR34\" citationid=\"CR32\" class=\"CitationRef\"\u003e32\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e35\u003c/span\u003e, \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e38\u003c/span\u003e, \u003cspan additionalcitationids=\"CR41 CR42\" citationid=\"CR40\" class=\"CitationRef\"\u003e40\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e43\u003c/span\u003e\u003c/sup\u003e. This consistent pattern of animal feeding practices illustrates the integral role of millet agriculture in supporting diverse animal husbandry strategies throughout the region's subsistence development in the Neolithic and Bronze Ages.\u003c/p\u003e \u003cp\u003eSubstantially more research efforts are still needed to comprehensively document the transition process of dietary complexity in prehistory of the PMSA, particularly during the critical period of early state formation from Neolithic to Bronze Ages. To address these research questions, a dual approach integrating both detailed case studies and broader regional comparisons is indispensable.\u003c/p\u003e \u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003eArchaeological background\u003c/h2\u003e \u003cp\u003eThe study site, MeiShan, is situated in RuZhou city, Henan province, on the left terrace of the Xi'Er River, a minor tributary of the BeiRu River within the upper Ying River Valley of the PMSA (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). Archaeological surveys have delineated an area of approximately 20 hectares, which was predominantly occupied from the Longshan culture through the late Shang or Yinxu periods. In 1995, an excavation spanning ca. 1200 m\u003csup\u003e2\u003c/sup\u003e unearthed dozens of trash pits and several house foundations dating back to the Longshan culture, Erlitou culture, and late Shang periods. Additionally, six earth-pit tombs from the Longshan culture were discovered as well. Typological examination of pottery indicates that the Longshan Culture period remains predominantly mirror the Wangwan Phase III cultural tradition, while also integrating non-local elements similar to the Qujialing-Shijiahe cultures of the middle Yangtze River catchment area in the south and the Dawenkou-Longshan cultures of Shandong Province in the east\u003csup\u003e\u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e44\u003c/span\u003e\u003c/sup\u003e.\u003c/p\u003e \u003cp\u003eDuring the 1995 field season, numerous fragmented human skeleton remains were identified within diverse contexts, such as strata and trash pits, etc., frequently intermingled with animal bones and artifacts\u0026mdash;a common phenomenon during the Longshan and Erlitou culture periods in the Central Plains. Zooarchaeological investigations at the MeiShan site revealed a diverse faunal assemblage comprising both domesticated and wild species. The domesticates included pigs, dogs, sheep, and cattle, while the wild fauna consisted of deer, porcupine, and various bird species. Quantitative analysis of the faunal remains showed that pig bones constituted the majority of the assemblage, accounting for approximately 53%, followed by deer (31%), dogs (6%), sheep (5%), and cattle (2.5%).\u003c/p\u003e \u003cp\u003eStable carbon and nitrogen isotope data of 4 individuals from late Longshan tombs have been published, showing a dichotomy in human diet at MeiShan. While three individuals exhibited diets dominated by \u003csup\u003e15\u003c/sup\u003eN-enriched C\u003csub\u003e3\u003c/sub\u003e food resources, the remaining individuals primarily relied on C\u003csub\u003e4\u003c/sub\u003e resources\u003csup\u003e\u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e45\u003c/span\u003e\u003c/sup\u003e. In this study, seventeen human bones from strata, trash pits of different periods were sampled for δ\u003csup\u003e13\u003c/sup\u003eC and δ\u003csup\u003e15\u003c/sup\u003eN measurement, allowing a paleodietary comparison of different contexts, prior to comprehensive analysis of subsistence transitions across the PMSA region. To establish a reliable isotopic baseline and investigate domesticated animal husbandry practices from the late Neolithic to the early Bronze Ages, both domesticated and wild animal bones were selected for stable carbon and nitrogen isotope analysis. The animal samples included 20 pigs, 5 dogs, 5 cattle, 8 sheep, and 8 deer. The majority of these samples were dated to the Longshan culture period, with the remaining specimens originating from contexts associated with the Erlitou culture and Shang periods, as detailed in Table\u0026nbsp;1.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e"},{"header":"Results","content":"\u003cp\u003eThe collagen yields, as well as carbon and nitrogen contents of the sampled bones from MeiShan, are summarized in Table \u003cspan refid=\"MOESM1\" class=\"InternalRef\"\u003eS1\u003c/span\u003e. The majority of the samples were well-preserved, with collagen yields ranging from 1.1% to 12.3%. Moreover, these samples yielded high quality collagen suitable for radiocarbon dating and dietary reconstruction\u003csup\u003e\u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e46\u003c/span\u003e\u003c/sup\u003e, with carbon contents ranging from 24.1% to 45.2% and nitrogen contents from 8.8% to 16.4%, and the atomic C:N ratios fell within the range of 3.1 to 3.3.\u003c/p\u003e \u003cp\u003eThe radiocarbon dating results are presented in Table \u003cspan refid=\"MOESM2\" class=\"InternalRef\"\u003eS2\u003c/span\u003e. The five samples ascribed to the Longshan culture period are concentrated within the range of ca. 4240 to 3720 BP, which is consistent with the chronological framework of the Wangwan Phase III Longshan culture. In contrast, the two samples from the Erlitou culture period date to ca. 3560 to 3390 BP, corresponding to the final stage of the Erlitou culture or slightly later.