To eat like Liangzhu: isotopic investigation of diets in the Lower Yangtze area prior to and during the Liangzhu period (5300-4300 cal. BP)

To eat like Liangzhu: isotopic investigation of diets in the Lower Yangtze area prior to and during the Liangzhu period (5300-4300 cal. 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BP) Pengfei Sheng, Edward Allen, Xiang Huang, Xiuwen Zheng, Michael Storozum This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-4185222/v1 This work is licensed under a CC BY 4.0 License Status: Published Journal Publication published 23 Jul, 2024 Read the published version in npj Heritage Science → Version 1 posted 11 You are reading this latest preprint version Abstract This essay analyzes a robust body of C and N stable isotopes from human and animal collagen and plant remains ( n = 423) in the area of Lower Yangtze River dating to 8000 − 4300 cal. BP, combined with new isotopic data recovered from the Zhelin site in Shanghai, to explore the dietary differences between coastal region the inland core area of the Liangzhu culture (5300 − 4300 cal. BP). Our results reveal that the periphery dwellers of the Liangzhu cultural area became increasingly reliant on a limited number of domesticated species over time, while maintaining a marked social differentiation of foodways that was potentially a result of the growing but unequal influence of populations from the core Liangzhu area. Diet Stable isotope Rice agriculture Lower Yangtze area Liangzhu Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 Figure 6 Introduction In recent years, archaeologists have made significant discoveries at Liangzhu period sites, including China’s earliest city and water management technologies at Mojiaoshan and Maoshan [ 1 , 2 ] . The Liangzhu culture, located in southern China near Shanghai, is arguably the first ancient society in the world to have relied on intensive rice agriculture [ 3 – 5 ] . Despite a long-lasting period of cultural development, archaeologists argue that the Liangzhu culture collapsed around 4200 cal. BP possibly due to the late Holocene climate event and widespread flooding [ 6 – 10 ] . Although the general chronology and some key accomplishments of the Liangzhu culture are widely known, many functional and ritual aspects of the Liangzhu economy remain poorly understood, partly because most archaeological investigations have focused on sites within the urban core at Liangzhu itself (Fig. 1 ). To counter this emphasis, we examine and compare sites within the municipal areas of Shanghai and Ningbo in coastal area of southeastern China, located on the coastal margin of the Liangzhu cultural sphere (Fig. 1 ). Scholars have argued that diet can be a potential vector to examine people’s self-conception [ 11 , 12 ] . Given that stable carbon ( δ 13 C) and nitrogen ( δ 15 N) isotope analysis of human bones are widely accepted as indicators of dietary practices [ 13 ] , this study reexamines existing isotopic data from archaeological sites at the Lower Yangtze area in light of new evidence concerning the Liangzhu culture as well as a newly reported Liangzhu cultural site, Zhelin (Fig. 1 ). Isotopic investigations of ancient diets over the past 30 years have made great progress in understanding dietary shifts in northern China, but major advances have been lacking in the area south of the Yangtze River, due to soil conditions and poor preservation of skeletal materials in this region [ 14 , 15 ] . As such, we extracted collagen from human ( n = 9) and dog ( n = 1) bones recovered from the tombs and pits at the Zhelin site located in southern Shanghai for stable isotope analysis. This would allow us to examine the foodways of Zhelin residents living at this small coastal Liangzhu site dating to 5200 − 4900 cal. BP [ 16 ] . Furthermore, we collected extant isotopic data on human ( n = 56), animal ( n = 361) and plant ( n = 3) isotopic results from the Lower Yangtze area, published in Chinese and English, in order to summarize their findings and explore how the phenomenon of the Liangzhu culture led to adjustments in regional dietary diversity. Specifically, using new stable isotopic evidence from the Zhelin site and previous isotope evidence of human and different kinds of foodstuff remains recovered from the Lower Yangtze area during 8000 − 4300 cal. BP, we aimed to examine how dietary practices and economic decisions differed between small coastal sites and core regions located further inland before and during the Liangzhu period (5300 − 4300 cal. BP). This study provides novel insights into how residents of these small coastal sites on the periphery of Liangzhu sphere made specialized economic decisions while participating within the broader Liangzhu culture. Archaeological background The Zhelin site (30°49′45″N, 121°27′56″E, and 4 m asl) is situated on the north bank of Hangzhou Bay, Fengxian District of Shanghai (Fig. 1 ). Archaeologists first discovered the site in 1973. The site is about 100 meters long from east to west and 70 meters wide from north to south and considered as a residential site of Liangzhu culture situated along the ancient Shanghai coastline. In 2019, archaeologists excavated the site, in order to better understand its size and type (Fig. 2 ). Excavations have already revealed at least twenty child and adult burials in varying states of preservation and some burial objects including jade, pottery and bone tools (Fig. 3 A, B). The burials appear to be located within a slightly elevated platform near the prehistoric coastline (Fig. 3 C). Zhelin experienced a subtropical monsoon climate, with an average annual temperature of around 17.9 degrees Celsius and an average annual precipitation of >1500 mm. Existing archaeobotanical and archaeological data suggests that ancient humans living at the eastern area of Taihu Lake had developed rice-based farming and animal husbandry since around the Majiabang cultural period (7000 − 6000 cal. BP) [ 17 , 18 ] . Moreover, paleoenvironmental evidence shows that the marshification process of shallow wetlands may have contributed to the expansion of prehistoric rice-farming societies in the Taihu Basin during the Liangzhu culture (5300 − 4300 cal. BP) [ 19 ] . Recent physical anthropological research on human bones from Zhelin has revealed dental calculus and metatarsal-phalangeal joint osteoarthritis among some Zhelin residents [ 20 ] . In addition, archaeologists from the Shanghai Museum have found similarities between pottery excavated from burial contexts at Zhelin and similar contexts at Fuquanshan site (Fig. 1 ), a high ranking Liangzhu burial mound further inland at modern Qingpu District, Shanghai (Fig. 4 A, B). These common grave goods include the double-nosed hu - jar (Fig. 4 C, D). Pending further archaeological evidence from Zhelin and Fuquanshan, archaeologists believe that Zhelin may have been a part of an extended sub-center of Liangzhu society that accommodated both sides [ 3 ] . In addition, unlike the north-south oriented burials found at Fanshan in Liangzhu City [ 21 ] , the burials at the Zhelin site are all oriented east-west. Materials and Methods Stable C, N isotope analysis The procedure for stable C, N isotope analysis used in this study is the same as described in Brock et al. (2010) [ 22 ] . Detailed information on archaeological contexts is listed in the online Table S1 . We modified the protocol slightly to include a final stage of ultrafiltration prior to lyophilization, as described in Brown et al. (1988) [ 23 ] . Bone (~ 0.5 g) was cleaned by sonication and then demineralized in a 0.5 M HCl solution at 4℃ for 1–2 weeks, with the HCl solution changed every 48 hr. The remains were washed with deionized water to neutrality, and reinsed in 0.125 mol/L cold sodium hydroxide for 20 hr, and washed again with deionized water. The residues were rinsed in 0.001 mol/L HCl, then gelatinized at 70℃ in a pH = 3 solution for 2 days. The heated solution was filtered into tube and frozen, then freeze dried for 48 hr. The purified collagen was measured at the Environmental Stable Isotope Laboratory (ESIL), Institute of Environment and Sustainable Development of Agriculture, Chinese Academy of Agricultural Sciences, using an Isoprime 100 IRMS (Elementar, UK) coupled with an Elementar Vario (Elementar, UK), and calibrated with USGS 40 ( δ 13 C VPDB =-26.39 ± 0.04‰, δ 15 N air =-4.52 ± 0.06‰) and USGS41a ( δ 13 C VPDB = + 36.55 ± 0.08‰, δ 15 N air = + 47.55 ± 0.15‰) reference materials. For every 12 samples, a laboratory reference-Gelatin from bovine skin ( δ 13 C VPDB = 14.7 ± 0.2‰; δ 15 N air = 6.9 ± 0.2‰) was inserted for calibration and to monitor stability. The isotope results were analyzed as the ratio of the heavier isotope to the lighter isotope ( 13 C/ 12 C or 15 N/ 14 N) and expressed as ‘ δ ’ in parts per 1000 or per mil (‰) relative to internationally defined standards [ 24 ] for carbon (Vienna Pee Dee Belemnite, VPDB) and air nitrogen. The measurement errors were less than ± 0.2‰ for both δ 13 C and δ 15 N values. Moreover, existing δ 13 C and δ 15 N isotopic data from archaeological sites from the Lower Yangtze area around Shanghai before and during the Liangzhu period (5300 − 4300 cal. BP) (regional chronology of cultures and the development of rice agriculture shown in Fig. 5 ), were collected to further assess differences in dietary pattern between these peripheral populations and core communities living in the Lower Yangtze area (Fig. 1 ). We searched for combinations of the search terms including ‘稳定同位素’ /‘stable isotope’, ‘食谱’/‘diet/dietary pattern’, ‘新石器时代’ /‘Neolithic period’ and ‘长江下游’/‘Lower Yangtze’ in Chinese and English literature using the China National Knowledge Infrastructure ( http://www.cnki.net ) and Google scholar, respectively. We extracted site information and relevant isotopic data from each relevent hit. Since some researchers only published scatter plots of stable carbon and nitrogen isotope values, we used software of GetData Graph Digitizer to capture the C, N isotopic values. The detailed information of these sites and data are listed in Table S2 and Table S3. Statistical data analysis was performed using the Student’s t -test using the Microsoft Office Excel, with the significance level set at P < 0.05. Results Isotopic evidence