\u003c/p\u003e \u003cp\u003eThe isotopic results of human and animal bone collagen are summarized in Table \u003cspan refid=\"MOESM1\" class=\"InternalRef\"\u003eS1\u003c/span\u003e and plotted in Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e. The isotopic values of animal bones exhibited a broad range, reflecting diverse dietary patterns. Deer displayed mean δ\u003csup\u003e13\u003c/sup\u003eC (-18.4\u0026thinsp;\u0026plusmn;\u0026thinsp;1.7\u0026permil;) and δ\u003csup\u003e15\u003c/sup\u003eN (3.8\u0026thinsp;\u0026plusmn;\u0026thinsp;0.8\u0026permil;) values, indicative of a C\u003csub\u003e3\u003c/sub\u003e plant-based diet. In contrast, domesticated animals exhibited higher δ\u003csup\u003e13\u003c/sup\u003eC values (ranging from \u0026minus;\u0026thinsp;18.9\u0026permil; to -6.0\u0026permil;), with mean δ\u003csup\u003e13\u003c/sup\u003eC values for pigs, dogs, cattle, and sheep being \u0026minus;\u0026thinsp;11.7\u0026thinsp;\u0026plusmn;\u0026thinsp;3.6\u0026permil;, -9.3\u0026thinsp;\u0026plusmn;\u0026thinsp;0.8\u0026permil;, -10.2\u0026thinsp;\u0026plusmn;\u0026thinsp;2.5\u0026permil;, and \u0026minus;\u0026thinsp;16.6\u0026thinsp;\u0026plusmn;\u0026thinsp;2.6\u0026permil;, respectively, suggesting varying degrees of C\u003csub\u003e4\u003c/sub\u003e fodder consumption. The mean δ\u003csup\u003e15\u003c/sup\u003eN values for dogs (7.4\u0026thinsp;\u0026plusmn;\u0026thinsp;0.7\u0026permil;), pigs (7.5\u0026thinsp;\u0026plusmn;\u0026thinsp;1.0\u0026permil;), cattle (6.8\u0026thinsp;\u0026plusmn;\u0026thinsp;2.4\u0026permil;), and sheep (3.9\u0026thinsp;\u0026plusmn;\u0026thinsp;1.1\u0026permil;) were all elevated in comparison to those of deer.\u003c/p\u003e \u003cp\u003eThe MeiShan inhabitants exhibited considerable variation in δ\u003csup\u003e13\u003c/sup\u003eC (-16.9\u0026permil; to -6.8\u0026permil;) and δ\u003csup\u003e15\u003c/sup\u003eN (7.5\u0026permil; to 12.3\u0026permil;) values, indicating dietary diversity. Their consumption of C\u003csub\u003e4\u003c/sub\u003e versus C\u003csub\u003e3\u003c/sub\u003e foods likely derived primarily from millet foods (i.e. millet grains and domesticated animals raised with millet byproducts) or rice and related food products. Among the 13 Longshan individuals, three displayed δ\u003csup\u003e13\u003c/sup\u003eC values no more positive than \u0026minus;\u0026thinsp;14.1\u0026permil; and δ\u003csup\u003e15\u003c/sup\u003eN values no less than 10.8\u0026permil;, while the remaining individuals exhibited less negative δ\u003csup\u003e13\u003c/sup\u003eC values (-13.4\u0026permil; to -6.8\u0026permil;) and lower δ\u003csup\u003e15\u003c/sup\u003eN values (7.5\u0026permil; to 9.5\u0026permil;). This pattern suggests differential access to food resources within the MeiShan community during the Longshan culture period. In contrast, the isotopic values (δ\u003csup\u003e13\u003c/sup\u003eC range = -10.9\u0026permil; to -7.1\u0026permil;, δ\u003csup\u003e15\u003c/sup\u003eN range\u0026thinsp;=\u0026thinsp;8.5\u0026permil; to 9.5\u0026permil;) indicate a primary reliance on millet-based nutrients with only minor supplementation from C\u003csub\u003e3\u003c/sub\u003e foods such as rice. The individual labelled MS01, lacking chronological context, displayed δ\u003csup\u003e13\u003c/sup\u003eC and δ\u003csup\u003e15\u003c/sup\u003eN values of -9.5\u0026permil; and 8.1\u0026permil;, respectively, suggesting a diet predominantly based on millet farming.\u003c/p\u003e"},{"header":"Discussion","content":"\u003cdiv id=\"Sec6\" class=\"Section2\"\u003e \u003ch2\u003eAnimal husbandry practices\u003c/h2\u003e \u003cp\u003eIn prehistoric farming communities of the Central Plains and adjacent regions, pigs and dogs, which are typical indigenous domesticates in East Asia, often exhibited comparable isotopic patterns. This suggests that their diets were similar, predominantly characterized by C\u003csub\u003e4\u003c/sub\u003e resources, such as millet byproducts, human leftovers, or/and even millet grains\u003csup\u003e\u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e35\u003c/span\u003e, \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e47\u003c/span\u003e\u003c/sup\u003e. The isotopic data (δ\u003csup\u003e13\u003c/sup\u003eC ranging from \u0026minus;\u0026thinsp;10.3\u0026permil; to -8.4\u0026permil;, δ\u003csup\u003e15\u003c/sup\u003eN values ranging from 6.3\u0026permil; to 8.3\u0026permil;, n\u0026thinsp;=\u0026thinsp;5) indicate that this dietary pattern likely applied to dogs at MeiShan from the Longshan culture to the YinXu periods. In contrast, pigs of the Longshan culture period at MeiShan displayed a wider range of δ\u003csup\u003e13\u003c/sup\u003eC values (-18.2\u0026permil; to -6.0\u0026permil;), with a relatively negative mean value (-12.0\u0026thinsp;\u0026plusmn;\u0026thinsp;3.8\u0026permil;, n\u0026thinsp;=\u0026thinsp;14). This implies a mixed C\u003csub\u003e3\u003c/sub\u003e/C\u003csub\u003e4\u003c/sub\u003e fodder and more flexible feeding practices. Similar isotopic data have also been identified at contemporaneous sites like WaDian\u003csup\u003e\u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e32\u003c/span\u003e\u003c/sup\u003e and XinZhai in the PMSA\u003csup\u003e\u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e34\u003c/span\u003e, \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e35\u003c/span\u003e\u003c/sup\u003e, where considerable amounts of rice remains were discovered alongside charred millets\u003csup\u003e\u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e18\u003c/span\u003e, \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e19\u003c/span\u003e, \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e22\u003c/span\u003e\u003c/sup\u003e. This has given rise to the hypothesis that C\u003csub\u003e3\u003c/sub\u003e fodder, presumably derived from rice, was employed for pig rearing at these sites, a practice which might have also occurred at MeiShan. During the Erlitou culture period, the mean δ\u003csup\u003e13\u003c/sup\u003eC value of pigs (-10.8\u0026thinsp;\u0026plusmn;\u0026thinsp;3.1\u0026permil;, n\u0026thinsp;=\u0026thinsp;5) increased slightly, ranging from \u0026minus;\u0026thinsp;14.0\u0026permil; to -6.8\u0026permil;, while δ\u003csup\u003e15\u003c/sup\u003eN values remained relatively stable (7.5\u0026thinsp;\u0026plusmn;\u0026thinsp;0.5\u0026permil;, ranging: 6.6\u0026permil; to 7.9\u0026permil;), indicating a decreased reliance on rice in pig diets. This trend appears to have been prevalent in the PMSA between approximately 4000 and 3500 BP, as pigs at WangGeDang, ErLiTou\u003csup\u003e\u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e40\u003c/span\u003e\u003c/sup\u003e and XinZhai\u003csup\u003e\u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e34\u003c/span\u003e, \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e35\u003c/span\u003e\u003c/sup\u003e exhibiting similar millet-dominated diet patterns.\u003c/p\u003e \u003cp\u003eThe introduction of domesticated sheep and cattle to the Central Plains can be traced back to no later than the Longshan period, as evidenced by the extensive distribution of their remains\u003csup\u003e\u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e14\u003c/span\u003e\u003c/sup\u003e. Sheep exhibited δ\u003csup\u003e15\u003c/sup\u003eN patterns (3.9\u0026thinsp;\u0026plusmn;\u0026thinsp;1.1\u0026permil;) similar to those of deer, while their δ\u003csup\u003e13\u003c/sup\u003eC values were more positive (-16.6\u0026thinsp;\u0026plusmn;\u0026thinsp;2.6\u0026permil;, range: -18.9\u0026permil; to -11.6\u0026permil;). The isotopic pattern has been observed at numerous Longshan culture period and Bronze Age sites in the Central Plains, and is attributed to the consumption of millet byproducts, particularly during winter\u003csup\u003e\u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e32\u003c/span\u003e, \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e47\u003c/span\u003e, \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e48\u003c/span\u003e\u003c/sup\u003e. At MeiShan, the five sheep from the Longshan culture period displayed δ\u003csup\u003e13\u003c/sup\u003eC values ranging from \u0026minus;\u0026thinsp;18.6\u0026permil; to -14.9\u0026permil;, while some sheep from the Erlitou culture (#MS48, -11.6\u0026permil;) and late Shang (#MS49, -15.0\u0026permil;) periods exhibited less negative δ\u003csup\u003e13\u003c/sup\u003eC values, indicating an increased intake of millet straws. All MeiShan cattle, including one from the Longshan and three from the Erlitou culture periods, exhibited elevated δ\u003csup\u003e13\u003c/sup\u003eC values (-11.7\u0026permil; to -7.0\u0026permil;), reflecting a focus on millet-related fodder. This pattern is consistent with cattle husbandry practices in other PMSA agricultural communities during the Longshan and Erlitou culture periods, indicating that cattle husbandry became increasingly dependent on millet farming\u003csup\u003e\u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e32\u003c/span\u003e, \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e47\u003c/span\u003e, \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e48\u003c/span\u003e\u003c/sup\u003e.\u003c/p\u003e \u003cp\u003eThe utilization of sheep and cattle offered novel opportunities for diversifying livestock husbandry practices. This was not only manifested in the provision of protein, wool, hides, bone materials, and traction power, but also in a more efficient utilization of agricultural byproducts, and grasses and stubble from fallow and marginal lands. As a result, the rearing of these ruminants contributed to a more productive land use model and agricultural system. From this viewpoint, it is contended that ruminant husbandry, especially cattle raising, had a positive influence on and became an essential part of the agricultural system as a supplementary subsistence practice since the end of the 3rd millennium BC.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e\n\u003ch3\u003eHuman diet complexity at MeiShan and contemporary sites in the PSMA\u003c/h3\u003e\n\u003cp\u003eAlthough there is a dearth of archaeobotanical data from MeiShan, flotation investigations at adjacent sites within the PMSA, dating from roughly 4400 BP to 3400 BP or later, have indicated that foxtail millet was the predominant constituent of the charred grain assemblages, with rice occupying the second or third position\u003csup\u003e\u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e49\u003c/span\u003e\u003c/sup\u003e. Consequently, it is hypothesized that rice cultivation significantly contributed to the subsistence economy and transformed the local cropping system during this period, which had previously been an agricultural regime primarily centered around millet\u003csup\u003e\u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e27\u003c/span\u003e, \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e50\u003c/span\u003e, \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e51\u003c/span\u003e\u003c/sup\u003e. Nevertheless, the role of rice farming in human subsistence and whether disparities in rice consumption existed among sites and individuals remain unresolved queries. Our study offers novel perspectives on these issues.