of dietary patterns New isotope data for Zhelin’s human and dog specimens ( n = 10) is listed in Table S1 . Due to the humid environment and acidic soils in Shanghai area, only two human specimens and one dog specimen produced collagen of suitable quality, with C:N ratios ranging between 2.9 to 3.6 for further analysis [ 25 , 26 ] . Our δ 13 C values for the human and dog samples ( n = 3) from Zhelin are − 19.2‰, -19.4‰ and − 19.6‰ (mean ± SD: -19.4 ± 0.2‰), respectively, indicative of similar diets significantly influenced by C 3 -based foods. The δ 15 N values of these three specimens were + 9.1‰, + 11.3‰ and + 8.6‰ (mean ± SD: +9.7 ± 1.2‰), respectively, suggesting they consumed foods at different trophic levels. All new and existing stable C, N isotope data analyzed in the present study are listed in Table S2 and Table S3. Previous studies have suggested that human bones are 15 N enriched by about 3–5‰ relative to the foods they consume [ 27 ] . As can be seen Fig. 6 and detailed in Table S3, carbon and nitrogen isotope values of charred plant samples ( n = 3) including the rice grains and acorns from the Tianluoshan site (7000 − 5300 cal. BP), located on the south bank of Hangzhou Bay, range from − 28.4‰ to -25.4‰ (mean ± SD: -27.1 ± 1.6‰) and + 4.1‰ to + 7.4‰ (mean ± SD: +5.3 ± 1.9‰), which are the lowest C and N isotopic values [ 28 ] , establishing an isotope baseline for primary producers in surrounding area. On the contrary, the highest δ 13 C (mean ± SD: -13.4 ± 1.1‰) and δ 15 N values (mean ± SD: +13.0 ± 1.6‰) derived tuna and whale bones (marine fish/mammal) from the Tianluoshan site [ 29 , 30 ] , revealing the highest enriched 13 C and 15 N isotopic signals for marine animals in this coastal area. Additionally, the largest amount of previously published isotopic data derived from terrestrial herbivores ( n = 194) including several kinds of deer and bovid (including water buffalo) from the sites (Fig. 6 ) of Kuahuqiao (8000 − 7000 cal. BP), Tianluoshan and Meirendi (5300 − 4300 cal. BP), which ranges from − 24.8‰ to -10.7‰ (mean ± SD: -17.8 ± 3.9‰) and + 3.3‰ to + 11.8‰ (mean ± SD: +7.4 ± 2.1‰), suggesting that these likely enjoyed a broad spectrum of diets derive from 13 C-enriched plant foods [ 29 – 34 ] . The second most abundant animal isotopic results belonged to terrestrial omnivores ( n = 139) dominated by pig and dog samples from Kuahuqiao, Jiangjiashan (7000 − 6000 cal. BP), Tianluoshan, Meirendi and our study site of Zhelin. The mean δ 13 C and δ 15 N values of these animals are − 20.6 ± 1.2‰ and + 6.4 ± 1.5‰, indicating that omnivore diet in the studied area was largely influenced by low-protein C 3 terrestrial grasses and shrubs [ 29 – 34 ] . The δ 13 C and δ 15 N values of freshwater animals including freshwater fish, frog, alligator, and turtle from Kuahuqiao, Tianluoshan and Meirendi ranged from − 23.5‰ to -18.8‰ (mean ± SD: -20.9 ± 1.4‰) and + 5.6‰ to + 12.3‰ (mean ± SD: +8.0 ± 1.7‰), the highest level for protein C 3 foods inland [ 29 – 32 ] . As for the isotopic data of human beings, plotted in Fig. 6 , in the coastal area around Shanghai, the Songze human bone samples ( n = 2) during Songze culture period (6000 − 5300 cal. BP) exhibited δ 13 C values of -20.2‰ and − 19.6‰ (mean ± SD: -19.9 ± 0.3‰) and δ 15 N values of + 9.7‰ and + 12.0‰ (+ 10.9 ± 1.2‰) during 6000 − 5300 cal. BP [ 35 ] . This reveals that individuals largely relied on C 3 -based diets with varying trophic levels. In another small coastal settlement in the Ningshao area (Fig. 1 ), Tashan site, the mean carbon and nitrogen isotope values of humans during Liangzhu period ( n = 2) are − 18.7 ± 0.3‰ and + 8.9 ± 0.5‰ during 5300 − 4300 cal. BP [ 36 ] . Dating before that time, one individual from the Tashan site during 5900 − 5600 cal. BP before Liangzhu culture period yielded 13 C and 15 N-enriched results ( δ 13 C = -18.0‰ and δ 15 N = + 9.8‰). Similarly, in the same area of Ningshao, the mean carbon and nitrogen isotope values of the Hemudu sample ( n = 2) during 7000 − 6000 cal. BP were − 16.7 ± 1.6‰ and + 11.4 ± 0.2‰, respectively [ 35 ] . However, the δ 13 C and δ 15 N values of human bone samples ( n = 20) from another Hemudu culture site of Tianluoshan ranged from − 21.9‰ to -20.1‰ (mean ± SD: -20.6 ± 0.5‰) and + 7.2‰ to + 10.7‰ (mean ± SD: +8.9 ± 0.9‰) during 7000 − 5300 cal. BP [ 29 , 33 ] . As for the isotopic evidence for diets in these inland sites at Lower Yangtze area prior to and during the Liangzhu period (5300 − 4300 cal. BP), carbon and nitrogen isotope values of Jiangjiashan people ( n = 1) living on the southwestern bank of Taihu Lake during 7000 − 6000 cal. BP (Fig. 1 ), were − 20.5‰ and + 10.2‰., respectively (Fig. 6 ). According to Hu et al (2007) [ 37 ] , the carbon and nitrogen isotope values of Sanxingcun peoples ( n = 19) ranges from − 20.4‰ to -19.6‰ (mean ± SD: -20.0 ± 0.2‰) and + 8.9‰ to + 10.3‰ (mean ± SD: +9.7 ± 0.3‰) indicative of a C 3 -based diet ate by these individuals. Given that the Sanxingcun site is located in the north of Taihu Lake at the Lower Yangtze area, where an unbroken rice agriculture had been practiced early Holocene period [ 4 , 38 ] , combined with the isotopic evidence of Sanxingcun people’s trophic level, it is safe to believe that the humans of Sanxingcun mainly fed on rice and terrestrial omnivores as well as a small number of terrestrial herbivores during the 6500 − 5500 cal. BP period. In addition, during the Liangzhu period (5300 − 4300 cal. BP), Liangzhu resident ( n = 9) δ 13 C and δ 15 N values recovered from the Liangzhu City area (Meirendi site) ranged from − 22.2‰ to -17.4‰ (mean ± SD: -19.9 ± 1.2‰) and + 9.7‰ to + 11.0‰ (mean ± SD: +10.3 ± 0.4‰), which is indicative of a C 3 -based diet with relatively high trophic levels. Discussion Just as at Liangzhu [ 1 ] , archaeological work around the world often prioritizes the great sites. However, to better understand the past and the role that these large sites had to play in shaping neighboring settlements, archaeologists should obviously also pay attention to these smaller sites. We argue that the recent studies at core inland Liangzhu sites can only be comprehensively understood within a broader regional context and with regard to peripheral coastal sites at Lower Yangtze area. Previous work by Dong and Yuan has suggested that existing zooarchaeological findings at the Liangzhu cultural area reveal different animal husbandry practices between the urban center of Mojiaoshan and small sized sites during Liangzhu period (5300 − 4300 cal. BP) [ 18 ] . In the present study, we first attempted to investigate previously unexamined dietary practices of Liangzhu cultural human societies at Shanghai area, as a geographic periphery of the Liangzhu core. More importantly, by integrating new and existing C, N isotopic data of humans ( n = 58), animals ( n = 362) and plants ( n = 3) recovered from sites around the Lower Yangtze area during 8000 − 4300 cal. BP, we provided a broad picture of the dynamic changes of diets and economic decisions of these coastal communities living at Shanghai and Ningshao areas before and during the Liangzhu phenomenon. Although rice agriculture began around 10000 cal. BP in the Lower Yangtze area [ 38 ] , archaeological evidence suggests that rice agricultural production in the region developed over thousands of years until before becoming more stabilized during the middle Songze culture through Liangzhu periods (5600 − 4300 cal. BP) [ 39 ] . Prior to this time, ancient humans may have consumed more wild animals and nuts for survival [ 40 ] . As shown in Fig. 6 , in the coastal area of Ningshao before Liangzhu time, stable carbon and nitrogen isotope evidence from the Hemudu site and Tianluoshan site suggests that diversified human subsistence, like exploitation of wild plants, seawater and freshwater resources as well as wild games, sustained a continuous parallel development with low-level rice farming for millennia. In another coastal area of Shanghai, δ 15 N results for human bones recovered from the Songze site, 6000 − 5300 cal. BP, yield an increase in δ 15 N values compared to the inland residents of Jiangjiashan (7000 − 6000 cal. BP) and Sanxingcun (6500 − 5500 cal. BP) (student’s t -test, P = 0.0056). 15 N enrichment levels in the Songze peoples (6000 − 5300 cal. BP) who lived in the coastal area of Shanghai at nearly the same time consumed a greater amount of freshwater foods. One important finding is that although different groups of Neolithic populations in Lower Yangtze region from 7000 − 5300 cal. BP have similar C 3 -based diets which were probably attributed to low-level rice farming and foraging of wild plant resources at large, according to varies of stable nitrogen values and zooarchaeological evidence from the study area [ 18 ] , there are significant differences in foodways between the populations who lived in the inland area (Sanxingcun, Jiangjiashan) and other coastal (Hemudu, Tianluoshan, Songze, Tashan) settlements (Fig. 1 , 6 ). This also shows that the food-related differentiation had appeared in inland and coastal societies at the Lower Yangtze River area during 7000 − 5300 cal. BP. According to previous archaeobotanical studies and archaeological finding of complex water management systems at ancient Liangzhu City [ 2 ] , during the Liangzhu period rice agriculture rapidly developed and rice foods eventually became a staple food incorporated into diets across the Lower Yangtze area [ 4 , 10 ] . At this time, more sophisticated water management and farming techniques allowed the majority of individuals in the core of the Liangzhu area had a relatively balanced diet including a large number of domesticated rice, and meat from domesticates like pigs and some wild games including a number of freshwater resources as well. We had compared the δ 13 C and δ 15 N values of human samples from the sites of Sanxingcun and Liangzhu City (Meirendi site). Our analysis reveals that both inland communities shared a similar C 3 -based dietary pattern, while the core Liangzhu culture humans from the Meirendi site had relatively high trophic levels than Sanxingcun peoples based on N isotopic evidence (student’s t -test, P = 0.0005). As for the foodways of human beings living at the peripheral coastal areas during Liangzhu