\u003c/p\u003e \u003cp\u003eThe isotopic results of 13 Longshan human samples from our study, combined with previously published data from 4 individuals, are discussed together to understand human diet and subsistence at MeiShan. Overall, the 17 individuals exhibited a wide spectrum of values, ranging from \u0026minus;\u0026thinsp;16.9\u0026permil; to -6.8\u0026permil; for δ\u003csup\u003e13\u003c/sup\u003eC and from 7.5\u0026permil; to 12.3\u0026permil; for δ\u003csup\u003e15\u003c/sup\u003eN, indicating a diverse intake among the subjects. While the majority of individuals followed a mixed C\u003csub\u003e3\u003c/sub\u003e/C\u003csub\u003e4\u003c/sub\u003e diet, some depended solely on either C\u003csub\u003e3\u003c/sub\u003e (rice-derived) or C\u003csub\u003e4\u003c/sub\u003e (millet-derived) nutrients, as shown by their δ\u003csup\u003e13\u003c/sup\u003eC values. The mean δ\u003csup\u003e15\u003c/sup\u003eN (9.6\u0026thinsp;\u0026plusmn;\u0026thinsp;1.4\u0026permil;, n\u0026thinsp;=\u0026thinsp;17) of these individuals was significantly higher than that of contemporary animals, including omnivorous pigs (7.5\u0026thinsp;\u0026plusmn;\u0026thinsp;1.2\u0026permil;, n\u0026thinsp;=\u0026thinsp;14) and dogs (7.5\u0026thinsp;\u0026plusmn;\u0026thinsp;1.0\u0026permil;, n\u0026thinsp;=\u0026thinsp;3), domesticated herbivorous cattle (5.5\u0026permil;, n\u0026thinsp;=\u0026thinsp;1) and sheep (3.7\u0026thinsp;\u0026plusmn;\u0026thinsp;0.8\u0026permil;, n\u0026thinsp;=\u0026thinsp;5), as well as deer (3.4\u0026thinsp;\u0026plusmn;\u0026thinsp;0.3\u0026permil;, n\u0026thinsp;=\u0026thinsp;4). Given that the δ\u003csup\u003e15\u003c/sup\u003eN range is 7.5\u0026permil; to 12.3\u0026permil;, some individuals with δ\u003csup\u003e15\u003c/sup\u003eN values comparable to those of domesticated omnivores might have had limited access to food resources with higher δ\u003csup\u003e15\u003c/sup\u003eN values, while others probably consumed more δ\u003csup\u003e15\u003c/sup\u003eN-enriched nutrients. The dietary diversity, as reflected by δ\u003csup\u003e15\u003c/sup\u003eN values, is primarily attributed to disparities in human trophic levels \u003csup\u003e\u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e52\u003c/span\u003e\u003c/sup\u003e, which might have resulted from differential access to animal products among individuals.\u003c/p\u003e \u003cp\u003eA previous study at WaDian identified two distinct dietary groups based on stable carbon and nitrogen isotope analysis. One group primarily consumed millet with some amount of domesticated animal protein, while the other exhibited a mixed C\u003csub\u003e3\u003c/sub\u003e/C\u003csub\u003e4\u003c/sub\u003e diet enriched in \u003csup\u003e15\u003c/sup\u003eN\u003csup\u003e\u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e32\u003c/span\u003e\u003c/sup\u003e. This finding has been corroborated by recent publications on WaDian, HaoJiaTai, MeiShan, Jiazhuang, and other Longshan culture sites\u003csup\u003e\u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e33\u003c/span\u003e, \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e45\u003c/span\u003e\u003c/sup\u003e. Zhou observed that elites at MeiShan and JiaZhuang, whose grave goods exhibited southern stylistic influences, primarily consumed rice-derived foods. These findings provide evidence for diffusion and popularization in PMSA of southern-oriented rice farming and materials. The high proportion of rice-derived foods in the diets of elites during the Longshan culture period raises questions about whether rice was a highly valued staple crop and whether access to rice was linked to social status.\u003c/p\u003e \u003cp\u003eWhile attributing δ\u003csup\u003e15\u003c/sup\u003eN enrichment to social disparities between rice-eaters and millet-eaters offers a straightforward explanation, it may not fully account for other potential factors. A pilot study on plant isotopes at XinZhai, WaDian, ChengYao, WangChengGang and other contemporary sites in PMSA indicates that rice exhibits higher δ\u003csup\u003e15\u003c/sup\u003eN values than millets, possibly due to nitrogen metabolism in paddy fields\u003csup\u003e\u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e53\u003c/span\u003e\u003c/sup\u003e. Fox example, at XinZhai, the mean δ\u003csup\u003e15\u003c/sup\u003eN values of C\u003csub\u003e4\u003c/sub\u003e millet and C\u003csub\u003e3\u003c/sub\u003e rice grains are 4.4\u0026thinsp;\u0026plusmn;\u0026thinsp;1.0\u0026permil; (n\u0026thinsp;=\u0026thinsp;15) and 8.0\u0026thinsp;\u0026plusmn;\u0026thinsp;1.4\u0026permil; (n\u0026thinsp;=\u0026thinsp;10), respectively, highlighting the disparity in δ\u003csup\u003e15\u003c/sup\u003eN between the two staple crops. Consequently, it remains unclear whether the lower δ\u003csup\u003e15\u003c/sup\u003eN values of millet-eaters reflect a more vegetarian diet. A linear correlation test between human δ\u003csup\u003e13\u003c/sup\u003eC and δ\u003csup\u003e15\u003c/sup\u003eN values at MeiShan provides further insights into human dietary preferences and potential causal factors. The correlation of δ\u003csup\u003e13\u003c/sup\u003eC and δ\u003csup\u003e15\u003c/sup\u003eN for all 17 Longshan culture residents at MeiShan (n\u0026thinsp;=\u0026thinsp;17, r\u003csup\u003e2\u003c/sup\u003e\u0026thinsp;=\u0026thinsp;0.69) suggests that individuals with a higher intake of \u003csup\u003e13\u003c/sup\u003eC-depleted or rice-derived nutrients exhibited elevated bone collagen δ\u003csup\u003e15\u003c/sup\u003eN values. Furthermore, the δ\u003csup\u003e13\u003c/sup\u003eC and δ\u003csup\u003e15\u003c/sup\u003eN values of MeiShan pigs showed a significant correlation as well (r\u003csup\u003e2\u003c/sup\u003e\u0026thinsp;=\u0026thinsp;0.62, n\u0026thinsp;=\u0026thinsp;20), indicating that the dietary intake of rice products by pigs does not necessarily imply a higher consumption of animal-derived materials. Instead, the correlation is probably related to the difference of δ\u003csup\u003e15\u003c/sup\u003eN values between rice and millet-related fodders. We argue consequently that the correlation among δ\u003csup\u003e13\u003c/sup\u003eC and δ\u003csup\u003e15\u003c/sup\u003eN values of MeiShan residents likely reflects differences in the consumption of millet versus rice.