period (5300 − 4300 cal. BP), previous work by Zhang et al. (2015) [ 36 ] , for example, has reported that Tashan people, occupying a small East China Sea coastal settlement in the period 5300 − 4300 cal. BP, likely exploited more foods from rice and terrestrial animals but less marine food resources than the residents of the same site for the period 5900 − 5600 cal. BP. Also, it can be seen that the Hemudu residents living in the Ningshao area during 7000 − 6000 cal. BP had higher stable nitrogen isotopic values than Tashan peoples during 5300 − 4300 cal. BP (student’s t -test, P = 0.0434). Considering that large-scale rice paddies have been found at the Shiao site in Ningshao area directly dating to the Hemudu culture and the Liangzhu culture periods [ 41 ] , we believe that the changes of diets and consumption habits of these peripheral populations at Tashan during the Liangzhu period were likely influenced by the expansion of massive rice cultivation in the Ningshao area from the Hemudu culture to Liangzhu culture periods (7000 − 4300 cal. BP). Nevertheless, we have also found that the δ 15 N values of humans of Tashan lower than those of the Liangzhu City during the same period (student’s t -test, P = 0.0052). This suggests that despite communication with the core Liangzhu culture, the Tashan societies reorganized their own sense of cultural identity along dietary lines and made specialized economic decisions to participate in a new social network, influenced by inland Liangzhu culture. Moreover, as can be seen in Fig. 6 , our new isotopic data suggests that the dietary lifestyle of the Zhelin people, living on a small Liangzhu site in the coastal area of the Lower Yangtze, was more similar to that of people from the core area of the Liangzhu culture compared to human beings from the Tashan site at Ningshao area during the Liangzhu period. Specifically, from Fig. 6 , the δ 13 C (student’s t -test, P = 0.5334) and the δ 15 N (student’s t -test, P = 0.8691) values of humans at the Zhelin site and the inland Liangzhu core (Meirendi site) express no clear differentiation. Mean δ 15 N values at Zhelin are, however, higher than those of the Tashan individuals during the same period (Fig. 6 ). Moreover, the diet of Zhelin remains largely consistent to previous dietary pattern of humans at the Songze site during 6000 − 5200 cal. BP ( δ 13 C: student’s t -test, P = 0.1982; δ 15 N: student’s t -test, P = 0.7225). Interestingly, the mean δ 15 N value of Zhelin humans decreased compared to inhabitants of Songze in nearby region prior to Liangzhu period (Fig. 6 ), perhaps suggesting a general decrease in the consumption of high-protein foods, such as aquatic meats, as subsistence economies became more agrarian during the Liangzhu period (5300 − 4300 cal. BP). In these cases, we argue that the economic decisions of the Zhelin people may be more dominated by Liangzhu core concerns, expressing a recognition of inland dietary lifeways of the core area of Liangzhu culture. Previous studies have suggested that intensive rice agriculture has underpinned increased human populations and massive social evolution at the Lower Yangtze area during the Liangzhu period (5300 − 4300 cal. BP) through a rapid expansion of wetland under rice cultivation and a greater reliance on domesticated foods like pigs and water buffalo [3–4,18−19,42–43] . As such, we believe that the diets and consumption habits of these peripheral Liangzhu populations at Zhelin and Tashan, perhaps influenced, to some extent, by their desired food choices under the rapid development of early urbanized and agricultural lifestyle originating in the core Liangzhu area. The differentiation in dietary practices between the populations from the coastal Zhelin and Tashan sites, further suggests that the two societies living at the periphery of the Liangzhu cultural sphere may have taken on different roles to integrated into a new social network of Liangzhu culture. In the long run, this transition to a more standardized man-made agricultural economic choice may have inadvertently locked Liangzhu society into a system that depended on a limited range of “taxable” domesticated species [ 44 ] , which may have been significantly affected by rapid climate changes at around 4200 cal. BP. Conclusion By integrating new C, N isotopes from Zhelin, Shanghai with previous isotopic studies done in the Taihu Basin and Ningshao areas during 8000 − 4300 cal. BP, we used the isotopic evidence to gain a better understanding of the dietary and culinary changes at the Lower Yangtze area prior to and during the Liangzhu cultural period (5300 − 4300 cal. BP). Our findings suggest that the dietary differences between the coastal periphery and more inland core cultural area of Lower Yangtze area likely existed through the Liangzhu period. Moreover, our study shows that the Liangzhu dependence on limited domesticated victuals became much more widespread around Lower Yangtze area, and more socially integrated terrestrial economic decisions appeared along with the development of Liangzhu cultural phenomenon in this coastal region. This study provides essential data for understanding the development of the Liangzhu cultural phenomenon at Lower Yangtze area in a more comprehensive manner. However, more C, N isotopic research is necessary for more robust comparisons between the developmental trajectory of foodways across different societies in the coastal reaches of Neolithic southeastern China. Declarations Acknowledgements The authors are grateful to the funds from Shanghai Planning Office of Philosophy and Social Science (2019ELS006) for financial support. We are thankful to Prof. Yi Guo at Zhejiang University and Dr Simei Zhu at Zhengzhou University for their help with the pre-treatment experiment of bone samples. Author contributions Conceptualization: P.S., E.A., M.S.; methodology: P.S., M.S.; Investigation: P.S., X.Z., X.H., M.S.; Funding Acquisition: P.S.; Project Administration: P.S.; Writing—original draft: P.S., E.A., M.S.; Writing—review & editing: P.S., E.A., M.S. All authors contributed to the article and approved the submitted version. Competing Interests The authors declare no competing interests. References Renfrew, C. & Liu, B. 2018. The emergence of complex society in China: the case of Liangzhu. Antiquity, 92(364): 975-90. Liu, B., Wang, N., Chen, M., Wu, X., Mo, D., Liu, J., Xu, S. and Zhuang, Y. 2017. 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Zhang, X.L., Wang, J.X., Xian, Z.Q. & Qiu, X.J. 2003. Isotopic study of ancient man’s diet. Archaeology, 158(2): 62-75 (in Chinese). Zhang, G.W., Jiang, L.P., Hu, Y.W., Si, Y., Lv, P., Song, G.D., Wang, C.S., Richard, M. & Guo, Y. 2015. Carbon and Nitrogen stable isotope analysis of human and animal bones recovered from Tashan site, Zhejiang. Huaxia Archaeology, 2: 138-46 (in Chinese). Hu, Y.W., Wang, G.F., Cui, Y.P., Dong, Y., Guan, L. & Wang, C.S. 2007. The study of human diets at Sangxingcun site, Jintan, Jiangsu. Chinese Science Bulletin, 52(1): 85-8 (in Chinese). Jones, M.K. & Liu, X.Y. 2009. Origins of agriculture in East Asia. Science, 324: 730-731. Lu, H.Y. 2017. New methods and progress in research on the origins and evolution of prehistoric agriculture in China. Science China Earth Sciences, 60: 2141-2159. Fuller, D.Q. & Qin, L. 2010. Declining oaks, increasing artistry, and cultivating rice: The environmental and social context of the emergence of farming in the Lower Yangtze Region. Environmental Archaeology, 15(2): 139-59. Wang, Y., Song, S., Zhang, Y., Mei, S., Lu, X., Zheng, Y. & Sun, G. 2023. Brief reports of the excavation of the ancient rice paddies at the Shiao site, Yuyao City, Zhejiang. Archaeology, 5: 3-21 (in Chinese). Qin, L., & Fuller, D. Q. 2019. Why Rice Farmers Don’t Sail: Coastal Subsistence Traditions and Maritime Trends in Early China. In Prehistoric Maritime Cultures and Seafaring in East Asia (pp. 159-191). Springer, Singapore. Zheng, H.B., Zhou, Y.S., Yang, Q., Hu, Z.J., Ling, G.J., Zhang, J.Z., Gu, C.G., Wang, Y.Y., Cao, Y.T., Huang, X.R., Cheng, Y., Zhang, X.Y. & Wu, W.X. 2018. Spatial and temporal distribution of Neolithic sites in coastal China: Sea level changes, geomorphic evolution and human adaptation. Science China Earth Sciences, 61: 123-133 (in Chinese). Scott, J.C. 2009. The Art of Not Being Governed: An Anarchist History of Upland Southeast Asia . New Haven : Yale University Press. Additional Declarations No competing interests reported. Supplementary Files SupplementaryTable.xls Cite Share Download PDF Status: Published Journal Publication published 23 Jul, 2024 Read the published version in npj Heritage Science → Version 1 posted Editorial decision: Revision requested 17 Apr, 2024 Reviews received at journal 16 Apr, 2024 Reviews received at journal 06 Apr, 2024 Reviewers agreed at journal 06 Apr, 2024 Reviews received at journal 03 Apr, 2024 Reviewers agreed at journal 02 Apr, 2024 Reviewers agreed at journal 01 Apr, 2024 Reviewers invited by journal 01 Apr, 2024 Editor assigned by journal 30 Mar, 2024 Submission checks completed at journal 30 Mar, 2024 First submitted to journal 28 Mar, 2024 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. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. 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Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-4185222","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":286820110,"identity":"c1315b46-4c28-41ca-bc12-8e429a4c06ef","order_by":0,"name":"Pengfei Sheng","email":"data:image/png;base64,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","orcid":"","institution":"Fudan University","correspondingAuthor":true,"prefix":"","firstName":"Pengfei","middleName":"","lastName":"Sheng","suffix":""},{"id":286820111,"identity":"bf5361b4-0784-4c36-9902-cf80016c2317","order_by":1,"name":"Edward Allen","email":"","orcid":"","institution":"Fudan University","correspondingAuthor":false,"prefix":"","firstName":"Edward","middleName":"","lastName":"Allen","suffix":""},{"id":286820112,"identity":"17e684ae-2f61-42cc-a5cb-0a47d420ad09","order_by":2,"name":"Xiang Huang","email":"","orcid":"","institution":"Shanghai Museum","correspondingAuthor":false,"prefix":"","firstName":"Xiang","middleName":"","lastName":"Huang","suffix":""},{"id":286820113,"identity":"a5673ef8-e5e8-4215-aacd-af96366c98d0","order_by":3,"name":"Xiuwen Zheng","email":"","orcid":"","institution":"Shanghai Museum","correspondingAuthor":false,"prefix":"","firstName":"Xiuwen","middleName":"","lastName":"Zheng","suffix":""},{"id":286820114,"identity":"2eae5e28-6ca1-4844-8f87-0510e41ce5ac","order_by":4,"name":"Michael Storozum","email":"","orcid":"","institution":"Newcastle University","correspondingAuthor":false,"prefix":"","firstName":"Michael","middleName":"","lastName":"Storozum","suffix":""}],"badges":[],"createdAt":"2024-03-29 02:44:14","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-4185222/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-4185222/v1","draftVersion":[],"editorialEvents":[{"content":"https://doi.org/10.1186/s40494-024-01374-3","type":"published","date":"2024-07-23T16:15:33+00:00"}],"editorialNote":"","failedWorkflow":false,"files":[{"id":53992124,"identity":"518ab5e5-c731-4578-a519-5d3380771f78","added_by":"auto","created_at":"2024-04-03 06:08:17","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":3402996,"visible":true,"origin":"","legend":"\u003cp\u003eMap of archaeological sites mentioned in the text, 1, Sanxingcun (6500-5500 cal. BP); 2, Fuquanshan (5300-4300 cal. BP); 3, Songze (6000-5300 cal. BP); 4, Zhelin (5300-4300 cal. BP); 5, Jiangjiashan (7000-6000 cal. BP); 6, Liangzhu (Meirendi site, 5300-4300 cal. BP); 7, Kuahuqiao (8000-7000 cal. BP); 8, Tianluoshan (7000-5300 cal. BP); 9, Hemudu (7000-6000 cal. BP); 10, Tashan (5300-4300 cal. BP).\u003c/p\u003e","description":"","filename":"Figure1.png","url":"https://assets-eu.researchsquare.com/files/rs-4185222/v1/db50cee251ed6b9a5a308f7c.png"},{"id":53992125,"identity":"c1082dda-1483-413a-a83d-ead508b64999","added_by":"auto","created_at":"2024-04-03 06:08:17","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":791059,"visible":true,"origin":"","legend":"\u003cp\u003eA) Map of the Zhelin site, B) Aerial photograph of the area around the Zhelin site, C) Excavation setting, D) Excavations in 2019.\u003c/p\u003e","description":"","filename":"Figure2.png","url":"https://assets-eu.researchsquare.com/files/rs-4185222/v1/cda7cbf8d49ef327dd50c6a0.png"},{"id":53992462,"identity":"a8d38f7e-342a-435a-a6ee-7ff77b21b1b8","added_by":"auto","created_at":"2024-04-03 06:16:17","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":621460,"visible":true,"origin":"","legend":"\u003cp\u003eA) Burial with associated grave goods found at the Zhelin site, B) Canine burial found in an excavated pit, C) Profile of stratigraphy at Zhelin: note the two separate shell layers (ZL-1, ZL-2) with what the archaeologists interpret to be an earthen platform (ZL-3) built on top.\u003c/p\u003e","description":"","filename":"Figure3.png","url":"https://assets-eu.researchsquare.com/files/rs-4185222/v1/83191d4de2859ea6c43baf8f.png"},{"id":53992463,"identity":"c938c8dd-b178-4e32-b5bc-e1d2d2ede1f7","added_by":"auto","created_at":"2024-04-03 06:16:17","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":1444178,"visible":true,"origin":"","legend":"\u003cp\u003eA) Map of the Fuquanshan site, B) Pottery double-nosed \u003cem\u003ehu\u003c/em\u003e-flask found at Zhelin (\u003cem\u003ephotographed by X. Zheng\u003c/em\u003e), Fuquanshan and Tashan, respectively.\u003c/p\u003e","description":"","filename":"Figure4.png","url":"https://assets-eu.researchsquare.com/files/rs-4185222/v1/5f470dc5b09e869c319916a1.png"},{"id":53992129,"identity":"bb4da96c-bcac-4866-aa62-ead14426f21b","added_by":"auto","created_at":"2024-04-03 06:08:17","extension":"png","order_by":5,"title":"Figure 5","display":"","copyAsset":false,"role":"figure","size":111385,"visible":true,"origin":"","legend":"\u003cp\u003eChronology of major cultures mentioned in this study and the density of rice seeds recovered from the sites at the eastern Taihu Basin (redrawn after\u003csup\u003e [19]\u003c/sup\u003e).\u003c/p\u003e","description":"","filename":"Figure5.png","url":"https://assets-eu.researchsquare.com/files/rs-4185222/v1/282376325cc6cf238f73ca71.png"},{"id":53992128,"identity":"21070b47-8133-42e6-af70-3bb261c25e33","added_by":"auto","created_at":"2024-04-03 06:08:17","extension":"png","order_by":6,"title":"Figure 6","display":"","copyAsset":false,"role":"figure","size":219593,"visible":true,"origin":"","legend":"\u003cp\u003eMean ± SD \u003cem\u003eδ\u003c/em\u003e\u003csup\u003e13\u003c/sup\u003eC and \u003cem\u003eδ\u003c/em\u003e\u003csup\u003e15\u003c/sup\u003eN values of human, animal and plant samples recovered from archaeological sites at the Lower Yangtze area before and during the Liangzhu period (Detailed information of these sites and isotopic data detailed in Table S2 and S3) (\u003cem\u003efigure by P. Sheng\u003c/em\u003e).\u0026nbsp;\u003c/p\u003e","description":"","filename":"Figure6.png","url":"https://assets-eu.researchsquare.com/files/rs-4185222/v1/a8ecea3e82ba1bc1fb579f14.png"},{"id":61597021,"identity":"60b121db-eb6f-46bd-88e6-eeea4ea03f6c","added_by":"auto","created_at":"2024-08-01 17:31:24","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":8405876,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-4185222/v1/8921d9e1-3e42-46c1-b990-16d2318a999c.pdf"},{"id":53992127,"identity":"68ab0126-5f6f-49a7-9253-f73a1d0ae954","added_by":"auto","created_at":"2024-04-03 06:08:17","extension":"xls","order_by":1,"title":"","display":"","copyAsset":false,"role":"supplement","size":103424,"visible":true,"origin":"","legend":"","description":"","filename":"SupplementaryTable.xls","url":"https://assets-eu.researchsquare.com/files/rs-4185222/v1/1f3e4b4f2f99a9595f243f96.xls"}],"financialInterests":"No competing interests reported.","formattedTitle":"To eat like Liangzhu: isotopic investigation of diets in the Lower Yangtze area prior to and during the Liangzhu period (5300-4300 cal. BP)","fulltext":[{"header":"Introduction","content":"\u003cp\u003eIn recent years, archaeologists have made significant discoveries at Liangzhu period sites, including China\u0026rsquo;s earliest city and water management technologies at Mojiaoshan and Maoshan\u003csup\u003e[\u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e1\u003c/span\u003e, \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2\u003c/span\u003e]\u003c/sup\u003e. The Liangzhu culture, located in southern China near Shanghai, is arguably the first ancient society in the world to have relied on intensive rice agriculture\u003csup\u003e[\u003cspan additionalcitationids=\"CR4\" citationid=\"CR3\" class=\"CitationRef\"\u003e3\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e5\u003c/span\u003e]\u003c/sup\u003e. Despite a long-lasting period of cultural development, archaeologists argue that the Liangzhu culture collapsed around 4200 cal. BP possibly due to the late Holocene climate event and widespread flooding\u003csup\u003e[\u003cspan additionalcitationids=\"CR7 CR8 CR9\" citationid=\"CR6\" class=\"CitationRef\"\u003e6\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e10\u003c/span\u003e]\u003c/sup\u003e. Although the general chronology and some key accomplishments of the Liangzhu culture are widely known, many functional and ritual aspects of the Liangzhu economy remain poorly understood, partly because most archaeological investigations have focused on sites within the urban core at Liangzhu itself (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). To counter this emphasis, we examine and compare sites within the municipal areas of Shanghai and Ningbo in coastal area of southeastern China, located on the coastal margin of the Liangzhu cultural sphere (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eScholars have argued that diet can be a potential vector to examine people\u0026rsquo;s self-conception \u003csup\u003e[\u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e11\u003c/span\u003e, \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e12\u003c/span\u003e]\u003c/sup\u003e. Given that stable carbon (\u003cem\u003eδ\u003c/em\u003e\u003csup\u003e13\u003c/sup\u003eC) and nitrogen (\u003cem\u003eδ\u003c/em\u003e\u003csup\u003e15\u003c/sup\u003eN) isotope analysis of human bones are widely accepted as indicators of dietary practices\u003csup\u003e[\u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e13\u003c/span\u003e]\u003c/sup\u003e, this study reexamines existing isotopic data from archaeological sites at the Lower Yangtze area in light of new evidence concerning the Liangzhu culture as well as a newly reported Liangzhu cultural site, Zhelin (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). Isotopic investigations of ancient diets over the past 30 years have made great progress in understanding dietary shifts in northern China, but major advances have been lacking in the area south of the Yangtze River, due to soil conditions and poor preservation of skeletal materials in this region\u003csup\u003e[\u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e14\u003c/span\u003e, \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e15\u003c/span\u003e]\u003c/sup\u003e. As such, we extracted collagen from human (\u003cem\u003en\u003c/em\u003e\u0026thinsp;=\u0026thinsp;9) and dog (\u003cem\u003en\u003c/em\u003e\u0026thinsp;=\u0026thinsp;1) bones recovered from the tombs and pits at the Zhelin site located in southern Shanghai for stable isotope analysis. This would allow us to examine the foodways of Zhelin residents living at this small coastal Liangzhu site dating to 5200\u0026thinsp;\u0026minus;\u0026thinsp;4900 cal. BP\u003csup\u003e[\u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e16\u003c/span\u003e]\u003c/sup\u003e.\u003c/p\u003e \u003cp\u003eFurthermore, we collected extant isotopic data on human (\u003cem\u003en\u003c/em\u003e\u0026thinsp;=\u0026thinsp;56), animal (\u003cem\u003en\u003c/em\u003e\u0026thinsp;=\u0026thinsp;361) and plant (\u003cem\u003en\u003c/em\u003e\u0026thinsp;=\u0026thinsp;3) isotopic results from the Lower Yangtze area, published in Chinese and English, in order to summarize their findings and explore how the phenomenon of the Liangzhu culture led to adjustments in regional dietary diversity. Specifically, using new stable isotopic evidence from the Zhelin site and previous isotope evidence of human and different kinds of foodstuff remains recovered from the Lower Yangtze area during 8000\u0026thinsp;\u0026minus;\u0026thinsp;4300 cal. BP, we aimed to examine how dietary practices and economic decisions differed between small coastal sites and core regions located further inland before and during the Liangzhu period (5300\u0026thinsp;\u0026minus;\u0026thinsp;4300 cal. BP). This study provides novel insights into how residents of these small coastal sites on the periphery of Liangzhu sphere made specialized economic decisions while participating within the broader Liangzhu culture.