\u003c/p\u003e \u003cp\u003eA comprehensive investigation integrating isotopic data and social status at WaDian indicated no distinct hierarchical disparities between millet-eaters and mixed C\u003csub\u003e3\u003c/sub\u003e/C\u003csub\u003e4\u003c/sub\u003e diet consumers. Instead, it was hypothesized that differences in social status existed within each group\u003csup\u003e\u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e33\u003c/span\u003e\u003c/sup\u003e. As previously noted, individuals with burial customs associated with the Shijiahe culture in the southern region typically presented more negative δ\u003csup\u003e13\u003c/sup\u003eC values at MeiShan and contemporary sites. This suggests that rice farming might have played a more important role in their subsistence economy, analogous to the communities of the Shijiahe culture. Nevertheless, more evidence is required to conclude that rice and rice-derived foods were highly valued during the Longshan culture period in the PMSA in the future.\u003c/p\u003e \u003cp\u003eComparatively speaking, the three MeiShan residents in the Erlitou culture period exhibited more clustered δ\u003csup\u003e13\u003c/sup\u003eC values (ranging from \u0026minus;\u0026thinsp;10.9\u0026permil; to -7.1\u0026permil;), indicating a focus on millet-derived nutrients supplemented with only a minor supplements of C\u003csub\u003e3\u003c/sub\u003e foods. Their δ\u003csup\u003e15\u003c/sup\u003eN values (ranging from 8.5\u0026permil; to 9.5\u0026permil;) exceeded those of pigs (7.5\u0026thinsp;\u0026plusmn;\u0026thinsp;0.5\u0026permil;, n\u0026thinsp;=\u0026thinsp;5), dogs (7.1\u0026permil;, 7.6\u0026permil;, n\u0026thinsp;=\u0026thinsp;2), cattle (7.2\u0026thinsp;\u0026plusmn;\u0026thinsp;2.7\u0026permil;, n\u0026thinsp;=\u0026thinsp;4) and sheep (2.6\u0026permil;, 5.9\u0026permil; n\u0026thinsp;=\u0026thinsp;2). This suggests that MeiShan residents during the Longshan culture period experienced diverse diets and lifestyles, while their successors in the Erlitou culture period relied more heavily on millets and millet feeding domesticates.\u003c/p\u003e \u003cdiv id=\"Sec8\" class=\"Section2\"\u003e \u003ch2\u003eSubsistence shift from the Neolithic to the early Bronze Age in PMSA\u003c/h2\u003e \u003cp\u003eTo gain a more comprehensive understanding of the development of subsistence economies in the PMSA from the Neolithic to the Bronze Age, we have systematically compiled and chronologically plotted the published δ\u003csup\u003e13\u003c/sup\u003eC and δ\u003csup\u003e15\u003c/sup\u003eN values of humans (Table \u003cspan refid=\"MOESM3\" class=\"InternalRef\"\u003eS3\u003c/span\u003e) in Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e. Isotopic analysis results indicate that millet-based foods constituted the primary component of the food resources for ancient peoples in the PMSA region from the Yangshao culture to the late Shang periods. Meanwhile, during certain periods, the dietary contribution of C\u003csub\u003e3\u003c/sub\u003e foods increased markedly among a small number of inhabitants. The dietary patterns largely reflect the diachronic developmental trends of the subsistence economy in the PMSA region.\u003c/p\u003e \u003cp\u003eArchaeobotanical evidence from Peiligang culture sites such as Peiligang\u003csup\u003e\u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e54\u003c/span\u003e\u003c/sup\u003e and Zhuzhai\u003csup\u003e\u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e8\u003c/span\u003e\u003c/sup\u003e indicates that millets and rice were cultivated in the PMSA as early as 8000 BP to 7500 BP. During the subsequent Yangshao culture period, the ubiquity and quantities of foxtail and broomcorn millets increased significantly as the primary crops, while rice cultivation served as the supplement farming practice\u003csup\u003e\u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e44\u003c/span\u003e, \u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e55\u003c/span\u003e\u003c/sup\u003e. By the Longshan culture period, the agricultural economy shifted from a focus on millet farming to a more complex system incorporating the cultivation of both millet and rice. The shift is evident at WaDian, HaoJiaTai, and MeiShan\u003csup\u003e\u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e32\u003c/span\u003e, \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e33\u003c/span\u003e\u003c/sup\u003e, where two distinct dietary groups were isotopically identified; one was reliant on millet, and the other consumed more rice-derived nutrients. The successful integration of diverse dietary and subsistence groups was crucial for the stability and subsequent development of early societies, which was a key factor in the formation of a unified early Chinese civilization.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eCompared to the Longshan culture period, the δ\u003csup\u003e13\u003c/sup\u003eC and δ\u003csup\u003e15\u003c/sup\u003eN isotope ratios of the Erlitou culture population in the PMSA were more concentrated, reflecting a greater reliance on millet-based agricultural products and a reduced contribution of C\u003csub\u003e3\u003c/sub\u003e foods, namely rice. These contracts with the proto-Shang, and late Shang populations at ZhengZhou ShangCheng\u003csup\u003e\u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e41\u003c/span\u003e\u003c/sup\u003e and the Erligang culture population at WangJingLou\u003csup\u003e\u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e37\u003c/span\u003e\u003c/sup\u003e, where some individuals exhibited more diverse diets with greater contribution from C\u003csub\u003e3\u003c/sub\u003e foods, despite millet farming remaining the dominant agricultural economy.