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e\n\u003ch3\u003eArchaeological background\u003c/h3\u003e\n\u003cp\u003eThe Zhelin site (30\u0026deg;49\u0026prime;45\u0026Prime;N, 121\u0026deg;27\u0026prime;56\u0026Prime;E, and 4 m asl) is situated on the north bank of Hangzhou Bay, Fengxian District of Shanghai (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). Archaeologists first discovered the site in 1973. The site is about 100 meters long from east to west and 70 meters wide from north to south and considered as a residential site of Liangzhu culture situated along the ancient Shanghai coastline. In 2019, archaeologists excavated the site, in order to better understand its size and type (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). Excavations have already revealed at least twenty child and adult burials in varying states of preservation and some burial objects including jade, pottery and bone tools (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003eA, B). The burials appear to be located within a slightly elevated platform near the prehistoric coastline (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003eC).\u003c/p\u003e \u003cp\u003eZhelin experienced a subtropical monsoon climate, with an average annual temperature of around 17.9 degrees Celsius and an average annual precipitation of \u0026gt;1500 mm. Existing archaeobotanical and archaeological data suggests that ancient humans living at the eastern area of Taihu Lake had developed rice-based farming and animal husbandry since around the Majiabang cultural period (7000\u0026thinsp;\u0026minus;\u0026thinsp;6000 cal. BP)\u003csup\u003e[\u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e17\u003c/span\u003e, \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e18\u003c/span\u003e]\u003c/sup\u003e. Moreover, paleoenvironmental evidence shows that the marshification process of shallow wetlands may have contributed to the expansion of prehistoric rice-farming societies in the Taihu Basin during the Liangzhu culture (5300\u0026thinsp;\u0026minus;\u0026thinsp;4300 cal. BP)\u003csup\u003e[\u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e19\u003c/span\u003e]\u003c/sup\u003e.\u003cdiv class=\"BlockQuote\"\u003e\u003cp\u003eRecent physical anthropological research on human bones from Zhelin has revealed dental calculus and metatarsal-phalangeal joint osteoarthritis among some Zhelin residents\u003csup\u003e[\u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e20\u003c/span\u003e]\u003c/sup\u003e. In addition, archaeologists from the Shanghai Museum have found similarities between pottery excavated from burial contexts at Zhelin and similar contexts at Fuquanshan site (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e), a high ranking Liangzhu burial mound further inland at modern Qingpu District, Shanghai (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003eA, B). These common grave goods include the double-nosed \u003cem\u003ehu\u003c/em\u003e-\u003cem\u003ejar\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003eC, D). Pending further archaeological evidence from Zhelin and Fuquanshan, archaeologists believe that Zhelin may have been a part of an extended sub-center of Liangzhu society that accommodated both sides\u003csup\u003e[\u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e3\u003c/span\u003e]\u003c/sup\u003e. In addition, unlike the north-south oriented burials found at Fanshan in Liangzhu City\u003csup\u003e[\u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e21\u003c/span\u003e]\u003c/sup\u003e, the burials at the Zhelin site are all oriented east-west.\u003c/p\u003e\u003c/div\u003e\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e"},{"header":"Materials and Methods","content":"\u003cdiv id=\"Sec4\" class=\"Section2\"\u003e \u003ch2\u003eStable C, N isotope analysis\u003c/h2\u003e \u003cp\u003eThe procedure for stable C, N isotope analysis used in this study is the same as described in Brock et al. (2010)\u003csup\u003e[\u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e22\u003c/span\u003e]\u003c/sup\u003e. Detailed information on archaeological contexts is listed in the online Table \u003cspan refid=\"MOESM1\" class=\"InternalRef\"\u003eS1\u003c/span\u003e. We modified the protocol slightly to include a final stage of ultrafiltration prior to lyophilization, as described in Brown et al. (1988)\u003csup\u003e[\u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e23\u003c/span\u003e]\u003c/sup\u003e. Bone (~\u0026thinsp;0.5 g) was cleaned by sonication and then demineralized in a 0.5 M HCl solution at 4℃ for 1\u0026ndash;2 weeks, with the HCl solution changed every 48 hr. The remains were washed with deionized water to neutrality, and reinsed in 0.125 mol/L cold sodium hydroxide for 20 hr, and washed again with deionized water. The residues were rinsed in 0.001 mol/L HCl, then gelatinized at 70℃ in a pH\u0026thinsp;=\u0026thinsp;3 solution for 2 days. The heated solution was filtered into tube and frozen, then freeze dried for 48 hr.\u003c/p\u003e \u003cp\u003eThe purified collagen was measured at the Environmental Stable Isotope Laboratory (ESIL), Institute of Environment and Sustainable Development of Agriculture, Chinese Academy of Agricultural Sciences, using an Isoprime 100 IRMS (Elementar, UK) coupled with an Elementar Vario (Elementar, UK), and calibrated with USGS 40 (\u003cem\u003eδ\u003c/em\u003e\u003csup\u003e13\u003c/sup\u003eC\u003csub\u003eVPDB\u003c/sub\u003e=-26.39\u0026thinsp;\u0026plusmn;\u0026thinsp;0.04\u0026permil;, \u003cem\u003eδ\u003c/em\u003e\u003csup\u003e15\u003c/sup\u003eN\u003csub\u003eair\u003c/sub\u003e=-4.52\u0026thinsp;\u0026plusmn;\u0026thinsp;0.06\u0026permil;) and USGS41a (\u003cem\u003eδ\u003c/em\u003e\u003csup\u003e13\u003c/sup\u003eC\u003csub\u003eVPDB\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;+\u0026thinsp;36.55\u0026thinsp;\u0026plusmn;\u0026thinsp;0.08\u0026permil;, \u003cem\u003eδ\u003c/em\u003e\u003csup\u003e15\u003c/sup\u003eN\u003csub\u003eair\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;+\u0026thinsp;47.55\u0026thinsp;\u0026plusmn;\u0026thinsp;0.15\u0026permil;) reference materials. For every 12 samples, a laboratory reference-Gelatin from bovine skin (\u003cem\u003eδ\u003c/em\u003e\u003csup\u003e13\u003c/sup\u003eC\u003csub\u003eVPDB\u003c/sub\u003e = 14.7\u0026thinsp;\u0026plusmn;\u0026thinsp;0.2\u0026permil;; \u003cem\u003eδ\u003c/em\u003e\u003csup\u003e15\u003c/sup\u003eN\u003csub\u003eair\u003c/sub\u003e = 6.9\u0026thinsp;\u0026plusmn;\u0026thinsp;0.2\u0026permil;) was inserted for calibration and to monitor stability. The isotope results were analyzed as the ratio of the heavier isotope to the lighter isotope (\u003csup\u003e13\u003c/sup\u003eC/\u003csup\u003e12\u003c/sup\u003eC or \u003csup\u003e15\u003c/sup\u003eN/\u003csup\u003e14\u003c/sup\u003eN) and expressed as \u0026lsquo;\u003cem\u003eδ\u003c/em\u003e\u0026rsquo; in parts per 1000 or per mil (\u0026permil;) relative to internationally defined standards\u003csup\u003e[\u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e24\u003c/span\u003e]\u003c/sup\u003e for carbon (Vienna Pee Dee Belemnite, VPDB) and air nitrogen. The measurement errors were less than \u0026plusmn;\u0026thinsp;0.2\u0026permil; for both \u003cem\u003eδ\u003c/em\u003e\u003csup\u003e13\u003c/sup\u003eC and \u003cem\u003eδ\u003c/em\u003e\u003csup\u003e15\u003c/sup\u003eN values.\u003c/p\u003e \u003cp\u003eMoreover, existing \u003cem\u003eδ\u003c/em\u003e\u003csup\u003e13\u003c/sup\u003eC and \u003cem\u003eδ\u003c/em\u003e\u003csup\u003e15\u003c/sup\u003eN isotopic data from archaeological sites from the Lower Yangtze area around Shanghai before and during the Liangzhu period (5300\u0026thinsp;\u0026minus;\u0026thinsp;4300 cal. BP) (regional chronology of cultures and the development of rice agriculture shown in Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003e), were collected to further assess differences in dietary pattern between these peripheral populations and core communities living in the Lower Yangtze area (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). We searched for combinations of the search terms including \u0026lsquo;稳定同位素\u0026rsquo; /\u0026lsquo;stable isotope\u0026rsquo;, \u0026lsquo;食谱\u0026rsquo;/\u0026lsquo;diet/dietary pattern\u0026rsquo;, \u0026lsquo;新石器时代\u0026rsquo; /\u0026lsquo;Neolithic period\u0026rsquo; and \u0026lsquo;长江下游\u0026rsquo;/\u0026lsquo;Lower Yangtze\u0026rsquo; in Chinese and English literature using the China National Knowledge Infrastructure (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://www.cnki.net\u003c/span\u003e\u003cspan address=\"http://www.cnki.net\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e) and Google scholar, respectively. We extracted site information and relevant isotopic data from each relevent hit. Since some researchers only published scatter plots of stable carbon and nitrogen isotope values, we used software of GetData Graph Digitizer to capture the C, N isotopic values. The detailed information of these sites and data are listed in Table S2 and Table S3. Statistical data analysis was performed using the Student\u0026rsquo;s \u003cem\u003et\u003c/em\u003e-test using the Microsoft Office Excel, with the significance level set at \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e"},{"header":"Results","content":"\u003cdiv id=\"Sec6\" class=\"Section2\"\u003e \u003ch2\u003eIsotopic evidence of dietary patterns\u003c/h2\u003e \u003cp\u003eNew isotope data for Zhelin\u0026rsquo;s human and dog specimens (\u003cem\u003en\u003c/em\u003e\u0026thinsp;=\u0026thinsp;10) is listed in Table \u003cspan refid=\"MOESM1\" class=\"InternalRef\"\u003eS1\u003c/span\u003e. Due to the humid environment and acidic soils in Shanghai area, only two human specimens and one dog specimen produced collagen of suitable quality, with C:N ratios ranging between 2.9 to 3.6 for further analysis\u003csup\u003e[\u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e25\u003c/span\u003e, \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e26\u003c/span\u003e]\u003c/sup\u003e. Our \u003cem\u003eδ\u003c/em\u003e\u003csup\u003e13\u003c/sup\u003eC values for the human and dog samples (\u003cem\u003en\u003c/em\u003e\u0026thinsp;=\u0026thinsp;3) from Zhelin are \u0026minus;\u0026thinsp;19.2\u0026permil;, -19.4\u0026permil; and \u0026minus;\u0026thinsp;19.6\u0026permil; (mean\u0026thinsp;\u0026plusmn;\u0026thinsp;SD: -19.4\u0026thinsp;\u0026plusmn;\u0026thinsp;0.2\u0026permil;), respectively, indicative of similar diets significantly influenced by C\u003csub\u003e3\u003c/sub\u003e-based foods. The \u003cem\u003eδ\u003c/em\u003e\u003csup\u003e15\u003c/sup\u003eN values of these three specimens were +\u0026thinsp;9.1\u0026permil;, +\u0026thinsp;11.3\u0026permil; and +\u0026thinsp;8.6\u0026permil; (mean\u0026thinsp;\u0026plusmn;\u0026thinsp;SD: +9.7\u0026thinsp;\u0026plusmn;\u0026thinsp;1.2\u0026permil;), respectively, suggesting they consumed foods at different trophic levels. All new and existing stable C, N isotope data analyzed in the present study are listed in Table S2 and Table S3.