\u003c/p\u003e \u003cp\u003eFrom the Yangshao culture to the late Shang period, the transitions in human diet, subsistence, and social complexity process appear to be chronologically aligned. The role of the agricultural economy in supporting or influencing the social complexity process in the PMSA requires further investigation. During the Yangshao culture period, the rapid development of millet farming in the Central Plains coincided with significant cultural changes, including population growth and the expansion of cultural influence to the surrounding areas. Millet-oriented agricultural economy probably provided the food security necessary for these developments. During the Longshan culture period, deteriorating climatic conditions and frequent floods may have prompted the adoption of rice farming in the Central Plains and surrounding areas, potentially influenced by interactions with archaeological cultures in southern lowlands, such as the Qujialing and Shijiahe cultures, as suggested by ancient DNA analysis\u003csup\u003e\u003cspan citationid=\"CR56\" class=\"CitationRef\"\u003e56\u003c/span\u003e\u003c/sup\u003e and artifacts with southern stylistic influences\u003csup\u003e\u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e9\u003c/span\u003e\u003c/sup\u003e. Our study provides insights into human adaptation, cultural integration, and population interaction in the PSMA during the Longshan culture period, topics that merit further research.\u003c/p\u003e \u003cp\u003eAt the beginning of the Bronze Age, the rise of Erlitou culture (ca. 3800 BP to 3500 BP) unified the Central Plains hinterland and surrounding regions, with its center in the Luoyang Basin of the northern PMSA\u003csup\u003e\u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e57\u003c/span\u003e\u003c/sup\u003e. While the contribution of rice to human diet declined, millets remained highly valued\u003csup\u003e\u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e27\u003c/span\u003e, \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e40\u003c/span\u003e\u003c/sup\u003e. Millet farming not only provided a staple food for ErLiTou communities but also served as feed for pigs, dogs, as well as cattle. The subsequent Erligang culture (ca. 3500 BP to 3350 BP), corresponding to the early period of the Shang Dynasty, centered on ZhengZhou ShangCheng, where some individuals exhibited more diverse diets with greater contributions from C\u003csub\u003e3\u003c/sub\u003e foods, possibly reflecting migration and cultural interactions during the transition from Erlitou to Erligang cultures. When it comes to the middle (ca. 3350 BP to 3210 BP) and late Shang periods (ca. 3210 BP to 3000 BP), dietary variations among individuals still persisted. Given the profound changes in social structure, namely the rise of large-scale state-level society, the implications reflected by dietary variations among different individuals can no longer be adequately explained solely through the lens of subsistence economy.\u003c/p\u003e \u003cp\u003eDuring the Neolithic Age, social division of labor had not yet manifested distinct specialized characteristics, and the majority of inhabitants at Neolithic sites still relied mainly on agricultural production for their livelihoods. Consequently, the proportion of C\u003csub\u003e4\u003c/sub\u003e foods (e.g., foxtail millet, broomcorn millet) to C\u003csub\u003e3\u003c/sub\u003e foods (e.g., rice) in the diets of inhabitants was directly linked to the cultivation scale and yield of different crops. This phenomenon may have been prevalent at the MeiShan site investigated in this study, as well as at other sites dating from the Yangshao to the Longshan culture periods, such as the WangGou, WaDian and HaoJiaTai sites. In contrast, for urban residents, whose livelihoods focused on non-agricultural activities at the capital and regional centers, the ratio of C\u003csub\u003e3\u003c/sub\u003e to C\u003csub\u003e4\u003c/sub\u003e foods in their diets was largely determined by food acquisition channels and distribution mechanisms developed by the early states. At the ZhengZhou ShangCheng, Erlitou and Wangjinglou sites, the capitals or regional centers of the Bronze Age, the dietary contribution of C\u003csub\u003e3\u003c/sub\u003e foods to urban inhabitants increased markedly, and the combination of food types became more diverse. This presumably suggests that they may have originated from subsistence economic zones distinct from those of the main population, possessed more extensive food acquisition networks, or occupied an advantageous position in the distribution of social resources\u0026mdash;allowing them to obtain non-local agricultural products from various regions via trade, tribute and other such means. Such dietary differentiation stemming from disparities in social roles and economic activities provides an important material cultural basis for our understanding of the process of social complexity, patterns of resource distribution, and urban-rural relations in ancient societies.\u003c/p\u003e \u003c/div\u003e"},{"header":"Conclusions","content":"\u003cp\u003eThe PSMA of the core area of the Central Plains of China featured an economic model dominated by millet farming alongside rice cultivation of a certain scale from the Yangshao period to the Early Bronze Age. Roughly from the 3rd millennium to the 2nd millennium BC, the Central Plains underwent a transformation of social structure, evolving from the polycentric settlement pattern of the late Longshan period into early large-scale state-level society with overarching authority. Meanwhile, domesticated cattle and sheep as well as wheat originating from West Asia were also introduced into this region. Although dietary patterns of humans at different sites and in different periods can reflect, to a certain extent, the developmental trends of the subsistence economy in this region, specifically millet farming versus rice farming, the subsistence economic conditions during the formation of early states, particularly the impact of early state formation on the dietary patterns of different populations, still warrant further research. To gain a deeper understanding of these issues, we conducted stable carbon and nitrogen isotope analyses on human and animal bones from the Longshan culture period, the Erlitou culture period, and the late Shang period at the MeiShan site in Ruzhou, Henan province. To facilitate a more rigorous and comprehensive examination of subsistence transitions and dietary differentiation across the PMSA region, the data from the MeiShan site are integrated with previous publications spanning from the late Neolithic to the early Bronze Ages.\u003c/p\u003e \u003cp\u003eIt is revealed that during the Longshan culture period, the dietary pattern of the inhabitants at the MeiShan site showed a strong signal of rice consumption, although millet-related foods were still widely consumed. However, during the Erlitou culture period, the contribution of rice to the human diet notably declined, whereas millet farming continued to be highly emphasized. This shift in agricultural practices was also reflected in the feeding practices of domesticated animals. From the Longshan culture to the Erlitou culture periods, the proportion of rice-related foods in the fodder of domesticated pigs decreased significantly, while millet-related feed increased. By integrating previous research findings, we propose that the subsistence economy in the PSMA, represented by the MeiShan site during the Longshan culture period, was likely influenced by the Shijiahe culture from the middle Yangtze River catchment. Under this influence, the inhabitants of MeiShan and nearby Longshan culture sites such as HaoJiaTai and WaDian consumed a notable amount of rice products. However, the trend toward agricultural diversification did not rapidly progress during the Erlitou culture period or the subsequent Erligang culture and late Shang periods. Instead, millet farming once again became the dominant form of local agricultural economy, with rice and wheat cultivation serving only as supplementary practices.\u003c/p\u003e \u003cp\u003eBy comprehensively examining the isotopic data of the inhabitants at the MeiShan site in conjunction with those from sites in this region spanning the Neolithic to the Early Bronze Ages, and contextualizing these findings within the macro backdrop of social transformation, we argue that the dietary patterns of Neolithic residents from the Yangshao to the Longshan periods likely primarily reflected the subsistence economic characteristics of the respective sites. The changes in the subsistence economy of the PSMA from the late Neolithic to the early Bronze Age were the result of the development of the traditional millet farming oriented agricultural economy in the area. At the same time, these changes were also related to the frequent cultural interactions between the Central Plains and the middle reaches of the Yangtze River, which began during the Longshan culture period. Besides, the dietary variations within the resident groups at early Bronze Age capital sites and regional centers may also have been associated with the social complexity phenomena driven by factors such as individual migration and food resource distribution during the urbanization process.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e \u003ch2\u003eCompeting interests\u003c/h2\u003e \u003cp\u003eThe authors declare no competing interests.\u003c/p\u003e \u003c/p\u003e\u003ch2\u003eAuthor Contribution\u003c/h2\u003e\u003cp\u003eX.C. collected the samples, analyzed the data, prepared all figures and tables, and wrote the manuscript; Y.Y. contributed to the interpretation of the data and revised the manuscript; G.Y. and Y.H. offered valuable suggestions and contributed to the revision of the manuscript.\u003c/p\u003e\u003ch2\u003eAcknowledgement\u003c/h2\u003e \u003cp\u003eWe would like to acknowledge the funding from The National Social Science Fund of China (No. 22BKG047), National Key R\u0026amp;D Program of China ( NO.2022YFF0903500), and Project of Key Laboratory of Archaeological Sciences and Cultural Heritage (No. S20250201).\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\u003cli\u003e\u003cspan\u003eLiu L, Lee GA, Jiang L, Zhang J. 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The tentative research of the spreading of rice in the Central Plains durng prehistoric period. \u003cem\u003eArchaeology in Western China\u003c/em\u003e 15, 211\u0026ndash;223 (2018).\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eHedges REM, Reynard LM. Nitrogen isotopes and the trophic level of humans in archaeology. J Archaeol Sci. 2007;34:1240\u0026ndash;51.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eWang X, Zhao Z, Zhong H, Chen X, Hu Y. Manuring and land exploitation in the Central Plains of late Longshan (2200\u0026ndash;1900 BCE) China: Implications of stable isotopes of archaeobotanical remains. J Archaeol Sci. 2022;148:105691.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eFirst Henan Fieldwork Team of IA-CASS, Zhengzhou Municipal Institute of Cultural Relics and Archaeology. \u0026amp; Xinzheng Municipal Bureau of Culture, Radio, Television, Tourism and Sports The excavation of the Peiligang Site of Xinzheng City, Henan,2018\u0026ndash;2019. Acta Archaeologica Sinica. 2020;4:521\u0026ndash;46.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eZhong H, Zhao Z. The preliminary study of the agricultural production mode of the Central Plains during the late Yangshao period. Agricultural History China. 2023;2:52\u0026ndash;61.