\u003cdiv class=\"BlockQuote\"\u003e\u003cp\u003ePrevious studies have suggested that human bones are \u003csup\u003e15\u003c/sup\u003eN enriched by about 3\u0026ndash;5\u0026permil; relative to the foods they consume\u003csup\u003e[\u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e27\u003c/span\u003e]\u003c/sup\u003e. As can be seen Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e6\u003c/span\u003e and detailed in Table S3, carbon and nitrogen isotope values of charred plant samples (\u003cem\u003en\u003c/em\u003e\u0026thinsp;=\u0026thinsp;3) including the rice grains and acorns from the Tianluoshan site (7000\u0026thinsp;\u0026minus;\u0026thinsp;5300 cal. BP), located on the south bank of Hangzhou Bay, range from \u0026minus;\u0026thinsp;28.4\u0026permil; to -25.4\u0026permil; (mean\u0026thinsp;\u0026plusmn;\u0026thinsp;SD: -27.1\u0026thinsp;\u0026plusmn;\u0026thinsp;1.6\u0026permil;) and +\u0026thinsp;4.1\u0026permil; to +\u0026thinsp;7.4\u0026permil; (mean\u0026thinsp;\u0026plusmn;\u0026thinsp;SD: +5.3\u0026thinsp;\u0026plusmn;\u0026thinsp;1.9\u0026permil;), which are the lowest C and N isotopic values\u003csup\u003e[\u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e28\u003c/span\u003e]\u003c/sup\u003e, establishing an isotope baseline for primary producers in surrounding area. On the contrary, the highest \u003cem\u003eδ\u003c/em\u003e\u003csup\u003e13\u003c/sup\u003eC (mean\u0026thinsp;\u0026plusmn;\u0026thinsp;SD: -13.4\u0026thinsp;\u0026plusmn;\u0026thinsp;1.1\u0026permil;) and \u003cem\u003eδ\u003c/em\u003e\u003csup\u003e15\u003c/sup\u003eN values (mean\u0026thinsp;\u0026plusmn;\u0026thinsp;SD: +13.0\u0026thinsp;\u0026plusmn;\u0026thinsp;1.6\u0026permil;) derived tuna and whale bones (marine fish/mammal) from the Tianluoshan site\u003csup\u003e[\u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e29\u003c/span\u003e, \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e30\u003c/span\u003e]\u003c/sup\u003e, revealing the highest enriched \u003csup\u003e13\u003c/sup\u003eC and \u003csup\u003e15\u003c/sup\u003eN isotopic signals for marine animals in this coastal area.\u003c/p\u003e\u003c/div\u003e\u003c/p\u003e \u003cp\u003eAdditionally, the largest amount of previously published isotopic data derived from terrestrial herbivores (\u003cem\u003en\u003c/em\u003e\u0026thinsp;=\u0026thinsp;194) including several kinds of deer and bovid (including water buffalo) from the sites (Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e6\u003c/span\u003e) of Kuahuqiao (8000\u0026thinsp;\u0026minus;\u0026thinsp;7000 cal. BP), Tianluoshan and Meirendi (5300\u0026thinsp;\u0026minus;\u0026thinsp;4300 cal. BP), which ranges from \u0026minus;\u0026thinsp;24.8\u0026permil; to -10.7\u0026permil; (mean\u0026thinsp;\u0026plusmn;\u0026thinsp;SD: -17.8\u0026thinsp;\u0026plusmn;\u0026thinsp;3.9\u0026permil;) and +\u0026thinsp;3.3\u0026permil; to +\u0026thinsp;11.8\u0026permil; (mean\u0026thinsp;\u0026plusmn;\u0026thinsp;SD: +7.4\u0026thinsp;\u0026plusmn;\u0026thinsp;2.1\u0026permil;), suggesting that these likely enjoyed a broad spectrum of diets derive from \u003csup\u003e13\u003c/sup\u003eC-enriched plant foods\u003csup\u003e[\u003cspan additionalcitationids=\"CR30 CR31 CR32 CR33\" citationid=\"CR29\" class=\"CitationRef\"\u003e29\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e34\u003c/span\u003e]\u003c/sup\u003e. The second most abundant animal isotopic results belonged to terrestrial omnivores (\u003cem\u003en\u003c/em\u003e\u0026thinsp;=\u0026thinsp;139) dominated by pig and dog samples from Kuahuqiao, Jiangjiashan (7000\u0026thinsp;\u0026minus;\u0026thinsp;6000 cal. BP), Tianluoshan, Meirendi and our study site of Zhelin. The mean \u003cem\u003eδ\u003c/em\u003e\u003csup\u003e13\u003c/sup\u003eC and \u003cem\u003eδ\u003c/em\u003e\u003csup\u003e15\u003c/sup\u003eN values of these animals are \u0026minus;\u0026thinsp;20.6\u0026thinsp;\u0026plusmn;\u0026thinsp;1.2\u0026permil; and +\u0026thinsp;6.4\u0026thinsp;\u0026plusmn;\u0026thinsp;1.5\u0026permil;, indicating that omnivore diet in the studied area was largely influenced by low-protein C\u003csub\u003e3\u003c/sub\u003e terrestrial grasses and shrubs\u003csup\u003e[\u003cspan additionalcitationids=\"CR30 CR31 CR32 CR33\" citationid=\"CR29\" class=\"CitationRef\"\u003e29\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e34\u003c/span\u003e]\u003c/sup\u003e. The \u003cem\u003eδ\u003c/em\u003e\u003csup\u003e13\u003c/sup\u003eC and \u003cem\u003eδ\u003c/em\u003e\u003csup\u003e15\u003c/sup\u003eN values of freshwater animals including freshwater fish, frog, alligator, and turtle from Kuahuqiao, Tianluoshan and Meirendi ranged from \u0026minus;\u0026thinsp;23.5\u0026permil; to -18.8\u0026permil; (mean\u0026thinsp;\u0026plusmn;\u0026thinsp;SD: -20.9\u0026thinsp;\u0026plusmn;\u0026thinsp;1.4\u0026permil;) and +\u0026thinsp;5.6\u0026permil; to +\u0026thinsp;12.3\u0026permil; (mean\u0026thinsp;\u0026plusmn;\u0026thinsp;SD: +8.0\u0026thinsp;\u0026plusmn;\u0026thinsp;1.7\u0026permil;), the highest level for protein C\u003csub\u003e3\u003c/sub\u003e foods inland\u003csup\u003e[\u003cspan additionalcitationids=\"CR30 CR31\" citationid=\"CR29\" class=\"CitationRef\"\u003e29\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e32\u003c/span\u003e]\u003c/sup\u003e.\u003cdiv class=\"BlockQuote\"\u003e\u003cp\u003eAs for the isotopic data of human beings, plotted in Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e6\u003c/span\u003e, in the coastal area around Shanghai, the Songze human bone samples (\u003cem\u003en\u003c/em\u003e\u0026thinsp;=\u0026thinsp;2) during Songze culture period (6000\u0026thinsp;\u0026minus;\u0026thinsp;5300 cal. BP) exhibited \u003cem\u003eδ\u003c/em\u003e\u003csup\u003e13\u003c/sup\u003eC values of -20.2\u0026permil; and \u0026minus;\u0026thinsp;19.6\u0026permil; (mean\u0026thinsp;\u0026plusmn;\u0026thinsp;SD: -19.9\u0026thinsp;\u0026plusmn;\u0026thinsp;0.3\u0026permil;) and \u003cem\u003eδ\u003c/em\u003e\u003csup\u003e15\u003c/sup\u003eN values of +\u0026thinsp;9.7\u0026permil; and +\u0026thinsp;12.0\u0026permil; (+\u0026thinsp;10.9\u0026thinsp;\u0026plusmn;\u0026thinsp;1.2\u0026permil;) during 6000\u0026thinsp;\u0026minus;\u0026thinsp;5300 cal. BP\u003csup\u003e[\u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e35\u003c/span\u003e]\u003c/sup\u003e. This reveals that individuals largely relied on C\u003csub\u003e3\u003c/sub\u003e-based diets with varying trophic levels. In another small coastal settlement in the Ningshao area (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e), Tashan site, the mean carbon and nitrogen isotope values of humans during Liangzhu period (\u003cem\u003en\u003c/em\u003e\u0026thinsp;=\u0026thinsp;2) are \u0026minus;\u0026thinsp;18.7\u0026thinsp;\u0026plusmn;\u0026thinsp;0.3\u0026permil; and +\u0026thinsp;8.9\u0026thinsp;\u0026plusmn;\u0026thinsp;0.5\u0026permil; during 5300\u0026thinsp;\u0026minus;\u0026thinsp;4300 cal. BP\u003csup\u003e[\u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e36\u003c/span\u003e]\u003c/sup\u003e. Dating before that time, one individual from the Tashan site during 5900\u0026thinsp;\u0026minus;\u0026thinsp;5600 cal. BP before Liangzhu culture period yielded \u003csup\u003e13\u003c/sup\u003eC and \u003csup\u003e15\u003c/sup\u003eN-enriched results (\u003cem\u003eδ\u003c/em\u003e\u003csup\u003e13\u003c/sup\u003eC = -18.0\u0026permil; and \u003cem\u003eδ\u003c/em\u003e\u003csup\u003e15\u003c/sup\u003eN\u0026thinsp;=\u0026thinsp;+\u0026thinsp;9.8\u0026permil;). Similarly, in the same area of Ningshao, the mean carbon and nitrogen isotope values of the Hemudu sample (\u003cem\u003en\u003c/em\u003e\u0026thinsp;=\u0026thinsp;2) during 7000\u0026thinsp;\u0026minus;\u0026thinsp;6000 cal. BP were \u0026minus;\u0026thinsp;16.7\u0026thinsp;\u0026plusmn;\u0026thinsp;1.6\u0026permil; and +\u0026thinsp;11.4\u0026thinsp;\u0026plusmn;\u0026thinsp;0.2\u0026permil;, respectively\u003csup\u003e[\u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e35\u003c/span\u003e]\u003c/sup\u003e. However, the \u003cem\u003eδ\u003c/em\u003e\u003csup\u003e13\u003c/sup\u003eC and \u003cem\u003eδ\u003c/em\u003e\u003csup\u003e15\u003c/sup\u003eN values of human bone samples (\u003cem\u003en\u003c/em\u003e\u0026thinsp;=\u0026thinsp;20) from another Hemudu culture site of Tianluoshan ranged from \u0026minus;\u0026thinsp;21.9\u0026permil; to -20.1\u0026permil; (mean\u0026thinsp;\u0026plusmn;\u0026thinsp;SD: -20.6\u0026thinsp;\u0026plusmn;\u0026thinsp;0.5\u0026permil;) and +\u0026thinsp;7.2\u0026permil; to +\u0026thinsp;10.7\u0026permil; (mean\u0026thinsp;\u0026plusmn;\u0026thinsp;SD: +8.9\u0026thinsp;\u0026plusmn;\u0026thinsp;0.9\u0026permil;) during 7000\u0026thinsp;\u0026minus;\u0026thinsp;5300 cal. BP\u003csup\u003e[\u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e29\u003c/span\u003e, \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e33\u003c/span\u003e]\u003c/sup\u003e.