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eCui Y, Fang Y, Zhou H. DNA analysis of ancient residents revealed at Longshan period, Wadian site, in Yuzhou, Henan. Huaxia Archaeol, 41\u0026ndash;6 (2019).\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eXu H, Erlitou. The origin of the tradition of non-fortified primary capitals in early China. Archaeol Res Asia. 2018;14:71\u0026ndash;9.\u003c/span\u003e\u003c/li\u003e\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":false,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"npj-heritage-science","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"hsci","sideBox":"Learn more about [Heritage Science](http://heritagesciencejournal.springeropen.com)","snPcode":"40494","submissionUrl":"https://submission.nature.com/new-submission/40494/3","title":"npj Heritage Science","twitterHandle":"@SpringerOpen","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"BMC/SO AJ","inReviewEnabled":true,"inReviewRevisionsEnabled":true},"keywords":"Late Neolithic, Bronze Age, MeiShan, Subsistence, Dietary Differentiation","lastPublishedDoi":"10.21203/rs.3.rs-8751401/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-8751401/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eThe Central Plains, a core region for early Chinese civilization, underwent profound sociocultural transformations during the third and second millennia BC, as well as the intensification and diversification of the agricultural economy. This study explores the subsistence changes through isotopic analysis of human and animal remains from the Pan Mount Song Area (PMSA), the hinterland of the Central Plains. We present new stable carbon and nitrogen isotope data from the MeiShan site in the PMSA, complemented by a comprehensive analysis of published datasets. Findings reveal distinct subsistence patterns across different periods. During the Longshan period (ca. 4400\u0026thinsp;\u0026minus;\u0026thinsp;3800 BP), millet remained the staple crop for both humans and domesticated animals, while some individuals consumed significant amounts of rice, indicating a notable shift in subsistence compared to the Yangshao culture period (ca. 6300\u0026thinsp;\u0026minus;\u0026thinsp;4500 BP), when millet farming was highly emphasized. From the Erlitou culture (ca. 3800\u0026thinsp;\u0026minus;\u0026thinsp;3500 BP) and Erligang culture (ca. 3500\u0026thinsp;\u0026minus;\u0026thinsp;3350 BP) to the late Shang periods (ca. 3210\u0026thinsp;\u0026minus;\u0026thinsp;3000 BP), millet-derived nutrients became predominant. When integrated with the published isotopic data focusing on PMSA, it is found that the dietary patterns of inhabitants from the late Neolithic to early Bronze Ages were predominantly millet-based during most periods. However, a relatively high proportion of C\u003csub\u003e3\u003c/sub\u003e or rice-derived foods could be found in the diets of some residents at several late Longshan sites, as well as the Early Bronze Age capitals and regional centers. The disparities in human dietary patterns at these sites are presumably ascribable to the increasing complexity of the agricultural economy. Furthermore, they may also be associated with human mobility and the disparity in resource allocation during the urbanization process, which are more prone to have exerted substantial impacts at the central sites of the Early Bronze Age.\u003c/p\u003e","manuscriptTitle":"Stable Carbon and Nitrogen Isotope Analysis for subsistence Shift and Dietary Differentiation from the Late Neolithic to the early Bronze Ages in the Central Plains, China","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2026-02-23 16:15:59","doi":"10.21203/rs.3.rs-8751401/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"decision","content":"Revision requested","date":"2026-03-31T14:25:48+00:00","index":"","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2026-03-31T07:28:55+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2026-03-10T11:27:34+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2026-03-03T01:09:49+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"198908155566680001755781410250599843980","date":"2026-02-26T17:38:29+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"169150434685919577434401070962647538182","date":"2026-02-22T11:03:42+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"299298865469574064360576112981511292865","date":"2026-02-20T02:48:38+00:00","index":"hide","fulltext":""},{"type":"reviewersInvited","content":"","date":"2026-02-19T00:39:33+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2026-02-06T04:24:40+00:00","index":"","fulltext":""},{"type":"checksComplete","content":"","date":"2026-02-06T04:23:51+00:00","index":"","fulltext":""},{"type":"submitted","content":"npj Heritage Science","date":"2026-01-31T16:22:26+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"npj-heritage-science","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"hsci","sideBox":"Learn more about [Heritage Science](http://heritagesciencejournal.springeropen.com)","snPcode":"40494","submissionUrl":"https://submission.nature.com/new-submission/40494/3","title":"npj Heritage Science","twitterHandle":"@SpringerOpen","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"BMC/SO AJ","inReviewEnabled":true,"inReviewRevisionsEnabled":true}}],"origin":"","ownerIdentity":"a4e82ae0-30bf-4b57-88b8-1eea743f904e","owner":[],"postedDate":"February 23rd, 2026","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"in-revision","subjectAreas":[],"tags":[],"updatedAt":"2026-03-31T14:39:57+00:00","versionOfRecord":[],"versionCreatedAt":"2026-02-23 16:15:59","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-8751401","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-8751401","identity":"rs-8751401","version":["v1"]},"buildId":"XKTyCvWXoU3ODBz1xrDgd","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}