\u003c/p\u003e\u003c/div\u003e\u003c/p\u003e \u003cp\u003eAs for the isotopic evidence for diets in these inland sites at Lower Yangtze area prior to and during the Liangzhu period (5300\u0026thinsp;\u0026minus;\u0026thinsp;4300 cal. BP), carbon and nitrogen isotope values of Jiangjiashan people (\u003cem\u003en\u003c/em\u003e\u0026thinsp;=\u0026thinsp;1) living on the southwestern bank of Taihu Lake during 7000\u0026thinsp;\u0026minus;\u0026thinsp;6000 cal. BP (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e), were \u0026minus;\u0026thinsp;20.5\u0026permil; and +\u0026thinsp;10.2\u0026permil;., respectively (Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e6\u003c/span\u003e). According to Hu et al (2007)\u003csup\u003e[\u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e37\u003c/span\u003e]\u003c/sup\u003e, the carbon and nitrogen isotope values of Sanxingcun peoples (\u003cem\u003en\u003c/em\u003e\u0026thinsp;=\u0026thinsp;19) ranges from \u0026minus;\u0026thinsp;20.4\u0026permil; to -19.6\u0026permil; (mean\u0026thinsp;\u0026plusmn;\u0026thinsp;SD: -20.0\u0026thinsp;\u0026plusmn;\u0026thinsp;0.2\u0026permil;) and +\u0026thinsp;8.9\u0026permil; to +\u0026thinsp;10.3\u0026permil; (mean\u0026thinsp;\u0026plusmn;\u0026thinsp;SD: +9.7\u0026thinsp;\u0026plusmn;\u0026thinsp;0.3\u0026permil;) indicative of a C\u003csub\u003e3\u003c/sub\u003e-based diet ate by these individuals. Given that the Sanxingcun site is located in the north of Taihu Lake at the Lower Yangtze area, where an unbroken rice agriculture had been practiced early Holocene period\u003csup\u003e[\u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e4\u003c/span\u003e, \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e38\u003c/span\u003e]\u003c/sup\u003e, combined with the isotopic evidence of Sanxingcun people\u0026rsquo;s trophic level, it is safe to believe that the humans of Sanxingcun mainly fed on rice and terrestrial omnivores as well as a small number of terrestrial herbivores during the 6500\u0026thinsp;\u0026minus;\u0026thinsp;5500 cal. BP period. In addition, during the Liangzhu period (5300\u0026thinsp;\u0026minus;\u0026thinsp;4300 cal. BP), Liangzhu resident (\u003cem\u003en\u003c/em\u003e\u0026thinsp;=\u0026thinsp;9) \u003cem\u003eδ\u003c/em\u003e\u003csup\u003e13\u003c/sup\u003eC and \u003cem\u003eδ\u003c/em\u003e\u003csup\u003e15\u003c/sup\u003eN values recovered from the Liangzhu City area (Meirendi site) ranged from \u0026minus;\u0026thinsp;22.2\u0026permil; to -17.4\u0026permil; (mean\u0026thinsp;\u0026plusmn;\u0026thinsp;SD: -19.9\u0026thinsp;\u0026plusmn;\u0026thinsp;1.2\u0026permil;) and +\u0026thinsp;9.7\u0026permil; to +\u0026thinsp;11.0\u0026permil; (mean\u0026thinsp;\u0026plusmn;\u0026thinsp;SD: +10.3\u0026thinsp;\u0026plusmn;\u0026thinsp;0.4\u0026permil;), which is indicative of a C\u003csub\u003e3\u003c/sub\u003e-based diet with relatively high trophic levels.\u003c/p\u003e \u003c/div\u003e"},{"header":"Discussion","content":"\u003cp\u003e \u003cdiv class=\"BlockQuote\"\u003e \u003cp\u003eJust as at Liangzhu\u003csup\u003e[\u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e1\u003c/span\u003e]\u003c/sup\u003e, archaeological work around the world often prioritizes the great sites. However, to better understand the past and the role that these large sites had to play in shaping neighboring settlements, archaeologists should obviously also pay attention to these smaller sites. We argue that the recent studies at core inland Liangzhu sites can only be comprehensively understood within a broader regional context and with regard to peripheral coastal sites at Lower Yangtze area. Previous work by Dong and Yuan has suggested that existing zooarchaeological findings at the Liangzhu cultural area reveal different animal husbandry practices between the urban center of Mojiaoshan and small sized sites during Liangzhu period (5300\u0026thinsp;\u0026minus;\u0026thinsp;4300 cal. BP)\u003csup\u003e[\u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e18\u003c/span\u003e]\u003c/sup\u003e. In the present study, we first attempted to investigate previously unexamined dietary practices of Liangzhu cultural human societies at Shanghai area, as a geographic periphery of the Liangzhu core. More importantly, by integrating new and existing C, N isotopic data of humans (\u003cem\u003en\u003c/em\u003e\u0026thinsp;=\u0026thinsp;58), animals (\u003cem\u003en\u003c/em\u003e\u0026thinsp;=\u0026thinsp;362) and plants (\u003cem\u003en\u003c/em\u003e\u0026thinsp;=\u0026thinsp;3) recovered from sites around the Lower Yangtze area during 8000\u0026thinsp;\u0026minus;\u0026thinsp;4300 cal. BP, we provided a broad picture of the dynamic changes of diets and economic decisions of these coastal communities living at Shanghai and Ningshao areas before and during the Liangzhu phenomenon.\u003c/p\u003e \u003c/div\u003e \u003c/p\u003e \u003cp\u003eAlthough rice agriculture began around 10000 cal. BP in the Lower Yangtze area\u003csup\u003e[\u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e38\u003c/span\u003e]\u003c/sup\u003e, archaeological evidence suggests that rice agricultural production in the region developed over thousands of years until before becoming more stabilized during the middle Songze culture through Liangzhu periods (5600\u0026thinsp;\u0026minus;\u0026thinsp;4300 cal. BP)\u003csup\u003e[\u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e39\u003c/span\u003e]\u003c/sup\u003e. Prior to this time, ancient humans may have consumed more wild animals and nuts for survival\u003csup\u003e[\u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e40\u003c/span\u003e]\u003c/sup\u003e. As shown in Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e6\u003c/span\u003e, in the coastal area of Ningshao before Liangzhu time, stable carbon and nitrogen isotope evidence from the Hemudu site and Tianluoshan site suggests that diversified human subsistence, like exploitation of wild plants, seawater and freshwater resources as well as wild games, sustained a continuous parallel development with low-level rice farming for millennia. In another coastal area of Shanghai, \u003cem\u003eδ\u003c/em\u003e\u003csup\u003e15\u003c/sup\u003eN results for human bones recovered from the Songze site, 6000\u0026thinsp;\u0026minus;\u0026thinsp;5300 cal. BP, yield an increase in \u003cem\u003eδ\u003c/em\u003e\u003csup\u003e15\u003c/sup\u003eN values compared to the inland residents of Jiangjiashan (7000\u0026thinsp;\u0026minus;\u0026thinsp;6000 cal. BP) and Sanxingcun (6500\u0026thinsp;\u0026minus;\u0026thinsp;5500 cal. BP) (student\u0026rsquo;s \u003cem\u003et\u003c/em\u003e-test, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.0056). \u003csup\u003e15\u003c/sup\u003eN enrichment levels in the Songze peoples (6000\u0026thinsp;\u0026minus;\u0026thinsp;5300 cal. BP) who lived in the coastal area of Shanghai at nearly the same time consumed a greater amount of freshwater foods. One important finding is that although different groups of Neolithic populations in Lower Yangtze region from 7000\u0026thinsp;\u0026minus;\u0026thinsp;5300 cal. BP have similar C\u003csub\u003e3\u003c/sub\u003e-based diets which were probably attributed to low-level rice farming and foraging of wild plant resources at large, according to varies of stable nitrogen values and zooarchaeological evidence from the study area\u003csup\u003e[\u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e18\u003c/span\u003e]\u003c/sup\u003e, there are significant differences in foodways between the populations who lived in the inland area (Sanxingcun, Jiangjiashan) and other coastal (Hemudu, Tianluoshan, Songze, Tashan) settlements (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e, \u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e6\u003c/span\u003e). This also shows that the food-related differentiation had appeared in inland and coastal societies at the Lower Yangtze River area during 7000\u0026thinsp;\u0026minus;\u0026thinsp;5300 cal. BP.\u003c/p\u003e \u003cp\u003eAccording to previous archaeobotanical studies and archaeological finding of complex water management systems at ancient Liangzhu City\u003csup\u003e[\u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2\u003c/span\u003e]\u003c/sup\u003e, during the Liangzhu period rice agriculture rapidly developed and rice foods eventually became a staple food incorporated into diets across the Lower Yangtze area\u003csup\u003e[\u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e4\u003c/span\u003e, \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e10\u003c/span\u003e]\u003c/sup\u003e. At this time, more sophisticated water management and farming techniques allowed the majority of individuals in the core of the Liangzhu area had a relatively balanced diet including a large number of domesticated rice, and meat from domesticates like pigs and some wild games including a number of freshwater resources as well. We had compared the \u003cem\u003eδ\u003c/em\u003e\u003csup\u003e13\u003c/sup\u003eC and \u003cem\u003eδ\u003c/em\u003e\u003csup\u003e15\u003c/sup\u003eN values of human samples from the sites of Sanxingcun and Liangzhu City (Meirendi site). Our analysis reveals that both inland communities shared a similar C\u003csub\u003e3\u003c/sub\u003e-based dietary pattern, while the core Liangzhu culture humans from the Meirendi site had relatively high trophic levels than Sanxingcun peoples based on N isotopic evidence (student\u0026rsquo;s \u003cem\u003et\u003c/em\u003e-test, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.0005).\u003c/p\u003e \u003cp\u003eAs for the foodways of human beings living at the peripheral coastal areas during Liangzhu period (5300\u0026thinsp;\u0026minus;\u0026thinsp;4300 cal. BP), previous work by Zhang et al. (2015)\u003csup\u003e[\u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e36\u003c/span\u003e]\u003c/sup\u003e, for example, has reported that Tashan people, occupying a small East China Sea coastal settlement in the period 5300\u0026thinsp;\u0026minus;\u0026thinsp;4300 cal. BP, likely exploited more foods from rice and terrestrial animals but less marine food resources than the residents of the same site for the period 5900\u0026thinsp;\u0026minus;\u0026thinsp;5600 cal. BP. Also, it can be seen that the Hemudu residents living in the Ningshao area during 7000\u0026thinsp;\u0026minus;\u0026thinsp;6000 cal. BP had higher stable nitrogen isotopic values than Tashan peoples during 5300\u0026thinsp;\u0026minus;\u0026thinsp;4300 cal. BP (student\u0026rsquo;s \u003cem\u003et\u003c/em\u003e-test, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.0434). Considering that large-scale rice paddies have been found at the Shiao site in Ningshao area directly dating to the Hemudu culture and the Liangzhu culture periods\u003csup\u003e[\u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e41\u003c/span\u003e]\u003c/sup\u003e, we believe that the changes of diets and consumption habits of these peripheral populations at Tashan during the Liangzhu period were likely influenced by the expansion of massive rice cultivation in the Ningshao area from the Hemudu culture to Liangzhu culture periods (7000\u0026thinsp;\u0026minus;\u0026thinsp;4300 cal. BP). Nevertheless, we have also found that the \u003cem\u003eδ\u003c/em\u003e\u003csup\u003e15\u003c/sup\u003eN values of humans of Tashan lower than those of the Liangzhu City during the same period (student\u0026rsquo;s \u003cem\u003et\u003c/em\u003e-test, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.0052). This suggests that despite communication with the core Liangzhu culture, the Tashan societies reorganized their own sense of cultural identity along dietary lines and made specialized economic decisions to participate in a new social network, influenced by inland Liangzhu culture.\u003c/p\u003e \u003cp\u003eMoreover, as can be seen in Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e6\u003c/span\u003e, our new isotopic data suggests that the dietary lifestyle of the Zhelin people, living on a small Liangzhu site in the coastal area of the Lower Yangtze, was more similar to that of people from the core area of the Liangzhu culture compared to human beings from the Tashan site at Ningshao area during the Liangzhu period. Specifically, from Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e6\u003c/span\u003e, the \u003cem\u003eδ\u003c/em\u003e\u003csup\u003e13\u003c/sup\u003eC (student\u0026rsquo;s \u003cem\u003et\u003c/em\u003e-test, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.5334) and the \u003cem\u003eδ\u003c/em\u003e\u003csup\u003e15\u003c/sup\u003eN (student\u0026rsquo;s \u003cem\u003et\u003c/em\u003e-test, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.8691) values of humans at the Zhelin site and the inland Liangzhu core (Meirendi site) express no clear differentiation. Mean \u003cem\u003eδ\u003c/em\u003e\u003csup\u003e15\u003c/sup\u003eN values at Zhelin are, however, higher than those of the Tashan individuals during the same period (Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e6\u003c/span\u003e). Moreover, the diet of Zhelin remains largely consistent to previous dietary pattern of humans at the Songze site during 6000\u0026thinsp;\u0026minus;\u0026thinsp;5200 cal. BP (\u003cem\u003eδ\u003c/em\u003e\u003csup\u003e13\u003c/sup\u003eC: student\u0026rsquo;s \u003cem\u003et\u003c/em\u003e-test, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.1982; \u003cem\u003eδ\u003c/em\u003e\u003csup\u003e15\u003c/sup\u003eN: student\u0026rsquo;s \u003cem\u003et\u003c/em\u003e-test, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.7225). Interestingly, the mean \u003cem\u003eδ\u003c/em\u003e\u003csup\u003e15\u003c/sup\u003eN value of Zhelin humans decreased compared to inhabitants of Songze in nearby region prior to Liangzhu period (Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e6\u003c/span\u003e), perhaps suggesting a general decrease in the consumption of high-protein foods, such as aquatic meats, as subsistence economies became more agrarian during the Liangzhu period (5300\u0026thinsp;\u0026minus;\u0026thinsp;4300 cal. BP). In these cases, we argue that the economic decisions of the Zhelin people may be more dominated by Liangzhu core concerns, expressing a recognition of inland dietary lifeways of the core area of Liangzhu culture.\u003c/p\u003e \u003cp\u003ePrevious studies have suggested that intensive rice agriculture has underpinned increased human populations and massive social evolution at the Lower Yangtze area during the Liangzhu period (5300\u0026thinsp;\u0026minus;\u0026thinsp;4300 cal. BP) through a rapid expansion of wetland under rice cultivation and a greater reliance on domesticated foods like pigs and water buffalo\u003csup\u003e[3\u0026ndash;4,18\u0026minus;19,42\u0026ndash;43]\u003c/sup\u003e. As such, we believe that the diets and consumption habits of these peripheral Liangzhu populations at Zhelin and Tashan, perhaps influenced, to some extent, by their desired food choices under the rapid development of early urbanized and agricultural lifestyle originating in the core Liangzhu area. The differentiation in dietary practices between the populations from the coastal Zhelin and Tashan sites, further suggests that the two societies living at the periphery of the Liangzhu cultural sphere may have taken on different roles to integrated into a new social network of Liangzhu culture. In the long run, this transition to a more standardized man-made agricultural economic choice may have inadvertently locked Liangzhu society into a system that depended on a limited range of \u0026ldquo;taxable\u0026rdquo; domesticated species\u003csup\u003e[\u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e44\u003c/span\u003e]\u003c/sup\u003e, which may have been significantly affected by rapid climate changes at around 4200 cal. BP.\u003c/p\u003e"},{"header":"Conclusion","content":"\u003cp\u003eBy integrating new C, N isotopes from Zhelin, Shanghai with previous isotopic studies done in the Taihu Basin and Ningshao areas during 8000\u0026thinsp;\u0026minus;\u0026thinsp;4300 cal. BP, we used the isotopic evidence to gain a better understanding of the dietary and culinary changes at the Lower Yangtze area prior to and during the Liangzhu cultural period (5300\u0026thinsp;\u0026minus;\u0026thinsp;4300 cal. BP). Our findings suggest that the dietary differences between the coastal periphery and more inland core cultural area of Lower Yangtze area likely existed through the Liangzhu period. Moreover, our study shows that the Liangzhu dependence on limited domesticated victuals became much more widespread around Lower Yangtze area, and more socially integrated terrestrial economic decisions appeared along with the development of Liangzhu cultural phenomenon in this coastal region. This study provides essential data for understanding the development of the Liangzhu cultural phenomenon at Lower Yangtze area in a more comprehensive manner. However, more C, N isotopic research is necessary for more robust comparisons between the developmental trajectory of foodways across different societies in the coastal reaches of Neolithic southeastern China.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eAcknowledgements\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe authors are grateful to the\u0026nbsp;funds from Shanghai Planning Office of Philosophy and Social Science\u0026nbsp;(2019ELS006) for financial support. We are thankful to\u0026nbsp;Prof.\u0026nbsp;Yi Guo at\u0026nbsp;Zhejiang University\u0026nbsp;and Dr Simei Zhu at Zhengzhou University\u0026nbsp;for\u0026nbsp;their\u0026nbsp;help with the pre-treatment experiment\u0026nbsp;of bone\u0026nbsp;samples.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003e\u003cstrong\u003eAuthor contributions\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eConceptualization: P.S., E.A., M.S.; methodology: P.S., M.S.; Investigation: P.S., X.Z., X.H., M.S.; Funding Acquisition: P.S.; Project Administration: P.S.; Writing\u0026mdash;original draft: P.S., E.A., M.S.; Writing\u0026mdash;review \u0026amp; editing: P.S., E.A., M.S. All authors contributed to the article and approved the submitted version.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003e\u003cstrong\u003eCompeting Interests\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe authors declare no competing interests.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n\u003cli\u003eRenfrew, C. \u0026amp; Liu, B. 2018. The emergence of complex society in China: the case of Liangzhu. Antiquity, 92(364): 975-90.\u003c/li\u003e\n\u003cli\u003eLiu, B., Wang, N., Chen, M., Wu, X., Mo, D., Liu, J., Xu, S. and Zhuang, Y. 2017. Earliest hydraulic enterprise in China, 5,100 years ago. Proceedings of the National Academy of Sciences, 114(52): 13637-13642. \u003c/li\u003e\n\u003cli\u003eQin, L. 2013. The Liangzhu culture. In A. P. Underhill (Ed.), A Companion to Chinese Archaeology (pp. 574-596). Blackwell Companions to Anthropology. Malden, MA: Wiley-Blackwell Press.\u003c/li\u003e\n\u003cli\u003eFuller, D.Q. 2020. 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New Haven : Yale University Press.\u003c/li\u003e\n\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":true,"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":"Diet, Stable isotope, Rice agriculture, Lower Yangtze area, Liangzhu","lastPublishedDoi":"10.21203/rs.3.rs-4185222/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-4185222/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eThis essay analyzes a robust body of C and N stable isotopes from human and animal collagen and plant remains (\u003cem\u003en\u003c/em\u003e\u0026thinsp;=\u0026thinsp;423) in the area of Lower Yangtze River dating to 8000\u0026thinsp;\u0026minus;\u0026thinsp;4300 cal. BP, combined with new isotopic data recovered from the Zhelin site in Shanghai, to explore the dietary differences between coastal region the inland core area of the Liangzhu culture (5300\u0026thinsp;\u0026minus;\u0026thinsp;4300 cal. BP). Our results reveal that the periphery dwellers of the Liangzhu cultural area became increasingly reliant on a limited number of domesticated species over time, while maintaining a marked social differentiation of foodways that was potentially a result of the growing but unequal influence of populations from the core Liangzhu area.\u003c/p\u003e","manuscriptTitle":"To eat like Liangzhu: isotopic investigation of diets in the Lower Yangtze area prior to and during the Liangzhu period (5300-4300 cal. 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